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SMITHSONIAN
MISCELLANEOUS COLLECTIONS
VOL. 103
“EVERY MAN IS A VALUABLE MEMBER OF SOCIETY WHO, BY HIS OBSERVATIONS, RESEARCHES,
AND EXPERIMENTS, PROCURES KNOWLEDGE FOR MEN *’—SMITHSON
(PUBLICATION 3731)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
1943
The Lord Baltimore Press
BALTIMORE, MD., U. 8 A.
ee
=
1s
ADVERTISEMENT
The Smithsonian Miscellaneous Collections series contains, since the
suspension in 1916 of the Smithsonian Contributions to Knowledge,
all the publications of the Institution except the Annual Report, the
annual volume describing the Institution’s field work, and occasional
publications of a special nature. As the name of the series implies, its
scope is not limited, and the volumes thus far issued relate to nearly
every branch of science. Papers in the fields of biology, geology,
anthropology, and astrophysics have predominated.
C. G. Asrot,
Secretary of the Smithsonian Institution.
(ii1)
i |
“Ny ae 7 , Py
a a ay
to
10,
CONTENTS
. WE cH, D’ALTE A. Distribution and variation of the Hawaiian
tree snail Achatinella apexfulva Dixon in the Koolau Range,
aha. 236 pp., 12 pls., 8 figs., Dec..16, 1942. (Publ. 3684.)
Snopcrass, R. E. The skeleto-muscular mechanisms of the
honey bee. 120 pp., 32 figs., Sept. 30, 1942. (Publ. 3688.)
Dertcnan, H. G. A revision of the Indo-Chinese forms of the
avian genus Prinia. 12 pp., Sept. I, 1942. (Publ. 3689.)
Roperts, FranK H. H., Jr. Archeological and geological in-
vestigations in the San Jon district, eastern New Mexico.
BO pp. 6 plsi,. 3) figs., Oct. 12, 1942. (Publ. 3692.)
ReEsseR, CHarLtes E. New Upper Cambrian trilobites. 136 pp.,
ai pls... (Oct. 21, 1942. (Publ. 3693.)
VALENTINE, J. MANson. On the preparation and preservation
of insects, with particular reference to Coleoptera. 16 pp.,
5 figs., Nov. 21, 1942. (Publ. 3696.)
Dorsey, CARL Kester. The musculature of the labrum, labium,
and pharyngeal region of adult and immature Coleoptera.
42 pp., 24 pls., Jan. 20, 1943. (Publ. 3697.)
AxpotT, C. G. The 1914 tests of the Langley “aerodrome.” 8 pp.,
ariies., Oct. 24, 1942. (Publ. 36099.)
PENNAK, RoBert W., and Zinn, Donatp J. Mystacocarida,
a new order of Crustacea from intertidal beaches in Massachu-
Bettsnand,) Gonnecticut, 11 pp, 2 pls.,-Feb, 23, 1943. (Publ:
3704.)
Kutimer, C. J. A remarkable reversal in the diStribution of
storm frequency in the United States in double Hale solar
cycles, of interest in long-range forecasting. 20 pp., 19 figs.,
10 storm-frequency year maps, Apr. 5, 1943. (Publ. 3729.)
(v)
“ie 5
mat iA
CG eae
oe Van
iv
i
i
i
a re
sv;
Hy 70 i ; x ; ,
Ri A es amy
HDA
Liege)
_ VOLUME 103, NUMBER 1:
_ KOOLAU RANGE, OAHU
we 12: Poates)
| ‘BY By
- DIALTE A. WELGH
(Pp UBLICATION 3684)
CITY OF WASHINGTON
‘ _DECEMBER 16, 1942
"SWITHSONIAN MISCELLANEOUS COLLECTIONS
d. DISTRIBUTION AND VARIATION OF THE.
\ HAWAIIAN TREE. SNAIL. ACHATINELLA
ey _APEXFULVA. DIXON” IN THe
sont PUBLISHED BY THE SMITHSONIAN INSTITUTION
if Ds
a S Mealy wh tn
lg ig Sa Anat
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 103, NUMBER 1
DISTRIBUTION AND VARIATION OF THE
HAWAIIAN TREE SNAIL ACHATINELLA
SeexPULYA DIXON IN THE
KOOLAU RANGE, OAHU
(WitTH 12 PLaTEs)
BY
D’ALTE A. WELCH
(PUBLICATION 3684)
GITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
DECEMBER 16, 1942
The Lord Baltimore Press
BALTIMORE, MD., U. S. As
DISTRIBUTION AND VARIATION OF THE HAWAIIAN
TREE SNAIL ACHATINELLA APEXFULVA DIXON
IN THE KOOLAU RANGE, OAHU
By p ALTE A. WELCH
(WirH 12 PrLates)
CONTENTS
PaGE
Wena INS Lit OTT MEPS cree ore cts pe ter ore te as: yc 1eis @, 4, Ais te ctus ceeho ty onan cleheGroncncrcushindh ones ads Noes I
PARE RTI WI OB arte ster: eevee hs Sl eet cera ee ue Nae mare ced a tide I
Bincemamesiorthe Moola Ranges... je sree ok ee ooo ation oe 2
Shimatolocicalmdatasands physiocraphiy.s eas ee nie eae eee ee 9
Table 1. Annual precipitation records in the Koolau Range...... 10
MctLetial man eIne MOUS ae. idols ate ee ecm ein ee eee hea iene Se aoe 12
ater se Dataom material .o.1<cee es me eset. 2. Sera Mah oases cae 15
SMS, CONGANE Bsus oh O SOIREE NiguT Sere AE ale 21
BRS UO AA Varner ects foro scolcye. oreo tapsieos @) «1, </ oy tele eating Sasso: Sele uel speistel sie rore Gees a ticlope 23
Synopsis of the subspecies of Achatinella apexfulva Dixon............. 23
SLOUpP Ole An el StUS IN EWCOMID: Se aegc tosses a esinatice tase ee soar 28
Groupeoniese as uiblata Reeves nse seer ie le hae ae aoe ae 2
Geouprok Aig. pilsbryi new! subspecies 0.0 0.6 cece sacs wame ses 52
Croup OLA a iurgidd = Neweomb: siiwcivesccee sconaaee ouelcieein< 57
Group) at A. a). polymorpha Gulick.. 2.03 s..s%0 25 vos Se eepe s Foo eon OI
Groupeot Asa, leucorraphe, Gulick’.2% 22% acc. cs ae dacencer sans IOI
Group ot A. a, wwint Pilsbry and Cooke l.....2..2.......25-+4- 115
CroupEOneAs as CONifOrmis: GUlCK Aye sae tice anette bio 126
GMI ROL eA, aanusracamn. Ganlichs i sd. cine Sloe samc de veya emda PORES 133
EON TOL As i. aprcata\ INE@w COM ioes.s-3 visio ss Gusts Saisie ale he viata sta 88 146
Gcoupinor.4.. a aiona Pilsbry and’ Cooke.0¢ ssc 22.0.8 ee sats 160
Catetipratts aanerpulve Dixoti si.) oe gss en Me oe wie) oa eaten 176
es SIE Ae Ra yt le oa bre aide al SOMES Eo aged SOs Sidhe d cos We clare 188
Sd GIA LIIC) Sah MMT RTC ern oars aes arses ese pe amtairas eb css Ree Meger ove, Senetta-nl Oe mise aa eee 206
RNC MCEG MRE Veter tote scl ays) aig le.0 casera sim evra san iaten ra trier ieae veguebr ae 208
Hoerape etl AtLO MO Tee AL@Stewsiehe eieesiere le cai c's. am e¥e cicasie ar onalsiain! stave cnsta aol eich Nieyo sya erste stots 208
Ete Meee OMe rte PTS terse fics 4s. cya dai seiees alate Gonaie eam tote elegeron a Soares 22
fidex fo sections, genera, species, and subspecies... .........-2.0.0.5.ce005 225
MS MaRL OUP LACE ANcITICS het eciet ielereies ic Sere eos che ois oeledciet ee cee tere ewes kiaey ee ene 229
INTRODUCTION
SCOPE OF
WORK
In a previous paper (Welch, 1938) the species Achatinella mustelina
is studied. The present paper continues the work on Hawaiian tree
snails and deals with the species Achatinella apexfulva, also of the
island of Oahu. Does the same variation or series of variations occur
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 103, No. 1
bo
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 103
in A. apexfulva of the Koolau Range as in A. mustelina of the
Waianae Mountains?
During the 7 years I spent in Hawaii studying Achatinella I was
greatly helped in acquiring a background for the present work by
Dr. C. M. Cooke, Jr. Working under Dr. Cooke, I also learned
many methods of studying and preparing material which greatly
facilitated my work. I am also indebted to Dr. E. A. Andrews,
Dr. Henry A. Pilsbry, Dr. H. B. Baker, and Dr. Ernst Mayr for
criticisms and advice in the preparation of the manuscript. Dr. F. Ray-
mond Fosberg was most helpful in identifying all genera of Hawaiian
plant names given by various collectors of shells. I also wish to
express appreciation to the following for various courtesies and
favors: Dr. Paul Bartsch, W. J. Clench, W. T. Calman, Col. A. J.
Peile, J. R. le B. Tomlin, G. S. Robson, Dr. Paul Galstoff, Dr. Gilbert
D. Harris, Dr. C. E: McClung, Dr. Peter H. Buck, Dr. Herbert
Gregory, the Trustees of the Bernice P. Bishop Museum, and the au-
thorities of the United States National Museum.
The ground work of this paper was done at the Bernice P. Bishop
Museum in Honolulu, the writing of the paper was carried on at
the United States Bureau of Fisheries at Woods Hole, the Zoological
Laboratory of the University of Pennsylvania, the Zoological De-
partment of the Johns Hopkins University, and the Zoological
Department of Barat College, Lake Forest, Ill. I wish to thank these
institutions for granting me facilities for work.
PLACE NAMES OF THE KOOLAU RANGE
Most of the names of the valleys or gulches of the Koolau Range
are well known, but the names of some have never appeared on a
Government map. Other gulch names have not been known, and
still other gulches, although their names are well known, contain large
subgulches that need to be designated by some name in order that
localities may be described exactly.
Pilsbry and Cooke (1912-1914, p. 277) published a rough dia-
grammatic sketch map of the northwestern half of the Koolau Range
in which they included the new names Kalaikoa, Ahonui, Main
Poamoho, Central Poamoho, West Poamoho, Kawaihalona, and
Luapo. This map was compiled from field notes made by Irwin
Spalding and is the first published map showing the approximate
regions occupied by colonies of Achatinella. Many of these names
do not exist on the advance sheets of the United States Geological
Survey topographic sheets of Oahu, Hawaiian Islands, 1/20,000,
which form the standard map I used for plotting all localities of
PO. OE ACHATINELLA APEXFULVA DIXON—WELCH 3
Achatinella and from which all maps published in this paper have
been compiled. Therefore a considerable amount of work has been
done to check the place names on the Pilsbry-Cooke map and to
ascertain the place names of all valleys not yet published.
I am indebted to the following people for help in this work: The
Bishop Estate for access to maps, Miss Jane L. Winne for place
ISLAND
OF
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PLANT ZONES A\Waimanolo
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D OHIA
E CLOUD
iG. t—Island of Oahu, showing the plant zones in Kipapa Gulch (after Hosaka) and the location
of rain-gage stations in the Koolau Range. Ridges outlined in solid lines.
names obtained from an old Hawaiian native to the district of
Maunalua; Manuel Baptista, a former cattle ranger native to the
districts of Kalauao and Waiawa; G. D. Robinson; William Holt, a
former cattle ranger in the regions of Wahiawa and Paala; Benny
Pulaa, a Hawaiian native to the districts of Kawailoa, Waimea, and
the northern section of Oahu.
The Koolau Range is divided into six large land grants or districts
which are numbered A to F on figure 2, p. 4. Within these large
103
VOL.
MISCELLANEOUS COLLECTIONS
SMITHSONIAN
“RTA Ul paurjjno saspry ‘osuey nejooy ay} Jo soweu aoejd oy} 0} deus Aay—z ‘OI
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NO. I ACHATINELLA APEXFULVA DIXON—WELCH 5
districts are smaller districts or subdistricts indicated by Roman
numerals, which may include a single valley or gulch with the same
name or several valleys with different names. The valleys or gulches
within a subdistrict are numbered in Arabic. In the discussion of
place names, all names not published on the United States Geological
Survey topographic advance sheets of the island of Oahu will have
in parentheses the names of the authorities after them. Main valleys
or gulches are often subdivided by the author for convenience into
north, south, east, west, or central. These main valley subdivisions
may be broken up again into north or south branches.
Figures 1 and 3-6 were drawn by Gordon Brett Littlepage from
_maps compiled by me. Figures 2, 7, 8 were drawn by me. All maps
are compiled from the United States Geological Survey topographic
advance sheets of the island of Oahu. Ridges on all maps are repre-
sented by lines.
Place names of the Koolau Range
A. Honolulu District.
IT. Maunalua.
1, Kealakipapa Valley; 2, Mauuwaii Valley; 3, Kalama Valley; 4,
Kamiloiki Valley; 5, Kamilonui Valley; 6, Hahaione Valley; 7,
Kaalakei Valley (Bishop Estate maps: J. Winne).
II. Kuliouou.
I, Kuliouou Gulch; 1a, East Branch (Welch); 1b, West Branch
(Welch) ; 2, Puu O Kona.
III. Niu.
1, East Niu Gulch; 2, West Niu Gulch. (1-2 Welch.)
IV. Wailupe.
1, Kului Gulch; 2, Laulaupoe Gulch; 3, Wailupe Gulch.
V. Waialae Iki.
1, Kapakahi Gulch (in this paper Waialae Iki Gulch will be used in
preference to Kapakahi Gulch).
VI. Waialae Nui.
1, Waialae Nui Gulch (spelled Waialaenui on U. S. Geol. Surv. topogr.
advance sheets).
VII. Palolo.
1, Waiomao Stream; 2, Pukele Stream.
VIII. Manoa.
1, Manoa Stream; 2, East Manoa Stream (Welch).
IX. Makiki. (This name is not on the U. S. Geol. Surv. topogr. advance
sheets, but it is a well-known name and has been in use by shell
collectors for many years.)
1, Maunalaha Stream; 2, Moleka Stream; 3, Kanealole Stream; 4,
Kanaha Stream; 5, Puu Ohia (Tantalus).
X. Pauoa.
1, Pauoa Valley.
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
XI. Nuuanu.
1, Glen Ada; 2, Glen Elm; 3, Lulumahu Valley; 4, Nuuanu Stream;
4a, Nuuanu Pali; 5, Hillerbrand’s Glen; 6, Maole Stream; 7, Waio-
lani Stream (Rook’s Valley, Cooke) ; 8, Puu Konahuanui; 9, Puu
Lanihuli. (1-3, 5, place names given by Dr. Cooke to certain un-
named places where shells were collected.)
XII. Kapalama.
1, Niuhelewai Stream; 2, Kapalama Stream.
XIII. Kalihi.
1, Kamanaiki Stream; 2, Kalihi Stream.
XIV. Kahauiki.
1, Kahauiki Stream.
XV. Moanalua.
1, Manaiki Stream; 2, Moanalua Stream.
B. Ewa District.
I. Halawa.
1, South Halawa Stream; 1a, South Branch (Welch); 1b, North
Branch (Welch) ; 2, Central Halawa Stream (Welch) ; 3, North
Halawa Stream.
II. Aiea.
1, Aiea Stream.
III. Kalauao.
1, Kalauao Stream.
IV. Waimalu.
1, Hanaiki Stream (Bishop Estate maps); 2, Waimalu Stream; 2a,
South Branch; 2b, Central Branch; 2c, North Branch; 3, Punanani
Gulch. (2a-2c Welch.)
V. Waiau.
1, South Waiau Gulch; 2, North Waiau Gulch. (1-2 Welch.)
VI. Waimano.
1, Waimano Stream; ta, South Waimano Stream; 1b, South Central
Waimano Stream; 1c, Central Waimano Stream; 1d, North Central
Waimano Stream; rte, North Waimano Stream. (1a-1e Welch.)
Ia-Ic are in the district of Waiau but for convenience have been
considered part of the Waimano Stream system.
VII. Manana.
1, Manana Stream.
VIII. Waiawa.
1, Waiawa Stream; ta, South Waiawa Stream (Welch); 1b, North
Waiawa Stream (Welch); tc, North Waiawa Stream, South
Branch (Welch); 1d, North Waiawa Stream, North Branch
(Welch) ; 2, Panakauahi Gulch; 2a, Panihakea Gulch (Baptista),
the upper part of Panakauahi Gulch. (Mr. Baptista tells me that the
Hawaiian name for South Waiawa Stream is Keahupuolo Stream.
However the shells in the collection are labeled South Waiawa
and I think it is advisable to use this name in preference to
Keahupuolo. )
NO. I ACHATINELLA APEXFULVA DIXON—WELCH
NI
IX. Waipio.
1, Kipapa Gulch; 1a, South Kipapa Gulch; 1b, Central Kipapa Gulch;
1c, North Kipapa Gulch, second North Branch; 1d, North Kipapa
Gulch, first North Branch. (G. D. Robinson tells me that only the
lower portion of the valley should be called Kipapa Gulch. The
upper portion is Waipio Valley. Kipapa was the crossing place in
the gulch where stones were placed together to form a road. A. F.
Judd also considers Waipio to be the correct name of the Gulch.
However, since the collections and the U. S. Geol. Surv. topogr.
advance sheets use the name Kipapa, I believe it would be confusing
to use Waipio.) ; 2, Waikakalaua Stream. (1a-1d Welch.)
C. Wahiawa District.
I. Waianae Uka.
1, South Kaukonahua Stream; 1a, South Kaukonahua, South Branch
(Welch) (this may be Gulick’s Kalaikoa District, or Kalaikoa may
have included all of South Kaukonahua Stream) ; 1b, South Kauko-
nahua, Central Branch (Welch); Ic, South Kaukonahua, North
Branch (Welch); 2, North Kaukonahua Stream. (Mr. Robinson
showed me a place where formerly stood a boulder called Oahunui.
The saying was that if anybody walked around this rock in a spiral
starting at the bottom and going up to the top he would have gone
around the island of Oahu. Ahonui may be a corruption of the
word Oahunui. The place shown me is in the South Kaukonahua at
an elevation of about 900 feet, below or opposite the North Branch
of the South Kaukonahua. Pilsbry and Cooke, on the authority of
Spalding, who in turn probably cbtained accurate information from
J. S. Emerson, an authority on Hawaiian place names, places
Kalaikoa and Ahonui of Gulick between the North and South
Kaukinehua (a misspelling for Kaukonahua), Kalaikoa being. in
the Central Branch, and Ahonui in the North Branch, of the
South Kaukonahua. This ties in with Mr. Robinson’s information.
Ahonui of Gulick, however, may have included this region as well
as North Kaukonahua Gulch. The Gulick shells represent a mix-
ture from over a considerable area.) ; 3, Poamoho Stream; ga,
South Poamoho Stream; 3b, Central Poamoho Stream (Main
Poamoho, Pilsbry and Cooke); 3c, Central Poamoho Stream,
North Branch; 3d, North Poamoho Stream (Central Poamoho?,
Pilsbry and Cooke) ; 3e, North Poamoho Stream, North Branch
(West Poamoho, Pilsbry and Cooke). (3a-3e Welch.)
If, Paala Uka.
1, Helemano Stream; 1a, South Helemano Stream (Welch) ; 1b, North
Helemano Stream (Welch) ; 2, Kawaihalona Gulch (Holt, Robin-
son, Pulaa, Pilsbry and Cooke).
D. Waialua District.
I. Kawailoa.
1, Opaeula Gulch; 2, Kawailoa Gulch; 2a, First South Branch; 2b,
Second South Branch; 2c, Third South Branch; 2d, Fourth South
Branch; 2e, First North Branch; 2f, Second North Branch (2a-2f
Welch) ; 3, Kawaiiki Gulch; 4, Kawainui Gulch; 4a, Puu Kaina-
puaa; 5, Kawaipapa Gulch (Pulaa) (this Gulch will always be
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. “103
referred to as Kawaipapa, Kawailoa Gulch, because this place name
occurs elsewhere in the Koolau Range); 6, Ukoa Gulch; 7,
Kukaiohiki Gulch; 8, Keamanea Gulch; 9, Kaluapo Gulch (Luapo,
Pilsbry and Cooke) ; 10, Kaalaea Gulch; 11, Kaluahole Gulch; 12,
Waoala Gulch (wrongly spelled Waiola on the U. S. Geol. Surv.
topogr. advance sheets); 13, Huluela Gulch; 14, Kawaikoele
Stream. (5-13 Pulaa.)
E. Koolauloa District.
I. Waimea.
1, Waimea River; 2, Kamananui Stream; 3, Kolokini Stream (Pulaa) ;
4, Namahana Stream (Pulaa); 5, Elehaha Stream; 6, Kawaieli
Stream (Pulaa). (Mr. Pulaa tells me that the Kawaieli Stream
is where the Elehaha Stream is marked on the map. However, for
‘convenience I shall not change the position of the Elehaha Stream
on the U. S. Geol. Surv. map but shall consider the stream to the
south the Kawaieli Stream instead of the Elehaha.); 7, Kauwalu
Stream.
II. Pupukea.
1, Kalunawaikaala Stream; 2, Pakulena Stream.
Ill. Paumalu.
1, Paumalu Stream; 2, Kaleleiki Stream; 3, Kawaipi Stream; 4, Aimu
Stream.
IV. Kaunala.
1, Kaunala Gulch.
V. Waialee.
1, Waialee Gulch.
VI. Pahipahialua.
1, Pahipahialua Gulch.
- VII. Opana.
1, Kawela Gulch.
VIII. Hanakaoe.
1, Oio Stream; 2, East Oio Stream.
IX. Kahuku.
1, Hoolapa Gulch; 2, Kalaeokahipa Gulch; 3, Ohia ai Gulch.
X. Keana.
1, Keaaulu Gulch.
XI. Malaekahana.
1, Lamaloa Gulch; 2, Hina Gulch; 3, Malaekahana Stream; 3a, North
Malaekahana Stream; 3b, South Malaekahana Stream. (3a-3b
Welch. )
XII. Laie.
1, Omao Gulch; 2, Kaluakauila Gulch; 3, Kahawainui Gulch; 4, Kaaoao
Gulch ; 5, [hithi Gulch; 6, Wailele Gulch; 7, Koloa Gulch; 8, Aakaki
Gulch; 9, Kokololio Gulch.
XIII. Kaipapau.
1, Kaipapau Gulch.
XIV. Hauula.
1, Waipilopilo Gulch; 2, Hanaimoa Gulch; 3, Kawaipapa Gulch; 4,
Maakua Gulch; 5, Papali Gulch; 6, Punaiki Gulch.
XV. Makao.
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 9
XVI. Kapaka. :
XVII. Kaluanui.
1, Kaluanui Stream; 1a, North Kaluanui Stream; 1b, Central Kaluanui
Stream; 1c, South Kaluanui Stream. (1a-1c Welch.)
XVIII. Punaluu.
1, Punaluu Stream.
XIX. Kahana.
1, Kahana Stream; 2, Kawa, Stream.
XX. Kaaawa.
1, Olona Gulch (Meinecke), or Weliweli Gulch (Oswald). (1 am using
the name Olona because I have heard from some source other than
Meinecke that that was the correct name, and I have heard of
Weliweli from only one source. However, I am not at all sure
which is correct.) ; 2, Kaaawa Stream.
F. Koolaupoko District.
I. Kualoa.
II. Hakipuu.
1, Hakipuu Stream.
III. Waikane.
1, Waikane Stream; 1a, North Branch; tb, South Branch; 2, Waikeekee
Stream; 3, South Waikane Stream (Welch).
IV. Waiahole.
1, Uwau Stream; 2, Waianu Stream; 3, Waiahole Stream.
V. Kaalaea.
1, Kaalaea Stream.
VI. Waihee.
1, Waihee Stream.
VII. Kahaluu.
1, Kahaluu Stream; 2, Ahuimanu Stream; 2a, East Ahuimanu Stream
(Welch).
VIII. Heeia.
1, Iolikaa Valley ; 2, Haiku Valley; 3, Keaahala Stream.
IX. Kaneohe.
1, Kaneohe Stream; 2, Luluku Stream; 3, Kamooalii Stream; 4, Kawa
Stream.
X. Kailua.
1, Kahanaiki Stream; 2, Palapu Stream; 3, Omao Stream; 4, Maunawili
Stream; 5, Ainoni Stream; 6, Makawao Stream; 7, Olomana
Stream.
XI. Waimanolo.
CLIMATOLOGICAL DATA AND PHYSIOGRAPHY
The main division ridge or backbone ridge of the Koolau Range
extends for 37 miles along a northwest and a southeast axis in the
eastern portion of the island of Oahu. Parallel to it are the Waianae
Mountains, which are of smaller extent and lie in the western part
of the island. The windward slope of the Koolau Range is made
up of high cliffs and short ridges usually less than 3 or 4 miles long.
IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
The leeward slope for the most part has long fingerlike ridges which
may be as much as 15 miles in length and is much drier than the
windward slope. The trade winds blow from the northeast and hit
the high wall of cliffs on the windward slope. The maximum rain-
fall resulting from the rising of the moisture-laden trade winds over
the mountain wall falls not on the crest of the range but leeward
of the summit. This is shown in table 1, where a series of precipita-
tion records obtained from a station at Nuuanu Pali, at the summit
of the Koolau Range, shows a lower rainfall than at Luakaha, below
the summit of the range. At a still lower elevation the rainfall drops
considerably at the Honolulu United States Weather Bureau station.
Table 1 also shows a greater precipitation at stations on the windward
TABLE 1.—Annual precipitation records in the Koolau Range
Precipitation in inches
Elevation ne
Station in feet 1916 1917 1918
Kaneohe merticne ceric skin: nbn 100 58.92 97.37
Ma gnawiline nr oes soaie 8 ine clacre ne ee 250 93.58 85.53 122.17
Vic ATIOIO. ' os on. c.tee cree ciate eee 25 55.80 63.19 66.90
Nera PAL | aco ost a eee one 1,136 123.30 119.00 136.00
Beak KCUPDEL) | eireses saMoctntat tune 1,028 196.99 159.19 202.61
Buakalia (lo wer)y << issn eres celta 881 175.50 146.07 187.88
Honolulu (U. S. Weather Bur.)....... III 44.96 46.64 37.40
Mia kcipttttmseaniercers tet tie cate eters 570 37.07 51.44 46.64
IA Wantkh (1.5 c sie tore brie untae id oars 1,185 121.94 105.08 107.32
Opaettlaitinn cacti woe arise ae 1,100 80.46 74.55 81.83
TeV atl Gal igh Suse ye Sisk che ie alten 185 38.31 46.90
side of the island, such as Kaneohe and Waimanolo, than at the
Honolulu station on the leeward side. Table 1 was compiled from
“Climatological Data,” published by the United States Department of
Agriculture.
Hosaka (1937) gives some interesting data on temperature, relative
humidity, soil moisture, and soil temperature in Kipapa Gulch, which
is in about the center of the Koolau Range. The general trend is a
decrease in temperature and an increase in humidity with increase
in elevation. The average annual rainfall taken over a period of 5
to 36 years in Kipapa Gulch is as follows: At 59.05 to 196.85 feet
(Haole Koa zone), 23.07 to 24.33 inches; at 377.28 feet (Haole Koa
zone) to 672.54 feet (Guava zone), 47.77 to 47.49 inches; at 738.16
feet (Koa zone), 79.05 inches; at 1,797.82 feet (Ohia zone),
199.01 inches. The atmospheric temperature records from March 19
to November 26, 1933, show an average temperature at 246.06 feet
NO. I ACHATINELLA APEXFULVA DIXON—WELCH ra
(Haole Koa zone) of 84.56° Fahrenheit; at 738.16 feet (Guava
mame), 77 > at 1,000.61 feet (Koa zone), 75°2; at 1,591.2 feet
(Ohia zone), 71°6; at 2,198.2 feet (Cloud zone), 70°7 (see below for
explanation of plant zones, and fig. I, p. 3).
The extreme southeastern section of the Koolau Range is ex-
ceedingly dry (see table 1, Makapuu). The main division ridge
rises toward the west to 2,200 feet above Kuliouou Gulch, at Puu O
Kona (fig. 7, p. 194). Between Puu O Kona and Konahuanui the
backbone ridge fluctuates in height from 2,200 to 2,600 feet. At the
head of Nuuanu Valley the division ridge rises to 3,105 feet at the
high peak of Konahuanui, descends at Nuuanu Pali to 1,186 feet,
and rises again to 2,700 feet at Puu Lanihuli. To the northeast of
Puu Lanihuli the backbone ridge fluctuates in height from 2,250 to
2,800 feet. The usual elevation would be roughly 2,400 to 2,500 feet.
Above Kawainui Gulch in Kawailoa the backbone ridge begins to
drop in elevation near Puu Kainapuaa. From Puu Kainapuaa to the
head of Ohia ai Gulch there is a gradual descent from 2,250 to 1,750
feet. From Ohia ai Gulch to the head of Pupukea the drop is a gradual
one from 1,750 to goo feet.
Achatinella apexfulva today occurs almost exclusively on the lee-
ward slope of the Koolau Range. The material in this paper comes
from 45 ridges and 11 valleys. Five of the ridges are on the wind-
ward slope of the Koolau Range. The upper limit at which A. apex-
fulva occurs is usually from 4 mile to 14 miles from the backbone
- ridge, but it may be farther on some ridges which have not been
collected at higher elevations. The lower edge of the collectable area
varies northwest of Nuuanu Valley to Kawailoa (fig. 7) from 24 to
5% miles in a straight line from the backbone ridge. In Nuuanu Val-
ley and on the Niu-Wailupe Ridge the localities are less than a quarter
of a mile from the summit of the backbone ridge. These two regions
are exceptions to the general rule. It is strange that this species is
not found on the backbone ridge as are other species. But as far as
the material from carefully plotted localities is concerned no speci-
mens are from the backbone ridge.
Hosaka (1937, pp. 179-180), in discussing the ecology of Kipapa
Gulch distinguishes six plant zones, which are copied on figure 1.
They are as follows: Maritime zone, Haole Koa zone, Guava zone,
Koa zone, Ohia zone, and the Cloud zone. Hosaka writes:
In the Guava Zone Psidium Guayava and Lantana Camara are most common.
Above this region is the Koa Zone, dominated by Acacia Koa and Gleichenia
linearis on the slopes, and by Aleurites moluccana in the gully bottoms. The
central portion of the native forest, the Ohia Zone, is dominated by Metrosideros
I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
collina (Ohia lehua), a tree towering 15 meters or more above the ground. The
cloud zone is characterized by low, dwarfed shrubs in more sheltered parts and
by mat-forming Panicum, Paspalum, Isachne, and mosses in exposed windswept
areas.
The maritime, Haole Koa, and Guava zones are mostly made up of
introduced plants, according to Hosaka; the Ohia and Cloud zones
contain the largest number of endemic species. The Ohia zone has
the largest number of indigenous plants.
Achatinella apexfulva is almost entirely confined to the Ohia zone
and the upper portion of the Koa zone. The width of the Ohia zone
obviously varies in different parts of the range. In the Nuuanu
Valley region the Ohia zone is comparatively narrow; to the west,
near the Poamoho Stream, it is undoubtedly wider than at Kipapa
Gulch.
MATERIAL AND METHODS
Over 11,302 shells were studied, the majority of which are from
287 localities. Out of this number 6,707 adult shells were measured
from lots containing 5 or more shells and having a length range of
at least 3 length classes. The total number of adults from plotted
localities is 3,687, of which all but 283 were collected by W. Meinecke.
Mr. Meinecke is the first person to make a large collection of
Achatinella from the Koolau Range with the localities plotted on a
good map such as the United States Geological Survey topographic
advance sheets of the map of Oahu. All his localities prior to 1933
were plotted from memory; after that date, up to 1937, he again
collected many of his former localities and many new ones, so that
many errors made in plotting from memory are checked and cor-
rected. Mr. Meinecke is such an unusually careful worker, with
such an extraordinary ability to work with maps, that I have the
greatest confidence in the exact plotting of his localities. This opinion
is further strengthened by a check of some of his localities, all of
which were found to be correctly plotted. A few of his localities
collected before July 1932 are not dependable—in fact, this comment
applies to all localities collected by anyone prior to 1932, with the
exception of the localities of Dr. C. M. Cooke, Jr., in Nuuanu
Valley which were mapped by sketch maps made in the field.
Other collectors who have supplied shells with locality data plotted
on the United States Geological Survey topographic advance sheets
and collected since 1932 are G. W. Russ, H. Lemke, and H. Lemke, Jr.
From localities that are not plotted on any map come 3,002 adult
shells; they are mostly localized from the Meinecke and other care-
NO. 1 ACHATINELLA APEXFULVA DIXON—WELCH 13
fully collected material. The exact locality of 350 of these, mostly
extinct shells from the Gulick collection in the Bishop Museum, is
unknown. The remainder of the shells were collected by W. D.
Wilder, O. H. Emerson, J. S. Emerson, L. A. Thurston, C. M. Cooke,
tm. A. Cooke, C. H. Cooke, D. D. Baldwin, E. D. Baldwin,
EK. Lyman, A. F. Judd, and I. Spalding.
Type specimens of all previously described forms were studied
whenever possible. Those in the British Museum were photographed
by E. J. Manley, those in the Academy of Natural Sciences of
Philadelphia by Miss Helen Winchester, those in the Museum of Com-
parative Zoology by the museum photographer, and those in the
Bishop Museum by Kenneth Emory. The remaining shells and the
holotypes of all new subspecies figured in this paper were photo-
graphed by E. Bafford, of Baltimore, and myself.
The type of A. swiftiti Newcomb and A. apexfulva Dixon were
among the few types not obtained for study. The type of A. a. swiftii
should be in the British Museum but could not be located. Possibly
if search were made again it would be found labeled A. a. turgida.
The lot labeled A. a. swift is not the type lot.
The data concerning localities are of four types. All localities that
were plotted from memory or prior to 1932 have an asterisk (*) after
the locality number. If the locality was collected after 1932, or if
an old locality prior to 1932 has been again collected and proved
correct, the locality number alone is given. A question mark (?) after
the locality number indicates that there is some doubt in my mind
as to the correct plotting of the material or that the material has been
localized from another collector’s material. All localities known only
as to approximate regions and plotted years after collecting also
have one question mark. Two question marks (??) after a locality
or area indicate that the locality is believed to be somewhere in the
approximate region, but there is no data to substantiate the plotting.
The plotting of the locality may be based on the form and color
pattern of the shell and what is known of shell variation in adjacent
localities.
As a general rule the greatest dimension of a locality collected
after 1932 is not over 100 or 200 yards, but some localities collected
prior to, and even some after, this date reach an extent of a quarter
of a mile or even as much as a mile. An attempt has been made to
plot on the locality maps (figs. 3-6) the exact location and extent of
each locality. Therefore the size of a locality and the distance be-
tween localities can be estimated from the scale of miles on each map.
14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Through necessity the circles denoting localities are drawn larger than
the actual scale size of the localities in order that they may be seen.
All shells collected prior to 1932 were live shells. After 1932 the
lots contained live and dead shells. The majority of shells measured
were live specimens.
In the list of localities found with each subspecies, all lots contain-
ing less than five adult specimens have the number of dextral and
sinistral specimens enumerated. If the number of adults in a lot is
five or more, the number of dextrals and sinistrals, the length range,
and the mean length of the shell are given in table 2, p. 15. All
locality place names in the text are put in boldface type. In the
Gulick lots only the type locality or localities which probably con-
tained the subspecies are put in boldface type. Other localities which
I consider doubtful are put in ordinary type.
When material is quoted under each form, the catalog numbers of
the lot are quoted only when necessary. Most of the Meinecke, Russ,
Cooke, and Lemke material has locality numbers, and is sufficiently
distinct so as not to require the quotation of catalog numbers. All
lots labeled Cooke, stand for Dr. C. M. Cooke. All lots not having
locality numbers have catalog numbers given. The Gulick lots have no
catalog numbers given because the Gulick lots have been selected
from lots containing a mixture of color forms or subspecies and are
not yet recataloged. However, the catalog numbers of the figured
shells are given in the explanation of plates.
Methods of collecting material and recording localities on map
tracings taken from the United States Geological Survey topographic
advance sheets, the manner in which the shells are graded into length
groups (table 2, p. 15), and the selecting of the typical or usual form
of the shell are the same as those already described for A. mustelina
(Welch, 1938). The only addition that should be made is the method
of counting the number of whorls, which is the same as that de-
scribed by Pilsbry (1939, p. xi). All holotypes of new subspecies
have the usual form and color pattern of the shell unless otherwise
stated.
All grading of shells into length groups for table 2, page 15, was
done by two W. P. A. workers. Many of the lots were remeasured
by me and found to be correct. Since the shells are all graded by the
measuring rod described for A. mustelina (Welch, 1938, p. 12), I
think there is little chance of error as far as has been ascertained
from remeasuring lots here and there at random.
NO: I
ACHATINELLA APEXFULVA DIXON—WELCH
15
The following abbreviations are used in citations of museum shell
lots, and in the references to the literature.
ANSP, Academy of Natural Sciences of Philadelphia
BM, British Museum
BBM, Bernice P. Bishop Museum
MCZ, Museum of Comparative Zoology
HL, Herman Lemke collection
Man. Conch., Manual of Conchology (second series)
The location of types is stated in the explanation of plates beginning
on page 208.
TABLE 2.—Data on Material
Subspecies Area
Recion I.t7
Region of ridge complex 1, zones I and II.
Collector]
A.a. forbesiana ...-.- 6?? Gulick
A. a. fuscostriata ..... 7°? ° Gulick
A. a. fuscostriata ..... 7?? Gulick
A. a. fuscostriata ..... 7°? Gulick
A. a. simulator .....4. ? Gulick
PIG. DUCNE vecaccee ae ? All lots
Ridge complex 1, zones II and III.
A. a. muricolor ...... 1 Meinecke, 1926
7A.a.muricolor ..... S I Meinecke
¢A.a.muricolor ...... I Meinecke
7A. a. waialaeensis .... 2 Meinecke, 1933
7A. a. waialaeensis .... 2 Meinecke
A. a. waialaeensis .... 2? O.H. Emerson, 1916
7A. a. simulator var.1.. 4 Meinecke
Ridge complex 2, zone III.
7A. a. vittata var. 1.... 8 Meinecke
7A. a. vittata var. 1.... 8 Meinecke, 1923
7A. a. vittata var. 1.... 8 Meinecke, 1925
TA. a. vittata var. 1.... 8 Meinecke, 1926
fA. a. vittata var. 1.... 8 Meinecke, 1927-31
qA. a. vittata var. 1.... 8 Meinecke, 1927-31
4A. a. vittata var. 2.... 9 C. M. & R. A. Cooke
WA. G. CINETER cuccecee 10 C. M. Cooke
7A. @. cinerea .....22- 10 C. M. Cooke
+A. a. cinerea ...seee. 10 Meinecke, 1911-16
A.d.cineread ...e.+e. 10? Wilder, 50504
PAS G: CINETEA «ccccees I0 R. A. Cooke, 58138
Region of ridge complex 3, zone II.
A.a.globosa .....008- ? All lots
Ridge complex 3, zone II.
fA. a. simulans ....... 16 Lemke
7A. a. simulans var. 1.. 17 Lemke & Anderson
+i For explanation of regions, ridge complexes, and zones, see pp.
+ Used for plotting on map, fig. 8.
{ Unless otherwise stated all catalog numbers are those of the Bishop Museum.
* Asterisk indicates localities that were plotted from memory or prior to 1932.
Locality Adults length
Waialae
Palolo
Waialae
Wailupe
Palolo
41-4*
41A-1*
41B-4*
s1Aa
50A*
62
goA*
g1A*
g1A*
g1A*
94A*
95A*
100-2*
100B-13
100B-12
100B-7?
121-2
I2I, 121-1
It
39
14
12
24
I!
IOI
It
16
16
16
15
12
37
57
92
12
53
32
17
62
29
20
8
10
Mean
17.0
16.5
17.0
17.0
17.0
17.0
17.5
18.5
18.0
18.5
18.0
19.5
18.0
17.5
18.0
18.0
18.0
18.5
18.0
18.5
19.0
19.5
19.0
19.5
20.0
17-5
17-5
17.0
Length Dex- Sinis-
range tral tral
mm.
16.5-18.5 10 10
15.5-18.5 Bao
15.5-18.5 30 26
16.5-18.5 ae 23
16.5-18.5 as
16.5-18.5 a. 21
15.5-19.5 170
16.5-19.5 ate 12
16.5-19.5 ae 6
17.5-19.5 bie 20
17.5-19.5 17323
18.5-20.5 ate 23
16.5-19.5 21
15.5-20.5 atc 44
16.5-19.5 Ae 72
16.5-19.5 ake gI
16.5-20.5
16.5-19.5 ee 10
17.5-19.5 ate 13
16.5-21.5 3C 75
16.5-20.5 ate 92
18.5-20.5 36 36
17.5-20.5 ae 25
17.5-21.5 ave 89
17.5-21.5 ore 31
16.5-18.5 23 ote
16.5-17.5 are Io
15.5-17.5 ate 17
189, 190, and 193, and fig. 8, p. 195.
? A question mark denotes there is some doubt concerning the correct plotting of a locality.
?? For two question marks, see explanation in text, p. 13.
2
16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
TABLE 2.—Data on Material—Continued
Mean Length Dex- Sinis«)
Subspecies Area Collector] Locality Adults length range tral tral |
Ridge complex 3, zone III. mm. mm.
FAL Gs TOE. <.<.ise'c nisi ia R. A. & C. M. Cooke, 100E-1* 16 18.5 16.5-20.5 oe arr
58284, 22119
{A. a. albofasciata .... 13? Wilder & Thurston 100A-1? 20° 19:0 11755-2005 4a) 4
A. a. albofasciata var. 1 14?? J. S. Emerson ereipia 16 19.0 17.5-20.5 ate 34
A. a. oliveri ......... 15? O.H.&J.S. Emerson 120-1? 35 §1010) 75-2765 a Ee
+A.a.rubidipicta ..... 11 Meinecke, 1921 I10* 119 19.5 16.5-21.5 aie ECGS
tA.a.rubidipicta ..... 11 Meinecke, 1922 I10* 65 “19:0 27.5-2015 0 3. tad
tA. a. rubidipicta ..... 11 Meinecke, 1927 r10* 76 19.5 16.5-21.5 +. 100
tA. a. rubidipicta ..... 11 Meinecke, 1929 110* 49: “19:0, 85<5-20)500 od
Ridge complex 4, zone II.
tA. a. simulans var. 2.. 19 Meinecke 151B* ¥2) 18:5) | -16/5=1ols ea
Ridge complex 4, zone III.
tA. a. rubidipicta var. 2. 18? Wilder Kahauiki 8 20.0 19.5-21.5 re 8
Ridge complex 4, probably zone ITI.
A. a. rubidipicta var. 1. ? Wilder Kalihi 18 19.5 18.5-20.5 9 “16
A. a. rubidipicta var. 3. ? Wilder Moanalua 75. 10:50) I7.5-2 0-5 en eR
Recion II.
Ridge complex 5, zone ITI.
TA. a. ovum var. 1..... 21 Meinecke 162B* T6) 8-5 16.5-19.5 36 a
A. a. ovum var. 1..... 21? Thurston Sacer 8 1850) 6-5-7055) ase
Ridge complex 5, zone III.
TAG. OVUM) ccsecccves 22 Meinecke 162C* 79190) -07.5=2055 eae 12
A, d. OVUM ...0.0056. 222? Wilder, 50525 serie 17 ©2050) "4825-22550 eto
tA. a. OVUM Var. 2...62 23 Meinecke 162D* 22) 19:5) 9 1755-20:5 eas
A. a. ovum var. 3..... 20 Meinecke & unlocal- 163* 9 19.0 £1£6.5-20.5 oer
izede yg Sy Ow Ee
Emerson lots
Ridge complex 6, zone II.
TA. Oe PSSOT YE cie:clewiclaie 26? Cooke & Judd 170-1? SLAs Sear eae ee
Ridge complex 6, zone III.
A.a. bruneola var. 2.. 25 Cooke & Judd se 61) —920;0) 5928.5-27.5) 77a
tA. a. bruneola var. 2.. 25 Lemke 170C-6* Al = (20.0) )016;5-20-5) 5 Ommete
fA. a. ovum var. 2..... 24 Meinecke 174* 14) (20:0. 5725-2255) ee ae.
Ridge complex 7, zone II.
TAG LOUTONE secceeus 27 Meinecke I91* 7) TS.5) et7 5219-51) eae ee
TA. a. baurant ...s00. - 27A? Cooke, Judd & Sheers mete MOY adeHetH Sin aa
Thurston
Ridge complex 7, zone III.
fA.a.bruneola ....... 28 Meinecke, 1926 182 9 19.0 17.5-20.5 PS yor
{A. a. bruneola var. 1.. 29 Meinecke, 1913 183? 19 20.0 18.5-22.5 2) 42
~A.a. bruneola var. 1.. 29 Meinecke, 1935 183 7 at, 19.5-20.5 aie 13
TA. a. aureola cecccsse 30 Meinecke, 1913 184 12 20.5, I9.5-22.5 I 47
FA. ad. aureola .cccecse 30 Meinecke, 1926 184 19 =. 20.5 18.5-23.5 2 40
TA Gi GUPeOlG. ..0s:0c0c1c 30 Meinecke, 1932 187? [29 9190:0))09.5-20:5) aro. I
A GNGULEOIE <aceccielcee ~30 Meinecke, 1933 187 3 ate 18.5-20.5 Al etre
Wis GHOUEOID:. <crstccir cists 30 Meinecke 185 10 820.0 18.5-21.5 Ae 16
{A. a. aureola var. 1... 31 Meinecke 190A* TO) Te.50 | 07.5= 20.5 se
Ridge complex 8, zone I.
fA. a. roseata .....22. 372 All lots eieieie 17 9TG.0 16.5-69.5) 55 eo
Ridge complex 8, zone II?
fA. a. parvicolor ...... 36?? Gulick aires 19 618.0) 16.5-19.5 24 9
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 17
TABLE 2—Data on Material—Continued
Mean Length Dex- Sinis-
Subspecies Area Collector§ Locality Adults length range tral tral
Ridge complex 8, zone III. mm, mm.
A a.turgida ..e...+. 35 Meinecke, 1913 202B Te xGGee 1745-2955)0) £20) 0
Galea. turgida ......-. 35 Meinecke, 1920 202B ST 9:0) t7.5-2tesn S58 ers
Midd. turGtda 3 ..cecese 35 Meinecke, 1934 202B 9 19.0 16.5-20.5 15 2
BEG. EUTGIAA .cecccee 35 Wilder, 50598 Ses 170 19.0 16.5-22.5 201 60
PIAL EUTGIAG «we ccceee 352 J.S. Emerson, 102218 .... 121 19:5 16:5-21.5)) 228).
A. a.turgida ...... -» 35? J.S. Emerson, 102218 .... TOF | XQ:5 os XOs5-22-5)) ee | 275
7A. a. meadowsi ...... 34 Meinecke 212b TENS LQlON ne EyA5-2 ies ere 17
7A. a. meadowsi ..... a4: Meinecke 213a meh Sia | SACHS Gor 18
7A. a. meadowsi var. 2. 33 Meinecke 200Ca* Pi2TZONETOu5-22es tcl 18
7A. a. meadowsi var. 2. 33 Meinecke 201C Rue 20st 20-5-22.5 aS II
7A. a. waimaluensis ... 32 Meinecke 202C 9) 20:5) loss-20. Serato PIE
7A. a. waimaluensis ... 32 Meinecke 203C @. (20:0) 9 38.5-20:5 ie T 4
Recions II?, IIIa.
Ridge complex 9g, zone II.
7A. a. perplera ....... 40 Meinecke 222* II 18.5 17.5-20.5 14 19
7A. a. perplera ....... 40 Meinecke 2214* T7)) VISs5 2625-20:58 924
74. a. perplexra ....... 40? Cooke dacee 4c) $2905) | 15.5-20-5, 54 4
7A.a. perplexa ...... - 40? Wilder iaieks 129 18.0 I5.5-21.5 94 63
7A. a. nigripicta var. 1. 44? Wilder Sask 27 18.5 15.5-20.5 26 7
A.a.nigripicta var. 1. 44? Cooke sree 23. 1735 16.5-19.5 41 I
A.a.nigripicta var. 1. 44? J. S. Emerson, 10717 .... 5I 18.0 16.5-20.5 105 ts
A.a.wigripicta var. 1. 44? J. S. Emerson, 10718 .... TIO. =6©19.0 ~=16.5-22.5 se 20%
A, a. chromatacme
SICA Ns) </a\oies\ o> so0o) Gis Meinecke 231" 68 185 16.5-20.5 13 102
tA. a.chromatacme ... 42 Meinecke 232 33. 19.0 16.5-20.5 I 73
7A. a. chromatacme ... 42 Meinecke 234* 35 19.0 17.5-20.5 ays 93
A.a.chromatacme ... 42 Meinecke 230B? TOWNIOLO LOSS 20.50 aon tars
+A.a.chromatacme ... 42 Meinecke 2358 Ss" ro:0° 17.5-21-5 69) 7
7A. a.chromatacme ... 42 Meinecke 236* 65 19.0 16.5-22.5 34 88
7A. a. chromatacme
Seitemteiicleleicieissis\cvis'e 43 Meinecke 237* 23 418.5 16.5-21.5 49 2
Ridge complex 9, zone III.
TA. a. cooket var. 1.... 39 Meinecke 211B 7 19.0 I17.5-20.5 << .27
A. a. cooket ......... 45? Wilder, 50594 erelete meg? | Bee) eA G5 SI
A.a.cooket ......... 45? Wilder, 50595 ee 02) ) E1O:0) 0725-205 ts
Ps COOREY nccercees 45? Baldwin & C. H. Cooke .... 8 20.0 18.5-21.5 12 ic
Gs. COOKE? secssccs - 45° ©! M. & R.A. Cooke Have 72) 20:0) 2725-2150 tr 6
Hea @. COOREL .ccens nena Meinecke 226* 6 20.0 17.5-21.5 15 ote
tA. a. simulacrum ..... 46 Meinecke 228a 8 20.0 18.5-21.5 A 13
TA. a. simulacrum ..... 46 Meinecke 228* 29 19.5 18.5-21.5 sree aA)
7A. a. simulacrum ..... 46 Meinecke 229* 17 20.5. 19.5-22.5 ag 31
WA. a. nigripicta ....+. 50 Meinecke 237b EON S65) 1 7a5=10.5) ero! ar
A. a. nigripicta ...... 50 Meinecke 237¢ 8 19.0 17.5-20.5 fe 13
7A. a. nigripicta ...... 50 Meinecke 237d GC) Fe) ECHR aa 2G
A.a. nigripicta ...... 50 Thurston seine 65 19.0 16.5-20.5 27 62
7A. a. cooket var. 2.... 49 Meinecke 2371 16) TQLO 8 1745-20-57 725 a
TA. a. cooket var. 2.... 49 Meinecke 237fa 7 19.5 18.5-20.5 8
7A. a. nigripicta var. 2. 48 Meinecke 237h 13 19.5 18.5-20.5 22 ora
7A. a. nigripicta var. 2. 48A Meinecke 238a-238b =—-:117 19.5 18.5-21.5 oe 27
7A. a.rubidilinea ..... 47 Meinecke 239a 7 20.5 18.5-21.5 .: 8
7A. a. rubidilinea ..... 47 Meinecke 220A? TE) t9Q:5)°) USt5-2105) ae ae
Recron IIIa.
Ridge complex 10, zone II.
7A. a. albipraetexta ... 63? Meinecke 231B* 22 18.0 16.5-20.5 cient 45
tA. a. albipraetexta ... 63? Meinecke 232B* 21 19.0 17.5-20.5 Seas
{A. a. albipraetexta ... 64 Lemke 233B-1* 14 19.0 16.5-20.5 13 6
Ridge complex 10, zone III.
7A. a. ewaensis var. 1.. 51 Meinecke 234B* 5 18.5 17.5-19.5 3 6
18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
TABLE 2.—Data on Material—Continued
Mean Length Dex- Sinis-
Subspecies Area Collector] Locality Adults length range’ tral tral
Recion III.
Ridge complex 11, zone II. mm, mm,
A. a. griseibasis ...... 652? J.S.&O.H. Emerson .... 21 17.5 15.5-20.5 46 I
A. a. polymorpha var. 1 ? J. S. Emerson ditene 15 18.0 16.5-19.5 18 5
td. a. flavitincta ...... 69 Meinecke 230G* 20 -17.0' | *TS.5-16.5 nos 6
tA. a. flavitincta ...... 69 Meinecke 231G? 23. (17.0) “Ta -S-1728) oats
TA. a. flavitincta ...... 69 Meinecke 231G 41 17.0 ©5.5-07.25) Os) 12
A. a. flavitincta ...... 69 Meinecke 232G* a7’ 56.55 ADE Gato sumer y eco
TA. a. linetficta ....... 70 Meinecke 232Ga 17 17.0 15.5-18.5 , 20
A. a. lineipicta ....... 70 Meinecke 233G* O°. t8ss) r745=20.5h One
fA. a. lineipicta ....++. 70 Meinecke 234G 25 70) O5.5-18.5) 33 I
tA.a.lemkeit var. 1.... 68? Lemke 233Ga? 24 18.0 16.5-20.5 23 29
tA. a.lemkei ......... 67? Lemke 230E-1? 46 18.0 15.5-21.5 7 50
A.a. fumositincta .... 71 Meinecke 240* 13 | t7.5° 16.8-10.5) " 0 40
tA. a. fumositincta .... 71 Welch 241-1* 8 16's, tS s5-17-50 ee eee
A. a. fumositincta .... 71 Russ 241-1? 8 16.5 15.5-17.5 1 ora
A. a. fumositincta .... 71 Russ 240-I-
241-1a? II 16.5 15.5°17.5 2 13
tA. a. fumositincta .... 71 Welch 241-2* 15 D720) 25.5=To.5 tad
tA. a. fumositincta .... 71 Welch 241-3* 6 | 06:5. 5S.5=17/5) ne 17
Ridge complex 11, zone III.
7A. a. ewaensis var. 3.. 52 Meinecke 243-2* Q 1825 _ 1655-19:5) 12 4
tA. a. ewaensis var. 3.. 52 Welch 243-4* 7 17.0 16.5-18.5 sie 9
7A. a. ewaensis var. 2.. 53 Meinecke 244* 1S) t6:5 wses-1Ss5 See 933
TA. a. ewaensis var. 4.. 54 Meinecke 244-3? 12) D725) | Seer ossya meee 16
{A. a. ewaensis var. 4.. 54 Meinecke 246* 7 17.5 16.5-19.5 a 13
Ridge complex 12, zone II.
fA. a. virgatifulva .... 73? Wilder Etioce 53: 18.0) +I6.6-2%-5) ¢20 ase
A. a. virgatifulva var.2 74 Meinecke 250* Guns 15.5°17.5 21 as
tA. a. virgatifulva var. 2 74? Wilder; 50571 Sena 32 17.0) 1 605.5-10.5) a4 ks
A. a. virgatifulva var. 2 74? Wilder, 50573 waas 32) “7.5 04.5-20:5) zoe
Ridge complex 12, zone III.
1A. a. ewaensis var. 5.. 55 Meinecke 240L-1-1a II 17.5 16.5-18.5 TAO
7A. a. ewaensis var. 5.. 55 Meinecke 240L-1 II 17.5 15.5°19.5 ve 17
+A. a. ewaensis ...... >. 56 Meinecke 240L-4 on 17.5 |fO.5-16.5) 02 4
TA. G. CEWAENSTS ceccece 56 Meinecke 240L-6 7a aa 15.5-18.5, 9 .
7A. a. ewaensis var. 6.. 57 Meinecke, 121935- 252-1-2? 29) ROWS, ) Ss5-17-S coe eed
121936 y
tA. a. ewaensis var. 6.. 57 Meinecke, 121937- 253? 31 07.0) | ets S-tors) eons
121938
tA. a. ewaensts var. 7.. 58 Meinecke 256 7 17.0 16.5-18.5 Ac 15
tA. a. ewaensis var. 7.. 58 Meinecke 257 18° (1.0) § 9X5-5-20ls ee Snr
Ridge complex 13, zone II.
+A. a. virgatifulva var.t 75? Thurston, 130725
aiere 16.5-18.5 ae 14
7A. a. tuberans var. 2.. 60 Meinecke 261*
18.0 16.5-19.5 4 10
a oo
-
uo
Ridge complex 13, zone III.
7A. a. ewaensis var. 8.. 59 Meinecke 261-262? 18 19.0 17.5-20.5 a 42
7A. a. ewaensis var. 8.. 59? Wilder isle 23 19.0 17.5-20.5 7 a5
Portion of ridge complex 14, zone II.
+A. a. tuberans var. 4.. 76? Meinecke, 1918 260A? 17 TSO) 1) )TS.8=21.5 22 2
7A. a. tuberans var. 4.. 76? Meinecke, 1924 260A? Il 18.0 16.5-19.5 oe 29
A.a.tuberans var. 4.. 76? O. H. Emerson ersiia’s 7 18.0 16.5-18.5 2 rs
A.a. tuberans var. 4.. 76? Wilder aie 27 18.0 15.5-20.5 20 19
A. a. tuberans var. 5.. ? Wilder, 50575 States 56) 00:5, 8 75 .5-18.5) OD eS
A.a.tuberans var. 5.. ? L. A. Thurston ratals 33 17.0 15.5-18.5 ae ee
NO. I ACHATINELLA APEXFULVA DIXON—WELCH Ig
Taste 2—Data on Material—Continued
Mean Length Dex- Sinis-
Subspecies Area Collector] Locality Adults length range tral tral
Recion IV.
Possibly in the region of ridge complexes 14, 15, zone I and lower zone II.
mm. mm.
A. a. CONtfOTMIS coese0 oe Gulick Ahonui &
Kalaikoa 14 18.0 16.5-20.5 14 ae
AG COMMJOTMIS .eeces oc Gulick Wahiawa 23° «17:5 %16.5-20.5 28 3
Ae. VEYSICOIOY ..c0e8 as Gulick Ahonui T4 17.0) °14.5-18.5 Pe 56
A.a. versicolor ..... Si hers Gulick Kalaikoa 12) 07:0) |) Ar 5.5-1055 as 28
A.a.gulickii ..... Rites ac Gulick Ahonui 8 18.0 16.5-18.5 19 9
AG GUICKW VAT. Looe cs Gulick Ahonui To) ess) | aeesrous ZY 14
nG: QUACK Var. 2... «+ Gulick Kalaikoa SoelOs5 ave DSaseSs5 se LOM er
A.a.flavida ..... eyaveres, tte Gulick Ahonui &
Kalaikoa 8 16.5 T5.5-D755 47 5
Ridge complex 14, zone II.
qa. a. steeli var. 1.0.06 79 Wilder rier TS! 56-0) 075-2005). 260) as
; TA. EISEEEIE. oi5 s<00 aies, 0.0 80 Russ 271 II 19.5 17.5-20.5 she 20
¥A.a.steeli ..... eoe-- S80? QO. H. Emerson Mors TOV SOLS eet 5-20. Se et eo
ae. SECEIS ws ccces -.. 80? Wilder, 50577 ee ATI TO'OUN et7es-20565 TS). 32
TA. a. steeli ......20. - 80? Wilder, so580 tee 13 19.0 17.5-20.5 ee 16
A. a. punicea ........ 81?? Wilder, 50581 sietets 6 919!0) | 1725-2055 3 4
FA. a. punicea ..ccseee 81?? Wilder, 50583 Rictece 56 20.0 17.5-22.5 OO
A.a. punicea var. 1... 82 Wilder, 50578 sveare E77 Sror8 17.5-21.5 2) ole
fA. a. tuberans var. 3.. 77 Lemke 274 15 19.5 17.5-21.5 19 we
A.a.tuberans var. 3.. 77 Wilder, 10448 ee 7 19.0 18.5-20.5 9 I
Ridge complex 14, zone III.
A. Q.irwint <2... see 61? Wilder ears T40) PISS L7-S-kOsse aa) 22
$A. Gd. WINE oc. eee ee 61? Meinecke 275? 13. 190.0 17.5-20.5 Aan Tl
+A. a. irwinit var. 1.... 62? Meinecke, 1918 277? 45 18.0 16.5-20.5 <f), , X00
A. a. irwinit var. 1.... 62? Meinecke, 1918 277? TG) 89:0; 7 17.5-20:5 19).
{A. a. irwint var. 1.... 62? Meinecke, 1923 277? 1o)6=—-«18.5 ~—_16.5-20.5 Be ers
Ridge complex 15, zone II.
PARTE ISLACEE <0. ie sieio -*83b?? Gulick Ahonui Te LO-O 725 -20ns ms .
A. a. lilacea .2...e-e. 83b¢? Gulick Kalaikoa 14 19.0 17.5-20.5 21 ‘
GA.a. lilacea vat. 1.... 83 Meinecke 280* LOM Nese Ges =2OuRn ls 6
7A. a. lilacea var. 1.... 83 Russ 280-1-280* 27 18.5 16.5-20.5 37 te
A.a.lilacea var. 1.... 83? Wilder Saas 23) elS.Sh el 7a5-0Os5 TA) hae
fA. a.lilacea var. 1.... 83a Meinecke 281-2* To 180 #8 16.5-20.5 tae
Ridge complex 15, zone III.
7A. a. lilacea var. 2.... 84 Meinecke 282-3 Ce LOLON tak 725-205500 | 5G
tA. a. poamohonesis ... 101 Meinecke 283-1-3-4 TOM OAS mr ie5=20.5 eis
7A. a. poamohoensis ... 101 Meinecke 284-2 POM TOS) kSs-2025.4 34 c
7A. a. roseipicta ...... 102 Meinecke 285-3-286 Ze TQ!0) ) ©7-5-19:5) 055
A,a.roseipicta ...... 102? QO. H. Emerson ayaa 29) 19:0) 7 17.5-21-5) 30
A. a. roseipicta ....-. 102? Wilder, 50563 welts 55 19.0 16.5-21.5 65 F
tA. a. roseipicta var. 1. 103 Meinecke 287-1-3 18 19.5 18.5-20.5 30
7A. a. roseipicta var. 1. 103 Meinecke 288 - Ge920:08) 1S-5-27.50) 10
7A. a. roseipicta var. 1. 103 Meinecke 289-1a 13) 1910 18.5-20,5) 23
WAL. GIOhd .s0ccereee LOO? Wilder ee 169 19.0 17.5-24.5 I91 :
fA. a.aloha .......... 1002 Thurston aoe 19 109.0 17.5-20.5 59
jA. a. aloha var. 1..... 100? Wilder sein 19 19.5 1S.5-20,5 130
Possibly in the region of ridge complex 16, zone I or lower zone II.
PAGS GPETGIDG .ccesse ss Gulick Ahonui P27.) O25 -0O85 mers *
A.a.apexalba ....... ot Gulick Wahiawa 31 17.0 14.5-19.5 26 ° 862
AG: WANIGWA scsccce + Gulick Wahiawa Us) 7.5) 16.§-18.5) 3S 3
xSee figure 7, page 194.
20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
TABLE 2.—Data on Material—Continued
Mean Length Dex- Sinis-
Subspecies Area Collector] Locality Adults length range tral tral
Ridge complex 16, zone II. mm. mm.
7A. a. suturafusca ...- 86 Meinecke 280K-1* 32) 10.0 17.5-210.5) ae teeag
A. a. suturafusca .... 86? Wilder ehevers 67° “19:0 =15.5-22.5)) 670 9
+A. a. glaucopicta ..... 87 Meinecke 280K-2 TO 19.0 £5.5-20.5 14 9
+A. @. DrUnosa .eeeeeee 88 Meinecke 280L-2 10 19.0 17.5-20.5 20 ate
+A. a. brunosa ...eeee6 88 Meinecke 280L-4 9 18.5 17.5-20.5 ms
+A. a. suturalba ....0+25 90 Lemke 290AA-8 22° (10.5) *t8.5220.5) Aree
7A. a. suturalba ...0022 90 Welch 290AA-8 13) LOLS * (18.8-20;500 aaleeene
A.a. suturalba ...ee2. 90? Wilder Raita 44 19.0 16,5=20-5) 5S 5 ete
+A. a. suturafusca var. 2 89 Meinecke, 1914 290* Sy 16:5 D7.e2r.5) vos 7
+A. a. suturafusca var. 2 89 Meinecke, 1917 290* 63 19.0 17.5-21.5 II 72
A. Gd. GPiCAEA .ecccccee 92? Gulick Helemano T4 19:0)" 517:5-20.5) cS
+A. a. Cervixnived ....2 93 Russ 300-4* 17 19.5 18.5-20.5 17 ave
A. Gd. CErViXNIVER .eeoe 93 Meinecke, 1917 300-6 Io 6. 18.0—S -16.5-19.5 26 9
A, @. CeErviznivea ...0e 93 Meinecke, 1916 300-6-301-2* 14 18.5 07.5-20.5 9 14) ern
+A. a. cervixnivea ....- 93? Wilder, 50631 124. 10,5 (17.5-22:5) 7a, ess
+A. a. cervixnivea ..... 93? Wilder, 50566 revere 19). 19:0) 17.5-2T Sid 8
+A. a. beata var. 1...2- 94 Meinecke, 1916 302* 19 19.0 16.5-20.5 25 6
+A.a. beata var. I..+4. 94 Meinecke, 1932 302* 8 19.0 17.5-19.5 9 5
+A. a. beata var. I...65 94 Meinecke, 1916-17 291-3-5* 83) {0:00 | -17.5-25-5 SOO mene
7A. a. beata var. Teese 94 Meinecke, 1932 291-3-5* 5 19:5 1£7.5-21.5 II a6
+A. a. beata var. I.-++. 94? Wilder, 50568-50569 aishe 59 19.0 17.5-20.5 88 3
+A. a. beata var. 1.....94-94a? Meinecke, 1914 291-2-302? 40 19.0 17.5-20.5 108 ne
Ridge complex 16, zone III.
+A. a. beata ..sccccecse 104 Meinecke 280L-6 8 18.5 17.5-19.5 6 9
7A. a. beata ..ccoccese 104 Meinecke 280L-9 US (20:0 {18.5-25.5) iss
4A. a. beata ..ccccceee 104 Meinecke 303 8) TONS) aS.5-20.5) eee
fA. a. beata ..seceeeee 104 Russ 303 17. TOS. |) N7.5-2l-s eat I
FA. a. DEAta .ecccceeee 104 Welch 304 Ze TOUS) erGss-20e sary
AnG: EGER ceccscccee 104 Meinecke, 1932 305-306? II 9.0 1&.5-20:5) 26
A. d. bedta .scccoccee 104 Meinecke, 1916 302-304? Of 10:0) n7c5-20.5) as
A. a. beata ...eeee0e6 104? Thurston solete 52 10:0) =17.5-20.5) 05
And: CEHtG. scacvcivcee 104? Walder erets 7e 19.5) | -S.5-25e5) ok Bi
A. a. roseipicta var. 2. 105 Meinecke 280F-2 8 720.0) 9 116.5-2755 eee
+A. a. roseipicta var. 2. 105 Meinecke 307-3-308-1 8 19.5 18.5-20.5 15
7A. a. roseipicta var. 2. 105 Meinecke, 1917 307-1a-1b 20 19.0 19.5°20.5 47
A. a. roseipicta var. 2. 105 Lemke 307-3 13 20.0 18.5-21.5 17
+A. a. roseipicta var. 2. 105 Russ, 1932 308-1 7 19.0 I17.5-21.5 8
Ridge complex 17, zone II.
A.a. suturalba var. 1... o1?? J. S. Emerson storie 32 18.5 16.5-20.5 40 6
A, a. suturalba var. 1.. 91?? Gulick ae 13) TSis) 775-1 o.s oe
tA. a. apicata var. I... 95 Meinecke, 1933 312-2-5 22 19:0 18.5-20:5 33 .
+A. a. apicata var. Ieee 95 Meinecke, 1932 310-312-4* 30 #£19.5 16.5-22.5 71 I
+A. a. paalaensis «secs 106 Meinecke 313-3 12 28:5 19.5-20.5 (15 AG
A. a. apicata var. 2... 96? Wilder, 50510 Retox 89 19.0 16.5-22.5 113 2
Ridge complex 17, zone III.
+A. a. paalaensis var. 1. 107 Meinecke 313-1 6 19.0 17.5-20.5 ne
A. a. paalaensis var. 1. 107 Meinecke, 1932 314-1 t= S.5) 0 D7s5-"Ols) esx he
7A. a. paalaensis var. 1. 107 Meinecke, 1933 314-1 TO) | 20u0) -1S55-27.5) 24
+A. a. paalaensis var. 1. 107 Meinecke, 1932 316-1-3 27 19.5 18.5-21.5 33 :
7A. a. paalaensis var. 1. 107 Meinecke, 1933 316-1-3 19 19.5 18.5-20.5 37
+A. a. paalaensis var. 1. 107 Meinecke, 1932 334* 2 20.0 18.5-21.5 40
7A. a. paalaensis var. 1. 107 Meinecke 336-1* 7 ‘ToLs 17.5=20.8) Pismo
A. a. paalaensis var. 1. 107 O. H. Emerson Bree 22) 191s 17.5-20.5) 57, ,
+A. a. beata var. 2..... 108 Meinecke 318 a7 tos 18.5-21.5 67 one
+A. a. beata var. 2..6+. 108 Meinecke 318a OQ) 20:0) ““AS.s-22:50 0
+A. a. paalaensis var. 2. 109 Meinecke, 1929 337-1-2? 12 20.0 18.5-22.5 16 :
+A. a. paalaensis var. 2. 109 Meinecke 337-3 9 20.0 18.5-20.5 14
A. a. paalaensis var. 2. 109 Meinecke, 1932 337-1-3? TS) IO.5) (ISie=20.5) ss
A. a. paalaensis var. 2. 109 Meinecke, 1932 337-1-3? 1S LOS) t7.5-2r.5 eo
A. a. paalaensis var. 2. 109 Meinecke, 1928 337-1-3? 12 19.5 17.5-21.5 17 te
+A. a. beata var. 3..+e+6 I10 Meinecke 337-4 13 19.5 17.5-20.5 26 ae
A. a. beata var. 3.ee06 110 Meinecke 330-330-1820 19:5 17.5-21.5 46
7A. a. beata var. 3..... 110 Meinecke 338 6a 2055 19.5-22.5 16 ate
+A. a. beata var. 3+... 110 Meinecke 339-1a* T5o0 20.0) 2 VSi5-20.5 Omics
A. a. beata var. 3..e6- I10 Meinecke 338? 6 ~2ofa) 1825-255) 36).
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 21
TABLE 2.—Data on Material—Continued
Mean’ Length Dex- Sinis-
Subspecies Area Collector] Locality Adults length range tral tral
Ridge complex 18, zone I. mm. mm,
7A. a. duplocincta ..... 97A?? J. S. Emerson ae ae EO eiS:O)0 1615-10:5))) Ox
Ridge complex 18, zone II.
4A. a.apexfulva ....+2 97 J. S. Emerson Opaeula Boo a e7AS- TOL Ess
Ridge complex 18, zone III.
fA. a. kawatiki ....... 99? Wilder & Thurston Shivers 9) To.5. 16.5-20.5 16
tA. a. vespertina ...... 98? All lots relate 220) TO Sag-2tes as
Somewhere between ridge complex 18 and 19, or in ridge complex 17 or 18.
Pie. DOREY wesc. ceees sve J. S. Emerson & Judd .... 12 7S PLS teaTO see ic: I
Probably ridge complex 19, zone I or lower zone II.
PAGs MAPUS .occecccce vs Gulick eynees TOM P1825) LOs5-20s8) TG
A.a.leucozona ....- «+ Gulick, ANSP 92656 .... 78 17.5 15.5-19.5 164
A.a.leucozona ...... we J. S. Emerson ssiete 8) 9725) -16.5-18i5) ro
Ridge complex 19, zone II.
tA. a. paumaluensis ... 111 Meinecke 431 TAS Loess TOss-20058 8 54)
7A. a. Otoensis ...... 56 pag Meinecke 460A Ou ty7es ee LOss<1S-5) 01g
TA. a. otoensis var. 2... 113 Meinecke 462* 20) 1S5)) )16s5-19:5) 56
A. a. otoensis var. 2... 113? Thurston eee 23 tS eeTOe5-1O.n nea,
tA. a. oioensis var. 1... 112 Meinecke 450 Be eises | atya5-LO.5) | 26
Ridge complex 20, zone I.
fA. a. wailelensis ..... 117 Russ & Welch Riis DAM LCs neta =2Oncy meni
Ridge complex 20, zone II.
TA. a. thithiensis ....+.. 116 Welch 510-2a 5 18.0 16.5-19.5 10
Ridge complex 20, zone III.
4A. a. kahukuensis .... 115? Wilder & Thurston areas 225) 10:5 D7a5cale Se Ss
SPECIES CONCEPT
The species concept of Achatinella apexfulva is the same as that
maintained for A. mustelina (Welch, 1938). If two forms are found
in the same locality or in adjacent localities and are not found to
hybridize they are considered distinct species. If hybrids are found
between two forms A and B one is considered a subspecies of the
other.
In the “Manual of Conchology” (1912-1914, p. 275) the various
forms of Achatinella are divided into three large sections: Section
Bulimella Pfeiffer, section Achatinellastrum Pfeiffer, and section
Achatinella sensu strictu. The forms of the third section are divided
by Cooke into eight species: A. lorata, A. cestus, A. vittata, A. tur-
gida, A. apexfulva, A. decora, A. valida, and A. mustelina. Pilsbry
added three more, A. leucorraphe, A. swifti, and A. concavospira,
making 11 species in all. However, Pilsbry states:
It was hoped that characters might be found in the reproductive organs which
would aid to indicate specific boundaries, but dissections of A. lorata, vittata
simulans, and mustelina show no tangible structural differences.
22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
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 suggested. (1) Series of
lorata, for A. lorata. (2) Series of A. apexfulva, for A. apexfulva, turgida,
swiftit, leucorraphe, vittata, cestus. (3) Series of decora, for A. valida, decora,
mustelina, concavospira.
Since the publication of the Manual large collections of Achatinella
from a great number of carefully plotted localities have been acquired
for study by the Bishop Museum. These collections contain many
hitherto unknown intermediate forms and greatly enlarge the species
concept that resulted from the scant amount of material available in
1914. It is interesting to note that Pilsbry’s alternative grouping of
forms of Achatinella sensu strictu is almost the same as that derived
from the study of additional material. The main difference is that
instead of having species grouped under a series, each series
is considered a single species, species complex, or Rassenkreise
(Rensch, 1938) because intergrades for the most part are found
existing between the various species.
In most places live specimens of A. apexfulva are comparatively
rare. A general idea of the scarcity of the species can be obtained by
looking at table 2 (p. 15). Most of the large lots were collected
prior to 1932 by J. S. Emerson, Wilder, Cooke, Thurston, and
Meinecke. In most cases they represent accumulations of shells
acquired by numerous visits to the same place. The more recently
collected Meinecke material consists mostly of small lots.
Achatinella apexfulva varies greatly in form and color pattern in
different parts of the Koolau Range. Large sectors, however, pro-
duce no forms of A. apexfulva, the shells having either died out or
escaped discovery. From Palolo to Pauoa Valleys (fig. 7, p. 194)
there is a big gap in localities of A. apexfulva. But from Nuuanu to
Opaeula Gulch a nearly complete series of forms is found from ridge
to ridge, the forms more or less intergrading. Where a great deal of
careful collecting has been done on certain ridges such as those
from Waimano to Waiawa and those between South Kaukonahua
Stream and Opaeula Gulch, certain areas are found to contain defi-
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 23
nite color forms at different elevations. Moreover, a color form be-
longing to a definite area is separated from the color form of an-
other area by a region containing shells which have an intermediate
color pattern or a mixture of the color patterns of the areas on
either side of it. The extent of these intermediate localities has not
yet been well determined, for the work in the Koolau Range is still
in the pioneer stage because of the rareness of the shells, the rough-
ness of the terrain, and the vast extent of the Koolau Range. The
shells in a border region between two areas may have a color pattern
which is a mixture of those of an upper and lower area and is also
intermediate between those of the shells occurring on parallel ridges
on either side of the area. From a study of the carefully collected
regions of the Koolau. Range and a study of the collection as a
whole, so that the trend in the differentiation of Achatinella is de-
termined, Achatinella sensu strictu is grouped into the following six
species: 1, A. lorata Férussac; 2, A. apexfulva Dixon; 3, A. turbini-
formis Gulick; 4, A. concavospira Pfeiffer; 5, A. decora Feérussac; 6,
A. mustelina Mighels. This paper concerns only A. apexfulva; a
discussion of species I, 3, 4, and 5 will be reserved for future papers.
Subspecific groups will be used in this paper to group together
the lower orders of forms of A. apexfulva, as is done for A. mustelina
(Welch, 1938). The term “variety” will also denote that a particular
form is closely related to a particular subspecies but has not enough
distinctive characters to warrant its separation into a definite sub-
species. All varieties will be numbered var. 1, var. 2, etc.
TAXONOMY
SYNOPSIS OF THE SUBSPECIES OF ACHATINELLA APEXFULVA DIXON
The following synopsis is purely an artificial grouping of forms
that seem to be similar. No great detail will be attempted because
so few really distinct characters exist beyond the shape and color of
the embryonic whorls and size of the shells. The color pattern of the
postembryonic whorls in some regions is of use, but very often it is
of little aid since the same pattern may occur again and again in widely
separated forms. Unless otherwise stated the color pattern of the
shell is the color of the last two postembryonic whorls.
A. Embryonic whorls bicolored. First embryonic whorl a dark color
(black, gray, light brown), remaining embryonic whorls white, cream
buff, or some shade of yellowish white. The dark band or line usually
fades out, on the first quarter or first half of the second embryonic
whorl, into the ground color or into a light yellowish-white band or
24. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
line at the lower edge of the whorl, which finally may fade out en-
tirely on the last quarter whorl.
a. Embryonic whorls pointed. First embryonic whorl light reddish
brown (pecan brown, army brown, walnut brown, sepia), remaining
embryonic whorls usually white. Exceptions are A. a. buena, A. a.
cinerea, and A. a. hanleyana, which have yellow or yellow-banded
embryonic whorls (such as pinkish buff, tilleul buff, cinnamon buff).
I. Small usually lowland forms, occurring mostly in zone II," re-
gions I, 1a, which may be white or buff, lined, banded, streaked, or
spotted with brown or reddish brown, or the patterns may be gray or
gray brown. The mean length ranges from 15.5 to 20.5 mm.
Group of A. a. cestus Newcomb
. muricolor, new subspecies
. waialaeensis, new subspecies
. forbesiana Pfeiffer
. fuscostriata, new subspecies
. nnotabilis Smith
cestus Newcomb
simulator Pilsbry and Cooke
buena, new subspecies
globosa Pfeiffer
hanleyana Pfeiffer
simulans Reeve
HH
FO Oe ia age sree he eee
SA RRRRA RR AD
eae aeegegeeagneaen
e
II. White obese forms lined with reddish brown having a mean
length of 18.5 to 20.5 mm., and occurring in zone III, regions I,
Ia. They are larger than shells belonging to the group of A. a. cestus.
Group of A. a. vittata Reeve
1. A. a. vittata Reeve
2. A. a. cinerea Sykes
3. A. a. albofasciata Smith
4. A. a. oliveri, new subspecies
5. A. a. rubidipicta, new subspecies
b. Embryonic whorls blunt, usually banded with yellow. Forms
occurring between North Waiawa Stream and the upper portion of
the North-South Kaukonahua Ridge.
I. The color pattern may have the white ground tinted or banded
with buff or gray, or the ground may be buff or a light shade of gray.
The shell is usually lined or banded with gray, reddish brown, or
1 For discussion of zones and regions see pp. 189, 190, and figs. 7, 8, pp. 194, 195.
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 25
brown and is found in zone II, region III, from Waiawa to Kipapa
DMN cis tows Sadek Oe Group of A. a. polymorpha Gulick
polymorpha Gulick
flavitincta, new subspecies
lemkei, new subspecies
lineipicta, new subspecies
fumositincta, new subspecies
a aS AS
oe a
Note——Some specimens of A. a. polymorpha and its varieties lack
the dark color, such as black or gray, on the first embryonic whorl.
Il. The pattern is a strongly axially streaked one, the color brown-
ish gray or light tan, impressed sutural band white, brown, or buff.
The forms are limited to zone II, region III, between Kipapa Gulch
and South Kaukonahua Stream. Exception, A. a. tuberans, which
has banded forms that may or may not be strongly axially streaked.
Group of A. a. leucorraphe Gulick
A. a. virgatifulva, new subspecies
A. a. leucorraphe Gulick
3. A. a. tuberans Gulick
III. White shells lined or banded with reddish brown, gray, and
sometimes yellow occurring in zone III, region III. :
Group of A. a. irwint Pilsbry and Cooke
how
I. A. a. ewaensis, new subspecies
2. A. a. irwini Pilsbry and Cooke
B. Embryonic whorls unicolored.
a. Embryonic whorls white.
I. Embryonic whorls very pointed to moderately pointed. Small
shells from the lowland zone or zones I and II, region II; mean
length from 17.5 to 18.5 mm...Group of A. a. pilsbryi, new subspecies
A. a. pilsbryi, new subspecies
a. roseata, new subspecies
a. laurani, new subspecies
A. a. parvicolor, new subspecies
Ade
oa
aoe Nic
A. a. laurani and A. a. parvicolor are intermediate forms between
a lower smaller race with very pointed embryonic whorls, occurring
in lower zone I, and a large obese race of shells inhabiting an upper
zone or zone III. I am merely putting them in the subspecific group
of A. a. pilsbryi because they are closer to the form of pilsbryi than
to the upper group of A. a. turgida.
26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
If. Large obese shells found mostly in zone III, region IT, and
zone II, region IIIa, between Moanalua and Waiawa Streams. Em-
bryonic whorls moderately pointed, usually white but sometimes
cream buff; mean length, 18.5 to 21.5 mm.
Group of A. a. turgida Newcomb
1. A. a. ovum Pfeiffer
2. A. a. bruneola, new subspecies
3. A. a. aureola, new subspecies
4. A. a. waimaluensis, new subspecies
5. A. a. turgida Newcomb
6. A. a. meadowsi, new subspecies
7. A. a. perplexa Pilsbry and Cooke
8. A. a. nigripicta, new subspecies
g. A. a. cookei Baldwin
10. A. a. simulacrum Pilsbry and Cooke
11. A. a. rubidilinea, new subspecies
12. A. a. chromatacme Pilsbry and Cooke
13. A. a. griseibasis, new subspecies
14. A. a. albipraetexta, new subspecies
Note.—A. a. chromatacme differs from the remaining subspecies
of this group in having shells with darker embryonic whorls of tan
or yellowish brown. A. a. griseibasis has more loosely coiled em-
bryonic whorls which are intermediate between the embryonic whorls
of the groups of A. a. turgida and A. a. leucorraphe.
b. Embryonic whorls white in a few forms, but usually they are
tinted with buff or yellow, or are a deep shade of yellow or yellowish
brown.
I. Small shells probably from zone I or lower zone II, region IV,
between South Kaukonahua Stream and North Helemano Stream.
All forms extinct as far as is known. Gray or gray-brown lined or
streaked color patterns, embryonic whorls white or tinted cream color.
Group of A. a. coniformis Gulick
1. A. a. coniformis Gulick
2. A. a. apexalba, new subspecies
3. A. a. versicolor Gulick
4. A. a. wahiawa, new subspecies
II. Shells with color patterns of light pastel shades of pink, yellow,
deep cream, or blue gray, usually lightly axially streaked with simi-
lar colors darker than the ground color. They occur in zone II and
probably zone I, region IV, between South Kaukonahua Stream and
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 27
North Kaukonahua Stream. Exceptions are A. a. steeli, A. a. gulickii,
which are banded with reddish brown or dark gray and strongly
axially streaked; A. a. flavidus, which may lack streaking or may be
Peed With DFOWM.. 6. ce ee a Group of A. a. lilacea Gulick
a. gulicku Smith
a. flavida Gulick
a. lilacea Gulick
a. glaucopicta, new subspecies
a. punicea, new subspecies
a. steeli, new subspecies
Am BW ND
SAR AD
III. Strongly axially streaked brown and gray color forms from
zone II and the lower part of zone III, region IV, between Poamoho
Stream and Kawainui Gulch. Embryonic whorls usually a dark yel-
lowish brown. Exceptions are A. a. brunosa and A. a. kawatiki, which
have lighter embryonic whorls..... Group of A. a. apicata Newcomb
. brunosa, new subspecies
. suturafusca, new subspecies
. suturalba, new subspecies
. apicata Newcomb
. paalaensis, new subspecies
. kawatiki, new subspecies
Am RW D 4
i ks
(“ye t=> {sy sts) ats} is)
IV. Not markedly axially streaked, white or light color patterns.
The color is mostly light pink, light gray, flesh tints, usually banded
over white or colors almost solid on the last two whorls. Exceptions
are the dark gray color pattern of A. a. cervixnivea, the reddish-brown
pattern of A. a. beata, which may resemble typical apexfulva, and
also A. a. kahukuensis which has a white spire and a yellow base.
These forms inhabit zone III and the upper part of zone II, region
IV, from North Kaukonahua Stream to the Malaekahana-Kaluakauila
ERR ve hess do sie alaie «= Group of A. a. aloha Pilsbry and Cooke
a. aloha Pilsbry and Cooke
a. roseipicta, new subspecies
. poamohoensis, new subspecies
. cervixnivea Pilsbry and Cooke
. beata Pilsbry and Cooke
vespertina Baldwin
. kahukuensis Pilsbry and Cooke
N Oo BP DW ND
PS RR RS AS AS AS AS
SS
e as
28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
V. Small lowland forms from zones I and II, region IV, north-
west of Helemano Stream. Embryonic whorls white, cream, or
yellow. Mean length, 16.5 to 18.5 mm.
Group A. a. apexfulva Dixon
. apexfulva Dixon
. duplocincta Pilsbry and Cooke
bakeri, new subspecies
. napus Pfeiffer
. leucozona Gulick
paumaluensis, new subspecies
. oloensis, new subspecies
. thiihiensis, new subspecies
. wailelensis, new subspecies
AP A
S 2 ea 8 Ss 2 Se
0 WN ANRYW No
Group oF A. A. CESTUS NEWCOMB
ACHATINELLA APEXFULVA FORBESIANA Pfeiffer
PLATE I, FIGURE 2; PLATE 4, FIGURES 5-6a
Achatinella forbesiana P¥rtrFER, Proc. Zool. Soc. London, 1855, p. 5, pl. 30,
fig. 16.
Achatinella cestus NEwcoms, Pilsbry and Cooke, Man. Conch., vol. 22, pp. 286,
288, pl. 52, figs. 11-11b, 13 (only), 1914.
The last two whorls are usually brown, lined with white; banded
with a white band at the edge of the periphery of the last whorl;
embryonic whorls pointed and bicolored.
The last two whorls of the lectotype (pl. 4, fig. 5) are lined with
white; penultimate vinaceous drab, last whorl vandyke brown and
rood’s brown, peripheral band white; lip and columella callus vina-
ceous buff. Length 18.4 mm., greater diameter 11.2 mm., spire
height 9.3 mm., number of whorls 64.
Distribution, area 6??: Waialae, Gulick. The race probably oc-
curred at a low elevation in either Waialae Nui or Waialae Iki
Gulch (fig. 3a, p. 29). Gulick also reports it from Wailupe, but this
may be an error due to mixture of material from different localities.
Lectotype in BM.
The usual form is sinistral (pl. 1, fig. 2) ; postembryonic whorls
pale pinkish cinnamon, streaked with sorghum brown and pale mouse
gray, last two whorls lined with pale pinkish cinnamon and white,
peripheral band white ; lip cinnamon ; impressed sutural band cinnamon
buff. Length 16.9 mm., greater diameter 11.0 mm., spire height
g.0 mm., number of whorls 6.
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 29
A dextral shell (pl. 4, fig. 6) has a light gray color pattern;
postembryonic whorls pale smoke gray darkening to ecru drab on the
last whorl, last whorl and a half lined and banded with white. Length
17.8 mm., greater diameter 11.3 mm., spire height 9.5 mm. A narrow
dextral (pl. 4, fig. 6a) measures: Length 17.3 mm., greater diameter
10.5 mm., spire height 10.0 mm.; color pattern similar to plate 1,
figure 2.
Manoa
Fic. 3——Eastern Oahu, leeward slope of the Koolau Range, including region I
(fig. 7, p. 194), showing localities of subspecies of A. apexfulva belonging to the
groups of A. a. cestus and A. a. vittata.
Fic. 3a.—Same as fig. 3, but exhibiting the area occupied by different subspecies of
A, apexfulva belonging to the groups of A. a. cestus and A. a. vittata.
ACHATINELLA APEXFULVA WAIALAEENSIS, new subspecies
‘PLATE I, FIGURE 3; PLATE 4, FIGURES 2-4
This race has a characteristic axially streaked color pattern, and
lacks the spiral lines and bands of A. a. forbesiana, although it has
the white peripheral band on the last whorl. A. a. forbesiana is an
extinct race which probably existed at a low elevation immediately
below the area occupied by A. a. waialaeensis.
The holotype of A. a. waialaeensis (pl. 1, fig. 3) has the first
fourth postembryonic whorl mikado brown, remainder of the first
30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
postembryonic whorl and penultimate whorl drab gray axially streaked
with benzo brown, the first half of the last whorl is drab axially
streaked with bone brown, tinted below the tawny impressed sutural
band with tawny, last half whorl changes from drab to tawny axially
streaked with bone brown, peripheral band white; lip and columella
callus army brown. Length 18.8 mm., greater diameter 11.5 mm.,
spire 10.5 mm., number of whorls 64.
Distribution, area 2: Waialae Iki-Waialae Nui Ridge, type lo-
cality 51Aa, elevation 1,250-1,350 feet, Meinecke, 1933; also 51A*,
elevation 1,000-1,350 feet, Meinecke, 6 sinistral 1926, I sinistral
1928; Waialae Nui Gulch, locality 50A*, elevation 1,400-1,450 feet,
Meinecke 1926, 1 sinistral 1929. Also collected in the region of
Waialae Iki-Waialae Nui Ridge by O. H. Emerson, BBM 103924,
1916, BBM 103926, 7 sinistral 1918; Waialae, Wailupe, Gulick.
Area 3: Waialae Iki-Waialae Nui Ridge, locality 50A-3, eleva-
tion 1,500-1,650 feet, O. H. Emerson, BBM 167275-167276, 6 sinis-
tral 1937 (figs. 3, 3a, p. 29).
The lightest color pattern (pl. 4, fig. 2) is gray; postembryonic
whorls pale ecru drab streaked with drab gray and ecru drab, tinted
on the last half whorl with sayal brown, and streaked with verona
brown, impressed sutural band mikado brown.
Plate 4, figures 2a and 3, show reddish-brown patterns of this
race which are peculiar to the shells of Waialae. The shell of figure 2a
has dark vinaceous brown postembryonic whorls streaked with drab
gray or deep olive buff. That of figure 3 has mikado brown post-
embryonic whorls darkening to cameo brown on the last whorl, at
the edge of the periphery a band of cinnamon buff.
In locality 50A-3 only dextrals occur. The typical form (pl. 4,
fig. 4) has the last two postembryonic whorls pinkish buff axially
streaked with saccardo’s umber and sepia, last half whorl snuff brown.
Length 18.2 mm., greater diameter 12.0 mm., spire height 9.6 mm.
ACHATINELLA APEXFULVA MURICOLOR, new subspecies
PLATE 1, FicguRE 1; PLATE 4, FIGURES I-Ib
Achatinella cestus NEwcoms, Pilsbry and Cooke, Man. Conch., vol. 22, p. 288,
pl. 52, figs. 9, 9a, 10 (only), 1914.
The shell resembles A. a. waialaeensis but differs in having the
dominant pattern always profusely lined with white over a gray
ground. No reddish-brown color forms occur in this race and the
characteristic white peripheral band is usually lacking, but may be
present.
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 31
The postembryonic whorls of the holotype (pl. 1, fig. 1) are drab
gray shading to drab, axially streaked on the last two whorls with
snuff brown and bister, spirally lined with white; impressed sutural
band cinnamon; lip and columella callus fawn color. Length 17.3 mm.,
greater diameter 11." mm., spire height 9.3 mm., number of
whorls 6.
Distribution, area 1: Niu-Wailupe Ridge, type locality 41-4",
elevation 1,659-1,750 feet, Meinecke, BBM 120184, 120177, 1926,
also BBM 120107, 6 sinistral 1931; Kului-Laulaupoe Ridge, Wai-
lupe, locality 41A-1*, elevation 1,800-2,000 feet, Meinecke, BBM
120216-120217, 1926; Laulaupoe-Wailupe Ridge, Wailupe, 41B-
4*, elevation 1,950-2,000 feet, Meinecke, BBM 120225, 1926 (figs. 3,
3a, p. 29).
The lightest color form of the shell (pl. 4, fig. 1) has the postem-
bryonic whorls pale drab gray axially streaked with light drab; 1m-
pressed sutural band pale pinkish cinnamon streaked with light pink-
ish cinnamon; last whorl lined with white. This color pattern re-
sembles the light color pattern of A. a. waialaeensis (pl. 4, fig. 2),
but muricolor is definitely more lined and lacks the strong white
peripheral band. An obese shell (pl. 4, fig. Ia) measures: Length
17.3 mm., greater diameter 11.8 mm., spire height 8.6 mm.; the post-
embryonic whorls strongly lined, last two whorls tilleul buff lined
and banded with wood brown, natal brown, snuff brown, and bister,
peripheral band white; lip and columella callus wood brown.
A rare dark color pattern (pl. 4, fig. 1b) occurs on 7 specimens out
of a total of 194 taken at various times from locality 41-4. The last
two whorls are drab lightly axially streaked with vinaceous and
sepia, faintly lined with pale pinkish buff.
ACHATINELLA APEXFULVA CESTUS Newcomb
PLATE I, FIGURE 5; PLATE 4, FIGURES Q-II
Achatinella cestus Newcomps, Proc. Zool. Soc. London, 1853, p. 132, pl. 22,
fig. 8—Puitssry and Cooke, Man. Conch., vol. 22, ). 286, pl. 20, fig. 8; pl. 55,
fig. 1 (only), 1914.
This form is characterized by having a light yellowish-white ground
splotched or rarely banded on the last two whorls above the periphery
with reddish brown, and by having a dark reddish-brown band below
the periphery; the base may be banded or splotched with reddish
brown.
The lectotype (pl. 4, fig. 9) is marked with an “x” by me. The last
two whorls are white streaked with splotches of walnut brown and
3
32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
chocolate, below the periphery a band of chocolate and a band of
white; lip and columella callus light brownish vinaceous. Length
17.7 mm., greater diameter 11.6 mm., spire height 9.6 mm., number
of whorls 6.
Distribution, area 5?: Palolo, Gulick; Waiomao Stream, Palolo,
Spalding, somewhere in the region of area 5? (figs. 3, 3a, p. 29).
Gulick also reports specimens from Wailupe and Waialae, but I be-
lieve these are errors due to a mixture, because the known highland
forms of this group from Waialae have distinctly different patterns
from those of Palolo, and the lowland forms, if not extinct today,
most probably would show a similar variation. Also collected by
D. D. Baldwin, BBM 54741. Lectotype BM.
The Spalding lot contains only specimens of A. a. cestus and no
specimens of A. a. fuscostriata, with which form A. a. cestus is
always mixed in the Gulick collection. Pilsbry (Pilsbry and Cooke,
1912-1914, p. 287) mentions the fact that the Spalding lot contains
dextral specimens of A. a. simulator which are undoubtedly A. a. simu-
lator var. I of area 4. Judging from the data at hand, I believe that
cestus is a distinct race from A. a. fuscostriata and probably grades
into the higher race of A. a. simulator var. 1.
Specimen A of the BM type lot (pl. 4, fig. 10) has the white ground
of the last whorl above the periphery streaked with zigzag lines and
axial streaks of chocolate, below the periphery banded with chocolate
and white.
In one BBM lot labeled Wailupe (pl. 4, fig. 11) a light color pattern
has the postembryonic whorls pale pinkish cinnamon, lightly streaked
with avellaneous, below the periphery of the last whorl a band of
russet and a band of buckthorn brown.
A sinistral (pl. 1, fig. 5) resembles the lectotype; postembryonic
whorls pale pinkish buff, splotched and streaked with russet, at the
edge of the periphery a band of white, below the periphery a band of
russet, a band of white, and a line of tawny.
‘ACHATINELLA APEXFULVA FUSCOSTRIATA, new subspecies
PLATE 1, FIGURE 4; PLATE 4, FIGURES 7, 7a
Achatinella cestus Newcoms, Pilsbry and Cooke, Man. Conch., vol. 22, pp. 287,
288, pl. 52, figs. 12, 12a, 13a, 14, 14a (only), 1914.
This extinct form resembles A. a. cestus but differs in not hav-
ing the characteristic dark reddish-brown subperipheral band. The
ground color is usually darker, being a yellowish cream instead of
the white or whitish cream color of A. a. cestus. Both forms have a
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 33
zigzag pattern and probably came from adjacent areas. Possibly
fuscostriata,came from some area below A. a. cestus in Palolo or
Waialae Nui somewhere in the region of area 7??.
The postembryonic whorls of the holotype (pl. 1, fig. 4) are pinkish
buff axially streaked with straight and zigzag lines of russet, last
whorl with a white peripheral band, base banded with white and
carob brown; lip and columella callus avellaneous. Length 16.4 mm.,
greater diameter 11.2 mm., spire height 8.4 mm., number of whorls 6.
Distribution, area 7??: Type locality Palolo, Gulick ; also Waialae,
Wailupe, Gulick (fig. 3a, p. 29). So much of the Gulick material is
undoubtedly mixed from several valleys that the true distribution of
the lowland races of Achatinella cannot be worked out from his data,
if all his labels are considered correct. I do not believe this form
ever occurred in Wailupe.
The color pattern may lack a white peripheral band on the last
whorl (pl. 4, fig. 7), postembryonic whorls pale pinkish buff streaked
with cinnamon, base banded with white. A dextral specimen (pl. 4,
fig. 7a) has the postembryonic whorls cream buff, axially streaked
with cinnamon, the last whorl banded above and below the edge of
the periphery with a band made up of broken lines of cinnamon, basal
band white and pinkish cinnamon.
ACHATINELLA APEXFULVA INNOTABILIS Smith
PLATE 4, Ficure 8
Apex innotabilis SmirH, Proc. Zool. Soc. London, 1873, p. 78, pl. 9, fig. 23
(not fig. 19).
Achatinella swiftii Newcoms, Pilsbry and Cooke, Man. Conch., vol. 22, pp. 306,
309, pl. 59, fig. 7 (only), 1914.
The holotype in the British Museum is marked with a red spot of
sealing wax. The embryonic whorls are similar to A. a. cestus;
postembryonic whorls are light pinkish cinnamon axially streaked
with a darker shade of the ground color, peripheral band pale pinkish
buff ; lip pale pinkish buff ; columella callus white. Length 18.1 mm.,
greater diameter 12.0 mm., spire height 10.6 mm., number of whorls 6.
This color form is undoubtedly a distinct race which is now ex-
tinct. Specimens of A. a. innotabilis are rare and are found only in
collections made prior to 1900. The form is closest to that of A. a.
fuscostriata. It was possibly a lowland race occurring below or near
fuscostriata. A. a. fuscostriata may be an intermediate race between
A. a. innotabilis and A. a. cestus. A. a. innotabilis possibly occurred
in Wailupe.
34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
ACHATINELLA APEXFULVA SIMULATOR Pilsbry and Cooke
PLATE I, FIGURE 6; PLATE 4, FIGURES I2-13a
Achatinella cestus color form simulator Pitspry and Cooke, Man. Conch.,
vol. 22, p. 287, pl. 55, figs. 2, 3, 4, 1914.
The holotype of A. a. simulator selected by Pilsbry and myself is
figured in the Manual of Conchology (pl. 55, fig. 2) and illustrated
in this paper on plate 4, figure 12. The postembryonic whorls are
white lined with chocolate except for a line of pecan brown on the
last whorl below the suture; sutural band white. Length 17.8 mm.,
greater diameter 12.2 mm., number of whorls 6.
Distribution. area?: Palolo, Gulick. Pilsbry (Pilsbry and Cooke,
IQI2-1914, p. 287), says: “A small lot; No. 42 coll. Irwin Spalding,
consists of dextral shells which occurred with the typical color form
of cestus.” This form is not typical simulator but is what I consider
to be A. a. simulator var. 1. Typical simulator is a lowland race now
extinct.
The color pattern of the holotype is a rare one occurring on only
one specimen in the Gulick lots in the Bishop Museum. The usual
color pattern and form of the shell (pl. 1, fig. 6) has white postem-
bryonic whorls, lined on the last whorl with pinkish cinnamon, about
the base a band of cinnamon; lip pale cinnamon pink slightly corru-
gated or roughened. Length 17.4 mm., greater diameter 11.5 mm.,
spire height 9.2 mm., number of whorls 6.
The shell may be more obese (pl. 4, fig. 13), length 17.2 mm.,
greater diameter 12.3 mm., spire height 8.5 mm., last whorl lined
with cinnamon about the periphery, and army brown about the base.
Dextral forms were also found by Gulick. A dextral specimen
is figured on plate 4, figure 13a.
ACHATINELLA APEXFULVA SIMULATOR var. 1
PLATE I, FIGURE 7
Area 4: Waiomao Stream, Palolo, locality 62, elevation 1,250-
1,350 feet, Meinecke, 8 dextral 1933, and 13 dextral 1914 (figs. 3, 3a,
p. 29).
The shell is very close to A. a. simulator and differs only in the
arrangement of the bands. Typical A. a. simulator always has a line
or band at the edge of the periphery, and the bands and lines are
never broken by axial streaks of a lighter color. The typical form of
this variety (pl. 1, fig. 7) has white postembryonic whorls, last
whorl above the periphery with a faint band of tilleul buff, below the
periphery the last whorl banded with mikado brown, which bands
NOS) T ACHATINELLA APEXFULVA DIXON—WELCH 35
are streaked with white, in the umbilical region a band of naples
yellow; lip and columella callus light vinaceous fawn. Length
17.0 mm., greater diameter 11.8 mm., spire height 9.2 mm., number
of whorls 6.
ACHATINELLA APEXFULVA BUENA, new subspecies
PLATE 1, FicurRE 8; PLATE 4, FIGURES 14, I4a
The shell is small, sinistral, and seems to be closest in form to
A. a. simulator but may be a race which occurred near A. a. hanleyana.
The yellow-banded embryonic whorls are different from anything
known from the region of Wailupe to Kalihi Valleys. However, be-
cause of the darkened embryonic whorls and the small size of the
shells, this distinct color form is placed under the subspecific group
of A. a. cestus. The first embryonic whorl of the holotype (pl. 1,
fig. 8) is army brown, next whorl, upper eighth white, lower seven-
eighths shades from avellaneous to chamois, last embryonic whorl,
upper half white, lower half chamois; postembryonic whorls white
banded just above the edge of the periphery with a band of russet
flecked with light buff, on last whorl supraperipheral band not so
intense, having more streaks and flecks of light buff in it, peripheral
band white except for a line of tawny on the lower portion of the
band, base tawny axially streaked with light buff, with a line of
white and a band of cream color in the umbilical region; lip and
columella callus vinaceous buff. Length 16.4 mm., greater diameter
10.6 mm., spire height 8.8 mm., number of whorls 53.
This subspecies has been named after Buena Blok, the wife of
Arthur Blok, of London, an ardent student of conchology.
Distribution, area?:. Nuuanu, west side, J. S. Emerson, BBM
102185, 33302.
This form appears in the Gulick collection in several lots. In one
lot from Palolo one specimen is found mixed with A. a. simulator.
Gulick has another lot labeled Waipio BBM 70382, and still another
Kalaikoa BBM 106736. It is probable that the Emerson and Gulick
shells all came from a single now extinct lowland locality, but the
exact location is not known. Another specimen was obtained from
A. Block, BBM 165793, and one from Sowerby and Fulton, BBM
165794.
A narrow specimen (pl. 4, fig. 14) measures: Length 16.6 mm.,
greater diameter 10.0 mm., spire height 8.9 mm.; the postembryonic
whorls are more heavily banded than the type, the bands are wood
brown flecked with vinaceous buff. A light color form (pl. 4, fig. 14a)
36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
has the postembryonic whorls white, banded on the penultimate above
the periphery with a narrow band of russet, last whorl lined just
above the periphery with a line of tawny, below the periphery ground
white tinted with cream color, base banded with tawny and lined
with cream color.
ACHATINELLA APEXFULVA GLOBOSA Pfeiffer
’ PLATE I, FIGURE 12; PLATE 4, FIGURES 21-24
Achatinella globosa PFEIFFER, Proc. Zool. Soe. London, 1855, p. 7, pl. 30, fig. 25.
Achatinella vittata Reeve, Pilsbry and Cooke, Man. Conch., vol. 22, pp. 289-2901,
pl. 30, fig. 25; pl. 57, fig. 2b (only), 1914.
The lectotype of A. a. globosa (pl. 4, fig. 21) in the British
Museum is marked “x” by me and is a deformed specimen. There is a
break in the shell at the second embryonic whorl, which accounts for
the depressed spire. The last postembryonic whorl is abnormally
rounded and enlarged; postembryonic whorls white lined with cinna-
mon buff, peripheral band cinnamon buff, subperipheral band white,
basal band hazel. On the aperture side of the last whorl, below the
periphery, there is another break in the shell which has been repaired
by the animal, and the pattern is lacking on approximately 2 mm. of
the shell. The shell is thin, sutural band white; the lip is little thick-
ened and colored pale vinaceous drab; the columella callus white.
Length 16.4 mm., greater diameter 12.8 mm., spire height 6.5 mm.,
number of whorls 54.
Specimen A of the type lot (pl. 4, fig. 22) more accurately repre-
sents the usual form of A. a. globosa. The postembryonic whorls
white lined with pinkish buff, last whorl banded with pinkish buff,
axially streaked with white, in the umbilical region a band of cinna-
mon buff, darkening near the edge of the lip to mikado brown.
Length 16.8 mm., greater diameter 11.4 mm., number of whorls 6.
Distribution: Nuuanu, Gulick, Baldwin, BBM 11884, J. S.
Emerson, BBM 102196; Kalihi, Gulick. Specimens also obtained by
Welch in London from Blok and Fulton.
A. a. globosa is a distinct subspecies which differs from vittata in
being a smaller shell, always dextral and having a different range of
color patterns. It occurs mixed with A. a. vittata in the Gulick col-
lections, and odd specimens identifiable as globosa are usually found
with typical forms of vittata in other collections.
The type lot of A. a. vittata, consisting of three specimens, has two
specimens which are color, patterns of A. a. globosa. One of these
marked A (pl. 4, fig. 23) is a light color pattern and a narrow form
On 1 ACHATINELLA APEXFULVA DIXON—WELCH 37
of the shell. The postembryonic whorls are white lined and banded
on the last whorl with hazel, in the umbilical region a patch of carob
brown; lip and columella callus vinaceous buff. Length 17.7 mm.,
greater diameter 11.2 mm., number of whorls 54.
The other specimen B has the usual A. a. globosa pattern similar
to plate 1, figure 12. The type lot of three shells of A. a. globosa
has one specimen marked B which is a typical A. a. vittata.
The usual form of the shell and color pattern of A. a. globosa
in the Bishop Museum (pl. 1, fig. 12) has white postembryonic
whorls banded and lined with hazel, base entirely hazel. Length
17.4 mm., greater diameter 11.7 mm., spire height 9.0 mm. A speci-
men from the Gulick collection (pl. 4, fig. 24) has white postembry-
onic whorls spirally banded with cinnamon buff, which darkens on the
last eighth whorl to mikado brown; impressed sutural band cinnamon.
ACHATINELLA APEXFULVA HANLEYANA Pfeiffer
PLATE I, FIGURE 16; PLATE 4, FIGURES 25, 26
Achatinella hanleyana P¥FetrreR, Proc. Zool. Soc. London, 1855, p. 202.
Achatinella lorata Férussac, Pilsbry and Cooke, Man. Conch., vol. 22, pp. 278,
281, IQT4.
To quote from Pilsbry and Cooke:
Mr. Sykes remarks that this is “related to the form of A. lorata described
as A. nobilis, and may prove to be an extreme variety.” Dr. C. Montague Cooke,
on examining the type in the British Museum considered it an artificially colored
lorata.
The lectotype in the British Museum is not an artificially colored
lorata but a normal shell which is a distinct subspecies and related
to A. a. globosa. A. a. hanleyana closely resembles A. a. globosa in
the shape of the bicolored embryonic whorls and in the size and color
of the shell. A. a. hanleyana, however, is not banded with white
but has a solid brown pattern above the periphery. No 4. a. lorata
has bicolored embryonic whorls similar to those of A. a. globosa.
A. a. hanleyana, however, can easily be confused with brown forms
of Partulina radiata Gould (see Pilsbry and Cooke, 1912-1914, p. 49,
pl. 13, fig. 8). The two forms are so close that I have been in doubt as
to whether or not to consider hanleyana a Partulina. However, I am
inclined to believe that it is an extinct lowland Achatinella, because
of its similarity to A. a. globosa and to a sinistral Achatinella (pl. 4,
fig. 26) obtained by Dos Santos in Kalihi Valley, which has embryonic
whorls very similar to those of A. a. hanleyana (pl. 1, fig. 16). A. a.
hanleyana, moreover, has an impressed sutural band, a columella cal-
38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
lus, and a rugose texture to the surface of the shell, not usual in
specimens of Partulina from Oahu. Partulina radiata is placed by
Pilsbry with Partulinas from the Island of Maui. In some collections
the shells are labeled Maui. However, radiata is typically an Oahu
shell and probably comes from the region of Nuuanu Valley.
The lectotype (pl. 4, fig. 25) has the first embryonic whorl colored
terra cotta, next whorl upper half cartridge buff, lower half shades
from terra cotta to pale vinaceous fawn and finally cartridge buff,
remainder of the embryonic whorls cream buff; the first half post-
embryonic whorl has the shell nacre worn off, remaining postem-
bryonic whorls pinkish buff, axially streaked with close-set lines of
hazel; lip and columella callus cartridge buff. Length 17.8 mm.,
greater diameter 12.0 mm., number of whorls 6. The surface of the
shell does not seem smooth but slightly rugose as compared with
A. a. globosa, but this rugosity of the surface may be an illusion
caused by the axial streaking.
From Arthur Blok I obtained a specimen (pl. I, fig. 16) which
greatly resembles the holotype of A. a. hanleyana in form and color
pattern. The postembryonic whorls ochraceous buff, axially streaked
with tawny, subperipheral band ochraceous buff; the shell has a
rugose appearance; impressed sutural band same as ground color of
the shell. Length 19.1 mm., greater diameter 12.4 mm., spire height
9.9 mm.
In the Thurston collection a sinistral shell (pl. 4, fig. 26) col-
lected by Dos Santos and reported from Kalihi has a ground color
that matches that of A. a. hanleyana. The main difference is that
the shell is heavily banded with dark bands. The postembryonic
whorls are warm buff axially streaked with ochraceous tawny and
banded with liver brown; lip and columella callus pale vinaceous fawn.
This shell closely resembles the form of A. a. simulans var. 1 on the
Kamanaiki-Kalihi Ridge. Probably it came from somewhere in the
Kalihi region at a similar elevation, possibly on the north side of the
valley. This sinistral shell may come from a colony of pure sinistral
hanleyana of which this is the dark color form. The dextral race is
probably from a colony of still lower elevation occurring directly
below the sinistrals.
ACHATINELLA APEXFULVA HANLEYANA var. 1
PLateE I, Figure 17; PLATE 4, FIGURE 32
A peculiar form of Achatinella apexfulva was procured from
Sowerby and Fulton in London. The shells were accompanied by no
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 39
data concerning their locality, but the fact that they have pointed
globosa-like embryonic whorls leads me to believe that they are a
lowland form of hanleyana which comes from somewhere in the
general region of Nuuanu to Kalihi. However, I may be wrong in this
belief and the shells may be lowland forms of Partulina radiata
Gould. The first embryonic whorl (pl. 1, fig. 17) is fawn color, next
whorl pale pinkish cinnamon, shading to pinkish buff ; postembryonic
whorls pinkish buff, axially streaked with mikado brown and spirally
lined on the last whorl above the periphery with pinkish buff; im-
pressed sutural band the ground color; columella callus white; lip
tilleul buff. Length 20.8 mm., greater diameter 12.5 mm., spire
height 10.8 mm., number of whorls 64.
The shell may have a darker pattern (pl. 4, fig. 32). Postembryonic
whorls clay color, axially streaked with mikado brown, spirally lined
with cinnamon buff, last whorl below the periphery with a band of
white, about the base a band of cream color; lip outer edge pinkish
buff ; columella callus and inner edge of lip white. Length 19.9 mm.,
greater diameter 12.6 mm., spire height 9.8 mm.
ACHATINELLA APEXFULVA SIMULANS Reeve
PLATE I, FIGURE 19; PLATE 4, FIGURES 33-35a
Achatinella simulans REEvE, Conch. Icon., vol. 6, pl. 2, fig. 15, April 1850.
Achatinella vittata simulans RrEEveE, Pilsbry and Cooke, Man. Conch., vol. 22,
p. 292, pl. 57, fig. 8 (figs. 9-11, 14a, pl. 57, not 4. simulans of Reeve 1850,
but A. a. rubidipicta, new subspecies), 1914.
The shell is a small lowland form, with a dark brownish band in
the umbilical region, one at or just below the edge of the periphery,
one just below the periphery; above the periphery on the last whorl
just behind the edge of the lip a patch of brown or group of brown
lines.
The type lot of A. a. simulans contains three specimens none of
which agree with Reeve’s original figure (pl. 4, fig. 33). One specimen
(pl. 4, fig. 34) in the type lot approaches the color pattern of the
original figure and is considered the lectotype. It differs from the
original figure in having a flat columella callus and in not having a
subsutural band on the last two whorls and three dark chocolate sub-
peripheral bands. The embryonic whorls are bicolored similar to those
of A. a. globosa, first embryonic whorl deep mouse gray, later em-
bryonic whorls white; postembryonic whorls white with a band of
cinnamon just above or at the edge of the periphery, above the
periphery on the last whorl just behind the edge of the lip a streak
or patch of mikado brown, below the edge of the periphery a band of
40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
warm sepia about the base, surface rough and pitted; impressed
sutural band white; lip and columella callus cartridge buff. Length
18.5 mm., greater diameter 13.2 mm., number of whorls 53.
Distribution, area 16: Kamanaiki-Kalihi Ridge, locality 121-2,
elevation 1,300-1,350 feet, H. Lemke and H. Lemke, Jr., 1934, BBM
114979, HL 128 (figs. 3, 3a, p. 29); Kalihi, Gulick, BBM 70464,
Wilder, BBM 50544. Lectotype specimen “x” type lot BM marked
by me.
Until shells were found in area 16, the only region containing shells
similar to A. a. simulans was that of area 11. However, no white
specimen in area II is found having a dark band about the umbilicus
plus dark bands on the last whorl similar in arrangement to those
on either the lectotype or the original figure. What Reeve called
A. decora closely matches a color pattern of A. a. rubidipicta
(area II).
Plate 1, figure 19, shows the usual form of the shell in the Lemke
lot from area 16, and a color pattern close to that of figure 34; the
postembryonic whorls white, last whorl just below the impressed
sutural band and below the periphery lined with cinnamon, basal
band in the umbilical region sayal brown. Length 18.0 mm., greater
diameter 11.7 mm., spire height 9.4 mm., number of whorls 64.
An obese specimen (pl. 4, fig. 35) looks more like the form of
figure 34; the postembryonic whorls white, banded on the first post-
embryonic whorl with cinnamon, on the penultimate and last whorl
lined with chocolate, last whorl below the periphery with a line of
chocolate, two lines of clay color, a line of chocolate, and a band of
chocolate in the umbilical region. Length 18.7 mm., greater diameter
12.3 mm., spire height 9.6 mm. One specimen (pl. 4, fig. 35a) has
the last two whorls colored pale pinkish buff streaked with cinnamon
buff and clay color and spirally lined with verona brown and warm
sepia.
The color patterns of figures 35 and 35a indicate that yellow
forms may occur at higher elevations. Quite possibly A. a. simulans
is an intermediate race between a form of A. a. albofasciata and the
lower form of A. a. simulans var. I.
ACHATINELLA APEXFULVA SIMULANS var. 1
PLATE I, FIGURE 20; PLATE 4, FIGURES 36, 36a
Area 17: Kamanaiki-Kalihi Ridge, locality 121, elevation 1,300-
1,350 feet, Lemke and Lemke, Jr., 1934, BBM 114976, HL 127;
121-1, elevation 1,300 feet, Anderson and Kondo, 6 sinistral 1935
(figs. 3, 3a, p. 29).
INOS (I ACHATINELLA APEXFULVA DIXON—WELCH AI
The color patterns of area 17 tend to be darker than those of A. a.
simulans, although individual shells in both areas 16 and 17 may
have very similar patterns. A shell with the usual form and color
pattern of the shell (pl. 1, fig. 20) has the postembryonic whorls
white, banded with warm blackish brown, last whorl lined and tinted
above the periphery with cinnamon buff, and a line of cinnamon buff
on the white subperipheral band, about the umbilicus a band of cream
color; lip and columella callus white, tinted with pale vinaceous fawn.
Length 17.6 mm., greater diameter 12.0 mm., spire height 9.1 mm.,
number of whorls 64.
Another specimen (pl. 4, fig. 36) has the usual form and a light
color pattern of the shell; postembryonic whorls white, lined with
cinnamon buff and cinnamon; impressed sutural band white, on the
penultimate whorl, just above the suture of the whorl below, a line
of warm sepia, last whorl lined at the edge of the periphery with
warm sepia, base banded with carob brown and a band of black in
the umbilical region; lip and columella callus white, streaked with
pale brownish vinaceous. Length 17.7 mm., greater diameter 12.3 mm.,
spire height 9.0 mm. This specimen of figure 36 resembles Reeve’s
original figure of A. a. simulans. The darkest color pattern (pl. 4,
fig. 36a) is a warm blackish brown on the postembryonic whorls, last
whorl banded at the periphery with a band of white which is lined
with two lines of mikado brown, below the periphery a line of cinna-
mon buff ; impressed sutural band sayal brown shading to pale pinkish
buff at the upper edge.
ACHATINELLA APEXFULVA SIMULANS var. 2
PLATE I, FIGURE 23; PLATE 5, FIGURES 5-5b
Area 19: Manaiki-Moanalua Ridge, locality 151B*, elevation
I,400-2,150 feet, Meinecke, 1932, also collected by O. H. Emerson,
BBM 103963, A. MacAuley, in J. S. Emerson collection, BBM
102208, Thurston, BBM 131112.
Area 19A: Moanalua Stream, locality 150B-2, elevation goo feet,
Welch and W. Giffard, 1935 (figs. 4, 4@, p. 53).
The usual form of the shell (pl. 1, fig. 23) in area 19 has the
postembryonic whorls white, lined with tawny, with a patch of close-
set ochraceous tawny lines on the first half postembryonic whorl;
impressed sutural band tawny; peripheral and basal bands mars
brown or chocolate, with the bands in the umbilical region seal brown
or black; lip and columella callus light vinaceous fawn. Length 18.5
mm., greater diameter 12.3 mm., spire height 9.9 mm., number of
whorls 64.
42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
The shell may lack the dark chocolate bands (pl. 5, fig. 5) and have
the last two postembryonic whorls white, lined with pinkish buff
above the periphery, at and below the periphery with sayal brown,
base tinted with cream color. The postembryonic whorls may be
almost entirely white (pl. 5, fig. 5a) except for a patch of sayal brown
on the impressed sutural band on the first and fourth postembryonic
whorl, and a band and a line of ochraceous tawny on the base. A
yellowish pattern (pl. 5, fig. 5b) is very much like the yellowish pat-
tern of A. a. simulans (pl. 4, fig. 35a), and also that of A. a. rubidi-
picta (pl. 5, fig. 2d), postembryonic whorls pinkish buff, lined and
banded with ochraceous tawny and tawny, about the periphery and
below the ground is white, banded below the periphery with russet
deepening to chocolate on the last half whorl.
This race has been considered a variety of A. a. simulans because
of its small size and similarity in form. Ina fossil bed, locality 150B-2,
a few specimens of apexfulva are found and may be considered
the extreme limit of this race of: simulans var. 2, although it is prob-
able that, were there more specimens obtainable retaining a sufficient
amount of the color pattern, it might prove to be another variety.
Group oF A. A. VITTATA REEVE
ACHATINELLA APEXFULVA VITTATA Reeve
PLATE I, FIGURE 13; PLATE 4, FIGURES 17-20)
Achatinella vittata REEVE, Conch. Icon., vol. 6, pl. 2, fig. 9, April 1850—PI_sBry
and Cooke, Man. Conch., vol. 22, p. 280, pl. 57, figs. I, 2, 2a, 2c (only), 1914.
The holotype of A. a. vittata in the British Museum has a spot
of red sealing wax on the aperture side of the shell (pl. 4, fig. 17).
Postembryonic whorls white, penultimate banded with avellaneous,
last whorl above the periphery with two bands of avellaneous and
a subperipheral band of pinkish buff, below the periphery banded and
lined with fawn color, and a band of army brown in the umbilical
region; lip and columella callus pale vinaceous fawn. Length 19.0
mm., greater diameter I2.7 mm., spire height 10.0 mm., number of
whorls 6.
Distribution, area 12: Nuuanu, Gulick; Hillerbrand’s Glen,
Nuuanu, locality 100E-1*, elevation 1,000-1,500 feet, R. A. Cooke,
BBM 58284, and C. M. Cooke, BBM 221109, 19916 (figs. 3, 3a, p. 29).
Dr. C. M. Cooke tells me that these shells come from somewhere along
the ridge on which locality 100E-1* is plotted. They were probably
collected on the lower portion of the ridge at an elevation of around
1,000-1,500 feet, because Meinecke reports the larger race of A. a.
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 43
rubidipicta near the head of this subridge on the Nuuanu-Kapalama
Ridge. Further collecting will establish the exact location of this
Face. ‘
The specimens of A. a. vittata from Nuuanu collected by Gulick
closely match the holotype. The shells may be dextral or sinistral.
A. sinistral specimen (pl. 1, fig. 13), is very similar to the color
pattern of the type; postembryonic whorls white, lined and streaked
with cinnamon buff or pinkish buff and lined with mikado brown.
A darker color pattern (pl. 4, fig. 18) has the penultimate whorl
banded with pinkish buff shading to avellaneous and fawn color on
the last whorl; impressed sutural band and subsutural band pale
pinkish buff.
The R. A. Cooke and C. M. Cooke, Jr., lots are not typical vittata
but contain patterns similar to those of typical vittata. The locality
is a wide one and probably represents a mixture of typical vittata
with typical rubidipicta patterns. In the C. M. Cooke lot, which may
represent a collection of shells taken mostly from the lower portion
of locality 100E-1*, 36 percent of the shells have a pattern similar to
that of plate 4, figure 19, penultimate whorl light buff banded with
white, the last whorl banded with white below the subsutural band
of light buff, at and below the periphery banded and lined with
ochraceous tawny and tawny axially streaked with light buff, base
banded with white, and lined and banded with mars brown. A rare
brown pattern (pl. 4, fig. 19a) has the last whorl and a half colored
light buff below the suture, just above and below the periphery the
ground is axially streaked and banded and tinted with tawny and
russet.
In the R. A. Cooke lot color patterns are closer to A. a. rubidipicta
and probably come from a region adjacent to area 11. The typical
form and color pattern of the shell in the R. A. Cooke lots (pl. 4,
fig. 20) has the penultimate whorl white, lined with cinnamon buff,
impressed sutural band cinnamon, last whorl above the periphery
pale pinkish buff lined with pinkish buff and verona brown, edge of
the periphery banded with vandyke brown and a band of white, sub-
peripheral band vandyke brown, remainder of the base white, sutural
band pinkish cinnamon. Length 18.7 mm., greater diameter 13.0 mm.,
spire height 9.5 mm. The shell may have an entirely white ground
(pl. 4, fig. 20a), the last whorl profusely lined with walnut brown
and chocolate; impressed sutural and subsutural bands light buff.
A dark color pattern (pl. 4, fig. 20b) has the penultimate whorl
warm buff, banded and streaked with ochraceous tawny on the upper
44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
half of the whorl, including the impressed sutural band, lower half
mars brown, last whorl the impressed sutural and subsutural bands
ochraceous tawny, remainder of the whorl carob brown, lined with
white.
ACHATINELLA APEXFULVA VITTATA var. 1
PLATE 1, FIGURE 9; PLATE 4, FIGURES 15-15)
Achatinella vittata REEvE, Pilsbry and Cooke, Man. Conch., vol. 22, p. 290, pl. 57,
figs. 3-3b (only), 1914.
Area 8: Pauoa-Nuuanu Ridge, locality 91A*, elevation 1,500-
1,800 feet, Meinecke, 1923, 1925, 1926, 4 sinistral 1931. Also
locality g90A*, elevation 1,400-1,450 feet, Meinecke, 1922; goA-1*, ele-
vation 1,400 feet, Lemke, 1932; 93A*, elevation 1,500-1,800 feet,
7 sinistral 1928; 94A*, elevation 1,700 feet, 1927, 1928, 1931; 95A*,
elevation 1,750 feet, Meinecke, 1927, 1931 (figs. 3, 3a, p. 29). Other
collectors are J. S. Emerson, BBM 102189, E. D. Baldwin, BBM
56644-50646, Wilder, BBM 50498, Thurston shells collected by
Dos Santos, BBM 131084, A. F. Judd, BBM 110294-110299.
The usual form of the shell is darker than A. a. vittata, both in
the shade of yellow on the early postembryonic whorls and in the
reddish-brown banding on the last whorl. Large series of shells of
A. a. vittata and A. a. vittata var. 1 are separable, but individual
specimens from each race are indistinguishable. The usual form of
vittata var. I (pl. 1, fig. 9) has the first half postembryonic whorl
white axially streaked with cinnamon buff, penultimate whorl white
axially streaked with cinnamon buff, spirally banded with dark
vinaceous brown and white, last whorl banded and lined with white
and dark vinaceous drab, below the periphery lines of dark vinaceous
brown; impressed sutural band shades from tawny, on the first whorl,
to cinnamon on the last; subsutural band on the last whorl also cin-
namon; lip and columella callus vinaceous fawn. Length 18.8 mm.,
greater diameter 12.5 mm., spire height 9.5 mm., number of whorls 6.
An elongate specimen (pl. 4, fig. 15) measures: Length 18.6 mm.,
greater diameter 11.6 mm., spire height 10.5 mm.; first postembryonic
whorl and penultimate whorl, white faintly tinted and axially streaked
with pinkish buff and lined with cinnamon, last whorl above the
periphery white, lined and streaked with pinkish buff, and a band of
snuff brown above the periphery, peripheral band white with a line of
chocolate at the upper margin of the band, below the periphery banded
and lined with dark vinaceous ‘brown; impressed sutural band changes
from tawny to cream buff.
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 45
An obese shell (pl. 4, fig. 15a) shows the lightest color form of
this race; postembryonic whorls white spirally lined with pinkish
buff, below the periphery of the last whorl a faint line of verona
brown, below which are faint bands of pinkish buff, on the last
fourth whorl the bands are darker above and below the periphery,
being warm sepia and mikado brown. Length 17.3 mm., greater
diameter 12.7 mm., spire height 8.9 mm. The darkest color pattern
(pl. 4, fig. 150) has the postembryonic whorls light ochraceous buff,
banded and axially streaked with hay’s brown and sorghum brown,
base lined with white. The color patterns of figures 15a and 15) are
rare patterns occurring only on a few specimens in area 8.
ACHATINELLA APEXFULVA VITTATA var. 2
PLATE I, FIGURE 10; PLATE 4, FiGuRES 16, 16a
Achatinella vittata REEVE, Pilsbry and Cooke, Man. Conch., vol. 22, p. 290, pl. 57,
figs. 4, 4a, 5-5b (only), 1914.
Area g: Glen Ada, Nuuanu, locality 100-2*, approximate eleva-
tion 1,200-1,500 feet, R. A. Cooke, C. M. Cooke (figs. 3, 3a, p. 29).
While series of shells from area 9 are quite distinct from typical
A. a. vittata or A. a. vittata var. 1, because of the dominance of light
color patterns, nevertheless the color patterns of area g are repeated
in other areas and mixed lots are difficult to separate.
The usual form of the shell (pl. 1, fig. 10) has white postembryonic
whorls, banded on the last half of the penultimate with two bands of
fawn color, last whorl above the periphery banded with two bands
of cameo brown, lined below the suture with cinnamon buff, the
lines being broken with axial streaks of white, subperipheral band
cameo brown; sutural band cinnamon buff; lip and columella callus
vinaceous buff. Length 18.9 mm., greater diameter 12.3 mm., spire
height 9.9 mm., number of whorls 6.
The lightest color pattern (pl. 4, fig. 16). has white postembryonic
whorls lined on the last whorl with walnut brown. An obese shell and
a rare dark color pattern (pl. 4, fig. 16a), found on only 6 percent of
the shells in the R. A. Cooke lot, has the postembryonic whorls white,
last whorl above the periphery banded with pinkish buff axially
streaked with cinnamon, banded at and below the periphery with
chocolate ; impressed sutural band shades from tawny to ochraceous
tawny. Length 18.5 mm., greater diameter 12.8 mm., spire height
9.7 mm.
40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
ACHATINELLA APEXFULVA CINEREA Sykes
PLATE I, FIGURE II; PLATE 4, FIGURE 37; PLATE 5, FIGURES I, Ia
Achatinella vittata var. cinerea SyKES, Fauna Hawaiiensis, p. 305, 1900.
Achatinella vittata cinerea SyKeEs, Pilsbry and Cooke, Man. Conch., vol. 22,
pp. 280, 201, pl. 57, figs. 6-7, 19014.
The holotype (pl. 4, fig. 37) has the first postembryonic whorl
buckthorn brown, penultimate whorl banded with avellaneous, white,
and dark olive buff, last whorl ground white banded with maroon or
hay’s maroon; impressed sutural band and subsutural band white;
columella callus vinaceous fawn; lip vinaceous fawn banded with
white. Length 18.7 mm., greater diameter 12.4 mm., number of
whorls 64.
Distribution, area 10: Nuuanu, locality 100B-7?, elevation 1,250-
1,400 feet, Meinecke, 1911, 1914, 1916; 100B-12, elevation 1,150-
1,250 feet; 100B-13, elevation 1,150-1,350 feet; 100B-14, eleva-
tion 1,250-1,500 feet, 1 sinistral, all collected by C. M. Cooke, Jr.
(figs. 3, 3a, p. 29). The Meinecke locality may be wrongly plotted one
ridge south of where it actually is, because Dr. Cooke, who carefully
worked over this section of Nuuanu and made sketch maps of the
region, does not report any apexfulva from the region of locality
100B-7? Other collectors of A. a. cinerea are Wilder, BBM 50504,
R. A. Cooke, BBM 58138. Holotype in BM, collected in Nuuanu by
Perkins.
The usual color pattern and form of the shell (pl. 1, fig. 11) has the
first embryonic whorl warm sepia, next whorl upper fourth pale
pinkish buff, remainder of the whorl shades from prout’s brown to
buckthorn brown, last embryonic whorl upper fourth white, lower
three-fourths warm buff finely lined with white; first postembryonic
whorl and first half of penultimate banded with a light shade of
buckthorn brown and warm buff, last half of penultimate, and first
half of last whorl above the periphery, drab gray, banded below the
white sutural band, with a band of wood brown shading to chocolate,
a band of white, and 2 bands of chocolate, last half of the last whorl
chocolate, banded and lined with white, below the periphery last
whorl white, banded with chocolate, in the umbilical region a patch
of naples yellow, banded with chocolate. Length 19.1 mm., greater
diameter 13.1 mm., spire height 9.3 mm., number of whorls 6.
The color pattern may be light gray (pl. 5, fig. 1); first post-
embryonic whorl and first half of the penultimate white faintly
tinted with pinkish buff, last whorl and a half shades from pale
smoke gray to smoke gray on the last whorl, banded, lined, and finely
axially streaked with white—the last half whorl is also lined above
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 47
the periphery with fawn color—below the periphery banded with a
band of warm sepia lined with smoke gray, the aperture side of the
last whorl has a line of warm sepia above the white basal band. An
elongate specimen (pl. 5, fig. 1a), length 19.8 mm., greater diameter
12.8 mm., spire height 10.0 mm., shows the darkest color form, the
last whorl banded with white and chocolate.
ACHATINELLA APEXFULVA ALBOFASCIATA Smith
PLATE I, FiGURE 14; PLATE 4, FIGURES 27-28)
Apex albofasciatus SmitH, Proc. Zool. Soc. London, 1873, p. 78, pl. 9, fig. 21.
Achatinella vittata simulans Reeve, Pilsbry and Cooke, Man. Conch., vol. 22,
Pp. 202, 293, pl. 57, figs. 12, 13, 14a, 14c (only), 1914.
The holotype of A. a. albofasciata (pl. 4, fig. 27) in the British
Museum is marked with a red dot of sealing wax. The postembryonic
whorls are cinnamon buff darkening to sayal brown on the last half
whorl, banded on the last whorl at the edge of the periphery with
white and in the umbilical region with white and light pinkish
cinnamon; impressed sutural band white; lip and columella callus
pale pinkish buff. Length 18.5 mm., greater diameter 12.6, spire
height 9.0 mm., number of whorls 53.
Distribution, area 13°: Kapalama Stream, locality 110A-1?, “N.
Bank opposite Waiolani Peak. Halfway from top to bottom of Gulch,
elevation 1,500-1,550, Herb. Alani [Pelea],” L. A. Thurston rgrt1,
BBM 131103-131106 (figs. 3, 3a, p. 29). Also collected in Kapalama
by W. D. Wilder, BBM 50543, D. Fraser, BBM 15838, and Dos
Santos for Thurston, BBM 131095-131096.
The Thurston lots contain 37 dextrals and 4 sinistrals. The usual
form of a dextral (pl. 1, fig. 14) has the last half of the first post-
embryonic whorl, and the first half of the penultimate whorl, pinkish
buff axially streaked with cinnamon buff, last half of the penultimate
pale pinkish buff banded with light pinkish buff, shading on the last
whorl to pinkish buff finely lined and streaked with white or pale
pinkish buff, at the edge of the periphery a wide band of white; im-
pressed sutural band pinkish buff; lip light vinaceous fawn splotched
with white. Length 19.8 mm., greater diameter 12.3 mm., spire height
10.8 mm., number of whorls 6}. A sinistral specimen (pl. 4, fig. 28D)
is very like the holotype, differing in being banded with white above
the periphery and about the base, last two whorls with bands of
cinnamon buff streaked with cinnamon. The Kapalama form of J. a.
albofasciata is very close in form to A. a. rubidipicta. From Thurs-
ton’s description it occupies an area just below A. a. rubidipicta (area
11). A few specimens of A. a. albofasciata are often found in lots
4
48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
of A. a. rubidipicta, and probably come from the border region be-
tween areas II and 13.
The darkest color form of the shell is shown in plate 4, figure
28, the last two whorls are tawny axially streaked and lined with
ochraceous buff, on the last whorl a wide band of white at the periph-
ery. The lightest color pattern (pl. 4, fig. 28a) is entirely white
on the last two whorls lined with ochraceous tawny.
ACHATINELLA APEXFULVA ALBOFASCIATA var. 1
PLATE I, FIGURE 21; PLATE 4, FIGURES 29, 20a
Distribution, area 14??: Kalihi, J. S. Emerson collection, BBM
102205-102207, collected by Gomes and W. B. Olson. The exact posi-
tion of the locality is unknown, but has been plotted by me on the
Kamanaiki-Kalihi Ridge opposite area 13? (fig. 3a, p. 29), because
of the similarity of this form to A. a. albofasciata. This form should
be collected again and the exact locality determined.
The shells have a lighter color pattern than A. a. albofasciata. The
shade of the color pattern of the Kalihi shells best matches the holo-
type of A. a. albofasciata, but the banding of the Kapalama shells is
closer to that of the holotype of albofasciata and for this reason
Kapalama is considered the region of the type locality. The Kalihi
shells are dominantly sinistral and composed of light color patterns.
Forms similar to plate 4, figure 28, are not found in this Kalihi lot.
Out of 34 shells, 9 are dextrals.
The typical form of the shell (pl. 1, fig. 21) is smaller than the
usual form of albofasciata from Kapalama, the postembryonic
whorls cinnamon buff faintly lined and streaked with white ; impressed
sutural band the color of the ground; last whorl below the edge of the
periphery white, lined with cinnamon buff; lip and columella callus
pale cinnamon pink. Length 18.1 mm., greater diameter 12.3 mm.,
spire height 9.7 mm., number of whorls 6.
The lightest color pattern (pl. 4, fig. 29) has the postembryonic
whorls white, lined with cinnamon buff. The dextral form of the
shell (pl. 4, fig. 29a) has the last two whorls light buff, axially streaked
and lined with ochraceous buff, last whorl at the periphery and below
the periphery banded with white. This form looks very much like
yellow forms of A. a. globosa (pl. 4, fig. 24).
ACHATINELLA APEXFULVA OLIVERI, new subspecies
PLATE I, FIGURE 18; PLATE 4, FIGURES 30-31
This form is intermediate between the more globose upper race of
A. a. albofasciata and the smaller form of A. a. simulans var. 1.
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 49
The first postembryonic whorl of the holotype (pl. 1, fig. 18) is pale
pinkish cinnamon spirally lined, banded, and tinted with sayal brown,
last two whorls sayal brown, last whorl below the periphery banded
and lined with white ; impressed sutural band white; lip and columella
callus light vinaceous fawn. Length 19.7 mm., greater diameter
12.5 mm., spire height 11.0 mm., number of whorls 63.
Distribution, area 15?: Kamanaiki Stream, type locality 120-1?,
O. H. Emerson, BBM 103962; also collected by J. S. Emerson, BBM
102203-102204, Dos Santos, BBM 74301, and Thwing, BBM 28544-
28546 (figs. 3, 3a, p. 29). The locality has not been found in recent
years. Only an approximation has been made by Dr. C. M. Cooke,
and Dr. Oliver Emerson which is somewhere in the vicinity of the lo-
cality. The shell is named after Dr. Oliver Emerson, who dis-
covered it.
A dark white-lined color pattern (pl. 4, fig. 31) has the last two
whorls banded with russet and lined with white. The darkest color
pattern (pl. 4, fig. 30) has the last two whorls liver brown axially
streaked with hazel, about the periphery of the last whorl a line of
white; impressed sutural band white. The lightest color pattern
(pl. 4, fig. 30a) is white, lined on the last two whorls with hazel.
This pattern occurs on only one shell in the type lot.
ACHATINELLA APEXFULVA RUBIDIPICTA, new subspecies
PLATE I, FIGURE 15; PLATE 5, FiGuRES 2-2d
Achatinella vittata simulans Reeve, Pilsbry and Cooke, Man. Conch., vol. 22,
p. 292, pl. 57, figs. 9-11, 14b (only), 1914 (not of Reeve 1850).
The shell is larger than A. a. simulans, having a characteristic
smooth surface, and usually having spiral bands which are generally
not broken by axial streaks. The form is closest to that of A. a.
cinerea, having an inflated last whorl. The holotype (pl. 1, fig. 15)
has the first half of the first embryonic whorl rood’s brown, last half
upper portion white, the lower half rood’s brown changing to mouse
gray on the next whorl, on the last half of the second embryonic
whorl and on the remaining embryonic whorls the lower band is a
faint shade of cream buff, upper band white; the postembryonic
whorls white, the first half of the first postembryonic whorl banded
on the lower fourth of the whorl with cinnamon buff, penultimate and
last whorls lined with cinnamon, last half of penultimate and last
whorl banded at the edge of the periphery with burnt umber, last
whorl below the periphery with a line of burnt umber; lip and
50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
columella callus pale vinaceous fawn. Length 19.5 mm., greater
diameter 13.2 mm., spire height 10.2 mm., number of whorls 63.
Distribution, area 11: Nuuanu-Kapalama Ridge, type locality
110*, elevation 1,750-1,870 feet, Meinecke, 1921, 1922, 1927, 1929;
also locality 111*, elevation 1,800-2,100 feet, Meinecke, I sinistral
1929 (figs. 3, 34, p. 29).
The color pattern of Ir percent of the shells (pl. 5, fig. 2) is a
more heavily lined one; the last two whorls may be white, lined and
banded on the penultimate with mikado brown, on the last whorl
banded with vandyke brown and lined with mikado brown, base
banded with seal brown, umbilical region tinted with cream buff.
Twenty-four percent are conspicuously banded ; last two whorls white,
(pl. 5, fig. 2a) banded with chocolate and lined on the penultimate
with a subsutural band of mikado brown, which splits into two lines
on the last half of the penultimate and fades out on the last half of
the last whorl. The darkest color pattern and elongate form of the
shell (pl. 5, fig. 2b), occurring on 2 percent of the shells, has the last
two whorls burnt umber axially streaked with cinnamon and banded
on the last whorl with white ; impressed sutural band chocolate shading
to cinnamon buff at the upper edge. Length 19.8 mm., greater di-
ameter 12.8 mm., spire height 9.9 mm. This shell resembles the dark
color pattern of A. a. cinerea (pl. 5, fig. 1a). The main difference is
that A. a. cinerea has the embryonic whorls banded with yellow and
the first two postembryonic whorls banded with yellow and gray.
The obese form and lightest color pattern of the shell (pl. 5, fig. 2c)
measures: Length 19.0 mm., greater diameter 13.8 mm., spire height
9.4 mm.; postembryonic whorls white, below the impressed sutural
band a line of cinnamon deepening to warm sepia on the last whorl,
the base lined and banded with chocolate. This white color pattern
occurs on 20 percent of the shells. Eight percent have a yellow color
pattern which looks like an intermediate form between rubidipicta
and albofasciata. One of these (pl. 5, fig. 2d) has the last whorl and
a half pale pinkish buff axially streaked and lined with cinnamon,
banded with white at the edge of the periphery, below the periphery
banded with amber brown, and a basal band of pinkish buff ; impressed
sutural band pinkish buff axially streaked with cinnamon. In some
collections specimens of A. a. albofasciata are found mixed with lots
of A. a. rubidipicta. The reason for this is that the shells were either
collected in a locality intermediate between areas 11 and 13 or all
the shells from areas 11 and 13 were lumped together in one locality.
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 51
ACHATINELLA APEXFULVA RUBIDIPICTA var. 1
PLATE 5, FIGURES 3-3)
Area?: Kalihi, Wilder, BBM 50544, 9 dextral, 16 sinistral.
The shells are similar to A. a. rubidipicta but have a different series
of color patterns. Some dextral specimens have a color pattern which
resembles A. a. albofasciata and differs only in having light reddish-
brown bands instead of yellowish-brown ones. One of these dis-
tinctive color patterns (pl. 5, fig. 3) is a white shell banded and
lined on the last two whorls with pecan brown, and a band of
ochraceous tawny below the periphery. The usual form of a dextral
shell (pl. 5, fig. 3a) measures: Length 19.7 mm., greater diameter
13.0 mm., spire height 10.0 mm.; last two postembryonic whorls
white, with a band of russet below the white sutural band and just
above the edge of the periphery, last whorl lined at the edge of the
periphery with two lines of burnt umber, below the periphery lined
and banded with burnt umber on the rest of the whorl. The sinistral
form of the shell (pl. 5, fig. 3b) is very like typical 4. a. rubidipicta
but the distribution of the chocolate bands over the white ground on
the last two whorls is different from anything found in locality 110.
ACHATINELLA APEXFULVA RUBIDIPICTA var. 2
PLATE I, FIGURE 22
Area 18: Kalihi-Kahauiki Ridge, locality 141*, elevation 1,650
feet, Meinecke, 2 sinistrals 1927 (figs. 3, 3a, p. 29). Also collected in
Kahauiki by Wilder, BBM 50519. The usual color pattern of the
shells in the Wilder lot is similar to that of the two Meinecke shells,
and they are believed to have come from the same locality.
The form is very close to 4. a. rubidipicta but has a slightly differ-
ent series of color patterns. The usual color pattern (pl. 1, fig. 22)
has the last two postembryonic whorls white, lined below the suture
with a line of mikado brown, last whorl banded with two chocolate
bands. Length 19.8 mm., greater diameter 13.4 mm., spire height
10.2 mm. The color pattern may be lined similar to plate 5, figure 30.
ACHATINELLA APEXFULVA RUBIDIPICTA var. 3
PLATE 5, FIGURES 4-4e
Area?: Moanalua, Wilder, BBM 50522.
The subspecies resembles A. a. rubidipicta but has a different
series of color patterns. The embryonic whorls are similar to those
of A. a. simulans var. 2, and are intermediate in form and color
52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
pattern between A. a. ovwm and A. a. rubidipicta. The shells prob-
ably came from above area 19 on the Manaiki-Moanalua Ridge.
The usual form of the shell and the color pattern of 52 percent
of the specimens (pl. 5, fig. 4) has the postembryonic whorls white,
lined with cinnamon and mikado brown, and banded with chocolate,
last whorl subperipheral band white, below which is a line of cinnamon
and cinnamon buff, base chocolate ; impressed sutural band cinnamon ;
lip and columella callus light vinaceous fawn. Length 19.6 mm.,
greater diameter 13.0 mm., spire height 9.8 mm., number of
whorls 63.
An elongate shell (pl. 5, fig. 4a) measures: Length 19.3 mm.,
greater diameter 12.5 mm., spire height 9.5 mm.; last two postem-
bryonic whorls pale pinkish buff, profusely lined and banded with
russet and mars brown, impressed sutural band cinnamon. An obese
form of the shell (pl. 5, fig. 40) with light color pattern found on
31 percent of the shells has the postembryonic whorls white, lined with
pinkish buff; impressed sutural band pinkish buff; last whorl banded
at the edge of the periphery with a band and below the periphery
with bands and lines of chocolate. Length 19.1 mm., greater diameter
13.4 mm., spire height 9.1 mm.
Seven percent have a lined spiral pattern (pl. 5, fig. 4c); post-
embryonic whorls above the periphery lined with russet, peripheral
band narrow, about 0.5 mm. instead of nearly 2 mm. as in figure 4,
banded below and at the periphery with chocolate. The darkest
pattern (pl. 5, fig. 4d) on 9 percent, is russet, streaked with cinnamon
buff on the first half of the penultimate, last half of penultimate ochra-
ceous tawny axially streaked with burnt umber, last whorl chocolate,
axially streaked with ochraceous tawny. A single specimen (pl. 5,
fig. 4¢) has a white ground lined with warm buff; impressed sutural
band ochraceous tawny fading to white on the last whorl. This
pattern is very similar to lined patterns of A. a. albofasciata.
Group oF A. A. PILSBRYI NEW SUBSPECIES
ACHATINELLA APEXFULVA PILSBRYI, new subspecies
PLATE I, FIGURE 29; PLATE 5, FIGURE 27
The holotype (pl. 1, fig. 29) is small, embryonic whorls white;
spire pointed, postembryonic whorls white; impressed sutural band
tawny ; last whorl below and at the periphery banded with tawny and
a basal band of kaiser brown; lip fawn color, tuberculate; columella
callus white. Length 17.6 mm., greater diameter 12.4 mm., spire
height 9.2 mm., number of whorls 63.
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 53
S>>
Kahaluu
239.
239b~ 2
32320 239a.k
4.2310 d—239B °2o,, f- 116-2
N. Waimano
2386
2300 ends : : a
Central
ST nls 22Ba~ g_o gbtlZA 162D-O
Kipapa 228 Qo. 222 szie8g
Gulch ae Pat o 5-176
3
O- 114
12 FA
Q-110 6-8
Woe “J
cc 150 B-2/
a Stream
Moanalua
Stream
Manaikj Stream
South Halaw
bay
&
®
5s
en
a
tral Halawa Stream,
Stream\ UO
Fic. 4—Southeast-central Oahu, leeward slope of the Koolau Range, including
regions Ia, II, IIIa, half of region III, and two valleys of region I (fig. 7, p. 194),
showing localities of subspecies of A. apexfulva belonging to the groups of A. a. cestus,
A. a. vittata, A. a. pilsbryi, A. a. turgida, A. a. polymorpha, and A. a. irwint.
iL Waikane sh = Kaneohe
iY
rn
w
$
Kahauiki Stream
Ss
v
mM
3
wn
s
=
&
oo
mlawa Streae ——
SS
Manaiki Stream
OF
y
Moanalua Stream
Central H
Zz
x
o
.
North
Halawa
Stream
Aiea
Stream
South H.
Panihakea
Fic. 4a.—Same as fig. 4, but showing the area occupied by different subspecies of
A. a. apexfulva belonging to the groups of A. a. cestus, A. a. vittata, A. a. pilsbryi,
A. a. turgida, A. a. polymorpha, and A. a. irwini.
54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Distribution probably somewhere in the region of area 26?: Aiea-
Kalauao Ridge, approximate locality 170-1?, Judd, BBM 110210,
and Cooke, BBM 72494 (figs. 4, 4a, p. 53). The subspecies is named
after Dr. Henry A. Pilsbry. The shell has not been found in many
years and may be extinct.
The form of a broadly banded juvenile shell (pl. 5, fig. 27) has
an elongate spire. Length 16.9 mm., greater diameter 11.6 mm.,
spire height 9.0 mm. Postembryonic whorls white, banded with
cinnamon, last whorl below the periphery cinnamon, lined with two
lines of white and a line of mikado brown. The shell is a dead speci-
men and the color has faded.
ACHATINELLA APEXFULVA ROSEATA, new subspecies
PLATE I, FIGURE 39; PLATE 6, FIGURES 25-25)
The shell is a small fossil form which looks like A. a. lilacea from
the region of North Kaukonahua Stream. The holotype (pl. 1, fig. 39)
has white, pointed, embryonic whorls; postembryonic whorls pale
vinaceous fawn axially streaked and faintly spirally lined with light
vinaceous fawn and vinaceous fawn; impressed sutural band white
or tinted pale pinkish buff; lip outer margin pinkish buff, inner
margin pale pinkish buff. Length 17.5 mm., greater diameter
11.5 mm., spire height 9.6 mm., number of whorls 63.
Distribution, area 37?: Waimano Stream. “Just below the bridge
on the plantation R. R.,’ C. M. Cooke, BBM 40679-40680, also col-
lected by Thaanum, BBM 40716-40717, and Wilder, BBM 51932.
The shells were found in fossil state under a boulder in the bottom
of the gulch. | have tentatively plotted the locality somewhere in the
region of area 37? (fig. 4a, p. 53).
The holotype is a little smaller than usual. The usual form should
be 18+mm., with a greater diameter of I11-++mm., but no specimens
with the usual form have the color sufficiently well preserved to be
used for the holotype. The darkest color pattern (pl. 6, fig. 25) ona
single specimen has the postembryonic whorls colored light vinaceous
fawn axially streaked and lined with vinaceous fawn; impressed
sutural band vinaceous buff; lip vinaceous buff. Length 19.1 mm.,
greater diameter 11.9 mm., spire height 11.2 mm.
The narrowest specimen (pl. 6, fig. 25a) measures: Length
18.1 mm., greater diameter 10.8 mm., spire height 11.0 mm. The
typical sinistral form of the shell (pl. 6, fig. 25b) measures: Length
18.0 mm., greater diameter 11.7 mm., spire height 10.4 mm.
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 55
ACHATINELLA APEXFULVA LAURANI, new subspecies
PLATE I, FiGuRES 37, 38; PLATE 5, Ficures 28-31; PLATE 6, FIGURE I
The embryonic whorls of the holotype (pl. 1, fig. 38) are white;
first postembryonic and penultimate whorls, avellaneous, finely axially
streaked with faint lines of bone brown, last whorl avellaneous axially
streaked with olive brown and natal brown, spirally lined below the
periphery with olive brown and cream buff, and banded with buffy
brown, last eighth whorl bone brown; impressed sutural band natal
brown; outer margin of the lip natal brown, inner margin fawn
color. Length 18.2 mm., greater diameter 12.4 mm., spire height
9.7 mm., number of whorls 6.
Distribution, area 27: Hanaiki-Waimalu Ridge, type locality
Ig1*, elevation I,200-1,250 feet, 1932; also 192, elevation 1,200-1,250
feet, Meinecke, 7 sinistral 1933 (figs. 4, 4a, p. 53). Also collected by
Thurston, BBM 130604, 9 sinistral, Thwing, BBM 28751-28759, 26
sinistral.
At a lower elevation on the same ridge, possibly in the region of
area 27A?, Thurston, 1910, BBM 130603, discovered a colony of
dextral forms living on maile (Alyxia), and lehua (Metrosidros).
Also collected by Cooke, BBM 72557-72558, 7 dextral, A. F. Judd,
BBM 110492-110494 and 110496, Thwing, BBM 28986, 1 dextral.
The lightest color pattern (pl. 5, fig. 31), on 8 percent of the
shells in the combined Meinecke-Thurston lots from area 27, has
the postembryonic whorls white; impressed sutural and subsutural
bands wood brown shading to bone brown on the last two whorls;
last whorl banded and lined below and at the edge of the periphery
with bister and snuff brown, about the umbilicus a band of bone
brown; lip and columella callus natal brown.
The obese form of the shell (pl. 6, fig. 1) has a lined pattern
which occurs on 13 percent of the shells; the first postembryonic
whorl fawn color banded with white, penultimate army brown axially
streaked with olive brown, and lined with white, last whorl above
the periphery banded with a band of olive brown, and one of white,
below the periphery a band about 5 mm. wide of cinnamon buff
lined and banded with olive brown, base banded with pale pinkish
buff, and a band of bister in the umbilical region. Length 18.8 mm.,
greater diameter 13.2 mm., spire height 9.5 mm.
A dextral shell of the usual form and color pattern from area
27A? (pl. 1, fig. 37) measures: Length 19.0 mm., greater diameter
12.4 mm., spire height 9.9 mm., number of whorls 64. The first
postembryonic whorl and a half shades from fawn color to army
56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
brown, axially streaked and finely lined with pale smoke gray, last
whorl and a half bay profusely lined and streaked with tilleul buff,
basal band warm sepia; outer margin of lip wood brown, inner margin
vinaceous buff, columella callus white.
A narrow specimen (pl. 5, fig. 28) measures: Length 18.7 mm.,
greater diameter 11.3 mm., spire height 11.0 mm.; postembryonic
whorls avellaneous, faintly axially streaked and spirally lined with
army brown, last whorl below and at the periphery lined and banded
with bister and snuff brown. An obese shell (pl. 5, fig. 29) shows
a white color pattern occurring on 16 percent of the shells in the com-
bined Cooke, Judd, and Thurston lots; postembryonic whorls white ;
impressed sutural band white, edged wth sayal brown; last whorl be-
low the periphery banded with light cinnamon drab, basal band
fuscous. A white banded color pattern (pl. 5, fig. 30), has the postem-
bryonic whorls white, banded with sayal brown, last whorl at and
below the periphery banded with snuff brown.
In the Thurston lot (pl. 5 fig. 30a) a shell with the usual obese
form has a pattern which resembles the form and type of banding
of A. a. pilsbryi, but the color of the bands are entirely different on
the two forms. The postembryonic whorls are white, last whorl
banded below the periphery with chocolate; impressed sutural band
chocolate. Length 18.4 mm., greater diameter 12.7 mm., spire height
9.2 mm. The presence of this chocolate-banded form in the
Thurston lot is of interest, because it may be a representative of a
still lower race, which is a chocolate-banded one and closely related
to A. a. pilsbryi.
ACHATINELLA APEXFULVA PARVICOLOR, new subspecies
PLaTe I, FIGURE 40; PLATE 6, FIGURES I5-I5c
Achatinella turgida ovum PFEIFFER, Pilsbry and Cooke, Man. Conch., vol. 22,
pp. 295, 208, pl. 56, figs. 3, 3@ (only), 1914.
This form is smaller than A. a. turgida and has a light color pattern
and never has the dark diamine brown or yellow color patterns of
A. a. turgida. The subspecies is not found today, being a race collected
by Gulick and identified by him as turgida. The embryonic whorls of
the holotype (pl. 1, fig. 40) are white; postembryonic whorls pale
pinkish buff, axially streaked with sayal brown, peripheral band on
the last whorl pale pinkish buff; edge of the lip fawn color. Length
19.2 mm., greater diameter 12.7 mm., spire height 10.3 mm., number
of whorls 64.
Distribution, area 36??: Waimalu?, Gulick. I have tentatively
placed the probable locality somewhere in Waiau and Punanani
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 57
Gulch, because this region is often referred to as Waimalu by
some collectors. The race may have come from somewhere near
area 35 because the shell looks so like forms of A. a. turgida (fig. 4a,
Pp. 53):
The lightest color pattern and extreme obese form (pl. 6, fig. 15)
has the postembryonic whorls pale pinkish cinnamon, spotted on the
first two postembryonic whorls and banded just below the impressed
sutural band on the last whorl with cinnamon buff, below the periph-
ery of the last whorl faintly lined with dots of cinnamon buff.
Length 17.2 mm., greater diameter 13.0 mm., spire height 9.0 mm.
An elongate dextral (pl. 6, fig. 15a) measures: Length 18.8 mm.,
greater diameter 11.5 mm., spire height 10.7 mm.; embryonic whorls
worn; postembryonic whorls pale pinkish cinnamon axially streaked
with sayal brown, banded on the last half whorl below the periphery
with a band of verona brown.
Plate 6, figure 15), shows an obese sinistral specimen with a color
pattern similar to plate 1, figure 4o. An elongate sinistral (pl. 6,
fig. 15c) with a color pattern similar to the holotype, measures:
Length 19.1 mm., greater diameter 11.8 mm., spire height 11.0 mm.
Group or A. A. TURGIDA NEWCoMB
ACHATINELLA APEXFULVA OVUM PFEIFFER
PLATE I, Figure 26; PLATE 5, FIGURES 9-II
Achatinella ovwm PretrFer, Proc. Zool. Soc. London, 1856, p. 334.
Achatinella ee ovum PFetrFrER, Pilsbry and Cooke, Man. Conch., vol. 22,
p. 297, pl. 59, fig. 17 (only), 1914.
The holotype (pl. 5, fig. 9) in the British Museum, marked with
a red dot of sealing wax, is entirely white except for the black lip.
Length 19.4 mm., spire height 10.3 mm., number of whorls 64. The
other two shells in the type lot are white dextral specimens. One is not
adult, the other (pl. 5, fig. 10) is fully developed and marked specimen
A; the lip is black. Length 17.8 mms, greater diameter 12.7 mm.,
Bain of whorls 61.
Distribution, area 22: North-Central Halawa Ridge, locality
162C*, elevation 1,400-1,450 feet, Meinecke, 1930 (figs. 4, 4a, p. 53).
Also collected by Wilder, BBM 50525.
The holotype probably came from area 22 because that area is
the lowest containing sinistral shells. The dextrals in the type lot
may have come from a lower locality somewhere near area 21. Only
six live shells are known to come from area 22. The colors are
equally divided between the typical white ovum pattern of plate 1,
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
ut
figure 20, and the lined pattern of plate 5, figure 11. The usual pat-
tern (pl. 1, fig. 26) and form of the shell from area 22 has the post-
embryonic whorls white; about 3 mm. behind the edge of the lip
the last whorl is ornamented with a streak of sayal brown and banded
with a band of brownish black; lip brownish black shading to cinna-
mon drab on the inner edge, near the columella callus the outer edge
is also cinnamon drab. Length 20.0 mm., greater diameter 13.7 mm.,
spire height 10.0 mm., number of whorls 63. The lined pattern (pl. 5,
fig. 11) has the embryonic whorls white shading to pale pinkish buff
on the first half of the first postembryonic whorl, last half pinkish
buff, last two whorls pale pinkish buff banded and lined with pinkish
buff and cinnamon buff, the last 2} mm. behind the edge of the lip
bone brown or a blackish brown; lip a blackish brown; tip of colu-
mella callus white, shading to vinaceous fawn, near the closed um-
bilicus. So few of the specimens were available for study that the
typical form was not determined.
ACHATINELLA APEXFULVA OVUM var. 1
PLATE I, FIGURE 25; PLATE 5, FicureEs 8, 8a
Achatinella turgida ovum PFEIFFER, Pilsbry and Cooke, Man. Conch., vol. 22,
p. 208, pl. 56, fig. 10 (only), 1914.
Area 21: North-Central Halawa Ridge, locality 160C*, elevation
1,000-1,100 feet, Meinecke, 1 dextral 1930; Central Halawa region
of 162B*, elevation 1,050-1,250 feet, Meinecke, 1914; also collected
by Thurston, BBM 130505 (figs. 4, 4a, p. 53).
The shells of area 21 differ from those of area 22 in being dextral,
smaller, and having an impressed sutural band of sayal brown on the
first postembryonic whorl, on the remaining whorls the sutural band
is white, fringed with sayal brown at the upper edge, or entirely
white.
The usual form of the shell (pl. 1, fig. 25) measures: Length
18.7 mm., greater diameter 12.5 mm., spire height 9.8 mm., number of
whorls 63. The shell is a dead specimen but retains most of the shell
nacre; postembryonic whorls white slightly tinted with cream color
just behind the edge of the lip; lip army brown.
In a fresh specimen (pl. 5, fig. 8) embryonic whorls a light shade
of cream color; first postembryonic whorl cream color shading to
white on the penultimate and first half of the last whorl, last half
whorl above the periphery ivory yellow, last whorl below the periph-
ery cream color lined and banded with avellaneous. Length 17.3 mm.,
greater diameter 12.4 mm., spire height 9.2 mm.
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 59
A narrow shell (pl. 5, fig. 8a) measures: Length 18.2 mm., greater
diameter 11.8 mm., spire height 10.2 mm.; the shell is white except
for the first postembryonic whorl and a half which is lined with
ochraceous fawn and has the impressed sutural band ochraceous
tawny ; lip army brown.
ACHATINELLA APEXFULVA OVUM var. 2
PLATE I, FIGURES 27, 28, 31; PLATE 5, FIGURES 12-14
Area 23: North-South Halawa Ridge, locality 162D*, elevation
1,500-1,750 feet; 164D*, elevation 1,500-1,650 feet, 5 sinistral;
165D*, elevation 1,400-1,500 feet, 4 sinistral, all Meinecke-collected,
1930. Also collected probably in the region of areas 22 and 23 by
J. S. Emerson, BBM 102211, by Dos Santos, BBM 74314, Thurston,
BBM 130506, Wilder, BBM 50523 (figs. 4, 4a, p. 53).
Area 24: Halawa-Kalauao Ridge, locality 174*, elevation 1,550
feet, 1930; 176-1, elevation 1,650 feet, 2 sinistral 1936; 176-2, eleva-
tion 1,700-1,750 feet, I sinistral 1936; 176*, elevation 1,650-
1,700 feet, I sinistral 1930 (figs. 4, 4a, p. 53) all Meinecke-collected.
Aiea, J. S. Emerson, BBM 102446, 6 sinistral.
The shells of areas 23 and 24 are characterized by having certain
yellow color patterns mixed with the usual A. a. ovum pattern of
area 22.
The usual form of the shell in area 23 (pl. 5, fig. 12) measures:
Length 19.4 mm., greater diameter 13.0 mm., spire height 9.5 mm.;
and has a color pattern similar to that of A. a. ovum (pl. I, fig. 26).
A distinctive yellow color pattern (pl. 1, fig. 27) has the embryonic
whorls white; first postembryonic whorl white, impressed sutural
band ochraceous tawny; first half of the penultimate whorl white, im-
pressed sutural band light pinkish cinnamon, last half of penultimate
and last whorl naples yellow, banded at the edge of the periphery with
white, impressed sutural band color of ground; lip seal brown or
bone brown; columella callus fawn color. Length 19.5 mm., greater
diameter 12.8 mm., spire height 10.5 mm.
In the Wilder lot (BBM 50523) of 30 shells, 50 percent are white
and resemble A. a. ovum (pl. I, fig. 26). The usual form of the shell
(pl. 5, fig. 13) measures: Length 20.1 mm., greater diameter 13.7
mm., spire height 10.0 mm. Four shells in the lot have a yellow pattern
(pl. 1, fig. 27), seven a yellow banded or lined pattern (pl. 5, fig. 11).
One of the yellow-banded shells, an elongate form, has unusually
dark bands (pl. 1, fig. 28). The postembryonic whorls are ochraceous
tawny, finely or faintly lined with white, last whorl below the
60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
periphery banded with a band of white, a line of ochraceous tawny,
base pale pinkish buff, darkening to warm sepia behind the edge
of the lip. Length 20.7 mm., greater diameter 13.1 mm., spire
_ height 11.1 mm.
In area 24 the usual color pattern and form (pl. I, fig. 31) isa
white shell with pale pinkish buff embryonic whorls; lip and columella
callus light seal brown. Length 19.2 mm., greater diameter 12.7 mm.,
spire height 10.0 mm. The shell is a little narrower than the usual
form. The yellow-lined pattern of this area (pl. 5, fig. 14) has the
postembryonic whorls white with a band of cream buff above the
periphery, last whorl banded at the edge of the periphery with a band
of white, below the periphery banded with chamois shading to white
in the umbilical region, and lined with two lines of cinnamon.
ACHATINELLA APEXFULVA OVUM var. 3
PLATE I, FIGURE 24; PLATE 5, FicurEs 6, 7
Area 20: Moanalua-Halawa Ridge, locality 163*, elevation
1,500-1,620 feet, Meinecke, 9 sinistral 1930. Also collected some-
where on the same ridge by O. H. Emerson, BBM 103964, I915 on
maile (Alyxia) and hoe (hoi?, Dioscorea); J. S. Emerson, BBM
102209, 1915 (figs. 4, 4a, p. 53). |
The usual color pattern of the shells in area 20 is similar to that of
A. a. ovum (area 22), but other patterns occur which do not resemble
those found in area 22. The embryonic whorls are very slightly bi-
colored but are closer to A. a. ovum than to A. a. simulans.
The usual form in the Meinecke lot from locality 163* (pl. 1, fig. 24)
measures: Length 20.2 mm., greater diameter 13.2 mm., spire height
10.5 mm.; the embryonic whorls are not usual, being more pointed,
first embryonic whorl light vinaceous fawn, remaining embryonic
whorls white shading to pale pinkish buff; first postembryonic whorl
white, lined with pinkish buff, remaining whorls white, 3 mm. behind
the edge of the lip a streak of verona brown, banded with warm
sepia; impressed sutural band pinkish buff shading to white on the
last whorl. A peculiar pattern (pl. 5, fig. 6) has the embryonic whorls
white, first embryonic whorl slightly darker than the rest ; postembry-
onic whorls above the periphery ochraceous buff deepening to cinna-
mon on last half whorl, and lined on the last two whorls with white;
impressed sutural band cinnamon, at and below the periphery the
base is white, with a central line of cinnamon.
A third color pattern (pl. 5, fig. 7) shown on the usual form of the
shell in the O. H. Emerson and J. S. Emerson lots has the postem-
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 61
bryonic whorls white, first postembryonic whorl banded with ochra-
ceous tawny, penultimate whorl with a sutural band and a band just
above the edge of the periphery of russet, last whorl lined at the
edge of the periphery with buckthorn brown, subperipheral band
russet, base banded with tawny; lip army brown. Length 19.5 mm.,
greater diameter 13.7 mm., spire height 10.0 mm.
ACHATINELLA APEXFULVA BRUNEOLA, new subspecies
PLATE I, FIGURE 32; PLATE 5, FIGURES 17-17)
Achatinella turgida ovum Preirrer, Pilsbry and Cooke, Man. Conch., vol. 22,
p. 208, pl. 56, fig. 8a (only), 1914.
The shell resembles A. a. ovum in form, but differs in having a
brown color pattern. Embryonic whorls of the holotype (pl. 1, fig. 32)
white ; first half postembryonic whorl shades from white to ochraceous
tawny, penultimate and last whorl sudan brown, at the periphery and
below the periphery of the last whorl the ground is argus brown,
lined and banded with apricot yellow; lip verona brown; columella
callus tilluel buff. Length 19.0 mm., greater diameter 13.0 mm., spire
height 9.7 mm., number of whorls 64.
Distribution, area 28: Kalauao-Hanaiki Ridge, type locality 182,
elevation 1,300-1,367 feet, Meinecke, 1926, 3 dextral 1929, 2 dextral
1933 (figs. 4, 4a, p. 53).
The holotype has the usual form of the shell, and a color pattern
which occurs on 33 percent of the shells. An elongate shell (pl. 5,
fig. 17) has the usual color pattern on 58 percent of the shells; post-
embryonic whorls ochraceous tawny spirally banded with russet,
below the periphery of the last whorl prout’s brown with a few lines
of yellow ochre; lip and columella callus mars brown. Length 19.3
mm., greater diameter 12.2 mm., spire height 10.7 mm. A squat
specimen (pl. 5, fig. 17a), with a similar color pattern to figure 17,
has a very concave spire and inflated last whorl; the postembryonic
whorls are profusely lined with russet. Length 19.0 mm., greater
diameter 13.4 mm., spire height 8.9 mm. The lightest color pattern
found on a single specimen (pl. 5, fig. 17>) has the first two post-
embryonic whorls pale ochraceous buff and lined with wood brown
darkening to fawn color, last whorl above the periphery pale ochra-
ceous buff lined with natal brown and buffy brown, at and below the
periphery the ground is chamois above and within the aperture, the
remainder of the whorl is cartridge buff; lip and columella callus
army brown streaked with white.
62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
ACHATINELLA APEXFULVA BRUNEOLA var. 1
PLATE I, FiGuRE 33; PLATE 5, Ficure 18
Area 29: Kalauao-Hanaiki Ridge, locality 183, elevation 1,300-
1,350 feet, Meinecke, 13 sinistral 1935; 183?, Meinecke, 1913 (figs. 4,
4a, p. 53).
The usual form of the shell is larger than typical A. a. bruneola,
the mean length 20+mm. instead of 19+mm. The color pattern
differs from typical bruneola by having the base banded with yellow ;
the shells are usually sinistral. The form is an intermediate one be-
tween the higher race of A. a. aureola and A. a. bruneola.
The usual form and color pattern of the shell (pl. 1, fig. 33) has
the embryonic whorls white or pale pinkish buff, last two postembry-
onic whorls amber brown, peripheral band and base light cadmium,
subperipheral band chestnut, below which are lines of amber brown
and a band about the umbilicus of a deep shade of mars brown,
behind the edge of the lip last whorl tinted with a streak of chestnut.
Length 20.4 mm., greater diameter 13.1 mm., spire height 10.7 mm.,
number of whorls 63. The color of a dextral shell (pl. 5, fig. 18)
shows the light color pattern of the shell; postembryonic whorls mars
yellow or a light shade of sudan brown, base cadmium with a sub-
peripheral band of chestnut.
ACHATINELLA APEXFULVA BRUNEOLA var. 2
PLATE I, FIGURE 30; PLATE 5, FIGURES 15-16
Achatinella turgida ovum PFEIFFER, Pilsbry and Cooke, Man. Conch., vol. 22,
p. 208, pl. 56, figs. 7-7d (not from Moanalua but from Aiea, according to
Dr. Cooke) and figure 6 (only), 1914 (figure 6 also matches color patterns
of A. a. ovum var. 2).
Area 25: Aiea-Kalauao Ridge, locality 170C-6*, elevation 1,550-
1,800 feet, Lemke, 1924 and 1932 (figs. 4, 4a, p. 53); also collected
in Aiea by Cooke, BBM 72501-72512, BBM 72514, BBM 16346-
16347, A. F. Judd, BBM r1o0211-110224, Wilder, BBM 50528, 15
sinistral.
The main difference between the shells of area 25 and typical
bruneola is that the dominant color pattern is a yellow-lined form in-
stead of the darker usual bruneola pattern of plate 5, figure 17. The
yellow shells are rare in area 28, whereas in area 25, 21 percent have
a definitely yellow pattern without strong brown bands (pl. 5, fig. 15).
The usual form and color pattern of 37 percent of the shells (pl. 1,
fig. 30) has the embryonic whorls white; first quarter postembryonic
whorl white, next quarter tawny lined with pale pinkish buff, penulti-
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 63
mate cream buff lined with tawny, last whorl antimony yellow lined
above the periphery with tawny, below and at the periphery with
chestnut ; lip warm sepia, columella callus white. Length 18.7 mm.,
greater diameter 12.5 mm., spire height 9.6 mm., number of
whorls 63.
Eight percent are dark brown above the periphery and yellow be-
low (pl. 5, fig. 15a); the postembryonic whorls tawny faintly lined
or tinted with warm buff, banded above and below the periphery
with chestnut, below and at the periphery the ground is naples yel-
low. The typical form of a shell of the 19+mm. length class (pl.
5, fig. 15) measures: Length 19.7 mm., greater diameter 13.6 mm.,
spire height 10.0 mm.; postembryonic whorls shade from white to
light buff on the last half of the penultimate, last whorl above
the periphery white, tinted with light buff, finely spirally lined with
faint lines of chamois, below the periphery ground is mustard yellow
within the aperture lightening to naples yellow, and spirally lined with
buckthorn brown, except behind the edge of the lip where the bands
and lines are russet; impressed sutural band russet. The lightest
yellow pattern (pl. 5, fig. 150) has the postembryonic whorls white up
to the last whorl which is mustard yellow with a line of ochraceous
tawny at the edge of the periphery.
The dextral race of bruneola var. 2 is probably a remnant of two
races. The lower, a brown race similar to typical A. a. bruneola, may
possibly have been a pure one and not mixed with the yellow or upper
form similar to A. a. aureola. Above the brown race yellow forms
existed, and locality 170C-6 is the point where the two races over-
lapped.
The Wilder lot, BBM 50528, may indicate that a yellow race
occurs between areas 24 and 25. The shells were probably collected
over a wide area inclusive of area 24 because of the presence of white
shells; and above area 25 because no dextrals are found in the lot.
One specimen has a brown color pattern (pl. 5, fig. 16) ; postembry-
onic whorls white, banded on the first postembryonic whorl with
tawny, last whorl banded with chestnut. Thirty-three percent have a
yellow pattern similar to that of plate 5, figure 15. Sixty percent have
whitish color patterns similar to those of plate 1, figure 31, and
plate 5, figure 6. The yellow shells are probably a sinistral race of
A. a. aureola, but because the locality is not definite, the range of
color variation is not known and these shells have therefore been
lumped with A. a. bruneola var. 2 until more data are obtainable.
In like manner the white forms are probably A. a. ovum var. 2.
~
o
64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
ACHATINELLA APEXFULVA AUREOLA, new subspecies
PLATE I, FIGURES 34, 35; PLATE 5, FIGURES 19-2Ia
Achatinella turgida ovum PretFreR, Pilsbry and Cooke, Man. Conch., vol. 22,
p. 208, pl. 56, fig. 8 (only), 1914.
The shell is obese and resembles A. a. bruneola in form, but differs
in the color pattern, having a yellow ground usually lightly banded
with brown. The embryonic whorls of the holotype (pl. 1, fig. 34)
are white; first postembryonic whorl pale pinkish buff axially streaked
with sayal brown, penultimate whorl pale pinkish buff axially streaked
with ochraceous tawny ; impressed sutural band ochraceous tawny, last
whorl above the periphery pale pinkish buff, banded at edge of the
periphery and above the periphery with a band 2.8 mm. wide, amber
brown above the aperture, the band is lined with mustard yellow,
and on the last half whorl breaks up into three bands of amber brown,
just behind the edge of the lip a streak of warm sepia, last whorl
below the periphery mustard yellow; lip army brown streaked with
white ; columella callus army brown shading to white at the tip of the
callus. Length 20.4 mm., greater diameter 14.1 mm., spire height
9.8 mm., number of whorls 63.
Distribution, area 30: Kalauao-Hanaiki Ridge, type locality 184,
elevation 1,350-1,450 feet, 1926, also 1913, 7 sinistral 1929, 8 sinistral
1935; 185, elevation 1,450-1,500 feet, 1933; 187, elevation 1,600 feet,
1933; 1877, 1932; 188, elevation 1,650-1,700 feet, I sinistral 1933, all
Meinecke-collected (figs. 4, 4a, p. 53).
The form of the shell in locality 184 may be more elongate (pl. 5,
fig. 20) and have a darker spire than the holotype. Length 20.2 mm.,
greater diameter 13.0 mm., spire height 10.6 mm. Last half of first
postembryonic whorl, penultimate, and last whorl above the periphery
tilleul buff, lined, banded, and streaked with a shade of color between
ochraceous tawny and avellaneous; impressed sutural band same
color; last whorl at the edge of the periphery banded with tawny,
below the periphery mustard yellow and with a line of tawny, behind
the edge of the lip a streak of black. A single specimen (pl. 5, fig. 19)
has a white spire; last whorl below and just above the periphery
naples yellow, banded with white at the edge of the periphery, below
' the periphery banded with amber brown, just behind the edge of the
lip a streak of black.
The lightest color pattern in locality 185 (pl. 5, fig. 20a) has white
postembryonic whorls, banded and lined with tawny and ochraceous
tawny, last half whorl banded at the edge of the periphery with mars
brown, behind the edge of the lip a streak of black or mars brown.
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 65
The dextral shells of locality 187 are not so obese as those of locality
184. The usual form and pattern of a dextral (pl. 1, fig. 35) measures:
Length 19.8 mm., greater diameter 12.4 mm., spire height 10.5 mm.,
embryonic whorls white; postembryonic whorls shade from cream
color to naples yellow on the last whorl, last whorl below the periphery
mustard yellow, banded at the edge of the periphery with a faint band
of ochraceous tawny, below the periphery with tawny, behind the
edge of the lip a streak of black.
The obese and banded form of the shell (pl. 5, fig. 21) measures:
Length 18.9 mm., greater diameter 13.3 mm., spire height 8.8 mm.
Postembryonic whorls mustard yellow, faintly lined on the last whorl
above the periphery with cinnamon buff; impressed sutural band
russet ; last whorl at the periphery and below banded with ochraceous
tawny, russet and mars brown about the base, behind the edge of the
lip a streak 2 to 3 mm. wide of black. A narrow elongate form and a
yellow color pattern (pl. 5, fig. 21a) has the postembryonic whorls
naples yellow deepening to yellow ochre on the last whorl, lined above
the periphery of the last whorl with naples yellow. Length 20.2 mm.,
greater diameter 12.0 mm., spire height 11.1 mm.
There is some doubt in my mind concerning the accuracy of some
of the material collected by Meinecke in localities 184 and 187. After
comparing the material with his more recently collected shells found
after 1933, I believe that the early lots of shells collected prior to
1933, and plotted from memory, have been misplotted. I have taken
the liberty of arranging the sequence of areas in a probably correct
order, taking the 1933 and later collected material to be correct. In
doing so I may have made an error, and I hope that some student of
Hawaiian tree snails may check the order of color variations on this
ridge. The change in form from narrow dextral shells in area 28 to
sinistral obese ones in locality 184 and finally to narrow dextrals
again in locality 187 is most interesting if true. On the variation of
form, one would be led to suppose that the sequence is area 28, locality
187, area 29, locality 184, and finally locality 185, locality 188, and
area 31.
In locality 185 the color pattern and form of the shells is similar
to that of locality 184, the only difference being that a few specimens
(3 out of 10) have a white ground in locality 185, while in area 184
no white color patterns occur. The single shell from 188 has an en-
tirely yellow color pattern. I have considered this locality the border-
line extremity of the typical aureola area with a few specimens show-
ing the lighter color patterns of area 31.
66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
ACHATINELLA APEXFULVA AUREOLA var. 1
PLATE I, FicuRE 36; PLATE 5, FIGURES 22-22b
Area 31: Kalauao-Waimalu Ridge, locality 190A*, elevation
1,750-1,800 feet, 1932; 191A*, elevation 1,600 feet, 5 sinistral 1932,
all Meinecke-collected (figs. 4, 4a, p. 53).
The color pattern of the shells in this sector are white shells banded.
or tinted with yellow or banded with brown. A usual pattern in
locality 1I90A* (pl. 5, fig. 22) has the postembryonic whorls white, the
last half of the first postembryonic whorl and the first half of the
penultimate faintly lined and banded with tawny, the last half of the
penultimate faintly banded and lined with cream buff, the first half of
the last whorl mustard yellow with a peripheral band and supraperiph-
eral line of buckthorn brown, last half whorl white tinted with mustard
yellow, with the peripheral band buckthorn brown, behind the edge
of the lip a streak of mars brown. A brown color pattern (pl. 5,
fig. 22a) has the postembryonic whorls above the periphery antimony
yellow axially streaked and lined with amber brown; impressed sutural
band chestnut; last whorl and a half at the edge of the periphery
and below the periphery white tinted with cream color, in the umbili-
cal region a patch of amber brown. A white color form (pl. 5,
fig. 22b) resembles A. a. ovum. The entire shell is white, tinted on the
last whorl with mustard yellow, last fourth whorl faintly banded with
mustard yellow, behind the edge of the lip mars brown.
In the higher locality of 191A*, the usual color pattern (pl. 1, fig.
36) is a white shell banded on the last whorl above the periphery and
all below the periphery with mustard yellow. The shells in the upper
region are all sinistral; in 190A* they are all dextral.
ACHATINELLA APEXFULVA WAIMALUENSIS, new subspecies
PLATE I, FIGURE 45; PLATE 5, FIGURES 25, 25a
The shell looks like a form of A. a. ovum or A. a. aureola and their
varieties, but is distinct from them in having the first postembryonic
whorl banded with brown so that the whorl stands out, being darker
than the whorls above and below it. The 4. a. ovum and A. a. aureola
forms have a color pattern which usually darkens consistently on the
later whorls.
The embryonic whorls of the holotype (pl. 1, fig. 45) are cartridge
buff ; first postembryonic whorl banded on the lower portion of the
whorl with ochraceous tawny, the band decreasing from three-fourths
the width of the whorl to half its width, on the white penultimate and
INO: EF ACHATINELLA APEXFULVA DIXON—WELCH 67
last whorl the band narrows to a line and fades from ochraceous
tawny to a light shade of ochraceous tawny; below the periphery of
the last whorl the shell is banded with a light shade of ochraceous
tawny, base cream color; lip russet; columella callus light vinaceous
fawn. Length 20.7 mm., greater diameter 13.1 mm., spire height
11.4 mm., number of whorls 63.
Distribution, area 32: Waimalu-South Central Waimano Ridge,
type locality 204C, elevation 1,660 feet, Meinecke, 6 sinistrals 1933;
also 202C, elevation 1,500-1,550 feet; 203C, elevation 1,600-1,660
feet; 206C, elevation 1,600 feet, Meinecke, 1 sinistral 1933 (figs. 4,
44, p. 53).
The darkest color pattern (pl. 5, fig. 25) has the first half post-
embryonic whorl banded on the lower half of the whorl with a light
shade of clay color lined with white and fading to white on the last
half of the whorl, the penultimate whorl is white, lined on the lower
portion just above the edge of the periphery with honey yellow
shading to chamois, the last whorl is chamois all below the periphery
and just above the periphery, the remainder of the shell is white, be-
low the periphery banded with tawny.
The lightest pattern (pl. 5, fig. 25a) has the last two whorls white,
tinted and faintly spirally banded with naples yellow, behind the edge
of the lip a streak or axial stripe of black.
ACHATINELLA APEXFULVA WAIMALUENSIS var. 1
PLATE I, FIGURE 44
Area 32A: South Central Waimano-Central Waimano Ridge,
locality 202D, elevation 1,400 feet; 203D, elevation 1,450 feet,
Meinecke, 1933 (figs. 4, 4a, p. 53). Only one live specimen has been
collected in area 32A. The other shells are such dead specimens that
the range of color patterns is not known.
The color pattern on the live shell (pl. 1, fig. 44) from locality
202D is a shell with a color pattern similar to A. a. waimaluensis
(pl. 5, fig. 25), except that the spire is banded. The first embryonic
whorl is broken off, the remaining embryonic whorls light buff;
postembryonic whorls white, banded with ochraceous tawny, lower
half of last whorl naples yellow with a line of ochraceous tawny at
the edge of the periphery, on the last half whorl three bands of warm
sepia; lip not formed; columella callus pale vinaceous fawn. The
color pattern is intermediate between A. a. simulacrum and A. a.
waimaluensis, but closest to waimaluensis.
68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
ACHATINELLA APEXFULVA TURGIDA Newcomb
PLATE I, FIGURE 41; PLATE 6, FicuREs 2-10
Achatinella turgida NeEwcoms, Proc. Zool. Soc. London, 1853, p. 134, pl. 22,
figs. 10-10a.—PiILsBry and Cooke, Man. Conch., vol. 22, p. 294, pl. 20,
figs. 10-10a; pl. 56, figs. I-2, 4, 11 (only), 1914.
Achatinella swiftii NeEwcoms, Proc. Zool. Soc. London, 1853, p. 133, pl. 22,
figs. 9, 9a.—PitssBry and Cooke, Man. Conch., vol. 22, p. 306, pl. 20,
figs. 9, 9a (only), 1914.
Newcomb’s description of A. a. turgida may refer to either of his
original figures. I shall choose his figure 10 (pl. 6, fig. 2) to be the
lectotype, because it has the most usual pattern. The original speci-
men (pl. 6, fig. 3) of Newcomb’s figure 10 has white embryonic
whorls ; penultimate and last postembryonic whorls pallid mouse gray,
axially spotted and spirally banded with hessian brown, on the last
whorl near the middle of the whorl there are three axial bands of
cinnamon, behind the edge of the lip an axial band of hessian brown,
in the umbilical region below the last hessian brown band the ground
is cream buff, mixed with pallid mouse gray; impressed sutural band
hessian brown; lip and columella callus light vinaceous fawn. Length
19.7 mm., greater diameter 13.4 mm., spire height 10.9 mm., number
of whorls 64.
Distribution, area 35: Waiau-South Waimano Ridge, locality
200B*, elevation 850-1,000 feet, 7 dextral, 1 sinistral 1929 ; 202B, ele-
vation I,000-1,150 feet, 1913, I dextral, 8 sinistral 1914, 1920, 7 dex-
tral, 2 sinistral 1929, 12 dextral, 1 sinistral 1932, 1934; 202Bb*,
elevation 1,050-1,150 feet, 5 dextral, 1 sinistral 1929, all collected by
Meinecke. Waimano, Wilder, BBM 50598; Waimalu, J. S. Emer-
son, BBM 102218. Pilsbry and Cooke report A. a. turgida to have
been taken in Waiawa by I. Spalding. This is an error, for no A. a.
turgida are found except in the region of Waimalu to Waimano
(figs. 4, 4a, p. 53). Holotype in BM marked with an x.
The usual form and color pattern (pl. 1, fig. 41) of A. a. turgida
in area 35 is very much like the lectotype in color. The shell measures:
Length 19.0 mm., greater diameter 13.2 mm., spire height 10.1 mm.,
number of whorls 63. The embryonic whorls are white; postem-
bryonic whorls diamine brown, splotched and banded with broken
bands of pale smoke gray tinted with chamois; impressed sutural
band diamine brown; lip and columella callus pale vinaceous fawn.
The darkest or reddish-brown color pattern and narrow form of the
shell (pl. 6, fig. 4) has the postembryonic whorls diamine brown and
looks like a sinistral specimen of A. a. apexfulva which has white
embryonic whorls instead of yellow ones. Length 19.5 mm., greater
diameter 12.6 mm., spire height 10.0 mm.
NO. I ACHATINELLA APEXFULVA DIXON—-WELCH 69
The usual yellow pattern and obese form of the shell (pl. 6, fig. 4a)
has the first half postembryonic whorl ochraceous tawny banded on
the lower half of the whorl with pale ecru drab, remaining whorls
chamois splotched with snuff brown, hessian brown, and saccardo’s
umber, supraperipheral band on the last whorl warm sepia, last
whorl below the periphery ornamented with zigzag streaks of snuff
brown; impressed sutural band warm sepia. Length 19.4 mm., greater
diameter 14.5 mm., spire height 9.7 mm.
The lightest or yellowish white color pattern (pl. 6, fig. 4b) has the
postembryonic whorls white, on the first quarter whorl a patch of
mars brown, remaining whorls streaked with ochraceous tawny; im-
pressed sutural band on the first postembryonic whorl and a half
diamine brown, last whorl and a half the sutural band is the color
of the whorl below; lip light vinaceous fawn.
The typical form of a dextral shell and a variant of the yellow
color pattern is figured on plate 6, figure 4c} first two postembryonic
whorls pale gull gray, axially streaked with zigzag lines of neutral
gray and deep neutral gray, and ochraceous tawny, last whorl
cartridge buff tinted with cream color streaked with white and with
axial splotches of ochraceous tawny, prout’s brown, and mars brown.
Length 19.5 mm., greater diameter 13.7 mm., spire height 9.4 mm.
A characteristic gray color pattern (pl. 6, fig. 4d) is also found in
all turgida localities; the postembryonic whorls are pale gull gray
spirally banded lined and splotched with diamine brown. The yellow
color pattern (pl. 6, fig. 4e) may lack or have very little axial
splotching ; the first two postembryonic whorls are cartridge buff tinted
with deep olive buff, last whorl chamois spirally lined and banded
with chestnut and faintly axially streaked on the last half whorl with
chestnut.
The size of the dextral shells varies from narrow specimens (pl. 6,
fig. 5) to obese forms (pl. 6, fig. 6). The shell of figure 5 is a
peculiar rare color pattern of the reddish-brown form, having a
subperipheral chamois band faintly streaked with tawny on the last
whorl, the remaining postembryonic whorls diamine brown; outer
margin of lip army brown shading to white on the inner margin;
columella callus white. Length 19.1 mm., greater diameter 12.5 mm.,
spire height 10.6 mm. The obese form of figure 6 measures: Length
19.2 mm., greater diameter 14.1 mm., spire height 9.6 mm.; the
postembryonic whorls diamine brown spirally lined with apricot
yellow on the penultimate, and banded and lined on the last whorl
with mars yellow, light buff, or antimony yellow, which bands are
axially streaked and lined with diamine brown or mouse gray.
7O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
The specimens used by Newcomb for his original figures of
A. swiftii were not located in the British Museum. One lot labeled
the type of A. swiftii contained shells which do not match the original
figures, and are forms of A. a. perplexa.
Newcomb’s figure 9 (pl. 6, fig. 7) closely matches yellow forms
splotched with brown of A. a. turgida (pl. 6, fig. 4c). In the New-
comb collection at Cornell University there is a specimen of A. a.
turgida (pl. 6, fig. 9) which closely resembles Newcomb’s figure 9.
Plate 6, figure 10, shows a specimen which looks very much like
Newcomb’s figure ga (pl. 6, fig. 8), the postembryonic whorls are
diamine brown, penultimate whorl is lined with white, last whorl
banded below the periphery with yellow ochre, with a line of diamine
brown near the center of the band; lip light vinaceous fawn.
Since A. swiftii so closely resembles patterns of A. a. turgida
I consider it to be a synonym of turgida. A. swiftii has page priority
over A. a. turgida but I am reluctant to make turgida a synonym of
swiftti since the type of swiftii has not been located and the type of
turgida has. Moreover, since the original figures do not match the
original description, there is some doubt about the exact identification
of A. swiftii. Therefore it is best to put swifti in the synonomy of
A, a. turgida.
ACHATINELLA APEXFULVA MEADOWSI, new subspecies
PLATE I, FIGURE 42; PLATE 6, FIGuRES II-I2
Achatinella turgida ovum PFEIFFER, Pilsbry and Cooke, Man. Conch., vol. 22,
p. 208, pl. 56, fig. 9 (only); pl. 56, fig. 12 (an intermediate form between
A. a. meadowsi and A. a. turgida but closest to meadowsi), 1914.
The shell of the holotype (pl. 1, fig. 42) is a yellow-brown form
of A. a. turgida differing mainly in color pattern; the embryonic
whorls pale pinkish buff ; postembryonic whorls yellow ochre heavily
banded on the first two whorls and above the periphery of the last
whorl with amber brown, below the periphery the amber brown
bands are faint so that the color is mostly yellow ochre, in the umbili-
cal region a band of black; impressed sutural band warm sepia or
black ; lip army brown; columella callus light vinaceous fawn; behind
the edge of the lip a streak of black. Length 19.6 mm., greater di-
ameter 13.7 mm., spire height 9.6 mm., number of whorls 6$.
Distribution, area 34: South Waimano-South Central Wai-
mano Ridge, type locality 212b, elevation 1,300 feet ; 213a-213b, ele-
vation 1I,350-1,400 feet, 17 sinistral; 213a, elevation 1,350 feet, all
collected by Meinecke, 1933. Meadows and Welch (BBM 107555-
NO. I ACHATINELLA APEXFULVA DIXON—WELCH fa
107550) collected this form on the Waiau-South Waimano Ridge
somewhere above area 35. The exact locality was not plotted, so is
not being used in this paper (figs. 4, 4a, p. 53).
A. a. meadowsi is found in the Meinecke lots of A. a. turgida
from area 35, but only a few specimens in the different lots, and
some lots entirely lack this color form. Area 35 of A. a. turgida is
probably a borderline area between A. a. meadovwsi and a distinct pure
race of A. a. turgida. The subspecies is named after E. J. Meadows,
of Honolulu.
The elongate form (pl. 6, fig. 11) measures: Length 20.0 mm.,
greater diameter I2.5 mm., spire height 11.0 mm.; postembryonic
whorls yellow ochre lightly streaked, banded, and lined with ochra-
ceous tawny, last whorl banded at and below the periphery with
warm sepia, or black; impressed sutural band warm sepia or black.
The darkest color pattern (pl. 6, fig. 11a) has the postembryonic
whorls carob brown banded on the last whorl and a half with a band
of yellow ochre which is lined and streaked with ochraceous tawny.
An obese specimen (pl. 6, fig. 12) with the postembryonic whorls
yellow ochre and very faintly lined and banded with ochraceous
tawny, measures: Length 19.0 mm., greater diameter 14.0 mm., spire
height 9.3 mm.
ACHATINELLA APEXFULVA MEADOWSI var. 1
PLATE 6, FIGURES I3-14a
In locality 213a (fig. 4, p. 53) two specimens are found which did
not have the usual glossy surface of typical A. a. meadowsi, but have
a very dull appearance. The postembryonic whorls (pl. 6, fig. 13) are
pale pinkish buff spirally banded and lined with ochraceous tawny ;
impressed sutural band warm sepia or black.
Wilder (BBM 50537) obtained from somewhere in the Waimano
region a series of the shells all of which are dull forms. The color
pattern ranges from dark patterns such as plate 6, figure 14, to light
ones such as plate 6, figure 14a. The dark color pattern has the
postembryonic whorls chestnut streaked with antimony yellow, sub-
peripheral band light buff with a central band of ochraceous tawny ;
impressed sutural band warm sepia. The light pattern (pl. 6, fig. 14a)
is colored on the first postembryonic whorl with pinkish buff; im-
pressed sutural band pinkish buff; banded in the umbilical region
with snuff brown.
This dull form may be a variety of A. a. meadowsi which occurs
on the Waimalu-South Waimano Ridge or somewhere between areas
72 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
34 and 35. The dull race as can be noted above has a wider range of
color pattern than typical A. a. meadowst.
ACHATINELLA APEXFULVA MEADOWSI var. 2
PLATE I, FIGURE 43; PLATE 5, FIGURES 23, 24
Area 33: Waimalu-South Central Waimano Ridge, locality
200Ca, elevation 1,200-1,350 feet, 1932; 200C, elevation 1,400 feet,
2 sinistrals 1933; 201Ca, elevation 1,250-1,400 feet, 10 sinistrals
1933; 201C, elevation 1,450 feet, 1933, all collected by Meinecke
(figs. 4, 4a, p. 53). This form also collected by J. S. Emerson (BBM
102229). .
From the dark brown meadowsi forms of area 34 the shells become
much lighter in shade the higher the locality. Area 33 is really an
intermediate one between the light color pattern of A. a. waimaluensis
and A. a. meadows, but is closest to meadows: in color pattern.
The usual color pattern (pl. 1, fig. 43) has the embryonic whorls
light buff lightening to pale pinkish buff lined with white; first post-
embryonic whorl pale pinkish buff lined with pinkish buff, penultimate
whorl cream color, subsutural band light ochraceous buff, last whorl
mustard yellow tinted with buckthorn brown; the impressed sutural
band on the first postembryonic whorl shades from mikado brown to
warm sepia, on the penultimate whorl the upper portion of the sutural
band is banded with warm sepia, the lower half is that of the ground
color, the width of the warm sepia band narrows continually so that
on the last whorl only a fringe of warm sepia exists on the upper
portion of the band; lip vinaceous brown; columella callus light
vinaceous fawn.
The lightest color pattern (pl. 5, fig. 24) is pale pinkish cinnamon,
tinted on the last whorl with faint lines of warm buff; impressed
sutural band russet lightening to avellaneous on the last whorl. A
pattern (pl. 5, fig. 23) that looks like a light form of A. a. meadowsi
also occurs; postembryonic whorls maize yellow spirally lined or
banded with a light shade of clay color or a yellowish shade of
ochraceous tawny.
ACHATINELLA APEXFULVA PERPLEXA Pilsbry and Cooke
PLATE I, FIGURE 47; PLATE 6, FiGuRES 16-10d
Achatinella turgida perplexa Pitspry and Cooxe, Man. Conch., vol. 22, p. 206,
pl. 56, figs. 5-5d (only), 1914.
To quote from Pilsbry and Cooke:
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
NO... ACHATINELLA APEXFULVA DIXON—WELCH 73
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 percent are sinistral.
Length 20.7 mm., diam. 13.6 mm., 64 whorls.
Length 18.0 mm., diam. 12 mm.
Lateral spurs, Badl northern ridge of Waimano Tatley: 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. Figures 5, 5a are typical patterns.
Distribution, area 40: Waimano- ee Ridge, locality 2214*
elevation 1,050-1,100 feet, 1926; 222*, elevation 1,100-1,150 feet,
1929, both collected by Meinecke (figs. 4, 4a, p. 53). Cooke’s type lo-
cality in Waimano probably extends over areas 40 and 44, because the
type lot (BBM 72723-72753) contains a mixture of the typical per-
plexa patterns of area 40 and the migripicta var. I patterns which in the
Meinecke collection are limited to area 44. Other collectors of this
form are Wilder, BBM 50550-50551, and J. S. Emerson, BBM
102243. The Emerson lot is dominantly from the upper region of
area 44, while the Wilder lots come mostly from the lower area 40.
Both lots have many intermediate forms the patterns of which would
identify them with either area.
Pilsbry’s plate 56, figure 5, is considered the lectotype and is re-
produced in this paper on plate 6, figure 16. The figure is slightly
larger in proportion to the other figures. All the typical perplexa
forms of the original Cooke lot are studied. The usual form (pl. 1,
fig. 47) and gray pattern on about 55 percent of the shells has the
embryonic whorls white; first half postembryonic whorl pale pinkish
buff axially streaked with cinnamon, remaining postembryonic whorls
white finely axially streaked and lined with light drab or a shade
between light drab and tawny olive, peripheral band on last whorl
white; impressed sutural band a faint shade of tawny olive; lip
and columella callus seashell pink. Length 18.4 mm., greater diameter
12.8 mm., spire height 10.0 mm.
A variant of the typical color pattern, which may be considered a
grayish pattern (pl. 6, fig. 17), has the first half postembryonic whorl
sayal brown banded with a line of pale pinkish buff, last half verona
brown; penultimate whorl drab streaked and tinted with benzo brown
on the first half of the whorl, lower fourth of the whorl banded with
pale gull gray, last whorl pale gull gray axially streaked with white,
74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
spirally banded above the periphery with a band of pale pinkish buff
axially streaked with drab, peripheral line fuscous, subperipheral
band warm sepia streaked with hair brown and smoke gray, basally
lined with sepia; spire concave in outline.
The grayish color pattern (pl. 6, fig. 18) may be more lined than
figure 17; first postembryonic whorl mikado brown, faintly lined with
pale pinkish buff, penultimate whorl ground pale pinkish buff covered
with close-set axial streaks of drab, cinnamon drab and benzo brown,
spirally lined with fuscous, last whorl pale pinkish buff, banded above
the periphery with a light drab band which is lined with olive brown,
banded at the periphery with olive brown, below the periphery with
hair brown, lined with fuscous, lip light vinaceous fawn. This pattern
collected by Meinecke in area 40 is also found in the Cooke lots.
Brownish color patterns are depicted on plate 6, figures 17a, 17D.
Figure 17a is an extreme elongate specimen, with the spire straight
in outline. Length 19.2 mm., greater diameter 11.8 mm., spire height
II.1 mm. Postembryonic whorls mikado brown, spirally lined and
faintly streaked with zigzag streaks of tilleul buff, peripheral band
tilleul buff lined with vinaceous buff. Figure 17) is a darker color
pattern and the obese form of the shell. Length 19.8 mm., greater di-
ameter 13.5 mm., spire height 11.0 mm. First postembryonic whorl
cream color, with impressed sutural band ochraceous tawny, last
two whorls army brown streaked with pale smoke gray.
Meinecke found a white sinistral specimen in locality 2214* (pl. 6,
fig. 18a) the postembryonic whorls white, first two postembryonic
whorls lined with pinkish buff, last whorl lined with cinnamon buff.
While the paratype Cooke lots are all dextral, 46 percent of the
shells in the Wilder collection are sinistral. These sinistrals have a
different color pattern range from those collected by Cooke. Yellowish
or pink color patterns occur. The usual form and color pattern of
50 percent of the sinistrals (pl. 6, fig. 19) has the embryonic whorls
pale pinkish buff; first half postembryonic whorl pale pinkish buff,
axially streaked with sayal brown, next half whorl streaked with
army brown, penultimate and last whorl light drab axially streaked
with hair brown, and banded below the suture with white, last whorl
below the periphery white with two lines of benzo brown. Length
18.1 mm., greater diameter 12.3 mm., spire height 9.4 mm.
A variant (pl. 6, fig. 19a) of the typical gray color pattern has the
first half of the first postembryonic whorl ochraceous tawny, last half
russet, penultimate tawny olive lined with smoke gray, last whorl
smoke gray spirally lined with tawny olive. It is an obese form.
Length 18.2 mm., greater diameter 13.3 mm., spire height 9.4 mm.
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 75
Twenty-two percent have a yellow ground (pl. 6, fig. 19b) the last
two whorls cream color with bands of buffy brown which are axially
streaked with natal brown. Eleven percent are pinkish patterns. The
darkest of these (pl. 6, fig. 19c) has the postembryonic whorls colored
pinkish cinnamon. The lightest pink pattern (pl. 6, fig. 19d) is sea-
shell pink axially streaked with onion-skin pink, and spirally banded
and lined on the last whorl with pale vinaceous fawn.
ACHATINELLA APEXFULVA COOKEI Baldwin
PLATE I, FIGURE 49; PLATE 7, FIGURES 7-10, 12-12d
Achatinella cooket BALDWIN, Proc. Acad. Nat. Sci. Philadelphia, 1895, p. 220,
pl. 10, fig. I5.
Achatinella turgida cookei BALpwin, Pilsbry and Cooke, Man. Conch., vol. 22,
p. 300, pl. 57, figs. 15-17, 1914.
Distribution, area 45: Waimano-Manana Ridge, locality 225%,
elevation 1,300-1,400 feet, 35 sinistral 1929; 225-1, elevation 1,500
feet, 2 dextral 1933; 226*, elevation 1,300-1,426 feet, 15 dextral
1929, all collected by Meinecke (figs. 4, 4a, p. 53). Type lot BBM
167352 of 8 dextral, collected by C. H. Cooke and labeled Ewa. The
D. D. Baldwin lot, BBM 546609, of 4 dextral, was obtained from C. H.
Cooke. Type locality on the Waimano-Manana Ridge was later
collected by C. M. Cooke, Jr., BBM 72697-72698, 4 dextral, 4 sinistral,
and R. A. Cooke, BBM 58340, 7 dextral and 2 sinistral. Probably
the later collections include a slightly different region than the type
locality, because the type lots contain no sinistrals. Also collected
by Wilder, in Waimano, BBM 50594-50595, 142 sinistral, and
13 dextral.
The exact location of the type locality has not been plotted, but the
color patterns in the type lot in the Bishop Museum closely match
those of the shells from area 45. The shell Baldwin used for his
original figure cannot be found in either the Bishop Museum or the
Academy of Natural Sciences of Philadelphia.
A shell in the Academy of Natural Sciences, figured on plate 7,
figure 8, more closely approximates the form of Baldwin’s figure re-
produced on plate 7, figure 7, but differs in size, thickness of the lip,
and in having a band of pinkish buff in the umbilical region. The
embryonic whorls of figure 8 are white shading to cartridge buff on
the last half embryonic whorl; postembryonic whorls pinkish buff,
last whorl spirally lined with lines darker than the ground, on the
last half whorl a band of white at the periphery, below the periphery
70 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
last whorl hessian brown; lip and columella callus light vinaceous
fawn. Length 19.4 mm., greater diameter 14.2 mm., number of whorls
6. While the shell of figure 8 does not match the original figure and
measurements in all details, it comes from what is supposed to be the
type lot and best matches the form of Baldwin’s original figure. This
shell is therefore considered the lectotype.
The usual form of the shell selected from the original type lot
collected by C. H. Cooke is shown on plate 7, figure 9; the embryonic
whorls white ; postembryonic whorls pale pinkish buff tinted with light
buff, lower half of last whorl a dark shade of diamine brown or dark
livid brown. Length 19.2 mm., greater diameter 13.4 mm., spire
height 9.8 mm. A narrow specimen (pl. 7, fig. 10) obtained in the
type lot has the postembryonic whorls white, with the base warm
blackish brown. Length 19.5 mm., greater diameter 12.8 mm., spire
height 10.4 mm.
_ Plate 1, figure 49, shows a dextral specimen from locality 225-1.
The postembryonic whorls are white, last whorl lower half warm
blackish brown. Length 19.5 mm., greater diameter 12.8 mm., spire
height 10.6 mm., number of whorls 63.
The range of color patterns in the Wilder collection is shown on
plate 7, figures 12-12d. All with the exception of the shell of figure
12d have the lower half of the last whorl a deep shade of diamine
brown or warm blackish brown. The usual form and color pattern
(pl. 7, fig. 12) has the postembryonic whorls white, tinted with
ochraceous tawny ; impressed sutural band ochraceous tawny. Length
18.3 mm., greater diameter 12.3 mm., spire height 9.7 mm. An elon-
gate shell (pl. 7, fig. 12a) with an entirely white spire measures:
Length 19.6 mm., greater diameter 12.3 mm., spire height 10.9 mm.
Thirty-six percent have a definitely yellow pattern. The usual
form and color pattern of one of the yellow shells (pl. 7, fig. 12b) has
the embryonic whorls cartridge buff; postembryonic whorls ochra-
ceous tawny with an impressed sutural band of tawny. Length
19.0 mm., greater diameter 12.4 mm., spire height 9.9 mm. An obese
shell and a lined form of the yellow pattern (pl. 7, fig. 12c) measures:
Length 18.2 mm., greater diameter 13.1 mm., spire height 9.5 mm.;
last two postembryonic whorls ochraceous tawny streaked with tawny
and finely spirally lined with white. Five shells of the yellow pattern
lack the typical dark cookei base. One of these (pl. 7, fig. 12d) has
the postembryonic whorls light buff, faintly axially streaked or tinted
ochraceous tawny; impressed sutural band ochraceous tawny.
NO. I ACHATINELLA APEXFULVA DIXON—-WELCH Wg)
ACHATINELLA APEXFULVA COOKEI var. 1
PLATE I, FIGURE 46
Area 39: North Central Waimano-North Waimano Ridge,
locality 211B, elevation 1,350-1,425 feet; 212B, elevation 1,400 feet,
I dextral, 3 sinistral; 213B, elevation 1,350-1,450 feet, 2 dextral;
214B, elevation 1,500-1,560 feet, 9 dextral, all Meinecke-collected,
1933 (figs. 4, 4a, p. 53).
The shells of area 39 have a series of color patterns similar to those
of the shells of area 45. The only reason for separating the two
areas is that a few shells occur in area 39 which do not have the
usual solid dark base of typical A. a. cookei and differ from the
patterns found in area 45. The shell may be white (pl. 1, fig. 46)
with a single band of black or warm blackish brown below the
periphery of the last whorl. One white specimen with a vinaceous
fawn lip and a patch of black behind the edge of the lip, below
the periphery, was found in locality 214 B.
ACHATINELLA APEXFULVA COOKEI var. 2
PLATE I, FiGuRE 56; PLATE 7, FicuRES II-11}
Area 49: Manana-Waiawa Ridge, locality 237f, elevation,
1,700 feet ; 237fa, elevation 1,500-1,600 feet, Meinecke 1933 (figs. 4,
4a, p. 53).
Sandwiched in between A. a. nigripicta (area 50), and migripicta
var. 2 (area 48) is a small region containing shells with A. a. cookei
patterns. The usual form and color pattern of the shell (pl. 1, fig. 56)
is more obese than the usual form in area 45, and the base is banded
instead of being a solid dark color. The embryonic whorls are pale
pinkish buff ; postembryonic whorls white, first half of last whorl lined
at the edge of the periphery with olive brown, subperipheral band
white, 2 mm. wide, tinted with pale mouse gray, on the last half
whorl, peripheral band warm blackish brown, subperipheral band
white, I mm. wide, base of last whorl warm blackish brown, with a
band of cream buff and a line of amber brown. Length 19.1 mm.,
greater diameter 13.9 mm., spire height 9.0 mm., number of
whorls 64.
The usual cookei base occurs on some specimens but the spire is
usually lined (pl. 7, fig. 11a); postembryonic whorls pale pinkish
buff spirally lined with ochraceous tawny. This shell is also the
obese form. Length 19.8 mm., greater diameter 13.9 mm., spire height
10.1 mm. A rare pattern (pl. 7, fig. 110), and an elongate shell found
78 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
on one specimen in locality 237f has the postembryonic whorls mars
yellow, lined and faintly axially streaked with light buff, last whorl
banded just below the edge of the periphery with pale pinkish buff,
the ground below this band light buff spirally banded with mars yellow
with a line of warm blackish brown; impressed sutural band russet.
Length I9.0 mm., greater diameter 12.7 mm., spire height 10.7 mm.
The darkest color pattern (pl. 7, fig. 11) found on only one specimen
has the postembryonic whorls banded with argus brown, lined and
axially streaked with light buff, last whorl just above the periphery
and all below warm blackish brown. The color of the lip and
columella callus is usually light vinaceous fawn.
ACHATINELLA APEXFULVA NIGRIPICTA, new subspecies
PLATE I, FiGURE 55; PLATE 7, FIGURES 3-4
Achatinella turgida simulacrum Pitssry and Cooke, Man. Conch., vol. 22, p. 300,
pl. 56, figs. 13-13b (only), 1914.
The shell of the holotype (pl. 1, fig. 55) has the embryonic whorls
white; postembryonic whorls pale pinkish buff banded with a very
deep shade of diamine brown or black; impressed sutural band same
black color ; lip and columella callus vinaceous fawn. Length 18.2 mm.,
greater diameter 12.2 mm., spire height 9.3 mm., number of whorls 6.
Distribution, area 50: Manana-Waiawa Ridge, type locality
237b, elevation 1,450-1,500 feet, 1933; also in locality 237c, elevation
1,350-1,400 feet; 237d, elevation 1,500 feet; 237e, elevation 1,600-
1,650 feet, 3 sinistral, all Meinecke-collected, 1933 (figs. 4, 4a, p. 53).
Another collector of this form is Vasconcellos in 1915, for Thurston,
BBM 130587-130588.
A variation (pl. 7, fig. 3) of the typical color pattern, and an
elongate shell, has a lighter colored spire; postembryonic whorls pale
pinkish buff spirally lined, and banded with ochraceous buff, and
lined with black, last whorl at and below the periphery black with a
band of pale pinkish buff just below the periphery. Length 18.8 mm.,
greater diameter 12.3 mm., spire height 10.5 mm. A lined pattern
(pl. 7, fig. 3a) has the embryonic whorls black or a deep shade of
diamine brown, lined with light buff. The color pattern (pl. 7,
fig. 3b) may be solid black or a deep shade of diamine brown, which
looks like a form of A. a. apexfulva; the last half of the first post-
embryonic whorl is lined with white. A rare pattern (pl. 7, fig. 3c)
on one or two shells has the postembryonic whorls white spirally
lined with tawny and black, banded below the periphery with black ;
impressed sutural band tawny. This shell also shows the usual form
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 79
of a dextral. Length 18.2 mm., greater diameter 12.5 mm., spire
height 9.3 mm. One white specimen (pl. 7, fig. 3d) has the impressed
sutural band edged with vinaceous cinnamon; last whorl below the
periphery lined with chocolate. Another light color pattern (pl. 7,
fig. 4) has the postembryonic whorls light buff spirally lined with
black.
ACHATINELLA APEXFULVA NIGRIPICTA var. 1
PLATE 1, Figure 48; PLATE 7, FicuRES 1-2g
Achatinella turgida perplexa Pitspry and Cooke, Man. Conch., vol. 22, p. 296,
pl. 56, figs. 5e, 5f (only), 1914.
Area 44: Waimano-Manana Ridge, locality 223, elevation 1,250-
1,300 feet, a few dextral juvenile shells, Meinecke, 1933 (figs. 4,
Aa, p. 53). A few specimens are found in locality 225 mixed with
typical A. a. cookei forms so that probably area 44 extends as far
as this locality. Other collectors of this form are Wilder, BBM
10449-10450, Cooke, BBM 72700-72722, and J. S. Emerson, BBM
10717-10718. Their lots are usually mixed with forms from the
lower region of area 40 and have been separated out from the
lots of A. a. perplexa. The shells of area 44 are so close to A. a.
nigripicta in form and color pattern that they can easily be con-
fused with them. This form of A. a. nigripicta var. 1 usually has a
more yellow ground color and no white color patterns. The placing of
the bands on the shells also differs. A. a. nigripicta occurs opposite
area 45 of A. a. cookei and not area 44 as one would suppose. All
this region needs to be re-collected very carefully to note whether
all areas have been accurately plotted.
The localized Meinecke material from area 223 is very scanty so
that form and color-pattern range is not determinable. The shell may
be dextral or sinistral. The embryonic whorls (pl. 1, fig. 48) are
white; last two postembryonic whorls and a half pale pinkish buff
spirally banded with a dark shade of diamine brown and faintly axially
streaked on the last whorl above the periphery with grayish olive ; lip
not formed; columella callus light vinaceous fawn. A lighter color
pattern (pl. 7, fig. 1) is pale pinkish buff on the postembryonic
whorls, spirally lined with sayal brown, and with warm sepia on the
-penultimate, last whorl lined with warm buff above the periphery,
warm sepia at the periphery, below the periphery, subperipheral
band pale pinkish buff, a line of warm sepia and a band of cream
buff about the base. A dark color pattern and a sinistral (pl. 7,
fig. 1a), has the embryonic whorls white; first postembryonic whorl
6
80 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
white, banded and lined with ochraceous tawny, penultimate and last
whorl above the periphery, upper half pale pinkish buff axially
streaked and spirally lined with ochraceous tawny, lower half a deep
shade of diamine brown or black, and banded about the base with
ochraceous tawny.
In the J. S. Emerson collection there is a large series of these
shells, with some A. a. perplexa specimens mixed from area 40. They
are made up of yellow and reddish-brown patterns. The usual form
and color pattern on 30 percent of the dextrals (pl. 7, fig. 2), has
the embryonic whorls white; postembryonic whorls warm buff,
spirally lined on the penultimate with verona brown, last whorl
banded and lined with verona brown and snuff brown, peripheral
band pale pinkish buff; lip and columella callus shell pink. Length
18.5 mm., greater diameter 12.7 mm., spire height 10.0 mm. The
color pattern may be lighter (pl. 7, fig. 2a), postembryonic whorls
warm buff, lined with ochraceous tawny; upper edge of impressed
sutural band russet, lower half warm buff ; base lined with warm sepia.
This yellow form may be lined and banded on the last two whorls
with diamine brown over a warm buff ground (pl. 7, fig. 2b). A
variant of this pattern has a white peripheral band on the last whorl
between the two diamine brown bands. Figure 2b shows an extreme
obese dextral. Length 18.6 mm., greater diameter 14.0 mm., spire
height 9.0 mm.
The reddish-brown pattern (pl. 7, fig. 2c) may be solid liver brown
on the last two and a half whorls, first half postembryonic whorl
ochraceous tawny and looks like a form of typical A. a. apexfulva.
The form and usual color pattern of a sinistral shell (pl. 7, fig. 2d),
has the last two whorls and a half liver brown, with a band of pale
pinkish buff at the periphery on the first half of the last whorl, and
a band or line of warm buff above the periphery on the last half of
the last whorl. Length 19.8 mm., greater diameter 13.6 mm., spire
height 10.4 mm. The shell may be narrower and more banded (pl. 7,
fig. 2¢). Length 19.2 mm., greater diameter 12.3 mm., spire height
10.1 mm. The first postembryonic whorl is light buff, last two
whorls a dark shade of diamine brown banded on the penultimate
with light buff, last whorl at and below the periphery banded with
white. An extremely lined form (pl. 7, fig. 2f) has the first half of
the penultimate whorl light buff spirally lined with ochraceous tawny,
last whorl and a half light buff, spirally lined above the periphery
with bone brown, base solid bone brown; impressed sutural band
amber brown. A rare pattern (pl. 7, fig. 2g) is diamine brown, lined
with pale gull gray.
NO. I ACHATINELLA APEXFULVA DIXON—-WELCH . 81
ACHATINELLA APEXFULVA NIGRIPICTA var. 2
PLATE 2, FIGURES I, 2; PLATE 7, FIGURES 5-60
Area 48: Manana-Waiawa Ridge, locality 237g, elevation 1,750-
1,800 feet, 2 dextral ; 237h, elevation 1,500-1,700 feet, Meinecke, 1933.
Area 48A: Manana-Waiawa Ridge, locality 238a, elevation
I,750-1,900 feet; 238b elevation 1,800-1,850 feet, Meinecke, 1933
(figs. 4, 4a, p. 53).
The shells of area 48 differ from A. a. migripicta in having very
obese dextral shells with usually a white ground instead of a yellow-
ish one. The usual form and color pattern of the shell (pl. 2, fig. 1)
measures: Length 19.4 mm., greater diameter 13.5 mm., spire height
9.5 mm.; embryonic whorls white; postembryonic whorls white
spirally banded and lined with a dark shade of diamine brown. An
obese form (pl. 7, fig. 5) with a white ground and black banded
postembryonic whorls measures: Length 20.3 mm., greater diameter
14.2 mm., spire height 9.8 mm.
In area 48a the color patterns are close to those of area 48. The
form of the shells shows no variation from those of area 50. The
darkest pattern (pl. 7, fig. 6a) has the postembryonic whorls colored
a dark shade of diamine brown, last whorls banded and lined at and
below the periphery with pale pinkish cinnamon. A lined color pat-
tern (pl. 7, fig. 6) has the postembryonic whorls white tinted with light
buff, lined with verona brown and black, subperipheral band white,
below which are two bands of black, in the umbilical region a band
of warm buff. The usual light color pattern (pl. 2, fig. 2) has the
postembryonic whorls white, lined on the last two whorls just above
the periphery with a line of mikado brown, at and below the periphery
lined and banded with black.
ACHATINELLA APEXFULVA SIMULACRUM Pilsbry and Cooke
PLaTE I, FIGURE 50; PLATE 7, FIGURES 13-15a
Achatinella turgida simulacrum Pitspry and Cooke, Man. Conch., vol. 22, p. 299,
pl. 56, figs. 14-14d (only), 1914.
The shell figured on plate 56, figure 14a, of the Manual of Conch-
ology is considered the lectotype of A. a. simulacrum and is repro-
duced in this paper on plate 7, figure 13. The embryonic whorls are
a light shade of cartridge buff, banded on the last half embryonic
whorl with a faint color of warm buff; postembryonic whorls upper
half white, lower half amber brown streaked and lined with chestnut,
82 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
at the edge of the periphery of the last whorl a band of white, below
the periphery a wide band of hessian brown, and a band of white
lined with hessian brown, and a band of antimony yellow; impressed
sutural band white; lip and columella callus vinaceous fawn. Length
18.9 mm., greater diameter 13.0 mm., spire height 9.5 mm., number of
whorls 6.
Distribution, area 46: Waimano-Manana Ridge, locality 2274,
elevation 1,550 feet, I sinistral 1933; 227b, elevation 1,600 feet, 3
sinistral 1933; 227c, elevation 1,700 feet, 3 sinistral 1933; 228*,
elevation 1,550-1,700 feet, 1929; 228a, elevation 1,750 feet, 1933;
229*, elevation 1,750-1,800 feet, 1929; 229a, elevation 1,600 feet,
4 sinistral 1933, all Meinecke-collected (figs. 4, 4a, p. 53). This area
also collected by H. Lemke. Lectotype, ANSP 108063. Type locality,
“Waimano-Manana ridge, along the summit trail, above the locality
of A. t. cookei (Spalding, Pilsbry, Merriam), types 108063 A.N.S.”
(Pilsbry and Cooke, 1914).
The type locality was probably somewhere in the region of locali-
ties 228 and 228a. The usual color pattern and form of the shell
(pl. 1, fig. 50) from these localities, has the embryonic whorls pale
pinkish buff shading to cream color; postembryonic whorls white
with a band of ochraceous tawny just above the edge of the periphery,
which band is lined with white, last whorl below the white peripheral
or subperipheral band warm blackish brown with three bands of pale
pinkish buff at the edge of the aperture; lip and columella callus
vinaceous fawn. Length 19.6 mm., greater diameter 13.5 mm., spire
height 9.5 mm., number of whorls 64.
The usual narrow form of the shell (pl. 7, fig. 14) is banded and
has a color pattern resembling plate 1, figure 50, except that the last
whorl is more completely banded on the base with pale pinkish buff.
Length 19.4 mm., greater diameter 12.5 mm., spire height 9.6 mm.
An elongate form (pl. 7, fig. 14a) has the spire ochraceous tawny ;
the impressed sutural and subsutural bands white; last postembryonic
whorl below the white peripheral band pale pinkish buff, banded with
warm blackish brown. Length 20.7 mm., greater diameter 12.5 mm.,
spire height 10.2 mm. The shell may lack a wide tawny band (pl. 7,
fig. 15) and be banded just above the edge of the periphery and all
below with warm blackish brown, above which is a line of ochraceous
tawny, remainder of the shell white. A white color pattern (pl. 7,
fig. 15a) has the postembryonic whorls white, banded and lined with
ochraceous tawny, last fourth whorl banded with black.
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 83
ACHATINELLA APEXFULVA SIMULACRUM var. 1
PLATE 5, FIGURE 26
Area 38: Central Waimano-North Central Waimano Ridge,
locality 212A, elevation 1,200-1,300 feet, 1 live sinistral, Meinecke,
1933; North Central Waimano-North Waimano Ridge, locality
215B, elevation 1,450 feet; 216B, elevation 1,450 feet; 218B, eleva-
tion 1,600-1,650 feet, I live sinistral, all Meinecke-collected, 1933
(figs. 4, 4a, p. 53). Since only two live shells are known from this area
the range of color patterns is not known.
The shells are light color forms very similar to A. a. simulacrum,
but differ in not having any dark-banded color patterns similar to
plate 7, figure 13. A color pattern (pl. 5, fig. 26) common to the two
live shells has the postembryonic whorls white spirally banded with
tawny, base pale pinkish buff tinted with massicot yellow; lip not
formed.
ACHATINELLA APEXFULVA RUBIDILINEA, new subspecies
PLATE I, FIGURE 51; PLATE 7, FicureEs 16-18
The shell of the holotype (pl. 1, fig. 51) has the embryonic whorls
white; postembryonic whorls white profusely lined with chocolate;
impressed sutural band russet ; lip and columella callus light vinaceous
fawn. Length 20.7 mm., greater diameter 14.0 mm., spire height
10.1 mm.
Distribution, area 47: Manana-Waiawa Ridge, type locality 239a,
elevation 2,000-2,050 feet, Meinecke, 1933; also 239b, elevation
2,050-2,250 feet, 6 sinistral ; 239c, elevation 2,100-2,300 feet, I sinis-
tral, Meinecke, 1933; North Central Waimano-North Waimano
Ridge, locality 219B, elevation 1,850-1,900 feet, 8 sinistral, Meinecke,
1933; Waimano-Manana Ridge, locality 220A?, Meinecke, 1929
(figs. 4, 4a, p. 53).
The form of the holotype may not be the usual shape of the shell.
Only seven adult specimens are had from the type locality, so a shell
that appears to be the most usual form and having the characteristic
color pattern is chosen for the holotype. The color pattern on a narrow
shell is lighter (pl. 7, fig. 18), entire shell white; banded on the first
postembryonic whorl just above the periphery with russet, shading
to mars brown, on the penultimate, last whorl lined above the periph-
ery with cinnamon buff, below the periphery with chocolate. Length
20.2 mm., greater diameter 12.8 mm., spire height 10.2 mm.
Each locality on a different ridge has shells with color patterns
showing a slightly different arrangement of bands and lines. The form
84 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
also shows a great range. All the shells of area 47, however, are
reddish-brown lined or banded shells, and the ground color is usually
pure white. The shells are the next race occurring above A. a.
simulacrum.
In locality 219B the usual form of the shell (pl. 7, fig. 16) has the
postembryonic whorls white tinted on the last whorl with pinkish buff,
banded on the last two whorls with chocolate. Length 19.1 mm.,
greater diameter 13.3 mm., spire height 9.5 mm. An obese specimen
(pl. 7, fig. 16a) measures: Length 20.5 mm., greater diameter
14.0 mm., spire height 10.9 mm. It has the lightest color pattern, which
is white lined with chocolate and tawny.
The region of locality 220A ? has dark banded forms such as plate 7,
figures 17 and 17a, which are banded or lined with chocolate, or light
forms which are banded and lined with kaiser brown (pl. 7, figs.
17b, 17c). The extreme obese form (pl. 7, fig. 17) measures: Length
20.6 mm., greater diameter 14.2 mm., spire height 9.7 mm. A narrow
shell, figure 17b, measures: Length I9.1 mm., greater diameter
12.7 mm., spire height 10.2 mm.
ACHATINELLA APEXFULVA CHROMATACME Pilsbry and Cooke
PLATE I, FIGURE 53; PLATE 6, FIGURES 20-23b
Achatinella swiftii chromatacme Pitspry and Cooke, Man. Conch., vol. 22,
p. 316, pl. 59, figs. 5-5b, 1914.
The subspecies A. a. chromatacme is closely related to A. a. per-
plexa, differing mainly in having dark-colored embryonic whorls, tan
or yellowish-brown color patterns that do not occur in area 40, and
the usual form having a wide light-colored band at the edge of the
periphery. In rare cases the embryonic whorls may be white (pl. 6,
fig. 22a), and the shell looks very much like a form of A. a. perplexa.
The shell of plate 59, figure 5, of the Manual of Conchology is
selected by Pilsbry and myself for the lectotype of A. a. chromatacme
and is reproduced in this paper on plate 6, figure 20. The first em-
bryonic whorl of the lectotype is pinkish buff shading on the next
whorl to cinnamon, which lightens on the upper half of the last
embryonic whorl to pinkish buff; first postembryonic whorl, upper
half pinkish buff lined with sayal brown, lower half sayal brown,
penultimate whorl, upper half pinkish buff, lower half ecru drab
axially streaked with rood’s brown, last whorl above the white periph-
eral band vandyke brown darkening to bister, axially streaked with
ecru drab, below the periphery warm sepia axially streaked with
chocolate; impressed sutural band pinkish buff; lip margin army
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 85
brown; columella callus light vinaceous fawn. Length 20.1 mm.,
greater diameter 13.1 mm., spire height I1.1 mm., number of
whorls 64.
Distribution, area 42: Manana-Waiawa Ridge, locality 232, ele-
vation 1,250 feet, 1929, 3 sinistral 1933; 233, elevation 1,350 feet,
1 dextral, 6 sinistral 1929; 234*, elevation 1,150-1,250 feet, 1929;
235*, elevation 1,150-1,250 feet, 1929; 236*, elevation 1,050-1,150
feet, 1929, all collected by Meinecke (figs. 4, 4a, p. 53). Lectotype,
ANSP 108804, collected in Waiawa by Kuhns for Thaanum.
Meinecke did a great deal of collecting with Kuhns and is familiar
with Kuhn’s localities. One or several of the above lots surely repre-
sents a series from the type locality. Meinecke, BBM 121835-121836,
also reports this form from locality 230B ?, elevation 1,000-1,150 feet,
on the North-South Waiawa Ridge, 1929, but I am inclined to believe
that the locality is wrongly plotted because of the wide difference be-
tween the color pattern of the shells of 230B* and the adjacent
localities of 231B* and 232B*.
The exact region of the type locality is not known to me, so I have
selected locality 235 for a typical one, and shall describe the color and
size variation of the shells from that place. In the usual form of the
shell (pl. 6, fig. 21) and a color pattern occurring on 53 percent of
the shells, the embryonic whorls shade from pale pinkish buff to tawny,
last embryonic whorl banded at the suture with white; on the first
postembryonic whorl and a half the impressed sutural band shades
from mikado brown to cinnamon, subsutural band extending to about
the middle of the whorl above the periphery, shades from cinnamon
buff to cinnamon, remainder of the whorl drab axially streaked with
cinnamon, last whorl and a half, impressed sutural and narrow sub-
sutural bands cinnamon, remainder of the whorl pinkish buff closely
axially streaked and finely spirally lined with drab, so that the color
looks at first glance to be solid drab, at the periphery a line and a
band of pale pinkish buff; lip and columella callus light vinaceous
fawn. Length 19.5 mm., greater diameter 13.2 mm., spire height
10.5 mm. A narrow shell (pl. 6, fig. 21a) shows the darkest color
form of the brownish or tan color patterns; the postembryonic
whorls shade from walnut brown to burnt umber axially streaked with
light buff; impressed sutural band light buff. Length 19.7 mm.,
greater diameter 12.7 mm., spire height 11.0 mm.
Forty-seven percent of the live shells in locality 235 have a gray
color pattern (pl. 6, fig. 21b) ; the embryonic whorls shade from pale
pinkish buff to russet; first postembryonic whorl shades from sayal
86 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
brown to verona brown, penultimate whorl benzo brown axially
streaked with pale drab gray, last whorl drab axially streaked with pale
drab gray; impressed sutural and subsutural bands white; lip and
columella callus light vinaceous fawn. Length 19.1 mm., greater di-
meter 13.7 mm., spire height 10.0 mm. The gray pattern may be
lighter (pl. 6, fig. 21c) ; postembryonic whorls white, banded on the
last whorl and a half above the periphery with benzo brown, pale
drab gray, and on the last fourth whorl natal brown, which bands
are axially streaked with white, last whorl below the white peripheral
band natal brown, axially streaked with drab and drab gray, in the
umbilical region a patch of white.
Below locality 235, in localities 232, 233, 234, the brown pattern
is more prevalent, and usually of a lighter color than the usual brown
color pattern in locality 235. Gray color patterns are rare in these
lower localities, occurring on one or two specimens in each lot; dex-
trals are also unusual. In the usual form and color pattern (pl. 1,
fig. 53), the embryonic whorls shade from pale pinkish buff to light
buff to pale pinkish buff; postembryonic whorls shade from light buff
to warm buff, spirally lined with white, banded at the periphery with
a line of snuff brown, and a band of white, base banded with saccardo’s
umber and pale pinkish buff; lip and columella callus light vinaceous
fawn. Length 19.3 mm., greater diameter 12.5 mm., spire height
10.6 mm. The usual obese form (pl. 6, fig. 23) measures: Length
19.7 mm., greater diameter 13.0 mm., spire height 11.1 mm.; post-
embryonic whorls ochraceous tawny, base saccardo’s umber tinted
with ochraceous tawny. An elongate specimen (pl. 6, fig. 23a)
measures: Length 19.4 mm., greater diameter 12.2 mm., spire height
11.5 mm.; postembryonic whorls light ochraceous buff with a central
band of light brownish olive, last whorl banded above and below the
cartridge buff peripheral band with light brownish olive. A rare
purplish pattern (pl. 6, fig. 23b) has the last two whorls dark vina-
ceous brown, axially streaked with white, peripheral band on last
whorl white; sutural and subsutural bands white.
Another common pattern in area 42 is a streaked form (pl. 6,
fig. 22); the postembryonic whorls, below the subsutural and im-
pressed sutural bands of light buff, are snuff brown axially streaked
with light buff, peripheral band on last whorl pale pinkish cinnamon.
The lightest color pattern (pl. 6, fig. 22a), and an unusual specimen
in that the embryonic whorls are pale pinkish buff and not darker,
has the postembryonic whorls white, last whorl lined at the edge of
the periphery with a faint line of sayal brown, banded below the
periphery with hair brown bands which are streaked with white.
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 87
ACHATINELLA APEXFULVA CHROMATACME var. 1
PLATE 1, FIGURE 52; PLATE 6, Figures 24-24f
Achatinella swiftii NeEwcoms, Pilsbry and Cooke, Man. Conch., vol. 22, p. 315,
pl. 50, figs. 4b, 4c (only), 1914. (While these two shells are not typically
A. a. chromatacme var. 1, they are very close to it in color pattern.)
Area 41: Manana-Waiawa Ridge, locality 230*, elevation 1,050-
1,150 feet, 5 sinistral, 1929; 231*, elevation 1,000-1,100 feet ; 230A*,
elevation 1,050-1,100 feet, 3 dextral, 1 sinistral 1929; 231A*, eleva-
tion 750-800 feet, 12 dextral 1929; 232A, elevation 550-600 feet,
I dextral, 2 sinistral 1933, all Meinecke-collected (figs. 4, 4a, p. 53).
The shells of area 41 can easily be confused with the shells of area
40 because the embryonic whorls may be white and the color pattern
similar to A. a. perplexa. They have been considered a variety of
A. a. chromatacme because some of the shells have dark embryonic
whorls and the range of color patterns is closer to A. a. chromatacme
than to A. a. perplexa. The form is characterized by most of the
patterns being spirally lined with white.
The usual form (pl. 1, fig. 52) has the first embryonic whorl pale
pinkish buff, next whorl pinkish buff, last half embryonic whorl
pinkish cinnamon with a central line of cinnamon; first postembryonic
whorl pale pinkish buff lined with cinnamon, penultimate whorl benzo
brown lined with pale pinkish buff, last whorl hair brown lined with
white; impressed sutural band white, tinted pinkish buff; lip and
columella callus light vinaceous fawn. Length 18.7 mm., greater
diameter 12.2 mm., spire height 9.5 mm. The color pattern may be
less lined (pl. 6, fig. 24) and look like A. a. chromatacme, first
embryonic whorl and a half pale pinkish buff shading to tawny on
the last embryonic whorl; postembryonic whorls pinkish buff, first
postembryonic whorl axially streaked with cinnamon drab, and lined
with the ground color, last two whorls lined and streaked with hair
brown.
The dominant color pattern on 25 percent of the shells is a white
lined pattern (pl. 6, fig. 24a) ; the postembryonic whorls white, first
two faintly lined with mikado brown, last whorl a line of cinnamon
above the periphery, at and below the periphery lined with sepia.
The bands and lines may be axially streaked with white (pl. 6,
fig. 24b) ; the shell is entirely white spirally lined with light drab, below
the periphery of the last whorl a band of hair brown.
Lined brownish forms also occur. The lightest brown pattern
(pl. 6, fig. 24c) has the first postembryonic whorl ochraceous tawny,
with a central line of pale pinkish buff, on the penultimate and last
88 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
whorl above the periphery sutural and subsutural bands ochraceous
tawny, below which is a thin band of pale pinkish buff, remainder of
the whorl ochraceous tawny streaked with snuff brown, last whorl
below the periphery pale pinkish buff, spirally lined with snuff
brown. This specimen also shows an obese shell; length 19.1 mm.,
greater diameter 12.8 mm., spire height 10.1 mm. A narrow shell
and dark brown color pattern (pl. 6, fig. 24d) has the penultimate
whorl burnt umber finely lined and streaked with tilleul buff, last
whorl bister axially streaked and spirally lined with snuff brown, and
lined with pale pinkish buff. Length 18.6 mm., greater diameter
11.7 mm., spire height 10.3 mm.
Plate 6, figure 24e is a light gray pattern and obese form of a
dextral shell; the postembryonic whorls pale pinkish cinnamon, the
lower half of the first half postembryonic whorl dotted with cinna-
mon, penultimate whorl banded on the lower half of the whorl with
a band of light drab shading to benzo brown axially streaked with
pale pinkish buff, last whorl light drab gray with a wide sutural and
subsutural band of pale pinkish buff, and a white peripheral band.
Plate 6, figure 24f, is a narrow dextral with a color pattern similar to
figure 24d, only the impressed sutural band is tawny.
ACHATINELLA APEXFULVA CHROMATACME var. 2
PLATE I, FIGURE 54; PLATE 8, FicureEs 16, 16a
Area 43: Manana-Waiawa Ridge, locality 237*, elevation 1,100-
1,200 feet, 1929; 237a, elevation 1,400-1,450 feet, 1933; 237ba, ele-
vation 1,300-1,400 feet, 1933, all Meinecke-collected (figs. 4, 4a,
Pp. 53).
This race is of interest because it occurs between chromatacme
and nigripicta. The shells are definitely more lined and have more
white color pattern than chromatacme and one would suppose they
were intermediate between a white-lined race and A. a. chromatacme.
But above area 43 the shélls have mostly a yellowish ground and are
banded with warm blackish brown or liver brown. The color patterns
in area 43 are so close to A. a. chromatacme that I think it inadvisable
to separate them, although the embryonic whorls are usually light
colored similar to A. a. perplexa.
In the usual form (pl. 8, fig. 16) the embryonic whorls shade from
pale pinkish buff to light buff ; postembryonic whorls pale pinkish buff
spirally lined and axially streaked with light buff, last whorl and a
half just above and all below the periphery warm sepia axially
streaked and spirally lined with warm buff; lip and columella cal-
lus light brownish vinaceous. Length 19.1 mm., greater diameter
12.8 mm., spire height 10.4 mm.
NO. I ACHATINELLA APEXFULVA DIXON—-WELCH 89
The usual color pattern and obese form of the shell is shown on
plate 1, figure 54, postembryonic whorls white; impressed sutural
band on the first postembryonic whorl light pinkish cinnamon, on the
remaining whorls the pinkish cinnamon color is confined to the upper
edge of the band and fades out on the last whorl; last whorl banded
above the periphery with hair brown, below the periphery hair brown _
streaked with white and lined with natal brown. Length 17.6 mm.,
greater diameter 13.7 mm., spire height 8.6 mm. The embryonic
whorls may be darker and the postembryonic whorls lighter (pl. 8,
fig. 16a) ; embryonic whorls pale pinkish buff shading to ochraceous
buff, banded at the suture on the last embryonic whorl with sayal
brown; first postembryonic whorl snuff brown lined with white,
penultimate whorl white, banded and lined with sayal brown, last
whorl white, lined with sayal brown and banded below the periphery
with warm sepia; impressed sutural band verona brown fading out to
sayal brown on the last half whorl where it occupies only the upper
edge of the sutural band.
ACHATINELLA APEXFULVA ALBIPRAETEXTA, new subspecies
PLATE 2, FIGURE 3; PLATE 8, FicurREs 18-21
The shell is closely related to A. a. chromatacme, but differs in the
gray banding of the postembryonic whorls, and by having white em-
bryonic whorls. The embryonic whorls of the holotype (pl. 2, fig. 3)
are white; first postembryonic whorl white, lined with russet; on the
last two whorls the impressed sutural and wide subsutural bands
are white, except on the last half whorl, where the lower edge of the
sutural band is edged with a line of wood brown; the remainder of
the shell shades from brownish drab on the first half of the penulti-
mate to drab gray axially darkened by streaks of light drab on the
remaining whorls; lip vinaceous fawn. Length 18.0 mm., greater di-
ameter 12.5 mm., spire height 9.3 mm., number of whorls 63.
Distribution, area 63?: North-South Waiawa Ridge, type locality
231B*, elevation 1,000-1,100 feet, 1929; also locality 232B*, eleva-
tion 750-850 feet, Meinecke, 1929. The Meinecke localities were
plotted in 1933 and may be plotted too low and should be checked.
Also collected in Waiawa by Thurston, BBM 130590.
Area 64: South Waiawa Stream, locality 237A-1*, elevation 750-
850 feet, 3 sinistral, H. Lemke, 1932; on a spur ridge of the North-
South Waiawa Ridge, locality 233B-1*, elevation 1,350-1,400 feet,
H. Lemke and H. Lemke, Jr., 1932. Area 63? is possibly wrongly
plotted and perhaps should be placed higher or nearer area 64
(figs. 4, 4a, p. 53).
go SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
The narrow form of the shell (pl. 8, fig. 19) measures: Length
18.4 mm., greater diameter 10.8 mm., spire height 10.2 mm.; the
color pattern is similar to plate 2, figure 3. An obese shell (pl. 8,
fig. 20) from locality 232B exhibits the tan color pattern, which
occurs on 22 percent of the shells in the type locality ; the embryonic
whorls are white; postembryonic whorls ochraceous buff, last half
of the penultimate and last whorl just above the periphery and be-
low the periphery mouse gray axially streaked with ochraceous buff ;
impressed sutural band the color of the ground except on the first post-
embryonic whorl, where it is tawny. Length 19.0 mm., greater di-
ameter I3.0 mm., spire height 10.1 mm. Two specimens of this tan
color pattern were found to have the entire sutural band russet.
A dextral shell from the Thurston collection is shown on plate 8,
figure 21. The color pattern is similar to plate 2, figure 3.
In locality 233B-1*, 48 percent of the 13 dextral shells have a gray
pattern similar to typical A. a. albipraetexta, except that the post-
embryonic whorls are not banded with gray. One of these (pl. 8,
fig. 18) has the embryonic whorls cartridge buff; first postembryonic
whorl cream buff shading to white, lined on the first half of the whorl
with tawny, the periphery faintly spirally banded or tinted with pale
drab gray, below the periphery the color is drab or verona brown
axially streaked with white, behind the edge of the lip is a band
from 2 to 3 mm. wide of warm sepia. Plate 8, figure 18a, shows the
tan color pattern that occurs on 52 percent of the dextral specimens,
a color pattern similar to that of plate 8, figure 20.
Thirty-two percent of the shells in 233B-1* are sinistral and differ
in the range of color patterns from the dextral shells in having lined
color patterns. The postembryonic whorls of the gray pattern (pl. 8,
fig. 18b) are white spirally lined or tinted on the last two whorls with
pale drab gray, just above or at the periphery a line of snuff brown,
last whorl below the periphery spirally banded with pale drab gray,
in the umbilical region banded with drab, pale pinkish buff, and
cinnamon. The tan pattern (pl. 8, fig. 18c) has the postembryonic
whorls white or pale pinkish buff, finely axially streaked or tinted
with pinkish buff, last whorl banded at the edge of the periphery
with white, below the periphery, spirally lined and axially streaked
with a light shade of sayal brown.
ACHATINELLA APEXFULVA GRISEIBASIS, new subspecies
PLATE 2, Ficure 6; Pate 8, Ficures 17, 17a
The shell is close to A. a. lemkei but differs in color pattern. The
embryonic whorls of the holotype (pl. 2, fig. 6) are white except for
NO. I ACHATINELLA APEXFULVA DIXON—WELCH gl
a line of chamois on the lower portion of the whorl just above the
periphery. This can best be seen under a low-power microscope.
First postembryonic whorl white, banded on the upper half of the
whorl with tawny, remaining whorls white, last whorl below the
periphery benzo brown axially streaked with pale drab gray; lip and
columella callus vinaceous buff. Length 17.5 mm., greater diameter
12.I mm., spire height 9.2 mm., number of whorls 6.
Distribution, area 65??: East Waiawa, “Ridge W. of Engineer’s
camp, + mi. mauka [toward the mountains] of the ditch trail from
large koa tree down toward camp, on koa [Acacia koa], pua
[ Osmanthus], guava [Psidium guajava],’ O. H. Emerson, BBM
103967, 1915; also J. S. Emerson, BBM 102248, 1915. The exact
location is not known. The shells were surely found at a low eleva-
tion, because they were collected on trees that occur at a low eleva-
tion. Area 65?? may possibly be the approximate region of the type
locality (fig. 4a, p. 53).
A variation of the typical pattern (pl. 8, fig. 17) is similar to
A. a. chromatacme var. 2 (pl. 8, fig. 16a); first embryonic whorl
white, tinted with army brown, next whorl white, faintly banded on
the lower half of the whorl with cream buff; postembryonic whorls
white, first half postembryonic whorl banded and streaked with
ochraceous tawny, last whorl below the periphery banded with
verona brown or snuff brown. The usual narrow form (pl. 8,
fig. 17a) measures: Length 17.6 mm., greater diameter 11.4 mm.,
spire height 9.5 mm.; embryonic whorls white; first half postem-
bryonic whorl tawny, finely lined with white, last half tawny, banded
on the lower two-thirds of the whorl with sorghum brown, penulti-
mate whorl white, banded on the first half of the whorl with sorghum
brown, last half with only a line of sorghum brown lightening to
sayal brown, last whorl above the periphery white, below the periph-
ery verona brown, finely lined and axially streaked with pale pinkish
cinnamon.
GrouP oF A. A. POLYMORPHA GULICK
ACHATINELLA APEXFULVA POLYMORPHA Gulick
Pate 8, Figures 24-28a
Apex polymorpha Gutick, Proc. Zool. Soc. London, 1873, p. 81, pl. 10, fig. 5.
Apex neglectus SmitH, Proc. Zool. Soc. London, 1873, p. 78, pl. 9, fig. 22.
Achatinella swiftii NEwcoms, Pilsbry and Cooke, Man. Conch., vol. 22, pp. 306,
307, 310, 315, pl. 50, figs. 4, 4a, 4d, 6 (only), 1914.
The holotype (pl. 8, fig. 27) has the first embryonic whorl white,
remaining two and a half embryonic whorls upper half white, lower
g2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
half chamois; first postembryonic whorl light buff, with a central
line of cinnamon buff, on the penultimate whorl, the subsutural band
light pinkish cinnamon, remainder of the whorl light brownish drab,
faintly spirally lined and axially streaked with pale pinkish cinnamon,
last whorl subsutural band avellaneous, below which is a line of
cartridge buff, supraperipheral band pale pinkish buff axially streaked
with brownish drab, basal band same color but also spirally lined with
brownish drab, peripheral band cartridge buff; lip cinnamon shading
to white within ; columella callus white ; impressed sutural band shades
from cameo brown on the first two whorls to wood brown on the first
half of the last whorl to cinnamon on the last half whorl. Length
16.8 mm., greater diameter 12.3 mm., spire height 8.4 mm., number
of whorls 6.
Distribution, area 66°?: Waipio, Gulick. Gulick also reports the
subspecies from Ahonui, Kalaikoa, Wahiawa. The district of Waipio
includes Waikakalaua and Kipapa Gulches. The holotype of A. a.
polymorpha surely did not come from north of Kipapa Gulch as
Gulick reports, because the range of color patterns of one race does
not extend over several gulches, although the same pattern may be
repeated in widely separated areas. The holotype probably came
from lower Kipapa or Panihakea Gulch in the district of Waiawa be-
cause it is similar to the shells existing in that region today. I have
tentatively plotted the possible region of the type locality, area 66??,
on fig. 5a, p. 105. Holotype, MCZ 39901.
A specimen (pl. 8, fig. 28) with a color pattern similar to the holo-
type has the embryonic whorls pale pinkish buff, banded on the lower
half of the last embryonic whorl with light buff, on the first embryonic
whorl at the edge of the suture a line of benzo brown; postembryonic
whorls pale pinkish buff, first postembryonic whorls spirally lined
with wood brown, last two whorls banded with drab, the bands lined
and streaked with cinnamon drab and benzo brown; impressed sutural
band russet shading to tawny on the last whorl; lip, outer margin
cinnamon buff, inner margin and columella callus white. Length
16.4 mm., greater diameter 11.7 mm., spire height 8.8 mm.
The color pattern on a sinistral shell (pl. 8, fig. 28a) is much lighter
and shows an elongate specimen; the first embryonic whorl and a half
bicolored, upper half white, lower half hair brown, remaining em-
bryonic whorls white, banded on the lower fourth of the whorl with a
faint shade of cream buff; postembryonic whorls avellaneous spirally
lined, banded, and streaked with white; impressed sutural band snuff
brown on the first whorl lightening to light pinkish cinnamon on the
last two whorls; lip not formed.
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 93
In the small series of shells in the Gulick collection in the Bishop
Museum that can safely be identified as A. a. polymorpha, the typical
pattern is a rare one and the range of color patterns is not determined.
Usually, in Waiawa where forms similar or closely related to
A. a. polymorpha occur, such as A. a. polymorpha var. 1, A. a.
flavitincta, and A. a. lemkei var. 1, the sutural band is tan or brown.
Exceptions to this rule are the shells of plate 8, figures 22b and 22c,
of A. a. lemkei. But these two shells may belong to a race occurring
at a higher elevation than A. a. lemkei. Since rare tan patterns occur
with the usual banded patterns in area 68 and 69, which are very
similar to the color patterns of the shell described as Apex neglectus
by Smith, I believe that neglectus is a rare light tan color form of
A. a. polymorpha.
There are four specimens of Apex neglecta in the type lot in the
British Museum. One specimen marked “x” by myself is considered
the lectotype, because it agrees best with Smith’s original figure re-
produced in this paper on plate 8, figure 24. The lectotype (pl. 8,
fig. 24) has the embryonic whorls cartridge buff banded on the last
embryonic whorl on the lower half of the whorl with cream buff;
first two postembryonic whorls finely lined with cinnamon buff, last
whorl pale olive gray axially streaked and faintly lined with light
olive gray; impressed sutural band shades from hazel to carob brown
on the last two whorls; subsutural band on the last whorl cinnamon
buff; lip and columella callus vinaceous fawn. Length 17.2 mm.,
greater diameter 12.0 mm., number of whorls 6. Another shell from
the type lot (pl. 8, fig. 25) has a convex spire, and a color pattern
similar to the lectotype.
A dextral specimen in the Gulick collection (pl. 8, fig. 26) has the
embryonic whorls white; postembryonic whorls pale pinkish buff
spirally banded and lined with cinnamon or cinnamon buff above the
periphery ; the impressed sutural band mikado brown.
ACHATINELLA APEXFULVA POLYMORPHA var, 1
Pirate 8, FIGURES 29, 29a
Area?: Waiawa, J. S. Emerson, BBM 102247, BBM 33312.
The usual color pattern of the shell (pl. 8, fig. 29) resembles
A. a. polymorpha except that the banding is a lighter shade of gray,
and the base is white; the shape of the last whorl is rounded and the
base is not flattened. Upper half of the first embryonic whorl white,
lower half cream color, remaining embryonic whorls white, banded on
the lower fourth of the whorl with warm buff; first half postembry-
onic whorl white, last half pinkish buff, first half of the penultimate
94 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
whorl pinkish buff spirally lined with light drab, last half pale mouse
gray, lined and streaked with light buff, last whorl below the im-
pressed sutural band light buff, banded or lined with drab and
cinnamon buff, just above and below the white peripheral band are
two bands of smoke gray, which shade to wood brown on the last
half whorl, base white faintly lined with smoke gray; impressed
sutural band amber brown shading to tawny on the last whorl; lip
and columella callus pale vinaceous pink. Length 18.0 mm., greater
diameter 12.2 mm., spire height 10.0 mm., number of whorls 6.
Five sinistrals are found in lot BBM 102247 of 23 shells. The
color pattern on a few shells differs from the usual one. The post-
embryonic whorls may be white or pale pinkish buff (pl. 8, fig. 29a),
lined on the first and penultimate whorls with fawn color or wood
brown, last whorl banded at the edge of the periphery with fawn
color, banded below the periphery with benzo brown axially streaked
with pale pinkish buff.
ACHATINELLA APEXFULVA FLAVITINCTA, new subspecies
PLATE 2, FicurE 7; PLATE 8, FIGURES 30-30)
The form is closely related to A. a. polymorpha but differs in having
the last whorl lined with dark lines of reddish brown. Plate 8,
figure 30a, looks like Smith’s Apex neglectus (pl. 8, fig. 24), except
for the embryonic whorls and the lined last half postembryonic
whorl.
The holotype (pl. 2, fig. 7) has the embryonic whorls bicolored, first
embryonic whorl upper half white, lower half walnut brown, remain-
ing embryonic whorls white ; first postembryonic whorl white, penulti-
mate whorl white with a faint line of light olive gray on the lower
half of the whorl, last whorl above the periphery white, finely lined,
axially streaked or tinted with chamois and banded above the edge of
the periphery with olive buff, below the periphery banded or tinted
with pallid mouse gray and lined with line and a band of benzo
brown or fuscous; impressed sutural band army brown on the first
whorl, on the penultimate avellaneous, on the last whorl ochraceous
tawny ; lip light vinaceous fawn. Length 17.2 mm., greater diameter
II.3 mm., spire height 9.5 mm., number of whorls 6.
Distribution, area 69: Waiawa-Panihakea Ridge, type locality
230G*, elevation 1,200-1,250 feet, 1932; also 230Ga*, elevation
1,300 feet, 2 dextral, I sinistral, 1932; 231G, elevation 1,350-1,400
feet, 1932, 5 dextral 1933; 232G*, elevation 1,300-1,400 feet, 1932,
all collected by Meinecke (figs. 4, 4a, p. 53).
The elongate form (pl. 8, fig. 30) and the usual lined color pattern
which occurs on 28 percent of the shells measures: Length 17.3 mm.,
NO: I ACHATINELLA APEXFULVA DIXON—WELCH 9
un
greater diameter 11.4 mm., spire height 9.6 mm., number of whorls 6;
postembryonic whorls white, penultimate whorl banded above the
periphery with light mouse gray, last whorl above the periphery light
buff faintly lined with white, and a line of sayal brown on the first
half of the whorl, 5 mm. behind the edge of the lip a streak or
tinting of tawny down to the supraperipheral band of light grayish
olive, last whorl banded at the periphery with white, below the periph-
ery ground white colored with a band of light grayish olive, which is
lined with two lines of fuscous ; impressed sutural band changes from
russet to avellaneous.
The obese form of the shell (pl. 8, fig. 30a) and a yellowish or light
variant of the typical color pattern has the first postembryonic whorl
and a half white, last postembryonic whorl and a half warm buff
axially streaked and spirally lined with white below the periphery the
ground is white or pale gull gray slightly tinted above the aperture, and
at the beginning of the last half of the whorl with spots of warm buff,
lined with warm sepia, on the last half of the whorl ; impressed sutural
band russet lightening to cinnamon buff on the last whorl. Length
16.2 mm., greater diameter 11.5 mm., spire height 8.1 mm.
The usual form of a sinistral (pl. 8, fig. 30b) and darker lined
variation of the lined pattern measures: Length 17.8 mm., greater
diameter 11.2 mm., spire height 10.1 mm.; postembryonic whorls
cartridge buff, spirally lined and axially streaked on the first post-
embryonic whorl with cinnamon buff, penultimate and last whorls
lined with bone brown, at the periphery banded with white, below the
periphery the ground is white with a central band of bone brown
which divides into three on the last half of the whorl; impressed
sutural band tawny on the first whorl, on the remaining whorls cinna-
mon buff.
ACHATINELLA APEXFULVA LEMKEI, new subspecies
PLATE 2, Figure 5; PLATE 8, FIGURES 22-22c
The color pattern resembles A. a. leucorraphe, but differs in not
having a white sutural band, and in having the spire tinted with
yellow ; first half embryonic whorl of the holotype (pl. 2, fig. 5) black,
next half of whorl upper half white, lower half black, last two em-
bryonic whorls upper half white, lower half chamois; first post-
embryonic whorl light buff, spirally lined with white, first half of the
penultimate light buff axially streaked with cinnamon drab; the im-
pressed sutural band on the first postembryonic whorl and a half
tawny, on the last whorl and a half the sutural and subsutural
7
96 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
bands light buff, the remainder of the whorl hair brown, chaetura |
drab, or warm sepia, axially broken by straight and zigzag streaks of
pale pinkish buff or white, edge of the periphery banded with white;
lip and columella callus vinaceous fawn. Length 17.2 mm., greater
diameter 12.0 mm., spire height 9.2 mm., number of whorls 6}.
Distribution, area 67?: North Waiawa Stream, locality 230E-1-2?,
approximate elevation 1,050-1,900 feet, “ridge back of tunnel-man’s
house,” found on lehua (Metrosideros), Lemke, 1933, also 1932. The
exact location of this race must be checked. Different lots from the
general region of area 67? contain varying proportions of different
color patterns which indicates slightly different localities (figs. 4, 4a,
p- 53).
The darkest color pattern (pl. 8, fig. 22) has the first postembryonic
whorl light buff finely axially streaked or tinted with ochraceous
tawny, impressed sutural band ochraceous tawny, last two whorls
bone brown, streaked with zigzag lines of light buff, sutural and
subsutural bands light buff axially streaked with ochraceous tawny
on the penultimate whorl, on the last whorl light buff, last whorl
banded at the edge of the periphery with white; lip and columella
callus light vinaceous fawn.
The usual color pattern of a dextral shell (pl. 8, fig. 22a) has the
first postembryonic whorl white, tinted with pinkish buff, sutural
and subsutural bands ochraceous tawny, penultimate whorl between
white and pale pinkish buff tinted and streaked with cinnamon buff,
sutural and subsutural bands cinnamon buff, just above the edge of
the periphery a band of benzo brown, streaked with zigzag lines of
white, last whorl above the white peripheral band pale cinnamon pink
tinted with pinkish buff, supraperipheral band and all below the
periphery natal brown spirally lined and streaked with zigzag axial
lines of pallid mouse gray and white.
I have a number of lots before me collected by Lemke at different
times from the region of the type locality. Lot 110, Lemke collection,
has seven specimens, all except one with the typical pattern of
plate 2, figure 5. In connection with lot 173, Lemke collection, Lemke
states that he collected over a wider range, going up a spur ridge to
the top of a larger ridge from which the spur branched. In this lot
45 percent of the shells have the typical patterns of A. a. lemkei;
the remainder have white, gray-lined color patterns which possibly
come from a locality at a higher elevation (see pl. 8, figs. 22b, 22c).
Although I am calling these two shells A. a. lemketi, they quite pos-
sibly belong to another race occurring above lemkei, the two forms
not being mixed except at the border of an upper and lower area.
nO. t ACHATINELLA APEXFULVA DIXON—WELCH 97
The shell of plate 8, figure 22), is white, last two postembryonic
whorls tinted with cinnamon buff and spirally banded with pale drab
gray deepening on the last half whorl to light drab and hair brown,
the bands axially broken with white. The spire may be almost en-
tirely white (pl. 8, fig. 22c), banded with drab below the periphery,
which bands are broken by axial streaks of white, the last whorl
above the periphery has a few faint lines of cinnamon buff.
ACHATINELLA APEXFULVA LEMKEI var, 1
PLATE 8, FIGURES 23-230
Area 68?: North Waiawa Stream, general region of locality
233Ga?, approximate elevation 1,300 feet, Lemke, 1932, on lehua
(Metrosideros), pua (Osmanthus), alani (Pelea) (figs. 4, 4a, p. 53).
The locality is a-wide one and extends up to locality 234G on the
Waiawa-Panihakea Ridge. The lots represent a mixture of two
races, A. a. lemkei var. 1 and A. a. lineipicta. Color patterns similar
to those found in locality 234G (A. a. linetpicta) are found mixed
with color patterns not occurring in locality 234G. For discussion I
shall take lot 113, Lemke collection, and describe only the unusual
patterns which markedly differ from locality 234G, and occur on
48 percent of the shells.
The usual form (pl. 8, fig. 23) and color pattern of the shell has
the embryonic whorls bicolored, first half embryonic whorl natal
brown, shading to white on the upper fourth of the whorl, next half
whorl upper half white, lower half pale drab gray, remaining em-
bryonic whorls white banded with honey yellow just above the
edge of the periphery; first postembryonic whorl light buff spirally
lined with tawny; impressed sutural band tawny; remaining post-
embryonic whorls light buff finely lined and axially streaked with
tawny, banded on the lower edge of the penultimate whorl with
bone brown, last whorl below the periphery light buff, closely axially
streaked, spirally lined and banded with bone brown; impressed sutural
band on last two whorls light buff; lip and columella callus vinaceous
fawn. Length 17.5 mm., greater diameter 12.1 mm., spire height
8.9 mm., number of whorls 6.
The color pattern may be darker (pl. 8, fig. 23a), the first half
of the penultimate whorl russet axially streaked with tawny, last
whorl and a half light buff almost entirely covered over by spiral
lines and axial streaks of fuscous or carob brown; impressed sutural
band light buff lined with tawny. An obese shell (pl. 8, fig. 23b),
length 18.7 mm., greater diameter 13.5 mm., spire height 8.9 mm.,
98 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
number of whorls 6}, has the postembryonic whorls light buff axially
streaked with cinnamon buff; impressed sutural band on the first
postembryonic whorl and a half tawny, on the remaining whorls
the sutural band has the ground color; just below the periphery of
the last whorl a band of white, the remainder of the base warm sepia
axially streaked with zigzag lines of light buff, the edge of the
periphery and just above banded with a band of warm sepia broken
by the ground color. This color pattern is very similar to the typical
lemkei pattern (pl. 2, fig. 5) except that the spire is yellow and not
banded or streaked with gray on the first half of the penultimate
whorl.
Although A. a. lemkei and its variety are not well localized, as far
as the material at hand is concerned, they are of great importance to
the addition of the knowledge of shell variation. Not only is there
a possibility of finding new color patterns and races at different ele-
vations on the long major ridges of the Koolau Range, but also
on short spur ridges local races may be found at different elevations.
ACHATINELLA APEXFULVA LINEIPICTA, new subspecies
PLATE 2, Ficure 8; PLATE 8, FIGURES I-3a
Achatinella swiftii Newcoms, Pilsbry and Cooke, Man. Conch., vol. 22, p. 314,
pl. 58, figs. 5-5b (only), 1914. Figure 5) matches A. a. Jemkei var. 1 (pl. 8,
fig. 23a) of area 68 which is an intermediate area between areas 67 and 70.
The shell is an intermediate lined form existing between the lower
race of A. a. flavitincta (area 69) and the upper race of A. a. ewaensis
var. 3 (area 52). The first embryonic whorl of the holotype (pl. 2,
fig. 8) fawn color, remaining embryonic whorls upper third white,
lower two-thirds cream buff darkening to chamois ; first postembryonic
whorl upper third banded with white, lower portion cinnamon buff
on the first half of the whorl, last half pinkish buff lined with verona
brown, last two whorls light buff above the periphery, lined with
seal brown, last whorl below the periphery pale gull gray spirally
banded with seal brown; impressed sutural band on first postembry-
onic whorl russet, on the remaining whorls light buff with a line of
tawny at the lower edge of the band; lip and columella callus
vinaceous fawn. Length 17.6 mm., greater diameter 12.3 mm., spire |
height 9.0 mm., number of whorls 64.
Distribution, area 70: Waiawa-Panihakea Ridge, type locality
232Ga, elevation 1,350-1,450 feet, 1933; 233G*, elevation 1,350-
1,450 feet, 1932; 234G, elevation 1,400-1,500 feet, 1933. All Mein-
ecke-collected (figs. 4, 4a, p. 53).
NO: I ACHATINELLA’ APEXFULVA DIXON—WELCH 99
The shell may be more obese and very lined (pl. 8, fig. 1), the
first half embryonic whorl fuscous, next whorl upper half white, lower
half light grayish olive fading to white on the last embryonic whorl
and a half; the first postembryonic whorl and a half white, banded
with mikado brown and lined with cinnamon buff, last postembryonic
whorl and a half pale gull gray lined with black, above the periphery
the ground is lined with cinnamon buff and tinted with pinkish buff ;
impressed sutural band russet up to the last whorl, where it fades to
pinkish buff. Length 17.8 mm., greater diameter 12.8 mm., spire
height 9.1 mm.
The spire may not be banded (pl. 8, fig. 1@), and the shell may be
narrow, postembryonic whorls white shading to pale gull gray on the
last two whorls, last whorl has a peripheral band of white, banded
and lined below the periphery with fuscous. Length 17.4 mm., greater
diameter II.0 mm., spire height 9.9 mm.
Each locality in area 71 appears to have similar patterns to the
type locality but the form and color pattern varies. A common form
and color pattern (pl. 8, fig. 2), in locality 233G* measures: Length
18.4 mm., greater diameter 13.1 mm., spire height 9.2 mm.; first
postembryonic whorl light buff, axially streaked with light buff,
penultimate whorl pale pinkish buff shading to pale gull gray on the
last whorl, last two whorls profusely banded and lined with seal
brown.
In locality 234G the usual form and color pattern (pl. 8, fig. 3)
measures: Length 17.8 mm., greater diameter 12.6 mm., spire height
9.9 mm. ; postembryonic whorls white darkening to pale gull gray, last
whorl spirally lined with seal brown or black, the lines above the
periphery broken by axial streaks of pale gull gray and the ground
tinted with light buff; impressed sutural band ochraceous tawny be-
coming a dilute shade of ochraceous tawny on the last whorl. In this
locality a dark color pattern (pl. 8, fig. 3a) is found on a single
specimen; first postembryonic whorl and a half light buff axially
streaked with chamois and lined with seal brown on the latter half
of the whorl, last whorl and a half seal brown, spirally banded and
lined with light buff or white.
ACHATINELLA APEXFULVA LINEIPICTA var. 1
PLATE 2, FicuRE 10; PLaTe 8, FiGuRES 4, 5
Area 72: Panihakea-Kipapa Ridge, locality 242-2*, elevation
I,350-1,400 feet, 13 sinistral; 242-3*, elevation 1,500-1,550 feet, 7
dextral, Russ and Welch, 1934, region of the upper portion of area
I0O0 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
71 and area 72, or locality 241-1-242-3?, Russ, 1931, BBM 134151
(figs. 4, 4a, p. 53). .
Area 72 is not very accurately plotted on a map. All of Welch’s
material consists of dead shells. The Russ material covers a con-
siderable area so that areas 72 and 71 overlap. However, from
the material on hand the shells appear to be intermediate between
the forms found in area 71 and 70. The usual pattern (pl. 2, fig. 10)
has the embryonic whorls similar to A. a. lineipicta; the first post-
embryonic whorl white, penultimate whorl white deepening in color to
pale pinkish buff on the last whorl, last two whorls lined above the
periphery with hair brown and fuscous, last whorl at and below
the periphery pale gull gray spirally lined and banded with olive brown
and mouse gray; impressed sutural band russet becoming dilute in
color on the last half whorl.
A few specimens have a gray color pattern (pl. 8, fig. 4), last two
postembryonic whorls pale gull gray finely axially streaked and
spirally lined with light gull gray. A single dark specimen (pl. 8,
fig. 5) from locality 242-2*, is white on the first postembryonic whorl,
lightly axially streaked with raw sienna, on the penultimate whorl the
subsutural band is warm buff axially streaked with sudan brown,
the remainder of the whorl seal brown lined with warm buff, last
whorl seal brown or black, subsutural band warm buff, lined with
russet; impressed sutural band russet; lip light vinaceous fawn.
ACHATINELLA APEXFULVA FUMOSITINCTA, new subspecies
PLATE 2, FigurE 9; PLaTe 8, FIGURES 31, 31a; PLATE 9, FIGURE I
Achatinella swiftii NeEwcomp, Pilsbry and Cooke, Man. Conch., vol. 22, p. 314,
pl. 58, figs. 4, 4a (only), 1914.
The shell has a-color pattern similar to A. a. polymorpha var. 1
(pl. 8, fig. 29) only it lacks the characteristic white peripheral bands
on the last whorl, differs in shape and has a different series of color
patterns which usually have a ground of smoke gray. The first em-
bryonic whorl of the holotype (pl. 2, fig. 9) cream buff, remaining
embryonic whorls cartridge buff, banded on the lower half of the
whorl with chamois; first postembryonic whorl white, banded just
above the periphery with tawny; impressed sutural and subsutural
bands light vinaceous drab; penultimate and last whorls light grayish
olive, tinted by fine axial streaks of smoke gray, last whorl banded
at the periphery and about the base with smoke gray; impressed
sutural band on the penultimate drab gray shading to avellaneous on
the last whorl ; lip and columella callus pale grayish vinaceous. Length
NO. I ACHATINELLA APEXFULVA DIXON—WELCH IOI
17.5 mm., greater diameter 12.5 mm., spire height 9.2 mm., number
of whorls 64.
Distribution, area 71: Panihakea-Kipapa Ridge, type locality
240*, elevation 1,300-1,350 feet, Meinecke, 1929; also locality 241-
ta*, elevation 1,350-1,400 feet, 5 sinistral; 241-1*, elevation 1,250-
1,450 feet; 241-2?, elevation 1,250-1,300 feet; 241-3*, elevation
1,300-1,350 feet, Welch and Russ, 1934; 240-1-241I-1a?, elevation
1,400-1,450 feet, Russ; 241-1?, Russ, 1934; all localities approxi-
mately correct. Also collected by Cooke, 1911, BBM 22826-22828,
22832 (figs. 4, 4a, p. 53).
The typical pattern occurs on 78 percent of the shells in the type
locality. One light gray shell (pl. 8, fig. 31) was found in the lot
which also shows a narrow specimen. Length 16.6 mm., greater di-
ameter II.2 mm., spire height 8.5 mm. The first postembryonic whorl
and a half are white, last whorl and a half pale mouse gray darkening
to drab gray on the last half whorl, tinted above the periphery with
pinkish buff, banded with white and faintly lined with benzo brown.
Two specimens had a chocolate-lined pattern (pl. 8, fig. 31a) ; post-
embryonic whorls pale smoke gray spirally lined, banded, and finely
axially streaked with chocolate; impressed sutural band chestnut.
The typical pattern (pl. 9, fig. 1) may be lined and may lack the
pronounced peripheral light gray band on the last whorl, last half of
first whorl and first half of penultimate whorl smoke gray spirally
lined and axially streaked with chocolate, last whorl and a half smoke
gray spirally lined with light grayish olive, last half whorl streaked
with fuscous, last whorl streaked with pale smoke gray.
Grourp or A. A. LEUCORRAPHE GULICK
ACHATINELLA APEXFULVA LEUCORRAPHE Gulick
PLATE 9, FIGURES 2-4
Achatinella leucorraphe GuticK, Proc. Zool. Soc. London, 1873, p. 79, pl. 10,
fig. 2—Pitspry and Cooke, Man. Conch., vol. 22, p. 301, pl. 59, fig. 8
(only), 1914.
The holotype (pl. 9, fig. 2) has the embryonic whorls bicolored,
first two embryonic whorls and a half, upper half white, lower half
hair brown or fuscous black, last embryonic whorl and a half white;
first postembryonic whorl pale drab gray axially streaked with benzo
brown, penultimate whorl pale drab gray axially streaked with zigzag
streaks of fuscous, first half of last whorl white axially streaked
with fuscous, and three streaks of dark vinaceous drab above the
columella, last half of the whorl drab gray axially streaked with hair
102 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
brown; impressed sutural band white; lip and columella callus white.
Length 19.0 mm., greater diameter 12.6 mm., spire height 10.2 mm.,
number of whorls 64.
Distribution, area?: Kalaikoa, Gulick. Holotype, MCZ 39903.
South Kaukonahua, South-Central Branch Ridge, general region
of locality 260A ?, elevation? (fig. 5, p. 105), probably the lower por-
tion of the locality, Meinecke, 2 sinistral 1928. Found mixed with
A. a. tuberans var. 4, BBM 121952. Also collected by Wilder, South
Kaukonahua Stream, BBM 50573, 1 dextral, found mixed with
A. a. virgatifulva, BBM 50572, and probably comes from the region
of Waikakalaua Stream.
One of the Meinecke shells (pl. 9, fig. 3) has the first embryonic
whorl slightly worn and colored army brown, remaining embryonic
whorls upper two-thirds white, lower third benzo brown, the benzo
brown fades out on the last half embryonic whorl; first postembryonic
whorl white, axially streaked with light drab, penultimate whorl
pale gull gray axially streaked with fuscous, last whorl spirally lined
and axially streaked with seal brown and fuscous, ground color pale
mouse gray, and sometimes appears as zigzag streaks of pale mouse
gray between the darker axial coloration, last half whorl drab,
axially streaked with seal brown and fuscous; lip and columella
callus a dilute shade of pale vinaceous fawn; impressed sutural band
white, lined on the last whorl with a line of fuscous on the lower por-
tion of the band; the shell is not quite adult. Length 17.5 mm., greater
diameter 12.1 mm., spire height 9.7 mm., number of whorls 64.
The other Meinecke shell (pl. 9, fig. 3a) has a pattern similar to
that of A. a. leucorraphe on the first half of the penultimate whorl,
while the remaining whorls look like some form of A. a. tuberans.
This shell (pl. 9, fig. 3¢) has the first half postembryonic whorl white
with a line of fawn color on the lower fourth of the whorl broken by
axial streaks of white, next three-fourths of the whorl pale gull gray
axially streaked with natal brown, next half whorl pale gull gray
lined with natal brown, the lines broken by the ground color, last
whorl darkening to drab on the last half, spirally lined and axially
streaked with white or pale gull gray.
The Wilder specimen (pl. 9, fig. 4) differs from typical leucorraphe
by having the embryonic whorls banded with yellow. The first em-
bryonic whorl natal brown, next whorl upper fourth white, lower
fourth natal brown shading to drab, last embryonic whorl shading
from drab to honey yellow; first two postembryonic whorls and a
fourth white, tinted on the penultimate with pale gull gray, last
whorl pale gull gray axially streaked with benzo brown; lip light buff,
columella callus white.
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 103
ACHATINELLA APEXFULVA VIRGATIFULVA, new subspecies
PLATE 2, FIGURE 16; PLATE 9, FIGURES 5-5¢
Achatinella leucorraphe irwini PirsBry and Cooker, Man. Conch., vol. 22, p. 304,
pl. 50, figs. 14-14b (only), 1014.
The shell resembles 4. a. leucorraphe, to which it is closely re-
lated, but differs from l/eucorraphe in having a brown sutural band,
the embryonic whorls lined on the last half embryonic whorl with
yellow, and the usual absence of zigzag streaks on the postembryonic
whorls. The holotype (pl. 2, fig. 16) has the embryonic whorls
bicolored ; embryonic whorls white, banded on the lower third of the
first embryonic whorl with natal brown, on the next whorl the band
fading to fawn color and finally honey yellow on the last half em-
bryonic whorl; postembryonic whorls pale pinkish cinnamon, axially
streaked with natal brown and drab; impressed sutural band chest-
nut; lip and columella callus white faintly tinted with pale vinaceous
fawn. Length 18.4 mm., greater diameter 12.2 mm., spire height
9.6 mm., number of whorls 64.
Distribution, area 73: Waikakalaua Stream, Wilder, BBM 50572,
exact location of Wilder’s type locality not known; Kipapa-Wai-
kakalaua Ridge, locality 250A-2, elevation 1,050-1,100 feet, 3 sinis-
tral, Lemke, 1936; North Kipapa Gulch, first North Branch, lo-
cality 240D-1a*, approximate elevation 900-1,000 feet, 2 dextral,
Spalding, BBM 117384. Wilder’s type locality probably came from
somewhere in the region of localities 250A-2 and 240D-1a*. Area 73
is considered the probable distribution of the race. This race occurs at
higher elevations but is usually not dominant and mixed with the
lighter tan form of A. a. virgatifulva var. 2 (figs. 5, 5a, p. 105).
In the Wilder lot the typical color pattern occurs on 58 percent
of the shells. A variation of this pattern, the lightest color pattern
and the obese form of the shell (pl. 9, fig. 5), has the first half post-
embryonic whorl white, axially streaked with cinnamon buff, the re-
maining postembryonic whorls pinkish buff axially streaked with
natal brown, army brown, and bone brown; impressed sutural band
russet shading to tawny on the last whorl; lip tilleul buff. Length
17.5 mm., greater diameter 12.5 mm., spire height 9.6 mm.
An elongate specimen (pl. 9, fig. 5@) with the usual color of the
darker color pattern occurring on 36 percent of the shells has the
postembryonic whorls pinkish buff, heavily axially streaked with
natal brown, the streaks broken by lines of pinkish buff; lip and
columella callus white shading to light pinkish cinnamon at the outer
edge. Length 19.2 mm., greater diameter 12.2 mm., spire height
104 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
1.0mm. The darkest form of this darker color pattern (pl. 9, fig. 5b)
is found on 5 specimens out of 26; first half postembryonic whorl
cinnamon buff, lined with white, last half of the whorl russet, lined
and streaked with pinkish buff, first three-fourths of the penultimate
whorl chocolate, last fourth of penultimate and last whorl pinkish
buff, spirally lined and heavily axially ornamented with closely set
axial streaks of bone brown; impressed sutural band chestnut; lip
and columella callus pale ochraceous buff.
On plate 9, figures 5c and 5d show the range of form of dextral
specimens. Figure 5c has a color pattern similar to plate 9, figure 5.
Length 17.7 mm., greater diameter 11.8 mm., spire height 9.8 mm.,
number of whorls 64. The shell of figure 5d has a color pattern
occurring on 3 percent of the shells and looks like typical A. a.
leucorraphe except for the brown sutural band; the last two and a
half postembryonic whorls on this specimen have the first whorl
white, and the last whorl and a half bone brown axially streaked and
spirally lined with pale smoke gray or white. Length 17.3 mm., greater
diameter 12.5 mm., spire height 8.8 mm., number of whorls 54.
On 3 percent of the shells the lined pattern of plate 9, figure 5e
is found, the last two postembryonic whorls pale pinkish buff axially
streaked with hair brown and spirally lined with fuscous, the lines
broken by the ground color.
ACHATINELLA APEXFULVA VIRGATIFULVA var. 1
PLATE 2, FIGURE 20; PLATE 9, FiIGuRE 6
Area 75: Waikakalaua-South Kaukonahua Ridge, locality 260%,
elevation 1,000-1,100 feet, Meinecke, BBM 121939, 3 dextral, 1 sinis-
tral, 1932. Also collected by Thurston, BBM 130725, 14 sinistral,
probably from area 75 (figs. 5, 5a, p. 105).
Only six shells are known from locality 260*. The shells have
been separated out from area 73 because the axial streaks appear to
be more closely set together and not as thick as in typical A. a.
virgatifulva. But then certain specimens of the two areas resemble
each other, so that I am undecided as to whether or not they are
distinct subspecies.
A common color pattern from locality 260* on a juvenile specimen
(pl. 2, fig. 20) has the embryonic whorls bicolored, upper half white,
lower half honey yellow shading to chamois; first postembryonic whorl
pale pinkish buff spirally lined and axially streaked with verona
brown, last whorl and a half pinkish buff, finely axially streaked with
olive brown and bone brown.
ACHATINELLA APEXFULVA DIXON—WELCH TO5
155 8
RI ok
Waikane
Punaluu
Fic. 5.—Northwest-central Oahu, leeward slope of the Koolau Range, comprising
regions IIIb and most of regions III and IV (fig. 7, p. 194), showing localities of
subspecies of A. apexfulva belonging to the groups of A. a. polymorpha, A. a. irwini,
A. a. leucorraphe, A. a. lilacea, A. a. apicata, A. a. aloha, and A. a. apexfulva.
N Kahana
Miles :
————] \I\Waikane,,.
0 1 Waiahole
(4
4
E
3
ov
~
§
nN
©
=
JE
=
Z=
s
a
o
Zz
\
1
o
Fic. 5a.—Same as fig. 5, but showing the area occupied by different subsperics of
Sek
A, apexfulva belonging to the groups of A. a. polymorpha, A. a. irwim,
leucorraphe, A. a. lilacea, A. a. apicata, A. a. aloha, and A. a. apexfulva.
100 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
One of the Thurston shells (pl. 9, fig. 6) has the upper half of
the first two embryonic whorls white, lower half natal brown or
army brown, last embryonic whorl upper two-thirds white, lower third
honey yellow; first postembryonic whorl pale pinkish buff, axially
streaked with pinkish buff, first half of the penultimate whorl pale
pinkish buff axially streaked with cinnamon buff, last half cinnamon
buff axially streaked with mikado brown, last whorl cinnamon brown
finely axially streaked with quaker drab and chocolate; impressed
sutural band chestnut; lip and columella callus pale vinaceous fawn,
lip outer edge light vinaceous cinnamon.
ACHATINELLA APEXFULVA VIRGATIFULVA var. 2
PLATE 2, FIGURE 17; PLATE 9, FIGURES 7-9
Achatinella leucorraphe irwint Pttspry and Cooke, Man. Conch., vol. 22, p. 305,
pl. 50, fig. 13 (only), 1914.
Area 74: North Kipapa Gulch, first North Branch, locality
240D-1, elevation 9g50-1,000 feet, 2 dextral, Lemke, 1935; Kipapa-
Waikakalaua Ridge, locality 251-1, elevation 1,300-1,350 feet, H.
Lemke, 5 dextral 1933; 250*, elevation 1,200 feet, Meinecke, 1932;
251-1-3*, Russ, 1931; 251-3, elevation 1,500-1,600 feet, 6 dextral,
Meinecke, 1933. Also collected by Spalding and Thurston (figs. 5, 5a,
ps Os,):
Shells with color patterns very similar to that of A. a. virgatifulva
var. 2 are found in lots of shells from Waikakalaua Stream, Wilder,
BBM 50571, and South Kaukonahua Stream, Wilder, BBM 50573
(probably the Waikakalaua-South Kaukonahua Ridge).
The shell is closely related to A. a. virgatifulva, but differs in having
a light tan streaked color pattern instead of a dark brown streaked
one, and a light impressed sutural band instead of a dark brown
sutural band. Specimens of A. a. virgatifulva are usually mixed with
this form. Although the color pattern is very distinct and the form
is undoubtedly a distinct one, I shall consider it a variety of A. a.
virgatifulva until a locality is found that contains shells of A. a.
virgatifulva var. 2 not mixed with A. a. virgatifulva. Most of the
localities at lower elevations in areas 74, 57, 75, and 60 are wide locali-
ties and little carefully plotted material is at my disposal from this
section.
The usual form (pl. 2, fig. 17) of the light color pattern has the
embryonic whorls bicolored, first embryonic whorl cinnamon drab,
next whorl upper half white, lower half shaded with natal brown, on
the last embryonic whorl the lower band fades out and the whorl is
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 107
white ; the first postembryonic whorl and a half white or pale pinkish
buff, axially streaked with pale pinkish buff, last whorl and a half
pale pinkish buff axially streaked with vinaceous buff, avellaneous and
wood brown, and spirally lined with pale pinkish buff; impressed
sutural band same as the remainder of the whorl; lip and columella
callus pale vinaceous fawn, outer edge of lip pinkish buff. Length
17.7 mm., greater diameter 12.3 mm., spire height 9.4 mm., number
of whorls 64.
The obese form (pl. 9, fig. 7) and light color pattern has the post-
embryonic whorls white, penultimate and last whorls shaded by axial
streaks of a dilute color of vinaceous buff. Length 18.5 mm., greater
diameter 13.1 mm., spire height 9.4 mm., number of whorls 6. An
elongate specimen (pl. 9, fig. 8) has the postembryonic whorls white
shading to tilleul buff on the last whorl, last two whorls axially
streaked with avellaneous. Length 17.7 mm., greater diameter
11.8 mm., spire height 9.8 mm.
In the Russ lot from the region of 250-251-3*, lined forms similar
to plate 9, figure 8a, are found; the postembryonic whorls are white,
penultimate faintly spirally lined or banded with cinnamon buff and
sayal brown which are broken by axial streaks of white, last whorl
spirally lined and banded with sayal brown and warm sepia, the lines
and bands broken by axial streaks of white. This form is probably
one from the upper edge of area 74 because it is similar to the form
of A. a. ewaensis var. 6 (pl. 8, fig. 7) found in locality 251-1-2.
This light color form may be sinistral (pl. 9, fig. 9) ; the postem-
bryonic whorls a yellowish hue of vinaceous buff, axially streaked
with white.
ACHATINELLA APEXFULVA TUBERANS Gulick
PLATE 9, FIGURE I2
Apex iuberans Guuick, Proc. Zool. Soc. London, 1873, p. 81, pl. 10, fig. 3.
Achatinella swiftii Newcoms, Pilsbry and Cooke, Man. Conch., vol. 22, pp.
306, 313, 314, pl. 58, figs. 6, 9b, 11a? (only), 1914.
The holotype (pl. 9, fig. 12) in the Museum of Comparative
Zoology has the embryonic whorls white, last embryonic whorl with
a central band of cartridge buff; first postembryonic whorl cartridge
buff, shading to tilleul buff on the last half of the whorl spirally
banded on the lower fourth of the whorl with clay color darkening to
benzo brown, penultimate whorl light mouse gray lined with mikado
brown, benzo brown, last whorl below the white subsutural band,
pale mouse gray axially streaked with light mouse gray, banded with
108 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
an upper band of mikado brown and a lower band of chocolate,
peripheral band white, below the periphery white lined with mikado
brown, sayal brown, and two bands of bone brown, below which is a
band of sorghum brown; lip and columella callus tilleul buff; im-
pressed sutural band shades from fawn color on the first half post-
embryonic whorl to vinaceous buff on the next whorl and a half, on
the last whorl white or tilleul buff. Length 19.6 mm., greater di-
ameter 13.8 mm., spire height 10.5 mm., number of whorls 6.
Distribution, area?: Kalaikoa, Gulick. Also reported from
Ahonui, Wahiawa, and Helemano by Gulick. These last three locali-
ties are probably erroneous.
The typical A. a. tuberans pattern is not plentiful in the Gulick
collection. Out of 30 specimens in the Bishop Museum marked
A. tuberans by Gulick coming from Kalaikoa and Ahonui, only six
can be said to have a pattern closely simulating the pattern of the
holotype. Quite probably Gulick obtained the few specimens of
A. a. tuberans from an entirely different place from the remainder of
his material and mixed them with shells from a different region.
Typical tuberans has not been located in recent years but the forms
collected today of A. a. tuberans are probably closer to typical A. a.
tuberans than most of Gulick’s specimens, which undoubtedly belong
to a different colony and will be dealt with under A. a. tuberans var. 1.
The type locality of the holotype probably was somewhere at a
low elevation in South Kaukonahua Stream.
ACHATINELLA APEXFULVA TUBERANS var. 1
PLATE 9, FIGURES 13-13¢
Achatinella swiftii NEwcoms, Pilsbry and Cooke, Man. Conch., vol. 22, p. 314,
pl. 58, figs. 9, 9a (only), 1914.
Area?: Kalaikoa, Gulick. This area was probably somewhere at
a low elevation in South Kaukonahua Stream.
The form is probably a race occurring near A. a. flavida because
the ground color of so many specimens is yellow. The usual color
pattern (pl. 9, fig. 13) on 45 percent of the shells has the embryonic
whorls white ; first postembryonic whorls white faintly axially streaked
with pallid mouse gray, penultimate whorl white axially streaked and
spirally lined with light mouse gray, last whorl above the periphery
pale olive gray axially streaked and tinted with colonial buff and
spirally lined or banded and axially streaked with natal brown,
below the periphery the ground is colonial buff axially streaked and
lined with dark livid brown; impressed sutural band vinaceous buff ;
NO. I ACHATINELLA APEXFULVA DIXON—-WELCH 10g
lip and columella callus white or a faint shade of pale vinaceous fawn.
Length 18.0 mm., greater diameter 12.7 mm., spire height 9.8 mm.,
number of whorls 64. The usual form is not determined because of
a lack of sufficient number of adult shells.
An obese shell (pl. 9, fig. 13a) and a variant of the usual pattern
which does not have the lip fully developed, measures: Length
17.1 mm., greater diameter 13.5 mm., spire height 8.6 mm.; penulti-
mate whorl white axially streaked with natal brown and spirally
lined with white, last whorl chamois spirally banded and axially
streaked with hay’s brown above the periphery, the peripheral band
is chamois axially streaked with avellaneous, below the periphery the
subperipheral band is hay’s brown, the remainder of the base is
dark olive buff, lined and streaked with hay’s brown. The light color
pattern on 17 percent of the shells (pl. 9, fig. 13h) is similar to
A. a. versicolor and has the last two whorls above the periphery
white, axially streaked and spirally lined with vinaceous fawn color
and army brown, on the last whorl below the periphery the ground
is colonial buff spirally banded with hay’s brown about the base and
faintly axially streaked and spirally banded with light drab.
Thirty-eight percent of the shells have a pattern similar to that of
plate 9, figure 13c, which shows a narrow shell; the first two post-
embryonic whorls are pale smoke gray, spirally lined and streaked
on the first postembryonic whorl with warm sepia, white, and cinna-
mon buff, penultimate whorl axially streaked with zigzag and straight
streaks of natal brown and army brown, and spirally banded with a
band of white below the suture and subsutural bands, last whorl
drab gray axially streaked and lined with benzo brown and hair
brown, and banded below the subsutural band and at the periphery
with white, tinted with tilleul buff; the sutural and subsutural bands
form a single wide band, which is natal brown axially streaked with
white on the first half of the penultimate, and wood brown axially
streaked with vinaceous buff on the last whorl and a half.
ACHATINELLA APEXFULVA TUBERANS var. 2
PLATE 2, FIGURE 21; PLATE 9, FIGURES IO-IIa
Achatinella leucorraphe irwini Pitspry and Cooxe, Man. Conch., vol. 22, p. 304,
‘pl. 59, fig. 9 (only), 1914.
Area 60: Waikakalaua-South Kaukonahua Ridge, locality 261%,
approximate elevation 1,450-1,529 feet, 1932; 260-261?, approxi-
mate elevation 1,100-1,529 feet, Meinecke, 1913. Locality 261 may
not be very accurately plotted but is probably somewhere between
localities 260 and 261 (figs. 5, 5a, p. 105).
Ifo SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
This race is apparently an intermediate between virgatifulva var. 1
(area 75) and ewaensis var. 8 (area 59). This race usually occurs
mixed with A. a. ewaensis var. 8, and I do not know whether it is a
pure race or not. Probably the reason for the mixture is that all the
collecting has been done over a wide area and the one locality that is
not of wide extent may be a region where the two races overlap.
The usual color pattern (pl. 2, fig. 21) has the embryonic whorls
bicolored; first half embryonic whorl hair brown, last half upper
half white, lower half hair brown, next embryonic whorl and a half
upper half white, lower half with two bands each covering a fourth
of the whorl, the lower one benzo brown, the upper honey yellow, last
half embryonic whorl upper half white, lower half chamois; first
one-half postembryonic whorl white, with a line just above the edge
of the periphery of chamois, next whorl white, lined with pinkish
cinnamon, just above the edge of the periphery a line of verona
brown, last whorl and a half white, shading to pale gull gray, spirally
lined and banded with deep brownish drab; impressed sutural band
white on the first postembryonic whorl and a half, on the last
whorl and a half the color darkening from cartridge buff to tawny
on the last whorl; lip and columella callus pale vinaceous fawn.
Length 18.4 mm., greater diameter 12.2 mm., spire height 9.8 mm.,
number of whorls 63. The usual form is not determined because
so few adult specimens of this race are at hand.
The color pattern varies. A dextral (pl. 9, fig. 10) has the post-
embryonic whorls white, first half of last whorl banded above the
periphery with a band of avellaneous and faintly axially streaked
with tilleul buff, below the periphery the axial streaks deepening to
vinaceous buff, last half whorl tilleul buff axially streaked with vina-
ceous buff, all below the periphery and just above the periphery the
base is lined, banded closely, axially streaked with fine streaks of
grayish olive and benzo brown, with a dark subperipheral band of
mouse gray; impressed sutural band on the last two whorls tawny;
subsutural band deep quaker drab. This specimen has the light streak-
ing of the lower race and the gray banding of the upper race.
The color pattern may be lighter (pl. 9, fig. 11); first half of
penultimate pale pinkish buff, spirally lined with cinnamon buff, last
whorl and a half tilleul buff, spirally lined and axially streaked with
wood brown and drab.
A specimen (pl. 9, fig. t1@) is found which has a pattern very
similar to the typical tuberans pattern (pl. 9, fig. 12). The shell is
not adult ; first postembryonic whorl white, spirally lined with mikado
NO. I ACHATINELLA APEXFULVA DIXON—WELCH III
brown, last whorl and a half white, spirally lined with mouse gray,
in the umbilical region a band of pinkish buff ; impressed sutural band
russet.
ACHATINELLA APEXFULVA TUBERANS var. 3
PLATE 2, FIGURE 28; PLATE 9, FIGURES 14-15
Area 77: North-South Kaukonahua Ridge, locality 274, elevation
1,600-1,747 feet, Lemke, 1932, Steel, 2 dextral 1934. Also collected
by Wilder, BBM 10448, and O. H. Emerson, BBM 103971, probably
in the region of area 77 (figs. 5, 5a, p. 105).
This form of A. a. tuberans resembles typical tuberans very closely
in color pattern. None of the specimens, however, have on the last
whorl the dark bands unbroken by axial streaks that are so char-
acteristic of typical tuberans. Area 77 is an intermediate one between
the subspecific group of A. a. lilacea and A. a. leucorraphe because
there is a mixture of both kinds of embryonic whorls in the locality.
A usual color pattern (pl. 2, fig. 28) has the embryonic whorls light
buff, last embryonic whorl and a half banded on the upper half of
the whorl with white; penultimate and last postembryonic whorl
light mouse gray spirally lined and banded with white and axially
streaked with dark livid brown and hair brown; impressed sutural
band ochraceous tawny with a line of bone brown on the lower half
of the band, on the last whorl the upper half of the band ochraceous
tawny and the bone-brown line is in the center of the band, lower half
light mouse gray axially streaked with dark livid brown; below the
sutural band is a line of bone brown; lip and columella callus pale
ochraceous fawn. Length 21.2 mm., greater diameter 14.2 mm., spire
height 11.4 mm. The usual form of the shell is not shown by
figure 28. The usual length of the shell is around 19.5 mm., and has
a greater diameter of 12.5 or 13.5 mm.
The color pattern may be much lighter (pl. 9, fig. 14) ; embryonic
whorls white; first two postembryonic whorls pale pinkish cinnamon
spirally lined with tilleul buff on the upper half of the penultimate
whorl, lower half banded smoke gray axially streaked with pale
pinkish cinnamon or white, last whorl light drab darkening to hair
brown on the last half whorl, axially streaked with benzo brown,
drab, and white and spirally banded and lined with white; impressed
sutural band tilleul buff ; lip not formed; specimen not quite adult.
The shells of area 77 seem to be an intermediate race between lower
punicea var. 1 and the higher race of irwini. The Lemke lot con-
tains shells with unicolored embryonic whorls, but one specimen
8
I12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
(pl. 9, fig. 14a) in the lot has embryonic whorls similar to A. a.
irwinit. The embryonic whorls are bicolored, first embryonic whorl
army brown, next embryonic whorl and a half upper half white,
lower half natal brown, last three-fourths embryonic whorl chamois
on the lower two-thirds of the whorl, upper third white; post-
embryonic whorls drab gray spirally lined with white and dusky
drab, and axially streaked with white, last half whorl above the
periphery almost solid dusky drab, at the edge of the periphery a
band of dusky drab, above and below which is a band of white.
The Wilder and O. H. Emerson shells probably came from near
area 77, the usual length of the shell is 18.5 mm., and the usual color
pattern (pl. 9, fig. 15) has the last two whorls white deepening to
pale gull gray on the last whorl, spirally lined above the periphery
with mouse gray, the peripheral band deep mouse gray, below the
white subperipheral band, the base mouse gray spirally banded with
benzo brown; impressed sutural band pinkish buff. Length 18.2 mm.,
greater diameter 12.8 mm., spire height 9.6 mm. The Wilder speci-
mens consisting of two shells are selected from a mixed lot contain-
ing shells from a lower locality with a different color pattern.
ACHATINELLA APEXFULVA TUBERANS var. 4
PLATE 2, FIGURE 23; PLATE 9, FicurREsS 16-16¢
Achatinella leucorraphe irwini Pitspry and Cooke, Man. Conch., vol. 22, p. 305,
pl. 50, figs. 16-16b (only), 1914.
Area 76?: South Kaukonahua South-Central Branch Ridge
in the general region of locality 260A ?, approximate elevation 1,550-
1,700 feet, Meinecke, BBM 121968, 1918, BBM 121952, 1924. Other
collectors of this form are Wilder, BBM 10442, O. H. Emerson,
BBM 103969, J. S. Emerson. The shells of area 76 are described
from the Meinecke lot, BBM 121968 (figs. 5, 5a, p. 105).
The usual color form of the shell (pl. 2, fig. 23), lot BBM 121968,
has the embryonic whorls bicolored, first half whorl army brown,
next whorl and a half upper half pinkish buff, lower half army brown
lightening to wood brown, next embryonic whorl pinkish buff lined
with white, last half embryonic whorl white; first quarter post-
embryonic whorl white, next quarter whorl light buff axially streaked
with warm sepia, last half of the whorl and first half of penultimate
warm buff spirally lined with and streaked with warm sepia, last half
of penultimate and first half of last whorl warm buff faintly lined
with wood brown and a line of fawn color, last whorl banded just
above the edge of the periphery with a band of sepia axially streaked
No. I * ACHATINELLA APEXFULVA DIXON—-WELCH 113
with snuff brown, below which is a band of pinkish buff, base spirally
banded with sepia and snuff brown; impressed sutural band warm
sepia lightening to verona brown on the last whorl; lip vinaceous
fawn; columella callus white. Length 18.0 mm., greater diameter
12.2 mm., spire height 10.2 mm. ;
Plate 9, figure 16, is a narrow specimen and light color pattern, the
last two postembryonic whorls tilleul buff spirally lined with natal
brown on the penultimate and hair brown and drab gray above the
periphery on the last whorl, base below the white subperipheral band
drab, banded with buffy brown and lined with white; impressed
sutural band wood brown darkening to cinnamon on the last whorl.
Length 18.0 mm., greater diameter 11.6 mm., spire height 10.2 mm.
The usual dark color form (pl. 9, fig. 16a) has the first whorl
vinaceous buff, axially streaked with army brown and natal brown,
last whorl and a half below the sutural band pale gull gray, lined
with hay’s brown, with a supraperipheral band of hay’s brown
axially streaked with deep olive buff, at the edge of the periphery
a band of white, below the periphery the base is avellaneous axially
streaked with hay’s brown and spirally lined with white; impressed
sutural band verona brown. Length 17.7 mm., greater diameter
13.0 mm., spire height 9.3 mm.
A variation of the dark color pattern (pl. 9, fig. 16b) found on two
specimens has the last two postembryonic whorls pale pinkish buff
axially streaked and spirally lined with hair brown and chaetura
drab, last whorl lined at the periphery with two lines of white, ground
color of the last half whorl drab gray; impressed sutural band russet.
One specimen (pl. 9, fig. 16c) is found to have a pattern similar to
that of typical A. a. tuberans, except that the embryonic whorls are
bicolored as in plate 2, figure 23; the last two whorls are pale gull
gray spirally lined and banded with bone brown and natal brown, base
on the last half whorl axially streaked with natal brown; impressed
sutural band russet lightening to tawny on the last half whorl. One
specimen similar to plate 9, figure 17¢, is also found in this lot,
differing only in that it has a peripheral yellow band.
ACHATINELLA APEXFULVA TUBERANS var. 5
PLATE 9, FIGURES 17-17¢
Distribution, area?: South Kaukonahua Stream, Wilder, BBM
50575. This lot of shells may have come from somewhere near or
along the South Kaukonahua South-Central Branch Ridge. However,
no material from this ridge has been carefully localized and no accu-
114 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103 |
|
rate information concerning distribution and variation has been avail- |
able to me. Every lot of shells collected from the South Kaukonahua |
shows a different set of color patterns. Wilder’s lot contained a mix- —
ture of A. a. tuberans var. 4 patterns, which were mixed with decid- —
edly smaller shells with a different range of color patterns. Therefore,
the var. 4 patterns are selected out and the remaining shells are con-
sidered tuberans var. 5, which may be the same or a lower race (fig. 5,
p. 105). Also collected by Thurston, BBM 130737.
The usual form of the shell (pl. 9, fig. 17) has the embryonic
whorls bicolored; first embryonic whorl natal brown, next whorl
upper half white, lower half natal brown, last embryonic whorl and
a fourth white; first half postembryonic whorl white, penultimate
whorl axially streaked by straight and zigzag streaks of warm sepia
and vinaceous buff, and spirally lined and banded with white, last
whorl above the periphery ornamented with a band of white and a
line of buffy brown and a band of vinaceous buff axially streaked
with olive brown and hair brown, below the periphery the ground
is white, banded with a band of bister and a band of drab axially
streaked with olive brown; impressed sutural band mikado brown.
Length 16.9 mm., greater diameter 11.6 mm., spire height 7.8 mm.,
number of whorls 6.
A narrow shell (pl. 9, fig. 17a) with lighter embryonic whorls,
has a more usual color pattern which occurs on 34 percent of the
shells, embryonic whorls bicolored, first embryonic whorl vinaceous
buff, next whorl upper half white, lower half vinaceous buff, last
embryonic whorl and a quarter white; first postembryonic whorl white
spirally lined with army brown, last two whorls pale gull gray, penulti-
mate whorl spirally lined and faintly axially streaked with fuscous,
last whorl above the periphery banded with olive brown, the bands
axially broken by streaks of pale gull gray, at the edge of the periphery
a line of bister, below the periphery a band of pale gull gray, below
this a line of bister and a band of drab axially streaked with natal
brown, the remainder of the base banded with bister and drab; im-
pressed sutural band chestnut lightening on the last half whorl to
tawny. Length 16.6 mm., greater diameter 10.8 mm., spire height
9.2 mm.
The obese form (pl. 9, fig. 17b) and color pattern occurring on
28 percent of the shells has the last two postembryonic whorls white
or pale gull gray spirally lined with natal brown and axially
streaked with pale gull gray, just above the edge of the periphery to
the edge of the periphery a band of light seal brown, base banded and
NO. I ACHATINELLA APEXFULVA DIXON—WELCH I15
lined with natal brown; impressed sutural band vinaceous_ buff.
Length 16.1 mm., greater diameter 11.8 mm., spire height 8.9 mm.
The lightest color pattern (pl. 9, fig. 17c) occurring on 31 percent
of the shells has the postembryonic whorls white spirally lined on the
last two whorls with light mouse gray and mouse gray, which lines
are broken by axial streaks of white; impressed sutural band white,
upper edge fawn color. This specimen also shows a sinistral shell.
Four percent have a pattern similar to that of plate 9, figure 17d;
postembryonic whorls white, last two whorls with a band of chocolate
just above the edge of the periphery, just below the edge of the
periphery a band of white, remainder of the base wood brown, lined
and banded with natal brown. Two specimens in the lot of 75 shells
have a color pattern resembling typical A. a. apexfulva. One of these
(pl. 9, fig. 17e) has the first embryonic whorl army brown, next
whorl upper two-thirds white, lower third natal brown, last embryonic
whorl white; last two postembryonic whorls chocolate faintly axially
streaked on the penultimate with orange cinnamon.
Group OF A. A. IRWINI PILSBRY AND COOKE
ACHATINELLA APEXFULVA EWAENSIS, new subspecies
PLATE 2, FIGURE 15; PLATE 7, FIGURES 23-24@
The shell is similar to A. a. irwini but differs in being smaller and
having a characteristic series of reddish-brown banded color forms
which differ entirely from A. a. irwint. The embryonic whorls of
the shells of A. a. ewaensis and its varieties are usually banded with
a deep shade of yellow such as dresden brown, chamois, or cinnamon
buff. A. a. irwini and its varieties usually do not have the embryonic
whorls banded with yellow. A few specimens of A. a. irwini do have
the embryonic whorls banded with tints of yellow, such as cartridge
buff and cream buff, but never intensely banded with yellow.
The embryonic whorls of the holotype (pl. 2, fig. 15) are bicolored,
upper half white, lower half banded with dresden brown shading to
buckthorn brown on the last embryonic whorl and a half ; postembry-
onic whorls white faintly tinted with faint lines of pale gull
gray, last whorl lined at and below the periphery with seal brown;
lip and columella callus tinted with light brownish vinaceous. Length
17.0 mm., greater diameter 12.1 mm., spire height 8.5 mm., number
of whorls 53.
Distribution, area 56: North-Central Kipapa Ridge, type locality
240L-6, elevation 1,750-1,900 feet; also locality 240L-2, elevation
1,550 feet, 2 sinistral; 240L-3, elevation 1,550-1,650 feet, 1 dextral,
116 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
3 sinistral ; 240L-4, elevation 1,750 feet, all Meinecke-collected, 1933
(figs. 5, 5a, p. 105).
The typical color pattern occurs on 40 percent of the shells in area
56. Thirty-three percent have a reddish-brown banded color pattern
(pl. 7, fig. 24a), which has the postembryonic whorls white, lined
with hay’s maroon; impressed sutural band russet. Half of these
reddish-brown banded shells are similar to plate 7, figure 23;
last postembryonic whorl and a half white, banded just above, at, and
below the periphery with hay’s maroon; impressed sutural band
russet. The light color form occurring on 27 percent of the shells is
shown by an elongate shell (pl. 7, fig. 24) ; the postembryonic whorls
white faintly banded with pale olive gray at the edge of the periphery,
last half whorl at and below the periphery lined with black. Length
17.7 mm., greater diameter 12.0 mm., spire height 9.2 mm.
ACHATINELLA APEXFULVA EWAENSIS var. 1
PLATE 2, FIGURE 4; PLATE 7, FIGURES 19-20
Area 51: North-South Waiawa Ridge, locality 233Ba, elevation
1,150 feet, 1 sinistral; 233B, elevation 1,450 feet, 14 sinistral; 234B,
elevation 1,500 feet; 235B, elevation 1,500-1,550 feet, 4 dextral;—
236B, elevation 1,500-1,650 feet, 1 sinistral; 237B, elevation 1,700
feet, 3 dextral, 9 sinistral; 238B, elevation 1,700-1,750 feet, 1 dextral ;
239B, elevation 1,700-1,750 feet, 1 dextral, 5 sinistral; North
Waiawa North-South Branch Ridge, locality 230D, elevation
1,650 feet, 13 sinistral; 231D, elevation 1,700-1,750 feet, I sinistral ;
232D, elevation 1,950-2,000 feet, 3 sinistral, all collected by Meinecke,
1933; Waiawa-Waiahole Ridge, Spalding, BBM 117388 (figs. 4,
44, p. 53).
The color pattern of many of the forms existing in area 51 are not
very strikingly different from those of the forms obtainable from
area 56. The main difference is that the typical form of the shell
from area 50 has the embryonic whorls more intensely banded with
yellow than the shells of area 51, and a few color patterns (pl. 7,
figs. 19, 19a) are not found in area 56.
The usual form and color pattern (pl. 2, fig. 4) has the first
embryonic whorl pale pinkish buff, remaining embryonic whorls upper
half white, lower half cream color; first half of first postembryonic
whorl white, banded just above the edge of the periphery with a
faint band of cream color, remaining postembryonic whorls white
with a line of ochraceous buff on the last whorl and a half just above
the edge of the periphery, last whorl at and below the periphery
NO. I ACHATINELLA APEXFULVA DIXON—-WELCH II7
lined and banded with seal brown; lip and columella callus pale
brownish vinaceous. Length 18.4 mm., greater diameter 12.2 mm.,
spire height 9.0 mm.
A dextral shell (pl. 7, fig. 19) has the postembryonic whorls
cartridge buff faintly tinted with lines and axial streaks of cream
buff, last whorl lined and banded with seal brown; lip and columella
callus vinaceous fawn.
An obese sinistral shell (pl. 7, fig. 19a), length 19.3 mm., greater
diameter 13.4 mm., spire height 9.1 mm., shows a gray color pattern
which is similar to the gray color patterns (pl. 8, figs. 6a, 8) found in
areas 55 and 58, but not found in area 56 or other areas containing
varieties of A. a. ewaensis. The embryonic whorls are white, banded
on the lower half with chamois; postembryonic whorls white, banded
on the last whorl above and below the periphery with light gull gray,
and lined at the periphery on the last whorl and at the base of the
last whorl with carob brown.
Four specimens in area 51 have a lined pattern (pl. 7, fig. 20) that
is similar to the typical A. a. ewaensis color pattern (pl. 2, fig. 15)
of area 56; embryonic whorls white, banded on the lower three-fourths
of the whorl with warm sepia on the first whorl and shading to
ochraceous tawny on the remaining embryonic whorls; postembryonic
whorls white, first postembryonic whorl banded on the lower half
with warm buff, last whorl and a half lined with seal brown. This
shell also shows an elongate form. Length 20.1 mm., greater diameter
12.3 mm., spire height 10.7 mm.
ACHATINELLA APEXFULVA EWAENSIS var. 2
PLATE 2, FIGURE 12; PLATE 7, FIGURES 21-21}
Area 53: Waiawa-Kipapa Ridge, locality 244*, elevation 1,500-
1,700 feet, Meinecke, 1929 (figs. 5, 5a, p. 105).
The shells in area 53 differ from those of the other areas of
varieties of A. a. ewaensis by having yellow banded color patterns
(pl. 2, fig. 12; pl. 7, fig. 21). The usual form and color pattern of
the shell (pl. 2, fig. 12) has the first embryonic whorl white banded
on the lower fourth of the whorl with bone brown, remaining
embryonic whorls upper half white, lower half cinnamon buff; post-
embryonic whorls white spirally banded with cinnamon buff; lip
and columella callus white faintly tinted with light vinaceous fawn.
Length 17.1 mm., greater diameter 11.5 mm., spire height 9.0 mm.
The color of the bands on the postembryonic whorls (pl. 7, fig. 21)
may be darker, being sudan brown, and the embryonic whorls may
118 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
be larger and less blunt than those of the typical form. Eight percent
of the shells are entirely white and are similar to plate 7, figure 21a.
A dark banded form (pl. 7, fig. 21b) occurs on 30 percent of the
shells; the postembryonic whorls shade from sudan brown axially
streaked with warm buff to carob brown on the last whorl and a half,
the edge of the periphery and below the periphery white, base banded
and lined with carob brown.
ACHATINELLA APEXFULVA EWAENSIS var. 3
PLATE 2, FIGURE II; PLATE 7, FIGURES 25, 26
Area 52: Waiawa-Kipapa Ridge, locality 243-2, elevation 1,500-
1,550 feet, Meinecke, 1933; 243-3-4*, elevation 1,500-1,620 feet,
Meinecke, 1929, 5 dextral, 5 sinistral; 243-4*, Welch, 1934 (figs. 5,
5a, p. 105).
Area 52 contains forms (pl. 2, fig. 11) similar to typical A. a.
ewaensis, but differing in having the embryonic whorls not as
markedly banded with yellow, and in having the last whorl banded
with lighter reddish-brown bands. With the typical pattern are mixed
color patterns (pl. 7, figs. 25, 26), which resemble the shells of
area 72.
The usual form and color pattern of the shell (pl. 2, fig. 11) has
the embryonic whorls bicolored, upper half or three-quarters white,
lower half or fourth of the whorl banded with yellow deepening to
chamois ; postembryonic whorls white, last whorl below the periphery
banded with chocolate; lip and columella callus pale grayish vina-
ceous. Length 18.7 mm., greater diameter 13.0 mm., spire height
9.3 mm., number of whorls 64.
Out of 16 specimens from locality 243-2, 12 are dextral, 8 of which
are similar to plate 2, figure 11. Three of the dextrals are similar
to the shell of plate 7, figure 25, which has the postembryonic whorls
white spirally lined and banded with olive buff, last whorl at the edge
of the periphery banded with white, below the periphery lined with
two lines of bone brown and banded with smoke gray, last half whorl
lined above the white peripheral band with a line of bone brown and
six lines of bone brown below the peripheral band. Length 19.0 mm.,
greater diameter 12.5 mm., spire height 10.2 mm.
The color pattern on three shells looks like a light form of the
shells from area 71 or 72. One of these three shells (pl. 7, fig. 26)
has the postembryonic whorls white, spirally lined with deep olive
buff; impressed sutural band amber brown, subsutural band ochra-
ceous tawny.
INO} E ACHATINELLA APEXFULVA DIXON—WELCH 119g
ACHATINELLA APEXFULVA EWAENSIS var. 4
PLATE 2, FIGURE 13; PLATE 7, FIGURES 22, 22a
Area 54: Waiawa-Kipapa Ridge, locality 246*, elevation 1,800
feet, Meinecke, 1929; 246a, elevation 2,000-2,100 feet, 2 sinistral,
Hosaka, 1935. Meinecke also collected a lot, BBM 118442, from the
general region of locality 244 to 244-37. But since the shells
differ from those in locality 244, I shall consider the shells to have
come from the region of locality 244-3? Since the lot was collected
before 1933, the shells may represent a series collected above area 53
(figs. 5, 5a, p. 105).
The yellow banded color patterns of area 53 do not occur in area 54.
The usual color pattern (pl. 2, fig. 13) occurring on 77 percent of
the shells in locality 246 has the embryonic whorls white, banded on
the lower fourth of the whorl with a band of russet which lightens
to chamois on the last half embryonic whorl; postembryonic whorls
white, lined and banded on the last whorl with chocolate. Twenty-
three percent have a color pattern similar to A. a. irwint var. I
(pl. 8, fig. 15a) only the embryonic whorls are not like irwini, being
banded with yellow.
The series of shells from locality 244-3? contains a mixture of
patterns. A very lined form (pl. 7, fig. 22) has the first embryonic
whorl upper half white, lower half verona brown, remaining em-
bryonic whorls honey yellow on the lower three-fourths of the whorl,
upper fourth white; first two postembryonic whorls shade from
honey yellow to ochraceous tawny lined with white, last whorl banded
and lined with carob brown. The last whorl of this shell resembles
the lined pattern on the last whorl of A. a. ewaensis (pl. 7, fig. 24a).
The postembryonic whorls (pl. 7, fig. 22a) may be white, banded
on the first two postembryonic whorls just above the edge of the
periphery with buckthorn brown, last whorl banded at the edge of the
periphery and below the periphery with liver brown; lip and
columella callus vinaceous fawn.
ACHATINELLA APEXFULVA EWAENSIS var. 5
PLate 2, Figure 14; PLate 8, Ficures 6-6)
Area 55: North-Central Kipapa Ridge, locality 240L-1-1a*, ele-
vation 1,450-1,550 feet, 1932; 240L-1, elevation 1,550 feet, 1933,
Meinecke (figs. 5, 5a, p. 105).
This variety of A. a. ewaensis is intermediate between A. a.
ewaensis and a lower gray race which is probably a form closely re-
lated to A. a. virgatifulva. The color patterns figured from area 55
are quite distinct from those of area 56, but mixed with these are a
I20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
few specimens with patterns similar to A. a. ewaensis patterns. Very
possibly the variety is distinct enough to be considered a subspecies,
but I am reluctant to so consider it until more material is available
for study.
The usual color pattern (pl. 2, fig. 14) on 51 percent of the shells
from locality 240L-1-1a* has the embryonic whorls bicolored, upper
third white, lower two-thirds honey yellow; postembryonic whorls
white, banded just above the edge of the periphery with honey yellow
on the first whorl, on the first half of the penultimate the band is
cinnamon buff axially streaked with mikado brown, on the last whorl
and a half the band is seal brown, below the periphery the base has
a band of seal brown; lip and columella callus light vinaceous fawn.
Length 17.2 mm., greater diameter 12.3 mm., spire height 8.5 mm.,
number of whorls 54.
Nine percent may be devoid of dark bands (pl. 8, fig. 6a), having
the postembryonic whorls white, tinted with pale mouse gray; lip pale
vinaceous fawn, number of whorls 6. An obese shell (pl. 8, fig. 6)
shows a color pattern on 17 percent; the postembryonic whorls white
tinted with gull gray, the base fuscous black axially streaked with pale
smoke gray, and pale pinkish buff; lip not developed. Length
17.4 mm., greater diameter 13.5 mm., spire height 9.3 mm. The
typical ewaensis pattern (pl. 2, fig. 15) is present on 21 percent of
the shells.
One specimen is found in both lots of shells from area 55 which
may be a stray shell belonging to a distinct lower race. The first
embryonic whorl of one of the shells having this peculiar pattern
(pl. 8, fig. 6b) is worn, the color fuscous, the next embryonic whorl
upper half white, lower half pale smoke gray, last embryonic whorl
white ; postembryonic whorls white axially streaked with zigzag lines
of bone brown, on the last whorl the white peripheral band has a
line of bone brown in it at the edge of the periphery, base bone
brown axially streaked with white. Length 17.2 mm., greater diameter
II.I mm., spire height 9.5 mm.
In locality 240L-1, out of 12 live shells approximately 42 percent
are similar to plate 8, figure 6a, 33 percent to plate 8, figure 6, 17 per-
cent to plate 2, figure 14, and 8 percent to plate 8, figure 6b.
ACHATINELLA APEXFULVA EWAENSIS var. 6
PLATE 2, Figure 18; Puiate 8, Ficures 7-7b
Achatinella leucorraphe irwini Pitspry and Cooke, Man. Conch., vol. 22, pp. 304,
305, pl. 50, figs. 12, 15, 15a (only), 1914. These patterns illustrated by
Pilsbry occur on the Kipapa-Waikakalaua Ridge either in area 57 or in a
region including the lower part of area 57 and the upper part of area 74.
NO. I ACHATINELLA APEXFULVA DIXON—WELCH I21
Area 57: Kipapa-Waikakalaua Ridge, locality 252-1-2?, approxi-
mate elevation 1,500-1,600 feet, 4 dextral; 253, elevation 1,650 feet,
I dextral; 254, elevation 1,800-1,850 feet, 7 dextral; 255, elevation
1,800-1,850 feet, 2 dextral, all Meinecke-collected, 1933. In 1932
Meinecke, BBM 121935-121936, collected a series of 49 shells of
this variety, which probably came from locality 252-1-2?. Another
lot, BBM 121937-121938, collected by Meinecke in 1932 probably
comes from locality 253?. Most of the collecting below locality 253?
on this ridge is not very reliable and should be carefully checked
(figs. 5, 5a, p. 105).
Although the usual form and color pattern of plate 2, figure 18, and
the peculiar color patterns (pl. 8, figs. 7, 7a) of area 57, are very
distinct, other patterns (pl. 8, fig. 7b) occur which are so similar
to some of the color patterns (pl. 8, figs. 1, 2) of area 70 that mixed
lots would be hard to separate with 100 percent accuracy. Usually
the shells from area 57 are smaller than shells from area 70, and as
long as the characteristic patterns of the region are present they are
easy enough to separate. The Meinecke lots from 251-1-2? form the
largest series available for study from area 57, and the following
discussion will be based on it.
The usual form and color pattern of the shell (pl. 2, fig. 18) on 67
percent of the shells has the embryonic whorls white, banded on the
lower half of the whorl with cream buff ; postembryonic whorls white,
last two whorls spirally lined with warm sepia; lip and columella callus
light vinaceous fawn; impressed sutural band natal brown lightening
to wood brown on the last half whorl. Length 17.1 mm., greater
diameter 11.7 mm., spire height 9.2 mm. The spiral lines on the last
whorl may be broken and the spire elongate in form (pl. 8, fig. 7) ;
postembryonic whorls white, last whorl lined with snuff brown, the
lines broken by axial streaks of white; impressed sutural band snuff
brown. Length 17.0 mm., greater diameter 11.5 mm., spire height
g.2 mm. An obese shell (pl. 8, fig. 7a) has the embryonic whorls
white with a single peripheral band of saccardo’s umber axially
streaked with white, last half whorl faintly lined with snuff brown.
Length 16.4 mm., greater diameter 12.5 mm., spire height 8.2 mm.
The usual form of a sinistral and an example of the dark lined
form of the shell is shown on plate 8, figure 7b, postembryonic whorls
white spirally lined on the last whorl and a half with bone brown
and natal brown. Length 16.6 mm., greater diameter 11.7 mm., spire
height 8.8 mm. This pattern may be darker and resemble plate 8,
figure 2, a shell pattern from area 70. Twenty-five percent have a
white spire and a basal band similar to typical A. a. ewaensis (pl. 2,
I22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
fig. 15) of area 56. Two specimens have a pattern resembling plate 8,
figure 6, except that the basal gray band is lighter, and two resemble
plate 8, figures 7, 7a.
ACHATINELLA APEXFULVA EWAENSIS var. 7
PLATE 2, Figure 19; PLATE 8, FicuRrEs 8-10
Area 58: Kipapa-Waikakalaua Ridge, locality 256, elevation
1,750-1,800 feet; 257, elevation 1,800-1,950 feet; 258, elevation
1,900-1,950 feet, 10 dextral, Meinecke, 1933 (figs. 5, 5a, p. 105).
Lined forms similar to those occurring in area 57 (pl. 8, fig. 7b)
are rare or absent in area 58. In locality 256 the ground color of the
shell is white, tinted with yellow or gray; above this locality the white
ground is not tinted with yellow. In the entire area the characteristic
color patterns have one dark basal band or line.
The usual color pattern in area 58 (pl. 2, fig. 19) has the embryonic
whorls honey yellow, upper half or three-quarters white; post-
embryonic whorls white, tinted with pale gull gray above the periph-
ery, at the periphery a band of white, tinted with cartridge buff,
below the peripheral band a band of seal brown, base cartridge buff ;
impressed sutural band the ground color ; lip and columella callus pale
grayish vinaceous. Length 17.4 mm., greater diameter 12.0 mm.,
spire height 8.6 mm.
A variation of the typical pattern (pl. 8, fig. 8) is a two-lined
pattern ; postembryonic whorls above the periphery white, tinted with
pale gull gray, last whorl below the periphery ivory yellow tinted with
colonial buff, lined at the edge of the periphery and below the periph-
ery by two lines of bone brown. The basal lines may be wider, be-
coming two basal bands. Another common pattern of this area is
the white color form of plate 8, figure 6, tinted with pale gull gray.
Usually the shells from area 58 have blunter embryonic whorls than
the shells from areas to the south.
In the higher localities of 257 and 258 the same general plan of
banding and color pattern occurs. Plate 8, figure 9, shows a dextral
with a color pattern similar to that of plate 2, figure 11, except that
there are two subperipheral bands instead of five. In locality 258
two very much lined specimens are found. These are the only two
shells with a much banded or lined pattern in the entire area. One
of these (pl. 8, fig. 10) has the first postembryonic whorl and a half
white, banded on the lower half of the whorl with chamois, last
whorl and a half white banded and lined with chocolate. This locality
should be collected again to ascertain whether these two shells with
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 123
the lined pattern really belong to area 58 or whether they are present
owing to a mixture. Shells of this pattern are more usual in areas
74 or 57.
ACHATINELLA APEXFULVA EWAENSIS var. 8
PLATE 2, FIGURE 22; PLATE 8, FIGURES II-12a
Area 59: Waikakalaua-South Kaukonahua Ridge, general
region of localities 261-262?, approximate elevation 1,450-1,529 feet,
1913; 262*, elevation 1,450-1,500 feet, Meinecke, 15 dextral, 4
sinistral, 1932, BBM 121945-121946. Also collected by Wilder, BBM
50574, probably in the immediate vicinity of area 59 (figs. 5, 5a,
p. 105).
In area 59 the shells have the first embryonic whorl and a half
banded on the lower half of the whorl with brown similar to those
of areas 61? and 62?, and the remaining embryonic whorls are strongly
banded with yellow, resembling A. a. ewaensis (area 56). Some speci-
mens in area 59 (pl. 8, fig. 12) have embryonic whorls and a color
pattern very much like typical ewaensis (pl. 2, fig. 15) except that
the bands are a lighter shade of reddish brown. The localities on this
ridge are not accurately plotted and should be carefully collected
again. Very probably area 59 can be divided into two areas, the
upper containing shells close to A. a. ewaensis (pl. 2, fig. 22), the
lower one having a distinct race with color patterns similar to the
shells of plate 8, figures 11 and 12a, which resemble A. a. tuberans
var. 2.
The usual form of the shell (pl. 2, fig. 22) and common color
pattern has large, flattened, bicolored embryonic whorls; first em-
bryonic whorl white with the lower third of the whorl wood brown,
next whorl upper half white, lower band wood brown, last embryonic
whorl white, faintly tinted on the lower third of the first half of the
whorl with a cream-buff band; postembryonic whorls white spirally
banded just above the periphery on the last two whorls with tawny
which shades to light grayish olive on the last half whorl, below the
periphery of the last whorl a single band of hay’s maroon; lip and
columella callus pale grayish vinaceous. Length 18.1 mm., greater
diameter 12.8 mm., spire height 8.5 mm., number of whorls 5%. The
last whorl may have all manner of variations; the subperipheral and
supraperipheral bands may be made of two smaller bands or lines,
or the band above the periphery may be hay’s maroon as well as the
lower band.
A gray color pattern (pl. 8, fig. 11) found on one or two specimens
in area 59, has the first embryonic whorl white, lower third natal
brown, remaining embryonic whorls banded on the lower third with
124 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
a band of wood brown shading to chamois; first two postembryonic
whorls banded just above the edge of the periphery with a band of
cream buff shading to pinkish cinnamon, last whorl above the periphery
white, lined and banded with a band of cinnamon buff, which band
is axially streaked with pale pinkish buff and mikado brown, last
half whorl bone brown axially streaked with pale pinkish buff, below
the periphery a band of white; impressed sutural band on the first
postembryonic whorl white, on the penultimate pinkish buff shading to
mikado brown on the last whorl.
In locality 262* (BBM 121946) collected in 1932 by Meinecke,
the shells are 78 percent dextral, and only 17 percent have a gray
pattern similar to that of plate 8, figure 11. The common color
pattern (pl. 8, fig. 12) has the embryonic whorls bicolored, upper half
white, lower half honey yellow; postembryonic whorls white, last
whorl banded with army brown, shape ovate. Length 18.6 mm.,
greater diameter 13.0 mm., spire height 9.0 mm. An elongate shell
with dark bands and ochraceous tawny lines (pl. 8, fig. 12a) has a
color pattern found on only one specimen; the postembryonic whorls
are white, spirally lined or banded above the periphery with ochra-
ceous tawny, last whorl banded at and below the periphery with hay’s
maroon. Length 19.7 mm., greater diameter 12.6 mm., spire height
10.8 mm.
ACHATINELLA APEXFULVA IRWINI Pilsbry and Cooke
PLATE 2, FIGURE 29; PLATE 8, FiGuRES I3-I4a
Achatinella leucorraphe irwini Pirspry and Cooke, Man. Conch., vol. 22, pp. 302,
303, pl. 50, fig. 11a (only), 1914.
The lectotype of A. a. irwini selected by Pilsbry and myself is
figured in the Manual of Conchology on plate 59, figure 11a, and is
reproduced in this paper on plate 8, figure 13. The embryonic whorls
of the lectotype are bicolored, first embryonic whorl warm sepia, re-
maining embryonic whorls upper half or fourth cartridge buff or
white, lower portion lightens from warm sepia to cream buff on the
last whorl; postembryonic whorls white, lined with one or two
faint lines of ochraceous buff, last whorl lined at the edge of the
periphery with a line of grayish olive, below the periphery a line of
light buff, below which are three lines and two bands of warm
blackish brown and a basal band of ochraceous buff, and one of warm
blackish brown; impressed sutural band warm sepia, shading to bister
on the penultimate and first half of the last whorl, on the last half the
impressed sutural band is white, the upper edge lined with a line of
sayal brown, the lower edge warm blackish brown; lip pale vinaceous
fawn. Length 19.0 mm., greater diameter 12.2 mm., spire height
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 125
9.6 mm., number of whorls 6. The color pattern of the lectotype is
a characteristic one of the North-South Kaukonahua Ridge.
Distribution, area 61?: North-South Kaukonahua Ridge, locality
275?, approximate elevation 1,650-1,810 feet, Meinecke, 1923 (figs. 5,
5a, p. 105). Also collected by Wilder, BBM 50576. Lectotype, ANSP
108778. Type locality, the “upper part of the Kaukinehua [Kauko-
nahua] ridge” (Pilsbry and Cooke, 1914), collected by I. Spalding.
In the Wilder collection there is a series of 22 shells, all sinistral,
which probably come from the region of the type locality or the
locality of the lectotype. Wilder collected with Spalding, and it is
probable that the Wilder lot is from the same place. The shells in
the Wilder lot have a uniform color pattern which may mean either
that Wilder selected out his color patterns, or that the lot is from a
localized area of fairly limited extent. I am inclined to believe it is
from a small area because the shells are very similar to those obtained
by Meinecke on this ridge in area 61?.
The usual form of the shell in the Wilder lot, BBM 50576 (pl. 2,
fig. 29), and the color pattern on 59 percent of the shells has the
embryonic whorls bicolored, first whorl cinnamon drab, next whorl
upper half white, lower half cinnamon drab, last embryonic whorl
white; postembryonic whorls white, lined just above or at the edge
of the periphery with sayal brown, last whorl and a half spirally lined
with sayal brown and cinnamon buff; lip and columella callus pale
vinaceous fawn. Length 18.5 mm., greater diameter 12.3 mm., spire
height 9.7 mm.
A variation of the typical pattern (pl. 8, fig. 14) found on three
shells is a light pattern; postembryonic whorls white, last whorl
at the edge of the periphery faintly banded with pale pinkish buff
and cinnamon buff. Another characteristic color pattern (pl. 8,
fig. 14a) occurring on 27 percent has the postembryonic whorls white,
last whorls faintly spirally lined and banded with pale pinkish buff, last
whorl lined at the edge of the periphery and banded and lined below
the periphery with hay’s maroon; impressed sutural band, on the
first postembryonic whorl and a half russet, on the last whorl and a
half light buff with a line of russet on the lower half of the band.
ACHATINELLA APEXFULVA IRWINI var. 1
PLATE 2, FIGURE 30; PLATE 8, FiGuRES 15-15)
Achatinella leucorraphe irwini Pirspry and Cooxe, Man. Conch., vol. 22 ,pp. 302,
303, pl. 59, figs. 10, 11, 11b (only), 1914.
Area 62?: North-South Kaukonahua Ridge, locality 277?, ele-
vation 1,750-1,900 feet, Meinecke, BBM 122671, 1923, BBM 121973-
121974, 1918 (figs. 5, 5a, p. 105).
126 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
The usual form of the shell and the form and color pattern of the
embryonic whorls are close to A. a. irwini. The usual color pattern
of the postembryonic whorls is almost identical with the color pattern
of the holotype of A. a. ewaensis. The two forms of A. a. ewaensis
are separable because this form does not have a yellow band on the
embryonic whorls and has a different range of color patterns.
The usual form (pl. 2, fig. 30) and color pattern on 50 percent of
the sinistral shells has the embryonic whorls bicolored, first two
embryonic whorls white, banded on the lower third of the whorl with
fuscous; postembryonic whorls white faintly tinted with pale gull
gray, last whorl banded and lined with seal brown; lip and columella
callus pale grayish vinaceous. Length 18.5 mm., greater diameter
12.6 mm., spire height 9.0 mm. A variation of the usual form is a
broad banded color pattern (pl. 8, fig. 15) with the same color as
plate 2, figure 30. Eighteen percent of the sinistrals have a white
color pattern tinted with gray similar to plate 8, figure 6a. Twelve
percent may have olive buff or yellow lines similar to A. a. irwint
mixed with the usual pattern.
The usual form of a dextral (pl. 8, fig. 15a) and color pattern on
50 percent of the dextral shells has the postembryonic whorls white,
last whorl faintly lined at the edge of the periphery with pale gull
gray, base lined with seal brown. Length 18.2 mm., greater diameter
12.5 mm., spire height 9.3 mm. Twenty-two percent have the banded
pattern of plate 8, figure 15, 11 percent have a white pattern similar
to plate 8, figure 6a. Seventeen percent have an unusual dark banded
pattern (pl. 8, fig. 15b); last two postembryonic whorls dark livid
brown banded with pale vinaceous fawn, the bands lined with wood
brown or vinaceous buff.
The dextrals do not contain any forms with olive buff or yellow
in them similar to A. a. irwint. For this reason I believe the dextrals
probably came from a higher locality than the sinistrals and prob-
ably occur as a pure race of dextrals. The sinistrals may also occur
as a pure white banded race without any olive buff or yellow lines on
the shell. This section of the North-South Kaukonahua Ridge is not
well known and should be carefully collected again.
Group OF A. A. CONIFORMIS GULICK
ACHATINELLA APEXFULVA CONIFORMIS Gulick
PLATE 2, FiGuRE 34; PLATE 9, Figures 18-20a
Apex coniformis GuLtcK, Proc. Zool. Soc. London, 1873, p. 80, pl. 9, fig. 17.
Achatinella swiftii NEwcoms, Pilsbry and Cooke, Man. Conch., vol. 22, pp. 306,
308, 309, 312, pl. 58, figs. 2, 2a, 7; pl. 59, figs. 1a, 1b, 2(?); pl. 60, fig. 10a
(only), 1914.
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 127
Achatinella apexfulva apicata (Newcomb) Pretrrer, Pilsbry and Cooke, Man.
Conch., vol. 22, pp. 325-326, 327, pl. 60, figs. 5(?), oa, 1914.
The holotype of A. a. coniformis (pl. 9, fig. 18) has the first
embryonic whorl and a half white, remaining embryonic whorls light
buff; postembryonic whorls pinkish buff axially streaked with
avellaneous, last whorl also spirally lined with avellaneous, last 6.4
mm. of the last whorl mikado brown; impressed sutural band russet ;
lip and columella callus pale cinnamon pink. Length 17.5 mm., greater
diameter 12.7 mm., spire height 8.7 mm., number of whorls 63.
Distribution, area?: Type locality Ahonui, Gulick. Holotype,
MCZ 39911. Also Kalaikoa, Wahiawa, Helemano, Gulick. These
last three localities are undoubtedly an error because all subspecies
of A. apexfulva found today at a low elevation in the region between
South Kaukonahua Stream and Helemano Stream are highly localized.
The area occupied by a single race usually covers less than a mile in
extent along the top of a ridge or bottom of a gulch, and never extends
over two main ridges. Gulick and Smith named a series of species
such as contformuis, flavida, versicolor, gulickui, and lilacea which are
undoubtedly distinct subspecies from highly localized localities. How-
ever, in the Gulick collection specimens of these very distinct forms
are reported from a number of widely separated localities. This indi-
cates that the Gulick collection is a mixture of shells from various
localities. This may be accounted for by the fact that most of Gulick’s
material was obtained from natives.
The typical color pattern of A. a. coniformts is exceedingly rare
in the Gulick collection in the Bishop Museum, only four juvenile
specimens being found. The lots from Ahonui containing these speci-
mens also include a diversity of patterns that belong to other sub-
species such as A. a. gulickii and A. a. lilacea.
An elongate form of the shell with a light shade of the typical color
pattern (pl. 9, fig. 19) has the embryonic whorls pale pinkish buff ;
postembryonic whorls a yellowish hue of avellaneous, spirally lined
above the periphery with tilleul buff and fawn color; impressed
sutural band mikado brown. A darker shell of the typical color
pattern (pl. 9, fig. 19a), also a juvenile, has the embryonic whorls
light buff darkening to pinkish buff ; postembryonic whorls avellaneous
spirally lined and axially streaked with cameo brown, last whorl
finely axially streaked with verona brown and spirally lined with
light pinkish cinnamon. A specimen that looks like a brown form of
A. a. gulicku has the first postembryonic and penultimate whorls,
above the wood brown supraperipheral band, light drab spirally lined
and finely axially streaked with white, last whorl wood brown spirally
9
128 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
lined with white or tilleul buff and axially streaked with benzo brown,
peripheral band white; lip and columella callus light pinkish cinnamon.
Mixed with other lots of shells of various subspecies from Ahonui,
Kalaikoa, and Wahiawa, a group of distinct color patterns occur which
resemble A. a. coniformis. All shells similar to coniformis are mark-
edly ornamented with fine spiral lines and weakly streaked with fine
axial streaks. These forms (pl. 2, fig. 34; pl. 9, figs. 20, 20a) prob-
ably came from a single locality or from the same general area as the
holotype of A. a. coniformis. The holotype is probably a rare color
pattern of the subspecies.
In the usual form and color pattern (pl. 2, fig. 34) of A. a.
coniformis the embryonic whorls shade from white to light buff;
first three-fourths postembryonic whorl white axially streaked with
brownish drab, next whorl and a fourth dark gray axially streaked
and faintly spirally lined with white, last whorl light drab, shading to
snuff brown on the last fourth whorl, axially streaked and spirally
lined with white; impressed sutural band on the first postembryonic
whorl the same as on the rest of the whorl, on the remaining whorls
white ; lip not completely thickened, color avellaneous ; columella callus
white tinted with pale vinaceous fawn. Length 17.8 mm., greater
diameter 12.5 mm., spire height 9.8 mm., number of whorls 64.
The color pattern may be more brownish (pl. 9, fig. 20) and the
form more elongate than the usual form; embryonic whorls white;
first postembryonic whorls, up to the last whorl, tilleul buff axially
streaked with pallid mouse gray, last whorl a yellowish hue of vina-
ceous buff spirally lined with white; impressed sutural band same as
rest of whorl; lip light vinaceous fawn. Length 18.3 mm., greater
diameter 11.7 mm., spire height 9.4 mm. An obese and strongly
lineate form (pl. 9, fig. 20a) has the first two embryonic whorls
white, last embryonic whorl cream color; first postembryonic whorl
vinaceous drab, lined with white, penultimate whorl light drab spirally
lined and axially streaked with white, last whorl drab spirally banded
above the periphery with a band of white which is lined with four
lines of drab, below the periphery ground lined and streaked with
white ; lip and columella callus light vinaceous fawn. Length 17.4 mm.,
greater diameter 13.2 mm., spire height 9.4 mm.
ACHATINELLA APEXFULVA APEXALBA, new subspecies
PLATE 2, FIGURE 35; PLATE 9, FiGuRES 26-26d
Achatinella swiftiti Newcoms, Pilsbry and Cooke, Man. Conch., vol. 22, pp. 308,
309, 312, pl. 58, figs. 1, 2b, 2c, 16; pl. 50, fig. 1 (only), 1914.
The shell is usually sinistral and resembles the form of A. a.
versicolor, but differs from A. a, versicolor by lacking a white periph-
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 129
eral band and having a different range of color patterns. The shell
has characteristic white embryonic whorls and contrasting dark brown
postembryonic whorls. This subspecies is usually found mixed
with A.-a. coniformis in the Gulick lots although some lots of
apexalba contain no forms of A. a. coniformis. I have separated
apexalba from coniformis because of the fact that apevxalba has
a crisscross color pattern in which the spiral bands are broken
or flecked by dark and light axial streaks somewhat resembling A. a.
versicolor. The shell of apexalba has a rough, coarse, or broken
pattern, whereas coniformis has a solid, finely axially streaked even
color. A few specimens are found which might be put in either
subspecies. These are probably intermediate forms between the two
races.
The embryonic whorls of the holotype (pl. 2, fig. 35) are pale
pinkish buff; first two postembryonic whorls light drab axially
streaked with hair brown and benzo brown spirally lined with a
central line of white, on the penultimate whorl a white subsutural
band broken by dark axial streaks, last whorl above and just below
the periphery spirally banded with light drab and drab axially streaked
with snuff brown, flecked and spirally lined with white, last half
whorl darkens to snuff brown, base snuff brown spirally banded with
avellaneous and tilleul buff; impressed sutural band chestnut on the
first two postembryonic whorls, on the last whorl impressed sutural
band the ground color except the upper edge, which is fawn color; lip
and columella callus pale vinaceous fawn. Length 18.5 mm., greater
diameter 11.7 mm., spire height 10.0 mm., number of whorls 64.
Distribution, area?: Type locality Wahiawa, Gulick. Also Kalai-
koa, Ahonui, Gulick. The last two localities are probably wrong, if
Wahiawa is correct.
The darkest color pattern and elongate form of the shell (pl. 9,
fig. 26) measures: Length 17.9 mm., greater diameter 11.3 mm., spire
height 10.8 mm.; first two postembryonic whorls drab spirally banded
with fawn color and axially streaked with verona brown, last whorl
saccardo’s umber, banded with fawn color above the periphery, be-
low the periphery a band of white and lines of white; impressed
sutural band russet lightening in color at the extreme upper edge to
tawny.
A light color pattern (pl. 9, fig. 26a) has the first postembryonic
whorl benzo brown with a central white line, penultimate whorl
white axially streaked with hair brown and spirally lined with a central
band of white, last whorl tilleul buff, spirally lined with verona brown
and axially streaked with drab, base banded with a hue between benzo
130 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
brown and hair brown; impressed sutural band warm sepia lightening
to sayal brown on the last half whorl, subsutural band shades from
benzo brown to hair brown and verona brown on the last half whorl.
An extreme obese specimen (pl. 9, fig. 26b) has the embryonic
whorls white; postembryonic whorls drab axially streaked with hair
brown, and on the last whorl also streaked with tilleul buff, on the
last two whorls just above the edge of the periphery two lines of
white broken by the axial ornamentation, base weakly lined with
white; impressed sutural band russet lightening to tawny on the
last half whorl. Length 17.6 mm., greater diameter 13.0 mm., spire
height 9.2 mm.
A lighter gray color pattern (pl. 9, fig. 26d) is strongly axially
streaked on the last two postembryonic whorls; penultimate whorl
benzo brown, lined with pinkish buff and white and streaked with
pale smoke gray, last whorl pale smoke gray lightening to pale cinna-
mon pink, axially streaked with light drab, spirally banded with drab
below the sutural band, and lined with white above and below the
periphery. The color may be a very light brownish hue (pl. 9,
fig. 26c), penultimate whorl similar to that of figure 26d, last whorl
avellaneous lined with white.
ACHATINELLA APEXFULVA VERSICOLOR Gulick
PLATE 2, FIGURE 33; PLATE 9, FIGURES 21-23
Apex versicolor GuLick, Proc. Zool. Soc. London, 1873, p. 80, pl. 9, fig. 18.
Achatinella swiftii NeEwcoms, Pilsbry and Cooke, Man. Conch., vol. 22, pp. 306,
310, pl. 58, figs. 14-14d, 15 (only), 1914.
The holotype (pl. 9, fig. 21) has the first embryonic whorl and a
half cartridge buff, remaining embryonic whorls cream buff, lined
with white; first quarter postembryonic whorl upper half pale gull
gray, lower half banded and lined with chamois, remaining post-
embryonic whorls pale gull gray axially streaked with mouse gray
on the penultimate, just above the edge of the periphery a band of
bone brown or clove brown up to the last whorl, last whorl pale mouse
gray axially streaked with benzo brown, banded above the pale
mouse gray peripheral band, and all below, with hay’s brown, streaked
with pale mouse gray; impressed sutural band the ground color;
lip and columella callus pale vinaceous fawn. Length 18.8 mm.,
greater diameter 13.0 mm., spire height 9.5 mm., number of whorls 6.
Distribution, area?: Type locality, Ahonui, Gulick. Also Kalaikoa,
Gulick. This race probably occurred somewhere in the region of the
North or South Kaukonahua Stream.
NOS 5 ACHATINELLA APEXFULVA DIXON—WELCH I31
The usual form of A. a. versicolor (pl..2, fig. 33) measures:
Length 17.2 mm., greater diameter 11.6 mm., spire height 9.2 mm.,
number of whorls 6; embryonic whorls light buff, lined or banded on
the last whorl with white; first half postembryonic whorl pale gull
gray, spirally lined with benzo brown below the impressed sutural
band of benzo brown, just above the edge of the periphery a line of
sayal brown, last half of whorl pale gull gray with an axial streak
of snuff brown, first half of the penultimate pale gull gray axially
streaked with cinnamon buff, last whorl and a half white or faintly
shaded with pallid mouse gray, axially streaked with snuff brown
and the ground color, spirally banded or lined with snuff brown and
bister, below the edge of the periphery a band of white, base bister
axially streaked with white.
A light color pattern (pl. 9, fig. 23) has the postembryonic whorls
white or faintly shaded with pallid mouse gray, first postembryonic
whorl and a half has a subsutural line of mouse gray, last postembry-
onic whorl and a half spirally lined and axially streaked with mikado
brown and sayal brown, subperipheral band white, base banded with
warm sepia and a line of white. A rare pattern (pl. 9, fig. 22) has the
last postembryonic whorl above the periphery white shaded with pale
gull gray, with a patch or streak of snuff brown above the aperture
and on the last half of the whorl, the band at the edge of the periph-
ery, and all below the white subperipheral band, drab axially streaked
with benzo brown.
ACHATINELLA APEXFULVA VERSICOLOR var. 1
PLATE 9, FIGURES 24-25)
Achatinella swiftii NEwcoms, Pilsbry and Cooke, Man. Conch., vol. 22, p. 314,
pi So, es. 3, 3a, 11; pl. 50, fig: 2, 1014.
Achatinella apexfulva apicata (Newcomb) PreirFer, Pilsbry and Cooke, Man.
Conch., vol. 22, p. 325, pl. 60, fig. 5a (only), 1914.
Area 78?: Somewhere at a low elevation below area 79 probably
in area 78? (figs. 5, 5a, p. 105), on the North-South Kaukonahua
Ridge, Thurston, BBM 130693-130694, Judd, BBM 110053-110054.
Also collected by Gulick and mixed with subspecies of A. apexfulva
from Kalaikoa and Ahonui.
A. a. versicolor var. 1 is a race of bluish-gray shells with certain
color patterns that resemble those of A. a. versicolor, but most of its
patterns are quite distinct.
A characteristic color pattern (pl. 9, fig. 25) has the postembryonic
whorls white, banded just above the periphery and on the last whorl
132 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
above and below the periphery with light mouse gray lined with white ;
impressed sutural band mouse gray with a line of verona brown along
the upper edge; lip and columella callus pale vinaceous fawn. Length
19.0 mm., greater diameter 11.7 mm., spire height 10.0 mm.
A dark color pattern and an elongate shell (pl. 9, fig. 25a) measures:
Length 19.4 mm., greater diameter 12.1 mm., spire height 11.0 mm.
The postembryonic whorls are hair brown axially streaked with benzo
brown, and spirally lined with pale pinkish cinnamon or white faintly
shaded with pallid mouse gray; on the last half whorl the light lines
or bands are pallid mouse gray.
A sinistral shell (pl. 9, fig. 25b) resembles A. a. versicolor (pl. 9,
fig. 22). The postembryonic whorls below the white subsutural band
are benzo brown axially streaked with light drab, last whorl just above
and all below the periphery wood brown axially streaked and spirally
lined with tilleul buff. The shell may have a lined pattern (pl. 9,
fig. 24) ; embryonic whorls naples yellow, lined with white on the last
half whorl; postembryonic whorls lined with natal brown; lip and
columella callus white, with the outer edge light vinaceous fawn.
The lightest color pattern and an obese dextral (pl. 9, fig. 24a)
measures: Length 18.0 mm., greater diameter 13.2 mm., spire height
9.6 mm. Postembryonic whorls white, banded at the suture and just
above the edge of the periphery with light drab, base drab gray,
lined with white.
ACHATINELLA APEXFULVA WAHIAWA, new subspecies
PLATE 2, FicurE 36; PLATE 11, Ficures 1-1b
Achatinella swiftti NEwcoms, Pilsbry and Cooke, Man. Conch., vol. 22, pp. 312,
313, pl. 58, fig. 8; pl. 59, fig. 2a (only), 1914.
Gulick called this race of shells A. pica. Pilsbry and Cooke correctly
consider the name pica a synonym of A. a. apexfulva. The form is
a distinct lowland one which probably occurred at some low elevation
below areas go and g1?? (fig. 5a, p. 105). A. a. wahiawa resembles
A. a. coniformis but differs in usually being more strongly axially
streaked, lacking pronounced fine spiral lines, and having white
embryonic whorls. The shell is usually dextral.
The holotype (pl. 2, fig. 36) has white embryonic whorls darkening
to pale pinkish buff on the last embryonic whorl; the first two and
three-fourths postembryonic whorls vinaceous buff axially streaked
with rood’s brown, last whorl avellaneous axially streaked and spirally
banded and lined with benzo brown and cameo brown; impressed
sutural band mikado brown darkening on the last half whorl to walnut
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 133
brown ; lip and columella callus white with a faint tint of seashell pink.
Length 17.8 mm., greater diameter 12.5 mm., spire height 9.6 mm.,
number of whorls 64.
Distribution, area?: Wahiawa, Gulick.
The spire may be more concave (pl. 11, fig. 1a), the embryonic
whorls more pointed, and the last whorl flattened and less rounded
at the edge of the periphery than the holotype; the color pattern is
similar to that of the holotype. A narrow specimen (pl. 11, fig. 1)
and light color pattern measures: Length 17.1 mm., greater diameter
12.0 mm., spire height 9.4 mm.; first half postembryonic whorl pale
pinkish buff axially streaked with warm buff, next whorl and a half
pale vinaceous fawn axially streaked with light drab, last whorl
seashell pink, axially streaked with vinaceous fawn and fawn color,
at the edge of the periphery and about the base a line of benzo brown,
in the umbilical region a patch of chamois.
A dark color pattern (pl. 6, fig. 1b) is figured, showing the dorsal
side of the shell, to exhibit the contrast between the white embryonic
whorls and the postembryonic whorls; the last half of the first post-
embryonic whorl and the penultimate whorl light cinnamon drab,
finely axially streaked with dark vinaceous brown, first half of last
whorl drab axially streaked with dark vinaceous brown, last half whorl
almost solid dark ‘vinaceous brown streaked with drab; impressed
sutural band walnut brown. Length 16.5 mm., greater diameter 12.6
mm., spire height 9.1 mm.
GrouP OF A. A. LILACEA GULICK
ACHATINELLA APEXFULVA GULICKII Smith
PLATE 10, FIGURES 14-15b
Apex gulickii SmitH, Proc. Zool. Soc. London, 1873, p. 78, pl. 9, fig. 19 (not
fig. 17).
Pee, apexfulva apicata (Newcomb) PFerrrer, Pilsbry and Cooke, Man.
Conch., vol. 22, pp. 325, 326, 327, pl. 58, fig. 10a (?); pl. 60, figs. 6, 6a, 12
(only), 1914. .
A. a. gulickit is characterized by having a white or light peripheral
band and a grayish or gray-brown color pattern. The holotype (pl. Io,
fig. 14) has the embryonic whorls cartridge buff; first half post-
embryonic whorl cinnamon buff darkening to cameo brown on the
last half of the whorl, penultimate whorl drab gray axially streaked
with benzo brown, last whorl pinkish buff above the peripheral band
of gull gray axially streaked with benzo brown and light drab, below
the periphery a band of benzo brown, below which is a band of pale
134 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
gull gray spirally lined with benzo brown, about the umbilicus a faint
band of cartridge buff; impressed sutural band on the first half
postembryonic whorl same as the ground color, on the remaining
whorls pecan brown lightening to cinnamon on the last whorl; lip
and columella callus light vinaceous cinnamon. Length 18.2 mm.,
greater diameter 12.3 mm., spire height 8.8 mm., number of whorls 6.
Distribution, area?: Type locality Kalaikoa, Gulick. Also reported
from Ahonui, Gulick. The type locality was probably a restricted
locality somewhere between North and South Kaukonahua Streams.
The color pattern of the holotype is an unusually dark gray pattern.
A more usual one in the Gulick lots (pl. 10, fig. 15) has cream-buff
embryonic whorls darkening to chamois on the last half embryonic
whorl; postembryonic whorls pale vinaceous fawn axially streaked
with vinaceous drab and pale brownish drab, peripheral band and
base white, subperipheral band light brownish drab ; impressed sutural
band pinkish cinnamon lightening on the following whorls to pale
ochraceous buff.
The darkest color pattern (pl. 10, fig. 15a) has the first two
embryonic whorls cartridge buff darkening to chamois on the last
embryonic whorl; postembryonic whorls pale quaker drab axially
streaked with anthracene purple, on the last whorl about the periphery
a line of white, last fourth whorl pinkish buff axially streaked with
cinnamon drab, lip not formed. Another shell has a yellowish pink
hue (pl. 10, fig. 155) ; postembryonic whorls up to the last whorl
vinaceous drab axially streaked with light pinkish cinnamon, last
whorl light pinkish cinnamon streaked with vinaceous drab, peripheral
band white ; impressed sutural band vinaceous cinnamon.
ACHATINELLA APEXFULVA GULICKII var. 1
PLATE 10, FIGURE 16, I6a
Achatinella apexfulva apicata (Newcomb) Preirrer, Pilsbry and Cooke, Man.
Conch., vol. 22, pp. 325, 326, pl. 60, figs. 11, 13 (only), 1914.
Area?: Ahonui, Gulick.
This light pinkish variety looks like an intermediate race between
A. a. gulicku and A. a. lilacea, and is considered a variety of A. a.
gulickw because of the light band just below or at the edge of the
periphery of the last whorl.
The usual form and color pattern (pl. 10, fig. 16) has the embryonic
whorls cartridge buff ; first postembryonic whorl and a half vinaceous
fawn axially streaked with pale brownish drab, last whorl and a half
pale ecru drab axially streaked with white and faintly banded above
NOS ACHATINELLA APEXFULVA DIXON—WELCH 135
the periphery with pale vinaceous fawn, on the last whorl a peripheral
band of white, base white, subperipheral band pale ecru drab; im-
pressed sutural band pale vinaceous fawn; lip and columella callus
white faintly tinted with pale vinaceous fawn. Length 19.0 mm.,
greater diameter 12.8 mm., spire height 10.4 mm., number of
whorls 63.
The color may be light pink (pl. 10, fig. 16a) and the color pattern
may resemble that of A. a. lilacea (pl. 10, fig. 10), although differing
in having the last whorl, just below the periphery and at the base,
banded and lined with white.
ACHATINELLA APEXFULVA GULICKII var. 2
PLATE 10, FIGURES 21-210
Achatinella swiftii Newcomes, Pilsbry and Cooke, Man. Conch., vol. 22, pp. 300,
312, 315, pl. 58, figs. 13-13); pl. 50, fig. 4e (only), 1014.
Area?: Kalaikoa, Gulick.
The form was described by Smith as a variety of A. a. gulickit.
It is probably an intermediate race between A. a. gulickii var. 1 and
A. a. flavida. The postembryonic whorls are of a pinkish color above
the periphery, last whorl at and below the periphery banded with
yellow.
The usual color pattern (pl. 10, fig. 21) on 37 percent of the
shells has the embryonic whorls white, shading to cream buff on
the last half embryonic whorl; first half postembryonic whorl vina-
ceous fawn with a central line of pale pinkish cinnamon, the remain-
ing postembryonic whorls vinaceous pink, lined and axially streaked
with pale pinkish cinnamon, the last whorl is tinted with chamois,
a band of fawn color below the pale pinkish cinnamon band just
below the edge of the periphery, base cream buff tinged with chamois ;
impressed sutural band pale pinkish cinnamon; lip and columella
callus a very dilute tint of a pink or tilleul buff.
Thirty-three percent have a lighter or yellow color pattern (pl. 10,
fig. 21a); the first postembryonic whorl and a half pale drab gray
axially streaked with light cinnamon drab, first half of last whorl
above the periphery pale drab gray tinged with chamois, last half
almost entirely chamois, base chamois below the peripheral band of
chamois, subperipheral band a faint shade of drab gray covered over
by the ground color; impressed sutural band pale pinkish cinnamon.
A sinistral specimen (pl. 10, fig. 21b) shows the dark form of
30 percent of the shells; postembryonic whorls above the periphery
fawn color; impressed sutural band fawn color; subsutural band or
136 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
line on the last two whorls white, shading to chamois on the last half
whorl; the last half whorl is tinted with chamois, last whorl below
the narrow peripheral band of white is chamois with a central band
of fawn color.
ACHATINELLA APEXFULVA FLAVIDA Gulick
PLATE 10, FIGURES 17-20
Apex flavidus Gutick, Proc. Zool. Soc. London, 1873, p. 80, pl. 10, figs. I, Ia.
Achatinella swiftti Newcoms, Pilsbry and Cooke, Man. Conch., vol. 22, pp. 306,
311, 312, 315, pl. 58, figs. 10, 12-12b (only), 1914.
The subspecies is characterized by having a light yellow color pat-
tern. The form of the shell resembles A. a. gulickw var. 2. The
holotype is a rare color pattern and not characteristic of the sub-
species. The holotype of A. a. flavida (pl. 10, fig. 17) has the first
embryonic whorls a very dilute shade of pale ecru drab, fading to
white ; postembryonic whorls white, first postembryonic whorl with
a cinnamon-buff supraperipheral line which widens on the last two
whorls to a band of mikado brown or walnut brown; the impressed
sutural band white on the first embryonic whorl, on the last two
whorls mikado brown darkening to cacao brown, base cartridge buff ;
columella callus pale pinkish cinnamon; lip cartridge buff. Length
19.8 mm., greater diameter 12.8 mm., spire height 10.7 mm., number
of whorls 64.
Distribution, area?: Type locality Kalaikoa, Gulick (probably
somewhere in South Kaukonahua Stream at a low elevation). Also
Ahonui, Gulick, which is probably erroneous if Kalaikoa is correct,
because the type locality was probably of limited extent.
Gulick’s var. a (pl. 10, fig. 18) has a more usual color pattern; first
embryonic whorl and a half white, shading to cream buff on the later
embryonic whorls; first two postembryonic whorls pinkish buff
axially streaked with cinnamon buff and mikado brown; below the
impressed sutural band of tawny, which darkens on the last two
whorls and a half to orange cinnamon, there is a band of white; on
the penultimate whorl there is another band of white just above the
edge of the periphery, last whorl chamois spirally banded with sayal
brown. Length 17.6 mm., greater diameter 12.2 mm., spire height
9.3 mm., number of whorls 6. This color pattern occurs on 34 percent
of the combined Kalaikoa and Ahonui lots, all but 7 of which are from
Kalaikoa.
The usual color pattern in the Bishop Museum lots of the Gulick
collection (pl. 10, fig. 19) has the embryonic whorls white, post-
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 137,
embryonic whorls white up to the last whorl, last whorl above the
periphery cream buff, with a band made up of fine drab-gray lines
which darken on the last half whorl to cinnamon drab, above the band
a line of avellaneous, below and at the edge of the periphery chamois,
banded about the center of the base with two bands or lines of natal
brown; impressed sutural band russet ; lip and columella callus a very
dilute shade of vinaceous fawn.
The shell may not be strongly ornamented with gray or brown
lines or bands (pl. 10, fig. 19a), the first postembryonic whorl white
spirally lined with warm sepia, penultimate whorl white, last whorl
mustard yellow faintly banded with bands slightly darker than the
ground; impressed sutural band on the first two postembryonic
whorls russet, on the first half of the last whorl white with upper edge
russet, on the last half whorl white. This color pattern occurs on
23 percent of the shells.
A variation of the usual color pattern (pl. 10, fig. 19b) has the first
postembryonic whorl and half army brown with a subsutural line of
white, and another line of white just above the periphery which
broadens on the last half of the penultimate to a band, so that the
whorl is white with a median band of army brown on the last half
of the penultimate, last whorl, above the white peripheral band or line,
banded with a white subsutural band, a band of army brown, and a
band of chamois, on the last half whorl subsutural band tinted with
chamois, below the periphery the ground chamois, with a central
line of army brown.
In other lots of shells from Ahonui and Kalaikoa sometimes marked
by Gulick “tuberans-tumefacta,” is a white shell (pl. 10, fig. 20)
which is banded on the postembryonic and on the impressed sutural
band with pecan brown. I believe that this rare color pattern is a
light color pattern of flavida which lacks the usual yellow color.
Pilsbry and Cooke (pl. 58, fig. 10) figured a sinistral shell with this
color pattern.
ACHATINELLA APEXFULVA LILACEA Gulick
PLATE 3, FIGURE 2; PLATE 10, Ficures 8-11
Apex lilaceus GuticK, Proc. Zool. Soc. London, 1873, p. 70, pl. 10, fig. 4.
Achatinella apexfulva apicata (Newcomb) Preirrer, Pilsbry and Cooke, Man.
Conch., vol. 22, pp. 325, 328, pl. 60, figs. 14, 14a (only), 1914.
The holotype of A. a. lilacea (pl. 10, fig. 8) has the embryonic
whorls white ; postembryonic whorls white axially streaked with light
vinaceous fawn darkening to vinaceous fawn on the last whorl;
138 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
impressed sutural band white; lip and columella callus light vinaceous
fawn. Length 20.2 mm., greater diameter 13.2 mm., spire height
10.7 mm., number of whorls 63.
Distribution, area 83b??: Type locality Kalaikoa, Gulick; also
Ahonui, Gulick. The type locality containing a pure race of pink
shells of the typical A. a. lilacea pattern has not been located in
recent years. Meinecke in locality 280* (fig. 5, p. 105), North Kau-
konahua-Poamoho Ridge, BBM 121982, found two specimens of
typical A. a. lilacea mixed with A. a. lilacea var. 1. This indicates
that Gulick’s type locality was probably somewhere in the region of
North Kaukonahua Stream which was all or part of Gulick’s Ahonui,
and not in Kalaikoa. The possible region of the type locality area
83b?? has been plotted on figure 7, (p. 194).
The usual form of A. a. lilacea (pl. 3, fig. 2) has the embryonic
whorls pale pinkish buff; postembryonic whorls seashell pink axially
streaked with buff pink; impressed sutural band white; lip and
columella callus shell pink. Length 18.7 mm., greater diameter 12.2
mm., spire height 10.0 mm., number of whorls 6. A narrow specimen
(pl. 10, fig. 9) measures: Length 18.0 mm., greater diameter 11.8
mm., spire height 11.0 mm. An obese specimen (pl. 10, fig. 10)
measures: Length 18.2 mm., greater diameter 12.8 mm., spire height
10.0 mm. The color pattern of figures g and Io is similar to that of
plate 3, figure 2, except that the impressed sutural band is the same
as the rest of the whorl.
One of the Meinecke pink shells (pl. 10, fig. 11) from locality 280*
has the embryonic whorls white; postembryonic whorls pale salmon
color ; impressed sutural band the same as the rest of the whorl.
ACHATINELLA APEXFULVA LILACEA var. 1
PLATE 3, FIGURE 3; PLATE 10, FIGURES 12-13
Area 83: North Kaukonahua-Poamoho Ridge, locality 280-1-
280*, approximate elevation 1,450 feet, Russ, 1929-1933 ; 280%, eleva-
tion 1,350-1,450 feet, Meinecke, 1932. Also collected by Wilder,
BBM 50585.
Area 83a: North Kaukonahua-Poamoho Ridge, locality 281-2*,
elevation 1,500 feet, 1932, BBM 121987, 121988; 281-1, elevation
1,500-1,550 feet, Meinecke, 5 dextral 1933 (figs. 5, 5a, p. 105). Lo-
cality 281-2*, plotted from memory, probably should be below locality
281-1.
The shell is very similar in form and color pattern to typical
A. a. lilacea, differing in having a bluish-gray hue instead of a pinkish
color pattern. This race is probably intermediate between A. a. lilacea
and A. a. lilacea var. 2.
NOT ACHATINELLA APEXFULVA DIXON—WELCH 139
The usual color pattern and form of the shell (pl. 10, fig. 12) has
the embryonic whorls white; postembryonic whorls pale vinaceous
fawn axially streaked with light vinaceous fawn on the first post-
embryonic whorl and light cinnamon drab on the penultimate, last
whorl ecru drab axially streaked with cinnamon drab and benzo
brown and faintly spirally lined and axially streaked with tilleul buff ;
impressed sutural band same as the rest of the whorl, upper edge
white; lip and columella callus pale grayish vinaceous. Length 19.0
mm., greater diameter 12.0 mm., spire height 10.9 mm.
An obese shell (pl. 10, fig. 12a) measures: Length 17.8 mm.,
greater diameter 12.7 mm., spire height 9.1 mm.; shows a very light
color form; first half postembryonic whorl pale pinkish buff, next
whorl and a quarter up to the last whorl light vinaceous fawn axially
streaked with a dilute shade of vinaceous fawn, last whorl pale ecru
drab axially streaked with cinnamon drab and tilleul buff on the last
whorl ; impressed sutural band white. White sutural bands occur on
one or two specimens in each lot.
In the Russ lot from locality 280-1-280*, 66 percent are pinkish
forms (pl. 10, fig. 12) and 36 percent bluish-gray forms (pl. 10,
fig. 12b; pl. 3, fig. 3). In the Meinecke lot from 280*, 56 percent are
pinkish forms and 44 percent bluish pink. Locality 281-2*, has 71
percent bluish forms, and 281-1 contains no pink forms. These last
two localities have been separated out into area 83a to emphasize the
trend toward bluish-gray color forms above area 83. More collecting
should be done in this region to determine whether bluish-gray shells
exist in a pure locality below area 84 where the streaked color pattern
changes but the color is definitely bluish gray.
The usual color pattern and form of a bluish-gray shell (pl. 3, fig. 3)
has the embryonic whorls pale pinkish buff ; first postembryonic whorl
pale pinkish buff, last two whorls pale mouse gray axially streaked
with quaker drab, sorghum brown, and tilleul buff, last fourth whorl
tilleul buff axially streaked with avellaneous ; impressed sutural band
tilleul buff axially streaked with vinaceous buff; lip and columella
callus light vinaceous fawn. Length 19.0 mm., greater diameter 12.0
mm., spire height 10.5 mm. The color of the sutural band is unusual.
The color pattern may be lighter, the form elongate and the im-
pressed sutural band white (pl. 10, fig. 12b), which is usual; embry-
onic whorls white; first half postembryonic whorl white, last half
white axially streaked with light vinaceous fawn, penultimate and last
whorl pale drab gray axially streaked with pale brownish drab and
light brownish drab; lip and columella callus light vinaceous fawn.
140 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Length 19.0 mm., greater diameter 12.0 mm., spire height 10.8 mm.
Plate 10, figure 13, shows a sinistral shell from area 83a which has
the same color pattern as plate 10, figure 12b.
ACHATINELLA APEXFULVA LILACEA var, 2
PLATE 3, FIGURE 4
Area 84: North Kaukonahua-Poamoho Ridge, locality 282-1,
elevation 1,600 feet, 3 dextral; 282-3, elevation 1,550 feet, 10 dextral,
all collected by Meinecke, 1933 (figs. 5, 5a, p. 105).
Out of 13 shells from area 84, 3 have pink color patterns similar
to the shells of area 83. The remainder have a bluish-gray color
pattern which differs from the forms of area 83a by having a yellowish
ground color and the last whorl more strongly spirally lined.
The usual color pattern (pl. 3, fig. 4) has the embryonic whorls
light buff; first half postembryonic light buff axially streaked with
cinnamon buff, next whorl light cinnamon drab, last whorl and a half
light buff axially streaked and faintly spirally lined with hair brown,
also streaked with benzo brown; impressed sutural band light buff ;
lip and columella callus white, lip edged with avellaneous.
ACHATINELLA APEXFULVA GLAUCOPICTA, new subspecies
PLATE 3, FIGURE 10; PLATE 10, FIGURES 22, 22a
Achatinella apexfulva cervixnivea pattern Pilsbry and Cooke, Man. Conch.,
vol. 22, pp. 324, 328, pl. 60, fig. 9 (only), 1914.
The shell resembles the blue-gray color pattern of A. a. lilacea var. 1
but differs in having a sutural band which is always white, more
bluish postembryonic whorls, stronger blue-gray axial streaks, and
the ground pale gray instead of pinkish.
The embryonic whorls of the holotype (pl. 3, fig. 10) are white,
postembryonic whorls white axially streaked with pale neutral gray
and deep neutral gray; impressed sutural band white; lip and
columella callus pale grayish vinaceous. Length 19.4 mm., greater
diameter 13.1 mm., spire height 10.4 mm., number of whorls 64. The
usual form was not determined for lack of specimens.
Distribution, area 87: Central Poamoho Stream, type locality
280K-2, elevation 1,250 feet, Meinecke, 1933; Central Poamoho-
Central Poamoho North Branch Ridge, 280L-1, elevation 1,450-
1,500 feet, 5 dextral and 1 sinistral, Meinecke, 1933 (figs. 5, 5a,
p. 105).
NO. I ACHATINELLA APEXFULVA DIXON—WELCH I4I
The darkest color pattern (pl. 10, fig. 22) has the postembryonic
whorls plumbeous axially streaked with white, dark violet gray, and
blackish violet gray, last whorl almost solid blackish violet gray
axially streaked by the above lighter shades of gray ; impressed sutural
band white; lip and columella callus light vinaceous fawn.
The usual form of a sinistral, shown on plate 10, figure 22a, has
a color pattern similar to plate 3, figure 10, except that the dark axial
streaks are finer and resemble those of A. a. lilacea var. 1 (pl. 3,
fig. 3).
ACHATINELLA APEXFULVA PUNICEA, new subspecies
PLATE 2, FIGURE 26; PLATE 10, Ficures 6-6¢
The shape is similar to the probably lower race of A. a. steelt,
but the color pattern is different. The color pattern resembles that
of A. a. lilacea, but the range of color pattern and the form of the
shell make it distinct from lilacea, which is a much smaller shell. The
embryonic whorls of the holotype (pl. 2, fig. 26) are white, shading
to pale pinkish buff on the last embryonic whorl; postembryonic
whorls vinaceous buff axially streaked with avellaneous and tilleul
buff ; impressed sutural band light buff, tinting to pale pinkish buff
on the last half whorl; lip and columella callus light vinaceous fawn
lightening to white within. Length 19.0 mm., greater diameter 13.3
mm., spire height 9.6 mm., number of whorls 6.
Distribution, area 81°?: Kaukonahua, Wilder, BBM 50583,
50581 ; also collected by O. P. Emerson on “Head gate road ridge
between N. & S. br. of Kaukonahua,” in the collection of the MCZ.
The type locality is not known exactly. Odd specimens with the
usual A. a. punicea pattern are found with A. a. steel in locality 272.
In the type lot a few blue-gray shells are found which are similar to
A. a. punicea var. 1 (area 82?). The subspecies A. a. punicea probably
exists somewhere between areas 80 and 82?, probably near the bottom
of North Kaukonahua Stream, in the region of area 81°? (fig. 5a,
p. 105).
An obese shell (pl. 10, fig. 6) measures: Length 19.4 mm., greater
diameter 13.5 mm., spire height 8.9 mm.; embryonic whorls light
buff ; postembryonic whorls wood brown faintly axially streaked with
tilleul buff, about the periphery a band of tilleul buff ; impressed sutural
band a hue between cinnamon and avellaneous ; lip and columella callus
pale vinaceous fawn. The pink color pattern of the holotype and
plate 10, figure 6 occurs on 59 percent of the shells.
The sutural band may be white, the form elongate, and the color
pattern a bluish pink (pl. 10, fig. 6a) intermediate between A. a.
142 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
punicea and A. a. punicea var. 1. The embryonic whorls are white ;
postembryonic whorls pale ecru drab axially streaked and faintly
finely lined with pale brownish drab; lip and columella callus pale
vinaceous fawn, the outer margin vinaceous fawn. Length 20.5 mm.,
greater diameter 13.0 mm., spire height 10.8 mm. This pattern is
present on 39 percent of the shells. One specimen in the type lot is
almost entirely white (pl. 10, fig. 6b), last whorl shaded with axial
streaks of pale salmon color. In lot BBM 50581, out of seven shells,
three are pinkish forms and three are bluish pink, and one has a
yellowish pattern (pl. 10, fig. 6c); embryonic whorls light buff ;
postembryonic whorls light buff axially streaked with a dilute tint of
pinkish cinnamon.
ACHATINELLA APEXFULVA PUNICEA var. 1
PLATE 2, FIGURE 27; PLATE 10, FiGuRES 7-75
Area 82°: North-South Kaukonahua Ridge, Lemke, BBM
115039, 1932, exact locality not accurately plotted but probably some-
where in the region of area 82?, but may have been collected at a
higher or a lower elevation; also collected by Wilder, BBM 50578,
but the shells have no locality label. Probably collected in or near
area 82?, or in North Kaukonahua Stream (fig. 5a, p. 105).
The shell resembles A. a. punicea in form and size but differs in
having a blue-gray color pattern similar to that of A. a. glaucopicta.
A shell with the characteristic color pattern (pl. 2, fig. 27) has the
first embryonic whorl and a half worn, colored cinnamon buff, remain-
ing embryonic whorls pale pinkish buff; first postembryonic whorl
white faintly axially streaked on the last half whorl with pale drab
gray, penultimate whorl pallid mouse gray, finely axially streaked
with white, and coarsely axially streaked on the last half with quaker
drab, last whorl pallid mouse gray axially streaked with deep quaker
drab, last fourth whorl hair brown axially streaked with fuscous;
impressed sutural band white. Length 20.5 mm., greater diameter
13.3 mm., spire height 11.1 mm. The usual form of the shell was
not obtainable owing to scarcity of material.
In the Wilder lot, BBM 50578, the lightest color pattern (pl. 10,
fig. 7) has a pinkish tint and occurs on 17 percent of the shells. The
postembryonic whorls shade from white to pale vinaceous fawn on
the last two whorls, faintly lined and axially streaked with pale drab
gray; impressed sutural band white. Length 18.5 mm., greater
diameter 13.1 mm., spire height 9.0 mm. A sinistral shell (pl. 10,
fig. 7a) has the embryonic whorls light buff; first postembryonic
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 143
whorl, below the white sutural and subsutural bands, light mouse gray,
last whorl and a half pale vinaceous fawn axially streaked and
faintly spirally lined with pale brownish drab; lip and columella
callus pale vinaceous fawn. Forty-seven percent of the Wilder lot
have yellow embryonic whorls. The darkest color pattern (pl. Io,
fig. 7b), found on two shells, has the embryonic whorls white; post-
embryonic whorls dark vinaceous drab axially streaked with drab
gray, last half whorl mostly drab gray axially streaked with dark
vinaceous drab; impressed sutural and subsutural bands white. The
subsutural band, as in all specimens of this variety, decreases in width
on each successive whorl so that it is lacking or merely a line on the
last whorl.
ACHATINELLA APEXFULVA STEELI, new subspecies
PLATE 2, FIGURE 25; PLATE 10, FIGURES 3-5c
Achatinella apexfulva apicata NEwcomps, Pilsbry and Cooke, Man. Conch., vol.
22, p. 325, pl. 60, figs: 5b, 10, 1014.
The color pattern of the shell resembles A. a. gulickii but differs in
being a larger shell with a dark reddish-brown color pattern. The
embryonic whorls of the holotype (pl. 2, fig. 25) are pale pinkish
buff; last half of the first postembryonic and penultimate whorls
chocolate axially streaked and spirally banded with cinnamon drab,
last whorl above the periphery light drab spirally lined and banded
with chocolate, subperipheral band tilleul buff, axially streaked with
light drab and on the last half whorl streaked with avellaneous, base
light drab axially streaked with white, spirally banded with natal
brown and bone brown; lip and columella callus fawn color lightening
to light vinaceous fawn on the inner margin. Length 19.3 mm., greater
diameter 13.0 mm., spire height 10.2 mm., number of whorls 6.
Distribution, area 80: North-South Kaukonahua Ridge, type
locality 271, elevation 1,450-1,532 feet, Russ, 1933, Steel BBM
129017-129018, 6 sinistral 1934; also 272, elevation 1,500-1,550 feet,
1 dextral, 7 sinistral, Steel and Welch, 1934. Also collected by Gulick,
O. H. Emerson, BBM 103972, Wilder, BBM 50577, 50580 (figs. 5,
5a, p. 105). This subspecies is named after Lt. Col. Charles L. Steel,
U.S. A., who helped me determine the distribution of certain forms
of Achatinella and generously gave me valuable material.
The obese form of the shell (pl. 10, fig. 3) measures: Length 19.3
mm., greater diameter 13.5 mm., spire height 10.0 mm.; embryonic
whorls pale pinkish cinnamon, lightening to white on the last embry-
onic whorl; first half postembryonic whorl cinnamon buff axially
10
144 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103 |
streaked with mikado brown, remaining whorls chocolate, faintly |
axially streaked with pinkish buff, spirally lined on the upper third —
of the whorl above the periphery with a line of light drab and cin- |
namon, on the last half whorl above the periphery the chocolate
ground color changing to buffy brown or olive brown axially streaked _
and banded with a band of bone brown, below the periphery a band |
of cartridge buff and a band of chocolate, the remainder of the base
cartridge buff spirally lined with drab gray and two bands of natal
brown in the umbilical region.
In the lot collected by Steel from locality 271 a gray color pattern
is found (pl. 10, fig. 4); postembryonic whorls light drab axially
streaked with hair brown, fuscous, benzo brown, and faintly spirally
lined with fine lines of fuscous and white, below the periphery of the
last whorl the ground white, shaded with pale gull gray and spirally
banded with a band and a line of light drab axially streaked with
fuscous, in the umbilical region a line and a band of fawn color.
In the Wilder collection, mixed with color patterns of typical
A. a. steeli, are a series of color patterns which have the base
white or banded with white below the periphery and contain dextral
as well as sinistral shells. The shells may be typical A. a. steeli or
they may come from a different area.
The usual form of a dextral shell and a characteristic color pattern
in this lot (pl. 10, fig. 5) has the embryonic whorls cartridge buff ;
postembryonic whorls pecan brown, spirally ornamented on the last
whorl and a half -with a white band or line, below the impressed
pecan brown sutural band and in the center of the whorl above the
periphery, edge of the periphery banded with a wide white band, at
the upper edge of which is a line of pecan brown, remainder of the
base white with a central band and a line of pecan brown, in the
umbilical region a patch of fawn color; lip and columella callus pale
vinaceous fawn, outer margin spotted with fawn. Length 20.0 mm.,
greater diameter 13.0 mm., spire height 10.6 mm.
The base may be entirely white (pl. 10, fig. 5a) faintly shaded with
vinaceous fawn and lined with a line of army brown in the umbilical
region; the postembryonic whorls above the periphery are similar
to plate 10, figure 5, only more axially streaked and spirally lined
with white. A narrow sinistral specimen (pl. 10, fig. 5b) measures:
Length 18.4 mm., greater diameter 12.2 mm., spire height 9.7 mm.
An obese shell (pl. 10, fig. 5c) measures: Length 18.0 mm., greater
diameter 13.6 mm., spire height 9.0 mm. The shells of figures 5b and
5c have a similar color pattern to the shell of plate 10, figure 3.
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 145
® ACHATINELLA APEXFULVA STEELI var. 1
PLATE 2, FIGURE 24; PLATE 10, FIGURES I-2
Area 79: North-South Kaukonahua Ridge, locality 270, eleva-
tion 1,450 feet, 3 dextral, dead specimens, Steel and Welch, BBM
129012, 1934; “Mauka [toward the mountains] of the Burnt District,
Kaukonahua,” Thurston, BBM 130674. The burnt district is in the
region of area 78?, on the North-South Kaukonahua Ridge. Also
collected by Wilder, BBM 10446, Lemke, BBM 115036, Gulick, BBM
10445 (figs. 5, 54, p. 105).
This variety may be a distinct subspecies separable from A. a. steeli
by having a distinctly lighter color pattern. The Welch lot of three
shells indicates this possibility, but in all other lots the variety is
found mixed with A. a. steeli. Therefore, until more careful collecting
is done in area 79 and this race is definitely proved to be a distinct
race and not a light color pattern of the shells of area 80, this form
is considered a variety of A. a. steeli.
The usual form and color pattern of the shell (pl. 2, fig. 24) has
the embryonic whorls pale pinkish buff; the upper half of the first
two postembryonic whorls including the impressed sutural band cameo
brown, lower half white, banded just above the edge of the periphery
with a band of vinaceous buff lined with army brown, this band
fading to white on the last half of the penultimate, the last whorl
white, sutural and subsutural bands chocolate, subsutural band lined
with pale pinkish cinnamon, below the subsutural band a band of
tawny lined with russet, above the peripheral line of tawny the
white supraperipheral band shaded with cinnamon buff, below the
periphery the ground white banded with army brown, base finely
lined with pale brownish drab; lip outer margin vinaceous fawn,
inner margin and columella callus white. Length 18.8 mm., greater
diameter 12.8 mm., spire height 9.8 mm., number of whorls 63.
A narrow form with a narrow sutural band (pl. 10, fig. 1) has the
embryonic whorls white; postembryonic whorls white, banded below
the burnt umber sutural band with a band of white, burnt umber, and
sayal brown, on the last whorl the sayal brown band lightening to
cinnamon buff and fading out on the last half whorl, the white ground
shaded by faint bands of pinkish buff, base lined with fawn color.
Length 18.3 mm., greater diameter 12.0 mm., spire height 10.4 mm.
A sinistral specimen (pl. 10, fig. 1a) resembles the pattern of plate 2,
figure 24, on the first two postembryonic whorls; last whorl pale
pinkish cinnamon, below the subsutural bands of chocolate and
pinkish cinnamon, spirally banded at and above the periphery with
146 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
light buff which deepens to warm buff on the last whorl, Below the
periphery base banded with a band of cinnamon drab, benzo brown
and cream color in the umbilical region.
A possible intermediate pattern between A. a. versicolor var. 1 and
A. a. steeli var. 1 (pl. 10, fig. 1b) is seen on three specimens in the
Wilder collection; embryonic whorls white; postembryonic whorls
banded at the suture and below the suture by a wide band of chestnut
brown covering the upper half of the whorl, which is solid or lined
with drab gray, just above the edge of the periphery a band of pale
ecru drab axially streaked with white and spirally lined with light
brownish drab, just below the edge of the periphery a band of white
and below this a band of natal brown, base white, shaded with ecru
drab and lined with a line of light brownish drab.
In the Thurston lot there is a very light color pattern (pl. 10, fig. 2).
The embryonic whorls are pale pinkish cinnamon ; first postembryonic
whorl pale pinkish cinnamon, banded with bone brown, and cinnamon,
penultimate and last whorl white, banded above or at the edge of the
periphery with a narrow band of bone brown lightening to army
brown, above the peripheral band a faint band of cinnamon buff,
below the periphery the ground white lined and banded with avella-
neous ; lip and columella callus avellaneous lightening to white toward
the inner margin. Before the Thurston collection was given to the
Bishop Museum the cabinets containing the collection were moved
during a fire. In moving the cabinets they were tilted and shells
from different localities were mixed. Therefore the exact location
of this light color form should be checked.
Group oF A. A. APICATA NEWCOMB
ACHATINELLA APEXFULVA BRUNOSA, new subspecies
PLATE 3, FIGURE I1; PLATE 10, FIGURES 26-28
This species is similar to A. a. lilacea var. 1 (area 83) in form, in
the width of the axial streaks on the postembryonic whorls, and in
usually having a white sutural band. It differs in having the last
whorl and a half gray or gray brown. A. a. brunosa also closely
resembles A. a. apicata, but differs in the shade of brown of the
postembryonic whorls, and in usually having a white sutural band.
The first two postembryonic whorls of the holotype (pl. 3, fig. 11)
are cinnamon pink, deepening on the last embryonic whorl to
ochraceous buff, first half postembryonic whorl army brown axially
streaked with white, next whorl axially streaked with pale gull gray
and deep neutral gray, next half whorl finely axially streaked with
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 147
hair brown, and smoke gray, last whorl drab finely axially streaked
with hair brown; impressed sutural band upper two-thirds white,
lower third the color of the whorl below; lip and columella callus
light vinaceous fawn. Length 18.9 mm., greater diameter 12.3 mm.,
spire height 10.8 mm., number of whorls 6.
Distribution, area 88: Central Poamoho-Central Poamoho
North Branch Ridge, type locality 280L-2, elevation 1,550 feet; also
280L-3, elevation 1,600-1,650 feet, 2 dextral, 5 sinistral, Meinecke,
1933; 280L-4, elevation 1,700 feet; 280L-5, elevation 1,650 feet, 18
sinistral, all Meinecke-collected, 1933 (figs. 5, 5a, p. 105).
An extremely narrow shell (pl. 10, fig. 26) shows a lighter color
pattern ; first half postembryonic whorl army brown axially streaked
with white, next whorl axially streaked with hair brown, benzo
brown, white, and pale pinkish cinnamon, first half of last whorl
the ground above the periphery pale pinkish cinnamon darkening on
the last half whorl and below the periphery to a yellowish hue of
wood brown, the entire whorl axially streaked with snuff brown and
warm sepia ; impressed sutural band white. Length 18.5 mm., greater
diameter 11.6 mm., spire height 10.5 mm.
On one specimen the sutural band is found to be mikado brown
on the last two whorls. This obese shell (pl. 10, fig. 26a) resembles
more the form and color pattern of A. a. suturafusca var. 2 (pl. 10,
fig. 25b). The first two postembryonic whorls and the last whorl
above the periphery pallid mouse gray or mouse gray axially streaked
with dark purple drab, last whorl below the periphery snuff brown
on the last half of the whorl, last whorl below the periphery sayal
brown axially streaked with bister or benzo brown. Length 20.0 mm.,
greater diameter 13.4 mm., spire height 11.2 mm.
The usual form of a sinistral and a grayish pattern (pl. 10, fig. 27)
has the penultimate and first half of last whorl drab gray axially
streaked with pale drab gray or white, last half of last whorl light drab
darkening to drab axially streaked with hair brown and snuff brown,
just above the edge of the periphery a line of white. Length 18.2 mm.,
greater diameter 12.1 mm., spire height 9.9 mm.
The color pattern of a sinistral (pl. 10, fig. 28) shows the darkest
brownish color pattern with the usual fine axial streaks, of the sub-
species. The first half of the first postembryonic whorl cinnamon buff,
faintly axially streaked with army brown, last half livid brown
lightly axially streaked with pallid mouse gray, penultimate whorl
light mouse gray deepening to mouse gray, and axially streaked with
148 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
deep mouse gray, last whorl drab axially streaked with hair brown
and fuscous, last fourth whorl sepia.
ACHATINELLA APEXFULVA SUTURAFUSCA, new subspecies
PLATE 3, FIGURE 9; PLATE 10, FIGURES 24, 24a
Achatinella apexfulva apicata (Newcomb) Prerrrer, Pilsbry and Cooke, Man.
Conch., vol. 22, p. 326, pl. 60, fig. 7b (only), 1914.
The shell resembles the form of A. a. glaucopicta but has a brown
sutural band and yellow embryonic whorls. The first two embryonic
whorls of the holotype (pl. 3, fig. 9) warm buff deepening to
ochraceous buff on the last embryonic whorl; first half of first post-
embryonic whorl white, next whorl and a half lilac gray axially
streaked with violet gray, last whorl pale mouse gray axially streaked
with deep mouse gray and tilleul buff, and faintly spirally lined with
deep mouse gray with a spiral band of tilleul buff just above the edge
of the periphery, last fourth whorl tilleul buff axially streaked with
army brown; impressed sutural band chestnut; lip and columella
callus pale vinaceous fawn. Length 19.1 mm., greater diameter 13.5
mm., spire height 9.3 mm., number of whorls 64.
Distribution, area 86: Central Poamoho Stream, type locality
280K-1*, elevation 1,150-1,250 feet, Meinecke, 1932 (figs. 5, 5a,
p. 105). Also collected by O. H. Emerson, BBM 102289, Wilder,
BBM 50586. The Wilder shells are labeled North Kaukonahua, prob-
ably an error for Central Poamoho Stream.
The lightest color pattern (pl. 10, fig. 24), found on only three
specimens, has the first two postembryonic whorls white, the penulti-
mate axially streaked with pale drab gray, last whorl a dilute shade of
pale drab gray or white axially streaked with drab gray and faintly
spirally banded, below the chestnut sutural band, with a wide band
of a dilute shade of pale quaker drab, and a band of tilleul buff at
the edge of the periphery.
The usual form and color pattern of a sinistral shell (pl. Io,
fig. 24a) also exhibits the dark color pattern of a dextral shell, embry-
onic whorls ochraceous tawny ; first postembryonic whorl light neutral
gray axially streaked with white, penultimate whorl light neutral gray
axially streaked with neutral gray and deep neutral gray and finely
spirally lined with lines of deep neutral gray, last whorl mouse gray
axially streaked and faintly spirally lined with deep mouse gray;
impressed sutural band chestnut. Length 18.1 mm., greater diameter
12.3 mm., spire height 9.6 mm.
NO. I ACHATINELLA APEXFULVA DIXON-—WELCH 149
ACHATINELLA APEXFULVA SUTURAFUSCA var, 1
PLATE 3, FIGURE 1; PLATE I0, FIGURE 23
Achatinella apexfulva apicata (Newcomb) Pretrrer, Pilsbry and Cooke, Man.
Conch., vol. 22, p. 326, pl. 60, fig. 7a (only), 1914.
Area 85: South Poamoho Stream, locality 280-7, elevation
1,300-1,350 feet, Welch and C. W. Isle, 8 sinistral 1935 (figs. 5, 5a,
Bp. 105).
The form is close to typical A. a. suturafusca, but differs in having
a mars brown impressed sutural band, and the postembryonic whorls
not colored with as bluish-gray streaks as A. a. suturafusca. More-
over, the dark color pattern (pl. 10, fig. 23) is not known to me from
area 86.
The light color form (pl. 3, fig. 1) has the embryonic whorls tawny ;
first fourth postembryonic whorl tawny, the remaining whorls tilleul
buff or pale drab gray axially streaked with deep mouse gray and
mouse gray, the last two whorls faintly spirally lined with deep mouse
gray; lip and columella callus light grayish vinaceous. Length 19.8
mm., greater diameter 13.1 mm., spire height 10.6 mm. So few
specimens are known from this area that the usual form and color
pattern is not determined.
The dark color pattern (pl. 10, fig. 23) has the embryonic whorls
amber brown; postembryonic whorls clove brown or black axially
streaked with drab gray, last eighth whorl verona brown; impressed
sutural band bone brown or black. Length 18.1 mm., greater diameter
12.7 mm., spire height 9.0 mm.
ACHATINELLA APEXFULVA SUTURAFUSCA var. 2
PLATE 3, FIGURE 13; PLATE 10, FIGURES 25-25d
Achatinella apexfulva apicata (Newcomb) Preirrer, Pilsbry and Cooke, Man.
Conch., vol. 22, p. 326, pl. 60, figs. 4, 7(?) (only), ror4.
Area 89: North Poamoho Stream, locality 290*, elevation
1,300 feet, Meinecke, 1914, 1917, 4 dextral, 8 sinistral 1926; Wahi-
awa, Gulick, ANSP 92628 (figs. 5, 5a, p. 105).
This race is very similar to A. a. suturafusca (area 86) but differs
in usually having a more purplish-gray or darker color pattern.
Brownish color forms similar to A. a. apicata also occur in area 89g but
not in area 86. The form also resembles A. a. suturalba but differs in
having a brown sutural band. This variety is an intermediate one
between A. a. apicata and A. a. suturalba.
150 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
The usual form and color pattern (pl. 3, fig. 13) has the embryonic
whorls ochraceous tawny ; first fourth postembryonic whorl cinnamon
drab axially streaked with tilleul buff, next whorl and a fourth gull
gray axially streaked and finely lined with vinaceous slate, last half
of penultimate and first fourth of last whorl gull gray heavily axially
streaked and finely lined with dusky drab, the second fourth of last
whorl dark livid brown, last half whorl cinnamon drab axially streaked
with deep brownish drab and faintly spirally banded or tinged with
a faint tint of dark heliotrope gray; impressed sutural band pale
ochraceous buff on the first two whorls spotted with cinnamon buff,
on the last whorl] the color is cinnamon; lip and columella callus light
vinaceous fawn. Length 20.0 mm., greater diameter 13.1 mm., spire
height 11.2 mm., number of whorls 6$.
The darkest color pattern (pl. 10, fig. 25) and the obese form of
the shell has the first three-fourths postembryonic whorl axially
streaked with gull gray and dusky brown, the next or last whorl and
a half is almost solid dusky brown with faint axial streaks and spiral
lines of light quaker drab which are almost obsolete; impressed
sutural band burnt umber. Length 18.8 mm., greater diameter 13.8
mm., spire height 9.3 mm. An extreme narrow shell and light grayish-
brown pattern (pl. 10, fig. 25a) has the first postembryonic whorl
and a half pale smoke gray axially streaked and faintly spirally lined
with light grayish olive, the last whorl pale smoke gray spirally lined
and axially streaked with hair brown, last one-fourth whorl tilleul
buff axially streaked with verona brown. Length 17.8 mm., greater
diameter 11.2 mm., spire height 10.2 mm.
Meinecke first collected locality 290* in 1914 and obtained 109
shells, 93 percent of which are dextral. In 1917 he again went to
the same region and collected 83 shells, 86 percent of which are
sinistral. Since all Meinecke’s plotting before 1933 is done from
memory it is probable that these two lots represent slightly different
localities. The sinistral lot appears to have more bluish forms than
the lot of dextral shells. The usual dextral color patterns resemble
typical A. a. apicata. It is possible that the dextral locality may be
one closer to the area of A. a. apicata than the sinistral lot and so
transitional specimens between the two forms are found.
The usual form of a dextral (pl. 10, fig. 25b) has the embryonic
whorls ochraceous tawny ; first postembryonic whorl and a half axially
streaked with natal brown and pallid mouse gray, last whorl and a
half hay’s brown, axially streaked with tilleul buff, and sorghum
brown and faintly spirally lined with benzo brown; impressed sutural
band cinnamon; lip and columella callus a very dilute tint of pale
NO. I ACHATINELLA APEXFULVA DIXON—WELCH I51
vinaceous fawn. Length 19.6 mm., greater diameter 13.1 mm., spire
height 10.8 mm. ;
A narrow dextral and light bluish pattern (pl. Io, fig. 25c) has
the first half postembryonic whorl pale ochraceous salmon, axially
streaked with pale vinaceous fawn, next whorl and a half light
neutral gray spirally lined and streaked with pale gull gray, on the
last half whorl ground color pallid neutral gray almost covered over
with closely set axial streaks of light purplish gray and purplish gray ;
impressed sutural band cinnamon, darkening to mikado brown on the
last whorl. Length 19.8 mm., greater diameter 12.7 mm., spire height
11.1 mm.
One specimen was found with a pinkish pattern (pl. 10, fig. 25d),
the last whorl and a half seashell pink streaked and spirally lined and
banded with light mouse gray. This color pattern is found in lots of
shells collected by Gulick, Thurston, and others. Possibly there is
an intermediate region between areas 89 and 93 where A. a. sutura-
fusca var. 2 intergrades with A. a. cervixnivea.
ACHATINELLA APEXFULVA SUTURALBA, new subspecies
PLATE 3, FIGURE 12; PLATE 10, FIGURES 29, 29a
Achatinella apexfulva apicata (Newcomb) Pretrrer, Pilsbry and Cooke, Man.
Conch., vol. 22, p. 325, pl. 60, fig. 4b (only), 1914.
The form resembles A. a. suturafusca but differs in usually having
a white sutural band instead of a brown sutural band. Gulick collected
this form, but his lot (ANSP) contains a mixture of A. a. sutura-
fusca var. 2 and A. a. suturalba. His material is probably a mixture
of shells from areas 89 and go together with some intermediate
localities between these two areas.
The embryonic whorls of the holotype (pl. 3, fig. 12) ochraceous
tawny ; first two postembryonic whorls axially streaked with pallid
mouse gray, white, deep quaker drab, and quaker drab, last whorl
axially streaked with dark vinaceous drab, pale drab gray, and on the
last fourth of the whorl deep brownish drab; lip and columella callus
white or faintly tinged with a pinkish shade, possibly pale vinaceous
fawn. Length 19.4 mm., greater diameter 13.2 mm., spire height
10.8 mm., number of whorls 62.
Distribution, area go: North Poamoho Stream, type locality
290AA-8, elevation 1,150-1,200 feet, Lemke and Welch, 1935; also
locality 290 AA-5, elevation 1,150 feet, 1 dead dextral, Welch, BBM
132753, 1935; 291A-2, elevation 1,150 feet, 1 dead dextral, Welch,
BBM 165914, 1935; Central Poamoho Stream, locality 280A A-2,
152 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
elevation 1,100 feet, 3 dead dextral, Welch, BBM 132771, 1935; also
collected by Wilder, BBM 50636, Gulick, ANSP 98628C (figs. 5,
5a, p. 105).
The lightest color pattern found on only two specimens is shown
on plate 10, figure 29; last two postembryonic whorls axially streaked
with white, neutral gray, and deep neutral gray; impressed sutural
band and also the subsutural band, which disappears on the last half
whorl, white axially streaked with seashell pink; lip and columella
callus light vinaceous fawn.
A narrow shell (pl. 10, fig. 29a) and a dark color pattern lacks
the usual white sutural band; postembryonic whorls chaetura black
axially streaked with white, or tilleul buff; impressed sutural band
tilleul buff streaked with avellaneous. Length 20.0 mm., greater
diameter 12.2 mm., spire height 12.1 mm. This form with a colored
sutural band occurs on five specimens in the combined Lemke and
Welch lots of 45 shells exhibiting the color pattern. Fifty-one percent
have a white sutural band, the rest a tinted sutural band similar to
that of figures 29 or 29a.
ACHATINELLA APEXFULVA SUTURALBA var. 1
PLATE 3, FIGURE 19
Area 91??: Helemano, J. S. Emerson, BBM 102298-102299, also
Gulick. Possibly this race occurred in the South Helemano opposite
area gO in area gI?? (fig. 5a, p. 105). No specimens have been found
in recent years.
This variety is similar to A. a. suturalba but has a wider band of
white about the suture and never has a dark or a pinkish sutural band
similar to that of plate 10, figures 29 and 29a. The usual form (pl. 3,
fig. 19) has the embryonic whorls ochraceous tawny ; postembryonic
whorls pallid neutral gray axially streaked and faintly spirally lined
with deep neutral gray, on the last half whorl the streaks are deep
purplish gray; sutural and subsutural bands white; lip and columella
callus tilleul buff. Length 18.8 mm., greater diameter 12.2 mm.,
spire height 10.6 mm.
ACHATINELLA APEXFULVA APICATA (Newcomb) Pfeiffer
PLATE 3, FIGURE 14; PLATE 11, FIGURES 2, 3
Achatinella apicata (Newcomb) Pretrrer, Proc. Zool. Soc. London, 1855, p. 210.
Achatinella apexfulva apicata (Newcomb) Pretrrer, Pilsbry and Cooke, Man.
Conch., vol. 22, pp 324, 325, 328, pl. 60, figs. 4a, 4c (only), 1914.
There are two specimens of A. a. apicata in the type lot. One of
them marked “A” by me (pl. 11, fig. 2) is considered the lectotype.
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 153
The embryonic whorls are ochraceous buff shading to ochraceous
tawny, penultimate whorl pale drab gray axially streaked with mouse
gray, last whorl vinaceous buff axially streaked with verona brown,
last half whorl streaked with russet, last whorl at the edge of the
periphery has a line of light vinaceous buff ; lip light vinaceous fawn.
Length 20.5 mm., greater diameter 12.8 mm., number of whorls 63.
Distribution, area 92: Helemano, Gulick; Poamoho-Helemano
Ridge, locality 300F, elevation 1,250-1,300 feet, H. Lemke; Wahi-
awa, Gulick (fig. 5a, p. 105). Lectotype, BM.
The usual form in the Gulick collection (pl. 3, fig. 14) has the
embryonic whorls ochraceous tawny; first three-fourths postembry-
onic whorl tilleul buff axially streaked with vinaceous fawn, remain-
ing whorls with ground color of pale pinkish cinnamon darkening
on the last whorl to light pinkish cinnamon, and pale pinkish cinnamon
on the last half whorl, axially streaked on the penultimate with army
brown, benzo brown, and shaded or streaked with pale drab gray,
the first half of the last whorl streaked and sparsely lined with
brownish drab, last half whorl strongly streaked with army brown
or walnut brown, just above the edge of the periphery of the last
whorl a white line or band; lip and columella callus vinaceous buff.
Length 19.7 mm., greater diameter 12.8 mm., spire height 11.2 mm.,
number of whorls 63.
On an obese shell with wide axial streaks (pl. 11, fig. 3) the embry-
onic whorls are ochraceous tawny; ground of the postembryonic
whorls pale ochraceous salmon deepening on the last whorl to light
ochraceous salmon, and axially streaked or shaded with army brown,
fawn, or light drab, on the last whorl above the periphery a faint line
of pale ochraceous salmon, base spirally banded with light buff;
impressed sutural band russet. Length 19.1 mm., greater diameter
13.3 mm., spire height 10.4 mm.
ACHATINELLA APEXFULVA APICATA var. 1
PLATE 3, FIGURE 20; PLATE 11, FIGURES 4-7a
Area 95: North-South Helemano Ridge, locality 312-2, elevation
1,550 feet; 312-3, elevation 1,500-1,600 feet, 6 dextral; 312-4, eleva-
tion 1,600-1,650 feet; 312-5, elevation 1,650 feet, Meinecke, 1933;
310*, elevation 1,400-1,450 feet, 1932; 311*, elevation 1,450-1,500
feet, 1932; 312-2-4*, 1932, all collected by Meinecke. All the 1932
localities are plotted from memory and may be too low (figs. 5, 5a,
p. 105).
The shell is similar to A. a. apicata but differs in having the usual
form narrower ; the usual color pattern finely streaked with grayish
154 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
brown instead of widely streaked with reddish brown; sutural band
the same color as the rest of the whorl; ground color not as pinkish
as some specimens of A. a. apicata (pl. 11, fig. 3). The embryonic
whorls of the usual form (pl. 3, fig. 20) are worn and are best
described from another specimen; postembryonic whorls tilleul buff
finely axially streaked with hair brown, benzo brown, fuscous, and a
few streaks of natal brown, on the last half whorl the ground color
shaded with mouse gray, streaked with fuscous; impressed sutural
band wood brown; lip and columella callus vinaceous fawn. Length
19.0 mm., greater diameter 12.0 mm., spire height 10.9 mm., number
of whorls 64.
An obese form (pl. 11, fig. 4) has ochraceous tawny embryonic
whorls; first fourth postembryonic whorl tawny faintly streaked
with ochraceous tawny, last half whorl pale pinkish cinnamon axially
streaked with drab gray and cinnamon drab, penultimate whorl tilleul
buff axially streaked with hair brown, benzo brown, and natal brown,
first half of last whorl tilleul buff, shaded with drab gray, axially
streaked with hair brown and natal brown, last half whorl drab
axially streaked with tilleul buff, fuscous, and dark vinaceous brown
at and below the periphery lined with pale drab gray; impressed
sutural band the same as the rest of the whorl. Length 19.4 mm.,
greater diameter 13.2 mm., spire height 10.3 mm.
A light brownish specimen (pl. 11, fig. 5) resembles A. a. apicata,
but differs in not having a brown sutural band distinct from the rest
of the whorl and in having a color pattern with a shade of brown not
found on A. a. apicata. The postembryonic whorls are pale ochraceous
buff, strongly axially streaked with tawny olive and a few streaks of
warm sepia.
Throughout area 96, odd specimens (pl. 11, fig. 6) occur of an
entirely different pattern. The shell has the usual embryonic whorls ;
postembryonic whorls light pinkish cinnamon banded and_ mottled
and lined with chocolate, the bands broken in places by the ground
color; lip and columella callus light vinaceous fawn.
One specimen (pl. 11, fig. 7) has the first two postembryonic whorls
with the usual color pattern of plate 11, figure 4, and the last half
similar to plate 11, figure 6.
Another shell (pl. 11, fig. 7a) looks as if it were diseased, the first
fourth postembryonic whorl cinnamon axially streaked with cinnamon
buff, the next whorl and three-quarters warm blackish brown faintly
streaked with pale drab gray, resembling A. a. apicata var. 1, last
three-fourths whorl white spirally banded or splotched with tawny
and might be taken for a faded color pattern of figure 6.
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 155
ACHATINELLA APEXFULVA APICATA var. 2
PLATE 3, FIGURE 23; PLATE 11, FIGURES 14-16d
Area 96: Kawaihalona Gulch, locality 324, elevation 1,450 feet,
4 dextral 1933; 325, elevation 1,500-1,600 feet, 4 dextral 1933 ; 326*,
elevation 1,600-1,650 feet, 3 dextral 1932, all collected by Meinecke.
Other collectors of A a. apicata var. 2 in Kawaihalona Gulch are
Wilder, BBM 50509-50510, Cheatham in Thurston collection, BBM
130851. The material is unlocalized and probably is a mixture of
several races (figs. 5, 5a, p. 105).
Very little accurate information is available on the shells of Kawai-
halona. From the data on hand it appears that the characteristic
pattern is similar to plate 3, figure 23. Embryonic whorls a dark
shade of ochraceous buff; first fourth postembryonic whorl tawny
axially streaked with ochraceous buff, remaining postembryonic
whorls pale pinkish buff darkening to light buff, strongly axially
streaked and spirally lined or banded with hair brown or fuscous ;
impressed sutural band the color of the rest of the whorl; lip and
columella callus pale vinaceous fawn. Length 19.2 mm., greater
diameter 13.2 mm., spire height 10.7 mm., number of whorls 6+.
A darker banded pattern (pl. 11, fig. 14) has the postembryonic
whorls dark vinaceous brown axially streaked and banded and lined
with pallid mouse gray changing on the last fourth whorl to tilleul
buff. The base of this specimen is flattened.
Besides the above patterns, which have been checked by dead
specimens collected’ by Meinecke in area 96, a color pattern (pl. 11,
fig. 15) similar to apicata var. 1 (pl. 11, fig. 4) is found in locality 326.
The embryonic whorls are light buff deepening to warm buff; post-
embryonic whorls pale vinaceous fawn axially streaked with cinnamon
drab and bone brown, last whorl with a line of pale vinaceous fawn
above, below, and at the edge of the periphery.
In the Wilder collection, lot BBM 50510 had no locality label on
the shells when it came into the Bishop Museum. The lot may be a
mixture, for shells with color patterns similar to A. a. suturafusca
var. 2 (pl. 3, fig. 13) and other patterns are found mixed in with what
I suppose may be shells from Kawaihalona, because the color pattern
of plate 3, figure 23, occurs in the lot, and because of the position of
the lot in the Wilder collection when first received by the Museum.
Although the locality data are faulty and should be checked, some
peculiar patterns occur in this lot which are worthy of note since
they are not recorded from any plotted locality today.
156 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Plate 11, figure 16, is a color pattern similar to plate 11, figure 15,
except that it is banded on the last whorl above and below the
periphery with a band of pale ochraceous buff which is the ground
color. The shell may have a zigzag pattern (pl. 11, fig. 16a), the
postembryonic whorl hay’s brown or sorghum brown, axially streaked
with straight on zigzag streaks of pale mouse gray. An extreme
narrow shell (pl. 11, fig. 16b) with a more marked zigzag pattern
measures: Length 19.0 mm., greater diameter 11.9 mm., spire height
10.6 mm.; postembryonic whorls warm blackish brown, axially
streaked with zigzag lines of warm buff. Another specimen (pl. 11,
fig. 16c) shows another form of a narrow shell with a flattened base ;
postembryonic whorls chestnut faintly axially streaked or splotched
with warm buff ; impressed sutural band warm buff. Length 18.8 mm.,
greater diameter 12.0 mm., spire height 10.6 mm.
But by far the most unusual pattern, with the exception of patterns
16b and 16c, is the shell of plate 11, figure 16d. The embryonic
whorls are a deep shade of ochraceous buff; postembryonic whorls
fawn color sparsely axially streaked with pale ochraceous buff;
impressed sutural band fawn color. This color pattern may be a
pure race somewhere, possibly in the North Helemano or the South
Helemano at a low elevation below area 95. It looks very much like
typical A. a. apicata.
ACHATINELLA APEXFULVA PAALAENSIS, new subspecies
PLATE 3, FIGURE 21; PLATE 12, FIGURE 20
The shell is related to A. a. apicata var. 1 but differs in having a
blue-gray color pattern instead of a brown one. The holotype (pl. 3,
fig. 21) has the embryonic whorls ochraceous tawny ; first half post-
embryonic whorl mikado brown axially streaked with white, remaining
whorls white axially streaked with light gull gray, and light neutral
gray; impressed sutural band the color of the ground, except on
the last half whorl where it is shaded with cinnamon drab; lip and
columella callus light vinaceous fawn. Length 18.1 mm., greater
diameter 12.1 mm., spire height 9.7 mm., number of whorls 6+.
Distribution, area 106: North-South Helemano Ridge, type
locality 313-3, elevation 1,600-1,700 feet, 1933; South Helemano
Stream, locality 301c, elevation 1,350-1,400 feet, 4 dextral, Meinecke,
1933 (figs. 5, 5a, p. 105).
The color pattern in area 106 may be darker (pl. 12, fig. 20) and
the form narrower than the holotype; last two and a half post-
embryonic whorls pale gull gray axially streaked with deep mouse
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 157
gray and dark mouse gray, below the periphery a line of dark mouse
gray, the entire last whorl faintly lined with a light shade of deep
mouse gray; outer margin of the lip vinaceous fawn, inner edge and
the columella callus white. Length 18.0 mm., greater diameter 11.8
mm., spire height 9.7 mm.
ACHATINELLA APEXFULVA PAALAENSIS var. 1
PLATE 12, FIGURES 21-22)
Area 107: North-South Helemano Ridge, locality 313-1, eleva-
tion 1,700-1,800 feet, 6 dextral 1933; 314-1, elevation 1,650-1,750
feet, 1932, 1933; 315-2, elevation 1,800-1,864 feet, 7 dextral 1933;
315-3, elevation 1,800 feet, 1 dextral 1933; 316-1, elevation 1,750-
1,800 feet, 1933, 4 dextral 1934; 316-3, elevation 1I,700-1,800 feet,
5 dextral 1933, 3 dextral 1934; 316-1-3, 1932, all Meinecke-collected ;
Kawaihalona-Opaeula Ridge, locality 332*, elevation 1,700-1,850
feet, Russ, 2 dextral 1931, Meinecke, 1 dextral 1932; Helemano-
Opaeula Ridge, 334, elevation 1,450 feet, 1932, 1934; 336-1*, eleva-
tion 1,850 feet, 1929; 336-2, elevation 1,800-1,850 feet, 1 dextral
1934, all Meinecke-collected. Also collected by O. H. Emerson, BBM
103977-103982, J. S. Emerson, BBM 102291 (figs. 5, 5a, p. 105).
Throughout this region the shells are a mixture of blue-gray
forms and pinkish blue-gray or pink color patterns. The usual color
pattern and form of the shell on the North-South Helemano Ridge
(pl. 12, fig. 21), has the embryonic whorls ochraceous buff ; first half
of first postembryonic whorl axially streaked with vinaceous cinnamon
and pale pinkish buff, next whorl and a half white axially streaked
with vinaceous brown, and dark vinaceous brown with a spiral band
of dark vinaceous gray on the last half of the penultimate on the upper
third of the whorl, last whorl white or pale vinaceous fawn, and
heavily streaked with light vinaceous drab, dark vinaceous drab and
faintly spirally lined and banded with light quaker drab; lip and
columella callus light vinaceous fawn. Length 19.9 mm., greater
diameter 12.7 mm., spire height 11.1 mm.
The shell may be bluish gray (pl. 12, fig. 21a) ; last two and a half
postembryonic whorls white or a very dilute tint of pallid quaker drab
axially streaked with mouse gray and deep quaker drab. A narrow
shell and light pink color pattern (pl. 12, fig. 21b) has the penultimate
whorl axially streaked with white and vinaceous cinnamon, first half
of last whorl axially streaked with white and buff pink and a faint
band above the periphery of light cinnamon drab, last half of last
whorl light russet vinaceous deepening to light brownish drab on the
158 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
last fourth of the whorl, axially streaked with a color lighter than
the ground and spirally banded and lined with light brownish drab.
Length 19.7 mm., greater diameter 12.7 mm., spire height 10.5 mm.
The shape of the usual form and color pattern (pl. 12, fig. 22)
of the shell on the Kawaihalona-Opaeula Ridge or the Helemano-
Opaeula Ridge is more elongate and the color pattern darker. The
embryonic whorls are ochraceous buff; postembryonic whorls vina-
ceous pink axially streaked and faintly spirally banded with benzo
brown; lip and columella callus light vinaceous fawn. Length 20.0
mm., greater diameter 12.5 mm., spire height 11.3 mm. An obese shell
with a light pinkish-gray pattern (pl. 12, fig. 22a) has pale vinaceous
fawn postembryonic whorls axially streaked and faintly banded with
drab gray or light drab. Length 19.6 mm., greater diameter 13.3 mm.,
spire height 10.4 mm.
In the lot collected at locality 334 in 1932 by Meinecke a few
narrow yellowish-brown shells (pl. 12, fig. 22b) are found. This
pattern possibly belongs to a lower race, but until more is known
about this color form it will be considered to be A. a. paalaensis var. 1.
The embryonic whorls are ochraceous buff; postembryonic whorls
light pinkish cinnamon axially streaked with light drab, and drab on
the first two whorls, last whorl axially streaked with saccardo’s umber
and snuff brown, on the last two whorls above the periphery a band
of white or a dilute tint of pale pinkish buff, below the periphery of
the last whorl a line of snuff brown; lip and columella callus light
vinaceous fawn. Length 19.6 mm., greater diameter 12.0 mm., spire
height 11.4 mm.
ACHATINELLA APEXFULVA PAALAENSIS var. 2
PLATE 3, FIGURE 24
Area 109: Helemano-Opaeula Ridge, locality 337-1, elevation
1,800-1,850 feet, 1934; 337-2, elevation 1,800-1,900 feet, 1934; 337-3;
elevation 1,750-1,900 feet, 1934, Meinecke ; 337-1-2?, Meinecke, BBM
122320-122321, 1929; 337-1-3?, Meinecke, BBM 122268-122269,
1928, BBM 122362-122368, 1932. Also collected by Wilder, BBM
50513-50517, O. H. Emerson, BBM 103985 (figs. 5, 5a, p. 105).
In area 109 the pink-streaked pattern of paalaensis var. 1 becomes
dominant over the bluish-gray pattern. The usual form and color pat-
tern (pl. 3, fig. 24) has the usual ochraceous buff embryonic whorls ;
first half postembryonic whorl axially streaked with ochraceous tawny
and pale cinnamon pink, next two whorls pale ochraceous salmon
axially streaked with russet vinaceous, last half whorl cameo brown
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 159
- with a few axial streaks of light vinaceous cinnamon and a faint
band of white at the periphery ; lip, sutural and subsutural bands light
vinaceous fawn. Length 20.0 mm., greater diameter 12.7 mm., spire
height 10.7 mm., number of whorls 63.
ACHATINELLA APEXFULVA KAWAIIKI, new subspecies
PLATE 3, FIGURE 29; PLATE 12, FIGURES 26, 26a
The shell is similar in form to the elongate shell of A. a. paalaensis
var. I (pl. 12, fig. 22b) but has a lighter gray color pattern and
usually tends not to have the yellowish embryonic whorls of figure 22D.
The holotype (pl. 3, fig. 29) has the first embryonic whorl and a half
white, last embryonic whorl and a half pale cinnamon pink; first half
postembryonic whorl pale cinnamon pink axially streaked with light
pinkish cinnamon, next two postembryonic whorls pale drab gray
axially streaked with light cinnamon drab, last half whorl tinted with
wood brown, last whorl with a line of white below the impressed
sutural band, and a band of white below the periphery ; impressed
sutural band pale pinkish buff; lip and columella callus vinaceous
buff. Length 19.9 mm., greater diameter 12.2 mm., spire height 11.6
mm., number of whorls 62.
Distribution, area 99: Kawaiiki-Kawainui Ridge, locality 350,
elevation 1,550-1,600 feet, 1 dextral; 351, elevation 1,600 feet, 2
dextral 1933; the type lot collected in 1913 from the same ridge was
localized from the 1933 material, all Meinecke-collected. Also col-
lected by Wilder, BBM 50506-50507, Thurston, BBM 130892 (figs. 5,
aa, pp: 105).
An extreme obese shell (pl. 12, fig. 26) has the embryonic whorls
light buff; postembryonic whorls pale drab gray faintly axially
streaked with drab gray, last whorl lined above and below the periphery
with white; impressed sutural band white; lip and columella callus
light pinkish cinnamon. Length 20.1 mm., greater diameter 13.7 mm.,
spire height 11.1 mm.
The shell may be more conspicuously banded (pl. 12, fig. 26a) and
have darker embryonic whorls; embryonic whorls warm buff; last
two and a half postembryonic whorls drab gray, axially streaked on
the penultimate with light cinnamon drab, last whorl streaked with
drab and light drab, last two and a half whorls above the periphery
have a band of tilleul buff, below the periphery three bands of tilleul
buff; lip and columella callus vinaceous buff. Length 19.0 mm.,
greater diameter 12.1 mm., spire height 10.8 mm. This form resembles
A. a. paalaensis var. 1 (pl. 12, fig. 22b).
II
160 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Group oF A. A, ALOHA PILSBRY AND COOKE
ACHATINELLA APEXFULVA ALOHA Pilsbry and Cooke
PLATE 2, Figure 31; PLATE 12, FiGures I-2a
Achatinella apexfulva aloha Pitspry and Cooke, Man. Conch., vol. 22, p. 330,
pl. 60, figs. 15-15a (only), 1914.
To quote from Pilsbry and Cooke:
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 vinaceous 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 vina-
ceous, the columellar fold paler.
Length 18.5, diam. 12.7 mm.; 64 whorls.
Length 19.2, diam. 11.9 mm.; 64 whorls.
Crest of the division ridge between the two branches of the Kaukinehua
[Kaukonahua] stream, above the Wahiawa head-gates cabin, the colony extend-
ing to within { mile of main ridge; on mokihaua [mokihana?], pua [Osmanthus],
maile [Alyxia], and alani [Pelea]. Cotypes in collections A.N.S.P. and Bishop
Mus., from No. 3813 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 pl. 60, fig. 16, and differs from the
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 (toward
the mountains) is shut in by the precipitous side of the main Koolau Range.
Distribution, area 100?: Kaukonahua, Wilder, BBM 50561;
North Kaukonahua Stream, Thurston, BBM 130740. Lectotype
BBM (pl. 12, fig. 1). Pilsbry and Cooke plot this locality in the region
of area 100?. This locality has not been collected in recent years. Since
their information was obtained from Spalding, I believe that the
Wilder and Thurston shells can be plotted as coming from the same
place because these men either collected together or told each other
about their localities so that they could visit them separately (fig. 5a,
p:iLO5)):
The usual form and color pattern (pl. 2, fig. 31) on 87 percent of
the shells in the Wilder collection has the first postembryonic whorl
and penultimate white, penultimate banded just above the edge of
the periphery with pale ochraceous salmon, last whorl pale ochraceous
salmon with a few faint broken lines of white, sutural and subsutural
bands white; lip tinged with pale grayish vinaceous; columella callus
white. Length 19.2 mm., greater diameter 13.2 mm., spire height
9.6 mm.
att ~ SS
nga NEE
No. I ACHATINELLA APEXFULVA DIXON—WELCH ° 161
A slightly more obese form (pl. 12, fig. 2) has a similar pattern to
plate 2, figure 31 except that it is banded on the last whorl with white
like the lectotype. Length 19.2 mm., greater diameter 13.5 mm., spire
height 9.5 mm. A narrow specimen (pl. 12, fig. 2a) shows the white
form of the shell occurring on 13 percent of the shells; entire shell
white, last whorl with lines slightly darker than the ground color;
outer margin of the lip ochraceous tawny, remainder of lip and
columella callus white. Length 18.9 mm., greater diameter 11.8 mm.,
spire height 10.0 mm.
ACHATINELLA APEXFULVA ALOHA var. 1
PLATE 2, FIGURE 32; PLATE 12, FIGURES 3, 3a
Achatinella apexfulva aloha Pirspry and Cooke, Man. Conch., vol. 22, p. 330,
pl. 60, fig. 16 (only), 1914.
Area 100?: Kaukonahua, Wilder, BBM 50559-50560; North
Kaukonahua Stream, Thurston, BBM 103741 (fig. 5a, p. 105). This
form of aloha has an entirely different color pattern from that of typi-
cal aloha but is found as a rare pattern mixed with typical aloha. It is
possible that this is a separate race occurring above or below the
typical aloha area. Future collecting will have to determine this point.
Pilsbry considered this variety to be a mutation of aloha. (See above
description of A. a. aloha, p. 160).
The usual form and color pattern (pl. 2, fig. 32) has white
embryonic whorls; postembryonic whorls above the periphery white,
banded on the last two and a half whorls just above the periphery with
claret brown, lined on the last half of the penultimate and the first
half of the last whorl below the white subsutural band with a line
of clay color, which widens and darkens to a band of claret brown
on the last half whorl, below this a line of claret brown on the last
whorl, peripheral band on last whorl white, with a central line of
claret brown on the last half whorl, base claret brown lined with
white within and just without the aperture; impressed sutural band
white ; lip and columella callus light vinaceous fawn. Length 18.9 mm.,
greater diameter 13.1 mm., spire height 9.9 mm., number of
whorls 64.
This form may be more heavily banded with claret brown so that
the last whorl is almost solid claret brown and lined or banded with
white.
The lightest color pattern and elongate form (pl. 12, fig. 3) is a
white shell banded just above the edge of the periphery with two
bands of claret brown on the last two whorls, on the last whorl a
162 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
central line of claret brown, between the two bands, last whorl at and
below the periphery white, banded and lined with claret brown; lip
and columella callus light vinaceous fawn. Length 20.8 mm., greater
diameter 12.8 mm., spire height 11.2 mm.
On 6 percent of the Wilder shells a gray lined color pattern occurs
(pl. 12, fig. 3a); embryonic whorls white; postembryonic whorls
white faintly banded above the edge of the periphery with a band
made up of lines of pale drab gray, which are so faint as to be almost
white on the penultimate, on the last half whorl the lines darkening to
hair brown and the ground color of the band shaded with pale olive
6! 9 oe eee ie
gray, the peripheral band of white shaded or faintly axially streaked —
with pale olive gray and outlined above by two lines and below by
three lines of bone brown which become a single band on the last
half of the whorl, base white shaded with pale olive gray and spirally
lined with army brown.
ACHATINELLA APEXFULVA ROSEIPICTA, new subspecies
PLATE 3, FIGURE 7; PLATE 12, FicuRES 4-4)
The form of the shell closely resembles that of A. a. beata. The
color patterns, however, tend toward pastel shades of pink and are
closer to those of A. a. aloha. The race seems to be intermediate
between A. a. beata and A. a. aloha. The embryonic whorls of the
holotype (pl. 3, fig 7) are pale pinkish cinnamon; postembryonic
whorls below the white impressed sutural and wide subsutural bands
are walnut brown faintly axially streaked or tinted with white and
a faint line of burnt umber just below the subsutural band, the ground
shading on the last whorl to vinaceous, russet, and kaiser brown on
the last half whorl with a faint band of white at the edge of the
periphery, which fades out on the last half whorl; lip and columella
callus pale grayish vinaceous. Length 19.5 mm., greater diameter
13.0 mm., spire height 10.0 mm., number of whorls 63.
Distribution, area 102: North Kaukonahua-Poamoho Ridge,
type locality 286, elevation 1,950 feet, 1933, 1936; also locality
285-3, elevation 1,850 feet, 1933; 285-4, elevation 1,750-1,900 feet,
1936; all collected by Meinecke (figs. 5, 5a, p. 105). Also collected
by O. H. Emerson, BBM 103984, on “Spurs north-east of head-gates,
Kaukonahua,” 1917. The head gates are probably in the North
Kaukonahua. Also collected by Wilder, BBM 50563, but the shells
have no original label and are marked Poamoho? by C. M. Cooke, Jr.,
who surmised the locality to be somewhere in that region from the
arrangement of this lot in the Wilder collection.
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 163
A lighter color pattern (pl. 12, fig. 4) occurs on 37 percent of the
shells in area 102; the last two postembryonic whorls and a half
pale congo pink faintly lined and banded with vinaceous pink, edge
of the periphery of the last whorl banded with a band of white tinted
with a very dilute tint of pale congo pink or a dilute pinkish tint of
pale vinaceous fawn, base vinaceous pink; subsutural and impressed
sutural band white. Length 19.8 mm., greater diameter 12.0 mm.,
spire height 11.6 mm. Forty-one percent of the shells have the color
pattern of the holotype.
Sixteen percent have a pinkish gray or gray color pattern shown
by an obese shell (pl. 12, fig. 4a), which has a concave spire, the first
two postembryonic whorls white, last whorl and a half below the
white sutural and subsutural bands are a very dilute tint of pale
grayish vinaceous spirally lined with pale grayish vinaceous, in the
umbilical region a patch of honey yellow. Length 19.7 mm., greater
diameter 13.8 mm., spire height 9.7 mm.
Six percent have a white color pattern (pl. 12, fig. 4b) ; which has
a spire straight in outline; embryonic whorls pale cinnamon pink
shading to white on the first half postembryonic whorl, the remaining
whorls white, in the umbilical region a dot or small patch of honey
yellow.
ACHATINELLA APEXFULVA ROSEIPICTA var. 1
PLATE 3, FicuRE 8; PLATE 12, FIGURES 5-6
Area 103: North Kaukonahua-Poamoho Ridge, locality 289-1a,
elevation 2,000-2,050 feet, 1933; 287-1-2, elevation 1,900 feet,
1933; 287-1, elevation 1,900 feet, 8 dextral 1936; 287-2, elevation
1,900 feet, 2 dextral 1936; 287-3, elevation 1,900-1,950 feet, 1936;
288, elevation 1,900-2,068 feet, 1933; all collected by Meinecke
(figs. 5, 5a, p. 105).
This variety is intermediate between A. a. aloha and A. a. roseipicta.
The banding resembles aloha but the color is much darker and is
closer to roseipicta. The usual form (pl. 3, fig. 8) and a color pattern,
found on 17 percent of the shells, has the embryonic whorls white;
postembryonic whorls white, banded and lined with avellaneous,
peripheral band white, outlined above and below by bands of cinnamon
rufous, base cinnamon rufous, banded with tilleul buff and a band
of white; impressed sutural and subsutural bands white; lip and
columella callus light vinaceous fawn. Length 19.1 mm., greater
diameter 13.0 mm., spire height 9.8 mm., number of whorls 64.
164 SMITHSONIAN MISCELLANEOUS COLLECTIONS , VOL. 103
Eight percent of the shells have a reddish-brown color pattern (pl. 3,
fis. 7).
The usual pink-lined color pattern on 67 percent of the shells and the
obese form of the shell (pl. 12, fig. 5), has the first two postembryonic
whorls tilleul buff below the white sutural and subsutural bands, last
whorl vinaceous buff, banded above and at the periphery with white,
below the periphery lined with fawn color. Length 19.4 mm., greater
diameter 14.0 mm., spire height 9.6 mm. The usual narrow. form of
the shell (pl. 12, fig. 5a) measures: Length 19.6 mm., greater diameter
12.7 mm., spire height 10.2 mm.; has an extremely light pink lined
pattern ; postembryonic whorls white faintly lined with a very dilute
tint of seashell pink darkening on the last half whorl to buff pink
lines or bands.
In locality 287-2 a dark gray color pattern occurs on two specimens.
The color pattern (pl. 12, fig. 6) is similar to that of A. a. poamo-
hoensis except that the gray ground color is broken up with light
lines and bands. The embryonic whorls are pale pinkish buff; post-
embryonic whorls below the white sutural and subsutural bands
pallid mouse gray spirally lined with benzo brown, peripheral band
on the first fourth of the last whorl pale smoke gray, last three-fourths
whorl bone brown or natal brown, lined and banded with light
vinaceous fawn. One shell, or 2 percent, has a white pattern similar
to that of plate 12, figure 4b. Another shell from locality 289-1a is
black lined and similar to plate 12, figure 3a.
ACHATINELLA APEXFULVA ROSEIPICTA var. 2
PLATE 3, FicureE 18; PLATE 12, FIGURES 9-I2a
Area 105: Central Poamoho Stream, locality 280F-1, elevation
1,550 feet, 5 dextral 1933; 280F-2, elevation 1,600 feet, 1933, 1936;
280F-3, elevation 1,600-1,650 feet, 4 dextral 1933, Meinecke;
Poamoho-Helemano Ridge, 306-4, elevation 1,700-1,950 feet,
Welch, 3 dextral 1935; 307-Ia-Ib, elevation 1,800-1,850 feet,
Meinecke, 1917, 5 dextral 1935; 307-1, elevation 1,800-1,850 feet,
C. W. Isle, 1 dextral 1935, Welch, 1 dextral 1935; 307-1-2, elevation
1,800-1,965 feet, Meinecke, 5 dextral 1934; 307-3, elevation 1,950 feet,
Lemke, 1934; 307-3-308-1, elevation 1,850-1,950 feet, Meinecke,
1934; 308-1, elevation 1,850-1,900 feet, Welch, 2 dextral 1935, Russ,
10 dextral 1931, Russ, BBM 134206, 120037, 1932; 308-2, elevation
1,850-1,900 feet, Welch, 1 dextral 1935, C. W. Isle, 1 dextral 1935;
also collected by Lemke above 307-3, BBM 10444, 1934 (figs. 5, 5a,
'p. 105).
|
eri aie
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 165
The usual color pattern (pl. 3, fig. 18) is lighter and not so sharply
banded as that of typical beata. The form resembles roseipicta var. 1
in shape and color pattern but differs in shade and range of the color
patterns. The race is intermediate between A. a. roseipicta and A. a.
beata. The embryonic whorls are cartridge buff ; first two postembry-
onic whorls white, penultimate whorl banded in the center of the whorl
with a band of vinaceous fawn, last whorl below the white sutural and
subsutural bands pale vinaceous fawn or a dilute shade of pale vina-
ceous fawn, banded with vinaceous fawn, in the umbilical region a
band of white ; lip a faint tint of pale vinaceous fawn, columella callus
white. Length 20.0 mm., greater diameter 13.1 mm., spire height
11.1 mm. The usual form is not determined by measurement because
of insufficient material. This shell is probably near the usual form.
An obese shell and light color pattern (pl. 12, fig. 9) has white
postembryonic whorls banded on the last whorl above the periphery
with a band of light vinaceous fawn darkening to vinaceous fawn on
the last half whorl, last half whorl below the periphery faintly shaded
or lined with a dilute tint of vinaceous fawn. Length 19.8 mm.,
greater diameter 13.5 mm., spire height 10.0 mm. The shell may have
pure white embryonic whorls.
In the higher localities of 307-3 and 308-1 a form (pl. 12, fig. 10)
occurs which resembles in color A. a. aloha (pl. 12, fig. 2) except
that the bands are a darker shade and a different hue. The shell of
figure 10 also shows an elongate form; postembryonic whorls white,
lined just above the edge of the periphery with two bands of pinkish
buff, a very dilute shade on the first whorl and darkening to a
stronger shade on the next two whorls, the lower band being pinkish
cinnamon on the last whorl, last whorl below the periphery banded
or lined with pinkish cinnamon the ground color pale pinkish
cinnamon.
A dark color pattern (pl. 12, fig. 11) has a grayish-pink color;
first half of the last postembryonic whorl below the white sutural
and subsutural bands ecru drab, tinted, faintly lined, and finely
axially streaked with light vinaceous fawn, last half whorl cinnamon
drab faintly tinted or finely streaked with benzo brown, at the edge
of the periphery a band of white, which fades out on the last half
whorl.
In locality 308-2 and in other localities at the upper limit of area 106
a gray pattern (pl. 12, fig. 12) occurs which resembles aloha var. 1
(pl. 12, fig. 3a), but does not have its obese appearance. The embry-
onic whorls are pale pinkish buff ; postembryonic whorls white, banded
166 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
on the last whorl and a half with pale drab gray below the white |
sutural band, above the white peripheral band banded with hair
brown, below the periphery base banded with hair brown and white;
lip and columella callus vinaceous fawn.
Lemke found a pink color pattern (pl. 12, fig. 12a) which looks
like A. a. roseipicta, the last whorl and a half vinaceous fawn, finely
axially streaked with light vinaceous fawn, on the last whorl at the
periphery a band of pale drab gray which fades out on the last fourth
whorl, base lined with light vinaceous fawn, a band of white in the
umbilical region.
ACHATINELLA APEXFULVA POAMOHOENSIS, new subspecies
PLATE 3, FIGURES 5, 6; PLATE 12, FiGuREs 7-8
The shell is similar in form to the higher race of A. a. roseipicta,
but differs in having a gray instead of a pink color pattern. The
embryonic whorls of the holotype (pl. 3, fig. 6) are white; post-
embryonic whorls white faintly banded on the penultimate with a
central band of pale drab gray, last whorl pale drab gray faintly
spirally lined with white ; lip and columella callus pale vinaceous fawn.
Length 19.7 mm., greater diameter 12.8 mm., spire height 9.6 mm.,
number of whorls 64.
Distribution, area 101: North Kaukonahua-Poamoho Ridge,
type locality 284-2, elevation 1,700-1,800 feet; also locality 283-3,
elevation 1,750 feet, 5 dextral, 1 sinistral; 283-1, elevation 1,650-
1,750 feet, 6 dextral; 283-4, elevation 1,600-1,650 feet, 4 dextral;
284-1, elevation 1,550-1,600 feet, 4 dextral; 285-1, elevation 1,750-
1,800 feet, 13 dextral; 285-2, elevation 1,750-1,850 feet, 1 dextral;
all collected by Meinecke, 1933 (figs. 5, 5a, p. 105).
The usual form of the shell and the dark color pattern (pl. 12,
fig. 7) has the embryonic and first postembryonic whorls light buff,
penultimate whorl below the white sutural and subsutural bands pale
drab gray finely and faintly spirally lined and finely axially streaked
with benzo brown, ground of last whorl drab gray, deepening to light
drab on the last whorl, finely axially streaked with benzo brown,
below the periphery the color shading to saccardo’s umber faintly
lined with benzo brown, in the umbilical region a patch of cream color ;
lip and columella callus tilleul buff, outer margin of lip avellaneous.
Length 19.3 mm., greater diameter 12.7 mm., spire height 10.8 mm.
The usual yellow pattern and the obese form of the shell (pl. 12,
fig. 7a) measures: Length 19.1 mm., greater diameter 13.0 mm., spire
height 9.5 mm.; embryonic and postembryonic whorls light buff,
oe
‘= eee
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 167
spirally banded above the periphery on the first half of the last
postembryonic whorl with a faint band of drab gray and a line of
cinnamon buff, below the periphery lined with pinkish buff, last half
whorl lined, banded, or tinged with pale drab gray ; impressed sutural
band the color of the ground.
A darker yellow color pattern and elongate shell (pl. 3, fig. 5) has
the embryonic whorls pale pinkish buff; postembryonic whorls warm
buff, last whorl, below the subsutural band of warm buff, drab gray,
and below this, just above the edge of the periphery, a band of warm
buff, which fades out on the last half whorl, base drab faintly axially
streaked or tinted with warm buff; impressed sutural band pale
cinnamon pink; lip and columella callus pale vinaceous fawn. Length
18.7 mm., greater diameter 12.0 mm., spire height 10.4 mm.
An exceedingly dark specimen (pl. 12, fig. 8) is found in locality
283-1; penultimate whorl light drab shading into rood’s brown on the
last whorl; impressed sutural band pale pinkish buff; subsutural
band shading from pale pinkish buff to light buff on the last whorl, on
the last half whorl narrowing to a line and fading out.
Below locality 284-2 in area 1o1 the shells have a yellow ground
color, and no shells with white sutural and subsutural bands are
present. In locality 284-2 only 6 shells out of 34 have a yellow ground
color. In localities 285-1 and 285-2 the impressed sutural and sub-
sutural bands are white, and pinkish gray and white color patterns
occur, so that this region appears to be intermediate between . a.
roseipicta and A. a. poamohoensis.
ACHATINELLA APEXFULVA CERVIXNIVEA Pilsbry and Cooke
PLATE 3, FIGURE 15; PLATE 11, FicuRES 8-10
Achatinella apexfulva cervixnivea pattern Pirspry and Cooke, Man. Conch.,
vol. 22, pp. 322, 328, pl. 60, figs. 8-8a (only), ror4.
The lectotype of A. a. cervixnivea is selected by Pilsbry and myself
from the figured cotypes (pl. 60, fig. 8) and reproduced in this paper
on plate 11, figure 8. The embryonic whorls are buckthorn brown ;
postembryonic whorls, below the broad white sutural and subsutural
bands, dark quaker drab or deep quaker drab, below the periphery of
the last whorl the ground is pale mouse gray, spirally banded, lined,
and axially streaked with quaker drab; lip and columella callus light
vinaceous fawn. Length 20.0 mm., greater diameter 13.6 mm., spire
height 10.0 mm., number of whorls 64.
Distribution, area 93: Poamoho-Helemano Ridge, locality 300-4*,
elevation 1,500-1,550 feet, Russ, 1932; 300-5, elevation 1,650 feet,
168 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
I sinistral 1933; 300-6, elevation 1,650-1,700 feet, 1917, 6 dextral
1932, 3 sinistral 1934; 300-7, elevation 1,500-1,700 feet, 4 dextral
1934; 300-8, elevation 1,550-1,700 feet, 4 dextral 1933, 1 dextral
1934; 30I-1-2*, elevation 1,650 feet, 9 sinistral 1932; 300-6-301-2*,
elevation 1,650 feet, 1916, all collected by Meinecke ; 300-10, elevation
1,550-1,600 feet, Russ, 4 dextral 1933; 301-1, elevation 1,650 feet,
H. B. Baker and Welch, 3 dextral and 6 sinistral 1935 ; 301-2, eleva-
tion 1,650 feet, H. B. Baker and Welch, I sinistral 1935. Also col-
lected by Wilder, BBM 50631, 50566 (figs. 5, 5a, p. 105).
The usual form and color pattern (pl. 3, fig. 15) has the embryonic
whorls clay color, first fourth postembryonic whorl pale pinkish buff
axially streaked or tinged with cinnamon buff, next half whorl pallid
quaker drab axially streaked and banded, below the white sutural and
subsutural bands, with a band of deep quaker drab, last whorl and a
half above the periphery and below the subsutural band vinaceous
slate deepening on the last half whorl to dark purple drab, just above
the edge of the periphery and all below, ground pale ecru drab finely
lined and faintly streaked with deep quaker drab, on the last half
of the whorl streaks and lines dark purple drab, in the umbilical
region a patch of pale pinkish buff; lip and columella callus light
vinaceous fawn. Length 18.8 mm., greater diameter 12.8 mm., spire
height 9.8 mm., number of whorls 6.
A light color pattern (pl. 11, fig. 9), has the last two postembryonic
whorls, below the white sutural and subsutural bands, pale vinaceous
fawn shading to light vinaceous fawn on the last half whorl, spirally
banded and faintly axially streaked with ecru drab, on the last half
whorl bands and streaks brownish drab, in the umbilical region a
band of pale pinkish buff.
The usual form of a sinistral shell (pl. 11, fig. 9a) has a color
pattern similar to that of plate 3, figure 15, except that the embryonic
whorls in this specimen are warm buff. Length 18.7 mm., greater
diameter 12.0 mm., spire height 10.0 mm.
In locality 300-4 the usual pattern has a more pinkish ground than
the typical pattern. Among these pinkish gray shells two extremely
light pink forms are found. One of these (pl. 11, fig. 10) shows an
obese shell; postembryonic whorls with a band of pecan brown below
the white sutural and subsutural bands, below which the ground is
buff pink streaked and lined on the last whorl with pecan brown;
lip and columella callus pale vinaceous pink. Length 19.3 mm.,
greater diameter 13.6 mm., spire height 10.2 mm.
From the data on hand cervixnivea runs from pinkish forms at a
low elevation to dominantly light gray shells. At the upper end of
j
\
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 169
area 93 near area 94 beata-like forms and the black forms of beata
var. I are found mixed with A. a. cervixnivea.
ACHATINELLA APEXFULVA BEATA Pilsbry and Cooke
PLATE 3, FIGURE 17; PLATE 12, FIGURES 13-16
Achatinella apexfulva beata Pitspry and Cooke, Man. Conch., vol. 22, p. 320,
pl. 60, figs. 17a-17c; pl. 55, fig. 5 (only), ror4.
Achatinella apicata var. alba Sykes, Fauna Hawaiiensis, p. 299, 1900. Not
Achatinella alba Nuttall in Jay’s Catalogue, ed. 3, p. 58, 1839 (name only),
which is Achatinella lorata Férussac.
Achatinella apexfulva alba SyKes, Pilsbry and Cooke, Man. Conch., vol. 22,
P- 324, 1914.
The shell of Pilsbry’s plate 60, figure 17a, reproduced here on plate
12, figure 13, is considered the lectotype. The embryonic whorls are
cream color; postembryonic whorls, below the white sutural and
subsutural bands, with a band of liver brown, below which is a
band of white or seashell pink, last whorl below the white sutural
and subsutural bands seashell pink, banded above, at the edge and
below the periphery with livid brown, below the periphery the bands
faintly axially streaked with seashell pink, in the umbilical region a
patch of naples yellow; lip pale vinaceous fawn, columella callus
white. Length 19.8 mm., greater diameter 13.0 mm., spire height
II.0 mm., number of whorls 6§. The pattern is dominantly a banded
one, and the streaks are faint or not prominent.
Distribution, area 104: Central Poamoho-Central Poamoho
North Branch Ridge, locality 280L-6, elevation 1,850-1,950 feet,
Meinecke, 1933; 280L-7, elevation 1,900 feet, Meinecke, 3 dextral
1933 ; 280L-9, elevation 1,850 feet, Russ, 1934; Poamoho-Helemano
Ridge, locality 303, elevation 1,700-1,800 feet, Meinecke, 1934, Russ
1934; 304, elevation 1,800-1,900 feet, Meinecke, 1 dextral 1933,
I dextral 1934, Welch, 1935; 305, elevation 1,850-2,300 feet,
Meinecke, 2 dextral 1933, Welch, 4 dextral 1935; 306-1, elevation
1,850-2,000 feet, Meinecke, 7 dextral 1933; 306-3, elevation 1,750-
1,850 feet, 1 sinistral 1934; also 302-3047, BBM 122106, 1916;
305-300-2?, BBM 122112-122113, 1932, less localized Meinecke
material plotted from memory; South Helemano Stream, locality
300H, elevation 1,500-1,600 feet, Meinecke, 1 dextral 1934 ; 300G, ele-
vation 1,600-1,750 feet, Meinecke, 6 dextral 1934; North-South
Helemano Ridge, locality 317, elevation 1,750-1,950 feet, Meinecke,
8 dextral 1934. Other collectors of A. a. beata probably from area
104 are Thurston, BBM 130772, Wilder, BBM 50633, Lemke, and
others (figs. 5, 5a, p. 105).
170 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
The usual form and color pattern (pl. 3, fig. 17) has the embry-
onic whorls warm buff; first half postembryonic whorl warm buff,
last half white with a central band of clay color, last two whorls
above the periphery white with a central band of hazel shading into
kaiser brown on the last whorl, last whorl below the kaiser brown
peripheral band naples yellow banded with kaiser brown; lip pale
vinaceous fawn, columella callus white. Length 19.8 mm., greater
diameter 12.8 mm., spire height 10.3 mm.
The lightest color pattern and obese form (pl. 12, fig. 15) has the
embryonic whorls light buff ; postembryonic whorls white, last whorl
above the periphery banded with light pinkish cinnamon, below the
edge of the periphery pale pinkish buff faintly banded on the last
half whorl at the edge of the periphery and about the base with
light vinaceous fawn; lip and columella callus light grayish vinaceous.
Length 19.3 mm., greater diameter 13.6 mm., spire height 9.1 mm.
The dark banded color pattern and narrow form (pl. 12, fig. 14)
has the first two postembryonic whorls white, banded about the
middle of the whorl above the periphery with a band darkening from
light pinkish cinnamon on the first postembryonic whorl to pecan
brown on the penultimate and last whorl ; sutural and subsutural bands
white, on the last whorl the white supraperipheral band is shaded
with seashell pink, the peripheral band is walnut brown deepening to
burnt umber on the last half of the whorl, subperipheral band light
vinaceous cinnamon fading out on the last half whorl, remainder of
base, except for a patch of pale pinkish buff in the umbilical region,
burnt umber faintly lined and axially streaked with light vinaceous
cinnamon. Length 20.0 mm., greater diameter 12.3 mm., spire height
11.2 mm.
The color pattern (pl. 12, fig. 16) is similar to plate 3, figure 17,
except that the embryonic whorls have a more snow-white ground,
and the last whorl below the periphery is massicot yellow faintly
banded and axially streaked with fawn color, last fourth whorl
streaked with army brown.
The holotype of A. a. alba Sykes in the British Museum is a white
shell similar to the white color forms of A. a. beata and which was
collected by Perkins in Kawailoa. A. a. beata would have to be put in
the synonymy of A. alba if the name alba were not already pre-
occupied. The name alba was used by Nuttall, in Jay’s Catalogue,
and placed in the synonymy of A. pallida Nuttall by Reeve (Conch.
Icon., vol. 6, pl. 1, species 2, 1850). No white shells similar to A. a.
beata have been taken in Kawailoa proper. Mr. Sykes’ shell may have
NO. I ACHATINELLA APEXFULVA DIXON—-WELCH 171
come from the Helemano-Opaeula Ridge, which is the division ridge
between the districts of Paala and Kawailoa. This ridge is the most
northern ridge known to contain white beata-like shells.
ACHATINELLA APEXFULVA BEATA var, 1
PLATE 3, FIGURE 16; PLATE II, FIGURES 11-13
Achatinella apexfulva beata Prtspry and Cooker, Man. Conch., vol. 22, p. 329,
pl. 60, fig. 17 (only), 1914.
Area 94: Poamoho-Helemano Ridge, locality 302, elevation 1,750
feet, Meinecke, 1916, 1932, Russ, 2 dextral 1933; 302-291-5, eleva-
tion 1,750-1,802 feet, Baker and Welch, 2 dextral 1935; North-
Central Poamoho Ridge, locality 291-3-5*, elevation 1,650-1,802
feet, 1916-1917, 1932; 291-5, elevation 1I,750-1,802 feet, 4 dextral
1934, all Meinecke localities. Also collected by Wilder, BBM 50568-
50569, and others.
Area 94-94a?: North-Central Poamoho Ridge, locality 2g1-2?,
elevation 1,600-1,700 feet, 6 dextral 1932; 291-3?, elevation 1,650-
1,700 feet, 12 dextral 1926; 291-2-302?, elevation 1,600-1,800 feet,
1914, all Meinecke-collected. The shells in this area contain a mixture
of color patterns of typical 4. a. cervixnivea and beata var. 1. All
the localities are plotted from memory and are of wide extent. Prob-
ably if more localized collecting were done in area g4a, the lower por-
tion of the ridge might contain only cervixnivea while the upper por-
tion of area 94 might contain a dominance of the dark color pattern
of beata var. 1 (figs. 5, 5a, p. 105).
Pilsbry also considered this race a form of beata; actually it is a
distinct race existing between beata and cervixnivea with a color
pattern distinct from both. However, this form is so close to the
typical apexfulva that it is almost impossible to tell the difference
between the forms. Typical apexfulva usually has forms with the
last whorl a light yellowish color and banded, lined, and axially
streaked with chocolate or black, while cervixnivea var. 1 usually has
a solid color or patterns intermediate with typical cervixnivea or beata.
The usual form (pl. 3, fig. 16) from locality 291-3-5 measures:
Length 19.4 mm., greater diameter 13.4 mm., spire height 10.3 mm. ;
the color pattern is a little lighter than usual; embryonic whorls light
buff; last two and a half postembryonic whorls chocolate, tinted or
lightened with streaks of mahogany red ; impressed sutural band warm
buff, shaded on the lower half of the band on the last whorl with
russet ; lip and columella callus white.
; 172 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
A more usual color pattern and the usual form in the Wilder col-
lection (pl. 11, fig. 12) has the last two whorls maroon; impressed
sutural band mahogany red, the upper portion of which is lined or
edged with pale pinkish buff; lip and columella callus pale vinaceous
fawn. Length 19.7 mm., greater diameter 12.8 mm., spire height
10.8 mm. In a lot of 69 shells, 11 percent have white sutural bands
similar to those of plate 11, figure 11. The postembryonic whorls on
this specimen are particularly dark, being seal brown.
One or two specimens in area 94 have a yellowish color pattern
(pl. 11, fig. 13), the embryonic whorls warm buff; first half post-
embryonic whorl warm buff lightly axially streaked with ochraceous
tawny, on the penultimate and last whorl above the periphery, the
upper third of the whorl, including the impressed sutural and sub-
sutural bands, amber brown faintly axially streaked with pale
orange yellow, below which the middle third of the whorl has a band
of dusky brown which is faint on the early whorls and does not really
start until the second half of the penultimate, just above the edge of
the periphery a band of pale orange yellow, tinted with amber brown,
which becomes narrower on the last half of the whorl, below the
periphery dusky brown faintly tinted or streaked with pale orange
yellow, in the umbilical region a band of mars yellow.
ACHATINELLA APEXFULVA BEATA var, 2
PLATE 3, FIGURE 22; PLATE 12, FIGURES 17, 18
Area 108: North-South Helemano Ridge, locality 318, elevation
1,800-1,900 feet, 1934; 318a, elevation 1,850-1,950 feet, 19343; 319,
elevation 1,850-1,900 feet, 5 dextral 1934; 319a, elevation 1,950-2,000
feet, Meinecke, 1 dextral 1934 (figs. 5, 5a, p. 105).
In area 108 the shells are dominantly white color forms mixed with
beata color patterns. The usual form of the shell (pl. 3, fig. 22) has
the embryonic whorls a dilute shade of light buff; remainder of the
shell white except for about 3.5 mm. behind the edge of the lip, where
the ground is tinged with naples yellow ; lip and columella callus white.
Length 19.4 mm., greater diameter 12.7 mm., spire height 11.0 mm.
An extremely obese shell (pl. 12, fig. 17) is pure white ; the embry-
onic whorls are worn and lack the shell nacre; the lip and columella
callus white, outer margin of the lip clay color. Length 19.2 mm.,
greater diameter 13.5 mm., spire height 9.8 mm. A narrow shell
(pl. 12, fig. 18) shows the usual light beata pattern; postembryonic
whorls white, last postembryonic whorl above the periphery with a
;
4
'
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 173
band of buff pink deepening to cacao brown on the last half of the
whorl, last whorl below the periphery white on the first half of the
whorl, last half banded at the edge of the periphery and on the base
with cacao brown, subperipheral band white tinged with vinaceous
pink. Length 19.7:mm., greater diameter 12.7 mm., spire height 11.2.
ACHATINELLA APEXFULVA BEATA var. 3
PLATE 3, FIGURE 25; PLATE 12, FIGURE 19-I19b
Area 110: Helemano-Opaeula Ridge, locality 337-4, elevation
1,900-2,000 feet, 1934; 338, elevation 1,950-2,000 feet, Meinecke,
1934; also less localized Meinecke material plotted from memory,
locality 338-339-1°?, BBM 128329-128333, 122335-122336, 1929;
339-1a *, elevation 1,950-2,000 feet, 1929; 338?, BBM 122267, 1928
(figs. 5, 5a, p. 105).
This race differs from A. a. beata in usually having lighter pink
color patterns and a different range of color patterns. The character-
istic pattern of this area (pl. 3, fig. 25) has the embryonic whorls
cream buff; first two postembryonic whorls white, first half of the
last whorl white faintly shaded with a very dilute color of pale grayish
vinaceous, lightly banded with light russet vinaceous, last half whorl
vinaceous brown, faintly lined or banded and axially streaked with
light russet vinaceous and banded with white in the umbilical region ;
lip and columella callus light grayish vinaceous. Length 19.6 mm.,
greater diameter 13.5 mm., spire height 10.3 mm. The aboral surface
of this shell is figured in order to show the abrupt change of coloration
from white to a dark almost solid pinkish color on the last half whorl.
This pattern is found only in area IIo.
A darker pattern (pl. 12, fig. 19) has the first half of last whorl
light brownish vinaceous, banded and shaded with russet vinaceous,
last half whorl almost solid vinaceous brown faintly lined and streaked
with a lighter tint.
A white specimen (pl. 12, fig. 19a) lacks color except for the
cream-buff embryonic whorls and a line of ochraceous tawny on the
outer margin or back edge of the lip. Plate 12, figure 19b, shows a
rare pink color form with the postembryonic whorls a very dilute
tint of pale flesh color deepening to flesh color on the last whorl,
3-5 mm. behind the edge of the lip the color deepening to vinaceous
pink,
174 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
ACHATINELLA APEXFULVA VESPERTINA Baldwin
PLATE 3, FicuRE 28; PLATE 12, FIGURES 23-25
Achatinella vespertina BAaLpwin, Proc. Acad. Nat. Sci., Philadelphia, 1895,
p. 210, pl. 10, fig. 14—Pitssry and Cooke, Man. Conch., vol. 22, p. 322, pl. 60,
figs. 2, 3, 1914.
The holotype in the Academy of Natural Sciences (pl. 12, fig. 23)
has the embryonic whorls cartridge buff shading to light pinkish
cinnamon on the last half embryonic whorl; postembryonic whorls
light pinkish cinnamon; impressed sutural band cartridge buff; lip
and columella callus cartridge buff. Length 20.6 mm., greater diameter
14.3 mm., number of whorls 63.
Distribution, area 98?: Kawailoa, D. D. Baldwin, type lot, BBM
57990, 5 dextral, also BBM 54685, 54737, 6 dextral, E. Lyman ex
Baldwin, BBM 167354, 9 dextral; Kamoku, J. S. Emerson, BBM
102310-102311; Waialua, J. S. Emerson, BBM 102309; “Kawaiiki
4 way down the valley, side north of 2nd Kamoku Plateau.” (O. P.
Emerson label in O. H. Emerson collection, BBM 103983.) “In a
secondary ravine of Kawatki Valley on the South side, not quite as
far up the valley as the hut back of Kamoku.” (O. P. Emerson
label in MCZ.) Also collected by Wilder, BBM 50605, and Spalding
in Kawaiiki Gulch. The Wilder lot has no label but is localized
from the Spalding collection. Spalding collected with Wilder, and
on his authority the approximate area 98? is marked on fig. 5a,
page 105.
The usual form and color pattern of the shell (pl. 3, fig. 28) has
been selected from the Baldwin and Lyman lots. The embryonic
whorls are light buff; postembryonic whorls pale ochraceous salmon
with a faint band on the last two whorls just above the edge of the
periphery of pale cinnamon pink, on the last whorl, below this band,
is a line of the same color; impressed sutural band the ground color ;
lip and columella callus pale ochraceous salmon. Length 19.6 mm.,
greater diameter 13.1 mm., spire height 10.8 mm.
In a lot in the Wilder collection the usual form (pl. 12, fig. 24)
has a more obese appearance, and also shows a pinkish-hued form
which does not occur in the type lots; embryonic whorls pale pinkish
buff ; first postembryonic whorl and a half pale pinkish buff faintly
streaked with pinkish buff, last postembryonic whorl and a half
vinaceous pink spirally banded and lined with pale vinaceous fawn;
impressed sutural band pale pinkish cinnamon; lip and columella
callus white or a dilute tint of tilleul buff. Length 19.1 mm., greater
diameter 13.4 mm., spire height 10.5 mm..
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 175
A narrow shell (pl. 12, fig. 25) measures: Length 19.5 mm.,
greater diameter 11.6 mm., spire height 11.5 mm.; embryonic whorls
white; first two postembryonic whorls a dilute tint of tilleul buff,
last whorl a very dilute tint of seashell pink spirally banded and
lined above and below the wide peripheral band with seashell pink.
ACHATINELLA APEXFULVA KAHUKUENSIS Pilsbry and Cooke
PLATE 3, FIGURE 34; PLATE 11, FIGURES 32-33a
Achatinella valida kahukuensis Ptrssry and Cooke, Man. Conch., vol. 22, p. 338,
pl. 52, figs. 17-17a, 1914.
To quote from Pilsbry and Cooke:
The shell is dextral, white above, yellow below the periphery, 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 perceptibly concave, the last whorl swollen. Lip
moderately thickened, white; columella very faintly rose-purple.
Length 20, diam. 13 mm.; 63 whorls.
Length 18.7, diam. 12.2 mm.; 64 whorls.
Oahu: Kahuku, at an elevation of 1,500 to 1,750 ft. (L. A. Thurston). Co-
types in collection A. N. S. and Bishop Mus., from Mr. Thurston’s Collection.
Distribution, area 115°: Malaekahana-Laie Ridge, Thurston
BBM 130973. From Thurston’s description of the locality, the ridge
is undoubtedly the Malaekahana-Kaluakauila Ridge. The exact
location is not known but tentatively may be placed in area 115?.
(fig. 6a, p. 185). Also collected by Wilder, BBM 50608, J. S. Emer-
son, BBM 102339.
The shell of Pilsbry’s plate 52, figure 17, is here reproduced on
plate 11, figure 32, and is considered the lectotype because it has the
usual color pattern of the shell. The color pattern of the lectotype
is similar to that of plate 3, figure 34, which is an obese form of the
shell, and has the first two and a half embryonic whorls worn and
cream color, last embryonic whorl white ; postembryonic whorls above
the periphery white, last whorl, below the peripheral band of burnt
umber, honey yellow with two central lines of cinnamon; outer
margin of the lip avellaneous, inner margin white, columella callus
light vinaceous fawn. Length 19.4 mm., greater diameter 13.5 mm.,
spire height 10.5 mm.
A lighter color pattern (pl. 11, fig. 33) has the embryonic whorls
light buff; postembryonic whorls white, shaded or faintly banded
with pale mouse gray, on the penultimate whorl a band of cinnamon
on the upper third of the whorl fading out on the last whorl, last
12
170 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
whorl banded at the periphery with a band of amber brown, below
the periphery the first half of the last whorl chamois shading to white
on the last half of the whorl, entire whorl lined with natal brown, the
last half of the whorl more profusely lined and banded than the first
half ; impressed sutural band wood brown; lip and columella callus
light vinaceous buff.
One shell in the Thurston lot (pl. 11, fig. 33a) has the post-
embryonic whorls white, banded just above or at the edge of the
periphery with natal brown, on the last whorl the band or line fades
to army brown and on the last half of the whorl to a line of cinna-
mon, within and just outside the aperture the shell callus honey
yellow; impressed sutural band white. Length 20.3 mm., greater
diameter 13.3 mm., spire height 10.8 mm.
A narrow shell measures: Length 18.0 mm., greater diameter 11.5
mm., spire height 10.3 mm.
Group oF A. A. APEXFULVA DIxon
ACHATINELLA APEXFULVA APEXFULVA Dixon
PLATE 3, FIGURE 27; PLATE II, FIGURES 17-19
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, 1780.
Achatinella apexfulva Dixon, Pilsbry and Cooke, Man. Conch., vol. 22, 317,
pl. 50, fig. 15; pl. 60, figs. 1-1b (only), 1014.
Turbo lugubris, etc., CHEMNitTz, Neues Syst. Conch. Cabinet, vol. 11, p. 278,
pl. 200, figs. 2059-2060, 1705.
Helix lugubris CueMNitz, Férussac, Tabl. Syst., Fam. Limagons, p. 56, 1821
(not Helix lugubris Gmelin, Syst. Nat., vol. 13, p. 3665, 1791).—FERUSSAC,
in Freycinet’s Voyage autour du Monde de l’Uranie et la Physicienne,
Zoologie, p. 479, 1824.—von Martens, Malakozool. Blatt., vol. 19, p. 40, 1872.
Achatinella lugubris CHEMNITz, Pfeiffer, Monographia Heliceorum Viventurum,
vol. 2, p. 230, 1848; vol. 3, p. 465, 1853; vol. 4 p. 542, 1859; vol. 6, p. 177,
1868.—REEvE, Conch. Icon., vol. 6, Achatinella, pl. 2, fig. 10a (not 10b),
April 1850.
Monodonta seminigra LAMARK, Hist. Nat. Anim. sans Vert., vol. 7, p. 37, 1822;
2d ed., Deshayes and Edwards, vol. 9, p. 181, 1845.—DeEtrEsseErt, Recueil de
Coq., Decrites par Lamark, pl. 37, figs. 2a-c, 1841.
Bulimus seminiger MENKE, Synops. Method. Moll., 2d ed., p. 26, 1830.
Achatinella pica SwAINson, Quart. Journ. Sci., Lit., and Art, January-March,
p. 84, 1828; Zool. Illustr., ser. 2, vol. 3, p. 99, pl. 99, fig. 1, 1832-1833.
To quote from Dixon:
eet tens the natives form necklaces, bracelets, 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,
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 77,
and is of a black-brown colour, 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 of Linnaeus, to which at first sight it appears
to be related. As I presume it to be a species hitherto undescribed, I have taken
the liberty to give it the trivial name of Apex Fulva, or the Yellow Tip. A
figure of it, in two views, is given in one of the following plates.
Specimens of this kind are in the Leverian Museum.
Distribution, area 97: “Picked before 1861 in Opaeula Valley on
the south side opposite ridge running down over the tunnel. Picked
on ieie [Freycinetia], kawau [keawau=Ilex], kopiko [Straussia],
kolea [koolea= Myrsine].” (O. P. Emerson label for his lot of A. a.
apexfulva in MCZ.) “Opaeula Gulch, Waialua,” J. S. Emerson,
BBM 102270, 102272; Kamoku, J. S. Emerson, BBM 33208;
Opaeula Gulch, locality 330-4, elevation 1,100 feet, Welch, K.
Emory, and W. Giffard, 15 dead dextral 1935. This locality is on
the south side of the Gulch and answers O. P. Emerson’s description
of his locality, and is probably the place where the Emerson brothers
got their material (see figs. 5, 5a, p. 105).
According to Murray (1904, p. 175) the Leverian Museum was
formed by Sir Ashton Lever (1729-1788) in the early years of his
life at his house, Alkrington Hall, near Manchester. In 1783 the
museum was sold by lottery. The lottery was won by James
Parkinson. In 1806 the museum was sold at auction. In a copy
of the sales catalog of the Leverian Museum, in my possession,
there are 10 lots which mention necklaces, bracelets, and other
ornaments, made of shells, from the Sandwich Islands. One of these
ornaments may be the one Dixon refers to, which contains the type
or paratypes of A. a. apexfulva. The lei (necklace) Dixon studied
may have been broken up and the specimens sold. Dixon’s original
figure is reproduced on plate 11, figure 17.
Pilsbry and Cooke (1914, p. 321) give such a good account of the
history of A. a. apexfulva that it seems unnecessary to review the
matter again. In discussing the origin of shell leis Pilsbry says:
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 obtained 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.
Pilsbry correctly considers Turbo lugubris, Monodonta seminigera,
and A. pica synonyms of A. a. apexfulva. The types of these forms
178 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
are not available to me and may be lost. It is possible that Jugubris
and seminigera come from the same lei Dixon used. If the shells
came from another lei there is a possibility that the shells are different
from apexfulva. But until the types are seen it is impossible to say
what the shells really are, and Pilsbry’s identification should stand.
Swainson’s shells were collected by Captain Byron of H. M. S. Blond.
These again could have come from elsewhere than the type locality
of apexfulva. Again until the type is studied it is difficult to say
whether the shell is or is not A. apexfulva. The Swainson shells may
be forms from area 94 of A. a. beata var. 1, which are impossible to
separate from typical A. a. apexfulva. Therefore, even were the
types available for all three forms they might be from different
localities and yet not separable from A. a. apexfulva, so that it is best
to consider them synonyms.
Since the holotype of 4. a. apexfulva has not been located, the
shells in the J. S. Emerson lot in the Bishop Museum, the Academy
of Natural Sciences, and the O. P. Emerson lot in the Museum of
Comparative Zoology will be considered typical. The usual form
and color pattern in the J. S. Emerson lot (pl. 3, fig. 27) has the
embryonic whorls light buff; postembryonic whorls hessian brown
or maroon; impressed sutural band the color of the ground with the
upper extreme edge pale pinkish buff; lip and columella callus pale
vinaceous fawn. Length 18.8 mm., greater diameter 12.3 mm., spire.
height 10.8 mm., number of whorls 64.
An extreme obese form (pl. 11, fig. 18) measures: Length 18.8
mm., greater diameter 13.3 mm., spire height 10.2 mm.; the color
pattern is similar to figure 27 except that the last whorl is faintly
streaked with tawny. Eight specimens in the Emerson lot of 38
specimens have a gray streaked pattern similar to plate 11, figure 18a;
last two postembryonic whorls maroon, streaked with pale quaker
drab, the streaks cut by lines of maroon. Length 19.1 mm., greater
diameter 12.3 mm., spire height 11.0. Pilsbry’s plate 60, figure Ic,
does not look like A. a. apexfulva, but rather like the lined form,
or A. a. aloha var. 1. No such color pattern is found in the J. S.
Emerson lots from Opaeula. The shell may be a rare color pattern
of an intermediate race which occurred between A. a. apexfulva and
A. a. duplocincta. Then again it may be a stray shell that was mixed
in with the shells of A. a. apexfulva.
An adult shell from locality 330-4 is shown on plate 11, figure 19.
eae ae
NO. I ACHATINELLA APEXFULVA DIXON—-WELCH 179
ACHATINELLA APEXFULVA DUPLOCINCTA Pilsbry and Cooke
PLATE 3, FIGURE 26; PLATE 12, Ficures 27, 28
Achatinella apexfulva color form duplocincta Pitspry and Cooke, Man. Conch.,
vol. 22, p. 323, pl. 55, figs. 6-8, 1914.
To quote from Pilsbry and Cooke:
The shell is dextral, white, encircled with two chestnut bands or group 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.”
Distribution, area 97A??: Type locality Kawailoa, Waialua, ‘“‘on
a group of 3 or 4 trees at the head of a little gulch beside and to the
north of the road to Kamoku, some two miles makai [toward the sea]
of Kamoku, collected by J. S. Emerson previous to the year 1863.”
(J. S. Emerson label, BBM 102312.) The original label on the O. P.
Emerson shells in the Museum of Comparative Zoology reads:
“Picked before 1861 in a secondary ravine branching S. from the
Kamoku reservoir valley below the narrows which is now a grassy
hollow. An old Akakea tree [Bobea sp.?] bore the shells—all now
extinct—and not found elsewhere by me.” The shell of Pilsbry and
Cooke’s plate 55, figure 8, here reproduced on plate 12, figure 27,
is selected for the lectotype. The label Wahiawa on the Cooke type
lot is surely erroneous, because the shells came from J. S. Emerson
who obtained his specimens from Kawailoa. From the above data of
the Emerson brothers, A. a. duplocincta was a lowland race that
occurred below the colony of A. a. apexfulva, which probably occurred
in Opaeula Gulch area 97 and was somewhere along the road on the
top of the Opaeula-Kawailoa Ridge. Area 97A?? is the probable
region of the type locality (fig. 7, p. 194).
The usual form and color pattern in the type lot in the J. S. Emer-
son collection (pl. 3, fig. 26) is a white shell, with a single line of
cinnamon above the periphery on the last whorl, and four lines of
mikado brown below the periphery; lip and columella callus light
pinkish cinnamon. Length 17.9 mm., greater diameter 11.8 mm.,
spire height 10.0 mm., number of whorls 63.
A narrow specimen and darkest color form (pl. 12, fig. 28) is
a white shell banded just above the periphery on the last two whorls
with two bands of mikado brown, above which on the last whorl and
180 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
.
a half are three faint lines of cinnamon, base banded with four bands
of burnt umber. Length 17.3 mm., greater diameter 10.7 mm., spire
height 9.6 mm. An obese shell measures: Length 17.1 mm., greater
diameter 12.6 mm., spire height 9.2 mm.
ACHATINELLA APEXFULVA BAKERI, new subspecies
PLATE 3, FIGURE 30; PLATE I1, FIGURES 20-20)
The shell is small, the last whorl is usually yellow, and has a
characteristic yellow subperipheral band. The form is extinct today.
It probably came from an isolated lowland locality. No intermediates
have been located by me connecting it to any other race of A. apex-
fulva. The strongly axially streaked color pattern is similar to
A. a. apicata or A. a. wahiawa. The presence of the subperipheral
band may cause this form to be confused with A. a. gulickit. But
A. a. bakeri mainly differs from both forms in the yellow color pattern
of the last whorl.
The embryonic whorls of the holotype (pl. 3, fig. 30) are light
ochraceous buff; first half postembryonic whorl vinaceous fawn
axially streaked with white, next half whorl pale drab gray axially
streaked with light brownish drab, last whorl and a half naples
yellow axially streaked with straight and zigzag streaks, speckled and
lined with mars brown, below the edge of the periphery a band of
naples yellow; impressed sutural band pale pinkish buff; lip and
columella callus tilleul buff. Length 16.4 mm., greater diameter 11.5
mm., spire height 8.5 mm. The usual form was not determined because
of the paucity of specimens.
Distribution, area?: Waimea, J. S. Emerson, BBM 102333, A. F.
Judd ex J. S. Emerson, BBM 162334. “Found on a Plateau north
of Waihalona [Kawaihalona?] Gulch below North Branch on pua
[Osmanthus| tree on a clump in open plain.” (O. P. Emerson
collection, MCZ.) This form is named after Dr. H. B. Baker, of the
University of Pennsylvania.
The shell may be narrower (pl. 11, fig. 20) and have a vivid
yellow ground; the first postembryonic whorl brownish drab axially
streaked with white or pale drab gray, last two whorls yellow ocher
axially streaked with natal brown, just below the edge of the periphery
a band of yellow ocher. Length 18.2 mm., greater diameter 11.5 mm.,
spire height 10.5 mm.
A lighter color pattern and an obese shell (pl. 11, fig. 20a) has the
first half postembryonic whorl axially streaked with light cinnamon
drab and white, penultimate whorl pale pinkish cinnamon or white,
INOS) I ACHATINELLA APEXFULVA DIXON—WELCH 181
streaked with light brownish drab, last whorl cream buff lightly
streaked with straight and zigzag streaks of verona brown, about the
periphery and base, bands of cream buff. Length 17.5 mm., greater
diameter 11.8 mm., spire height 8.7 mm.
The last whorl may not have a wide yellow peripheral band (pl. 11,
fig. 20D) ; first postembryonic whorl axially streaked with brownish
drab or deep brownish drab and pallid mouse gray, penultimate whorl
streaked with deep quaker drab and pallid mouse gray, last whorl
pale pinkish buff shaded with cream buff axially streaked with sepia
and warm sepia. Length 17.3 mm., greater diameter 11.0 mm., spire
height 9.3 mm.
ACHATINELLA APEXFULVA LEUCOZONA Gulick
PLATE 3, FIGURE 31; PLATE 11, Figures 28-31
Apex leucozonus GULICK, Proc. Zool. Soc. London, 1873, p. 83, pl. 10, fig. 6.
Achatinella valida cinerosa PretrFer, Pilsbry and Cooke, Man. Conch., vol. 22,
PP. 337, 338, pl. 55, figs. 9-16, 1914.
The embryonic whorls of the holotype (pl. 11, fig. 29) are cartridge
buff, last half embryonic whorl banded on the lower fourth of the
whorl with a band of a light tint of isabella color; postembryonic
whorls pale drab gray, first two postembryonic whorls axially streaked
with benzo brown and lined with drab, last whorl banded and axially
streaked with snuff brown darkening to verona brown on the last half
whorl. Length 18.2 mm., greater diameter 12.7 mm.
Distribution, area?: Type locality Waialei [Waialee], Gulick,
BBM, ANSP 92656; Waialee, J. S. Emerson, BBM 102337. Holo-
type, MCZ 39906.
A. a. leucozona is either a highland or a lowland form of A. a. napus.
The shell of plate 11, figure 28, looks like an intermediate form be-
tween A. a. napus and A. a. leucozona but closest to the color pattern
of A. a. leucozona. The embryonic whorls are light buff; first post-
embryonic whorl light buff, streaked with mikado brown, penultimate
whorl a yellowish tint of avellaneous streaked with wood brown, and
lined and banded with pinkish buff, last whorl above the periphery
banded and lined with pinkish buff and a narrow and a wide band
of buckthorn brown, at the edge of the periphery a band of light buff,
and a subperipheral band of raw sienna, base raw sienna banded
with a wide band of a light tint of antique brown axially streaked with
antique brown, below which is a line of argus brown.
The usual form of the shell (pl. 3, fig. 31) in the Gulick ANSP
lot has the embryonic whorls white; first two postembryonic whorls
182 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
light drab faintly spirally lined with white and strongly axially
streaked with cinnamon drab, last whorl wood brown axially streaked
with straight and zigzag streaks of wood brown, fawn color, and
light pinkish buff, and faintly spirally lined with white and wood
brown, just below the edge of the periphery a line of white ; impressed
sutural and subsutural bands white ; lip pale ochraceous buff, columella
callus white. Length 17.7 mm., greater diameter I1.4 mm., spire
height 9.9 mm.
The lightest color pattern and obese form of the shell (pl. 11,
fig. 30) has the first postembryonic whorl pale drab gray, banded
and streaked with light cinnamon drab, penultimate whorl pale ecru
drab banded and streaked with ecru drab, first half of the last whorl a
light tint of pale ecru drab, lined and streaked with ecru drab, last half
whorl pale vinaceous fawn, lined and faintly streaked with light
vinaceous fawn; impressed sutural and subsutural bands white.
Length 17.7 mm., greater diameter 12.0 mm., spire height 9.3 mm.
A darker color pattern than the usual one and a narrower form of
the shell (pl. 11, fig. 30a) has the postembryonic whorls snuff brown
axially streaked with russet and mars brown, and lined with a line
of mars brown on the upper third of the whorl, about the periphery a
band of white, lined or banded with mars yellow; impressed sutural
band light ochraceous buff lightening to light buff on the last whorl.
Length 17.7 mm., greater diameter 10.8 mm., spire height 9.6 mm.
The embryonic whorls of the darkest color pattern (pl. 11, fig. 31)
are light buff shading to ochraceous salmon; postembryonic whorls
warm sepia lightened by occasional axial streaks of mikado brown, last
whorl banded just below the edge of the periphery with a band of pale
cinnamon pink lined and streaked with mars yellow, on the first half
of the last whorl a basal band of ochraceous orange ; impressed sutural
band pale pinkish buff darkening to light ochraceous buff on the last
half whorl; lip and columella callus pale pinkish cinnamon deepening
to pale vinaceous fawn on the outer margin. The color pattern is a
rare one in the Gulick ANSP lot, but the usual one in the J. S.
Emerson lot.
ACHATINELLA APEXFULVA NAPUS Pfeiffer
PLATE II, FIGURES 25-27
Achatinella napus PFreIrFER, Proc. Zool. Soc. London, 1855, p. 5, pl. 30, fig. 19.
Achatinella mustelina sordida Newcoms, Pilsbry and Cooke, Man. Conch., vol.
22, p. 349, pl. 30, fig. 19 (only), 1914.
The holotype (pl. 11, fig. 25) in the British Museum is marked
with a red spot of sealing wax. The embryonic whorls are white;
NO: I ACHATINELLA APEXFULVA DIXON—WELCH 183
first two postembryonic whorls cartridge buff, last whorl above the
periphery vinaceous buff, at the edge of the periphery a band of
cartridge buff or white, below the periphery banded with vinaceous
buff, basal band about the umbilicus cartridge buff or white with
faint lines of vinaceous buff ; lip cartridge buff banded with vinaceous
buff. Length 19.4 mm., greater diameter 10.8 mm., spire height
12.0 mm., number of whorls 63.
Distribution, area?: Waialei [Waialee], Gulick.
The shell may have a yellow base (pl. 11, fig. 26) embryonic whorls
white; postembryonic whorls pale cinnamon pink; suture and sub-
sutural bands slightly more dilute tint of cinnamon pink; peripheral
band white, below the periphery chamois; lip and columella callus
tilleul buff. Length 19.9 mm., greater diameter 12.0 mm., spire
height 11.6 mm.
A larger specimen (pl. 11, fig. 27) shows a darker pink color
pattern and a more inflated last whorl. The postembryonic whorls
are pale vinaceous fawn, banded on the lower half of the whorl
above the periphery with a band of light vinaceous buff which darkens
to vinaceous buff streaked with light vinaceous buff on the penultimate,
and on the last whorl buff pink streaked with vinaceous buff, at the
edge of the periphery a band of white, base yellow ocher lined with
buckthorn brown; lip vinaceous buff, columella white. Length 20.0
mm., greater diameter 12.5 mm., spire height 11.5 mm.
ACHATINELLA APEXFULVA PAUMALUENSIS, new subspecies
PLATE 3, FIGURE 32; PLATE I1, FIGURES 21, 21a
The subspecies is a highland gray race closely related to A. a. napus.
The embryonic whorls of the holotype (pl. 3, fig. 32) are pale pinkish
buff with a line of pinkish buff in the center of the lower third of the
whorl; first postembryonic whorl upper half white or pallid mouse
gray, lower half sorghum brown shading to pale mouse gray axially
streaked with zigzag lines of benzo brown, remaining whorls spirally
banded with fuscous or fuscous black, ground pale mouse gray up
to the last half whorl which is shaded with light vinaceous fawn,
below the periphery last whorl isabella color banded with fuscous ;
upper third of the impressed sutural band white or tilleul buff,
remainder of the band shading from vinaceous buff to vinaceous fawn
and fawn color on the last half whorl. Length 18.3 mm., greater
diameter 12.2 mm., spire height 10.0 mm., number of whorls 64.
Distribution, area 111: Paumalu-Kaunala Ridge, type locality 431
elevation 1,000-1,050 feet, Meinecke, 1933 (fig. 6, p. 185). Alsa
collected by Thurston, BBM 1309015.
184 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
The shell may be more strongly banded (pl. 11, fig. 21) and more
obese ; postembryonic whorls pallid mouse gray faintly axially streaked
with mouse gray, above the periphery spirally ornamented with faint
lines and a band of warm blackish brown, base naples yellow banded
with warm blackish brown; the impressed sutural band vinaceous
fawn on the first whorl, on the penultimate the ground color, on the
last whorl the base of the band shaded with warm blackish brown.
Length 18.8 mm., greater diameter 13.0 mm., spire height 9.5 mm.
One specimen (pl. 11, fig. 21a) has the postembryonic whorls warm
blackish brown, banded on the first whorl with white, the remaining
whorls with extremely faint lines of white or pallid mouse gray.
Length 18.0 mm., greater diameter 12.1 mm., spire height 8.8 mm.
4
ACHATINELLA APEXFULVA OIOENSIS, new subspecies
PLATE 3, FIGURE 33; PLATE 11, FIGURES 22, 22a
The shell is a close relative of A. a. pawmaluensis, but having a
series of different color patterns; the embryonic whorls of the holo-
type (pl. 3, fig. 33) are pale pinkish cinnamon ; postembryonic whorls
above the periphery diamine brown, below the edge of the periphery
of the last whorl ochraceous buff finely axially streaked with ochra-
ceous tawny, spirally banded with a band of diamine brown and fine
lines of russet, in the umbilical region a patch of black or diamine
brown; impressed sutural band diamine brown; lip and columella
callus fawn color. Length 18.0 mm., greater diameter 11.5 mm.,
spire height 10.3 mm.
Distribution, area 114: Oio-Oio East Branch Ridge, type
locality 460A, elevation 1,300 feet, Meinecke, 1933 (figs. 6, 6a,
p. 185).
The color pattern may lack the yellow base (pl. 11, fig. 22) ; post-
embryonic whorls liver brown, last whorl and a half faintly banded
above the periphery, last whorl just below the edge of the periphery
with a band of mars yellow almost obscured by axial streaks of liver
brown; lip and columella callus pale vinaceous fawn, outer edge of the
lip bone brown; impressed sutural band the color of the ground. The
lightest color pattern (pl. 11, fig. 22@) has the postembryonic whorls
light buff deepening to ochraceous buff on the last whorl, axially
streaked with tawny, last whorl and a quarter at the edge of the
periphery and about the base banded with diamine brown; impressed
sutural band black. Length 17.0 mm., greater diameter 12.3 mm.,
spire height 8.7 mm.
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 185
ACHATINELLA APEXFULVA OIOENSIS var. 1
PLATE I1, FIGURES 23-23b
Distribution, area 112: Pahipahialua Gulch, locality 450, eleva-
tion 1,250 feet, Meinecke, 1933 (figs. 6, 6a, below) ; also collected by
Wilder, BBM 50612.
The color patterns of area 112 at first glance seem the same as
those of typical oioensis (area 114), but close examination shows that
°3) Koloa Sra
Gulch
Namahana
Stream
Fic. 6—Northern Oahu, windward slope of the Koolau Range, the northwestern
half of region IV (fig. 7, p. 194), showing localities of subspecies of A. apexfulva
belonging to the groups of A. a. apexfulva and A. a. aloha.
iw
Hoolapa >
Namahana
Stream
Fic. 6a.—Same as fig. 6, but showing the areas of distribution of the subspecies of
A. apexfulva belonging to the groups of A. a. apexfulva and A. a. aloha.
slight variations occur. Usually the yellow ground color is lighter in
area 112. A close reproduction of the patterns of plate 3, figure 33,
and plate 11, figure 22a, is not found in area 114. Plate 11, figure 23,
resembles figure 22a, but the characteristic wider sutural band of the
yellow patterns of area 112 separates it from the Oio Stream forms
of area 114. Moreover, dull brown forms with faint zigzag streaks
occur in area 112 and do not occur in area I14.
The usual color pattern and the obese form of the shell (pl. 11,
fig. 23) has the embryonic whorls pale pinkish buff; postembryonic
whorls light buff very finely lined and streaked with ochraceous
186 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
tawny, at the edge of the periphery a thin band of dresden brown,
base seal brown or black; sutural and subsutural bands verona brown
darkening to seal brown or black on the last two whorls; lip fawn
color, columella callus pale vinaceous fawn. Length 17.3 mm., greater
diameter 12.6 mm., spire height 8.9 mm.
A zigzag pattern and elongate form of the shell is shown on
plate 11, figure 23a; last two postembryonic whorls are verona brown
faintly axially streaked with zigzag and straight streaks of tawny
olive; impressed sutural band the ground color. Length 19.0 mm.,
greater diameter 12.3 mm., spire height 10.4 mm.
Only four live specimens are known from locality 450. A dark color
pattern (pl. 11, fig. 23b) seems to be an intermediate color form
between figures 22 and 23a of plate 11. It may be fairly close to the
usual form; the last two postembryonic whorls are seal brown, at
the edge of the periphery a line of yellow ocher, below the periphery
a band of ochraceous buff axially streaked or spotted with seal brown,
the seal brown or black base is also streaked with patches or streaks
of ochraceous buff. Length 18.0 mm., greater diameter 12.5 mm.,
spire height 9.2 mm.
ACHATINELLA APEXFULVA OIOENSIS var. 2
PLATE II, FIGURES 24-24d
Area 113: Pahipahialua-Oio Ridge, locality 460, elevation 1,150-
1,200 feet, 3 dextral; 461, elevation 1,250-1,300 feet, Meinecke, 33
dextral 1933; Kaunala-Oio Ridge, locality 462*, elevation 1,300-
1,376 feet, Meinecke, 1914 (figs. 6, 6a, p. 185). This form also col-
lected by J. S. Emerson, BBM 102335-102336, O. H. Emerson, BBM
103986, Thurston, BBM 130954, 130950.
The shells resemble those of area 112 but have a different range
of color patterns. The usual form and color pattern of the shell
(pl. 11, fig. 24) has the embryonic whorls pale pinkish buff; first
postembryonic whorl haematite red axially streaked with straight and
zigzag streaks of pale olive gray, the last whorl and a quarter above
the periphery lighter in color, being more streaked with pale olive
gray and banded at the edge of the periphery with a band of pale
olive gray tinted on the first half of the whorl with chamois, the
base warm sepia lined with chamois; impressed sutural band mikado
brown; lip and columella callus pale vinaceous fawn. Length 19.7
mm., greater diameter 12.8 mm., spire height 10.6 mm.
The color pattern may be lighter and the base yellow (pl. 11,
fig. 24a); first postembryonic whorl and three-quarters cinnamon
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 187
drab faintly axially streaked with drab gray, below the impressed
sutural band of fawn color two lines of tilleul buff, on the last whorl
the ground above the periphery tilleul buff faintly axially streaked
with straight and zigzag streaks of avellaneous, base mustard yellow
with a band of avellaneous just below the subperipheral mustard
yellow band, umbilical region lined with tawny.
An elongate shell (pl. 11, fig. 24b) and a variant of figure 24a has
the postembryonic whorls white, tinted, faintly lined, and faintly
axially streaked with tilleul buff or vinaceous buff; sutural and wide
subsutural bands and the line below the subsutural band wood brown ;
on the last whorl a thin band at the edge of the periphery of drab,
below which the ground is chamois banded with wood brown. Length
18.2, greater diameter 11.8 mm., spire height 10.3 mm.
The lightest color pattern (pl. 11, fig. 24c) has the white post-
embryonic whorls above the periphery lined with a faint line of
naples yellow on the last whorl, below the periphery naples yellow
with a central line of cinnamon, and a line and a band of russet in
the umbilical region ; sutural and wide subsutural bands chocolate.
A yellow color pattern (pl. 11, fig. 24d) has the first two post-
embryonic whorls pale pinkish buff, last whorl light buff finely spirally
lined and axially streaked with ochraceous tawny, on the base a thin
band of russet and faint lines of tawny; impressed sutural band and
thin subsutural band mars brown. The wide subsutural banded form
of figure 23 also occurs in this area.
ACHATINELLA APEXFULVA IHIIHIENSIS, new subspecies
PLATE 3, FIGURE 35; PLATE 11, FIGURE 34
The form is closely related to A. a. kahukuensis but differs in color
pattern. The color pattern of the holotype resembles that of A. a.
oioensis, but thithiensis has a base colored a lighter shade of yellow.
The embryonic whorls of the holotype (pl. 3, fig. 35) are cream color ;
postembryonic whorls warm blackish brown; impressed sutural band
cinnamon; edge of the periphery and all below the periphery of the
last whorl mustard yellow, banded with a central band of warm black-
ish brown and two lines of mikado brown; lip and columella callus
light vinaceous fawn, lip streaked with white. Length 17.7 mm.,
greater diameter 11.6 mm., spire height 9.6 mm., number of whorls 6.
Distribution, area 116: Thiihi-Kahawainui Ridge, type locality
510, elevation 1,250-1,300 feet, Meinecke, 12 dextral 1932-1934;
also locality 510-2a, elevation 1,150-1,200 feet, Welch, 10 dead
specimens, 1935 (figs. 6, 6a, p. 185).
188 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
The shell may have a lighter color pattern (pl. 11, fig. 34) ; embry-
onic whorls light buff, faintly lined on the last half whorl with
ochraceous tawny; first postembryonic whorl white, shaded with
pale gull gray and banded with a central band of mars brown in the
center of which is a black line, a lower band just above the periphery
bone brown, divides into two lines on the last half of the penultimate,
the ground on the last half of the penultimate is faintly shaded with
mustard yellow, last whorl mustard yellow, banded above, below, and
at the periphery with a band of bone brown at the lower edge of
which is a band of mikado brown, just above the edge of the periphery
are two lines, an upper of mikado brown, and a lower of bone brown,
both of which fade to a lighter tint on the last half whorl; the im-
pressed sutural band darkens from mikado brown on the first whorl
to warm sepia on the last whorl. Length 18.5 mm., greater diameter
12.6 mm., spire height 9.5 mm.
ACHATINELLA APEXFULVA WAILELENSIS, new subspecies
PLATE 3, FicuRE 36; PLATE I1, FIGURE 35
The shell is not closely related to any of the known forms of
A. apexfulva. The locality occurs far away from all the known
localities of A. apexfulva and no intermediates between wailelensis
and any other form are known. The shell, however, has the usual form
of A. apexfulva and resembles A. a. roseata of Waimano Stream. The
holotype (pl. 3, fig. 36) has the embryonic whorls white; first post-
embryonic whorl fawn color, last two whorls vinaceous fawn; im-
pressed sutural band pinkish buff; lip pinkish buff, columella callus
white. Length 17.3 mm., greater diameter 12.0 mm., spire height
10.3 mm., number of whorls 6.
Distribution, area, 117: Wailele Gulch, locality 529-1, elevation
20 feet, collected by Mr. and Mrs. G. W. Russ, and Welch 1933. The
shells are found in a fossil state. in a stone outcropping near the
Kamehameha highway (fig. 7, p. 194).
An obese form of the shell (pl. 11, fig. 35) measures: Length 17.1
mm., greater diameter 11.9 mm., spire height 8.7 mm.; postembryonic
whorls shell pink axially streaked with buff pink. A narrow specimen
measures: Length 17.6 mm., greater diameter 11.2 mm., spire height
9.4 mm.
CONCLUSIONS
Gulick (1905), Pilsbry and Cooke (1912-1914) considering various
species of Achatinella from Hawaii, and Crampton (1916, 1932) in
ee
NOSE ACHATINELLA APEXFULVA DIXON—WELCH 189
a study of the species of Partula on Tahiti and Moorea, have pointed
out that tree snails vary from valley to valley in various character-
istics such as size, color pattern, and form of the shell. Welch (1938)
pointed out that A. mustelina from the Waianae Mountains of Oahu,
Hawaii, not only varied from valley to valley in different character-
istics, but also at different elevations on the same ridge or in the same
valley.
Before discussing the variation of Achatinella apexfulva 1 will
explain what is meant by a highland and lowland form. On a basis
of a group of characteristics such as color pattern of the embryonic
and postembryonic whorls, shape and size of the shell, I have divided
the material into highland and lowland forms. On figure 7 (p. 194)
two broken lines drawn across the Koolau Range outline the three
zones of shell variation. Zone I contains extreme lowland forms and
is probably the region where many of the extinct Gulick forms were
collected. The zone occupies all the region below the lowest broken
line, which extends from just above areas 117, 97A??, 37?, to just
below areas I and 2.2 Zone II contains the usual lowland forms.
Gulick collected or obtained material from the lower part of this zone.
Zone II occupies all the region between the two broken lines. Zone
III is inhabited by highland forms and extends from above zone II
to the backbone ridge of the Koolau Range. Along the border of
zones II and III forms occur in certain areas which can be classed
as either highland or lowland shells on a number of characteristics.
The placing of these borderline areas into highland or lowland zones
has been a matter of judgment and might be done differently by
another worker. Areas 5??,6??, 7??, 147?, 65??, 667?, 78?, found on
figures 3, 4, and 5, have been omitted from figures 7 and 8 because I
am uncertain concerning the exact location of these areas.
In the following account of shell variation in each area, the usual
form * and color pattern of the shell, generally exhibited on plates 1,
2, and 3, is taken as a basis for discussion. In some cases it was not
possible to obtain a shell having both the usual form and color pattern.
In that case only one of the group of characteristics is shown on one
of the first three plates, such as the usual form of the shell or the
usual color pattern, and the other characteristic is shown on another
plate. In one case (pl. 1, figs. 27, 28) rare color patterns of a race
2 An area is made up of a single locality or a group of localities containing
similar forms or the same subspecies.
3 See Welch (1038) for an account of how the usual form of the shell is
selected.
Igo SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I03
are shown, which distinguish the area containing them from another
area of similar shells.
A. HorizonTaL VARIATION
I. COLOR PATTERN OR COLOR OF THE EMBRYONIC WHORLS
On the color of the embryonic whorls the subspecies of A. apexfulva
are segregated into four regions. Regions I and III have shells with
bicolored embryonic whorls, and regions II and IV have unicolored
embryonic whorls (fig. 7, p. 194). These four regions are further
subdivided into intermediate regions Ia (area 19), Ila (areas 48,
48A, and the localities of area 47 occurring on the Manana-Waiawa
Ridge), IIa (areas 41, 42, 43, 63, and 64), IIIb (areas 61? and 627),
which contain intermediate or border forms between those of the
main regions. The embryonic whorls of these border forms have a
color pattern similar to that found on the shells at a similar elevation
on the next parallel ridge to the southeast. This is also true of areas
40, 49, and 50.
In region IIIa the embryonic whorls are white, cream buff, or tan,
faintly lined or banded with a lighter or darker color. They are
intermediate between the white unicolored embryonic whorls of the
group of A. a. turgida (region II) and the yellow banded bicolored
embryonic whorls of the group of A. a. polymorpha (region III).
Region IIIb and areas 40, 49, and 50 definitely have the same
colored embryonic whorls as found on the adjacent parallel ridge to
the southeast. Region Ia contains shells with embryonic whorls which
are still bicolored similar to the group of A. a. simulans (region 1)
to the southeast but are approaching the unicolored condition of the
shells in region II. Region Ila has shells with slightly bicolored
embryonic whorls intermediate between region II and III, but closest
to the unicolored embryonic whorls of region II.
2. COLOR PATTERN OR COLOR OF THE POSTEMBRYONIC WHORLS
Many difficulties are encountered in distinguishing subspecies using
color pattern for a criterion, because the same color pattern occurs
again and again in colonies of shells in widely separated areas. For
example, the dark typical apexfulva pattern from area 97 or one very
similar to it, may be found in area 35 (A. a. turgida) (fig. 7, p. 194),
area 94 (A. a. beata var. 1), area 111 (A. a. paumaluensis), area 114
(A. a, oioensis). Pink patterns such as A. a. lilacea var. 1 in area 83
and A. a. lilacea possibly in area 83b?? may also be found in area 81? ?
NO. I ACHATINELLA APEXFULVA DIXON—WELCH Ig!
(A. a. punicea), area 37° (A. a. roseata), and area 117 (A. a.
watlelensis). Somewhere near area 40 Wilder found specimens of
A. a. perplexa (pl. 6, fig. 19d) which have a pattern similar to that of
A.a. lilacea. The various forms of A. a. rubidipicta can easily be taken
for specimens of A. a. rubidilinea. White color patterns of A. a.
ovum, (area 22), A. a. cookei var. 1 (area 39), A. a. beata var. 2
(area 108), and A. a. roseipicta (area 102), are very similar, differing
only by a band or in the color of the lip. However, the range of color
patterns of any one area always differs from that of another. There-
fore, in considering horizontal variation not only the usual color
pattern of the shell must be considered, but also the color pattern
of the colony as a whole.
As has been pointed out before, A. apexfulva exhibits valley-to-
valley or ridge-to-ridge variation, and the color patterns of widely
separated localities are usually more strikingly different than those
found on shells from adjacent areas. However, as in A. mustelina
(Welch, 1938), A. apexfulva also exhibits vertical as well as horizon-
tal variation. Therefore, when studying horizontal valley-to-valley
variation it is important that both localities or areas under considera-
tion are from approximately the same elevation or in the same low-
land or highland zone such as zones I, IJ, or III.
I do not like to use the word elevation because two localities on
opposite ridges may be at an equal distance from the backbone ridge
and yet because of a hill on one ridge and a depression on the other,
one locality may be definitely higher than the other. Again, one
locality may be in a valley, the other on a nearby ridge, the two
showing a wide difference in elevation. The distance a locality is
from the backbone ridge or in what zone a shell is found is of greater
importance than differences of several hundred feet in altitude.
Therefore, zones and not differences in elevation will be discussed.
I have already noted above that the color of the embryonic whorls
varies in regions I, II, III, IV. The color pattern of the shell not
only shows marked variation between each of the four regions, but
also less variation within each region. For instance a series of areas
in region II zone II will have rather similar color patterns. In like
manner a group of areas in region III zone II will have another
series of similar patterns but which are markedly different from those
found in region II zone II. In most areas occurring at similar eleva-
tions or in the same zone on opposite ridges in a certain region the
color pattern of the shells are similar. However, in the region of
Waimano Stream the color patterns of such opposite areas—for
example, areas 44 and 43—are not similar. Area 44 has patterns
13
192 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
similar to those of area 50 above area 43 (fig. 4, p. 53). Also, the
shells of area 45 on the Waimano-Manana Ridge more closely
resemble the shells of area 49 above area 50, although area 50 is
opposite area 45. In like manner the color patterns found in any
zone in region III will definitely differ from those found in region
IV. The same holds true for regions I and II.
In between the regions of major differentiation intermediate or
border areas occur, such as regions Ia, IIa, IIIa, IIIb, and areas 4o,
49, 50. Region Ia contains forms which have a color pattern inter-
mediate between regions I and II, zone II, but closer to the color
pattern of the shells to the southeast, or region I. Region Ila and
IIIb and areas 49 and 50 have shells the color pattern of which is
very similar to that of the shells to the southeast and entirely different
from those to the northwest in the same zone. Region II]la and area
40, zone II, contain gray color patterns which are close to the color
patterns of region III, zone II, and very different from those to the
southeast.
As a general rule more horizontal variation occurs in zone I] than
in zone III. This can be explained by the fact that usually the lower
reaches of a ridge fans out into several long ridges. Consequently,
there are more ridges and valleys at low elevations or in a lowland
zone, which increases the possible number of isolated colonies in
zones I and II as compared with zone III.
Today the isolation of the lowland colonies is more complete than
it was originally because of the dying out of the forest and the spread
of the staghorn fern (Gleichenia linearis). In this manner groups
of trees are isolated from other groups of trees so that snails cannot
possibly migrate. This is more especially true of lowland forms in
zone II and the lower portions of zone III. In many places trees
exist only in the bottom of a gulch the sides of which are covered with
staghorn fern, while the tops of the intervening ridges are covered
with grass. Again the trees on the side or summit of a ridge may also
be surrounded by staghorn fern.
In zone III, which includes the upper part of Hosaka’s (1937)
Ohia Zone and all of the Cloud Zone (fig. 1, p. 3), the forest is more
continuous, although in many regions large sections are covered with
staghorn fern. One would expect more horizontal migration here
because of the relatively larger amount of flora, greater rainfall, and
the connection of the backbone ridge with the main ridges of the
Koolau Range, which would afford a road for migration. On the
other hand the peaks of the backbone ridge might also serve as
effectual barriers. But A. apexfulva does not occur at the backbone
NOS ACHATINELLA APEXFULVA DIXON—WELCH 193
ridge. When it is collected on a ridge, the highest locality is usually
a mile or more below the backbone ridge. An exception to this is
area I, which is nearer the backbone ridge. Area 115? is only an
approximate locality and so cannot be considered (fig. 7, p. 194). The
Nuuanu localities of area 10 are on subridges in the valley separated
from the backbone ridge by a high precipice. On the other hand,
although the forests are fairly continuous in zone III, migration from
one ridge to another would be impeded by the gulches which contain
water.
Whether there is any horizontal migration in zone III is not known.
The forms in regions I, II, III, and IV, zone III, are certainly
distinct from each other but have color patterns that are more similar
to each other than are the color patterns in the same regions in zone II.
I do not believe that the more extreme horizontal variation in zone II
is entirely due to isolation which has come about in the last 50 or 100
years, because forms of A. apexfulva seem to colonize limited areas
and to break up into different varieties or subspecies at different eleva-
tions or at short distances from an adjacent colony. This is especially
true of zone II, which has less rainfall than zone III and consequently
would tend to discourage migration. Nevertheless, the present-day
extreme isolation and inbreeding of certain groups of genes un-
doubtedly accentuates the former horizontal valley-to-valley variation.
The characteristic of A. a. apexfulva of breaking up into varieties in
limited areas differs from what Crampton found (1916, 1932) in
species of Partula, which apparently migrate into various adjacent
areas and do not change color pattern in doing so.
3. SIZE
Table 2 (p. 15) gives the statistics of length range, mean length of
shell, and number of dextral and sinistral shells found in each locality
having a series of five shells or more with a length range of at least
three sizes such as 18.5 mm., 19.5 mm., 20.5 mm. The means of the
shells are grouped into length classes. For example, everything be-
tween 17.76 mm. and 18.25 mm. is considered 18.0 mm. and everything
between 18.26 mm. and 18.75 mm., is taken to be 18.5 mm. This was
also done in the paper on A. mustelina Mighels (Welch, 1938). In the
following discussion on size variation the mean length of the shell
will be referred to without needlessly repeating the words “mean
length of shell.”
The various localities or areas containing forms of A. apexfulva
are grouped for convenience into 20 ridge complexes or ridge groups
in order to demonstrate size variation. A ridge complex such as
3 (fig. 8, p. 195) may be a single ridge which fans out near the
VOL. 103
MISCELLANEOUS COLLECTIONS
SMITHSONIAN
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ACHATINELLA APEXFULVA DIXON—WELCH
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196 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
terminus into four main ridges and includes not only all localities on
these ridges but also all localities in valleys adjacent to or in between
these ridges. Again if only one or two areas are reported from one
ridge (4, 5, 12, fig. 8, p. 195) they are included with the ridge complex
of the adjacent ridge, or with an area in an upper or lower portion of
an adjacent ridge. With the exception of ridge complexes I, 19, 20,
which are groups of ridges, no ridge complex includes more than two
main ridges, which arise from the backbone ridge of the Koolau
Range. In compiling table 2, all forms from a ridge complex are
grouped together. Those from localities at the lowest elevations
(zones I and II) are given first and followed by forms from localities
at successively higher elevations (zone III). The exact localities of
most of the extinct Gulick forms are unknown, and they are placed
in table 2 next to the ridge complex which probably contained them.
On figure 8 (p. 195) size variation of A. apexfulva in the Koolau
Range is illustrated. The data given in table 2 are used for compiling
the map. Where space permits, the length of the shell for each
locality is plotted with the appropriate symbol. Usually there is not
enough room to plot each individual locality, so that a symbol may
stand for several localities or an entire area having the same shell
length. The shell length of the majority of localities in an area is
plotted where there is not enough room to plot the shell length of one
or more localities having different shell lengths. If three lots from
a single locality have three different lengths such as 17+, 18+, 19+,
the middle length of 18.5 is chosen for plotting. All data from
localities collected after 1932, or from localities I believe to be
reliably plotted, are used in preference to less reliable material such
as Wilder’s and that of other collectors which has been localized
by being matched with reliable material. Material which has been
localized from localized data is included because in some cases it
contains large series which furnish additional data as to the possi-
bilities of size variation of a form. In some cases the localized material
is represented by such small lots that it is of little statistical value.
Shells in highland localities (zone III) show as much horizontal size
variation as those in lowland localities (zone II). The shells in zone I
cannot be considered because of insufficient data. Little is known
about zone I, and the three widely separated localities on the map in
figure 8 are the only ones in zone I concerning which there are any
data. They show no horizontal variation. If more were known about
the location of the Gulick localities more variation would be noted in
zone I, The Gulick shells came from zone I and lower zone II. Only
NO. £ ACHATINELLA APEXFULVA DIXON—-WELCH 1Q7
the Bishop Museum’s portion of the Gulick collection is measured,
with the exception of one Gulick lot of A. leucozona in the Academy
of Natural Sciences of Philadelphia. This is approximately only 22
percent of the entire Gulick collection, which was broken up into 20
separate collections by Gulick. Many specimens were also traded
with people all over the world by Gulick, so that 22 percent may be
too high an estimate.
The Gulick material used, therefore, cannot be statistically signifi-
cant, but does give an approximation of size variation of at least
those series which are measured. In the region of ridge complexes 1
and 2 (table 2) 17+ shells are dominant in the Gulick lots, whereas
in the region of ridge complexes 14, 15, and 19 the mean length
ranges from 16+ to 17+. The 19+ forms from the region of ridge
complex 15 are probably from zone II.
Zone II shows marked size variation in certain regions. Groups of
ridge complexes have similar shell lengths and the change in some
regions from large to small shells is abrupt. Half the ridge com-
plexes (1, 4, 5, 7,4 8, 9, 15, 18, 19, and 20) contain mostly 18+ shells,
22 percent (ridge complexes 10, 14, 16, and 17) have mostly larger
19+ shells, and 27 percent (ridge complexes 3, 6, I1, 12, and 13) con-
tain small 17+ shells but may also have 18+ shells. Small 16+ shells
occur only in ridge complex 11.
Twenty-two percent of the ridge complexes (1, 10, 11, and 12) in
zone IIT have small 18+ or 17+ shells. The remaining 78 percent of
the ridge complexes, including number 2 (fig. 8, p. 195) have 19+
or 20+ shells, and in ridge complex 8, 21+ shells also occur. Only
the uppermost localities in ridge complex 2 contain 19+ shells, the
lower localities having small 18+ or 17+ shells. If size alone were
considered, these lower localities would be placed in zone II. In ridge
complex 3 (fig. 8, p. 195) what appears to be the highest area
contains 18+ shells. Actually this area (area 12, fig. 8) is on the
lower portion of a small spur ridge in ridge complex 3 and is at a
lower elevation than the other localities. Ridge complex 7 also has 18+
shells in zone III (area 31, table 2). This may be due to the
small number of shells measured; possibly if a larger series were
obtained, the mean length would be greater.
*Area 27A?, the lowest area in ridge complex 7, has a mean length of
18.77 mm. and is therefore represented in figure 8 by the symbol for the
19 + class shells. However, it really is very close to the 18.5 class and could
be considered an 18+ area.
198 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
B. VERTICAL VARIATION
I. EMBRYONIC WHORLS
No vertical variation in the color of the embryonic whorls occurs
in regions I, II, Ila (fig. 7, p. 194). Vertical variation occurs between
regions IIIa and the portion of region III immediately above IIIa,
and on the North-South Kaukonahua Ridge between regions IV and
IIIb. The lower regions have shells with unicolored embryonic
whorls (region IV) or with embryonic whorls that are intermediate
between the bicolored and unicolored condition (region IIIa). The
upper regions IIT and IIIb on the other hand have shells with bicolored
embryonic whorls (fig. 7, p. 194).
In area 77, between regions IV and IIIb, shells of A. a. tuberans
var. 3 have either unicolored or bicolored embryonic whorls. In
the Gulick collection specimens of A. a. polymorpha have either
bicolored or unicolored embryonic whorls. This may indicate that
at a low elevation (zone I) in the region of Waiawa and Kipapa
Gulch shells with unicolored embryonic whorls existed below the
present-day forms with bicolored embryonic whorls in zone II of
region ITI.
Region IV contains only shells with unicolored embryonic whorls
but northwest of North Kaukonahua Stream to Opaeula Gulch in the
lowest areas (areas 85, 86, 89, 90, 91??, 92, 93, 95, 96, 106, 107, 109,
fig. 5a, p. 105), the shells have dark embryonic whorls which are
some shade of brownish yellow such as ochraceous tawny or buck-
thorn brown. The upper localities have lighter embryonic whorls
with such colors as white, light buff, or pale pinkish buff. An excep-
tion to this rule is area 88, which contains shells with darker embryonic
whorls than lower area 87. Area 97 of Opaeula Gulch (ridge complex
18, fig. 8, p. 195) has shells with darker embryonic whorls than those
of area 99 above it. But area g7A ??, which is below area 97, has shells
with white embryonic whorls. So that in the region of Opaeula and
Kawailoa Gulch three distinct types of embryonic whorls occur : High-
land white or buff, lowland brownish, and finally white in the extreme
lowland shells. Northwest of Kawailoa Gulch no vertical variation
is noted in the color of the embryonic whorls in shells found today.
2. COLOR PATTERN OF THE POSTEMBRYONIC WHORLS
The postembryonic whorls show more vertical variation than the
embryonic whorls. At different elevations along a ridge different
colonies or areas with distinct color patterns are encountered. The
NO. I ACHATINELLA APEXFULVA DIXON—WELCH I99
vertical variation between two areas is usually more marked than
the horizontal variation. In studying the patterns of any area or
locality in relation to other forms, not only must the shells in areas
on adjacent ridges on either side be considered, but also those above
and below the area on the same ridge.
The color patterns of the Koolau Range can be divided into light
and dark. The light patterns are: 1, white strongly banded with
reddish brown; 2, white lightly banded or lined with reddish brown;
3, white banded with yellow or tan; 4, pink or flesh color, which may
be banded with white. The dark color patterns are: 1, gray mixed
with pink; 2, brown; 3, grayish brown or gray; 4, reddish brown;
5, reddish brown banded with yellow; 6, yellowish or tan; 7, gray
mixed with yellow.
The division of color patterns into light and dark forms in the
Koolau Range, while very close to the division of patterns between
zones II and III, is not the same. A dotted line has been drawn
(fig. 7, p. 194) showing the boundary between highland light color
patterns, usually present in zone III, and lowland darker patterns not
dominantly banded with white, which exist below the line in zone II
and lower zone III. With the exception of areas 30 and 33
(fig. 7, p. 194), which contain dominantly yellow patterns, all forms
above the dotted line are white forms banded with various colors,
usually reddish brown or yellow. In all areas below the dotted line,
with the exception of area 97A??,° not only are the shells darker in
color but also the patterns are axially streaked. Area 70, below the
dotted line, has white color patterns which are distinct from the
highland forms in being tinted with gray and in not having the white
ground color found on shells above the dotted line. Pink shells from
lowland areas (37?, 83b??) differ from pink shells found above the
dotted line in not being banded with white and in lacking white color
patterns.
Therefore, the color pattern of shells coming from areas of high
humidity (most of zone III) tends to be dominantly white and
banded, while shells with darker patterns which are banded and
streaked occur in dryer situations (lower zone III, zone II, and zone
I). Although banded patterns are not characteristic of highland or
lowland forms, streaked patterns with the exception of area g7A??,
seem to be characteristic of shells inhabiting dry situations. This cor-
5 Area 97A?? contains white color forms lightly lined and banded and not
markedly axially streaked. This locality is an exception to the general rule
concerning the color pattern of lowland color forms.
200 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
relation between pattern and elevation in A. apexfulva was noted by
Pilsbry and Cooke (1912-1914) and Welch (1938a). No correlation
exists between a particular pattern and a definite altitude.
The shells of area 1 (fig. 8, p. 195) are near the backbone ridge and
would be expected to contain white shells and be part of zone III.
However, area I contains lowland dark streaked gray forms although
it is at a higher elevation than area 3, where white shells occur. As re-
gards moisture conditions, area 1 is found to be in a section of rela-
tively low humidity as compared with most localities to the northwest
in zone III. Northeast of Nuuanu near the backbone ridge the rainfall
decreases the farther one goes to the northeast (compare Luakaha
(upper) with Makapuu, table 1, p. 10).
Region I, northwest of Palolo, shows less variation than the other
regions. This is probably due to the fact that the lowland localities
are all wiped out and only the upper less variable forms exist today.
However, from what is known of shells collected by Gulick and older
collectors in the region of Nuuanu-Kalihi, there existed a number of
small dark-colored races of shells below the present-day white races.
Area 15 is the only area in the Nuuanu-Kalihi region with markedly
dark color patterns.
Little vertical variation is shown in the Halawa section of region II.
Area 21 has white shells, above which in areas 22 and 23 the dominant
pattern is white but contains yellowish banded and lined forms which
are darker than those found in area 21. Two of these rarer darker
patterns from area 23 are shown on plate 1, figures 27 and 28.
The shells in area 26 are brown, banded with white, and are lighter
in color than those higher up the ridge, which are yellow banded with
brown in area 25. Above area 25, the usual color pattern is white.
North of Kalauao to Waimano, the lowland areas have shells with
brown patterns grading into yellow at a higher elevation and finally,
in area 31, into white. In Waimano, in area 34, brown patterns occur
which also grade to yellow and finally to white in the upper areas. But
below area 34, in area 35, reddish-brown patterns are found, and these
occur also in areas 44 and 50. Above these last two areas the shells of
the upper areas have patterns that contain more and more white and
are less heavily banded with reddish brown.
North of Waimano, in regions III, IIIa, and IV, the predominant
lowland patterns, zone II and lower zone III, are brown or gray with
here and there an area of reddish-brown, yellow, or pink shells.
Above these in zone III the color patterns are all white. Region IV
is the section of the Koolau Range which shows maximum vertical
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 201
differentiation. The contrast between lowland streaked and highland
banded patterns is very marked. On some ridges as many as 7 or 8
distinct color races are found (see pl. 2, figs. 24-30; pl. 3, figs. 1-8,
12-18, for examples of forms occurring on three ridge complexes).
North of Kawailoa Gulch no particular zoning is known to occur
today in ridge complex 19. But in Gulick’s day there must have been a
division between A. a. napus and the darker reddish-brown forms that
occur today in areas III, 112, 113, I14.
3. SIZE
Thirteen ridge complexes (2-6, 8-9, 13-15, 17, 18, and 20, fig. 8,
p. 195) show size increase in the majority of localities with increase
of elevation. The smaller lowland shells, with the exception of area 2,
are in zones I and II and are usually 17+ or 18+ mm., with here and
there a locality with 19+ shells. In area 2 all forms are in zone III,
but all the lowest localities have small 18+ or 17+ shells, similar
in size to those usually found in zone II, while the highest locality
contains 19+ shells. In all the remaining 12 ridge complexes showing
size increase in zone III, the shells are 19+ or 20+ mm. Although
there is a contrast between zones II and III the increase in size is
not a gradual one. The highest localities do not contain the largest
shells. Northwest of ridge complex 7, 19+ shells occur above 20+
shells in many localities. In ridge complex 8, 20+ shells occur above
21+ shells. Ridge complex 3, area 12 (table 2), has 18+ shells
and appears on figure 8 (p. 195) to be at a higher elevation than
the 19+ shells. However, the shells were collected on a small spur
ridge of ridge complex 3, probably at a lower elevation in Nuuanu
than the 19+ shells. The exact extent of the locality is uncertain.
The 18+ sinistral shells from area 62?, ridge complex 14 (table 2),
are reported from the same locality as the larger 19+ dextral shells,
and are separated because of the size difference. If both dextrals and
sinistrals are lumped, together, the mean length is 18.0 mm. The
locality was plotted 13 years after being collected and probably
represents two localities, one of dextral and one of sinistral shells.
On figure 8, ridge complex 14, 18+ shells are plotted above 19+
shells, which may be an error. Whether the dextral or sinistral shells
occur in separate localities, or one above the other, is of no great
importance. The small 18+ shells are of interest because they are
another exception to the general trend.
Out of the seven ridge complexes not showing altitudinal size
variation, two can be disregarded. One of these, ridge complex 19, is
202 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I03
all in zone II; the other, ridge complex 10, is represented by only
one locality in zone III, which contains only five specimens. The
remaining five ridge complexes show no marked variation in the
majority of localities between zones II and III. Ridge complex 1
contains only 18+ shells, regardless of altitude. If the Gulick shells
from this general region, probably zone I and lower zone II, are
compared with present-day forms, size variation .can be noted. The
lower Gulick forms are 16+ or 17+, shells. However, if all the
Gulick shells in institutions outside of the Bishop Museum collection
were measured and considered together, the mean length might be
greater, probably 18+ instead of 17+. The highest locality in ridge
complex 7 (area 31, table 2, p. 15) has 18+ shells, while the lowest
locality (area 27A?) has shells with a mean length of 18.77 mm.,
or 19+ shells. In between these localities, 19+ and 20+ shells occur.
The most remarkable exceptions to altitudinal size increase are
ridge complexes 11 and 12, which have small 17+ shells occupying
the majority of localities in both zones IT and III. In ridge complex
11, zone II], 16+ shells also occur. Localities of 18+ shells are
found here and there in both zones II and III in both these ridge
complexes.
As with A. mustelina (Welch, 1938), no clear-cut correlation can
be drawn between increase of moisture and size variation. The
highest localities in ridge complex 1 near the backbone ridge range
in elevation from 1,650-2,000 feet, while those in ridge complex 2
range from 1,150-1,500 feet. However, the rainfall in upper ridge
complex 2 is undoubtedly greater than in upper ridge complex 1, for
although there are no rain-gage stations in this region, south of
Nuuanu and Manoa the backbone ridge undoubtedly receives less
rainfall (compare Luakaha (upper) with Makapuu, table 1, p. 10), so
that in this case a correlation can be drawn between increase of
size and increase of moisture. In a similar manner the localities in
area 8, which is a much dryer section of Nuuanu Valley than area Io,
have smaller shells than the upper locality even though the elevation
of both areas is about the same (compare Luakaha upper and
Luakaha lower, table 1, p. 10). While a correlation can be drawn
_ between an increase of size with an increase in moisture conditions
in ridge complexes I and 2, it is not possible on the same basis to
explain the small shells of ridge complexes 11 and 12, zone III, or
the larger 19+ shells of ridge complexes 7, 16, and 17, zone II.
Even though exceptions occur at random, 65 percent of the ridge
complexes show a size change between zone II and III. A correla-
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 203
tion, then, can be drawn in the majority of ridge complexes between
increase of size with increase of moisture and lower temperature.
This is the reverse of what was found for A. mustelina, which species
shows a correlation between decrease of size and increase of moisture.
Therefore, these two different species react very differently to similar
environmental conditions, the reaction probably depending not only
on the genetic make-up of the individual but also on the influence of
the environment on the hereditary factors.
4. SHAPE
If the usual form of the shell from each area is considered and
the ratio of the length to the greater diameter is obtained, the usual
form of the shell is found to vary at random. Only in ridge complex
I5 is any vertical variation shown; here the lower areas contain
narrower shells than the highland areas. This general condition is
similar to that found in the case of A. mustelina (Welch, 1938).
In most areas in the Waianae Mountains A. mustelina showed no
vertical variation. Only in the northern section of the mountains is
the shape of the shell found to change from elongate lowland to more
squat highland forms.
C. VARIATION WITHIN A LOCALITY or AREA
I. COLOR PATTERN OF EMBRYONIC AND POSTEMBRYONIC WHORLS
Each subspecies in a given locality has a range of color patterns
from light to dark. The dark color patterns of shells from some areas
in zone III in regions I and II are often very different from the usual
white color form of the shell. In area 8 (pl. 4, fig. 15), area 12
(pl. 4, fig. 20b), area 11 (pl. 5, fig. 2b), area 10 (pl. 5, fig. 1a), and
area 46 (pl. 7, fig. 14@) dark reddish-brown forms occur which would
never have been expected in an area of white shells. One of the
most striking examples of this occurrence of two widely differing
color patterns from the same locality is to be seen in the region of
the North Kaukonahua area 103 of A. a. roseipicta var. 1. The domi-
nant color pattern (pl. 3, fig. 8) is a pinkish one banded with white, and
the other color patterns are white ones banded with pink. With these
patterns, a few rare dark gray color forms occur (pl. 12, fig. 6) which
are strikingly different. The same thing occurs in area 105. In area
100? the strikingly banded patterns of A. a. aloha var. I may be an
example of the same thing, but in this locality the patterns are not
only different but show a series of variants which may indicate a
separate area containing special color patterns.
204 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
2. FORM
A considerable amount of trouble has been taken to show the range
of form variation within each area. The extreme narrow and obese
forms of the shell are generally figured for each form and the measure-
ments given in the text. For example, various forms of A. a. turgida
have been figured on plate 6. The extreme obese form of the shell
(pl. 6, fig. 4a) looks very different from the narrow form of the
shell. The spire may be concave in outline (pl. 6, fig. 4e) or straight
in outline (pl. 6, fig. 5). The contrast between the narrow (pl. 5,
fig. 17) and the obese form (pl. 5, fig. 17a) of A. a. bruneola is
enough to lead to the consideration of the specimens, disregarding
color, as two different forms. In like manner specimens of A. a.
parvicolor (pl. 6, figs. 15, 15a) also show marked variation, as do
many other subspecies. Just as with color pattern, the shape and size
of the shell cannot be used as a criterion in determining a species,
unless a series of forms are available so that something is known
about the range of variation.
D. Size VARIATION IN THE SAME COLONY OVER A PERIOD OF YEARS
There is little information on this question. A. a. rubidipicta (area
11, ridge complex 3) collected from 1921 to 1929 shows little varia-
tion over a period of years (see table 2, p. 15). A. a. turgida (area 35,
ridge complex 8) collected from locality 202b from 1920 to 1934
shows no variation in the mean length of the shell over a period of
14 years. However, a lot collected in 1913 (table 2) is smaller than
the lots collected in 1920 to 1934. This difference in size of the 1913
lot might be interpreted to be a size change or evolutionary change
occurring in a period of 7 years, but I believe it is more likely the
result of an error due to the small number of shells or to the collecting
of the shells in a locality slightly different from, or lower than, that
where later collections were made.
As in the case of A. mustelina Mighels (Welch, 1938, p. 142), I do
not believe any evolutionary change has brought about a difference in
the size of shells of A. apexfulva collected 20 years ago and those
collected in the same locality today. This differs from Crampton’s
(1916, 1932) findings in his study of Partula of the Society Islands.
E. DEXTRALITY AND SINISTRALITY
No order is found in the occurrence of dextral and sinistral forms
either horizontally or vertically. Ridge complexes 15-20 are domi-
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 205
nantly dextral, while ridge complexes 1-14 contain both dextral and
sinistral shells with the exception of ridge complexes 2 and 4, which
have only sinistral forms.
F. THe Errect oF THE HABITAT ON SUBSPECIATION
Gulick (1905) does not believe that natural selection due to external
conditions explains the diversity of species in Achatinella. He points
out that climate, soil, and vegetation are essentially the same from
valley to valley. He discounts the effect of differences in the amount
of rainfall in a locality on speciation, because forms from the south-
east slope of the Koolau Range show more divergency among them-
selves than those on opposite sides of the Koolau Range, even though
the rainfall is greater on the northwestern or windward slope than
on the leeward slope. Gulick collected most of his material at a low
elevation below 1,500 feet. He had no material that showed any
possible variation between a lowland dry locality and a highland more
humid locality in the same valley or on the same ridge. The diversity
of forms on opposite sides of the Koolau Range is certainly as great
as, if not greater than, that found between forms on the same side
of the mountains.
Crampton (1932, p. 208), in his study of Partula from the Island
of Moorea, states:
With reference to the problem of the possible effects of the environment upon
structural or other qualities the only conclusion warranted by the facts is that
congenital factors are solely responsible for the diversities exhibited by the
several varieties, by the numerous colonies, and by the individual components
of the colonies.
Dobzhansky (1937, p. 136) believes that while it is difficult to prove
that a given trait is not, or has not been, influenced by adaptation to
the environment, nevertheless the facts given by Crampton and
Gulick are explainable on the assumption that racial differences are
merely due to random mutations and to random changes of gene
frequencies in isolated populations. In discussing Wright’s papers on
evolution Dobzhansky (1937, p. 134) explains how species may be
differentiated into subspecies. Such an explanation applied to A. apex-
fulva would satisfactorily account for valley-to-valley variation.
However, it does not explain why there is a definite vertical change
in color pattern along a definite line such as the dotted line in figure 7
which divides highland dominantly white color patterns from lowland
dark color patterns. If variation is purely a matter of the random
combination of genes and mutations within a locality, both highland
206 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
and lowland localities have an equal chance of having dark or light
color patterns and should occur in both areas at random. Possibly,
then, other factors such as difference in moisture conditions, tempera-
ture, differences between a highland and lowland flora, and various
other ecological factors are playing a role. The highland set of
external factors may be favorable to white color patterns, while the
lowland ones favor the survival of dark patterns.
These data indicate that subspeciation within the species A. apex-
fulva is due both to the effects of random variation in partly isolated
populations and to selective factors in the environment.
SUMMARY
1. The various forms of A. apexfulva are organized into subspecies.
New subspecies to the number of 43 are recognized from a total of
78 forms.
HORIZONTAL VARIATION
2. The embryonic whorls may be either bicolored or unicolored.
Shells with bicolored embryonic whorls occur in regions I, III, and
IIIb; those with unicolored ones occur in regions II and IV. Regions
Ia, Ila, and IIIa have embryonic whorls which are intermediate be-
tween the bicolored and the unicolored condition.
3. The color pattern of the postembryonic whorls shows marked
differentiation between regions I, II, III, and IV, and less variation
within each region. More horizontal differentiation occurs in zone II
than in zone III. No horizontal migration is noted of one form
invading the territory of another, because A. apexfulva, like A. muste-
lina (Welch, 1938), tends to break up into subspecies at different
elevations and at short distances away from a given colony.
4. Highland forms occurring in zone III show as much horizontal
size variation as lowland forms in zone II. Half the ridge complexes
in zone II have 18+ shells, 22 percent have 19+ shells, and 27 percent
have a mixture of 18+, 17+, and 16+, shells. In zone III 22
percent of the ridge complexes have 18+ or 17+ shells, while 78
percent have 19+, 20+, or 21+ shells.
VERTICAL VARIATION
5. Vertical variation occurs in the color of the embryonic whorls
between regions IIIa and III, and between regions IV and IIIb.
Regions III and IIIb have shells with bicolored embryonic whorls,
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 207
region IV has shells with unicolored embryonic whorls, and region
IIIa has embryonic whorls intermediate between the bicolored and
unicolored condition. In ridge complex 18, area 97A ?? (fig. 8, p. 195)
has shells with white embryonic whorls; above this area (area 97)
shells with yellowish-brown embryonic whorls are found; and in
areas 98 and 99 at a still higher elevation shells with lighter buff em-
bryonic whorls occur.
6. The color pattern of the postembryonic whorls varies at different
elevations, and this variation is usually more marked than the hori-
zontal variation. Shells from areas having a high rainfall tend to be
white and to have banded patterns ; those occurring in dryer situations
(lower zone III, zone II, and I) tend to have darker patterns which
are banded and streaked. The region of maximum vertical variation
is region IV.
7. Out of 20 ridge complexes, 13 show size increase with increase
of altitude between lower zone II and higher zone IIJ. The remaining
ridge complexes are exceptions to the general trend. Although, the
correlation is not so clear-cut as for A. mustelina, a definite tendency
toward size increase with increase of moisture and lower temperature
exists in A. apexfulva, just the reverse of the correlation for A.
mustelina,
FORM AND COLOR
8. The shape of the shell varies at random both vertically and
horizontally.
g. No order in the distribution of dextral and sinistral shells is
ascertained.
10. Color variation within a locality is often very extensive. This
makes it necessary to study large series in order to ascertain whether
a form is a distinct race or merely a color pattern of a known sub-
species. The same color pattern may occur again in a number of
widely separated localities. The embryonic whorls may be unicolored
or bicolored in the same locality.
I1. Form varies a great deal within a locality. A subspecies may
have narrow or obese shells. The spire may be straight, concave,
or convex in outline. Therefore, form is no criterion for the naming
of a species or subspecies until*the range of variations of a series
ot specimens is studied.
12. Over a period of years no size variation was noted in shells
from a definite locality. However, little data was available on the
subject.
14
208 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
SUBSPECIATION
13. Subspeciation in A. apexfulva is believed to be caused by both
the effects of random variation in partly isolated populations and
selective factors of environment.
REFERENCES
Crampton, H. E.
1916. Studies on the variation, distribution, and evolution of the genus
Partula. The species inhabiting Tahiti. Carnegie Inst. Washington,
Publ. No. 228, pp. 1-331.
1932. Studies on the variation, distribution, and evolution of the genus
Partula. The species inhabiting Moorea. Carnegie Inst. Wash-
ington, Publ. No. 410, pp. 1-335.
DoszHANSKY, THEODOSIUS.
1937. Genetics and the origin of species. New York.
GULICK,, Jeu:
1905. Evolution, racial and habitudinal. Carnegie Inst. Washington, Publ.
No. 25, pp. 1-269.
Hosaka, E. Y.
1937. Ecological and floristic studies in Kipapa Gulch, Oahu. Occ. Pap.,
Bernice P. Bishop Mus., vol. 13, No. 17, pp. 175-232.
Murray, D. M.
1904. Museums, their history and their use. 3 vols., Glasgow.
Pivspry, Henry A.
1939. Land Mollusca of North America (north of Mexico). Acad. Nat.
Sci. Philadelphia, Monogr. No. 3, vol. 1, pt. 1.
Pitsspry, Henry A., and Cooke, C. MonrTaGue.
1912-1914. Man. Conch., ser. 2, vol. 22, Achatinellidae, Philadelphia.
RENSCH, BERNHARD.
1938. Some problems of geographical variation and species-formation.
Proc. Linn. Soc. London, 150th Sess. (1937-8), pt. 4, pp. 275-285.
WELcH, p’ALTE, A.
1938. The distribution and variation of Achatinella mustelina Mighels in
the Waianae Mountains, Oahu. Bernice P. Bishop Mus. Bull. 152,
pp. I-164.
1938a. Some problems of distribution and variation in the Hawaiian tree
snail Achatinella. The Collecting Net, vol. 13, No. 6, pp. 131-133;
also Biol. Bull. vol. 75, No. 2, pp. 342-343.
EXPLANATION OF PLATES
PLATE I
Page
1. A. a. muricolor Welch. Holotype, Niu-Wailupe Ridge, area 1, 41-4,*
Meinecke, BBM 10401). vincsiccs.ccasited valk aide ee eee 30
2. A. a. forbesiana Pfeiffer. Waialae, area 6??, Gulick, BBM 70367...... 28
3. A. a. waialaeensis Welch. Holotype, Waialae Iki-Waialae Nui Ridge,
area 2, 51Aa, Meinecke, BEM 10402: .......5.. 4.2405 5cnee ee 29
NO. I ACHATINELLA APEXFULVA DIXON—WELCH
4. A. a. fuscostriata Welch. Holotype, Palolo area 7??, Gulick, BBM
err oe rye io sins fie yada a eth gus rede elure wcapaks caret ied ters oe
5. A. a. cestus Newcomb. Palolo, area 5?, Gulick, BBM 7o42r...........
6. A. a. simulator Pilsbry and Cooke. Palolo, Gulick, BBM 70378.......
7. A. a simulator var. 1. Waiomao Stream, Palolo, area 4, 62, Meinecke,
ENE NIMSTEAO SID ecrrrer seis caste scnehotenets cuccecteracshials OcuatasGiet stole Ole rale Gite Ranerole a elas
8. A. a. buena Welch. Holotype, Nuuanu, west side, J. S. Emerson,
ENVIRO OAM ere sceisya io ciciei aa one Sits wlave aye aiotaven eine e shave Shere oa teete el ieteheers
g. A. a. vittata var. I. Pauoa-Nuuanu Ridge, area 8, 91A,* Meinecke,
EE SIMD NE ath eek oe he ol eee kee Mee cas eeurtes
10. A. a. vittata var. 2. Glen Ada, Nuuanu, area 9, 100-2,* R. A. Cooke,
EMESIS (IL © Ltn tee Car Niet aica lec sna as7s pe Papateic Ruste eat e eho Ah Beye o oes
11. A. a. cinerea Sykes, Nuuanu, area 10?, Wilder, BBM 50504............
12. A. a. globosa Pfeiffer. From the Blok collection in London, BBM
CO GAEKS MMREPE TTL T SE SSIS es sores os capers alle Ave ra SGED ahaha desi olen BRAS Gatvarsre snares ieye
ite womeitrata. keeve, Nuuanu, Gulick, BBM 7o37i.c.....0-s--e ee oon:
14. A. a. albofasciata Smith. Kapalama Stream, area 13?, Thurston, BBM
HS:
106.
17.
EZTEOS) es
eee ee ewe ee ee we we wee eee etre een eee eeeesreseeeereseeeeeseee
MUON leimecke aus EM TOADS: scree sicceie ole alias siietas. vjemersleclemeias wees
A. a. hanleyana
BBM 165646
BBM 16530
Pieiffer. From the Arthur Blok collection in London,
eee eee eee em me eee ew emer eee eee eee eee eeeeeeeeneeeeeseesne
A. a. hanleyana var. 1. Bought from Sowerby and Fulton, of London,
CC
18. A. a. oliveri Welch. Holotype, Kamanaiki Stream, area 15?, O. H.
RATIELS Ota Es Ey NAG oT OA OO sy kertreit ayer, fopelascs ale eters ci stalin aie! ov as otetoienecavotopatefaiaetess
10.
20.
21
BBM 114979
BBM: 114976
A. a. simulans Reeve. Kamanaiki-Kalihi Ridge, area 16, 121-2, Lemke,
weld n).wile) =) W/o rae! 'e|0/ 6) ee) eels ee ele ;.6\ 9.0 a (=) eX 6 ¢'s v\ ee vs) ee) »ile.elerm) ©. e)6. =
A. a. simulans var. 1. Kamanaiki-Kalihi Ridge, area 17, 121-1, Lemke,
eee ee ww eee wee ewe ee eee Oe wee eee eee eee EE
. A.a. albofasciata var. 1. Kalihi, area 14??, J. S. Emerson, BBM 102206.
22. A. a. rubidipicta var..2. Kalihi-Kahauiki Ridge, area 18?, Wilder,
oa
24.
BBM 505190
Bi wl@ a) 0) 66, © 6).6 8) 4\0)\ 0 © 0:6 0.00) 0) 6 0 0 8 u 6 6.0 6 00 06 0 0 6 0 6 ee eve ee ee see <
A. a. simulans var. 2. Manaiki-Moanalua Ridge, area 19, 151B,*
Metts Kemmis Nik sT 200 7.5) te srarctc cise ats cieee eons tose sineee cuctamel ove cloieroiere octets
BBM, 121037
A. a. ovum var. 3. Moanalua-Halawa Ridge, area 20, 163,* Meinecke,
Ce
A. a. ovum var. 1. Central Halawa, area 21, 162B,* Meinecke, BBM
oie) s) 6.90] val © 6 eve a oe) a) «(00s 6 ©) 6/0 e 66 eh0\16)6)0)e 0 010.0 s)0)0: Diwie ve is) ees ela
A. a. ovum Pfeiffer. North-Central Halawa Ridge, area 22, 162C,*
IMCineclce wes Eine TST OLO crt ..yacssse (ie Sais staaecoie ee area reels etetaeies
A. a. ovum var. 2. North-South Halawa Ridge, area‘ 23, 162D,*
Neineckenp sis Ween TOA nie clow'clrareltsetistme me syei ic miee iek wieseloineioetemle
25 Elalaway area 23s, Walder, DBM: 50523) ree se cele
A. a. pilsbryi Welch. Holotype, Aiea-Kalauao Ridge, area 26?, A. F.
addReB PIM OqO7 es er siesce st the te dae cue sihe eile de eee seine tala ce
25.
T20003> <0. 6.
26.
27.
28. A. a. ovum var.
20.
30.
BBM 114988
31.
BBM 121135
A. a. bruneola var. 2. Aiea-Kalauao Ridge, area 25, 170C-6,* Lemke,
ee ey
A. a. ovum var. 2. Halawa-Kalauao Ridge, area 24, 174,* Meinecke,
eee eee ee eee eee anee ese oeeeseeeeeeee ee eeeeeeeeeeeeeeene
209
Page
44
45
46
36
42
47
A. a. rubidipicta Welch. Holotype, Nuuanu-Kapalama Ridge, area 11,
49
37
38
48
39
40
48
51
4I
60
58
57
59
59
210 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Page
32. A. a. bruneola Welch. Holotype, Kalauao-Hanaiki Ridge, area 28,
182, Meinecke, BBM 10408. 0 i ccc basics as cao cnet eee ooe 61
33. A. a. bruneola var. 1. Kalauao-Hanaiki Ridge, area 29, 183°, Meinecke,
BBM. R23 50> \.sjersietsic'h + w clnete lane lgaesiee chetegas ete oul ae a eee 62
34. A. a. aureola Welch. Holotype, Kalauao-Hanaiki Ridge, area 30, 184,
Meinecke, BIBM: 10400: x6 s:s-st1e 0s ores sicvelescyerevovel breiere nine eiate oe SUT tere 64
35. A. a. aureola Welch. Kalauao-Hanaiki Ridge, area 30, 187, Meinecke,
BBM "r225&o «: < isias'3.ss 5-0's bales dia eleaia-pis ayavb ate. avales. Bia acare ie etter eee 64
36. A. a. awreola var. 1. Kalauao-Waimalu Ridge, area 31, 191A,*
Meinecke; “BBM T2210) s:. sis/sicinc'sierescicioisleiers «ate e'clore elatetetorctoraietertictet ate 66
37. A. a. laurani Welch. Ridge North of Aiea (Hanaiki-Waimalu Ridge),
area @7A?, Cooke; BBM 725574. 0 tshiscans ones bee cucewe en cneeiee ne 55
38. A. a. laurani Welch. Holotype, Hanaiki-Waimalu Ridge, area 27, 191,*
Meinecke; “BBM 21041022 .uGiaccektodees ooa seme seeentiee oes een 55
39. A. a. roseata Welch. Holotype, Waimano Stream, area 37?, Wilder,
BBM LOTR: )iwisis dd, cei ewes sich aea State. atetalavora totetatetevateravereioictae te arevanetalere 54
40. A. a. parvicolor Welch. Holotype, Waimalu?, Gulick, BBM r1o0412.... 56
41. A. a. turgida Newcomb. Waiau-South Waimano Ridge, area 35, :202B,*
Meinecke; (BBM: L2E3TOi 6055s .c0ls eielertie ote oterete teehee Deter oe Ree ae 68
42. A. a. meadowsi Welch. Holotype, South Waimano-South Central
Waimano Ridge, area 34, 212b, Meinecke, BBM 10413.............. 70
43. A. a. meadowsi var. 2. Waimalu-South Central Waimano Ridge, area
33,,200Ca;; Meineckes BBIMeIZrdaee na ciercteetore ote crereia terete rterteiore 72,
44. A. a. waimaluensis var. 1. South Central Waimano-Central Waimano
Ridge, area 32A, 202D, Meinecke; BBM 121502... 656.20. cs eww va ce 67
45. A. a. waimaluensis Welch. Holotype, Waimalu-South Central Waimano
Ridge, area 32, 204C, Meinecke, BBM 10484... 60..005500¢000e eres 66
46. A. a. cookei var. 1. North Central Waimano-North Waimano Ridge,
atea 30; 2118) MeinecketBBMe121602)).\.4c501 <6) eee eee cere: Ta
47. A.a. perplexa Pilsbry and Cooke. Waimano, Cooke (Waimano-Manana
Ridge:Vatea\ 40)" BBM.-7274 ht jon ctow pomals a cciiee octet eee cerereters 72
48. A. a. nigripicta var. 1. Waimano-Manana Ridge, area 44, 223, Meinecke,
BBM, 229738 (0354. v ccc sie sale sedis neat Sh a SA Cate ae Bk Oe eet eee 79
49. A. a. cookei Baldwin. Waimano-Manana Ridge, area 45, 225-1,
Meinecke: "BBM Gr2t67, crc ciclo sone aan oe on se ieee Dee 75
50. A. a. simulacrum Pilsbry and Cooke. Waimano-Manana Ridge, area 46,
228 * Meimecke, BBM 121773 jcc sisi cocie vnioe cain eee a eeetine 81
51. A. a. rubidilinea Welch. Holotype, Manana-Waiawa Ridge, area 47,
230a,° Meiriecke:, BBM GTOADE ss cvs sleseistets aynreis-nieictole stcrerevettion lokeiersbefoiott terete 83
52. A. a. chromatacme var. 1. Manana-Waiawa Ridge, area 41, 231,*
Meinecke, BBM. 121806. .0jcce0 ence ccdieas ach ginfereta elajuie ein in ajatalatateia eter tangs
53. A. a. chromatacme Pilsbry and Cooke. Manana-Waiawa Ridge, area 42,
234," Meinecke, BBM: I27B46 i, occcrasecateecraetels Oss aca gia enn ee 84
54. A. a. chromatacme var. 2. Manana-Waiawa Ridge area 43, 237,*
Meinecke; BBM) t21814socc once cote tase se uniec heen oa eee 88
55. A. a. nigripicta Welch. Holotype, Manana-Waiawa Ridge, area 50,
237b; Meinecke, (BBM, 10416: 2c 0 sae cie oe ce th oe eee ae 78
56. A. a. cookei var. 2. Manana-Waiawa Ridge, area 49, 237fa, Meinecke,
BBM. 324840. ic icce vances ena ees wea a iNercareronatate cies lera eiers ate 77
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 2IiI
PLATE 2
Page
1. A. a. nigripicta var. 2. Manana-Waiawa Ridge, area 48, 237h, Meinecke,
EV Te2A OG BaAPae te erareh cite fe forescatore) aici ereeersta (ote ayers oh Sereesuay® ole elo etstuns neers 81
2. A. a. nigripicta var. 2. Manana-Waiawa Ridge, area 48A, 238b,
PP THE GENEL NSS TOA GUO LOM oe Gili b.ciate css ah ais acoiatoreiate wd el nanlecle Cd ohs 81
3. A. a. albipraetexta Welch. Holotype, North-South Waiawa Ridge, area
Haeecnn,)Meinecke, BEM) LOATH: Jie ca cd end ones e 8d co velba e's bee's 89
4. A. a. ewaensis var. 1. North-South Waiawa Ridge, area 51, 237B,
Wettecicers sis Vigl24 SOQ yacccian tec nits oracles oe erheiatele os Ho sbi ee eek oan 116
5. A. a. lemkei Welch. Holotype, North Waiawa Stream, area 67?,
moka ermiice mts NEw TOAT Os «cc aa ieieie ents se 8 wleeeled $410 v'o hebwie ee ok 95
6. A. a. griseibasis Welch. Holotype, East Waiawa, area 65??, O. H.
MSLSONMEEs Is VasTOATO Sats operrocloaers << /sielel sissies ajo cualevcre era: siels Gan odiv te 90
7. A. a. flavitincia Welch. Holotype, Waiawa-Panihakea Ridge, area 60,
2BO GreaMieInec keris BI MeETOAZO S.1aciots stave No oriare's ociteiare caters ae avemieie 04
8. A. a. lineipicta Welch. Holotype, Waiawa-Panihakea Ridge, area 70,
Daa aw MCIHECKE 4 Eyibs Mig TOAD Tiayer tatsie ers Ss ayeheverciocs icles ore eiere wiclore ersrers 08
9. A. a. fumositincta Welch. Holotype, Panihakea-Kipapa Ridge, area 71,
pe atenteckce: WIEN TOAZ2) . bviscis ecu sluts ces tpembateecveces ns 100
10. A. a. lineipicta var. 1. Panihakea-Kipapa Ridge, area 72?, 241-1-242-3?,
FRAG SHES Es IV ee TESTING Tr ct aeacibel Pah cate icteveressferereicia se ccniaie Gore ein stove ralgreleeitte. elerere 99
II. A. a. ewaensis var. 3. Waiawa-Kipapa Ridge, area 52, 243-2, Meinecke,
EESIVIMITET. OAS aera vetcss) cls toteroeraie te ave were icle eae os etea Ore d claictorate cal meahele Aa avels 118
12. A. a. ewaensis var. 2. Waiawa-Kipapa Ridge, area 53, 244,* Meinecke,
EEUVIGET AT OO Opp raeccr ain toce Sclele Whe aie Aik iehatiaier are ice SOaorenad cso a roles 117
13. A. a. ewaensis var. 4. Waiawa-Kipapa Ridge, area 54, 246,* Meinecke,
EN ES IVE WeSC) Almere repetae epaisusie tatoo eee te ravecers ays forse pears iav ale Sayetelsioreoreeelen ears 119
14. A. a. ewaensis var. 5. North-Central Kipapa Ridge, area 55, 240L-1-1a,*
[en ETDS 45 OM SUB) (ie (0 OR a ee a 119
15. A. a. ewaensis Welch. Holotype, North-Central Kipapa Ridge, area 56,
BAe te MGIICE KE: ENE TOADS... bee av cccte os Haat elnece a Cele eee es 115
16. A. a. virgatifulua Welch, Holotype, Waikakalaua Stream, area 73?,
NVA Gime sis MieaT O4DA lta ty sevolelave (aleve ters cleleieiels.«, « apace nile aiicleisies eiuieatels 103
17. A. a. virgatifulva var 2. Waikakalaua Stream, Wilder, BBM 50571.... 106
18. A. a. ewaensis var. 6. Kipapa-Waikakalaua Ridge, area 57, 252-1 -2 ?,
MEINE CKe mE IMB i 2TOS5 chee. sare se oh itae ee,6 sate datise-clalale me auc outst ere 120
19. A. a. ewaensis var. 7. Kipapa-Waikakalaua Ridge, area 58, 256,
Meine ckeres BINS e040 4. chylseie cceisceisiele eaves isicies « Sieh ntere tieleleke, avers 122
20. A. a. virgatifulva var. 1. Waikakalaua-South Kaukonahua Ridge, area
FE ee2bOmeNVeIneckes BM) T2TO3O. acinjeccins cisco cals 01s lc8e slelemioel cere 104
21. A. a. tuberans var. 2. Waikakalaua-South Kaukonahua Ridge, area 60,
OM ENIEINeC KEM ID D MOOT2LOATS cc, ctoeve Stevsisvatevn ters a6 wintere mn vere eerie sis feiss 109
22. A. a. ewaensis var. 8. Waikakalaua-South Kaukonahua Ridge, area 50,
Peto o te MeTREE KG Has Bs Wis F2IQ27.0i3aiticls ois oi vine s Sealdie or e'eisideart eaets ee 123
23. A. a. tuberans var. 4. South Kaukonahua South-Central Branch Ridge,
afea Zor, 200A.?, Meinecke, BBM 121068 6.2506 vclss sacs slew vieenese vc 112
24. A.a. steeli var. 1. Kaukonahua, Wilder, BBM 10446....'...........--- 145
25. A. a. steeli Welch. Holotype, North-South Kaukonahua Ridge, area 80,
SIS ESE NI IOADE oy ul sie ee <3 ec Uerale oe Six meee eine gia ees atic ey 143
26. A. a. punicea Welch. Holotype, Kaukonahua, Wilder, BBM 10426.... 141
bo
_
30.
ar
32.
33.
34.
35.
30.
10.
II.
12;
¥3;
14.
15.
16.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Page
A. a. punicea var. 1. North-South Kaukonahua Ridge, area 82?, Lemke,
BBM H215080 0's vw iclc'dias is enw einve sone byeik cp ote atten p ett lace Sin a ea ee 142
A, a. tuberans var. 3. North-South Kaukonahua Ridge, area 77, 274,
Lemke, BBM 2115045 )0\0\s «icc 'ss o'vieleosiw sles halo wiv aus Sealer salah ee III
A. a. irwini Pilsbry and Cooke. Kaukonahua, Wilder (probably North-
South Kaukonahua Ridge, area 61), BBM 50576.................. 124
A. a. irwini var. 1. North-South Kaukonahua Ridge area 62?, 277?,
Meinecke,. BEM 222107.3 08:25 25:< siete atersichele etelete aietelaeiarsteretere tae ieteranetetee 125
A. a. aloha Pilsbry and Cooke. Kaukonahua, Wilder, BBM 5o56r1..... 160
A. a. aloha var. 1. Kaukonahua, Wilder, BBM 50560................. 161
A. a. versicolor Gulick. Ahonui, Gulick, BBM 70380................ 130
A. a. coniformis Gulick. Wahiawa, Gulick, BBM 70368................ 126
A. a. apexalba Welch. Holotype, Wahiawa, Gulick, BBM 10427...... 128
A. a. wahiawa Welch. Holotype, Wahiawa, Gulick, BBM 10428....... 132
PLATE 3
A. a. suturafusca var. 1. South Poamoho Stream, area 85, 280-7, Isle,
BBM) (13484005 56 8isle bcd wine wien er eect mie Siete aia le Oe ee iene tereer ster 149
A. @, lilacea Gulick, Ahonut, (Gullick) BBM 107037214.) .-)s selena 137
A. a. lilacea var. 1. North Kaukonahua-Poamoho Ridge, area 83, 280-1-
280% Russ BIB Mi wr2003Scroi ts ce crtecct carcuciceete en eete okie dare reset erate 138
A. a. lilacea var. 2. North Kaukonahua-Poamoho Ridge, area 84, 282-3,
Meinecke; BIBIM sy 124 034s yan cre cotrcetomcie cla cicne ch uavavse aelaeie nie reece 140
A. a. poamohoensis Welch. Paratype, North Kaukonahua-Poamoho
Ridge, area 101, 284-2, Meinecke, BBM 124151, the usual pattern of
ECA TOD eis oe al Lae wcatthow cal aie Sie wi is ane arco ree eta ta tana caaney cee eee 166
A. a. poamohoensis Welch. Holotype, North Kaukonahua-Poamoho
Ridge, area 101, 284-2, Meinecke, BBM. 10420; 4...) 52 (o.sscceesm ee 166
A. a. roseipicta Welch. Holotype, North Kaukonahua-Poamoho ridge,
area 102200; Meinecke! BM 10480 ene rece eer ete aeeree 162
A. a. roseipicta var. 1. North Kaukonahua-Poamoho Ridge, area 103,
280-Ta,, Meinecke;, (BBM: 5247360): cciey cinic civtss'c sleyeloinievelereyeercverspelareieteietele 163
A. a. suturafusca Welch. Holotype, Central Poamoho Stream, area 86,
280K -1:* Meinecke; BBM: 1G43% ccisccies or 'sietdc.vis sete uraete cadeetceren et 148
A. a. glaucopicta Welch. Holotype, Central Poamoho Stream, area 87,
280K=2, Meinecke, BEM T0492 5.)2 5 cacsns.s 0.0.0 0/0; chalet io cceeteretin nsec 140
A. a. brunosa Welch. Holotype, Central Poamoho-Central Poamoho
North Branch Ridge, area 88, 280L-2, Meinecke, BBM 10433....... 146
A. a. suturalba Welch. Holotype, North Poamoho Stream, area 90,
200A:A=8, ‘Lemke: BE MnT0agart) ieiiescvais s.cate vam vee teenie oe ISI
A. a. suturafusca var. 2. North Poamoho Stream, area 89, 2090,*
Meinecke, ) BBM (122089). 5b Bota averse se 6 t's x obe evcetoete Rarer arareiee tte 149
A, a. apicata Newcomb. Helemano, area 92, Gulick, BBM 106700.... 152
A, a. cervixnivea Pilsbry and Cooke. Poamoho-Helemano Ridge, area
03; 300-0-30I-2,% Meinecke, BBIM 122104. cise ciclcte retiree ee titers 167
A. a, beata var. 1. North-Central Poamoho Ridge, area 94, 291-3-5,*
Meinecke, BBM’ \121002). $200 cess GRGAM SER ES See eee eee 171
7. A. a. beata Pilsbry and Cooke. Poamoho-Helemano Ridge, area 104,
303; Meinecke,.. BBM 11283726 iv diiscted acu nes eee eee ace 169
NO. I ACHATINELLA APEXFULVA DIXON—WELCH
18
10.
. A. a. roseipicta var. 2. Poamoho-Helemano Ridge, area 105, 307-1a-1b,
.Meinecke, BBM 1220090.......... OPEC ARTs RIT Panett he Wee
A. a. suturalba var. 1. Helemano, area 91??, J. S. Emerson, BBM
PRR MME era oe tcla te Naas tis Clk wi ata: Sra Bek a Aa OS BEA Ow oa oe as
. A. a. apicata var. 1. North-South Helemano Ridge, area 95, 312-4,
Micinecenmaia MinnT26g22eir 2a. 58 ok Seino pea ie eee el eee ee
. A. a. paalaensis Welch. Holotype, North-South Helemano Ridge,
aLeamloomera-seNleinecke, BIBI T0435e..-2--06:me nents comes eee oe
. A. a. beata var. 2. North-South Helemano Ridge, area 108, 318,
NCIHEC Camm Es Es Veal 2843 Oleret ao oj dis. vis heya secre sats ale le nia eto ek
23. A. a. apicata var. 2. Kawaihalona Gulch, area 96?, Thurston, BBM
TESTE” cath chi ap 0 eg a Spa rene Pr es
24. A. a. paalaensis var. 2. Helemano-Opaeula Ridge, area 1009, 337-3,
NEMEC kemmE IV leps28 77 On vents ars 8 As eae a Oe a ee ees nee
25. A.a. beata var. 3. Helemano-Opaeula Ridge, area 110, 337-4, Meinecke,
PaCS Ete ORC Pee he o Siars Sore nih ciciaee oc Cae Oe ak Bites ae
26. A. a, duplocincta Pilsbry and Cooke. Kawailoa, Emerson, BBM 102312.
27. A. a. apexfulva Dixon. Opaeula Gulch, J. S. Emerson, BBM 102270..
28. A.a. vespertina Baldwin. Paratype, Kawailoa, E. Lyman, BBM 167354.
29. A. a. kawatiki Welch. Holotype, Kawaiiki-Kawainui Ridge, area 90,
IMieiitecice wo Mil O43O Sere eitere: toe cee cele isisle rine ae blots Sinan
30. A. a. bakeri Welch. Holotype, Waimea, J. S. Emerson, BBM 10437...
31. A. a. leucozona Gulick. Waialee, Gulick, ANSP 92656...............
32. A.a. paumaluensis Welch. Holotype, Paumalu-Kaunala Ridge, area 111,
Asie einteckens BIS MM TOASS so Fae aly Se eeucels osha slcsclansibe iol eieatdigeleions
33. A. a. oioensis Welch. Holotype, Oio-Oio East Branch Ridge, area 114,
GIy ame LemIeeice Eds VE TOAZO® S 5.2.0 /bier sere ois ais ers tes Wea eres elo nee wcrele's
34. A. a. kahukuensis Pilsbry and Cooke. Malaekahana-Laie Ridge, area
NaH mola SLO ees SMe P3073 cis: cress icin so cieiciele sieiaia sista < sieteteiei ae al ete
35. A. a. thiihiensis Welch. Holotype, Thiihi-Kahawainui Ridge, area 116,
EMOMVICINECKE WE MN TOAA Oni ac deveined store etna cee te ies cl aloretetee intestate
36. A. a. wailelensis Welch. Holotype, Wailele Gulch, area 117, 529-1, Russ,
FEMI IRE OMA Tem nr eee tinted itso ohatstaleiie he ec etalatcna als che aie eae oxe snob umtebcheteeeis ic ie
PLATE 4
1-1b. A. a. muricolor Welch. Paratypes, Niu-Wailupe Ridge, area 1,
AeA eNlemecker BIB: T20177- TZOUSA se ciecle che cles ces cine rss
2-2a. A. a. waialaeensis Welch. Waialae Nui, area 2, 50A,* Meinecke,
NIM O2 Dieta tic oth ote rcie grote ello ata oie tie. dre 'otelel suetaue eratenste Pavetan ae
3. A. a. watalaeensis Welch. Waialae Iki-Waialae Nui Ridge, area 2,
OlMEIenla mesons Ms, TOZG20 2. -tas stiicvare oo eA cre a tie cl oreaiere reuse
4. A. a. waialaeensis Welch. Waialae Iki-Waialae Nui Ridge, area 3,
SOA Aw ON Eessmersonn Bib MialG727.5% sclera shai sais tenets ciel rere ote ale
Ea a fororsiana Pieimer. Holotype, BM. 26.5. e iccecevers seecce sees
6-6a. A. a. forbesiana Pfeiffer. Waialae, area 6??, Gulick, BBM 70367...
7-7a. A.a. fuscostriata Welch. Palolo area 7??, Gulick, BBM 70421.....
Sree ee tnnorauses Snnth. tolotype, BM... .oiattet.saiea nen sconces as
ieeAen Ce sit seNew. Comp!) cectoty,pe,) Mis s\vsierie s+ «sles clalalel sialerele estes ole ios
10. A. a. cestus Newcomb. Specimen A, type lot, BM........... cece eeee
213
Page
173
179
176
174
159
180
181
183
30
29
20
29
28
28
32
33
214 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Page
11. A. a. cestus Newcomb. Wailupe, Gulick, BBM 70383................. 31
12. A. a. simulator Pilsbry and Cooke. Holotype, ANSP 92502.......... 34
13-13a. A. a. simulator Pilsbry and Cooke. Palolo, Gulick, BBM 106960,
$37 Gi eins arsisieinieye.acejaeslonetee viele iolerein al eFecers Gabe meen ecient ee eee 34
14-14a. A. a. buena Welch. Paratypes, Nuuanu, west side, J. S. Emerson,
BBM. 202285) s.isie/nie.s'sia/ais 2's ba.e han wie bien eaioee eee ete nae Mieey 8
15-15). A. a. vittata var. 1. Pauoa-Nuuanu Ridge, area 8, 91A,* Meinecke,
BBM 8207360 5:82 ieis.0ccteidiereltesoe acre cole sleteite areca eee 44
16-16a. A. a. vittata var. 2. Glen Ada, Nuuanu, area 9, 100-2,* R. A. Cooke,
BEM. S8EGE\. kissd vierais bie Pek es accndscieie archer ae oh eI nanan 45
I7.:A. 4d. uitiata Reeve: Holotypeiy BM wissen ccenee eee ee eee 42
18. A. a. vittata Reeve. Nuuanu, Gulick, BBM 70371...... ait ahaa a ORES 42
19-19a. A. a. vittata Reeve, Hillerbrand’s Glen, Nuuanu, area 12, 100E-1,*
GC. M, Cooke, BBM, 19016. scduas oc Hel ee se eee eee 42
20-20b. A. a. vittata Reeve. Nuuanu, area 100F-1,* R. A. Cooke, BBM
BO285, BOLO) BO2GA wis are rere wcelele.xtnre aterm nie etarelerave ats lave epee eee 42
21. A. a. globosa Pfeiffer. Lectotype, BM............... Vet Rae! 36
22. A.a. globosa Pfeiffer. Specimen A of type lot of A. a. globosa Pfeiffer,
BM a eciaudt Tia B'S 5h wry cl See WW ts ese tap ce ale ae Ae devel TO yr ne 36
23. A.a. globosa Pfeiffer. Specimen A, type lot of A. vittata Reeve, BM.. 36
24. A.a. globosa Pfeiffer. Nuuanu, Gulick, BBM 107003.............+++- 36
25 Ana nonievana bteltterm Lectotype Mieeeeenee nee nee 37
26. A. a. hanleyana Pfeiffer. Kalihi, Thurston collection, collected by
Dos, Santos; BBM s131009F bccn eis eee eee Ite eee: 37
27 ANG AGO Tasciataromithy Holotypeab Minna. veseeete eee een tne 47
28-28b. A. a. albofasciata Smith. Kapalama Stream, area 13?, Thurston,
BBM. 131006, IZL104, 131005. ...0.1024s0% sea eee ee eee 47
29-29a. A. a. albofasciata var. 1. Kalihi, area 14??, J. S. Emerson, BBM
TO2ZOG, - TOZZOS, <6. a!b ainisata! ec ate aie aeeehente ane mene clamor eee eee 48
30-30a. A. a. oliveri Welch. Paratypes, Kamanaiki Stream, area 15?, O. H.
Emerson, “BBM<1030625. 2... cena tee oe tee eee ene 48
31. A. a. oliveri Welch. Kamanaiki Stream, area 15?, J. S. Emerson, BBM
TO2QO4 oo ins 1055 ates aye ekeva ssa doe aeleis Wile atell oe eres Sear ete me eee 48
32. A. a. hanleyana var. 1. Bought from Sowerby and Fulton, of London,
BBM “TG5GAG,. Aa5 bo oles Siete tyre orcteuw bea alo ite wae ee ete 38
33. Aa. similans Reeve, Copy of original Redre. {1702.60 eee 39
34. “Aas simulans Reeve, a Lectotype sbi ence ene cee meee ae eee 39
35-354. A. a. simulans Reeve. Kamanaiki-Kalihi Ridge, area 16, 121-2,
Lemke, BBM, 124070 os % sian teicacinlea sgn hve seieene ceroemnae area 39
36. A. a. simulans var. 1. Kamanaiki-Kalihi Ridge, area 17, 121-1, Anderson
and ‘Kondo, BBM 166904)42.2 scene cehuin Bee antaoaeen ines eee 40
36a. A. a. simulans var. 1. Kamanaiki-Kalihi Ridge, area 17, 121, Lemke,
BB M2i4076einicis.ct Be dtaeinte teenie ei ota ee ae waletep ing eens 40
37: .A.0. ‘cimerca Sykes,, Holotype BM: soso in ccenscss pee eee eee 46
PUATESS
I-1a. A. a. cinerea Sykes. Nuuanu, area 10?, Wilder, BBM 50504........ 46
2-2d. A. a. rubidipicta Welch. Nuuanu-Kapalama Ridge, area 11, 110,*
Meinecke;; BBM. 120858. co eee retie ata a ieistone oneriorsteree 49
3-3b. A. a. rubidipicta var. 1. Kalihi, Wilder, BBM 50544............... 51
Oey) T ACHATINELLA APEXFULVA DIXON—WELCH 215
Page
4-4e. A. a. rubidipicta var. 3. Moanalua, Wilder, BBM 50522............ 51
5-50. A. a. simulans var. 2. Manaiki-Moanalua Ridge, area 19, 151B,*
Meinecke sss DIMM 20675 nsiieacterineres ta erred card Woe miei dcaics Al
6. A. a. ovum var. 3. Moanalua-Halawa Ridge, area 20, 163,* Meinecke,
IBIBIMESTOTOS Ts yore ka Mere cee eras aah paote bets etaleleralakortverss oso ra ts Rede use Se 60
7. A. a. ovum var. 3. Moanalua-Halawa Ridge, area 20?, O. H. Emerson,
IB BNR ST O3 OO 4 ieiacrc de te: ous vaves cyapaxaraterctahetanec eter ola ne accestedeterare telcoiekt Oe Bie iaca ao 60
8. A. a. ovum var. 1. Central Halawa, area 21, 162B,* Meinecke, BBM
TAO Shanes Sie sac y steve love ot arokatsioin ore’ ceaue laiaeols.cuebt ere eens eels ohana sc eeieleies 58
8a. A. a. ovum var. 1. Halawa, Thurston, BBM 130505................. 58
PANG OVUM se teliter., Eloloty.pe; BiIMi.s aecc.ne. coe nisia ssid eesjomieiers ots oe arene 57
10. A. a. ovum Pfeiffer. Specimen A, type of the lot, BM................. 57
11. A. a. ovum Pfeiffer. North-Central Halawa Ridge, area 22, 162C,*
Meinteckese Bs MisI2r OU Os sce: ays ciorererctr acl eetrereie ort oertinis arenes 57
12. A. a. ovum var. 2. North-South Halawa Ridge, area 23, 162D,*
icinPee ENT) P21O42 a) tos oo alee aes eel a eo yd Le ee 59
13. A. a. ovum var. 2. Halawa, area 23?, Wilder, BBM 50523............. 59
14. A. a. ovum var. 2. Aiea, area 24? J. S. Emerson, BBM 102446........ 59
15-15b. A. a. bruneola var. 2, Aiea-Kalauao Ridge, area 25, 170C-6,* Lemke,
EE META GSO IAS 2 eiTAOG0. i aaipeiee citieraeioieie aka aitresianrcierere 62
feed. ce bruncola var. 2. Aiea, Wilder, BBM 50528... .6026..000c.c00cees 62
17-17b. A. a. bruneola Welch. Paratypes, Kalauao-Hanaiki Ridge, area 28,
ToO2 se Viemneckes SIDING T2ATIOON MAMI 7h. seyeteeriels oe eiteeieete eels 61
18. A. a. bruneola var. 1. Kalauao-Hanaiki Ridge, area 20, 183?, Meinecke,
BB INEPT 2 TS One cranes beorererecsyarc ree eietaccis recatele eis icwe ores ansiceass chaldimnae 62
19. A. a. aureola Welch. Paratypes, Kalauao-Hanaiki Ridge, area 30, 184,
Meineckes BB Miet2 tO 2c martyrs s soe Geet ere cee amine sions 64
20-20a. A. a. aureola Welch. Kalauao-Hanaiki Ridge, area 30, 185,
INGIEEKe HEE My T2100) ST 212035. weate sc oie ot oe tis eee oe eislomiwe eet 64
21-21a. A. a aureola Welch. Kalauao-Hanaiki Ridge, area 30, 187,
NEEINICE One Dy DARE SE UGD eT otha ceils ey otaak oe teahaee elo Moniaree cle 64
22-22b. A. a. aureola var. 1. Kalauao-Waimalu Ridge, area 31, 190A,*
IMeinecKerb Bi Mig TOT2O72 lac erksie os Vis cies rele ke aoe ein ds a ects 66
23. A. a. meadowsi var. 2. Waimalu-South Central Waimano Ridge, area
3332200 CaseMieinecke, IBIBIMe TATE S Oi aria «cine eiereisn oo taekrel or obe oie ane 72
24. A. a. meadowsi var. 2. Waimalu-South Central Waimano Ridge, area
BawezOl ew \eimeckes PBI TaTAs sn. an ee clace oroths ae vee 72
25-25a. A. a. waimaluensis Welch. Waimalu-South Central Waimano
Ridge; area’ 32, 203C, Meinecke, BBM 121460. .......0.0.<.000 66
26. A. a. simulacrum var. 1. North Central Waimano-North Waimano
Ridge, area 38, 216B, Meinecke, BBM 121631...............00. 83
27. A. a. pilsbryi Welch. Paratype, Aiea-Kalauao Ridge, area 26?, Cooke,
IB EBIMBS 724 OA aerate as ferniaciore eee Ie ciate IO si SIRE ie lon ain Pelee 52
28. A. a. lawrani Welch. Ridge North of Aiea (Hanaiki-Waimalu Ridge),
aheak27- wi COOke ME BME Z2E57 trebles waves sie cee evel eualorarciieiatelaiaiele 55
29. A. a. lawrani Welch. Ridge North of Aiea (Hanaiki-Waimalu Ridge),
ALCAee 7 Nor (Gd EMME TOA OO saree is sloth eiaisateeeieie clare ieie moieie 55
30-30a. A. a. lawrani Welch. Hanaiki-Wiamalu Ridge, area 27A?, Thurston,
IMIR TG OGO SW erent At cette stret ieee one oooh oe oreo eran aieve crated 55
31. A. a. laurani Welch. Paratype, Hanaiki-Waimalu Ridge, area 27, 191,*
MeiniEcKey PISA TORTOOL Gi. Lae (eta bs oe eb swear beta cele te winleten 55
216
_
wh
-4e,
WAH
10. A
II-I1a.
ey wale
13. A
14-I4a.
15-15¢.
16. A,
17-17).
18-18a.
19-19d.
20. A.
A,
A.
A.
A,
A.
A,
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL.
PLATE 6
a. laurani Welch. Paratype, Hanaiki-Waimalu Ridge, area 27,
16r,* Meinecke; BBM ‘t2rtgSnaccieee cate Vee ee cee ee
a. turgida Newcomb, Newcomb’s original figure 10...............-
a. turgida Newcomb. Lectotype; BMiii. cede... ove oe nk wens wate
A. a. turgida Newcomb. Waiau-South Waimano Ridge, area 35,
202B, Meinecke, BBM 121307, 121319, 121387, 121303, 121320...
a. turgida Newcomb. Waimano, Wilder, BBM 50508..............
a. turgida Newcomb. Waimalu, J. S. Emerson, BBM 102218........
a. turgida Newcomb. Copy of the original figure 9 of A. swiftii
Newcomb” .A.Qccca sides Se eae eh Bee ek Ghee eee ene
. a. turgida Newcomb. Copy of the original figure oa of A. a. swiftti
INGwGomb! <2 eee OA xk ie rR et ict rae a arte
. a. turgida Newcomb. A specimen collected by Newcomb, from the
Newcomb collection at Cornell University, and matched with
plate:G, mire 7 Pires oe asic Savien wate aie ale Says lale aed allele Ghat ge
a, turgida Newcomb. Waimano, Wilder, BBM 50508, matched with
plate G; figure Sis sso. pokes ca were eactes horace meteor Me ome
A. a. meadowsi Welch. Paratypes, South Waimano-South Central
Waimano Ridge, area 34, 213a-213b, Meinecke, BBM 121546....
a, meadowsi Welch. South Waimano-South Central Waimano Ridge,
area 34,2134, Meineckes ISB M 123588: 5 a ucscns sce cie mes eiseee
a. meadowsi var. 1. South Waimano-South Central Waimano Ridge,
2i8a.Mieineckes 5 BiNastes5C0ma aria ae ence cee eee
A. a. meadowsi var. 1. Waimano, Wilder, BBM 50537............
A. a. parvicolor Welch. Paratypes, Waimalu?, area 36??, Gullick,
BBM :-7O488H 00728) auntie ruse oto et ene eee Doe eee
a. perplexa Pilsbry and Cooke. Lectotype, Waimano, C. M. Cooke,
BBM: -Qo7Ag niick6, estas aa taclhe ae datieeee ake Dae Skea eee
A. a. perplexa Pilsbry and Cooke. Paratypes, Waimano, Cooke,
BBM 92745272728) Foro oven siaunte chaste celts ett ce ee eee
A. a. perplexa Pilsbry and Cooke. Waimano-Manana Ridge, area
40, 2214-* ‘Meinecke. (BBM I21718; T2T7201 s.. Jeo cee ene ee
A. a. perplexa Pilsbry and Cooke. Manana, Wilder, BBM 50551...
a. chromatacme Pilsbry and Cooke. Lectotype, Waiawa, ANSP
108804
© a a\je) 0! (a eo. 6) 0) a) Wm) 0) le a ae) see) 6) 6 # 16a 6) 61 e 6) 610, a) ne) /a, 2 1s) 0) vim 6) eis 8 Sele evens
. A. a. chromatacme Pilsbry and Cooke. Manana-Waiawa Ridge,
area 42, 235,* Meinecke, BBM 121853
. A. a. chromatacme Pilsbry and Cooke. Manana-Waiawa Ridge,
area 42, 234,* Meinecke, BBM 121845
©) Se) es eXn, are "she. a6 10 oe o.0lw swe elsilals
. A, a. chromatacme Pilsbry and Cooke. Manana-Waiawa Ridge,
area 42, 232, Meinecke, BBM 121810
. A. a. chromatacme var. 1. Manana-Waiawa Ridge, area 41, 231,*
Meinecke, BBM 121806, 121808
6) ) \¢.8\'0)'O..6 18)e 16 ee) wells © ovw een Le eee lb) elie. sass
. A. a. roseata Welch. Paratypes, Waimano Stream, area 37?,
Wilder, BBM 51032
eCoeetoesce eee oes e een ese seen eee eet eee ses ees
25b. A. a. roseata Welch. Paratypes, Waimano Stream, area 37?, Thaanum,
BBM 40717
©1009 (69 010) 6 ee a 8b is) © 60 6 soln) 6. 6in, © ale, 0.16 (ee, 8) 0 wie) es S818) 6) 6 ale ise
103
7O
~
71
Fil
N
to
to
NI
NI
bw b
NI
NO. I ACHATINELLA APEXFULVA DIXON—WELCH 217
PLATE 7
Page
I-1a. A. a. nigripicta var. 1. Waimano-Manana Ridge, area 44, 223,
Miemeckes sp BiIMit21 738) 1217485 tan naetcioerd ee hoses Nake 79
2-29. A. a. nigripicta var. 1. Mamnana, area 44?, J. S. Emerson, BBM
OZ MOL OFT Gree erate mer nicks Whe he csi eee ERA ee ee eT he 79
3-3d. A. a.-nigripicta Welch. Paratypes, Manana-Waiawa Ridge, area 50,
237 Dee Meineckes Eb) Mat24q780n 02477 Oasis sen aes aerate ale nce 78
4. A.a.nigripicta Welch. Manana-Waiawa Ridge, area 50, 237d, Meinecke,
IBIBIMMT2A 7 ORS Sits cio tic hatte ores Oe Grae aie ae ey thd 78
5. A.a. nigripicta var. 2. Manana-Waiawa Ridge, area 48, 237h, Meinecke,
Exo RE DABOG Sia cL leo thes URNS Ma ake ree gina cee SEI 8 81
6-6a. A. a. mgripicta var. 2. Manana-Waiawa Ridge, area 48A, 238b,
iIMemeckesE BIM@I 24800 mr 2A820 ees ye cite seach le Leas 81
7. A. a. cookei Baldwin. Copy of Baldwin’s original figure.............. 75
Pia. Cooke: Baldwin. Lectotype, ANSP 65602. ......00cs<lacscecceces 75
9. A. a. cookei Baldwin. Paratype, Ewa, C. H. Cooke, BBM 167352...... 75
10. A. a. cookei Baldwin. Paratype, Ewa, Baldwin, BBM 54669........... 75
11-11b. A. a. cookei var. 2. Manana-Waiawa Ridge, area 49, 237f, Meinecke,
Sa AEA ZOU. eet « Mnrcd tot ncucia's ake SPINS Pee ei ois seated ig Miatanets Pam boiciss whe Ta
12-12d. A. a. cookei Baldwin. Waimano, Wilder, BBM 50594............. 75
13. A.a. simulacrum Pilsbry and Cooke. Lectotype, ANSP 108063........ SI
14-14a. A. a. simulacrum Pilsbry and Cooke. Waimano-Manana Ridge,
aLearA Gy 226. = Meineckem sy 12th ene sieeiaceeeicnian ce: 81
15-15a. A. a. simulacrwm Pilsbry and Cooke. Waimano-Manana Ridge,
atedsA0,220," Meinecke. BBM 921777).5 =. acs cc cien Ss cee leld ewe nce 81
16-16a. A. a. rubidilinea Welch. North Central Waimano-North Waimano
Ridge, area 47, 219B, Meinecke, BBM 121630, 121641........... 83
17-17c. A. a. rubidilinea Welch. Waimano-Manana Ridge, area 47, 220A ?,
IMeineckexss bi MimleT7SOnes me tactic nire tices ose TE 83
18. A. a. rubidilinea Welch. Paratype, Manana-Waiawa Ridge, area 47,
Zee MeCN EE ME 2AS 39) Fra raias atin Hatake saisieneieie Ss Seis a ss 83
19-19a. A. a. ewaensis var. 1. North-South Waiawa Ridge, area 51, 237B,
Meineke EB Mi 124880, 124870 otter ciaicie:« e cxohe plata) a) icles a a/heNayn o's 116
20. A. a. ewaensis var. 1. North-South Waiawa Ridge, area 51, 234B,
IMeimecker Es BiMier 2A Silene cae seca irene oockestorni ols los tiataes Aioweee 116
21-21b. A. a. ewaensis var. 2. Waiawa-Kipapa Ridge, area 53, 244,*
INMeinecke SB MaiieTOOO ss yrvrse.chercree ain cele ate orctesis ore ei emieicys sani 117
22-22a. A. a. ewaensis var. 4. Waiawa-Kipapa Ridge, area 54, 244-3?,
IMfemeckenp Sis Matt GAA Sets tee wae nate pk im AIS 4 dy J re a 119
23. A. a. ewaensis Welch. Paratypes, North-Central Kipapa Ridge, area
G6 240L-0;. Meimnecke, BBM): T24040i 6.1.0 028. 0266 case cane cee 115
24-24a. A. a. ewaensis Welch. North-Central Kipapa Ridge, area 56,
240L.-4,. Memecke; BBM 124034, 124035). hccac-e2s.0 oe cole ems vine & 115
25. A. a. ewaensis var. 3. Waiawa-Kipapa Ridge, area 52, 243-2, Meinecke,
Ss NMP Ate hath.) em reraiatls bata Ria se baro nl tret Spake Aeneas Sears een eas 118
26. A. a. ewaensis var. 3. Waiawa-Kipapa Ridge, area 52, 243-2, Meinecke,
BB MUs rT S437 itrcts tates ote ot « averoreisioatetec sya eal she ctehcrs os texeiatchavo craters eve vorens 118
218 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
PLate 8
Page
1-1a. A. a. lineipicta Welch. Paratype, Waiawa-Panihakea Ridge, area 70,
232Ga, Meinecke, BBM) 11S4qgera an. eters oie ere ree aie eee 98
2. A. a. lineipicta Welch. Waiawa-Panihakea Ridge, area 70, 233G,*
Meinecke, "BBM: 121884). 2yscrWats cictotos there oi kere eaten nie 98
3-3a. A. a. lineipicta Welch. Waiawa-Panihakea Ridge, area 70, 234G,
Meinecke; BBM: TISAsi.c cic. Severe restarts Glo cre taleteterlete orale ieee 98
4. A. a. lineipicta var. 1. Panihakea-Kipapa Ridge, area 72?, 241-1-242-3?,
Russ; BBM T4057 i .ccisréjeclenv te staienitiors ree eteae Pace eee 99
5. A. a. lineipicta var. 1. Panihakea-Kipapa Ridge, area 72, 242-2,* Welch,
BBM} 120003) <5 Src.5 .rc15.c'm Site arvaele Oencleue ve, olshe exslolelola eka teeietes toler 99
6-6b. A. a. ewaensis var. 5. North-Central Kipapa Ridge, area 55, figure 6,
240L-I-1a,* Meinecke, BBM 121914, figures 6a, 6b, 240L-1,
Meineckes: BBM) 12402T:.5stenvetyererevovobeca oie ieveloreiciao oreiorayotereiele senciate 119
7-7b. A. a. ewaensis var. 6. Kipapa-Waikakalaua Ridge, area 57, 252-1-
2°, Meinecke; BBM 120035, 121030. .:0'ccc s)-ysc hers tire miter eee 120
8. A. a. ewaensis var. 7. Kipapa-Waikakalaua Ridge, area 58, 256,
Meinecke: BBIMMt23650vmaloadites nor eaters cnrole hee ometeriston icine 122
9. A. a. ewaensis var. 7. Kipapa-Waikakalaua Ridge, area 58, 257,
Meinecke; "BBM, 123058: sc:ac.dieraats slotere tern cietaversten rere eitaieteer eine 122
10. A. a. ewaensis var. 7. Kipapa-Waikakalaua Ridge, area 58, 258,
Meinecke BIBIMrT2367A hvac rcsvsttrecciisratoee mela reteie sicher ernie 122
11. A. a. ewaensis var. 8. Waikakalaua-South Kaukonahua Ridge, area 509,
261-262?) Memecke, BBM t21027 <4 ccsisac0 boc creas sienna 123
12-12a. A. a. ewaensis var. 8. Waikakalaua-South Kaukonahua Ridge, area
Bo, 262," Meinecke; BBM 121946. cen ane os anee sae eee oie 123
13. A. a. irunni Pilsbry and Cooke. Lectotype, ANSP 108778............. 124
14-14a. A. a. irwini Pilsbry and Cooke. Kaukonahua, probably area 61?,
Walder: BBM 50570 0s tance ote ee abe slant Cero eere aac irce eres 124
15-15b. A. a. irwint var. 1. North-South Kaukonahua Ridge, area 62?,
27773 Meinecke BB MelZlO73yl2iGO7Aaee ies conic crore 125
16-16a. A. a. chromatacme var. 2. Manana-Waiawa Ridge, area 43, 237,*
Meinecke WB BIMeTaremae science cisterersisecterele cine. clientes 88
17-17a. A. a. griseibasis Welch. Paratypes, East Waiawa, area 65??, O. H.
Emerson BBM)’ 103067 os. Stee oe ecipe oie iste sores Oakce eee oe neta oteiere 9o
18-18c. A. a. albipraetexta Welch. North-South Waiawa Ridge, area 64,
233B-1,* Lemke, BBM 115022, 115020, 115023, II502I......... 890
19. A. a. albipraetexta Welch. Paratype, North-South Waiawa Ridge,
atea/637, 231B;*\ Meinecke, BBM TerS40.), dc.» ove cee es onars 89
20. A. a. albipraetexta Welch. North-South Waiawa Ridge, area 63?,
232B,* Meinecke, BBM.1 21842. 0.cnama« 44cebicnmenteeaiinare cue 89
21. A. a. albipraetexta Welch. Waiawa, Thurston, BBM 130590.......... 890
22-22c. A. a. lemkei Welch. Paratypes, North Waiawa Stream, area 67?,
220h-1P. emke, BBM 115025, .01AGOO) iTS O20.) cies elteiecie eae 95
23-23b. A. a. lemkei var. 1. North Waiawa Stream, area 68?, 233Ga?,
Lemke; BBM 01 4009's sche arore.« 5: ovevare icicle chars terete, are fotos sNctaleretoueteretore tee 97
24. A.a. polymorpha Gulick. The lectotype of Apex neglectus Smith, BM.. 91!
25. A. a. polymorpha Gulick. A specimen in the type lot of Apex neglectus
Smith, “BIM fo 5 ccsaie cc overseers aie ree melo atole ote te) eutvetesster nee nerereters ot
26. A. a. polymorpha Gulick. Waipio, Gulick, BBM 70384.............+.. gl
27. A. a. polymorpha Gulick. Holotype, MCZ 39901... .............0000- gl
me
MNO: 5 ACHATINELLA APEXFULVA DIXON—WELCH 219
Page
28-28a. A. a. polymorpha Gulick. Waipio, Gulick, BBM 70364, 70384.... 91
29-29a. A. a. polymorpha var. 1. Waiawa, J. S. Emerson, BBM 102247.... 93
30-30b. A. a. flavitincta Welch. Paratypes, Waiawa-Panihakea Ridge,
area 00, 230G,*) Meinecke BBM) 1218750006 066 ods leoeu ais oss 94
31-31a. A. a. fumositincta Welch. Paratypes, Panihakea-Kipapa Ridge,
near 210 ee Meinecke, BIBIME mi 2lQO0rmr tian cies ie ace aac 100
PLATE 9
1. A. a. fumositincta Welch. Paratype, Panihakea-Kipapa Ridge, area 71.
PAQorNeineclker bs WuL2ROOO se sae eferaiererele crouiesere Oia cisleks els staxeterel als 100
ear inanlencorrapne Gulick, Holotype, MCZ 39003)... 2..-055-+00+.005 + cre 101
3-3a. A. a. leucorraphe Gulick. South Kaukonahua South-Central Branch
Radges200 Ave Viemecke a bIMe TO72T seme cmeacieie ceecicins caiclee cle IOI
4. A. a. leucorraphe Gulick. South Kaukonahua Stream, Wilder, BBM
SG INS erence Oct ae SR Oe tae un eer eae we’ sin dea IOI
5-5e. A. a. virgatifulva Welch. Paratypes, Waikakalaua Stream, area 73?,
Walderare Bi Maes O52 pisces om cscerte semis elec e ceive echoes rhein hers 103
6. A. a. virgatifulva var. 1. Waikakalaua-South Kaukonahua Ridge, area
Poy Gok > Materiel IBIBIL Higoye ooonsanouleboadnoobobdouonoc 104
7. A. a. virgatifulva var. 2. Paratype. Waikakalaua Stream, area 74?,
WalderaB BIMENSOS ZI 0 cree ce iets cre nieene ayers aie iaiete gras seas 106
8-8a. A. a. virgatifulva var. 2. Kipapa-Waikakalaua Ridge, area 74,
2rieal= ase RUSS a Via TOA Te Owl 34TOOna. telacisetiee selene eels 106
9. A. a. virgatifulva var. 2. Kipapa-Waikakalaua Ridge, area 74, Spalding,
REIN THTE7S Bremer hterce ea sct teres eve an ere anGtA coer os Sree alu ker ss aeaiel oe ictter akon oars 106
10. A. a. tuberans var. 2. Waikakalaua-South Kaukonahua Ridge, area 60,
Ag.» WMeinecke/s DBM) T2LQAO sae cak cian ee sid eames ele shure. Oe ais 109
11-I1a. A. a. tuberans var. 2. Waikakalaua-South Kaukonahua Ridge, area
60, 260-261?, Meinecke, BBM 121923, 121022...........s-c002: 109
rowan tuberans Gulick. seloloty pes) MiGZi 300008: eee ane eee weieiecieels © aici 107
13-13c. A. a. tuberans var. 1. Kalaikoa, Gulick, BBM 70414, 106734, 167043. 108
14-14a. A. a. tuberans var. 3. North-South Kaukonahua Ridge, area 77,
By Ameeimken io Mest O4An lt O42 eeincttercteie cies cevaciereisieea tse see III
15. A.a. tuberans var. 3. South Kaukonahua, O. H. Emerson, BBM 103971. I11
16-16c. A. a. tuberans var. 4. South Kaukonahua South-Central Branch
Ridge, area 76°, 260A ?, Meinecke, BBM 121068............2.000: 112
17-17e. A. a. tuberans var. 5. South Kaukonahua Stream, Wilder, BBM
RSs SMM eee resets v aretes ech ian onthe 4 « gtebvsnel A ato aie \eyel sucaearecer tee ws hae hare 113
fo. 4. 0. comformis Gulick. Holotype, MCZ 30011... .\.. 000 0scc0eeceess 126
19-19b. A. a. coniformis Gulick. Ahonui, Gulick, BBM 70459, 107030,
OZ OAM re tenat ater Se aisle eoe sie) ehcuh ch aefor whales aie ciel ota gle ete vee) ecwa avorene. Hale Loorsvetals 126
20-20a. A. a. coniformis Gulick. Wahiawa, Gulick, BBM 70368, 107058.... 126
aie A. @. versicolor Gulick. Holotype, MCZ 30007... ....0..cccccceseccees 130
22. A.a. versicolor Gulick. Ahonui, Gulick, BBM 70380..............0005 130
23. A. a. versicolor Gulick. Kalaikoa, Gulick, BBM 7o450................ 130
24-24a. A. a. versicolor var. 1. Kaukonahua, Thurston, BBM 130693,
TS OC OA NW Me veetde SPE eT ato be eee darn era poate ae oon Mel Setar ep 131
25-25b. A. a. versicolor var. 1. Kaukonahua, A. F. Judd, BBM 110054,
TT OO SSM eer eee ste a MSIL! ciate oreeetc in PASO RS helenae 131
26-26d. A. a. apexalba Welch. Paratypes, Wahiawa, Gulick, BBM 70444,
A OAAG ST OAT2Y crise re acta: cis, eveveveie eral ala iia ate ters ose taleke cevacealeyalecataranctaie ib: overs als 128
220 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL.
PLATE 10
1-1b, A. a, steeli var. 1. Welch. Paratypes, Kaukonahua, Wilder, BBM
TOAAG Si. Sats ee oe Oh Oe
2. A. a. steeli var. 1. Kaukonahua, Thurston, BBM 130674..............
3. A. a. steeli Welch. Paratype, North-South Kaukonahua Ridge, area 80,
271, Russ, BBM 134174" <0. oes i eee eee
4. A. a, steeli Welch. North-South Kaukonahua Ridge, area 80, 271, Steel,
BBM. 120008 sor ed vias Ms cts'e gael eee ee ‘
5-5¢. A. a. steeli Welch. Kaukonahua, Wilder, BBM 50577..............
6-6c. A. a. punicea Welch. Paratypes, Kaukonahua, Wilder, BBM 50583,
BOBO © x: tssidusecs occ a's ate hati veratae a ieas OITA ee ee ee
7-7b. A. a. punicea var. 1. Probably from the North-South Kaukonahua
Ridge; (Welch), Wilder: (BBM: s0s78*;). 5: a2. achiecaccs eee
8. A. a. lilacea Gulick: Holotype, MCZ 30008: oc. 02-0 eo ces see oees:
9. Axia. Hilacea Gulick, Kalatkoa) BEM igoqr7cue. ce ee ae ee cee
10. A. a. lilacea Gulick. Ahonui, Gulick, BBM 70372..............eseeee-
11. A. a. lilacea. North Kaukonahua-Poamoho Ridge, area 83, 280,*
Meinecke,. “BBM (12T682.05 sac tice poste eaten Oe een eee
12-12b. A. a. lilacea var. 1. North Kaukonahua-Poamoho Ridge, area 83,
280-1-280;* (Russ; ;BE M cESATeD ace us tenn. teehee oe ane
13. A. a. lilacea var. 1. North Kaukonahua-Poamoho Ridge, area 83a, 281-2,
Meinecke, “BRM 1oTgsy7: 4. kas ecto etek ce ae ee ee
14. A. a: gulickn Smith, Holotype, MCZ 30834~.,~... sens: s aueee eee
15-15b. A. a. gulickii Smith. Ahonui, Gulick, BBM 70377, 70413.........
16-16a. A. a. gulickii var. 1. Ahonui, Gulick, BBM 10443, 70426...........
17, Aig. flavida Guiick, /Holotypes MCZ so004. 23.4icde.Necs oe eee
16. 430. flavideGulick: “Paratype, MCZ go004. ele ee
19-19b. A. a. flavida Gulick. Kalaikoa, Gulick, BBM 70418...............
20. A. a. flavida Gulick. Kalaikoa, Gulick, BBM 70458...............+--:
21-21b. A. a. gulickii var. 2. Kalaikoa, Gulick, BBM 70416...............
22-22a. A. a. glaucopicta Welch. Paratypes, Central Poamoho Stream, area
87, 280K-2, Meinecke, BBM 124565, 124564..........sececeeees
23. A. a. suturafusca var. 1. South Poamoho Stream, area 85, 280-7, C. W.
Isle; BBM engaBagic cua eee ates tule Sela ace ae ee
24-24a. A. a. suturafusca Welch. Paratypes, Central Poamoho Stream,
area 86, 280K-1,* Meinecke, BBM 121907.........00eeeeee eee
25-25d. A. a. suturafusca var. 2. North Poamoho Stream, area 89, 290,*
Meinecke, ‘BBM. ‘122093,, 1220229... Wont otis hose eee eee
26-26a. A.a. brunosa Welch. Paratypes, Central Poamoho-Central Poamoho
North Branch Ridge, area 88, 280L-2, Meinecke, BBM 124587,.
27. 1. a. brunosa Welch. Central Poamoho-Central Poamoho North
Branch Ridge, area 88, 280L-4, Meinecke, BBM 124605.........
28. A. a. brunosa Welch. Central Poamoho-Central Poamoho North
Branch Ridge, area 88, 280L-3, Meinecke, BBM 124501........
29-20a. A. a. suturalba Welch. Paratypes, North Poamoho Stream, area 90,
200A A-8," Lemke: BRM 1528Tat ot cence eee eC eae ree
103
Page
143
143
141
142
137
137
137
137
138
138
133
133
134
136
136
136
136
135
140
149
148
149
146
146
146
eNO. I
3. A.
9-9a.
10. A.
mT: 4,
20 RA.
m3: A.
mAs A.
rs. A.
16-16d.
Mofo. All
18-184.
mo; A.
20-20).
21-214.
22-224.
23-23b.
24-24d.
25. AA.
26: A.
27 LNs
28: A.
29. A.
30-304.
ACHATINELLA APEXFULVA DIXON—WELCH
PLATE II
. A, a. wahiawa Welch. Paratypes, Wahiawa, Gulick, BBM 70423,
PAA SS OAD Ti eaictatons syeteen severe & cieiris! ereiriets Hace ey uae M Mle ato Mat BY spon) sve heae, Golan
maapicata Newcomb... ectoty pey Misr cee ci eo st-foicisarels selves «ia e's cre
. a. apicata Newcomb. Helemano, area 92?, Gulick, BBM 107021.....
. a. apicata var. 1. North-South Helemano Ridge, area 95, 312-4,
Menmiccke: (BEM roe san) (2 emesis 2 ae ccrha wa sie <a kyeis thos peihig ain Cate
. a. apicata var. 1. North-South Helemano Ridge, area 95, 312-3,
IMcinecke ws ED Vieenae sO acme setereriaters nici cccrele icici el crerersieeretere fare
. a. apicata var. 1. North-South Helemano Ridge, area 95, 311,*
Mieineckeh BB Me T22T AO thc miecrae tetas ters msn ahelerelae’e slat ete a.
A. a. apicata var. 1. North-South Helemano Ridge, area 95, 311,*
Memecke-s BiIBiIM stoone Cito Oita nmcccucterctateeloke ol aist ole ele ats ie tie ¢ cre
a. cervixnivea Pilsbry and Cooke. Holotype, ANSP 108807........
A. a. cervixnivea Pilsbry and Cooke. Poamoho-Helemano Ridge,
area 93, 300-6-301-2,* Meinecke, BBM 122104................--
a. cervixnivea Pilsbry and Cooke. Poamoho-Helemano Ridge, area
O25 400-A.* Riss; |b Wl) TaA2O7. oo ericsson od ots la nid o's w 510 6
a. beata var. 1. North-Central Poamoho Ridge, area 94, 201-3-5,*
IMeineckessb B Mate roozi.. qattis rater ee eaiaiae nar derevaarionsic ers
M@ubeatavar-ei) /eoamoho, Wilder BBIMl SO500n- 242) crc asi < stein ces
a. beata var. 1. North-Central Poamoho Ridge, area 94-94a?, 291-2-
SOA eVeinecice Es Migs 2 01 Oleic crcletc clerere ere erste cists oleracea) sicletelorsi
a. apicata var. 2. Kawaihalona Gulch, area 96?, Thurston, BBM
TES OSGHEN | eres St oa cae aa sv Tommi OE oer oaie seats clofuyeare eTmelereteie sieidle eis
a. apicata var. 2. Kawaihalona Gulch, 326*, Meinecke, BBM 122184.
A. a. apicata var. 2. Kawaihalona Gulch, Wilder, BBM 50500-
BOS Ome ca eiveleyhaposeh shersvorepalclsneater etek ol anstnte taaia) cieVelstclaiiatstwieraletelciele!s\e7 ale. e\siis
paperiiliva: Dixon: CopyeOt OLiginal figures. soem eee ee -erelols wees
A. a. apexfulva Dixon. Opaeula Gulch, J. S. Emerson, BBM
REVS SEN SNL eey, SECM tt AGS iA y Weds 1 Ais Coe Aer BO SOE a itta sikw tos
a. apexfulva Dixon. Opaeula Gulch, area 97, 330-4, Welch, BBM
MPRA OMNIS te Soe aie oe eer ee rae a orca aie ops le ove ai Shecevederalsieregonake
A. a. bakeri Welch. Paratypes, Waimea, J. S. Emerson, BBM
LOLS SIME ete ee Nera eyes cata heratra ice wale ey aieare iplerarers alates
A. a. paumaluensis Welch. Paratypes, Paumalu-Kaunala Ridge,
area III, 431, Meinecke, BBM 123868, 123866.................-
A. a. oioensis Welch. Paratypes, Oio-Oio East Branch Ridge,
area 114,)400A, Meinecke; BIBIMl 125000)... 3 .)fec.. ecle cee. ones
A, a. oioensis var. 1. Pahipahialua Gulch, area 112, 450, Meinecke,
BIB Mist 23805 1238005 29 GO Sa aera cicicvel rots otenatatee ate eta\evel' olor
A. a. oioensis var. 2. Kaunala-Oio Ridge, area 113, 462,* Meinecke,
EIMIRT 22 RO som cet: Sues eae eierevterereten ss canis aretah cscrebei state feucie ioyeteleyovsis
MRAP U SEE er nel OL OLY Perk Es NUearctarefeieik ators ele ieis nietalleta shake «3 eke sahae
a. napus Pfeiffer. Waialee, Gulick, BBM 106716..................
a. napus Pfeiffer. Bought from Sowerby and Fulton, London, BBM
IOVS AEAIE Tease tant Le ee ty ah de ach eT tlt 2M ks pI IE win eo iy
a. leucozona Gulick. From Arthur Blok, of London, BBM 165627...
a. leucozona Gulick. Holotype, MCZ 39906..............-.+e00-05
A. a. leucogona Gulick. Waialei, Gulick, ANSP 92656...........
221
Page
186
182
182
182
I8I
181
222 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Page
31. A. a, leucozona Gulick. Waialee, J. S. Emerson, BBM 102337......... 181
32. A. a. kahukuensis Pilsbry and Cooke. Lectotype, BBM 73085......... 175
33-33a. A. a. kahukuensis Pilsbry and Cooke. Malakahanae-Laie Ridge,
akeasli ot, Lhurston, bb Mensog7s eure ere een aoe 175
34. A. a. thithiensis Welch. Paratype, Ihiihi-Kahawainui Ridge, area 116,
5TO, Meinecke BBM CTeecO8s. cocmare cine smectite nace eee 187
35. A. a. wailelensis Welch. Paratype, Wailele Gulch, area 117, 529-1,
Russ and Welch, BEM 1604445 vo vcaedsccanersecsee sacle ee cnet 188
PLATE I2
1. A. a. aloha Pilsbry and Cooke. Lectotype, BBM 73087............+. 160
2-2a. A. a. aloha Pilsbry and Cooke. Kaukonahua, Wilder, BBM 50561.. 160
3-3a. A. a. aloha var.1. Kaukonahua, Wilder, BBM 50550............06- 161
4-4b. A. a. roseipicta Welch. Paratypes, North Kaukonahua-Poamoho
Ridge, area 102, 286, Meinecke, BBM 124780, 166133, 124683.... 162
5-5a. A. a. roseipicia var. 1. North Kaukonahua-Poamoho Ridge, area
103, 289-1a, Meinecke, BBM 124736, 124737......000ccccccesevs 163
6. A. a. roseipicta var. 1. North Kaukonahua-Poamoho Ridge, area 103,
287-2, Meinecke, BE My 1O0TAO. xs «soe et nto oe eee ee Cee 163
7-7a. A. a. poamohoensis Welch. Paratypes, North Kaukonahua-Poamoho
Ridge, area 101, 284-2, Meinecke, BBM 124152, 124151.......... 166
8. A. a. poamohoensis Welch. North Kaukonahua-Poamoho Ridge, area
VON) 203-1,9 Meinecke EB Vi siGOle7- ean lenucioni ciara 166
9. A. a. roseipicta var. 2. Poamoho-Helemano Ridge, area 105, 307-1a-1b,
Meineke BB Mi 122000} waits Kener ure tee acon ose cars wae 164
10. A. a. roseipicta var. 2. Poamoho-Helemano Ridge, area 105, 307-3-
S06-1,) Meinecke -SB BM. -IeS38O.c% noc on talene «cc ak vaetee certet 164
11. A. a. roseipicta var. 2. Poamoho-Helemano Ridge, area 105, Lemke,
BSS MSuiGAaaa: oc staves & 5.8 eiayorala tee oC a etacainie e rare oie eee 164
12. A. a. roseipicta var. 2. Poamoho-Helemano Ridge, area 105, 308-2, Isle,
EBM G1G63773) cia vatcelery sec trate a wterei este bn wtih ke chee Oe ora ee eee 164
12a. A. a. roseipicta var. 2. Poamoho-Helemano Ridge, area 105, Lemke,
BBM road. ise ek we 5 seats oc SR eed Oo rn cee eee 164
13. A. a. beata Pilsbry and Cooke. Lectotype, ANSP 108809............ 169
14. A. a. beata Pilsbry and Cooke. Poamoho-Helemano Ridge, area 104,
303, Russ, BBM 134egaus 034 Cae -ctavebow wecadie sent Meee meneen 169
15. A. a. beata Pilsbry and Cooke. Central Poamoho-Central Poamoho
North Branch Ridge, area 104, 280L-9, Russ, BBM 134234...... 169
16. A. a. beata Pilsbry and Cooke. Poamoho-Helemano Ridge, area 104,
304; Meinectke, “BBM rg20gt ens: ; aye stsicd Seen ine ete 169
17-18. A. a. beata var. 2.. North-South Helemano Ridge, area 108, 318,
Meinecke, (BEM. 128430, 128Aggti 7 eae cece mee ene etce 172
19-19b. A. a. beata var. 3. Helemano-Opaeula Ridge, area 110, 337-4,
Meinecke, (BEM 125778; 128770, 128776... . sabes we tee eee 173
20. A. a. paalaensis Welch. Paratype, North-South Helemano Ridge, area
106, 313-3, Meinecke - BBM: 122206; ch.vku1 aacscs co eet ee eee ae 156
21-21b. A. a. paalaensis var..1. North-South Helemano Ridge, area 107;
figs. 21, 21a, 316-1-3, Meinecke, BBM 122175; fig. 21b, 316-1,
Meinecke; BBM 122367.) oseack dts tdanee een ee eee 157
NO. I ACHATINELLA APEXFULVA DIXON
WELCH 223
22-22b. A. a. paalacnsis var. 1. Helemano-Opaeula Ridge, area 107; figs.
22, 22b, 334, Meinecke, BBM 122397, 122374; fig. 22a, 336-1,
Memecke ub Mb ie2sT Ae, ane rte ec ean eer aay atest cna oeats Cesc 157
23. A. a. vespertina Baldwin. Holotype, ANSP 656090...............-008: 174
24. A. a. vespertina Baldwin. Kawaiiki Gulch?, Wilder, BBM so605...... 174
25. A. a. vespertina Baldwin. Waialua, J. S. Emerson, BBM 102309....... 174
26-26a. A. a. kawaiiki Welch. Kawaiiki-Kawainui Ridge, Wilder, BBM
ES OGOOMESOOOTE te erieets teres aisnoter sto alee et cr Ta oe OIE en ha aes 159
27. A.a. duplocincta Pilsbry and Cooke. Lectotype, Kawailoa, BBM 72831. 179
28. A. a. duplocincta Pilsbry and Cooke. Paratype, Kawailoa, J. S. Emer-
SOME EM OSSD ato \ia rar tetans Rin ro eRe tah ACHE iw sese aa ToC cleo ia ae teers 179
aan
er en
Peat
VOL. 103, NO. 1,
SMITHSONIAN MISCELLANEOUS COLLECTIONS
20 MM
SUBSPECIES OF ACHATINELLA APEXFULVA
(For explanation, see pages 208-210.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLES 1035-NOo 1 PEs2
SUBSPECIES OF ACHATINELLA APEXFULVA
(For explanation, see pages 211-212.)
A
HSONIAN MISCELLANEOUS COLLECTIONS OE OSw NOs TPES
SUBSPECIES OF ACHATINELLA APEXFULVA
(For explanation, see pages 212-213.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 1, Pam
eo Cot! eee
e ~—jeare in
Ping gl, a > & =.
wet de te
20 MM
SUBSPECIES OF ACHATINELLA APEXFULVA
(For explanation, see pages 213-214.)
J THSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 1, PL. 5
SUBSPECIES OF ACHATINELLA APEXFULVA
(For explanation, see pages 214-215.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 1, BB
SUBSPECIES OF ACHATINELLA APEXFULVA
(For explanation, see page 216.)
ITHSONIAN MISCELLANEOUS COLLECTIONS
SUBSPECIES OF ACHATINELLA APEXFULVA
(For explanation, see page 217.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 1,5
SUBSPECIES OF ACHATINELLA APEXFULVA
(For explanation, see pages 218-219.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOE. 103;,) NO= 1, PE. 9
SUBSPECIES OF ACHATINELLA APEXFULVA
(For explanation, see page 210.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 17m
ee
A
aXe
A
& <
Lng 29°
SUBSPECIES OF ACHATINELLA APEXFULVA
(For explanation, see page 220.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLE. 1033) NOs, (PEs id
SUBSPECIES OF ACHATINELLA APEXFULVA
(For explanation, see pages 221-222.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 1, PL. 12
SUBSPECIES OF ACHATINELLA APEXFULVA
(For explanation, see pages 222-223.)
ine TO SECTIONS, GENERA, SPECIES AND SUBSPECIES
(Names of synonyms printed in italics.)
PMA ASE nine eT ELiheI: wards lars clsiecuewiebecrsteesre'e eercicce ad e.ecsjejeuaveial oven the ways) 21
Fee Pere LatPro ed Shaler ch calle cca che Penersionetots Rusieiatereicie/e sais. @sishele eisiaa ies 169, 170
ELS PII Ae hee’. lately salah sea bcs ty xisieieha ia. ibinie alas give wd, sa ue ws ea 169, 170
Pate Me Gli eaneawOS ih TOA a nats aroc/s valecie Meise aoe 5 sss vs decciels wpe sie’ 89
eB oetetacete AMS TRILt Ine theese erat. 3) cyanea vals sjcieietela S a.ce cicadas a ie idc wala e.e 47
Plbnartca lett ee Pa AT Ce aR LAUR a, tsie cle jecarenaiesajsreral ara. «/S.5) ous lovelies wef so /a.eya/e sp sie 6rs 48
alge shiv andi @OOkes area, TOO! . c-..)c)s as tse oss cise slerele.a t10)4 0 ere 0 sieie 160, 161
eM ht TMC LAM TCT Cel eT OSes Val acai acl aie) sfaea tetas averal cveliens! ate Gravel cca suis ab she stereos aes 161
See eM EDIE CTEM sae Lee iaicig, ely) ee bor, cing si nyejhl siti sles aisis’weemeitane c's 128
ap lemsy tlio Ox OTINM UL Cals 7/5) fay ete arereiayc (overs cies cielo.cisie-o-e eiel oie ei ele cieideueraisiere @ eleieve's 176
per emectmay) (Pieter, ALE OZ... 06s .n<vceee es ced eevee sean sennce 152
apicata {Newcomb) Pfeiffer, Pilsbry and Cooke....... 127 erates suansAs
137, 143, 148, 149, 151
Apa here AMMEN LoemB eta Ic cU gO) Sienehatelralctele) «/ <) faye) -\ici(eiiere! eke s1/<1 6/'5)/<) 6/0) ifm) c1.6,.01°¢) «1 =, 101/0)ral(sy'en) ale. 01.51 67.810) 6 153
APIGALARV AT 2 HATICA) QO. oss cis /a civics vine cuie ¢ Pe eee et aca vs setae 155
Pence LGlammclh CNS On snrd a misrelomysielc, «jolts aieras) a5 Sore. di6ieicbe.clqie.e <eievei dierele eaves coe 64
PRP AMSASUY MAN Gol Feu LMR ATC Cet Teste selec Pele erected sere, sia 3) oy ava 5 Sic. e3¢ er8ls w ciesays)e/G:enaya-aievete ela 66
rel est VALG CLONE See rac tfaley ccyayaveere, ais care 616 os el dielsisi eve d 6diStea' ei aire eae 180
Demat s nian COOKE areal TOA cs coisas 6 siete so.cre oo <0 eciavers, dye viele so.s ie o's 169
Peer icetyeand Cooke, not from atea 104.2... 2... cc... cece cc eee cceee 171
Hecieimnd iam lPEATCAS OAM OA-OA AIS cst clcleteinlsusisicoie a Sale Calssigatelesicssmabld sess 171
Hear MR CEUIOMMEAT Cra MMT Oo ea rteme sie ran oie, cosy T ss wes Sued sue cigar ey ci duss oes ares: </euw ere ois) a. esene 172
eee METRE IANA DUO a3 cf)s)as/a(cc ciclo v0 ecard manic se whe see deeenvaciecmesa 173
mn MeMMEN E et ree oso ofn Pe diel iovujarateysie's. visv Waals cielo da diewie esas aaeie 61
PUMA CO MRVEAI MIMM LT CoO Per iele etic: oe sicvars sii neie cin ea cies sioa a decesecheew nace 62
EINE RULE OMA L CAMO Geert crane eh SR ee dale s 62
RPGC SAMY NLC CHIMEAT CARS Ocectanrtrss earelciie eines eines Rice oeaie w.ciaiece ee sigies oils wise cletae 146
PSIUSINCAMN VLE GAMER eye Metra erect teeta eh dich a arars Mies ahh coat cha-oiave eels Giwlarel 6 etdroconde saints 35
Ee Satie am a eT tC Tome Ben Gite ey sesere oss ea iciel seers Giese eG Sa ialh ciel S101 ois Svea ones 21
Peeremmednetishiry did Cooke) area O31... 020k. ci ieee scccccesaccesaeees 167
Peaermevea patterime isbry atid Cooke... 0.6.02... 0.00 ce ean ees ce nencees 140
RLSM NCC OTT Mem CclatG | aie ayeletc/o) aie) 9.0) a'eieforersier Pass a: ic oicvei nia valeliveleile shave leis 'etsieve s 3I
Pasnew oomb, Pilsbry and Cooke. i. ...4..5..06.0% sce ccs ceens 28, 30, 32
Binomatacinemelsbiy) andl Cooke, areav42: sv. kss. besos cscs sodetos os cee 84
Ree enemy Aen rare eRe ALS Le tte het ecinbcac ees ocneelnsh ae ent 7
IER CRT ROP INCA AGLS hv.) s 5.00 ipielsihie were Sis bus hs be ab ee chic eae Seema 8&
IGHC HES Ie SRALCAMUOM RE Circ RiGee he clemies| eamielele ur aeeiiain & Sine Move dicoime cemelns 46
SRO SM EZCIINE Gham emer ee LM SRA fe bihe, kA fevaltssh cts aici g © exe oie dere anerele sea lsio oh 181
SETA TCO Tal See Cnt Cheat eRe epee ce ciracts eres wie Psteyes ets beagle letoiocaleters) sratanaveteiee chensie. 126
UG ele all Wllee he AuA Gr yeiepae cays ciciers saison Rake aio wie s alee Siders nie calc 75
TOMY Ale MLA EAS SORT ie ae htc sult soe IRM alae mimibre aie crarera a Rralstcret auras oie i
OCIA Line eR UC eel Ore RRR Raye A iodo So sceieiest aetolateretortte bakers teint cee isle Geom eate 77
16 225
226 INDEX TO SECTIONS, GENERA, SPECIES AND SUBSPECIES
CECOLAa WER USSAGs Frio tate os, eis so nateanelar seco ra raleee eeeterayasc cists ete eat Re RO ots 23
MCCOFG! REEVES aids sciasidk aa lo-ateoie e ore leven tronctarmintabaiern) galelele RICO a ore meee oe 40
duplocincta Pilsbry and Cooke, area 97A??.........-ee00: Meteenae 179
ewaensis” Welch areasSGincs.clcscars eleva sterere leon ovate oem area ete creer Saye cwte einer 115
BWaclisisivan,, I, ahea) Hlves-csc vce schon om care tas ts ae ee cee a.oictereee 116
EWACNSIS TV ATs: 2st ATA! Solo cicrerercleiesasavere cioiespreieieye clase lei ereture a eleva nae eee 117
CWAENSIS VATIL3; ALCA: 52. cisieai/ee sais os atatecal jaley ave Ul etar eters te evek ete nea here Pete LLG
EWwaensis vat: 45 area S54... ocsac ss ee Sr aie ey ER eee aia Ugrottre sree ters 2 ee ALLO
GWAGHSIS VAL Sy ALCAN SS isrcrerate oiere evs aimeueetetclaralevateoarel chatter aeiaieie erates ts roe eeaeLO
GWaeNnSiS Var, pareaiS7 sion naw cereceinn saci biieses ayes atetere.sdaaleaeete tenes ait 120
ECWACUSIS Ware 7), ATCA: HS isisise chen e'ss piv key susie wee minloieie arate ote reer renee 122
EWACHSIS Vat TO. sALE Ay SOS tres cletstare wise aie iavelo aiomeceueturera fete ciare clers erie ait ioc tenets 12
fa VId PAG ULICle Sys ae act. fetes canes ee pole ene ernie Miele ers sd tiNerstelaste teraele 136
HavitinctawVelchwarearOOnemstcceeie sins eevee ce cnietrecn sid Been On eee 04
forbesiana Pteiffer, area 67? ..........-+.- Ses eee NED ero tS vs ; 28
fumositincta Welch, area 71..... gi acctiatorstaneie ors Be TS core ten Bier OP eter . 100
fUSCOStriata VWWelGH, \aALCa Ghia ccele nicer sharetalea wists @ elate Pea eraye roe Slojeiaierete 32
glaticopictal Welch: areaya7 esc sci aaeleciers Sie cispakciele rece leigate om Oaktree cae Ne 140
globosa: Preifier oss ces eemroaiegetestl ett Danis ieatae nearer ip alele fe aloe oetenees 36
griseibasis Welch, area 65??......... Re ects Fn OL ISO beaks : go
WUC MO MINE ties os, cores atace aioe Barer arayeeete cease tetere bie nci ateteee rors Senko 133
oultekit vate Waly... “els sere eo Selene Steere ice eiaterstehtateteiete a beid-abars tetera cremate 134
UIC KIM Walle 2 einere < eiaralieieieteteisiatete eis shetahelstereteretoiohe tiers Ricoh abes siettiaee hele ee
hanleyana Pfeiffer ........ Soko ete Bro eeiiere evslheievemcareeistens elistayahsterejare 37
hanleyana vat Wess. ss etatal eaeetansicte eee siete Baia ease Soe Satehiorie oe SS
ihihiensis Welch; area 116....5. <2.) . SUL eo eon Ae ee ‘lavas pyavasein aces Riese pero
IMMOtADIS GS masth cor. eds yerolls woe ee tees ies aieralbia.avanete oa ttaley wets ialasare erereyane Oieites teem
irwini Pilsbry and Cooke, area 61?........... Satetersiotnetelstene ate Sica atelier Led
irwint Pilsbry and Cooke, not from area 61?...... 103, 106, 109, II2, I20, 125
iTwint Var. 1, Catea 62°... 65. ere< SoRid ira (utaeetentata ees Te piaiers Shafeseee savers a ete nben
kahukuensis Pilsbry and Cooke, area 115?........+-- Soevialasle Orehelevelte ates 175
kawaiiki Welch, area 99..... AT Mee tie inte orereiiate eantets eStart aU pis eteiscoe aS
lauranie Welch} area 27; 27Ab cassie Pace eatin grtke sso cueneatarcebitrete noe 55
lemker Welch; area (67:t-scler etetasoueretore atime aiocune aaveterd are teaeiele avotiels tree nee 95
lemkei ‘var 41 sareaO8 2 's.ccic,o%c1s oe icreubiscote eipieue ete cteterelalere oie ie eratoleretore tcteterstersloreromnchzd
leucorraphe Gulick ......... Dee cree anes AV ahs Nose eanee Nal etera ctoietewsios ke aRnOL
leucozona Gulick ......52.... Bere tec ieie I aikaierelersrone teteies eles sw tid so leyararmre rete toetoreye 181
laces (Gulick sarea ‘Sabi Pin) .cc si. semicewene oes wef paths reas heaeiele sativa 137
lilacea var. I, area 83, 83a...... Clare Dae wrotuevatereen chaser tare a Atcvete cep re See ISS
lilacea.. var.\2, area’ B4ancsusck)s see e cece eee cteceececenseceesseseceeseseess 140
lineipicta. Welch; areai7O,...05 cis iaveectesars ieee sii eisteseralateorsi oa lesnickatelonsicrsetetleteetPmEOS
lineipictaivare Ts arearg72.e ie ag Mailaras cfovetabe abadetarayers ture) ete teers « Shalavesate(aierete cleresep OO)
INDEX TO SECTIONS, GENERA, SPECIES AND SUBSPECIES 227
RE REAMMICE IIS SAGE a seenreyk eies bycitis inverse eho iaisttenha Suacsapeetes Sahel aie ciereies Beato 2375 LOO
PME MG Te SING PTA RII Ais fin Slee bw kietecvin wipe FAAS AICHE. 2a SiGe inte isis 176
BEIM VISION CL CH sh AL ay SAX here ave) s\eisiei ake picts ay nlensls Weleretoratabece el alenche Matepe eos ean eees 70
RSTO SIME AL nT ssley cos sie oisre ss St cid clecers sa telalaie siete seam ottat ut diane oaneniced Quail alata 71
SERN SIV DLS EO aT CAS 3 cc Seem ns ere ale ae er ore asiere ake eee recs have Gai oysters eye Saver 2
Eee OGRE Ca manea ste store chogiepe ht ceeds TIME Saat cece ater 30
MmTe Rea cM Eto Nel Sy atop sysdova svcue cer tau eve fuy stars execs ooelred gape iin escent ie te li, Oy aia a
PAINE GMM SLM IRE Tang eccee’ataccx etnies VS e eiate eseuaiores olds ais allen covets otetevateve catty sharon reve teutes acta ote 182
SE REIMER SSTTIIEL Dooce Wain, toc. baie aie Si dus Gis She havens Bsns BGI nel «eB er Shake oe QI
op cinmarciah’ VAAGIGT Ite One eae aa oe eid carer a aie oor a arcane 78
ae Min TALC TC AAO ea tee crayei einai ayes bi ob Gon Wer aaln ARES oe see 79
RP UTIGt mV Attn eA CARA Of 4 SACS cecvelaye tec nian eters Oitieciele ehaueteleioiehare naarvias lela at nese 8
Re MESCC HILO Ieee een Toe re ota ieee oicicic me tvoiste teeta Melee casks oR eI oh cee eee 37
PEAT SISMV VLE Ceci aly iA ec eva crcte sisi «cid eve od -atecs Tore sane nn og GE echo UE. toeourkal ah neta exe 184
Ere mPM ATCA PUD yc ies tie or om uate anes Wee a eee Signin < nak eae Oe nmisthts 185
MPR ETE STS 1A RAL Cr yeL Use erica as gta Tels carcvcvansit ov aneee eee rete ne Ey orale arte acer 186
Bra (ELC KG ALCA TUS iicniees ct ises chain.c fs ie sschs oy eatin oben testes Seeks PRE nee 48
PeUTRIM TeV OT VaALGA see eve cep raise eve cheicicvuyehs aisha sere re whol asia nae sha Doel e 57
mine eiciier lilsbry and '@ookess.....2-o6.66+sece 56, 58, 61, 62, 64, 70
Dea CUL Pe ACCALO Dor, fi uaiareitr cits ale) s/arenelsicssreis ie elves Glee 6 Glace wida ce ak eteie soe RntTG 58
MTHS A OT AAG SP nn eek, ate claccie Som eS aratanp ated « 2 nesters tees < weno 50
SEM ARY AIRC MAGE ALO crete Neer 0h ne ci ieee rolel siete ryene oleae O ovetsio oie oe ewel ate Cyer ass eropare 60
SEMEN VCE aL Gay LOO i aty site rewid nodes aha paie bys o aye 4 aphasia eyecare ay she cs 150
PEGE CALA Oia eey- ciate SAA aycces & arotaba Sets wi nisercleei she sibi es & Ae ee 157
MELAS TSTEV ERT eile LOO) siertcteien 6S ccs boal Saf Siovewierow goa Se whe toe wiles s Sess wa 158
Bea TERE ICI LEA CONT Sit. oi als) alelic o clo shnsssvovaiois wi ciate ee. Dieceanteug idcha apie ote es 56
Beta ENSISMVV El Closest ea pI Ts c., ot. erty ait casters te icyaronn oists soos desicie eee eomarae eee 183
Peaiexaee tishrysand) @CoOke, Area. AO. .cs)octacscesconey oes aniwnaged.etage 72)
eave ro Pilsbry and Cooke, not from area 40...... 00.0500... 0000se0 088 79
SEN LLIESO ihe RR RoE ct Sco cle arta hee sac den tities DEH b ARO tees 176
BSI el ChigaTcaec iss cic ite keke tc Pn e anne ive See ns Se 2
SE MAIR SAV VCLCH ALCAGTON 6.4 0°55 0's. 200d os ass cities ihe FAR ore eee Bae 166
BoPtirTeniy pede Gn Ill Gk, malt eta Os einer sietavatede cies. cre ts ee ittawtonstete coke a can rata alerts QI
RTA ALL sey are che ASG isis oye wicisiaystsd me he Aiclers eae ieee aun wens oem eae 93
ECMO IE KIRCET et COME te 2 pdt paren ah ie 318 cra sic care ous, ow aE Re On later ROE 141
PIC OAMV ATE RAT CARG2 Gieicta ctr area oye uiars, articustalav simone tetonon aber icte wivin Goa Wo ors ane 142
Eadtata Gould, Partulina........:.. av A ae Pa eae ae ate Re kt ts Nee ae ee SSO
EEA RUNG MRO eet sah 5 cites J aigetntca iota aimelte giao) eee ea eee 54
researc CMR LEA NOD Eid '5:0'2 a bie are dl aint paca RPA E aie tas ise See ee ee esl 162
RACE eM ATCA STOR hs ai nioseis.5 Sie ieee) fee tears Bava a Shain se Sole nie 163
_ LE ULE I SARE tet ic a a ad ate em at Ae 164
MMRDA ClCH marear Ayn tect t 4 tek Ass nh Re Boer een oem la ee 83
SMERANCEEVY CUCU NAT EAA T Ne, u5.2)3 205k Seis Acide ne cael dehy precios Ae os a te 49
RR TRINGSS CLAGV ANT Perr ste a ni erise TAs ch ts AE adh RUN ALS ly ARV ed Any Veron teres Es ch 51
MBM EtA Aker Om ALCA DUO cen 2s Fre tt caret eee oe ale a Re 51
MRE ROE CUM tical rapes eevee tyme tres te ute SR cael eden UIE oie an Mabie A 51
228 INDEX TO SECTIONS, GENERA, SPECIES AND SUBSPECIES
SEMMNIG ET MERKE,. BAlHUs iors a otic sinsele ons aveiaves tts Qratele biclore On Mice e RET ee 176
SemumMgra amark. monogonta: coq aacicis besa acle ei eiace oe eee Cee 176
simulacrum Pilsbry and Cooke, area 46...... ae Aae cicbyeate heen See es oo. wOL
simulacrum Pilsbry and Cooke, not from area 46.............+0c0eeece0e 7
simulacrum var. I, area 38...... ato Ne ata Release: sere ietee alae ole Mie aCo nT 83
simttlans. REEVE) (area IG. 6 occa xcwiels ic io ieibrete ws een Niele aoe tea 39
SHMMANS Reeve, -EsISDEV sand) GCOOKe. J ..stchis crecsies serene ninten ee raere 30, 47, 49
Simtllanspvated) atea 7 yee emer cee Sans heite tere CaM eastatdic GES AC tree ea 40
simulans var. 2: ated 19; IOAK occ cec cee eater a aetne sheeted etna eee ae 41
simulator Pilsbry and Cooke.......... Steg cas ais [ovaeisia ares otewarere Cesta Set
simulator var. I, area 4..... SYA naar aavevere clavenstarata sheave etal icra oa eam 34
SOLdiday. ING WCOMD: i eile cute « sciste.scelsie be oie ees mipveretovele tee nit |store coe ney ae teee «eros
Steet Welles. aren 80 eis ioa's vase woe win, ore ota aontegerata See vos oe eae ER Aa 143
steelnivarel, area JO sew anconetoccta ise cite reas Serene st dre wire eee 145
suturarusca: Welch. area SOs. weve cees «cmetioweneee oe aes rare acohdte renee ee 148
SULUITALUSCA Vaio I, area G55 pi sleet cae aistsiaic lata s Be aiaue eee Ectarsieys sietatones act epete 149
suturafusca var. 2, area 89......... Sdetac Ne wots Mulla Ove GES LED Clee ee een . 149
Suturalbay Welch areaiQOsssceoseie einer e cee cs REE ee LAK ak ae eee ISI
suturalba: var. 1, AreacQhiel ss oc0.dsscd sacks peramy es 42 ae aA oa serosa 152
Seiji INewconibes. «cree cseeneeesewoes SRI hare a SeseiC a era Otao ekereere aKQe 68, 70
swiftiti Newcomb, Pilsbry and Cooke........ ee 68, 87, 91, 98, 100, 107,
108, 126, 128, 130, 131, 132, 135, 136
tuperans: Gulick= sia. os cele sole cemeteries ideo eet hee gana everoin bieteia aS 107
{uberags vans Tsclch.c dca cates Soa e eee eRe een eet 108
fuberans.vat- 2area GO... «2 scl serie sraasirckels Sd NOG ey eR niche too SEC eG 109
tuberans: vate srarea 77 tin ccc ce nctee ten te en ete SR APPA cy ke Ser ete ae III
tuberans: vari AyareavZGs 5. cdicien wtaews'ss sae reas « Sate eet en ene eh eee 112
tODErANS: Vals Saiseicsowe ese seas ml avcrstecapAraebchale oa. etansane test alaeeetar anal eee 1T3
“tuberans-tumefacta” .......... iardislohaTal ale ble eer SIN eee eee OED eee 137
turgida Newcomb, area 35........ wi ev eyeh ny otiet ake oke felonete eke ceria oiste eeo eee eee 68
Walia Plesiier so.5) tiki ccnsia) orererere tole pacae eletare eraibee nie io ee ee ee 21, “22
versicolor Gullick sic venice nk vase Drala fe ii acere eke cate eRe nee eee 130
versicolor var. I, area 78?..... stesscuietevs Sis ire erore ehetve eae See Nel GES ORE Se 131
yespertina Baldwin; area OS? .....%<0ceaseacpase a AccafSisjeost es apes ws ape dovera eth eG « te
virgatifulva Welch, area 73...... Sonu Siaiaicio ore torse ocarshei che nie ete eter ee OERIOR era 103
Vit@atilithva: Vatovyt, ateay7 51s. oa osicalactisaanwe cee heen mein tes i aarestete is ie Od
virgatifulva var. 2, area 74..... Sausremias ate she wraccune Guia eioteh teow els Pom deeeiee 106
Vittata Reeve; abea-I2...cshaececcesee alin! Sata torecehe Sie etsionele Sai eroreareie tere Sheter PAS
viltata Reeve, Pilsbry and Gaake SA eee Sisralstete a Gave inetiiie etre .-30, 44, 45
Viktata Wars ALOR, Gon tei nccctee oe weet sc evs S.Gs ove wale aT hie'».cabhai ielione, esStRee rape eat amet
Vittatal val. 2 aLedl Queenie Aetad otote wah Set a late ener lated a eiatatay aero ee ees 45
wahiawa Welch ....:. Ga Jara lets Sivas Sinn ee here ala otare va ih x GRRE are sfyre shew eh WEES
waialaeensis Welch, area 2, 3........ a eetetdieteraietert are insane deters prereset ora Perce eee
wailelensis Welch, area 117...... o laiteaiee tae aie hetistei ae sievotereve siete User ee oe
waimaluensis Welch, area 32...... pede ere lennecae ohigeee mates oe Mises OG
waimaluensis var. I; area $2A.. 00. ee ce BSS teceve [Sie eisalovnae avanele eter eh Senators 67
INDEX TO PLACE NAMES
(An asterisk after the number of the page indicates that the name is on a
map on that page. On figure 2, page 4, all place names are referred to by a
number. Reference to page 4 will be omitted from this index since reference
to figure 2 is given. A few place names found on pages 5-9 which are sub-
divisions to certain streams not used in the body of the paper are not included
in the index.)
MeeetremGatcia(iigs 2.15. SORE. B) (ide cicvs cae cacielsse eects ee Ad Uses 8
ETOn it OTStmce: Gilicks oc cjl cise o aeiesersers sie ererels Zh op. TO Oy Vis ia, lWetse
120) ZO 130. T34y 30.0375 13S
EMUNITITTIcATIeS tearm (es) 2b WTA Bi) eee racetetpeiereiecteresaie ciate icrarsiete cies cictceieters 9
Beers II Clee cherie crn So niclan ci retste icles la ete eT a cee Ree Uren ae 54, 62
MECN TE CRG, Se SESS UES IT Vocus, 4s: ci oor aieard are wie a eye ere desta wareim dard oie MiseIR ey 6, 53"
Peeamsmudistrict (ne: 2 BT). cc. gsc. cs desc a we ees sack 6, 59, 62, 194*, 195*
Peememeenetrite (i) 20 TD Al) eee sg asus it a era ayes mtcceke a OS es oe eee 8
EMIMOTMES Cheariin (ior ain Oar emai yn cuts (icree etree. WLU It Miah yi An arian Mansy cal 9
Seen TE SURE NT CTs. Bas boris op he eclan dat eicleiaate ois ems oie altel Geek SOLs 8, 185*
MMOS EEE IO uD, Es) ote Alas oo ciaisitiec toe siete oem Meas mistelen Meee keels be 675
EMER em tle ose ACNE POS RIS a cel oa hes whet owed gale yeas haga eam OA 6°45
PRN AMET eT SD) tr 28 tare gat coche rs eprieo tase ave face akwobeke IRI D. hy WER S IE 6
mri low tigs Oo A. LO Gyo Ay Aienn acmedags se cheniee aa timsnsayed aii 5
eM euma Chie. Vek WEE, oe oe Na Fay aes ae oe ee hy os 9
RR eMMN EE SELeat tC fire 2 ON TGS ID )ix eer tea teaccune oder Noida ey domed vale eee avn te 9
mein sepcistrict (fie. 2) Fy DE) sci sic wie ssa siete a oy9'e esse awleuninleeneiaeielane the 9
BMT Act HIT RIDE) cad close vars is wicicrs ore FRE ev ates Se wed hd BOLO aS secs SR 59
Pe udeleicee.. Northi-Centralis.. o/5.<2 sorccle-a cutee 21a. ese,orsale, walla aid # vinsascyviays 57, 58
ERT GE EDN OEE N= OUI. 5/575 5:0 5.c.0's ibe are 5 ore eed wi egebuahe- kia 6 we arapa geste eee Rs 59
Halawa Stream, Central, or Central Halawa (fig. 2, B, I, 2)........6, 53*, 58
MerEaEunS Tea NOL thie( irr. 2) 15, Ty 3 ys scale orev oyasaye is eas aheveval ona Srouwrar tial a vente Oa53F
lawamS trea South: Chew 2. Boule wla)ra ses fae dosrs crs cio in ele ag omer ae 6, 53*
Soetbrm ae SeDCISErict. (fie. 02h By D)is. ik wees sige alelne ons-o% elvatnn 6, 194*, 195*, 200
SemnemEsreai(tien sD. \i cs crs, Seip aewbicin ele ete 0.6 Mavens Ws Reo nae 6, 53*
Sema MMR UN CAAT TUTE ERAGE NS Voces cvs ov nfele sce eiateie A, oS aia ahe elebansts tape te Meee 11S AG
semceeantiaerrye GITEL) (ie OE WD) acs.) o) uisrars Hehchels. 6 bole ueeine he aeineuieds 8
menace Sundistrice Cha: 2. Ee VLIT iit ouk vcleurtadee am saicibicolsokinete ake aaa 8
marae eSUnICE (tips 2) Ee. OX LV lecciaerecicic oe Mialela S ocorssera. coined Soeaume urmedwiee 8
FESTA I PRM se ests a sean as ie Fre AM ae) oe ES Re 188, 189
Rete Subaisenict (id 12, 0h, VIDE) caters to Sacesieccaws yee pa dace cee cs 9, 53*
Relbee AaR II NOUN eres ef oa Sealer eile a a4 So! hath ieiat etree kigearns BED, TOG Ler e TS eae
Ree EETANO “WON DACTIAan ICP E: oa) sca. cic nessa aoe ad debe ah sails acs mawTAu Ute Loe Ow ln
Peeemanoiciage,. Notth=Sotiths <6. 6.5 gs sels ces ow ose NS3e T5OmS 7a LOO, 172
Peteniano, stream (fe. 2, DE, 1) sits eaceics cose seca cu 728027) 1OAty OS®
faeiemanorstream, North. (he. 2 C, IU; tb)... oes. k kee. ges ce eas Fae 2O uO st
220
230 INDEX TO PLACE NAMES
Helemano Stream; South (fig. 2; C, I, ta).....2...026.-s 7, 105", 152) 196; 65
Hallerbrand's:'Glen (fies 2A, EB) dens cose oak eat ee 6, 42
Ping Gulely (hig. 22:B, Moe). od aoe soe be ote ue cee ee 8, 185*
Honolula: District (ig, 2, A) .is sse-decne ye otc cane eee ce oa ee 5
Honolulu United States Weather Bureau Station (fig. 1).............. ato
Hoolapa ‘Guich (fie. 2, By 1X, B)o<ohe lee cae a eee 8, 185*
Hutuela--Guich (fie: 2, 1),-T, 33). conc ed not Stake ee eee ae ee 8, 185*
Tasihs:Goleh’ (Cie: 2) EP GD, 16) o.s cs See tonto te ae a 8, 185%
Thith-K absent deid re) 5c die saa. eas cae ae kao ee 187
Tolikaa Valley: \(fig.02, 58, VEIT, Dic. udavak finan ee eee eee 9
Kasawa Streata \(iig>2) We» SeXs 2). f cde tree ec e s ooh ee eee 9
Kanaawa subdistciet (fig. 2), XO) Jess acdc cas Ges cee oe ee eee 9
Kaalaea Guich, Kawailoa (hig 1D) 55°10)... .005 4 een a obec ee eee 8
Kaalaea ‘Stream, -Kaalaen (he: '2- By Nee 1) 22 cca ts es ee eee ee 9
Kaalaea ‘Subdistrict Gigi?) Fy). usc) chee cee en oe co ee 0
Kaalakei Valley (fig. 2) Ao 1:9) ah. eee acne eee wee eee ee 5
Kaaoao' Gulely Gig. 2: Gy) ane ie. tadae cas bocca Ocoee hee eee 8
Kanal’ Stream ‘(ae 2, Rev Li a).0. o. ic hate oo ee ee ee 9
Kahalou Subdistrict (fie. 2) 7F, WD) ~..cssen cs dceks cae eee eee 9, 53*
Kahana Stream (fie;-3) Ie, SCE 9B) saiise dd eaictes cewek event aoe eee 9
Kahana Subdistrict” (figs 2) JB cI ics oc gee ae caace oaoe ee are Are Oe 9, 105*
Kahanaila\ Stream, Katlaa ((fie.-2.0F, XT iw. annie. sosces «mae eee 9
Kahauiki Stream (figs 2) Ay SOI Vs) ack ncuron cee tale oe See ee 6, 53*
Kahanika(Sabdistrict <(igs.2) Ay Ooi) eae ccce sues cele ucee oes 6, 16, 20*,5 58
Kahawainun Gulch (feo GUE a) .c akon e as dena cee ee 8, 11, 185*
IcahiglenSubdisteret (tie: 2, Feel is. so oaciace ote ee ee eee 8
Katlua Soebdistriet (here. bs Xo) s.r can weiner nettas See ee ee ee 9, 29*
IKaipapaw' Guich- (fies 2; "By XDD Tas a0. woes ens austere Ao rae eran 8
Kaipapan-Sabidistrict(igs2) bos PU) sas ts se scegs.aie etter ae 8, 105*, 185*
KalacokahiparGulehs(hg. 2,5, IX,)2)+.e 2b eececeut ees once meee eee 8
Kialaikoa District, (Gulieksi.4 cc aceuen ase ae 2) 7 -A1O 135.02) LOS IOS mer
128, 120, 130) 13h 134) 145s 130) 137 emis
Iealama: Valley (hes2 AT, S)iccscscck waesteos cee pana crein Dele | aera 5
Walawuao=Hang ike siRidge: a... vols sch Sees Se aes Sete esa Te ne a eee 61, 62, 64
Kalauao Stream “(ties 2: sBy DED yi cc oe oe soa eae eee a ee oe 6, 53*
Kalaune, Subdistrict + (ag 2) BUST) < anc, ociewniees heres wee 3, 6, 194*, 195*;, 200
Walauao=-W dialee Ridge. cassis ase Foie cd cdi » oo ecis se ae 66
Kaleleiks Stream (he. 2.0 IN a) 5 ask vasesc coe se ne ee a ee 8
Kalsha-Kahttiker Wid Ge: oad icloee oan Sa pha Mie a Nistor d Cte ea ote a 51
Kahhi' Stream: (ie. 2; (A> OU 2) i:2 bs ae. Se cee tec oe ciate pee eee 6
Kalihi Subdistrict, or Kalihi Valley (fig. 2, A, XIII).....6, 16, 29*, 35, 36,
37, 38, 39, 40, 48, 51, 194*, 195*, 200
Kalvahole Gulch (figt2; D, Tp 2h )iek ests «depen ces caer ce at eee 8
Kalwakauila: Gulch) (fig. 2) Ell 2) ne nicg cee ne see ee eee 8, 185*
Kaluanui Stream (fig. '2,\B; SOV IL, acc. danate can come eae sea eee 9
Kalnanit. Sabdistriet (Cig. so: Ey 2S VED) ire cones bose een ete 9, 105*
Kaluapo; Guleli (figs 72, TD) Ty Gin crece sig ceeute taco teats oy 8
INDEX TO PLACE NAMES ail
Realinawalkaalanstheani (ties ach Ts Tejas caters cisteteievciete =) spots nccisiavoeies 8
Beenie Ki Cab MERIC OE) soya e cy alone) sieves erevenateis fo Rasico Meuse she meal aerate ok 38, 40, 48
euchiaikig Streamate) 2, At MOI: Tye. a Urars ineuseonsse cee a onele ed siete o Aone the 6, 40
eriamamtiteSEreanie (ier a2) Msn oly Biles foamy aa er siatvene sm &, Hicleveue crane etomictn steieeanne 8
Mee Ure tame Ed LL Waly; Pye oe er eerie scvereverne rinses eae nectar oalsenicele eu cre Sabin ries 188
erin otis leyn Chime 2p Alec): 15 cts costs eketnete ohn re are nets eo ouel ale ae ah E
HeannOnuimevalleya (hier 2 NG MAG ink nici: dec cies citexcisa meets Revers ard Wiles stad ayers: 5
ESTE ear Phe oj ic sich a's ok ale hapa ee Sane eid Suet alee ahyaed Ss Smokes 174, 177; 170
BETO OAMiMS treat ies Seb: TEX cS) Me. © aes cha coe rnie aySar cbs eiacaeace se nietavaisio rote tert 9
Srey Enea Tilia Olea ota Avast ENG eH) - hspotersesnes overe io siciecatcacterorts ayahons avciteleseneys eucaieis 5
BetealOlem@Sereaitin (11s 20) At ol Xow) ar a eier crane caste efeune eeere oc omic nie e 5
Re MCOMEMS CAttOI (ul Ortsl)) aise traeyarerien clo ce secre ehorgeate cud aco iene enn es Si elO
emeOneRStccainn (lotro, Hi aulONe DT ern « ciciscois alorstsre tore wieloe ae a cmneleie sls bievers o tole 9
Berale Supaistrice (he, 2. NB TX) 5s close cee dsc teaihawcderesaie eet 9, 29*, 53°
Bernas bdisthict: (hes i2s Bs OVID) ..caccie aniersisetsie og be asia eas sate isiouclaclealee 9
agelkalol (Gieln = Wweareieve 1st (Gnilente gneoccosebnoaconncenncsodsenbouor 5
Se OTS EL Carmi Ose 2 ANG RENCI Un 2) savicycyssctas ee ha caeinieicllsimcr ends oot ismer ae 61) 247
ianalamarsupdistrict (ie: 2, Ay XML); oc. qant vse es 6, 47, 48, 104*, 195*
Heme LNA Lest TA COL) (1A p= EC AUT OTANI LAA toys ain ese favev el asad ele scar atcdonovee we lacs) ote iaoieiccsaetiehae a rstararetes oe m5
ev MapiiIaMr eels ace aye teictet sien eos Te tae aren pa Acie I4I, 160, 161, 162
Beiaiconali ass unite District «serie sms cre cers oreo lenis cera aiaiail Setar aieemieeries = 145
Kaukonahua-Poamoho Ridge, North.................. 138, 140, 162, 163, 166
Kaukonahua Ridge, North-South.......... Pay With, WAR, Ue TIy TID aeleh Tesi, O'S)
Kaukonahua, South-Central Branch Ridge, South................ 102) 112103
Kaukonahua Stream, Central Branch, South (fig. 2, C, I, tb).......... 7, 105*
Kaukonahua Stream, North, or North Kaukonahua (fig. 2, C, I, 2)...7, 27,
Bah, HORA, UO, MSY, WSIS, Ikan ile,
148, 160, 161, 162, 194*, 195*, 198, 203
Kaukonahua Stream, North Branch, South (fig. 2, C, I, 1c)............ 7, 105*
Kaukonahua Stream, South (fig. 2, C, I, 1)....7, 22, 25, 26, 102, 105*, 106,
HOS la el 7A One ane OMNIA mL OAS nO St
ankonahtia Stream, South Branch, South (fie. 2, ©, I ta)...........- 7, 105*
arr mcet eta imeraee Ey AT Vint ine See tre con, Vai nid galalenale seh a a Adve vis sae sho 8
Genie OOM Gries ao elena as cise orate Aa eI eet teincayeie cha veksieisters 186
Perino aS UDCisttctm (ioaeonn EV) eer cinereus o deities ah einittlar spouses 8, 185%
Pemreer ese StneAialiag iim.) sel \ia Fe 521. o Beka ashe: sie wl Onde od. oe Sates on 8
Grn Seca ILI met tel Inca ticom (iT alt eNO LENG 12) ene topericiiece ee ci peraie easy steers ey aeie a eters 9
AE SuGedtit pINATE ONE! (il@e 2.7 Ela loX AN) anaes aheteebsicieyeo ete toe olsteretole isle 9
See EME LER ELE IITNG Ie cian Dt )) acc yas eotyeie te aint oi) ae Seder hl Cie Wissatim ae sli 8, 185*
ianailiallomney (Greilelm (Gites, 2 (CG) ly 2) goscocuoonouemosss nosey 7h UOSs TSG, alsto
HeawalicalonasO) pact aeinid Soma mre sete. cman he tlereloseere octal Meme asa oon tee areal LS aLso
awake Gulch or Kawai Valley (ic, 2) Dil 3)id.0. 25.5050 45- Fi, MOSS, yp
ercclite cy ATT Ute d Sey arnt ce nie ween termes bya fake’. c. hbsiepabaths hice eaiiers 159
HEMT bests ted OITA (eT rare Te) Raeeke chavs oe hss cle ohclcacoroueoe eccsetc arenas & aoe eee Beto
See aac Ole MAMEEM)) 5, shia oyn cls spel aNetaiy wei ae ele le ane Qe vone asieghesh asesaemoes 8
Esenveii ocin Gil chin (tiga 25 es ollai2))iavere s:oyeis clerste cakelacdvere eles 7, 194*, 195*, 198, 201
Ree ALS Pe EUULGN (AAAATO) Pcs ces sc do pated Sie eed aie Bia Gas So ae note dee 3-10
iawailoa Subdistrict, or Kawailoa (fg. 2, D,,1)....3, 7, 11, 170) 171, 174, 179
Kawainui Gulch, or Kawainui (fig. 2, D, I, 4)..7, 11, 27, 105*, 185*, 194*, 195*
Reem ep ACHNC Haagen Eyl WV) Ride arc Seeds, £E apes eeaixca tla whee oa es 8
>
32 INDEX TO PLACE NAMES
Kawaipapa Gulch, Kawarlea.:Cfir. 2, Dy 1<8))..\ie ad ce eronieeenern eee 9
Kawaipi Stream (fig. 2, EB, TED, 3)... cnc acces on do ees cates se ee nae 8
Kawela; Guich! (hes 23 By VTE a) eric tae erste dolstevele hice ate eae nae eee 8, 185*
Keaahala (Stream) ((fig.. 2, Be VTS 3) ecyre certnrncterstaet ceil tastier etter aes 9
Keaaula Guleb Cig, 2.55, 36,20) as sastucae oes anna SP es oes oe 8, 185*
Keahupuolo Stream-South Watawa Stream. <i oc cies tee ves a seca 6
Kealakipapa Valley. Gigs 2; Asus 1) his an fer ctoele ic voce eter ebels steed erence 5
Keammea ‘Gulch (fir. 2: DT, 8) Siccdscn cide ee eis wie ev ng ne eee eee 8
Keana Subdistrict’ (fig. 2; Ek) kc dats Boleidis ercteteverscorpiarajstcte ns mais ae ieee 8
Kinapas Gillchin(ipe 2p by les mel) eet eets 3;.7, 10, Ii, 12, 25,92, To4*, 195% Toe
Kapapa, Gulch Gentralin(ign.2 9 le Tb) irarsererersist-telnieratsiere nie eters 7. 53%) Toss
Kipapa Gulch, first North Branch, North (fig. 2, B, IX, td)........ 7, 103, 100
Kipapa Gulch, second North Branch, North (fig. 2, B, IX, Ic)........ 7, 105*
Kapapa:Guich South" (digs 25. b eXs Tal) aca tacks erate ioe eeistereee eee ele 7
Kipapay Ridge; North=Gentiallin..c+:scuien recom oe eho eenieee II5, 119
Kipapa-\Walkakalatta «Rad ge cies e/s aeleyetinteleteistereie s)erevelierereietele le 103, 106, I2I, 122
Kokololio Galch Chg. 2 “BOTT. 5) oe nciee shave aie orectateee ener clasieyeieete ore 8
Ito} Coral CibdCe8 ol (GO evemee tal OL GN DoD erin maaan cco amo abomacmmiccdoon. 8, 185*
Kolokint Stream( (fig: 2; TE; p23 ))ies sachets sepsis nierate mec kere aeieie echt 8, 185*
Kooldu Ranger. seem Pei A Oy AKO Wie tebe), Ae, Ay, Ge). Gis
185, 189, 192, 194*, 195*, 196, 199, 200, 205
KoolauloasDistrict (hig t2' FE) eis ahlentenin caste sereelcie ise SORT eee eee 8
Koolaupoko Districth (essa. ul )ictc ce c/cieecrecsiete eretete sie erento nial aerate 9
Kualoas Subdistrict (howto. Raw iin ae ncrrea ctocticrters caterctona cree rte eee icons 9
Kukaiohiks ‘Gulch. (he: 2 Des 9) 2 aa. co cate etree oe ee eon oe eee 8
Kulouous Gulch (fig 2A si ies tar everereterene tehcveioneele cre ueheiie oie eee tae Ses Ten
Kaloo SUDGISthiCh (diesen RAY wll) eerie tere\ereie micro cee entree 5, 194*, 195*
KeuluiiGulchis (hig 2: AG Viste) ic tererperavesars.ccs si avonevertiere ev aetuon ayecoven mle atone eter 5
Kulii-aulaupoe. Ridge:4 i242. wacociaccios ont aie ete ee ee eee 31
aie Subdistrict: (hee 2 Boel D ister cieeve ier siotien tec eee a eee 8, 105*
Bamaloa. Guleh iGiig. 24, Fax, “1a ives b toarcie s screstretneis tran ee aerate 8, 185*
LaulaupoesGuleh= (fig::2) A. V5 2) ie accis cterrccvae sere cceie siete ere heim tenets 5
Laulaupoe=Weailttipe Ridge sca c.screscstacs svatave secre aire ctale stare (ohake etevaten tear terete 31
Luakaha (Gower): Station (Gig. 0)’ ns. hs aes eeaatee ee aoe ie See BLO
BuakahanGuppem)r station (i1e-0 WW) iaiecn neice sie iets 3*, 10, 200, 202
apo ==: Waluapove sya strain ernie area eee ataecie a eerie aera Neste oravenaio gts 2
LulukuyStream! (Cig. 25) FIX 2) is osaretecitersiciwlcrsicroleeiecs ois cris eratorstareneereteeiee 9
Ealumahu Valley (Gtig.)2) Ay SXEo <3 acacia televise crecesareictatocierep nye eee ict eererene 6
Maakua ‘Gulch c(fig.t2, Ei, X1EVy 4)tc sana sires ches cn vats s cee alee deters 8
Main ‘Range = Koolan ‘Range yi. \jeulssinies news ais sae seg sé sleet 22
Makao' Subdistrict” (hiwices By XV) cect ctistare orate Os ce ties ene aie eee 8
Miaka parttt s cievcvelssc-crebipiai aheiaice uel atere 'e a tort onse form vore Siticsenuatetarcis cycle ei ciate aie renee ane 194*, 195*
Makaput Station: (figs or) ac sesemereae aise eetomierstee al ietecierster 3* 10} I1, 200) 202
Makawao:Stream: (hig 2:0, 050) acacia scciereistaiene siotete crc ereter mementos erent 9
Makikt: SubdistrictGiig- 2: Ase 1X) ee oes ctecreertneiee silo eio ior eaep ener 5, 20*
Malaekahana-Kaluakanilay Feidger sence testcrerseiteraie ere vate ents ius ee teete 27 IGS
Malaekahana-Lare® Ridoeoiis caiciee ciefe sees. eleva s ies eustateverareeiee choi el agate seteay catch 175
eS ar Ee Pee ae eet
ME ee. ee
OD ae ee re
INDEX TO PLACE NAMES ane
Meera itty Steam (aie, re Fes DL, 1S) ore cards. 6, sy0/suchs:'0.6.ciw wissaiove's Gsleis ic aie 6°) 8
Malaekahana Stream, North (fig. 2, E, XI, 3a)...........scccecceees 8, 185*
aiaekanana Stream, South (fig. 2, E, XI, 3b)... 6.00.0. .00csceen ans 8, 185*
Merereiana Suidistrict. (igs 2, Be MY) i. a5 se Sie etlcse tba teen ea ee ss 8
Serta item Chie. 2 AW OSV, I )is.cjacs cele cane of icine cn 6, 29*, 53*, 194%, 105*
DONE ee srag RASA CANT UIs RTC OO as Se coeterirel peices fre enone ekskavavey cit ob. Shave totais onoloi elle ofa age Ade eas2
etre Wo Ps. VIBE) a Scia tiem 'e saleveeioa a desing eee-t a cietaie aratelen Gust
Sea atistthict "(hie DS 1s,, VEL isi. ccs cx nes esate eas omens 6, 194%, 195*
Srinana-VWalawa RIGZE o. 2.6. ce einen eee 77, 78, 81, 83, 85, 87, 88, 190
me MER CeeT et OAS WV LETT) isa cc ciao. 0'e a cloja'asnie din aeein, oho aid wayo\ ayes oa alate 5
Peremennetin ast teem) AG VLE Dic ooo cise o's cies e mals 66.66 slain mame elnla a's 5
menon suudistrct (fig. 2, A, VIIT).....ccnceescsncsaas 5, 20*, 194*, 195*, 202
MN ETET EAC IG PNG) NCO Noo acceiniaho)<-oha iain e's 1¢,msisinje sae oe witha cies) o'r ale s/he 6
Neat MRT Pe Pet cay Stree to ets as'stels she's) ste ota aii coy vit1 = fesexeta/SiQPeVS are wiate yee oe aLealeha\e 38
nt aLne SPE eatia (CE. 2, Ay LM, 5) jy a tsisin i cw tia oe wore diate nies Dale« Baie ein © ae 5
MMR AES TISCEICE. CIE: 2) A Lida a ats wes wiolnie tienes meee 6 ae ose es sie Be 5
Rea SC AETENY (FGF. T= uv, Sie ei dis'nie ais ei sib alnve icin died oa 2 ays iso bivuels sels 3°) 10
MOET AWTS ERE alii (pest 2 kev os A) la\kcsi over «:histereiaicyercisi are ea etave alelaccierais!<(nie laieiers 9
RMT REELS fim -P NG be 2). rosea Ss ainteieiaraie w stare'e 32s Mieicinie eielats) armies on 8% 5
Me ME etl het RIE epee oie em oie 0 arose iss isl oin a laim les eiela oie ne Atel miwrsra stale 60
Moanalua Stream (fig. 2, A, XV, 2))..........-. 6, 26, 20*, 41, 53*, 194*, 105*
eaiinusinpaistrict (fie. 2) A; OV). .2.-sieesccccssducaotiecd sce 6, 16, 51
a eet NET TITEL 2 At LOK 2) fof s'e a ws chara) alae slerd, sieve oi6ls © oie ai afeie! evslole inva o/ip 5
MONG SCc1 PO Passes ee ona oe w otegares eis Sel ole sisi ave) aie cists sioisiahnicisrsustote-<iersjetenseveie 189, 205
iianmiabiatrarstheam (hips 2) E154) 2 sic.c)< cise ctacte cies vis ale) ateis sie ciatere icles ayes 6-2 8, 185*
Carine Lema Se tte oO AC UTT LT) Bo teres as crciece o/accis'as, aaa s)slei@ictavs.ayeisiae/ee's,oi5 6 5
Rerun ests ig. SN TEI 2). 5c vat ceo vinjae. sles isct acer aroun asee bole 5
mrmmrreersteset. (fe 2. A; TED)! fc:ci'alo'e cleo sheng s 4 ae-sisl ey eisicje a's 5, 194*, 195*
SRN MIMGEVIIORE! 10.) c122).. 5-0 .- woo inlea soe a dle pigiain oe clo eaerelcticwe sce ses LiaesT
rr betewarestreatns (igs 25 Aer Mobi. ot) /o:.rslchereze aveioy stovsieve cle evs sicicisis\ciele\scielele oe 6
MBE ALI— Can alamaRid@el . siccs eicters « (cveicis actetois co's sis leevele's ortiaicveferse'e ehalels «ies 43, 50
meer ti eG pe NT AD) dia cian a Se'ss a crates, cyeteb nie ate 6, II, 194*, 195*
SERIE ALE StAtiOn (oT) esacie a) cats wiaielvig aoa dS eee 'c wie'e cic ae cicle vere gk “YO
Ranariatate Stearn, (fie sce, As NL, oA) cinistere stereve a teileta) cies "a; otelm 6 0)s) abs le'eioccigyele sate « 6
Nuuanu Subdistrict, or Nuuanu Valley (fig. 2, A, XI)..6, 11, 12, 22, 29%,
35, 36, 38, 39, 42, 43, 45, 46, 193, 194*, 195*, 200, 201, 202
BERLIN TEASERS PEA eis crstchor ato eels edie e aielee wleee seus Teper ese CLOW T2520 MES Guta yoloo
MBICREISTLTILIP ET erase aye ye saya ces aie we ouetels fo sace Sincere, Oa ola eb ts Slate o vellevaveictel els 7
Mebitaeate Gt lchr (for 2s DEX 3) ncs.clotevacua(tieccieine Maas cheisreis 8, 11, 185*, 194*, 195*
Be MEEASE OD TATICH RIGS. or. ns0ic0'o oc <ic'e's six wa cen clade nceces Mena ee sete 184
MSE Festa reyrinee = Blo WP NNT) java 2 fais, een eo 0 clo: ave, Siain\ x) ay tel stacaiesg «tons setae ldvatare 8, 185*
Peemireti Past, (fo 2.0 WILT, 2) ice cis s scot sec sslacmec ioe tecsiews 8, 185*
MOMENI AMS ERCAIING (125. See bN 7)): fe lovaler a sve elevererais aie ©: crsysie. sieve a Neiessieicleuciarove late 9
AER ETC INC GO Fe eC E') w ra cinscisinintelpieigatec’s «el cet alate aieiaiais aleoisieine Eig 9
SEEM IC Essar a Be OT Le T))) c'a chayo.c:o5c,a.ale Wie aie) o:Sseie wie, ik wie se/eisin gale aeelee s 8
Pa IACHE SELECT CLE es Ee ON 9)) 5 sare; ove afelss «eerste arete/sse\ejerere Store steve ibiere eueelensiare: a 9
Opaeula Gulch (fig. 2, D, I, 1)..... 7, 21, 22,105, 177, 178, 179, 104*, 195*, 198
BINACiaa Na walloamRddeean sacc ac qee'se Ge erela cules sajersie scinie ere ersteiena Glerelajere’s 179
234 INDEX TO PLACE NAMES
QOpaeula Station! (Gig. 1) suc cok os cctrgumile ttre cata ee eon eee Cis 1G
OpanarSubdistrict (fir. 2) Bs AVL) cel ies seeoe oe eenctea enka eee eee 8
Paala Ukay Subdistrict or eadla (igs 2G ms) eerie reenter Be ole
Pahipahialia Gulch (fies2, ES VEC). t csc bee ae eee ee See ee 8, 185*
Pahipabialua-Oio Ride@e: ss jc/ie evens w cieseye a give rsa eens we etnias ee ee 186
Pahipahialua Subdistrict (fig. 2, E, VI)...... atin sie eisteteraela sae ee 8
Pakulena; Stream (figw2 EU 2)caAce eee ee oe ee eee 8
Palapus Stream Gig.i2, Be Xo) es tec cere chine taht cio ree eee 9
Palolo Subdistrict, or Palolo Valley (fig. 2, A, VII)....5, 15, 22, 20*, 32,
33, 34, 35, 194*, 195*, 200
6
Panakayaht'Guleh (hig: 2, By VILL 2))52 sa. «taolcc pee ate | coe een
Bantha Kea Grrl eli (hte e2) es Vac) te eee ee ene ee 6, 53*, 92, 105*
PanthakeasKipapa (Ridge). 27.ctee a @anw ce tn seem ae esc enews tee 99, IOI
PapaliiGuleh Gio.) TVG) Seer sens ccckoie sition oe ote cece eee 8
PaumalusKaunala Ridge. s a-acscce arene cnouet os cron tee Le eee 183
PatimaludStream: (figs 24 Vet) ei fk care Sores care ela be heaner eae ee 8
Paumalie Subdistrict Cio.: 2) Dae UU) iss yeserremievser ein cect eee 8, 185
Panoa=Nuuang! Ridges tos. de dee 6 ca euin so ecg Pe ees ee 44
Pauda. Subdistrict: \(iig 20 As xa) 2 os ie arene crate alsin a ener 5, 194*, 195*
Patoa, VialleyaGhgs2) As Xe D)inc.ee 2 tae see eee ee eee Sao
PO amos ei aiisis staid cin, attepestuetete vofecac ens tors racine eit ara Ce ee ee ee ee eee 162
Poamoho, Central, Pilsbry and Cooke = North Poamoho Stream?....... Dit Ser
Poamoho-Central Poamoho North Branch Ridge, Central........ 140, 147, 169
PoamahosHelemano hkidseaeeiemn see eee 153, 164, 167, 169, 171
Poamoho, Main, Pilsbry and Cooke = Central Poamoho Stream........ DA eT,
PoamohowRidge: North-Central e-ice teeee cee oe ete nee oer 171
BoamohorStreamin(dreseoin Gre lenceria eral iret rie ete 7, 12, 27) 1o4*) Tose
Poamoho Stream,’ Central (hig: '2) Col, gb) oo. eee 7, 105*, 140, 148, I5I, 164
Poamoho Stream, North, or North Poamoho (fig. 2, C, I, 3d)..7, 105*, 149, 151
Poamoho Stream, North Branch, Central (fig. 2, C, I, 3c)................ 7
Poamoho: Stream, North Branch, ‘North (fig: 2, (G6). 46) 2. eee 7
Poamoho Stream, South, or South Poamoho (fig. 2, C, I, 3a)...... 7, 105*, 149
Poamoho, West, Pilsbry and Cooke= North Poamoho Stream, North
Bianchi Seine cS ee co oes aE Sie aoe I Ae a ee ee Say
PukeleStream: (figs; sAgl Valle 2) etic nee nee ease ee Ore rae 5
Puna Gulch? (fig. 2:1) EV 86) ce iccaerical cade eset «oats 8
Panalun Stream (fig; 2 EO SONATL or) eee ous du coments ibn eke es ate 9
Panalus Subdistrict, (ig 2,96, SVL). Soe ne ew ate ee esiote ee g, 105*
Punanany Gulch, (ig 2.Bs PVE B)i.n tes eaten ne oceania ie 6, 53%, 56
Pupukea Subdistrict: (fic. /2. UnL)..: oc dec n orn aie aoa 8, 11, 194*, 195*
Poo Kainapyad. (ieso. 1, Laake satis wa caween. aa eee 7, II, 194*, 195*
PudRonahuanan, Ciel Aw X18) os amet or ceny cites kee 6, II, 194*, 195*
Pou. Lanihuls, (Cnet 2; As CoO) on cacicutee actiee fe ee ee eee 6, II, 194*, 195*
PuniOhia— Pantaltse (tes 25 cAy Axes) eercieyarterasciicene eee oer ae 5
Puuti@, Kona: hie, ALM) ocak ew acneek ao ter aac ce eee ieee 5, Il, 104%, 105m
Rook's (Valley: Wiatolani+Streain:.,.c.cscvacen ncttac.cats ate ee ee acer: 6
Society Tskards: picts c.0h ors eesic, «ob iglese eiata¥a etecanye o ce hee tans ret ne ea eee 204
Wn iii Sx emer al etme een vate cl aa aA tai ite Nai lala alii a a acer Rt a iat ane
:
|
pa we -
INDEX TO PLACE NAMES 235
Be MITA Tae apse Pe tera eos des cram Ci crei mise Sead a sheial edo at ar ieidl at's) Svar ¥ og Sis cudvereraicieleiels 189
err — Eis Oar (tase 2) A ES Ge as iii vv alsa le e's’ eleiste deleleia ed dose 5
aber AMRIT CII Oli ae) ihete Ol) ecvaisslatersieiercheharnl a etovereR ere. oa ielares vgn eta cetal: 8
TERING LIE) 2 ET, SEN rT) ja ovs5 0-5 bn vale ce Sw Gin cleroren e's bene acentine pl RW acy
RerthitanuctaMsstrict,. (hisa2) VG) seis vc aa, cilelte aiclevale ei sitio BELO Oey lOc Mie 7
128, 129, 133, 149, 153, 179
See iOem oT neamim (ie i aealivc TV .99))Votmea iche aia ara stcles ee elects o.sitveigvocfas oSismohe 9
Seeehiolems~apaistnict, (hei 2. He TV) vous eos vies ai bie tee sees elise On 53h, 005"
ReeeeGues: 7, 6) —— Walalae’ Nits). 2c. ose d0 nt sinc sce sislcc abs a0 194*, 195*
Waialae, Gulick probably includes Subdistricts Waialae Iki and Waialae
MEE ore 2 a ra UA LS oe oie) a ot hg Sa wae cae ome es E20.) 30) 32.0 ae
See camber Cla Chi g ey 2 Ati eV) OT) oe sim atectsin telestec wate neck ae eo sletoee 5, 28
een emlKi a SUpdistnict, (hg. 25 Ay WV )ic ot te derscicd caceesicsiessd eee cess 5, 20%
ME rcidem Kt VVaialae?-NUl RISES «Sac iacree aie oe sce swe wsceiey ouieiete s erdelate storie 30
eres rae tits Grete ines ee A Wile 1) sc scresc nove svi save ee ocaroe nib isle arora B28 30
Bee temoii subdistrict (fe. 2, A, VI). i 3: fs ceu cael so oicls ne soe 20 45
Seeonataent Guich = Waidlae Nui Gulch... 0.60.0... 50 c eens cece one se cess 5
Babee rict cement tions (bn ee EAN CTO) IA. os Siarelcie ciecaises: arajnume tatcla eudee 3 waves wie dian eters 8
Peeeernstnmistrien (hie. 2. BV) Soc. eee olicce evs weeeve ccueen 8, 181, 185*
ME eM MAILS Secchi ene ita. sags oo yal Enis 01s Nw Qa RS che veein Bios SA RES 181, 183
Serer Am ISEEICE A (hSe), 2", DD) esr. sta cya ato srchcine nis sceiielen te oe Deis 7, 174, 179
EMCI CRPNVLOIINICAIN Se tania t Moritse sree cis coe ee sie sek ddie 6 Danaea 2, 9, 180, 203
Beene ange — W atanae Mountains... 0.26.06 s<2 coe ccedse seve ccuctes 22
eemaremelicdastpcistricn Chis. 2.05) D's. kis ba oc so o's vieerle ve oils vse ee os 7
MRM Sretiaa UIE) 2. ata UW, Direc catatcs/arels wiles avleiels wm as ba occadng saloeles 9
Serene Ginicha North o(hig:. 2 1B Vy, 2))lc cas cies ad res stoic ols are oe tuts eelse sila 6
Dee IER eet (oes 26. VT) se oicn ce lc Genes cscigas wie odie. eevee ean 6
tae MAAEEN Vi AIMIANIO IGE. cy ciclo elon’ cacec.ec's oad ooe4s Saws duere se 68, 71
Seem mmrMbmicthich? (He.-2. 1: Vit sior sa celds God cb cewwcendisdcees 6; 53F) 56
LE. TE See lal Wop Ee Or
MRI PERE ADR MIRIGIO «2/0 vista se stash cc cinee Ge ue Dass eed bela Go vene FIZ, TiS. 17O
Paiaw. North-South: Branch Ridge, North...............cc000sseeceee 116
eet eee RIPE. b. Sci a)!s aime Sicko datalale\e ole sje s,c/b0e Ces seed wes 94, 97, 98
Berman etic, NOP SOU. o2.0(cs coun ccd aces ccvucecccceewces 85, 89, 116
MmMCM Rett CHE so ES. WEES D).c ok cc cys aw siecle cle nncdewelee cbeuecad 6, 26
WV aiawa: Stream, North (fig. 2, B, VIII, 1b).......... 6, 24, 53*, 96, 97, 105*
Waiawa Stream, North Branch, North (fig. 2, B, VIII, 1d)............ 6, 53*
meaawat stream, South (fig: 2 B, VIIT, ta) .....:...ccccccsceccee 6, 53*, 80
Waiawa Stream, South Branch, North (fig. 2, B, VIII, 1c).............. 6
Waiawa Subdistrict (fig. 2, B, VIII)......... 3, 6, 22, 25, 53*, 68, 85, 8o,
92, 93, 194*, 195*, 108
SMM eM AT EE ME IUD Ech. ce, ays Sats ote aeee. dase en « Deak eda oa ORs we 116
SMES ALS Cee Etc Pee Seo. hash asateiareinseie ade alttre ie aisles a hated hehe, Weaken elk 180
MIE REM CCISIEIOTI ON 21H WT iE) rains cate a: eis o oluialos io Pie's ais oitia be die sce 9
EMRE SIS EIU ffigee, 1, Eos t ML) oka coca syale-a ecb stain cl toh aiee adie ealiical fe 9, 53*
Waikakalaua-South Kaukonahua Ridge.................... 104, 106, 109, 123
Waikakalaua Stream, or Waikakalaua (fig. 2, B, IX, 2)....7, 92, 102, 103,
105*, 106, 194*, 195*
230 INDEX TO PLACE NAMES
Waikane ‘Stream: (fig: 2°F; Wi, Bs. Dic ticnteoes eh 5 wee ee oe eeerenie nto ares 9
Waikane Stream;)South (fio: 2, eR) WITS a) tecteye stele tos steetereeiete ctatemantere . 9
Watkane Subdistrict: (fig? 2, P,. TLDS vc. casas Seas Cee 9, 53*, 105*
Waikeekee: Stream-(fig2 2; JP SUT, 2): Soe nace ata ano meron ee eee 9
Wailele 'Guich® (hig: 2)ySXdL 16) sc wate acetee lon ee emer er Ia 8, 185*, 188
Warlupes Gulch ' (Gigs 2) °As DV3c3)) iis <ietereate eresercareeteaeerctatetaretess eshte eee 5
Wailupe Subdistrict, or Wailupe Valley (fig. 2, A, IV)....5, 15, 28, 20%,
31, 32, 33, 35, 194*, 195*
Waimalu-South Central Waimano Ridge..............ecececeeecvecece G75: 72
Waimalu-South’Waimano: Ridge. 30.057. fs wun os doen aes te ae ee ee 71
Waimalui\Stream (he: 2) Be V pe) eee se eee ro cineicti nian re ene ee 6535
WiaimalusSupdistrict: (irene 2 Vie) nie neers Serene te tame eee 6, 56, 57, 68
Waimano-Manana Ridge arnecideiteckii iat ice 73° 75, 70) 82) 183) 192
Waimano Ridge, Central Waimano-North Central...................:00e 83
Waimano Ridge, North Central Waimano-North..................000- 7. Og
Waimano Ridge, South Central Waimano-Central.....................+- 67
Waimano Ridge, South Waimano-South Central................:0..000- 70
Waimano Stream, or Waimano Valley (fig. 2, B, VI, 1)..6, 53*, 54, 71, 73, IO
Wraimano Stream; Central (gee. Bs Vile) ecm seine eine teeta 6, 53*
Waimanon Stream: NorthinGitie; 25 byl, he). eiyerenfeidon ier crate tere 6;.53%
Waimano Stream, North Central (is. 29B eVilevid) ern ote 6, 53*
Waitmano Stream, South (he. 2,-B; VI, ta))cs it pace oe Raieigeigs hRetnes OR 6, 53*
Waimano Stream, southiCentral (Gigs) By) Vil, 1b) emia eee ret etenies 6
Waimanol Subdistrict | (fiecr2) mise) = 6, 22, 68, 73, 75, 194%, 195*, 200
WiarmanoloyStatrone (iam) irre eetetlioe wheel OL Dharma heete, Mee eeeee Br, 10
Waimanolo: Subdistrict: (fig:.2, -F,. XE) et cis ean i ee ee eric nee 9, 20*
Waimea River: (figs 2. Be a) Saieters aad here clone ae io etaeenaee cle ccs aan ey are ee 8
Were selnosuptae (Giem Aalas ID Aneonndodadccs do soddesodcdoeeoand 3, 8, 180
Waiola \Gulch== Waoalay Gullcinicicctom cia ctercteicis oie = eitiente cranan erie eltrerretenateiate 8
Waiolani Peak—at the head of Waiolani Stream on the Nuuanu-Kapalama
RIMES ois aca Gionya sie's Peal tia gm v ovel Ae nien hernia ala tl CUD MGR eter a ee ante ee 47
Wratolant Stream ((hes(2* As exe 7.) raver. rela ss cienaiendieanoaiate cv aiene ree pertains een iG
W aiomaoy Stream: (Gig. 2) As sVilem )iciuarccisystersrsnotete crertitcre ceetor cir ieweee tiers 5; 32) 44
Waipilopilo’ Gulch (Cfizy 2.05 XIV 52) Su te dac: ee asocule sae anes ee ete eee 8
Waipio Gulch, or Waipio Valley = Kipapa Gulch.: 51-25)... cen ees oe 7
Waipio: Sabdistrict Gig. 2, U0) Saran ght tees onitid aks Saag aet 7.225). O2
Waoald Gulch: (fig, 2) DyiT) te) nik Se ete terete dt Se alee eee en 8
Weliweli: ‘Guich = Olona: “Gulehii. SoS Scar ns wines ioe eds cake ee eee 9
eRe E. SNODGRASS _
‘aread or Entomology. and Plant. Gdaminint
(uh U. Ss. perenne of | er Re
Fa) LF is a sy
“ cry OF WASHINGTON
"PUBLISHED BY THE SMITHSONIAN INSTITUTION
| SEPTEMBER: 30, 1942 ,
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 103, NUMBER 2
- THE SKELETO-MUSCULAR MECHANISMS
ODS TONE eB ET
BY
R. E. SNODGRASS
Bureau of Entomology and Plant Quarantine
U. S. Department of Agriculture
(PUBLICATION 3688)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
SEPTEMBER 30, 1942
<r
7
The Lord Battimore Press
BALTIMORE, MD., U. S. A.
Maine SKELETO-MUSCULAR MECHANISMS OF THE
HONEY BEE
By R. E. SNODGRASS
Bureau of Entomology and Plant Quarantine, U. S. Department of Agriculture
CONTENTS
PAGE
Imtroduction ........ GPS TS evel ere see U aie eral ea Nee ace tar lle SHUT tape rele aianete 2
fepithicahiead: and the teedine appatattise cemisas sccm eraseciesisi)sioleie! «eis ais 4
BIMinem reclame yn se occu cae oy ric cock teense epee ve Seat eieharnece chavaraee 7)
Mh ewATLLEMN ACI tc a stetene ciciee ier aialareicue Siotoerer ne roioversdetaerernretemie rors ate 9
MIBENe Bled Gitirtpvsv cts eye cies ermine wie eve lore averscave teva Pete riaic te levevelcveiehelevereaaie eicace 10
SIRTEME DIP a TV AUK ys creketats eehceter coches aoual cucaay ois tevevsie lasena avers el averaloversvorsie eve 10
BINTemrmOUtliene eb eyes rere te eyersioneleteraeynveies aie ieioiavcvole fences tonstotors crete wielews’s 10
MblveM TrialTlClt yl Star. racers paper ier svete CePA ay uceeu syave Poy ex sheove erases Ssasheto dave rehclicisneker ons II
sibliemiaaa scl Aelia: vaste sraratcicratsropsrncieys eveim are is! sloiecshexele evocererevie) sterats 12
Miheslabiimeandelnypophanyarxmeieeie cece eiciioieecce ceticiis certs 18
ihespreoralefoodichantnell ie t.ivsiitaeionaeisksiaite tie ce side biosiae le 27
plbem si chinioa pUtripy ane tees sie esac tve ic chara ele neie ous iale ere wicierars ns eieleronel 30
PEM em DEOL OMAR Maya setts fos vot ahote sec atever es Geeiovel de Gieksroneust Sr Asl shove ale /avelenes's 32
pp eM MHONOCUM Las aisyoeie a ’clerters eiee ce atiatel « oMieraslsials sie «eines cine seis 32
PPM Ee prope CtSats nc ais Aciny~ cere rsyenecsnavcte reves sset eiorare te nie Oltaxsherevevercievee steers 2
Mheaprothoracie, end OStELMUtmnet. clasiiciies celeleiceiclercisile eiokcie elaletoreis 33
si tempi Oth OraCicutnlUSGlesnyrtmitetorerclocrole aiaie siekeversiclenete siecle le sini ahs 35
PRN RSET Ot Olcexs sate eusiey iim cdeyaronarcavevataus lee inuetace’e scste a evniouare ie aiers are chelates 30
PIPTE MH TTIESONOLIIIIY iy cieic tcl a crelade peiayorecok sievele. wid orci aioreicrsists iia oie slater evenaiattens AI
sMliemtMe Catto tlt eresstecrere.cis seosieccievncrale sieve arene orelsiciceig ens ne citer 42
MER HO POCME Uta cersl cyst a syavots crovave yea Suess: sysiovepane Sheke tiesnieeaiw:a'ajs leyepnevelels che 43
iheEpterothonacieapleutas, qactctesite coe vette eres cee c-ccnee eetee se 43
fibesthoraciorspiracles. csi sca. stolen tecieise horace eran a sla aie) alalar stele 46
silies ptecotharacic istertiall “atedsin.. scs,esi< 1s cls nic'sis-e «oie /etdvola Soo a lobe 48
sGheapterothoracic endosternttin= casein ole aia a cea ctoerniere iets 49
TAVere li rem wtttoisteser sean epee sree siny’ ch lois Stu seramete ss encleca ee clavsieisie attains cceiecns 49
PN eETOLE Win om wee iy senate he eae mere a el atone cat ereneaa ete 51
RIB eETINAD OVATE ae bye te ker. Be eres Med Ne RO Le aly) Sra enciepenea Ar 56
Ehemeniplemnitesmars sn ris = Soiree ee ER Oe Cate os 56
sBheamechanisinmonatiehiteecicrret itera terior 57
Hiexion and extension of the wins .css<< «<< siesis sa sees se seme 62
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 103, No. 2
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 1035
PAGE ©
Wins De ees esi isiciw's nin oe: vio viecv aie ey wind Hw Sai re Rie lore pes eee eee 64 —
The fore’ leesinics cic sactbivw’s awelne niente hee eee ee 65
The: middle-legs:. 5 j.ic.ccShoet de eck cet anee ah nae coe eee 66
The hind) dege's s..55 cides tosis ntecyopiecicttte alae tiers eiara oracle MoE iat 66
The coxae and their muscles... 3, 124. snare «ta o> ae cee tee 69
The: coxo-trachariteral joint. ote oe.cse sews 42 oe mete he eee 72
‘The trochantero-femoral joints... senior ite ee einer nie ere 73
The) femoro-tibtal) joint. 5.14). ic 30:5 ostames sae ae eee oe 73
‘The tibiostarsal: joint. isc oes ce eee as catecre cei acto ee erator 75
The tarsal: jounts 1..4.0-5:hciats eh wo ne ole seve be secs eee ee 76
Phe “pretarsis 2:0 sagieeiege eee wale alaiarols werearetesielg een ne 77
VEE“ The:--abdontrial’- petiole: sic cies.cices cae ca senior ceteree aiemiom ee eee 80
Wille Thevabdoinien cis toe sets cae ie are late at er tee 84
General istructure of the-abdomien. <0... occ. +. steowicsemee oe eee 84
herabdominalmmiusculattiness ee aan Coe eects 85
Thennale-genital organi. sissc ss oceeee Laas otses eee eee eee 89
(PNEs StI 53 Fok eek acai doses cae ad neue sales ete enter ee 94
VILL, Annotated Wlist\:or: amuseles) cts 215.0% <mi- anisysetais as rie ter tied ere 100
Abbreviations used jon’ the fieuress... essai sic eres eae 115
References: 5 ches elec deans isa Sa.t cieiers alciee Seek 6 eters oleae 119
INTRODUCTION
An insect is preeminently a miniature machine; its mechanical
elements are the skeleton and the muscles. The skeletal structure,
therefore, cannot be fully understood without observing its correla-
tion with the muscular system and the part it plays in the body
mechanisms. And yet, almost our entire code of insect anatomy has
been built up on a study of the external pattern of the skeleton, done
in about the same way that a casual student of geography studies a
map of the world. The results of our charting of the surface
structure, however, have not been so far wrong as might be sup-
posed, because all the lines that are commonly said to “divide” the
insect exoskeleton into sclerites have a mechanical significance, and
many of them are of fundamental importance in the insect mechanism.
The common mistake is the assumption that the sclerites themselves
are the fundamental elements of the skeletal organization, that the
sclerites have been handed down from a remote ancestor, and that
the primitive sclerite pattern has been somehow altered without
essential changes to fit the varying needs of modern insects.
The diversifications of the insect skeleton are far too extensive to
be reduced to any one formula, but in general it may be said that
sclerites which are defined by grooves, or “sutures,” in a continuously
sclerotized region are subdivisions of the skeleton that are merely
incidental to the formation of internal strengthening ridges, while
NO. 2 HONEY BEE—SNODGRASS 3
sclerites isolated in otherwise membranous areas are themselves func-
tional units. Most sclerites, therefore, instead of being primitive
elements of the exoskeleton, are more probably recent developments
accompanying the evolution of skeletal mechanisms.
With regard to the homology of sclerotized areas the question
often arises as to whether or not an area of sclerotization represents
necessarily a primary morphological area. As, for example, if a so-
called pleural plate that ordinarily is confined to the side of the
thorax is found in some species to be continued down upon the
venter, are we to assume that the original pleural integument has
accompanied the sclerotic extension into the venter at the expense
of the primitive sternal integument, or is it merely that the physio-
logical process of cuticular sclerotization has spread beyond the usual
limits? Until this question can be answered, confusing problems in
nomenclature will always intrude themselves into practical anatomical
descriptions.
A similar question concerns the significance or morphological value
of muscle attachments. Do muscle attachments necessarily imply
homology in the skeletal parts, or can muscles shift their attachments
from one place to another? In answer to this question there is
abundant evidence that the so-called insertion points of muscles
seldom show radical changes, while, on the other hand, the points
of origin may be at widely different places on the skeleton in different
species. Evidence of homology, therefore, is pretty sound when
based on the working end of the muscle, though the work performed
by the muscle may be entirely different in two species by reason of a
change in the position of the muscle or some alteration in the skeletal
mechanism. We do not know, however, whether a muscle arising at
different places in two species has migrated during the evolution of
one species or the other, or whether a muscle, during its formative
stage of development, may evoke an entirely new attachment on the
body wall by some hormonal effect at the point where its end comes
in contact with the epidermis.
The musculature of the head and the feeding organs appears to be
more nearly standardized than that of any other part of the insect,
regardless of the great variation in structure and function of the
mouth parts. The muscles of the thorax, the legs, the abdomen, and
the ovipositor conform fairly well with an underlying plan, but the
number of muscles is far from constant. The musculature of the
male genitalia, however, shows no prevailing pattern and may be
entirely different in different orders of insects. The contrast between
the stability of the mouth part musculature and the variable nature
4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
of the male genital musculature may mean that the external genital
organs of male insects, as compared with the mouth parts, are new
structures having no antecedents going far back in hexapod ancestry.
The wings are examples of new structures with a highly consistent
musculature, but the wing mechanism has been adapted to the thoracic
musculature present before wings were developed.
A study of insect mechanisms, aside from its morphological value,
is important from the standpoint of understanding the functional
activities of insects, and opens a wide field in entomological research
to students who may be mechanically inclined. Insect mechanisms
have little resemblance to man-made machines, for one reason, be-
cause animals cannot have anatomical wheels, and for another, be-
cause the insect cuticle has properties of flexibility and elasticity that
react with muscle tension to produce movements that cannot be
imitated in the usual rigid materials of human workmanship. The
insect machine, therefore, depends on the use of levers and springs
and hydraulic apparatus, but still it is not impossible that inventors
might get a few hints from its study.
I. THE HEAD AND THE FEEDING APPARATUS
The hymenopterous head and the organs of feeding, by comparison
with these parts in such orders as Lepidoptera, Diptera, and
Hemiptera, show relatively little deviation from the generalized plan
of structure in fundamental ways; but, on the other hand, they
present numerous minor specializations and many finely adjusted
mechanical alterations by which they are adapted to methods of feed-
ing and to nonfeeding uses not found in any of the other insects.
The structures directly concerned with the intake of food in the bee
include the labrum, a large preoral epipharyngeal lobe on the under
surface of the clypeus, the mandibles, a maxillo-labial complex, or
proboscis, the mouth, and an internal sucking pump. Intimately as-
sociated with the ingestive organs, moreover, is a salivary ejection
pump, or syringe, for the discharge of the saliva from the salivary
duct. The hypopharynx does not appear as an independent organ in
the bee; its distal part is incorporated into the base of the labium
and its proximal part forms the floor of the mouth.
The natural food materials of the bee are principally pollen and
nectar, both being carried to the hive for storage, where the nectar is
converted into honey. The eating of pollen involves little more than
a use of the mandibles in the primitive manner of biting and chewing
insects, and modification of the mandibles in the bee pertain chiefly
i
NO. 2 HONEY BEE—SNODGRASS 5
to the worker caste, in which the jaws serve as tools for handling
wax. The ingestion of nectar from the depths of flower corollas or
of honey from the cells of the comb, on the other hand, necessitates
a special apparatus for reaching the liquid and for drawing it up to
the mouth. Hence the principal structural modifications of the feed-
ing apparatus affect those parts that contribute to the formation of
the proboscis and the sucking pump. These same organs are able to
function also in reverse for the regurgitation of nectar and honey.
The principal “new” feature in the mouth parts of the bee is the
salivary channel of the tongue, but this channel is probably only an
elaboration of a groove along the line of union of the glossae. Finally,
the need of having the proboscis out of the way when comb-building
is in progress, or when the mandibles are otherwise in use, has been
met by adaptations in the free parts of the proboscis that enable the
latter to be folded and temporarily stored in a receptive cavity on
the back of the head, from which the parts can be again extended
and reassembled for functional purposes.
The so-called proboscis of the bee is hardly to be termed an “organ,”
since as a functional unit it is improvised when the bee would feed
on liquids by bringing together the various free parts of the
maxillae and the labium (fig. 3 A) in such a way as to form a tube
(C) through which nectar, honey, or water may be drawn up to the
mouth. When not in use the parts are disassembled and folded back
behind the head (fig. 3B), where they are held in place by the
mandibles (Md) clasped beneath them. The major parts of the
maxillae and labium are adapted in form to the roles they play in the
functional position; the flexing mechanism depends on smaller
modifications and minute adjustments between the skeletal elements
and certain muscles. All the specialized movements involved in the
operation of the proboscis of the bee, however, are produced by
muscles that can readily be identified with the usual muscles of the
maxillae and the labium in generalized insects.
The adjustable components of the proboscis are the broad, bladelike
galeal lobes of the maxillae (fig. 3A, Ga), the long labial palpi
(LbPIp), and the slender median tongue formed of the united
glossae (Gls). When these elements are assembled to form a tube,
the galeae and the palpi are brought alongside the tongue and, with
the last, enclose a temporary channel. The wide galeae (C, Ga)
form the sides of the tube and overlap each other above the tongue to
form also the roof; the labial palpi (LbPIp) lie beneath the galeae
and close the tube below by underlapping the tongue, while marginal
fringes of hairs on the galeae and palpi prevent leakage between
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
the approximated parts. The pointed ends of the galeae converge
over the protruding distal part of the tongue (fig. 3 C), but the
small terminal segments of the labial palpi diverge laterally beyond
the galeae and serve as sensory outposts. Back-and-forth movements
of the tongue draw the food liquid into the canal of the proboscis,
through which it ascends probably at first by capillary attraction, but
before reaching the mouth the liquid is received into a preoral food
channel on the base of the labium (fig. 9 A) between the bases of
the maxillae, closed anteriorly by the epipharynx (B). Here at
least the action of the sucking pump must become effective in finally
drawing the liquid food into the mouth.
The proboscis as a whole is retractile and protractile by swinging
back and forth on the long, suspensory rodlike cardines of the maxillae
(figs. 3 A, 4 A, Cd), which are hinged to the sides of the cavity on
the back of the head that lodges the bases of the maxillae and labium,
but the tongue and the paraglossae themselves are deeply retractile
into the end of the basal part of the labium (fig. 7 D, E).
The sucking pump of the aculeate Hymenoptera lies entirely within
the head (fig. 10 A), and is clearly a combination of structures that
in generalized insects constitute two distinct regions of the alimentary
tract. The first is the true pumping apparatus (Cb); it is derived
from the cibarial chamber of the preoral “mouth cavity,’ which
primitively is a food pocket between the epipharyngeal wall of the
clypeus and the base of the hypopharynx that is distensible by
dilator muscles arising on the clypeus. The second part (Phy)
represents the usual postoral pharyngeal dilatatton of the stomodaeum.
In the bee, as in most other sucking insects, the cibarial chamber has
been enclosed within the head and converted into a sucking mecha-
nism; but, while in Hemiptera and Diptera the pump is entirely a
cibarial structure, in the Hymenoptera and Lepidoptera the cibarium
and the pharynx are united to form a single pumping organ, in
which, however, the cibarial and pharyngeal components may be
identified. With the worker bee egestion is an important function
of the feeding apparatus in the preparation of honey from nectar
and in the feeding of the brood, the queen, and the drones; it is
probable, therefore, that the sucking pump is also the principal
regurgitative organ.
The salivary ejection apparatus of the bee is merely the salivary
pocket, or salivarium, between the base of the hypopharynx and
the base of the labium, into which commonly opens the salivary duct.
The salivarium is converted into a closed chamber by the union of
the hypopharynx with the anterior wall of the labium, and its
NO. 2 ; HONEY BEE—SNODGRASS 7
hypopharyngeal and labial muscles become antagonistic dilators and
compressors of the salivary chamber thus formed. The saliva pre-
_ sumably is conveyed around the base of the tongue within the para-
Fic. 1—The head and cranial structures of the worker.
A, anterior view of head. B, posterior view of head, with bases of labium
and maxillae. C, right half of head, mesal view, showing right tentorial bar.
D, foramen magnum and associated structures, posterior view. E, horizontal
section of head through upper part of foramen magnum and below bases of
antennae, dorsal view.
a, shelflike plate on lower margin of foramen magnum; b, hollow bar from
posterior tentorial pit (pt) to plate a on lower margin of foramen.
glossal lobes and carried out to the tip of the tongue through the
salivary channel on the under side of the latter.
The head.—On the facial aspect of the bee’s head (fig. 1 A) the
significant feature is the unusual size of the clypeus (Clp), which
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
latter extends upward from the mandibles almost to the bases of
the antennae beneath the strongly arched epistomal sulcus (es). The
expansion of the clypeus is directly correlated with the great number
of muscle fibers attached on its inner surface, which constitute the
dilator muscles of the internal sucking pump (fig. 10 A, C).
On the back of the head (fig. 1 B) the opening into the neck, the
foramen magnum (For), occupies a central position, and below it is
the deep excavation of the cranial wall known as the fossa of the
proboscis (PF), having a membranous floor in which are implanted
the bases of the maxillae (x) and the labium (LD). To the inflected
sides of the fossa are articulated the rodlike maxillary cardines (C,
Cd). Between the foramen and the fossa the broad postgenal walls
_of the cranium are connected by a subforaminal bridge. The median
part of the bridge clearly is of hypostomal derivation, but the
hypostomal element (D, HB) is constricted between a pair of
mesally directed postgenal lobes (PgL). In Vespidae the postgenal
lobes themselves are united in a postgenal bridge, and the more
primitive hypostomal bridge is reduced to an internal ridge connect-
ing an exposed subforaminal remnant with the hypostomal margin
of the fossa. Thus, as stated by Duncan (1939), the subforaminal
bridge of Vespula “consists chiefly of postgenal structures, though
it does include a hypostomal component.” In Apis, on the other
hand, the bridge is chiefly hypostomal, but includes postgenal
intrusions.
At the sides of the foramen magnum are the long posterior ten-
torial pits (fig. 1 B, D, pt), from which there extends forward and
downward in the head a pair of strong tentorial bars (C, E, AT)
attached anteriorly on the ridge of the epistomal sulcus (C, ER),
with their roots marked externally by a pair of pits in the groove
(A, E, at). These bars, except for their extreme posterior ends,
represent the anterior arms of the tentorium as this structure is
developed in more generalized insects. Just within the foramen
magnum the longitudinal tentorial bars are connected dorsally by a
narrow, arched rod, or tentorial bridge (D, E, TB), and shortly
before the bridge each bar gives off a slender threadlike branch
representing the usual dorsal tentorial arm (C, E), which extends
forward but disappears before reaching the facial wall of the head.
The tentorium of the bee, besides bracing the cranial walls, gives
attachment to muscles of the antennae, the maxillae, the labium, the
pharynx, and the oral plate of the sucking apparatus. From the
deeper part of each posterior tentorial pit there extends into the
head cavity a slender hollow rod (D, b), which is attached mesally
NO. 2 HONEY BEE—SNODGRASS 9
to a small plate (a) inflected from the lower lip of the foramen
magnum.
The antennae.——The antennae arise close together near the center
of the face (fig. 1 A, Ant), where each appendage is implanted in a
small membranous area with a slightly elevated rim, known as the
antennal socket. Each antenna (fig. 2A, G) is divided by an elbow
into two major parts, a basal stalk, or scape (A, Scp), and a long
distal arm subdivided into 11 small sections in the female (A) and
ew
|
NB)
Wize
Fic. 2.—The antenna.
A, right antenna of worker and cranial muscles inserted on base, mesal view.
B, two consecutive subsegments of flagellum pulled apart to show necklike
membranous connection. C, inner rim of base of right antenna, showing position
of muscle insertions. D, base of right antenna and section of ‘antennal “socket”
through antennafer (af), posterior view. E, joint between scape and pedicel,
ventral view. F, rim of right antennal socket, anterior view. G, antenna of
drone. H, proximal part of antenna of worker, showing muscles in scape
inserted on pedicel.
I2 in the male (G). The proximal piece of the arm is usually
distinguished as the pedicel (Pdc), and the rest termed the flagellum
(Fl), but in the bee there is no pronounced differentiation between
these two parts. The flagellar subdivisions are not true segments
since they are not articulated upon each other nor provided with
muscles; the rounded base of each (B) fits into a distal depression
of the one proximal to it, and the two are connected by a narrow,
cylindrical, necklike membrane. The joint between the pedicel and
the scape, on the other hand, has a dicondylic leglike articulation (FE),
and hence forms a definite transverse axis of movement, on which
the pedicel and flagellum are turned up or down by levator and
10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
depression muscles arising respectively on the dorsal and the ventral
wall of the scape (H, 6, 7).
The scape of the antenna is articulated to the head by a distinct
basal knob (fig. 2D), which is pivoted on a small articular process,
the antennafer (af), arising ventrolaterally from the rim of the
antennal socket (f/f). The antenna is thus freely movable in all
directions except as it is limited by the socket membrane. To provide
for its movement four muscles arising on the concave dorsal surface
of the tentorial bar of the same side of the head are inserted on the
base of the scape (C), two above the level of the pivot and two
below it.
The labrum.—tThe labrum of the honey bee (fig. 1 A, Lm) is a
simple, transversely elongate flap with well-sclerotized, closely ap-
pressed outer and inner walls, and is freely suspended from the
lower edge of the clypeus. It contains no compressor muscles (fig.
10 C, Lm), and is movable by only a single pair of extrinsic muscles
arising on the frontal area of the head between the antennal bases,
which are adductors since they are inserted by long tendons attached
laterally on the posterior margin of the base of the labrum. In the
wasps the labrum is retracted beneath the clypeus and the clypeal
margin forms the lower edge of the face (fig. 9 E).
The epipharynx.—tn both the wasps and the bees the epipharynx
(figs. 9E, 10C, Ephy) is a preoral outgrowth of the inner, or
“epipharyngeal,’ surface of the clypeus—not of the labrum. The
epipharynx of the honey bee is a large, soft structure with a prominent
median keel and padlike lateral lobes (fig.g A, Ephy). It is retractile
by a group of divergent muscle fibers arising on the lower part of
the clypeus (fig. 10 C, 25). Just above and behind the base of the
epipharynx is the wide, oval mouth opening (figs. 9g A, 10 C, Mth)
that leads into the cavity of the sucking pump. The epipharynx is
richly provided with sense organs, but mechanically it serves, to-
gether with the lacinial lobes of the maxillae, to close the food channel
on the base of the proboscis (fig. 9 A, B).
The mouth—The wide aperture behind the epipharynx that leads
into the sucking pump (figs. 9g A, 10 A, Mth) is the mouth of the
bee in a functional sense, but since the anterior part of the pump
(fig. 10 A, Cb) represents the primitively preoral cibarium lying
between the clypeus and the hypopharynx, the true mouth, in a
morphological sense, is between the cibarial and pharyngeal sections
of the pump. In the Hymenoptera, therefore, as in various other
sucking insects, the functional mouth is a secondary constriction of
the food meatus between the inner face of the clypeus and the
eae
ee ee
NO. 2 HONEY BEE—SNODGRASS II
anterior wall of the hypopharynx (fig. 10 A, Mth). In some of the
Hymenoptera the mouth is guarded by a pair of long, valvelike lips,
the upper one being the epipharynx, the lower one a fold from the
lower edge of the mouth. In the bee the suboral fold hangs like a
bib from the lower lip of the mouth (fig. 9 A, bib).
The mandibles——The mandibles of the honey bee differ in shape
and relative size in the three castes (fig. 3 F, G, J). In the worker
(F) each mandible is elongate, thick at the base, narrowed through
the middle, and widened again distally in a flattened expansion. The
inner face of the expanded part is somewhat concave and traversed
obliquely by a channel (d) fringed on both sides with hairs, which
is continuous with a groove (e) that runs upward to the orifice of
the mandibular gland (f) at the base of the jaw. The mandible is
suspended from the lower lateral angle of the cranium (fig. 1 A, C),
and has the usual two points of articulation, one with the lower part
of the clypeus (fig. 3 E, c), the other (a’) with the subgenal margin
of the cranium. The hinge line of the jaw between the two articula-
tions slopes downward from in front, so that when the mandible is
adducted its apex turns not only mesally but also posteriorly.
Each mandible has only two muscles, an abductor and an adductor.
The fibers of the abductor spread over the lower part of the side wall
of the head behind the compound eye (fig. 3 I, 8) and converge
upon a stalked apodeme arising in the articular membrane at the
outer side of the mandibular base (EK, 8Ap). The large adductor
muscle (H, 9) has several groups of fibers, some of which arise
behind the eye above the abductor fibers, others on the back of the
head below the level of the foramen magnum, while a small group
(9a), inserted on a long slender branch of the apodeme, takes its
origin on the top of the head. The strongly stalked adductor apodeme
arises from the articular membrane at the inner side of the base of
the mandible (fig. 1 C, 9Ap), and is closely applied to the mesal wall
of the mandibular gland (fig. 3 F).
The worker bee makes various specific uses of its mandibles, in-
cluding the collecting and breaking of pollen grains for food, manipu-
lation of wax in comb-building, supporting the base of the extended
proboscis while feeding on liquids, and holding the flexed proboscis
in place when the latter is folded behind the head (fig. 3B). The
provision of each mandible with but two muscles gives the jaws
active movements of abduction and adduction only, but there is
sufficient flexibility at the articulations to allow of some play of the
mandibles on each other. When the mandibles are used to hold the
base of the extended proboscis, the channels on the inner surfaces
12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I03
of their expanded ends fit exactly on the midribs of the maxillary
galeae and allow the latter to slide between them. When the proboscis
is folded (fig. 3B) the mandibles are crossed behind the labrum
against the bases of the flexed galeae, and thus hold the proboscis
snugly against the head.
Passively the mandibles, when partly opened, form a conduit for
the discharge of nectar or honey and of brood food from the mouth.
According to Park (1925), “the honey is forced out over the dorsal
surface of the folded proboscis between the mandibles which are
held well apart’; and in a personal communication Frank E. Todd
of the United States Bureau of Entomology and Plant Quarantine
says, ‘‘feeding of royal jelly, and of honey and pollen, to the larvae
is done through the mandibles.”
The mandible of the queen (fig. 3 J) is of about the same length
as that of the worker (F), but is bilobed distally and much wider at
the base; it has a concavity on the inner face of the apical lobe, but
there is no groove leading up to the orifice of the mandibular gland.
The posterior surface is clothed with more numerous and longer
hairs than the worker mandible, but in both castes of the female the
mandibular hairs are unbranched. The mandible of the drone (G) is
relatively and absolutely smaller than that of either the worker or
the queen ; its distal part is narrow, provided with a small apical tooth,
and has a mesal depression from which a faintly marked groove leads
up to the base of the jaw. The hairs of the drone mandible are par-
ticularly long and numerous, and, in contrast to those of the female
mandible, are nearly all of the plumose variety.
The mandibular gland of the worker (fig. 3 F, MdGld) and the
queen is a large sack lying between the facial wall of the head and
the apodeme of the adductor muscle of the mandible. The gland in
the worker extends upward to the level of the antennal bases, and
in the queen is even larger. In the drone the mandibular gland is
but a small vesicle at the base of the jaw. The secretion of the
mandibular glands of the bee is said to serve for softening wax, but
corresponding glands in other insects presumably have a “salivary”
function in connection with feeding.
The maxillae——The two maxillae (fig. 3 A) lie at the sides of the
median labium rather than before it. The long stipital sclerites (St)
of the maxillary bases are implanted proximally in the membrane of
the proboscis fossa on the back of the head, and are suspended from
the hypostomal margins of the fossa by the slender cardines (Cd),
which also are contained in the fossal membrane. The free distal
part of each stipes bears a long, broad, tapering galea (Ga), a large,
HONEY BEE—SNODGRASS 13
/
LbPlp
Gls B
|
Fic. 3—The mandibles, maxillae, and labium.
A, maxillae and labium of worker with parts artificially spread out behind
the mandibles, suspended by maxillary cardines from margins of proboscis
fossa (PF) on back of head. B, mandibles and free parts of maxillae and
labium folded beneath head. C, distal parts of maxillae and labium assembled
to form a proboscis, anterior view. D, lateral view of retracted and folded
labium and left maxilla. E, left mandible of worker and adjoining parts of
head, lateral view. F, right mandible of worker and mandibular gland, mesal
view. G, right mandible of drone, mesal view. H, adductor muscle of right
mandible of worker, arising behind compound eye. I, abductor of right mandible
of worker. J, right mandible of queen, mesal view.
a’, posterior articulation of mandible; a”, cranial articulation of maxillary
cardo; c, anterior articulation of mandible; d, mesal depression of mandible:
e, mesal groove of mandible; f, orifice of mandibular gland.
14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
soft, cushionlike lobe arising mesad of the base of the galea, and a
very small, 2-segmented, lateral palpus (MxPlp). A V-shaped |
sclerite, known as the lorum (Lr), which holds the basal plate of the
labium in its apical angle, has its arms articulated with the distal
ends of the maxillary cardines.
Each maxilla is provided with four extrinsic muscles arising within |
the head (fig. 4 A), one inserted on the cardo (10) ; the other three ©
(11, 12, 13) on the stipes. The single cardinal muscle (70) is the
usual promoter of the maxilla, which in generalized insects arises
dorsally on the head wall and is inserted on the cardo anterior to the
cranial articulation of the latter. In the bee, however, this muscle
arises postertorly on the postgenal inflection at the side of the
proboscis fossa (fig. 1 C, Pge), and is inserted on a short lever arm
of the cardo that projects above the articular condyle (fig. 4A). By
this shift in its point of origin the cardinal muscle becomes an effective
protractor of the maxilla. The movement of the two maxillary
cardines, however, affects both the maxillae and the labium because
the three appendages are yoked at their bases by the V-shaped lorum —
(fig. 3 A, Lr). The muscles of the cardines in the bee, therefore, are
cardinal protractors of the proboscis.
The long stipital muscles of the maxilla (fig. 4 A, 17, 12, 13) are —
the primitive adductors, or functionally, protractors of a generalized |
maxillary appendage, that arise on the tentorium. In the bee these
muscles have their origins on the extreme anterior ends of the longi- ¢
tudinal tentorial bars (AT), and hence at first sight appear to arise
|
|
on the facial wall of the head. In most insects the stipital muscles act
as protractors of the maxilla because their mesal pull (adduction)
flattens the angulation between the cardo and the stipes and in this —
way protracts the appendage. The muscles evidently produce the
same action in the bee, and become stipital protractors of the
proboscis. Though the slant of the muscles in the fully protracted —
condition of the maxilla might suggest that the stipital muscles now
become retractors, it is probable that the retraction of the proboscis —
is effected principally by the long cranial muscles of the labium (fig.
FEO ed Ae
The tapering bladelike galea of the maxilla is attached to the stipes
by a prolongation of its base that forms a triangular plate, or subgalea —
(fig. 4D, G, Sga), implanted laterally on the anterior surface of the —
distal end of the stipes. From the subgalea a strong midrib runs —
through the length of the galea to its tip. In the functional, protracted —
position of the proboscis the galeal blade extends straight out from
a
NO. 2 HONEY BEE—SNODGRASS 15
MxPlp
Fic. 4.—The maxilla.
A, right maxilla of worker and maxillary muscles, mesal view. B, left maxil-
lary palpus and muscle, lateral view. C, lacinial lobe and lever of right maxilla
of drone, mesal view. D, base of left maxilla of worker, lateral view. E, right
maxilla with galea extended, mesal view. FF, same, galea partly deflexed.
G, stipes and base of galea of right maxilla, anterior view.
a”, cranial articulation of cardo; a-b, c-d, lines of flexure in base of galea;
e, stipital arm in base of mesal wall of lacinial lobe; f, articular point of galea
on e; g, leverlike marginal ridge on base of galea articulating with articular
point (f) of stipes.
16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
the stipes (fig. 4E), its broad outer part being vertical and the
narrower mesal part horizontal, as the two galeae overlap to form the
roof and the sides of the temporary proboscis canal (figs. 3 C, 9 C).
When the proboscis is retracted the galea is bent back and folded
upside down below or behind the stipes (fig. 3 D).
The folding of the galea is produced by a single muscle arising in
the stipes (fig. 4 F, 15), which evidently represents the usual galeal
flexor of other insects. The action of the muscle in the bee, however,
depends on a special device in the base of the galea. Where the galea
joins the subgalea it is abruptly narrowed, and the lateral margin of
the narrow part is thickened to form a small leverlike ridge (E, g)
that articulates proximally on the tapering, laterally curved end (f)
of a long narrow sclerite (e) in the mesal wall of the stipes. On the
middle of the lever ridge is attached the tendon of the flexor muscle
of the galea (F, 15). The tension of the muscle in contraction first
depresses the galea and turns it backward and outward because of
the obliquity of the line of bending (E, a-b) between the galea and
the subgalea; but a second line of bending (c-d) beyond the first and
oblique in the opposite direction soon counteracts the lateral move-
ment, so that the continuing pull of the muscle finally turns the galea
straight back and folds it up against the stipes. The extension of
the galea evidently results automatically from the elasticity of its base
and its basal continuity with the firmly affixed subgalea (G).
The large, soft, cushionlike lobe of the stipes that arises at the
mesal side of the subgalea (fig. 4A, D, F, G, Icl) is a prominent
feature of the bee’s maxilla. On its proximal end is a small sub-
sidiary lobe (D, G, Lc) bearing a few small setae, which, by com-
parison with other Hymenoptera (fig. 5), evidently represents the
maxillary /acinia. In a vespoid wasp (fig. 5 A, B) the lacinia is a
well-developed lobe (Lc) arising from the stipes mesad of the base
of the larger galea (Ga). In Bombus (D) and Xylocopa (E) the
lacinia is a small setigerous or spiny lobe overlapping a large mem-
branous area of the stipes. It would appear, therefore, that the major
part of the large cushionlike lobe of the maxilla in Apis (fig. 4 D, Icl)
is a development of the sublacinial membranous area present in
Bombus and Xylocopa, and that the small setigerous lobule (Lc)
on its base is a remnant of the lacinia proper. The entire structure in
Apis is here termed the lacinial lobe; it plays an important part in the
closure of the food channel on the base of the proboscis.
In the steeply declivous proximal wall of the lacinial lobe is a
small curved sclerite (fig. 4 E, jvr) with a tapering distal arm and a
thickened base, which latter is connected by a fulcral point with the
NO. 2 HONEY BEE—SNODGRASS 17
proximal end of the narrow mesal sclerite (e) of the stipes. The
lacinial sclerite serves in the bee as a lever for keeping the lobe erect.
On the upper point of its base is inserted a muscle from the stipes
(F, 16), the contraction of which revolves the lever on its fulcrum
and thus exerts a tension on the lobe. The lacinial lobe is the “Segel-
halter’ of Wolff (1875), who notes the lever sclerite in its base, and
terms its muscle the “Spanner des Segelhalters.” The muscle repre-
sents the flexor of the lacinia in generalized insects; the lever ap-
paratus is present also in other Hymenoptera. In Vespula (fig. 5 A,
Fig. 5—Examples of the maxillary lacinia in Vespidae and Apoidea.
A, left maxilla of Vespula maculata (L.), with well-developed lacinia, dorso-
lateral view. B, right maxilla of same, ventromesal view. CC, detached lacinia
of left maxilla of same, lateral view, showing mesal lever arm (/vr). D, base
of right maxilla of Bombus americanorum (F.), with small lacinia attached to
large membranous area, mesal view. E, right lacinia and lever sclerite of
Xylocopa virginica (L.), mesal view.
B, C) a small sclerite (Jur) in the base of the lacinia, giving insertion
to the lacinial muscle (A, B, 76), curves around the proximal end
of the lacinia to articulate on the stipes (B). In Bombus (D) the
base of the lacinia itself articulates with the stipes, but in Xylocopa
(E) there is a distinct lever sclerite (Jur) giving insertion to the
lacinial muscle (16).
The minute, two-segmented palpus of the maxilla (fig. 3 A, MxPlp)
arises from the end of the stipes laterad of the base of the galea, and
is provided with a large muscle (fig. 4B) arising in the stipes
(F, 14). It is not evident why so small and seemingly unimportant
an organ should have a muscle of such size. Wolff (1875) believed
that the palpus muscle serves as an extensor of the galea, and Mori-
son (1927) regarded it as a muscle of the galea, which, he says
18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I03
mistakenly, is attached “near” the base of the palpus. There can be
no question that the muscle is inserted directly on the base of the
palpus (fig. 4 B), and it is not clear how it could act as an extensor
of the galea. Duncan (1939) says there are two muscles for each
maxillary palpus in Vespula inserted on the palpal base.
The labium and hypopharynx.—tThe labium of the bee presents
externally all the parts of a generalized labial appendage (fig. 3 A).
The lengthened base contains a small, triangular postmental sclerite
(Pmt) and a long trough-shaped premental plate (Prt). Diverging
from the end of the prementum are the slender, four-segmented
labial palpi (LbPIp), each supported on a small basal’ stalk, and
between the latter arise the ligular lobes, including the long, median,
so-called tongue (Gls), which is probably the combined glossae, and
a pair of short lateral paraglossae (Pgl). The postmentum and the
proximal part of the prementum are suspended in the membranous
floor of the fossa of the proboscis (PF) on the back of the head,
and are membranously united also with the bases of the maxillary
stipites. The apex of the postmentum is embraced by the angle of
the lorum (Lr), and turns in the latter when the proboscis is pro-
tracted or retracted. In the completely retracted position the apex
of the lorum rests in the depression of the hypostomal bridge (HB).
The premental sclerite covers only the posterior and lateral walls
of the premental area of the labium, the anterior wall being entirely
membranous (fig. 6 A, 7D). The membranous anterior wall ends in
a transverse inflection at the bases of the ligular lobes (fig. 6 A), and
in this groove, as may be seen by separating the paraglossae (B), is
the opening of the salivary ejection apparatus (S/O). The apparent
anterior wall of the prementum, therefore, represents the hypopharynx
(B, Hphy) adnate upon the labium, since the salivary duct in
generalized insects opens between the hypopharynx and the labium.
The hypopharyngeal surface in the bee, furthermore, extends up
to the functional mouth (fig. 10 A, Hphy), and includes the oral
plate (ofl) on the floor of the mouth cavity at the entrance to the
sucking pump and the biblike fold (bib) that hangs down from the
margin of the plate. The ascendent hypopharyngeal surface is flanked
by a pair of slender suspensorial rods (figs. 7D, 9 A, s) that extend
from the base of the prementum to the mouth angles, and probably
represent lateral hypopharyngeal sclerites of generalized insects.
The ligular lobes of the labium have a common base which is con-
tinuous with the prementum (fig. 6 A, B, C), but is supported on the
latter anteriorly by a pair of ligular arms (A, B, h), and posteriorly
by a triangular subligular plate (C, k). The ligular arms are sclerotic
bands arising from the anterior margins of the premental plate; at
NO. 2 HONEY BEE—SNODGRASS 19
am
i
Fic. 6.—The labium of the worker.
A, end of prementum with ligula and bases of labial palpi, anterior view,
paraglossae in natural position. B, same, paraglossae spread apart exposing
salivary opening (S/O). CC, same, posterior view, showing base of glossal
rod (rd) and entrance to salivary channel (sc). D, two tiers of glossal hairs,
anterior view. E, base of glossal rod exposed by eversion of salivary channel.
F, ligula and salivary syringe, anterior view, anterior wall of syringe removed.
G, end of glossal tongue, posterior view. H, diagram of base of protracted
tongue. I, same, tongue partly retracted by contraction of glossal muscles (20).
h, ligular arm of prementum; 7, pivotal process of base of tongue; j, basal
plate of tongue; k, subligular plate of prementum; /, canal of glossal rod;
m, membranous wall of salivary channel; n, elastic posterior rod of labial
palpiger; rd, glossal rod; sc, salivary canal of tongue.
20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
the base of the ligula they are inflected laterad of the salivary opening
(B), and are then continued as a pair of slender processes (7) that
support the bases of the paraglossae laterally, and finally curve in-
ward to form a pair of fulcral points articulating with the base of
the tongue (j). The subligular plate (C, k) is directly continuous
from the prementum, but is flexible on the latter, and its apex is
produced into the rod (rd) that traverses the channel of the tongue.
The long, cylindrical, densely hairy glossal tongue of the honey bee
has a closely ringed structure, in which hard circles bearing the hairs
alternate with membranous smooth areas (fig. 6 D), so that the organ
is both flexible and contractile. The posterior, or under, surface
presents a narrow median groove (C, sc) that expands within the
tongue into a wide thin-walled channel. The inner wall of the channel
is traversed by a slender, elastic rod (rd), which is itself grooved
on its outer surface (FE), and the groove is fringed with small hairs
directed distally. The lips of the glossal channel spread apart at the
base of the tongue (C) and expose the proximal end of the rod,
which is here seen to be directly continuous with the apex of the
subligular plate (C, E, k). Distally the rod ends in a small, freely
projecting, spoon-shaped terminal lobe of the tongue, known as the
flabellum (G, Fbl). The convex under surface of the flabellum is
smooth, but the distal margin and the upper surface are covered
with small branched hairs. The base of the tongue is somewhat
thickened; anteriorly it is covered by a hard, bonnet-shaped sclerite
.(F, H, 7) decurved abruptly before the salivary orifice and produced
distally in a tapering plate on the tongue surface. The basal sclerite
is embraced laterally by the fulcral processes (7) of the ligular arms
of the prementum, which hold the tongue firmly in place, but allow
it to revolve freely in a vertical plane on the transverse axis between
their opposing points.
The paraglossae are thin, elongate lobes arising from the common
ligular base at the sides of the tongue (fig. 6 A, B, C, Pgl), where
they are attached to the sides of the distal process of the ligular arms
(B, 7) that form the fulcral points of the tongue. The concave mesal
surfaces of the paraglossae clasp the base of the tongue (A) and
underlap its posterior surface, thus covering the proximal end of the
tongue channel. The paraglossae automatically assume this position ;
they are not individually movable, since no muscles are attached
directly on them.
The salivary ejection apparatus, which may be termed the salivary
syringe, opens by a wide aperture situated, as already noted, anteriorly
at the distal end of the prementum just behind the declivous base of
NO. 2 HONEY BEE—SNODGRASS 21
the tongue (fig. 6B, S/O). The aperture leads into a wide, flat, tri-
angular chamber (F, Syr, exposed here by removal of a part of the
anterior wall), into the inner end of which opens the salivary duct
(SID). The floor of the chamber is a rigid, slightly concave plate
continuous at its distal angles with the inflected ends of the ligular
arms of the prementum (/), which flank the salivary orifice (B). The
roof is flexible and elastic, and gives insertion to a pair of convergent
dilator muscles (fig. 7A, 23) arising on the anterior edges of the
premental sclerite. A pair of much larger muscles arising in the base
of the prementum (figs. 6 F, 7 A, C, 24) and inserted on the lateral
margins of the syringe evidently act as expulsors of the saliva by
flattening the dilated chamber. The salivary muscles are effective
only in the protracted condition of the proboscis; as they are shown
in figure 7 A the muscles are slack and nonfunctional because of the
retraction of the ligula into the prementum.
The salivary syringe of the bee is an elaboration of the usual
salivarium, which in generalized insects is a mere pocket between
the hypopharynx and the base of the labium. The union of the
hypopharynx with the labium has converted the salivary pocket into
a closed chamber, and the ordinary hypopharyngeal and labial muscles
of the salivarium become, respectively, dilators and compressors of
the syringe.
The saliva, forcibly expelled from the salivary syringe, encounters
at once the steeply declivous base of the tongue, which lies im-
mediately before the salivary orifice (fig. 6B). It must, therefore,
be deflected in two divergent streams past the sides of the tongue
into the concavities of the mesal surfaces of the paraglossae, and
by the latter conveyed to the posterior surface of the tongue. Here,
presumably, the saliva enters the tongue channel and runs through
it to the tip of the organ, where it flows out on the under surface of
the apical flabellum (fig. 6G). It must be admitted that, so far as
known to the writer, no direct observations have been made on the
course of the saliva in a living bee; the relations of the structural
parts concerned, however, would seem to leave no other course avail-
able than that described above, except, perhaps, that the actual
conduit of the saliva through the tongue may be the groove of the
glossal rod instead of the larger channel that opens on the surface
of the tongue. The hair-fringed groove of the rod clearly must have
some specific function.
It is possible that the same apparatus may be used also in the
process of “ripening” nectar into honey, during which process, as
described by Park (1925), nectar extruded from the mouth is said to
22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
flow over the upper surface of the proboscis and to swell into a
globule in the posterior angle of the proboscis between the recurved
free distal parts and the basal parts of the latter. The nectar globule
is several times sucked back into the mouth and redischarged. Though
recorded observations are not specific as to the exact course of the
nectar around the base of the tongue, it is much to be suspected that
the passageways between the tongue and the paraglossae, and likewise
the glossal channel itself, the wall of which is eversible, serve to
conduct the nectar in both directions.
The principal movements of the labium are (1) the back-and-forth
motion of the labium with the maxillae when the entire proboscis is
retracted and protracted, (2) an independent movement of the labium
between the maxillae, (3) individual movements of the tongue during
feeding, and (4) the backward flexion of the tongue and palpi when
the proboscis is not in use. The flexion of the tongue accompanies
the retraction of the proboscis, and involves a retraction of the base
of the ligula into the end of the prementum. Other minor movements,
probably depending on muscle tension, adjust the various parts to
their functional positions.
The extrinsic musculature of the bee’s labium consists of two pairs
of muscles corresponding with the anterior and the posterior labial
adductors of generalized insects inserted on the prementum. The
anterior labial muscles in the bee (fig. 7 D, 17) are the longest muscles
of the head; they arise on the dorsal wall of the cranium, go down-
ward through the head, traverse the prementum (A), and insert by
tendons on the distal ends of the ligular arms (A, D, h) of the pre-
mentum. These muscles are labial retractors of the proboscis, since
their contraction affects not only the labium but the maxillae as well.
Also they take part in the retraction of the ligula into the end of the
prementum (A, E), which causes the deflexion of the tongue. The
posterior labial muscles (D, 7S) arise on the anterior ends of the
tentorial bars of the head and converge posteriorly to a single median
tendon attached on the proximal extremity of the prementum. These
muscles probably act as protractors of the labium by flattening the
angle between the postmentum and prementum, thus giving the
labium an independent movement between the maxillae, and in addi-
tion they draw the prementum toward the mouth during feeding.
The posterior labial muscles may, therefore, be termed protractor-
adductors of the labium.
The movements of the tongue during feeding, or when the bee is
exploring a food source, consist of lengthwise extensions and con-
tractions of the organ, and lateral movements of the part projected
NO. 2 HONEY BEE—SNODGRASS 23
beyond the galeae and palpi. The movements are so varied and
rapid that the tongue itself seems to be endowed with mobility. Only
two muscles, however, account for the activity of the tongue, and
they are inserted on the base of the axial rod. These muscles (fig.
Fic. 7—The mechanism of ligular retraction, and the salivary syringe.
A, lengthwise section of prementum of worker with ligula and _ salivary
syringe retracted by pull of muscles 17 and 79 on ends of ligular arms (h),
salivary muscles inoperative. B, detail of base of retracted tongue. C, salivary
syringe and muscles, posterior view. D, prementum, with ligula protracted.
E, same, ligula retracted as at A.
h, ligular arm of prementum; 7, pivotal process of h supporting base of tongue;
k, subligular plate of prementum supporting base of glossal rod; s, suspensorial
rod of hypopharynx.
6H, 20), arising in the base of the prementum (fig. 7A), un-
doubtedly represent the usual flexors of the two glossal lobes of
generalized insects. The rod of the bee’s tongue is continuous distally
with the flabellum, it traverses the inner wall of the membranous
glossal channel, proximally it turns backward (fig. 6 H, rd) and is
24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
attached to the apex of the subligular plate (k) at the end of the
prementum. The two muscles of the rod (20) are attached by tendons,
not directly on the rod, but immediately at the sides of its recurved
basal part. It is clear that the contraction of the muscles increases the
basal curvature of the rod (I), and therefore pulls the rod back
through the tongue as far as the membranous wall of the enclosing
channel will permit. The tongue is thus shortened, owing to the
compressibility of its ringed wall, particularly in its distal part. On
relaxation of the muscles the tongue is again extended, evidently by
the elasticity of the rod and the compressed tongue wall, since there
is no specific extensor mechanism. Furthermore, the attachment of
the muscles at the sides of the rod enables the muscles by acting
singly and alternately to revolve the rod, and thus probably are
effected the varied lateral movements of the tip of the tongue.
We may now examine the mechanism by which the base of the
ligula is retracted into the end of the prementum and the tongue turned
back behind the head in the position it assumes when the proboscis
is retracted (fig. 3B, D). In the functionally protracted condition
of the labium (fig. 7 D) the tongue and the paraglossae are extended
from the end of the prementum and their bases are fully exposed.
In the retracted state (E) the base of the ligula is deeply sunken into
the end of the prementum, and the tongue is sharply bent backward
behind the latter. It is to be seen also that the ligular arms of the
prementum (D, h) have been completely folded inward, and that the
subligular plate is angularly bent upward (B, k). The position of
the parts concealed by retraction is best seen in a sectional view (A).
The retraction of the ligula is brought about by two pairs of
muscles inserted on the distal ends of the ligular arms of the pre-
mentum (fig. 7A). One muscle of each pair is the long retractor
of the labium (77) arising in the top of the head; the other muscle
(19) arises in the base of the prementum, and probably represents
the paraglossal muscle of other insects. The pull of these muscles
on the ends of the ligular supports (/) inflects the latter into the
membranous distal end of the prementum, and consequently retracts
the attached tongue, paraglossae, and salivary syringe, while the
subligular plate (k) is bent forward and upward beneath the base of
the tongue.
The base of the retracted tongue (fig. 7 A, B), held by the pivotal
processes (1) of the ligular arms, is separated by a considerable
distance from the end of the subligular plate (k), to which the tongue
rod (rd) is affixed. As a consequence the rod has been drawn in a
wide loop far out of the base of the tongue. Since the rod traverses
NO. 2 HONEY BEE—SNODGRASS 25
the posterior part of the tongue in the ample membrane of the glossal
channel, the contractile and flexible tongue is not only shortened by
the retraction of the rod, but is sharply curved in a posterior direction.
Thus the retraction of the proboscis by the retractor muscles of the
labium effects also the retraction of the base of the ligula into the
prementum, and the latter action produces automatically the shorten-
ing and posterior flexion of the tongue. The salivary syringe also is
pulled inward by the inflection of the ligular arms, and in the re-
tracted condition its muscles are slack and temporarily functionless.
Inasmuch as there is no muscular mechanism for the protraction of
the ligula, it must be assumed that the parts are extended by the
elasticity of the inflected ligular arms and the subligular plate when
the retractor muscles relax.
The long, 4-segmented labial palpi (fig. 8 A) arise from the distal
end of the prementum, where each palpus is supported on a small
basal lobe (Plg) commonly regarded as being a palpiger, i.e., a palp-
supporting lobe of the prementum, rather than a basal segment of the
palpus. In nonconformity with this idea, however, the single muscle
of the palpus (21) is attached within the basal lobe. The palpus itself
contains only one muscle (22), which arises in the long first segment
and is attached mesally on the base of the succeeding segment ; it flexes
the distal part of the palpus, which, when released, springs back to
the extended position by its own elasticity.
The flexing mechanism of.the entire palpus, by which the palpus is
turned back behind the prementum when the proboscis is retracted,
is in the small palpiger. Each palpiger is flattened from side to side,
and its walls are membranous except for a narrow sclerotic bridge
in the anterior wall, which connects the first palpus segment with the
prementum (fig. 8A), and a slender, flexible and elastic rod in the
posterior wall (A, D, 1) that is continuous from the prementum to the
midrib of the palpus. The basal muscle of the palpus (B, 21) is
attached by a long tendon on the posterior rod a little beyond the
middle of the latter. When the palpus is extended (B) the rod is
slightly curved into the rear wall of the palpiger; when artificially
' flexed (C) the palpus is turned back with its lateral surface posterior,
and the elastic rod of the palpiger (”) is strongly curved into the
posterior wall of the palpiger. There can be little question, therefore,
that the flexing of the palpus results, conversely, from an incurving
of the rod by the pull of the muscle attached on it, and, in fact, the
action can be demonstrated in a dead bee by pressing the rod so as
to bend it into the palpiger. The exact movements of the palpus,
26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
however, depend on the obliquity of lines of bending at the ends of
the palpiger (B, a-c, b-c, and b-d).
The movements of the palpiger can be well illustrated with a ‘piece
of stiff paper cut into the form shown at E of figure 8 and creased
Pie Prot
AC Q{E_E
Ty XY SSS
d
Wn a f
Fic. 8.—The labial palpus.
A, right palpus, palpiger, and prementum, mesal view, showing muscles.
B, base of palpus in position of extension from prementum. C, same, palpus
flexed. D, base of palpus, posterior view, showing elastic rod (7) in posterior
wall of palpiger, and attachment of flexor muscle tendon. E, F, diagrams of
flexing of palpus on prementum, illustrated with piece of paper cut and creased
as at E, and folded as at F.
a-c, b-d, lines of folding at ends of palpiger; b-c, line of diagonal folding of
palpiger; b-e, line of infolding at base of flexed palpiger; f, distal point on
palpus; , elastic rod in posterior wall of palpiger.
along the lines a-c, b-c, b-d, and b-e. By folding outward along the
diagonal (b-c) through the “palpiger” the triangle abc is turned over
and directed downward (F) ; the ‘“‘palpus” can now be bent backward
in a horizontal position, but its alignment with the “prementum”
necessitates a compensating fold inward along the lines b-d and b-e.
NO. 2 HONEY BEE—SNODGRASS 27
The distance from c to d is thus shortened, and, by reversing the
sequence of the movements, the approximation of c and d would
produce the folding. In the palpiger of the bee (B) this shortening
results from the inward curvature of the rod (m) effected by the
pull of the muscle on its convex side, and the palpus automatically
turns over and bends backward (C). Extension of the flexed palpus
evidently depends on the elasticity of the palpiger, since there is no
extensor muscle.
The preoral food channel_—The membranous anterior, or hypo-
pharyngeal, wall of the labium extends upward from the salivary
orifice at the distal end of the prementum to the mouth, and forms
the floor of a deep preoral food channel between the maxillary bases
(fig. 9 A, FC) that is continuous with the closed food canal of the
distal part of the proboscis (C, fc). The wide aperture of the mouth
(A, Mth) at the upper end of the channel lies’ behind the base of the
epipharynx (Ephy), and the epipharynx is covered by the projecting
labrum. On the floor of the mouth is the broad oral plate (fig. 10 A,
B, ofl), from which depends the large, thin, double-pointed, tri-
angular fold (bib) that hangs like a bib from the lower lip of the
mouth over the adoral hypopharyngeal surface (fig. g A, bib). At
the sides of the bib in the hypopharyngeal membrane are the slender
hypopharyngeal suspensory bars (s), which taper downward to the
sides of the prementum.
When the proboscis is depressed or retracted, the mandibles are able
to close before the mouth between the labrum and the bases of the
maxillae. If the mandibles are spread apart when the proboscis is
protruded the preoral food canal is widely exposed (fig. 9 A) and
the lacinial lobes of the maxillae (/cl) are prominently displayed on
the mesal margins of the stipites. When the proboscis is fully ex-
tended in the functional position, the mandibles are opened, allowing
the base of the proboscis to swing up between them, and they now
clasp the bases of the galeae. In this position, the proboscis extends
straight out from before the mouth; the formerly vertical floor of the
preoral food channel becomes more nearly horizontal, and the cushion-
like lacinial lobes of the maxillae are pressed into the lateral con-
cavities of the epipharynx, with the median keel of the latter closing
the space between them (fig. 9 B), while the labrum clamps down
on the epipharynx. The preoral food channel on the base of the
proboscis is thus converted into a closed passageway, by which the
food canal of the distal part of the proboscis is put into communica-
tion with the mouth. The approximation of the labium to the mouth
bows the suspensorial rods (A, s) outward and accentuates the
28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
depression between them into a deep heart-shaped cavity. The bib
dependent from the lower lip of the mouth, however, has now
assumed a nearly horizontal position with its points resting on the
hypopharyngeal surface of the prementum, and thus forms a ramp
from the floor of the food channel to the mouth, bridging the cavity
behind it.
The cavity temporarily formed behind the suboral fold in the bee
by the infolding of the adoral hypopharyngeal surface (fig. 10 A, C,
IbS) evidently corresponds with the permanently differentiated
infrabuccal sac of certain other Hymenoptera. In Vespidae the sac
is a well-developed structural feature of the preoral cavity, completely
closed except for a narrow, transverse slitlike opening (fig. 9 D, JbS)
beneath a broad suboral fold (bib) corresponding with the bib of the
honey bee.
The preoral feeding mechanism of the wasps, as described by
Duncan (1939) in Vespula, is similar to that of the bee, except for
the more highly developed infrabuccal sac. The hypopharyngeal rods
are shorter and thicker in Vespula (fig. 9 D, E, s) than in Apis, and
are termed by Duncan the “labral tractors” because when the
proboscis is extended they pull down on the long basal angles of the
labrum (FE, Lm) and thus open the mouth. The labrum of the wasp,
however, is entirely detached from the edge of the clypeus (C/p) and
is retracted beneath the latter. The epipharynx of the wasp, accord-
ing to Duncan, has no muscles; in the bee it is provided with a group
of muscle fibers arising on the clypeus (fig. 10 C, 25). A short branch
of each hypopharyngeal rod in Vespula (fig. 9 D, E, x), the “trac-
toral suspensor” of Duncan, goes laterally to the base of the mandible,
and is suggestive of being the mandibular arm of the lateral hypo-
pharyngeal sclerite of generalized insects, on which the hypopharyngeal
muscle of the mandible, when present, is attached.
Just within the mouth on the floor, or posterior wall, of the entrance
to the sucking pump is the broad oral plate (fig. 10 A, ofl), which is
commonly known as the “pharyngeal plate.” There can be little
doubt, however, that the plate in question is a hypopharyngeal struc-
ture, since from its proximal angles two long arms (A, B, C, y)
extend upward and anteriorly in the lateral walls of the pump and
give attachment to muscles arising on the frons (B, C, 32, 33).
Similar though shorter rods giving insertion to frontal muscles are
present in most insects, and usually are prolongations from hypo-
pharyngeal sclerites; the rods, together with their muscles, in fact,
constitute the adductor apparatus of the hypopharynx. In some cases
the pharyngeal rods are processes of a plate on the base of the
NO. 2 HONEY BEE—SNODGRASS 29
hypopharynx, but the structure as developed in the bee (B) is char-
acteristic of the Hymenoptera.
In the honey bee the food glands that furnish the so-called royal
jelly open by pores on the distal angles of the oral plate (fig. 10 B, 0).
It may be noted incidentally, therefore, that these glands are hypo-
pharyngeal glands and not “pharyngeal” glands as they are generally
Fic. 9.—The preoral food channel and the mouth region in Apis and Vespula.
A, base of protracted proboscis of worker bee, anterior view, exposing the
epipharynx, mouth, bib, and food channel between lacinial lobes of maxillae.
B, same, base of proboscis retracted, food channel closed by lacinial lobes pressed
against epipharynx. C, cross section through distal part of proboscis. D,
mouth region of Vespula maculata (L.), anterior wall of sucking pump removed,
showing oral plate (0f/), suboral bib (bib), and closed entrance to infrabuccal
sac (JbS). E, same in natural condition, ventral view, mouth closed to a
narrow slit (Mth) above closed entrance to infrabuccal sac (JbS), overhung
by labrum (Lm).
c, anterior articulation of mandible; s, suspensorial rod of hypopharynx;
a, mandibular branch of s; y, pharyngeal arm of oral plate.
termed. Their secretion discharged upon the hypopharyngeal bib
evidently accumulates in the food channel on the base of the proboscis,
where it becomes accessible to other adult bees. The open food
channel thus serves as a feeding trough for the queen and the drones
when the latter are being fed by the workers. The muscles attached
on the pharyngeal arms of the oral plate would appear to give move-
ments to the oral plate and the bib that might facilitate the expulsion
of the royal jelly into the feeding trough, but the plate with its arms
30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
and their muscles is present in the drones and the queen of the
honey bee, and also in other Hymenoptera that do not have food
glands.
The sucking pump.—tThe organ by which the bee draws liquids
through the closed canal of the proboscis and into the mouth and
passes them on to the oesophagus is a large sack with strongly
muscular walls lying in the head. This sack, the sucking pump (fig.
10 C), extends from the mouth upward and posteriorly (A) to the
level of the foramen magnum, where its narrowed upper end rests
on the tentorial bridge (TB) and passes into the slender oesophagus
(Oe), which goes posteriorly through the foramen.
The sacklike sucking pump (fig. to A, C), though its outlines are
continuous, is differentiated by structural features into two parts, a
larger upper part (A, Phy) ensheathed in circular muscle fibers,
and a smaller lower part (Cb) with a strong musculature of dilator
and compressor fibers on its anterior wall (C). The upper part
unquestionably is the pharynx: the frontal ganglion (A, FrGng)
lies against the lower part of its anterior wall, the arms of the oral
plate (¥) traverse its lateral walls, its dilator muscles (dlphy) arise
on the frontal region of the cranium and are inserted above the
frontal-ganglion connectives—all of which characters, together with
the ensheathment by circular muscle fibers, are diagnostic of the
pharynx. The lower part of the organ (A, Cb) has five pairs of
large bundles of dilator muscle fibers attached on its anterior wall
(C, 26-30), and these muscles arise on the clypeus. This part of the
pump, therefore, must be derived from the cibarium of generalized
insects, which is a pocket of the preoral cavity between the under
surface of the clypeus and the base of the hypopharynx. The hypo-
pharyngeal floor of the cibarial section of the pump in the bee is
represented by the oral plate (op/). In addition to the dilator muscles
the cibarium is provided with thick bands of compressor fibers (C,
31) running obliquely between the dilator bundles from the side of
the oral plate to the median area of the anterior wall, where they turn
dorsally. In addition a strong anteriormost group of compressor
fibers arches over the mouth. In Hymenoptera, therefore, the sucking
pump is a cibario-pharyngeal structure; in Hemiptera and Diptera
the pump is purely cibarial.
The capacious pharyngeal section of the pump has no effective
dilator apparatus, since only three pairs of small muscles (fig. 10 C,
34, 35, 36) attach on it from the head wall. On its posterior surface
two groups of long slender fibers (38), arising laterally on the ten-
torial bridge (TB), spread downward, going beneath the circular
NO. 2 HONEY BEE—SNODGRASS 31
fibers to attach on the arms of the oral plate; and finally, a slender,
fusiform, unpaired median muscle (37), arising by a long tendon on a
median point of the tentorial bridge, stretches along the posterior wall
and attaches on the oral plate itself.
Slv(Syr)
“bib
-Gils
Fic. 10—The sucking pump.
A, median vertical section of the head, diagrammatic, showing sucking pump
composed of cibarial and pharyngeal sections (Cb, Phy), and salivary syringe
(Syr) in base of labium. B, oral plate, its pharyngeal arms and their muscles,
and the suboral bib, flattened, anterior view. CC, the sucking pump and its
muscles, lateral view.
0, orifice of brood-food gland on distal angle of oral plate; y, pharyngeal arm
of oral plate.
3
32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
A discussion of the working mechanism of the sucking pump with-
out observations on the action of the organ in a living bee could be
but little convincing, and will not be attempted here, except to point
out that the cibarial section, with its strong equipment of dilator and
compressor muscles, must be the active pumping apparatus, or true
suctorium. With the bee, however, an important function of the feed-
ing mechanism is regurgitation. It is probable, therefore, that the
pump is capable of reversing its action according as the occasion
demands an intake or an output of food material.
II. THE PROTHORAX
The prothorax of the bee, as of most other clistogastrous Hymenop-
tera, is not an anatomical unit because of the close association of
the pronotum with the pterothorax, and the independence of the
prothoracic pleurosternal complex, or propectus, which serves as a
suspensorium for the first legs and as a support for the head.
The pronotum.—The notal plate of the prothorax is a collarlike
sclerite closely fitted on the anterior end of the mesothorax (fig. 15,
N,). Since its tapering lower ends meet on the venter, though they
do not unite (fig. 11 A), the pronotum forms a sclerotic annulus inter-
polated between the pleurosternal parts of its own segment and the
mesothorax. The lateral areas of the pronotum are produced pos-
teriorly into a pair of spiracular lobes (spl) that cover depressions
of the mesothorax and conceal the first spiracles on their inner
surfaces. The posterior pronotal margin dorsally is sharply and
deeply inflected with the anterior margin of the mesonotum (fig.
11 F); from the latter depends a small prephragma (1Ph), but the
pronotum is separated from the base of the phragma by a narrow
line of intersegmental membrane (Mb).
The propectus—The pleurosternal region of the prothorax is
merged with the neck in a prenotal cone (fig. 15), which is partly
overlapped posteriorly by the pronotum and tapers forward to its
connection with the foramen magnum on the back of the head. The
dorsal wall of the cone is the dorsal neck membrane, which is un-
interrupted from the head to the pronotum; the lateral and ventral
walls contain the pleural and sternal plates of the prothorax, which
together constitute the propectus (fig. 11 G).
Each prothoracic pleuron is an elongate triangular plate (fig. 11 C),
the anterior apical part of which tapers into a mesally bent occipital
process (C, E, e) that articulates with the corresponding occipital
condyle of the head (B), and which, therefore, would appear to
NO. 2 HONEY BEE—SNODGRASS 33
represent a cervical sclerite fused with the pleuron. On the lower
posterior angle of the pleural plate is articulated the coxa of the first
leg (C, Cx). From the coxal articulation there extends upward close
to the posterior pleural margin a groove (pls), which is the pleural
sulcus, since it runs into the base of the large internal pleural
apophysis (PIA). The groove sets off a very narrow posterior
epimeral strip (Epm) of the pleuron from a large anterior episternal
region (Eps).
Internally the episternum presents close to its upper margin a wide,
shelflike horizontal apodeme (fig. 11 E, c) inflected from a sub-
marginal groove on its outer surface (C, b). The horizontal apodeme
expands anteriorly into a large mesal lobe (C, E, d), termed by
Duncan (1939) in the wasp the “posterior ramus of the occipital
process,’ but which, since it gives attachment to several muscles that
are in most insects inserted on a distinct neck sclerite, may ap-
propriately be named the cervical apodeme. The cervical apodeme and
the pleural apophysis form prominent points of muscle attachments
at opposite ends of the pleuron.
The prosternum lies ventrally between the pleural plates and the
bases of the coxae (fig. 11G). It is differentiated by a narrow
constriction into a wide, triangular, anterior basisternal region (Bs)
flanked by the episterna, and a narrow, elongate, posterior furcasternal
region (Fs) lying between the coxae. The furcasternum bends up-
ward at a steep angle from the horizontal basisternum (1), and carries
an elaborate endoskeletal structure, the prothoracic endosternum.
Externally the furcasternum is featured by a median groove (K, f),
* and by a pair of posterior pits (g) that mark the roots of the sternal
apophyses.
The prothoracic endosternum.—The large endoskeletal structure
supported on the sternum of the prothorax occupies much of the in-
terior of the segment (fig. 11 B, E, Endst) and gives attachment to
various muscles (fig. 12 B, C). Structurally the endosternum includes
a supporting base on the furcasternum, a pair of laterally divergent
dorsal wings, and a horizontal bridge between the wings. The sternal
support consists of a vertical median plate (fig. 11 I, h) inflected
from the median groove of the furcasternum, which posteriorly is
reinforced by a pair of lateral thickenings (7) arising from the
posterior furcasternal pits, and anteriorly tapers into a low ridge that
runs out on the basisternum (j). The endosternal wings are a pair of
plates with thickened anterior margins diverging from the basal ridge
(J, k, k) in a forward direction because of the upward slant of the
furcasternum in the natural position (I). The bridge is a wide,
Fic. 11—The prothorax (drone).
A, pronotum, anterior view. B, pleurosternal parts of prothorax and horizontal section
of attached head, dorsal view. C, propleuron and base of first leg, left side. D, articula-
tion of lateral arm of endosternum with propleuron, dorsal view. EE, pleurosternal and
endosternal skeleton of prothorax, dorsal view. F, median section of pronotum and ad-
jacent part of mesonotum, showing first phragma (Ph) to be a prephragma of mesonotum,
separated from pronotum by narrow membrane (Mb). G, pleurosternal parts of pro-
thorax and bases of first legs, ventral view. H, prosternum and endosternum, posterior
view. I, same, natural position, lateral view. J, same, anterodorsal view. K, same,
ventral view, furcasternum horizontal.
b, submarginal groove of episternum; c, horizontal apodeme of episternum along line
of external groove b; d, cervical apodeme of episternum; e, occipital process of e isternum;
f, median groove of furcasternum; g, posterior pit of furcasternum; h, median basal plate
of endosternum; 4, thickened posterior edge of wing plate of endosternum; j, median ridge
of basisternum; k, k, divergent wing plates of endosternum; /, lateral extremity of en-
dosternum; m, supraneural bridge of endosternum; mn, apodeme of muscle 5; ’, base of n,
cut off; 0, attachment lobe of muscle 52.
(2a)
a
NO. 2 HONEY BEE—SNODGRASS 35
scoop-shaped plate, broad anteriorly, tapering and rounded posteriorly
(1, J, m), set approximately horizontally on the upper edges of the
wing plates. The space between the wings is thus converted by the
bridge into a short channel, which gives passage to the nerve cords,
and hence may be termed the neural foramen (H, J, K, nf). The
lateral extremities of the bridge (/) are closely articulated with the
posterior angles of the horizontal apodemes of the episterna (D, E).
The prothoracic endosternum of the bee does not have the structure
of an ordinary Y-shaped sternal “furca,” in which the divergent arms
are the primary sternal apophyses carried inward by a median sternal
inflection that forms the stalk of the fork. In the bee the posterior
lateral thickenings that brace the median plate (fig. 11, H, I,7) appear
to be independent ingrowths from the furcasternum and evidently
represent the usual pair of sternal apophyses, which, while retaining
their bases on the sternum, have united with the posterior end of the
median plate. The supraneural bridge may be regarded as a secondary
formation produced by the union of flat mesal outgrowths from the
divergent wing plates.
Various aspects of the prothoracic endosternum are shown in figure
11. At B and E the endosternum is seen in place within the prothorax,
with its lateral extremities articulating against the pleura. H gives a
posterior view looking through the neural foramen, only the posterior
end of the upturned furcasternum being visible. At I the endosternum
and sternum are shown from the left side, the basisternum being
horizontal, the furcasternum steeply ascending. J gives an antero-
dorsal view of the same parts, showing the endosternal wings diverg-
ing upward from the median basal ridge and united by the supraneural
bridge. At K the sternum and endosternum are shown from below
as seen when the furcasternum is turned horizontally.
The prothoracic muscles—The musculature of the prothorax com-
prises (1) muscles that move the head directly, (2) dorsal and ventral
intersegmental muscles between the prothorax and the mesothorax,
(3) muscles that move the propleura, or the entire propectus, which
probably effect indirectly movements of the head, and (4) muscles
of the legs.
The direct muscles of the head are attached on the margins of the
foramen magnum, not in the usual manner on a postoccipital ridge.
They include four pairs of levator muscles and one pair of depressor
muscles. Two pairs of the levators take their origins on the pre-
phragma of the mesothorax (fig. 12 A), one pair (40) being median,
parallel, and inserted on the head laterad of the apical notch of the
foramen magnum (B, 40), the other pair (A, 41) lateral in origin,
36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
convergent beneath the first, and inserted in the apical notch of the
foramen (B, 41). The third pair of levators are large three-branched ©
muscles arising by broad bases on the episterna (D, 42), one branch
of each muscle (42a) above the horizontal apodeme of the episternum
(c), the second (42b) on the side below the apodeme, the third (42c)
on the ventral expansion; the three branches converge to a strong
tendon attached in the laterodorsal notch of the foramen (B, 42a,
Fie. 12—Muscles of prothorax (drone).
A, dorsal muscles, ventral view. B, ventral and lateral muscles, dorsal view.
C, muscles of endosternum and fore coxae, posterior view. D, branches of
cephalic muscle 42 arising on inner face of proepisternum.
c, horizontal apodeme of episternum; d, cervical apodeme of episternum;
e, occipital process of episternum; m, supraneural bridge of endosternum.
only one branch, on the left side, shown in this figure). The fourth
pair of head levators arises posteriorly on the dorsal surface of the
supraneural bridge of the endosternum (B, C, 43), the fibers of each
muscle converging to a long, thick tendon attached on the head just
laterad of the pleural levator (B, 43). The depressors of the head ~
are a pair of large two-branched muscles (B, 44) arising on the ~
NO. 2 HONEY BEE—SNODGRASS 37
supraneural bridge beneath the endosternal levators (43), and inserted
by strong tendons on the lower margin of the foramen magnum.
Duncan (1939) describes the two branches of this muscle in Vespula
as first and second depressors of the head, but adds that the two
might be considered subdivisions of a single muscle.
Fic. 13.—The prothoracic leg (drone).
A, left half of propleuron, and left fore leg, lateral view. B, trochanter and
base of femur of left leg, posterior view. C, base of right leg and coxal muscles,
mesal view. D, base of right coxa with muscles, dorsal view. E, base of left
trochanter with trochanteral muscles, dorsal view. FF, base of right femur,
anterodorsal view. G, left coxa with lateral muscles, lateral view. H, right
trochanter and base of femur with muscles, anterior view.
c-d, axis of pleurocoxal articulation; e, occipital process of episternum;
f, anterior coxo-trochanteral articulation; g, posterior coxo-trochanteral articu-
lation; h, dorsal trochantero-femoral articulation; 7, base of depressor apodeme
of trochanter.
38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
The dorsal and ventral intersegmental muscles of the prothorax
are, respectively, intertergal and intersternal. The dorsal muscles are
two large bundles of fibers (fig. 12 A, 45) arising anteriorly by broad
bases on the pronotum (fig. 11 F), and tapering somewhat to their
posterior attachments on the prephragma of the mesonotum (1Ph).
The contraction of these muscles would appear to depress the pro-
notum in the bee, though in Vespula two pairs of corresponding muscles
are regarded by Duncan (1939) as retractors of the mesonotum in
opposition to the dorsal muscles of the mesothorax. The ventral in-
tersegmental muscles (fig. 12 C, 52) have their anterior attachments
on a pair of small lobes of the’ apophyseal components of the pro-
thoracic endosternum (fig. 11 H, 0), and their posterior attachments
on two small anterior tonguelike processes of the supraneural bridge
of the pterothoracic endosternum (figs. 22C, 24A, 52). These
muscles are evidently retractors of the propectus.
The muscles of the propleura comprise tergal and endosternal
muscles inserted on the anterior parts of the episterna, and tergal
muscles inserted on the extremities of the pleural apophyses. The
largest of the propleural muscles (fig. 12 A, 48) arise close together
on the posterior inflection of the pronotum and diverge forward above
the dorsal muscles of the head to the anterior ends of.the horizontal
apodemes of the episterna. A second, slenderer pair (46) arises
laterally on the prephragma of the mesonotum and converges an-
teriorly to the cervical apodemes (d). Inserted close to the latter
muscles on each side is another small muscle (47), which may be a
branch of 48 since its fibers appear to follow those of this muscle
from the pronotum. A fourth pair, two large sternopleural muscles
(fig. 12 B, 5r), arises from the long, straplike apodemes of the
supraneural bridge of the endosternum (fig. 11 E, 1), and diverges
forward to the cervical apodemes of the pleura (fig. 12 B, d). The
tergopleural muscles inserted on the pleural apophyses include two
fan-shaped muscles on each side (fig. 12 C, 49, 50) arising anteriorly
on the side of the pronotum, and converging posteriorly to the upper
extremity of the apophysis. These muscles clearly are protractors
of the propectus, their antagonists being the ventral intersegmental
muscles (52).
The strength of the anterior pleural musculature of the prothorax
suggests that movement of the pleural plates has some important
purpose. The head, being articulated on a transverse axis between the
occipital processes of the episterna (fig. 11 B), can have only a tilting
movement on the thorax by the action of its own levator and depressor
muscles ; but the pleural plates, by reason of their lack of fixity on the
NO. 2 HONEY BEE—SNODGRASS 39
sternum, their posterior articulations on the endosternal arms, and the
shortness of the occipital axis between their anterior ends (fig. 11 B,
e), are well adapted to give partial rotary movements to the head by
a differential, or antagonistic, action of the right and left sets of
muscles attached on their anterior parts.
The prothoracic leg muscles will be described in connection with the
structure of the legs.
B G an\H
Fic. 14.—The antenna cleaner and the tibio-tarsal joint of the fore leg.
A, left fore leg of worker bee, showing antenna cleaner at base of first
tarsomere (Btar). B, antenna cleaner open, anterior view. C, antenna cleaner
closed. D, tibio-tarsal joint with muscles exposed by removal of anterior wall
of tibia. E, base of first tarsomere with muscles, anterior view. F, antenna
cleaner of Halictoides calochorti Ckll. G, same of Vespula maculata (L.).
H, same of Trogus vulpinus (Grav.). I, same of Orussus sayi Westw.
a, closing lobe (fibula) of antenna cleaner; b, tarsal notch of antenna cleaner ;
c, basal angle of notch; d, distal angle of notch; e, anterior lobule of closing
lobe; f, stop-process of tibia; m, tibial articular process; m, tarsal articular
process.
Ill. THE PTEROTHORAX
The wing-bearing section of the trunk in the clistogastrous
Hymenoptera (fig. 15) is composed of the three body segments that
are the mesothorax, the metathorax and the first abdominal segment
of other insects. Though the notal plate of the prothorax (N;) is
closely associated with the pterothorax, and looks like a collar on the
40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I03
front of the mesothorax, its connection with the latter is for its own
security, since the pronotum serves principally for the attachment of
prothoracic and head muscles, and takes no part in the pterothoracic
mechanism. The pterothoracic segments, on the other hand, are
intimately united to form a mechanical unit supporting the wings,
the first and second legs, and the abdomen. Between the third ptero-
thoracic segment, known as the propodeum, and the abdomen is a
deep constriction, which allows the abdomen to be freely movable on
the thorax.
On the dorsum of the pterothorax the segmental plates are well
defined, and behind the pronotum (fig. 15 Ni), the mesonotum (N2),
the metanotum (N;), and the propodeal tergum (JT) can readily
be identified. On the venter (fig. 16F) the sternal areas of the
mesothorax (.S.) and the metathorax (S;) are separated by a distinct
groove between the bases of the middle legs, and the propodeal
sternum (JS) is an independent sclerite behind the third legs, but
laterally there is no demarcation between the sternal and pleural areas
in either the mesothorax or the metathorax (fig. 15). The structure
of the pleural region of the pterothorax is difficult to interpret, be-
cause neither the grooves nor the areas distinctly set off by the latter
appear to conform with the pleural pattern of generalized insects.
The mesonotum.—The back plate of the mesothorax (fig. 15, Ne)
covers the strongly convex upper and anterior parts of the ptero-
thorax, and is completely divided by a transverse fissure (sf) into two
plates. For practical description the two plates are commonly called
the “scutum” and the ‘“‘scutellum,” but the line of division does not
conform with the sulcus that separates the scutellum from the scutum
in generalized insects. The true scutoscutellar sulcus in the bee is
marked internally by a strong, arched notal ridge (fig. 16D, NR),
which on the middle of the back is coincident with the groove between
the two mesonotal plates, but laterally follows a groove (fig. 15, ns)
that curves posteriorly and sets off before it on each side an elongate
oval or lenticular area (sct) which properly belongs to the scutum.
The mesonotum of the bee and of related Hymenoptera, therefore, is
secondarily divided into two major parts without regard to the original
differentiation into scutum and scutellum by the external sulcus (ns)
of the notal ridge. The true scutellum forms the prominent swelling
on the highest part of the back (Scl.) and tapers laterally to the
posterior angles of the mesothoracic wing bases; the scutum (Sct. and
sct) covers the entire anterior rounded surface of the back between
the notal sulcus (ms) and the pronotum, and supports the greater
NO. 2 HONEY BEE—SNODGRASS 4I
part of the wing bases on its sides. In a study of the mechanism of
the thorax rather than its morphology, however, it will be better to
follow the mechanical pattern of the structure, in which the meso-
notum consists of an anterior notal plate and a posterior notal plate.
The reason for the division of the notum into these two plates will
be clear when we study the wing mechanism.
Fic. 15.—The thorax and the base of the abdomen (drone), left side, wings
removed.
é, occipital process of proepisternum; Epm, epimeron of mesopleuron; Epsi,
prothoracic episternum; Eps: mesothoracic episternum; 1, groove between
mesopleuron and metapleuron; //S, sternum of first abdominal segment; JIT,
IIIT, tergal plates of first and second abdominal segments; JT, tergal plate of
propodeum; /, groove between metanotum and propodeum; Li, Le, Ls, legs;
m, n, groove between metapleuron and propodeum; Ni, pronotum; N2, meso-
notum; N;, metanotum; ns, primary notal sulcus between scutum and scutellum;
Pl:, pls, upper and lower parts of metapleuron; f/sz, pleural sulcus of meso-
thorax; S:, mesosternum; S3, metasternum; Scl2, scutellum of mesonotum;
Sctz, scutum of mesonotum; sct, posterolateral subdivision of mesoscutum;
sf, scutal fissure; spl, spiracular lobe of pronotum; Tg, tegula.
The anterior mesonotal plate (figs. 15, 16 C, Sct) is a large, shield-
shaped sclerite (fig. 16B) convex dorsally and strongly declivous
anteriorly. Its front margin is deflected into the relatively small
prephragma (rPh), and on its lateral margins are the anterior notal
wing processes (ANP). The posterior, scutoscutellar mesonotal
plate (figs. 15, 16C, Scl, sct) is somewhat crescent-shaped as seen
from above (fig. 16A) or below (D) with the concave margin
anterior; internally it is braced by the strong scutoscutellar notal
ridge (D, NR). The lateral scutal areas of the plate bear the
posterior notal wing processes, and the tapering ends of the scutellar
42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
area give attachment to the axillary cords of the wing bases. The
posterior margin is deeply inflected laterally in the groove before the
metanotum, and is here connected with the arms of the mesothoracic
postphragma (fig. 16C, D, G, 2Ph).
The phragmata of the insect thorax are primarily infoldings or in-
growths of the body wall on the intersegmental lines between the
successive notal plates, but in the bee the first phragma is solidly
united with the anterior margin of the mesonotum, and thus becomes
a mesothoracic prephragma, while the close association of the second
phragma with the posterior margin of the mesonotum gives this
phragma the status of a mesothoracic postphragma. A third phragma,
usually present between the metanotum and the first abdominal seg-
ment, is absent in the bee. The phragmata serve to give increased
surfaces of attachment for the intersegmental dorsal muscles, but
when two successive phragmata are united with opposite ends of the
same segmental plate, the muscles become intrasegmental in their
action.
The prephragma of the mesothorax of the bee is a simple, crescent-
shaped ingrowth from the deflected anterior margin of the scutum
(fig. 16B, zrPh), separated by a narrow line of intersegmental
membrane (fig. 11 F, Mb) from the overlapping pronotum. The
postphragma, on the other hand, is a large U-shaped sclerite (fig.
16 D, 2Ph) projecting far back under the metanotum into the cavity
of the propodeum. Its only connections with the mesonotum are
the attachments of its basal arms with the lateral ends of the scutellum.
The base of each arm splits into an outer and an inner lamella (G) ;
the inner lamella (v) is continuous with an infolding of the meso-
scutellar margin, the outer lamella (w) is reflected back into the
anterior margin of the metanotum. The postphragma thus maintains
its intersegmental relations, but there is in the bee no representative
of the usual postscutellar supporting plate, or postnotum, other than
the exposed parts of the inner lamellae of the phragma where the latter
join the mesoscutellum.
The metanotum.—tThe back plate of the metathorax is a narrow
transverse sclerite (fig. 16E), constricted mesally and expanded
laterally, closely interpolated between the mesonotum and the pro-
podeum (fig. 15, N;). The metanotum shows no division into
scutum and scutellum, but its widened lateral parts form a prominent
triangular area on each side. From the lateral margin of each tri-
angular area there is deflected an irregular, semidetached sclerite
(fig. 21 E, m3), and before the latter is a smaller independent sclerite
(d), both of which sclerites give support to the wing base, as will
NO. 2 HONEY BEE—SNODGRASS 43
be described later. The metanotum is separated from the propodeum
by a prominent groove (fig. 15, 7), which forms a low ridge internally,
but there is no metathoracic postphragma.
The propodeum.—The dorsal region of the propodeum (fig. 15,
IT) is a large, strongly convex plate covering the posterior part of the
pterothorax, containing the spiracles of its segment laterally, and
posteriorly descending steeply to the petiole of the abdomen. This
plate, commonly termed the “propodeum,” is really the propodeal
tergum, the sternum of the propodeal segment being a narrow ventral
plate behind the bases of the third legs (fig. 16 F, JS).
The pterothoracic pleura.—The pleuron of a wing-bearing segment
in most insects is differentiated into an anterior episternum and a
posterior epimeron by a groove, the pleural sulcus, which extends
upward from the coxal articulation to the wing base, and forms in-
ternally a strong pleural ridge, which bears an apodemal process, or
pleural apophysis, associated with the extremity of the endosternal
arm of the corresponding sternum. In the bee this structural pattern
of the pleuron is obscured in both the mesothorax and the metathorax.
The mesopleural area of the honey bee is well defined anteriorly
and posteriorly by deep grooves separating it from the prothoracic
notum in front and the metathoracic pleuron behind (fig. 15), its
“free” upper margin borders the subalar membrane, ventrally, how-
ever, as already noted, the lateral sclerotization of the segment is
continuous to the midline of the venter (fig. 16 F), and there is noth-
ing to mark the transition from pleuron to sternum, except the
ventral attachment of the huge dorsoventral muscle of the meso-
thorax (fig. 16 C, 72), which is assumed to arise on the sternum.
The upper part of the mesopleuron is crossed by an oblique groove
(fig. 15, pls2), which appears to be the pleural sulcus, separating a
small upper epimeral area (Epm) from a large lower episternal
area (Eps). The groove contains near its posterior end a conspicuous
pit (fig. 17 A, pla) that forms internally a small pleural apophysis
(B, PIA) intimately associated with the mesothoracic arm of the
pterothoracic endosternum (G, PIA, SA;). Furthermore, the line
of the posterior part of the groove is marked internally by a strong
ridge (B, PIR) bearing the apophysis. These several features almost
certainly identify the oblique groove (A, pls) as the true pleural
sulcus of the mesothorax. Beyond the apophysis the sulcus merges
with the intersegmental groove (fig. 15,7) that runs downward to the
pleural articulation of the middle coxa. A second groove of the
pleuron (fig. 17 A, e) arises near the middle of the pleural sulcus,
runs upward and posteriorly through the epimeral region, cutting the
Fic. 16.—Dissection of the pterothorax (drone).
A, mesothorax and wing bases, dorsal view, with most of scutal area removed exposing
ends of longitudinal and dorsoventral muscles. B, major scutal plate of mesothoracic notum,
ventral view. C, pterothorax with pleural and propodeal walls removed on left side, showing
phragmata and muscles. D, posterior notal plate of mesothorax and attached second phragma,
ventral view. E, metanotum, dorsal view. F, ventral surface of thorax, legs and propectus
removed. G, left lateral arm of mesoscutellum and corresponding stalk of second phragma,
lateral view. H, ventral half of pterothorax, dorsal view, showing endoskeletal structures.
f, posterior marginal ridge of mesopleuron; 1, conjunctiva between mesopleuron and meta-
pleuron; &, anterior inflection of metapleuron; m, groove between metapleuron and propodeum};
n, line of external groove n (fig. 15); ns, notal sulcus; 0, ridge of external groove m (fig. 15);
p, process of mesosternal ridge giving attachment to muscle 58 (fig. 22C); q, process of supra-
neural bridge of endosternum giving attachment to muscle 52 (fig. 22C); r, lobe of ridge o
giving attachment to basalar muscle of metathorax (fig. 21H, Ior); s, median ridge of
metasternum; sf, scutal fissure; ¢, median process of propodeum giving attachment to
muscles 96 (C, and fig. 27C); u, v, outer and inner lamellae of basal stalk of second phragma;
w, articular process on stalk of second phragma for axillary lever.
(AA)
Anaya ie
c
i ae
aa
NO. 2 HONEY BEE—SNODGRASS 45
latter into two parts (Epm, epm), and then turns downward close
to the posterior epimeral margin, and, along with the pleural sulcus,
finally becomes lost in the intersegmental groove. This recurrent
groove also forms a strong internal ridge (B, f), and it too has been
regarded as the pleural sulcus (see Snodgrass, 1925), but it has none
of the distinctive features of the latter. Duncan (1939) gives the
same interpretation of the pleural structure in Vespula as is given
here for Apis. The obliteration of the lower part of the mesepimeron
makes it appear that the middle coxa (fig. 15) has an intersegmental
pleural articulation, though its ventral articulation is with the
mesosternum (fig. 16 F).
Anteriorly the mesopleuron presents a deep concavity (fig. 17 A,
c), which is covered by the spiracular lobe of the pronotum (fig. 15
spl) ; its dorsal margin forms a long elevation, on the anterior part of
which (fig. 17 A, d) articulates the basalar sclerite (fig. 19 B, Ba),
and on the posterior part (WP) the second axillary of the wing base
(2Ax). Behind the wing process (WP) the descending margin of
the epimeron is inflected mesally in a wide plate (fig. 17 B, Epm’)
supported on the ridge (f) of the recurrent suture. The inner edge
of this inflected plate carries an elongate sclerite (G, h), attached
posteriorly to the base of the postphragma (fig. 19 1), which supports
the small subalare (Sa).
The inner surface of the mesopleuron presents an unusual structure
(fig. 17B). From its anterior and upper margin, between the first
spiracle and the wing process, there is reflected posteriorly a large
plate (g), the lower part of which is bridged to both the pleural
ridge (PIR) and the recurrent ridge (f) by a strong arch. Between
this plate and the outer pleural wall is a flat pocket open above and
below the bridge (as indicated in the figure by an arrow), which
lodges one of the muscles of the third axillary sclerite of the wing
base (E, 76a). Behind the upper part of the pocket is seen the in-
flected external plate of the epimeron (B, Epm’) united with the re-
current ridge (f). The thick pleural ridge (P/F) ends posteriorly in
the small pleural apophysis (PIA).
The metathoracic pleuron is a narrow, oblique plate on the side of the
pterothorax (fig. 15), extending from the wing base above to the hind
coxa below, between the mesopleuron and the propodeum. It is di-
vided by a groove into a larger upper part (P/,) and a smaller lower
part (pl;), but the latter is continuous ventrally with the metasternum
(S3). The dorsal plate (fig. 21 C, Pl,) supports on its upper margin
the basalare (Ba), the second axillary (24x), and the small subalare
(Sa) of the metathorax; it includes, therefore, in its upper part at
46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
least, both the episternum and the epimeron, though a separating
sulcus is not present. The anterior pleural margin is inflected in a
high apodemal ridge (figs. 16H, 21 C, k); posteriorly the upper
pleural plate is limited by a deep groove (fig. 15, m) separating it
from the propodeum, which forms a strong internal ridge (fig. 16 H,
0) with which the endosternal arm of the metathorax (SA;) is fused.
From the lower part of the groove m (fig. 15) a weak impression (7)
goes posteriorly to the coxal articulation. It is evident, therefore,
that the epimeron of the metathorax, as that of the mesothorax, has
been mostly eliminated, and that the apparent intersegmental groove
(m, n) before the propodeum includes the lower part of the pleural
sulcus extending to the coxal articulation. The position of the
posterior pleural muscles behind this groove and its internal ridge
confirms the above interpretation.
The thoracic spiracles—Of the three pairs of spiracles on the
thorax those of the first pair are of particular interest because of
their size, position, and structure. They are contained in the mem-
branous inner walls of the spiracular lobes of the pronotum (fig. 17 C,
tSp), and therefore open toward the body into the depressions of
the mesopleura (A, c) beneath the lobes, the entrances to which are
guarded by dense fringes of plumose hairs on the lobe margins. The
reflected lower margin of each pronotal lobe is supported on a small,
rectangular cartilagelike plate (D, a) arising from a lobe of the
anterior margin of the mesopleuron (A, B, b). The plate is a thicken-
ing of the intersegmental membrane, and serves to prevent movement
of the spiracular lobe. The orifice of each spiracle (C, F) is a simple
opening in the spiracular membrane leading into a large tracheal trunk
(Tra) going forward into the prothorax. Above the aperture is an
oval plate (F, Op), the ventral edge of which forms the upper lip
of the aperture ; the lower lip is merely the soft edge of the surround-
ing membrane. The plate can be pulled down over the aperture by a
slender muscle (73) and thus serves as a closing lid, or operculum,
of the spiracle. The occlusor muscle arises from a small knob on
the anterior margin of the mesopleuron (C, 73), and is inserted by a
tendon in the membrane close to the anterior margin of the spiracular
orifice (F), not directly on the operculum. The simple closing ap-
paratus of this spiracle would appear to be more effective in blocking
the entrance of air into the trachea than for preventing its egress,
but the soft lower lip of the closed aperture is easily pushed inward,
and probably gives little resistance to parasitic mites seeking an
entrance.
NO. 2 HONEY BEE—SNODGRASS 47
The second spiracles lie below the wings in the membrane concealed
between the upper edges of the mesothoracic and metathoracic pleura.
They are minute apertures without a closing apparatus, and have been
WP
/
Fic. 17—The mesopleuron and the first spiracle (drone).
A, left mesopleuron, lateral view. B, right mesopleuron, mesal view. C, right
first spiracle and adjoining parts of prothorax and mesothorax, mesal view.
D, right spiracular lobe of pronotum, mesal view. E, mesopleural muscles
arising on episternum, right side, mesal view. F, right first spiracle and con-
nected trachea, mesal view. G, posterior pleural muscles of mesothorax, right
side, mesal view.
a, platelike thickening of intersegmental membrane connecting spiracle lobe
of pronotum with mesepisternum; b, lobe of mesepisternum supporting a;
c, depression of mesepisternum covered by spiracular lobe of pronotum; d, lobe
of mesepisternum supporting basalare; e, recurrent groove of mesopleuron;
f, internal ridge of recurrent groove; g, mesal inflection of mesopleuron con-
tinuous with pleural ridge (PIR); h, epimeral plate supporting subalare; ¢, point
at which mesoscutum is supported on pleuron.
observed by the writer only in the worker. The third spiracles are
those of the propodeum, and have the structure of the other ab-
dominal spiracles. Each is provided with a short occlusor muscle
(fig. 27 B, 122) stretched between the two ends of the spiracle, and
4
48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
with a long slender dilator muscle (123) arising ventrally from a small
knob on the pleural margin of the metacoxal cavity just before the
pleural coxal process (CxP3).
The pterothoracic sternal areas —The ventral surface of the ptero-
thorax is entirely sclerotized except in the propodeal region (fig.
16 F), and the sternal areas of the mesothorax and the metathorax
(S2, S;), though separated from each other by a groove between the
bases of the middle legs, are in each segment continuous with the
pleural areas (fig. 15). The lateral extent of the mesosternum can
be judged only by the areas occupied by the ventral ends of the great
pillars of dorsoventral muscles (fig. 16C, 72), which muscles in
generalized insects have their ventral attachments on the sternum;
in the bee the bases of these muscles extend well up on the sides of
the pleurosternal areas. In the metathorax there are no corresponding
tergosternal muscles, but the sternal area of this segment may be
regarded as that part of the venter lying between and before the
hind legs (F, S;). Both the mesosternal area and the metasternal
area are marked each by a median groove, the two grooves being the
lines of inflection of a large endosternal structure (H, Endst). The
sternum of the propodeum is a narrow, weakly sclerotized transverse
band (F, JS) lying in the ventral membrane between the hind legs
and the abdominal petiole. Laterally it connects with the lower
margins of the propodeal tergum behind the legs.
According to a recent interpretation of the ventral sclerotization of
the insect thorax by Ferris (1940), the apparent sternal areas of the
mesothorax and the metathorax of the bee (fig. 16 F, S:, S;) would
probably be regarded as ventral continuations of the pleura, the true
sternal parts being supposed to be entirely folded in to make the
base of the endosternum. The evidence presented in favor of this
interpretation, however, is argumentative, and does not include a con-
sideration of the dorsoventral muscles, which appear to be tergo-
sternal in their attachments among generalized insects. It also seems
to assume that morphological regions of a segment necessarily follow
shifts in the sclerotization, rather than the converse, that sclerotized
areas may expand or contract in a continuous and otherwise stationary
integument. In the chalastogastrous family Xyelidae, among the
Hymenoptera, the episternal sclerotizations of the mesothorax are
extended downward onto the ventral surface, where they are con-
tinuous across the median line in a narrow or wide bridge. Behind
the bridge, however, is a large, triangular basisternal plate with a
strong median groove, and on this sternal plate are attached the lower
ends of the dorsoventral muscles. In most other Hymenoptera the
NO. 2 HONEY BEE—SNODGRASS 49
muscle-bearing areas of the ventral sclerotization are continuous with
the episternal areas, as in the honey bee, but in some forms, as in
the tenthredinoid Arge, the two are separated by distinct sutural lines.
The pterothoracic endosternum.—tThe sternal areas of the meso-
thorax and the metathorax support together a large endoskeletal
structure similar to the endosternum of the prothorax, except that the
sides are produced into two pairs of divergent arms (fig. 16 H,
Endst) and the entire structure arises from the median inflections
of the two sterna. The pterothoracic endosternuum, therefore, appears
to be composed of two ordinary Y-shaped sternal furcae united at
their bases, which latter are stretched out anteriorly and posteriorly
in the median ridges of the supporting sternal plates. The supra-
neural bridge, however, spans the space between the bases of the
anterior arms, and thus appears to belong to the mesothoracic com-
ponent only. The long anterior, or mesothoracic, arms (SA,.) extend
laterally and upward to the sides of the thorax, where each arm ends
in a thin, forward-curved expansion that gives attachment to a large
group of short muscle fibers (79) arising on the posterior epimeral
region and on the small pleural apophysis of the mesopleuron (fig.
17G). The shorter posterior, or metathoracic, arms (fig. 16H,
SA;) go laterally and posteriorly over the bases of the second legs,
just beyond which each arm unites solidly with a thick ridge (0) fol-
lowing the external suture (m) between the metapleuron and the
propodeum (fig. 15 m). The pterothoracic endosternum gives attach-
ment to the ventral intersegmental muscles between the prothorax
and the mesothorax (fig. 22 C, 52, 58), and between the metathorax
and the second abdominal segment (fig. 27 E, rz8, 119), to muscles
of the second and third legs (figs. 22 C, 24 A), to the muscles of the
axillary levers of the mesothoracic wings, and to the tergosternal
muscles of the metathorax (fig. 21 F, 97, 98, 90).
IV. THE WINGS
The movements of an insect’s wings fall into two distinct categories:
first, there are the movements of flight when the wings are extended,
and second, the movements by which the wings are turned posteriorly
over the body when at rest, or again brought into the extended posi-
tion preliminary to flight. The action of the wing muscles that pro-
duce the movements of flight depends chiefly on structural adapta-
tions in the skeletal parts of the thorax; the effect of the muscles
concerned in the flexion and extension of the wings, on the other
hand, is mainly dependent on structural details in the wing bases.
50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
In other words, both sets of wing movements are produced by primi-
tive muscles that have become motors of the wings by modifications
in the skeletal elements, the primary changes that made flight possible
being mostly in the thoracic skeleton, the secondary adaptations for
flexion and extension being largely in the bases of the wings
themselves.
In the bee the principal muscles of flight are the huge longitudinal
and dorsoventral muscles of the mesothorax (fig. 16C). For this
B
Fic. 18.—The wings.
A, B, fore and hind wing of drone. C, hook of hind wing of worker. D, hook
of hind wing of drone. FE, section of interlocked wing margins.
f, deflected fold on posterior margin of fore wing; h, hooks on anterior
margin of hind wing.
reason probably the two wings on each side, when extended, are
attached to each other. The coupling mechanism consists of a series
of small hooks on the anterior margin of the hind wing (fig. 18 B, h)
and a fold on the posterior margin of the fore wing (A, f). The
hind-wing hooks, or hamuli (C, D), curve upward and backward
from the marginal vein of the wing with a twist in the direction of
the wing tip. The fold of the fore wing is a rounded deflection of the
posterior part of the elevated wing area behind the last vein, and
has a narrowly upturned margin (E, f). When the wings are ex-
tended preparatory to flight the fore wing is drawn over the upper
NO. 2 HONEY BEE—SNODGRASS 51
surface of the hind wing, and the hooks of the latter become engaged
in the fold of the former (FE).
The fore wing—The mesothoracic wing of the bee is much larger
than the hind wing (fig. 18) and has a stronger and more elaborate
venation. The four veins that arise at the base of the wing are
probably costa (A, C), radius (R), media-cubitus (MCwu), and an
anal, or vannal, vein (A). It is difficult to identify the branches of
these main veins in the coarse reticulation of the distal part of the
wing, but it will not be necessary for a study of the wing mechanism ;
an understanding of the basal structure of the wing, however, is of
much importance.
The base of the fore wing is overlapped by a large scalelike lobe
known as the tegula (figs. 15, 19 A, Tg), which is flexibly attached
to the margin of the scutum just before the wing. In the wasp
Vespula Duncan (1939) finds a small depressor muscle of the tegula
arising on the scutum, but the writer has observed no muscle con-
nected with the tegula in Apis.
In the naturally extended position of the fore wing, the wing base is
rolled on itself (fig. 19 A), the posterior part being turned downward
so that little of it is visible when viewed from above. If the basal
region is artificially spread out flat (D) the posterior part is seen to
be an extensive area widening toward its attachment on the body.
The whole region of the wing base presents at first sight a confusion
of small irregular parts, some of which are expansions of the wing
veins and others independent sclerites in the basal wing membrane.
A closer study, however, soon shows that there is an orderly arrange-
ment of the parts and a mechanical interrelationship between them,
which latter becomes more evident when the wing is moved.
The anterior part, or humeral area, of the wing base (fig. 19 D)
is occupied by a large irregular sclerotization, here termed the humeral
complex because it appears to include parts derived from the bases of
the first three veins and also a remnant of the subcosta, which is
suppressed in the distal part of the wing. The posterior part of the
wing base, or axillary area, contains several discrete sclerites, which
are the first, second, third, and fourth axillaries (1Ax, 2Ax, 3Ax,
4Ax), and a prominent, transversely elongate median plate (m). All
these sclerotizations are held in a basal wing membrane, bordered
posteriorly by a corrugated marginal thickening known as the axillary
cord (AxC), which is continuous on the one hand with the posterior
margin of the wing, and on the other with the lateral extremity of
the scutellum of the mesonotum (F, J, dxC).
52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
The first axillary (fig. 19 E, rAx) is an elongate sclerite with a
large anterior head (b) turned laterally on a curved neck, posterior
to which is a wide lateral projection (f) and a smaller articular lobe
(c) on the mesal margin. The sclerite is attached to the side of the
mesonotum in such a manner (J) that its neck rests on the anterior
lobe (~) of the notal wing process (I), and its articular lobe is closely
hinged to the posterior lobe (q) of the wing process, while the
tapering posterior end rests and turns in a concave surface on the
lateral margin (7) of the scutal area of the mesonotum (J. sct), where
normally (D) it is concealed in a pocket of the wing membrane be-
neath the fourth axillary (4d). In the natural position of the
extended wing (A) the first axillary (74x) stands almost vertical;
in the flattened wing base (D) it is seen that its head abuts against
the humeral complex, and the lateral projection underlaps the median
plate (m). The first axillary is the anterior hinge plate of the wing.
The second axillary (fig. 19 E, 24x) is a thick sclerite standing
vertically in the wing base. Its exposed upper surface (7) appears
as a small oval plate (D) lying in the notch between the head and
the body of the first axillary. Ventrally the second axillary articulates
by an anterior knob (E, B, 7) with the wing process of the pleuron,
and by a posterior arm (f) is associated with the subalar sclerite (B,
Sa) on the upper edge of the pleuron. The second axillary is the
pivotal sclerite of the wing base, since it is the only axillary sclerite
that articulates with the pleuron and gives the wing a solid support
from below. In the bee the second axillary has no direct connection
with any of the wing veins, though in most insects it is connected
with the base of the radius.
The third axillary (fig. 19 E, 3A) is an elongate sclerite lying
close along the axillary cord (D), with its tapering distal end as-
sociated with the enlarged base of the single anal vein of the wing.
Proximally the third axillary bears a large lobe (E, /) projecting
anterior to its axis, on which are inserted three muscles arising on
the mesopleuron (H). The small, flat uppermost muscle (H, 76a)
arises in the pocket of the inner wall of the pleuron (fig. 17 B) be-
neath the reflected plate (g) ; the other two larger muscles (fig. 19 H,
70b, 76c) arise ventrally on the episternum. The third axillary is
the only axillary sclerite of the wing base provided with muscles; it
is the effective skeletal element of the wing-flexing mechanism, and
may be termed the flexor sclerite.
The fourth axillary (fig. 19 E, 44x) is a small, irregularly tri-
angular sclerite lying transversely in the posterior angle of the wing
base (D), articulated mesally on the lateral margin of the scutal area
NO. 2 HONEY BEE—SNODGRASS 53
Gaz
Fic. 19.—Details of the mesothoracic wing base, its articulations, and pleural
muscles (drone).
A, tegula (Tg), and wing base in natural position of extended wing. B, upper part of
mesopleuron, showing pleural articulation of basalare and second axillary. C, humeral area
of wing base, ventral view. D, base of right wing with axillary region flattened, dorsal view.
E, axillaries and axillary lever of right wing. F, left margin of mesonotum in region of wing
attachment. G, right basalare and its muscle, mesal view. H, right third axillary and its
muscles, mesal view. I, right subalare and its muscle, with supporting sclerite (h) on edge
of epimeron, mesal view. J, left lateral part of mesonotum, with first axillary in natural
position.
a, articular surface of first axillary with humeral complex of wing base; b, head of first
axillary; c, notal hinge-lobe of first axillary; d, lobe of mesopleuron supporting basalare; e,
recurrent suture of mesopleuron; f, lateral process of first axillary; g, posterior end of first
axillary; h, supporting plate of subalare; 1, exposed dorsal surface of second axillary; j,
pleural articular process of second axillary; k, posterior ventral process of second axillary;
l, muscle lobe of third axillary; m, median plate of wing base; m, distal lobe of median plate;
0, articular process of axillary lever; p, anterior lobe of anterior notal wing process supporting
neck of first axillary; g, posterior lobe of anterior notal wing process articulating with hinge-
lobe of first axillary (c); r, posterior notal wing process; s, exposed head of axillary lever;
t, point at which mesoscutum is supported on anterior angle of pleuron.
54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
of the posterior plate of the mesonotum (F, J, 7), above the posterior
end of the first axillary (D). Its narrow outer end is closely pressed
against the axillary cord not far behind the proximal end of the third
axillary. The fourth axillary is the posterior hinge plate of the wing,
but in the bees it has an accessory function in connection with the
flexion of the wing by reason of its close association with a leverlike
sclerite (D, azlv) lying immediately behind it and connected with
the arm of the postphragma.
The axillary lever is an elongate triangular sclerite, as seen from
the side (fig. 20 B), lying against the mesal surface of the stalk of
the mesothoracic postphragma (A, axlv). By a small process (0) on
the ventral angle of its base, directed anteriorly, the lever articulates
with the narrowed extremity of the phragma stalk, and on its decurved
distal end is attached a relatively large, fan-shaped muscle (78) aris-
ing ventrally on the mesothoracic arm of the pterothoracic endo-
sternum. The sclerite is an internal structure except for the end of
the thick dorsal angle of its base (s), which comes to the surface in
the membrane of the posterior angle of the wing base, and appears
here externally as a small sclerite lying immediately behind the
fourth axillary (fig. 19 D, axlv). The function of the lever will be
described in connection with the mechanism of the wing.
So far as yet observed the axillary lever is a free sclerite only in
the Apoidea. In other Hymenoptera it is represented by an im-
movable arm or lobe of the postphragma, on which, however, there
is always attached a muscle from the mesothoracic endosternum.
In Sphecius (fig. 20C) and Vespula (D), for example, the lever
of Apis is so closely imitated by a long decurved arm (¢) arising by
its widened anterior end from the mesal surface of the stalk of the
phragma that there can be no question of the homology of the two
structures. In the ichneumonid Megarhyssa lunator (E) the cor-
responding arm (ft) is simply a backwardly directed process of the
phragma, and in the chalastogastrous genera Sirexr (F), Pteronidea
(G), and Cimbex (H) it is a mere process or lobe (¢) of the lateral
part of the phragma giving attachment to an endosternal muscle
(78). In these last forms the muscles evidently pull downward on
the phragma and probably depress the phragma-bearing postnotal
plate (PN.). According to Duncan (1939) the muscle of the phragma
arm in Vespula “resists the tendency of the indirect depressor muscle
of the forewing (attached on the phragma) to lift the mesopost-
phragma, and helps to retract the phragma to the position of rest
following a contraction of the indirect depressor muscle.” The de-
velopment of this structure into an accessory of the wing-flexing ap-
NO. 2 HONEY BEE—SNODGRASS 55
paratus in the bees is a highly interesting example of the evolution of
a special mechanism from a structure having originally a quite
different function.
Fic. 20.—Evolution of the axillary lever.
A, Apis mellifera L., drone, right stalk of second phragma, mesal view,
showing axillary lever (axlv) and associated fourth axillary of wing base.
B, same, left axillary lever, lateral view. C, Sphecius speciosus (Drury), right
stalk of second phragma with mesal process (t) corresponding with axillary
lever of Apis. D, Vespula maculata (L.), same parts as in Sphecius (C).
E, Megarhyssa lunator (F.), right stalk of phragma with simple muscle process,
mesal view. F, Sirex abbottii Kirby, second phragma and lateral muscle lobes
(t), anterior (ventral) view. G, Pteronidea ribesii (Scop.), mesothorax and
second phragma, lateral view, showing muscle lobe (t) with muscle from furcal
arm (S42). H, Cimbex americana Leach, posterior end of mesonotum with
second phragma, showing lateral muscle processes (¢) connected by short
muscles (78) with arms of mesosternal furca.
h, supporting sclerite of subalare; o, articular process of axillary lever;
5, exposed head of axillary lever associated with fourth axillary sclerite;
t, muscle-bearing process of second phragma of non-apoid Hymenoptera corre-
sponding with axillary lever (ax/v) of Apoidea.
The large, transversely elongate median plate in the base of the
fore wing (fig. 19 E, m) lies close behind the humeral complex
(A, D), and forms a link between the first and second axillaries at
its mesal end, and the enlarged base of the fourth wing vein at its
56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
distal end. In most insects the median plate is associated with the
bases of the veins media and cubitus, but in the Hymenoptera these
two veins appear to be combined, and united basally with the radius.
The hind wing.—The small metathoracic wing of the bee (fig. 18 B)
has a more simplified venation than the fore wing, but it has the
same vein stems in its proximal part (fig. 21 A) except that there
are two veins (7A, 24) in the vannal region. The basal structure of
the hind wing also is simpler than that of the fore wing, and there is
no metathoracic tegula. The humeral complex supports only the first
two veins; a median plate and a fourth axillary are absent.
The relatively small first axillary of the hind wing (fig. 21 A, B,
1Ax) articulates on the lateral plates of the metanotum (E), its
posterior process being received in a depression (f) of the lower
angle of the semi-detached plate (73), its head resting on a small lobe
(e) of the free plate (d). Laterally the large head of the first
axillary abuts against the broad mesal end of the humeral complex
(A). The irregular second axillary (B, 24%) lies laterad of the first
axillary (A) and articulates by a large ventral process in a notch on
the dorsal margin of the metapleuron (C). The third axillary is a
large, elongate sclerite (B, 3Ax) lying obliquely transverse behind
the second axillary (A) between the edge of the metanotum and the
base of the fourth wing vein. The large, fan-shaped, three-branched
muscle of the wing-flexing third axillary (B, F, G, H, 100) arises
on the anterior infold of the metapleuron (C, H, k) and is attached,
not directly on the third axillary, but on a minute sclerite of the wing
membrane (A, B, a) close to the base of the axillary, and hence
pulls indirectly on the latter through the intervening membrane.
The epipleurites—The epipleural sclerites associated with the under
sides of the wing bases include in each segment the basalare lying
before the pivotal second axillary, and the subalare lying behind it.
The sclerites are articulated on the upper edges of the pleura; they
give insertion to muscles that deflect them inwardly and thus pull
on the wing bases by reason of the close membranous connection of
the epipleurites with the latter. The specific effect of the epipleurite
muscles on the wing movements will be described later.
The basalare of the mesothorax (fig. 19 B, Ba) is an elongate
sclerite hinged by its entire length on a wide lobe (d) of the dorsal
margin of the mesopleuron, which supports posteriorly also the
second axillary (24%). On the inner surface of the basalare is
attached by a strong apodemal tendon a large, flat, fan-shaped muscle
(G, 77) arising on the anterior part of the pleuron (fig. 17 EF), where
it partly overlaps internally the second and third muscles of the third
NU. 2 HONEY BEE—SNODGRASS 57
axillary (76b, 76c). The contraction of this muscle turns the basalare
inward on the supporting pleural lobe, and the movement is trans-
mitted to the wing because of the close connection of the basalare
with the humeral angle of the wing base.
The subalare of the mesothorax is a small triangular sclerite (fig.
19 1, Sa) but little visible in lateral view (B) beneath the posterior
part of the wing. Its broad base is supported on a longer sclerite (I,
h), which in turn rests on the inner margin of the inflected dorsal
plate of the epimeron (Epm’), and is linked posteriorly with the
anterior extremity of the stalk of the postphragma (2Ph). On the
upper angle of the subalare is attached the tendon of a long muscle
($2), which arises ventrally on the mesothoracic coxa (figs. 17 G,
22 C, 82). The subalare is closely associated with the posterior basal
process of the second axillary sclerite (fig. 19 B), so that the pull
of its muscle is indirectly transmitted to the pivotal sclerite of the
wing base.
The epipleural sclerites of the metathorax have the same relations
to the hind wing as do those of the mesothorax to the fore wing. The
triangular basalare (fig. 21 B, Ba) sits on a marginal elevation of
the metapleuron before the second axillary (C, Ba), and the small
subalare (B, Sa) is attached to the concave pleural margin behind
the second axillary (C, Sa). A single large basalar muscle (H, ror)
arises on a lobe (7) of the ridge in the metapleural area behind the
middle leg (fig. 16 H, r), and tapers upward to a strong tendon at-
tached mesally on the basalar sclerite (fig. 21 B, H, Ba). The small,
inconspicuous subalare (B, Sa) has an elongate apodemal process
on which are inserted two muscles, one a long fusiform muscle (105),
corresponding with the subalar muscle of the mesothorax, which
arises ventrally on the hind coxa (fig. 24 A, 105), the other a broad
thick muscle (fig. 21 B, C, H, ro2) arising on the upper plate of the
metapleuron, and apparently represented in the mesothorax by a
similar pleural muscle (fig. 17 E, 75) attached dorsally on the margin
of the posterior scutal plate of the mesonotum (fig. 19 F, 75). The
subalare is intimately associated with the second axillary of the wing
base (fig. 21 B, 24x) by a close membranous connection (b) with
the posterior basal arm of the latter.
The mechanism. of flight—To understand the movements of an
insect’s wings it is necessary to observe first that the wings are sup-
ported from below on the second axillary sclerites of their bases, and
that these sclerites rest on the upper edges of the pleura, while the
flexible attachments of the wings to the notal plates lie a little mesad
of the pleural supports. As a consequence a slight depression of the
58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I03
Fic. 21.—Details of the metathorax, base of the hind wing, and the lateral
metathoracic muscles (drone).
A, base of hind wing, dorsal view. B, axillary, basalar, and subalar sclerites
of hind wing, dorsal view. C, inner face of upper part of right metapleuron,
showing pleural articulations of basalare, second axillary, and subalare, and
muscles of subalare. D, upper part of right metapleuron and adjoining part
of propodeum, anteromesal view. E, lateral part of metanotum, left side.
F, lateral muscles of metathorax, right side, mesal view. G, same, with third
tergosternal muscle (99) removed. H, same, with notum and first, second, and
third tergosternal muscles (97, 98, 99) removed, exposing basalar and subalar
muscles (ror, 102, 105) and muscle of third axillary (100).
a, nodule of wing membrane giving insertion to muscle of third axillary;
b, “ligament” of wing membrane connecting second axillary with subalare;
c, process of metathoracic endosternal arm (SA;) giving attachment to third
tergosternal muscle (99); d, free marginal sclerite of metanotum; e, lobe of d
supporting head of first axillary; f, posterior articular socket of first axillary;
k, anterior inflection of metapleuron; m, groove between metapleuron and pro-
podeum ; :, semidetached lateral plate of metanotum; o, internal ridge between
metapleuron and propodeum, r, lobe giving attachment to basalar muscle (zor).
NO. 2 HONEY BEE—SNODGRASS 59
lateral margins of the back plates of the wing-bearing segments
causes the wings, in the manner of a pump handle, but involving the
interaction of many structural details, to turn upward, and an eleva-
tion of the notal margins tilts the wings downward. The up-and-
down movements produced in this way, however, are not sufficient for
progressive flight; in addition the wings must have a partial rotary
movement on their long axes accompanied by a forward and rearward
motion, by which the anterior margins are turned downward and
forward during the downstroke, and reversed during the upstroke.
This compound motion gives a sculling movement to each wing, but
the result of the wings on opposite sides of the body acting together
is that a current of air is driven backward, and the insect propelled
forward. In the bee the two wings of each side act in unison as a
single pair of flight organs because of their connection with each
other, but the unity of action is further secured by a reduction of the
metathoracic musculature and its subordination to the action of the
highly developed mesothoracic muscles, which latter thus become the
chief motor elements for both pairs of wings.
The cavity of the pterothorax is largely occupied by the huge dorso-
ventral and dorsal longitudinal muscles of the mesothorax (fig. 16 A,
C). The thick pillars of dorsoventral muscles (72) have their upper
attachments (A) on the lateral parts of the anterior plate of the
mesonotum (B), and go downward and backward (C) to their lower
attachments on the ventral and iower lateral walls of the pleuro-
sternal region of the mesothorax. The longitudinal muscles (77) are
attached anteriorly on the prephragma and the median area of the
anterior notal plate (B), and extend posteriorly through the meta-
thorax into the posterior end of the propodeum, where they are at-
tached on the postphragma (2P/). Though the postphragma of the
bee is extended through the metathorax far back into the propodeum.
to accommodate the great length of the mesothoracic dorsal muscles,
it is morphologically an ingrowth between the mesonotum and the
metanotum. A pair of small fan-shaped muscles (C, 96), arising on a
median process of the posterior end of the propodeal tergum (/T)
and spreading to their insertions on the rear surface of the phragma,
must represent, therefore, the usual longitudinal dorsal muscles of
the metathorax with their posterior attachments transposed backward
on the propodeum. A pair of short external dorsal muscles of the
mesothorax (D, 70) traverses the scutellar area of the mesonotum.
The vertical component of the upstroke of the fore wings, resulting
from a depression of the lateral margins of the mesonotum, is effected
principally by the great dorsoventral muscles (fig. 16C, 72), but
60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
probably important adjuncts are the large pleuronotal muscles arising
on the sides of the pleuron (fig. 17 E, 75) and attached on the ends of
the posterior scutal areas of the notum (fig. 19 F, 75). The wing
bases, hinged to the back by the first and fourth axillary sclerites, are
pressed downward proximal to the fulcral second axillaries by the
descending margins of the notum, and thereby the distal parts of
the wings are thrown upward. The vertical component of the down-
stroke, conversely, results from the contraction of the longitudinal
muscles, which restores and increases the curvature of the meso-
notum, and thus turns the wings down on the fulcral sclerites.
In most insects the wing-bearing plates of the back are sufficiently
thin and flexible to respond by changes in shape to the alternating
pull of the vertical and longitudinal muscles. Such movements, how-
ever, are not possible in the thick and rigid mesonotum of the bee and
related Hymenoptera, and it is for this reason that the mesonotum is
cut by a transverse fissure (fig. 16C, sf) into an anterior and a
posterior plate. The fissure widens on each side into an open cleft
(fig. 19 J, sf), the edges of which are connected by an infolded
membrane. The closed dorsal part of the fissure acts as a hinge be-
tween the two plates. Pressure on the top of the back depresses the
notal plates at the hinge line and widely opens the lateral clefts, the
anterior notal plate here moving forward, the posterior plate back-
ward. The divergent movement of the two plates, however, ac-
centuates the downward movement of their lower margins at the
points just before and behind the lateral clefts where the wings are
articulated to the back by the first and fourth axillary sclerites. This
action of the mesonotum is easily demonstrated by manipulation of a
dead bee, and could be well illustrated with half of a hollow rubber
ball having a meridional slit on each side.
In the chalastogastrous Hymenoptera the posterolateral areas of
the mesoscutum form deep concavities of the scutal surface at the
sides of the elevated median area of the scutellum, and are separated
from the main part of the scutal plate before them by the lines of
abrupt deflections of the notal surface. These lines coincide with the
scutal clefts of the Clistogastra, and, in fact, each is split laterally
by a short cleft. The response of the relatively flat mesonotum of a
tenthredinid to gentle compression between its two ends is most
pronounced on the sides; the anterior wing processes spread laterally
and move upward and posteriorly, the lateroposterior margins of the
anterior scutal areas at the lateral clefts folding backward above the
anterior edges of the posterolateral areas. In the densely sclerotic and
strongly convex mesonotum of the Clistogastra a much deeper splitting
NO. 2 HONEY BEE—SNODGRASS 61
of the scutum has become necessary for carrying out the mechanical
function of the notum. The scutal clefts of the Hymenoptera in any
case, are simply extensions of the usual lateral emarginations of a
wing-bearing notum that are bridged by the first axillary sclerites of
the wing bases.
The opening of the scutal clefts by pressure on the back causes a
posterior displacement of the postphragma in the propodeum because
of the connection of the phragma with the scutellum (fig. 16 C), and
this movement of the phragma must stretch the longitudinal muscles
preparatory to their succeeding contraction. The contraction of the
longitudinal muscles (77), extending from the anterior notal plate
to the phragma, not only closes the lateral clefts of the notum, but
causes the edges of the posterior plate to slide beneath those of the
anterior plate, thus increasing the curvature of the back to give greater
effectiveness to the downstroke of the wings. The small propodeal
muscles (96) attached on the back of the phragma may be supposed
to contribute to the movement of the phragma, though their narrowed
and convergent posterior ends (fig. 27 C, 96) would suggest that
they pull on the propodeum. The function of the pair of delicate
muscles in the mesothoracic scutellum (fig. 16D, 70) is not clear.
The metathorax contains no muscles corresponding in function
with the dorsal muscles of the mesothorax, since the only interseg-
mental dorsal muscles that can be referred to the metathorax are the
greatly displaced pair between the postphragma and the propodeum
(fig. 16 C, 96). The dorsal muscles of the mesothorax (77), there-
fore, must cause the downstroke of both pairs of wings when the
wings of each side are connected. Probably in the same way the
dorsoventral muscles of the mesothorax are largely responsible for
the upstroke of the metathoracic wings, since as long as the wings are
connected pressure on the mesonotum alone lifts both pairs of wings,
though it has no effect on the hind wings when the latter are un-
hooked. However, the metanotum itself is amply provided with de-
pressor muscles, which very probably in the living bee play an im-
portant part in the elevation of the wings.
In each side of the metathorax there are three large muscles at-
tached on the notum, which arise ventrally on the metathoracic arm
of the pterothoracic endosternum. The first two of these muscles
(fig. 21 F, G, 97, 98) arise by broad bases on the endosternal arm
and taper upward to their dorsal attachments on the lower edge of
the semidetached lateral plate of the notum (FE, 3). The third and
largest muscle (F, 99) arises from a small lobe (c) of the endosternal
arm, and its fibers spread upward to their attachments on the lateral
62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
triangular area of the dorsal part of the notum. A downward pres-
sure on this region of the metanotum at once turns the hind wings
upward, leaving little doubt that the metanotal muscles are wing
elevators. .
The horizontal and rotary components of the wing motion, by which
the wings are turned forward with the costal margins deflected during
the downstroke, and reversed during the upstroke, are produced
chiefly by the muscles of the basalar and subalar sclerites. It is these
movements that convert the wings from mere flaps into organs of
propulsion. The mechanism involved is equally developed in each
wing-bearing segment of the bee. The epipleural sclerites, as already
explained (figs. 19 B, I, 21 C, Ba, Sa), are hinged to the upper °
edges of the pleura in such a manner that the contraction of their
muscles turns them mesally in the subalar membranes. The move-
ments of the sclerites are effective on specific points in the wing bases
because a tract of the membrane intervening between each sclerite
and the wing is so tightly stretched that it acts as a direct union, and
often gives a superficial appearance of being a connecting ligament.
The basalare in each wing of the bee is thus connected with the
humeral complex of the wing base, and the subalare with the posterior
lever arm of the second axillary (fig. 21 B). Contraction of the
basalar muscle during the downstroke of the wing, therefore, revolves
the basalare inward on its pleural hinge, and the tension exerted on
the wing base turns the descending wing forward on the pivotal
second axillary and deflects its costal margin. Similarly, during the
upstroke, the subalar muscle acting on the subalare pulls on the lever
arm of the second axillary and thus tilts the wing downward pos-
teriorly, so that the costal margin turns upward and moves backward,
relative to the body, while the wing is ascending.
Flexion and extension of the wings——When the bee alights after
flight, or when the organs of flight are not otherwise in use, as in
“fanning,” the wings are flexed posteriorly, the hooks on the hind
wings automatically loosing their grasp on the fore wings, and both
pairs of wings are laid over the back, the fore wings on top of the
hind wings. The mechanical problem involved in the horizontal flexing
of an insect’s wing may be illustrated with a piece of paper firmly
held flat at one end and bent horizontally on itself; the bending
necessarily produces a fold across the base. Conversely, if the paper
is folded in the same way, it automatically bends backward at a right
angle to its flat position. The flexing apparatus of the wing acts on
the second principle—by the production of a basal fold the wing is
turned horizontally backward. The fold-producing mechanism is the
NO. 2 HONEY BEE—SNODGRASS 63
third axillary sclerite and its muscles, but in the fore wing of the bee
the action of the third axillary appears to be assisted by the fourth
axillary and its lever.
The basal region of the fore wing of the bee in the natural position
of extension (fig. 19 A) is rolled on itself in such a way that most
of the axillary area is deflected. There is hence an oblique fold in
the wing base extending from the head of the first axillary (1A),
behind the second axillary and the median plate, to the upwardly
directed distal end of the third axillary ( 3Ax). The pull of the third
axillary muscles (H), therefore, simply revolves the sclerite mesally
and somewhat forward on its base and thus accentuates the fold al-
ready present, with the result that the distal part of the wing turns
backward. If the wing base were flat (D), as it is in most insects,
the action would not be essentially different, as can be demonstrated
by revolving the third axillary in the wing of a dead specimen.
The part that the fourth axillary plays in the flexing of the fore
wing of the bees can be shown by depressing its lever (fig. 19 D,
axlv). The fourth axillary (44%) is movably articulated on the
margin of the mesonotum and its distal end is pressed against the
axillary cord (AxC). The head of the lever lies in the wing mem-
brane immediately behind the fourth axillary. A downward pull on
the lever muscle (fig. 20 A, 78) depresses the lever on its fulcral con-
nection (0) with the end of the postphragma, and causes the head (s)
to pull backward on the fourth axillary (44x). The axillary turns
on its notal articulation and its outer end pulls on the axillary cord.
The wing gives a quick response with a slight backward movement.
Slight though this movement of the wing is, the elaborateness of the
mechanism that produces it suggests that the movement is one of
considerable importance. It may be seen, in fact, that the backward
jerk on the extended wing brings the third axillary into a more
nearly vertical position in which it becomes quickly responsive to the
action of its own muscles. The fourth axillary and its lever in the
fore wing of the bee, therefore, appear to be a starting apparatus for
the flexor mechanism.
The flexing of the hind wing is produced entirely by the large third
axillary sclerite and its muscle (fig. 21 A, B, 34%), since there is
no fourth axillary or lever accessory. The axillary region of the
hind wing, however, unlike that of the fore wing, is approximately
flat when the wing is extended (A). It is probable, therefore, that
on relaxation of the extensor muscle the wing automatically turns
backward sufficiently to raise the third axillary into a position in
which its muscle can become effective.
5
64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
The extension of the flexed wings preparatory to flight is in general
produced by the basalar sclerites and their muscles. In a dead speci-
men a slight pressure on the basalare of either wing, or a pull on its
muscle, brings the wing out at right angles to the body and deflects
its anterior margin. The fore wing of the bee, however, is provided
with a small pleuroalar muscle (fig. 19 C, 74), which arises on the
anterior marginal carina of the mesopleuron (fig. 17C, 74) just
below the attachment of the spiracular muscle (73), and extends
dorsally and posteriorly to its insertion by two branches attached
ventrally on the humeral complex of the wing base (fig. 19 C). This
muscle evidently must be an accessory wing-extensor. In some insects
a depression of the second axillary, produced by the pull of the subalar
muscle, helps to extend the wing, but an extensor action of the second
axillary cannot be demonstrated in the bee.
VV. THE:DEGS
The legs of Hymenoptera have the usual six segments of an insect
leg, namely a coxa, a trochanter, a femur, a tibia, a tarsus, and a
pretarsus. A leg segment is to be defined as a section of the limb
independently movable by muscles attached on its base, except where
there is evidence of the union of segments or a suppression of muscles.
The tarsus is usually divided into subsegments, or tarsomeres, which,
though movable on each other, are not “tarsal segments” since only
the proximal piece is provided with muscles, The flexible areas be-
tween segments or subsegments are the joints of the leg. The
proximal tarsomere, especially when differentiated by size or other
features, as in the bee, is designated the basitarsus. The pretarsus
either terminates in a single claw, or dactyl, or it bears a pair of
lateral claws, or ungues, and usually a median lobe, the arolium.
The three pairs of legs of the bee are attached at somewhat different
angles on the thorax so that in their action they are radially distributed
at the sides of the body. The coxae of the fore legs hang downward
on transverse axes of movement (fig. 11 G, Ca), and hence the
first legs swing forward and backward in the transverse planes of the
coxae. The middle coxae are transversely elongate and set obliquely
into the under wall of the thorax (figs. 15, 16 F), and the trochanters
are attached at their mesal ends; the middle legs, therefore, are
limited in their up-and-down motion, and swing forward and back-
ward by the revolution of the coxae on the body. The hind coxae
also have obliquely transverse axes on the thorax (figs. 15, 16 F),
but they are directed posteriorly and hence give the hind legs a more
NO. 2 HONEY BEE—SNODGRASS 65
backward position at the sides of the abdomen. Each leg has char-
acteristic features, and, in the worker particularly, various parts of
the legs are developed into specific structures for special uses.
For descriptive purposes it is customary to name the leg surfaces
according to the positions they assume when the limb is extended
straight out from the body. In such a position the upper surface is
dorsal, the under surface ventral, the preaxial surface anterior, and
the postaxial surface posterior. The actual directions of the leg
surfaces in normal positions, however, may be quite different.
Inasmuch as the legs differ principally in relative size, form, and
outer structure, the general external features will be described
separately for each pair, while the mechanism of the joints, and the
muscles, which are but little different in the three pairs of appendages,
will be treated collectively in order to avoid unnecessary repetition.
The fore legs—tThe fore legs (fig. 13 A) are smaller than the
others and have somewhat less freedom of motion because of the
vertical position of the coxae (fig. 11 G), but, in compensation, the
femur of each fore leg has an anterior twist (fig. 14 A) by which the
tibia acquires a forward movement, and, when fully flexed, lies
against the anterior surface of the femur and trochanter. The long,
cylindrical, bristly first tarsomere is used as a brush for removing
pollen or other particles from the head and fore parts of the body,
but the most important special features of the first leg is the antenna
cleaner at the base of the tarsus.
The antenna cleaner (fig. 14 B) consists of a deep semicircular
notch (b) on the inner (ventral) face of the proximal end of the
basitarsus (Btar), and of a large flattened spur (a) that serves as a
clasp, or fibula, projecting from the inner angle of the distal end of
the tibia (Tb). The posterior margin of the notch bears a comblike
fringe, or pecten, of fine, closely set, spinelike hairs. The fibula is a
broad, thin, movable appendage with a narrowed base, a tapering
distal point, and a strong, spatulate accessory lobe (e) on its anterior
surface. When the tarsus is flexed by the depressor muscle (FE, 68)
the proximal lobe of the basitarsal notch (C, c) glides under the
accessory lobe (¢) of the fibula, and the distal point of the tarsal
notch (d) is caught in the angle between the two fibular lobes. The
tarsal notch is thus closed to a semicircular aperture by the accessory
lobe (e) of the fibula, and the larger flat lobe (a) overlaps the aper-
ture posteriorly. When the organ is used by the bee, the tarsal notch
is placed around the base of the antennal flagellum by movements of
the leg, and then, by flexion of the tarsus, the antenna is pressed
against the accessory lobe of the fibula, which latter resists the
66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
pressure because of a small process of the tibia (f) that abuts against
its base. The antenna is thus held securely and is now drawn upward
through the cleaner, the tarsal comb brushing its sensory outer sur-
face, the thin lobe of the fibula scraping its inner surface.
The antenna cleaner is not a gadget peculiar to the bees; it is
developed in some measure in most of the clistogastrous Hymenoptera
(fig. 14, F, G, H, 1), though the tarsal notch is usually shallower than
in the honey bee, and may be but a sinuosity of the tarsal margin.
The fibula of the cleaner is simply a special development of an or-
dinary movable spur usually present on the end of the tibia, such
as is present on the middle leg of the adult bee (fig. 22 A, B), and
also on the hind leg of the pupa.
The middle legs—The second legs are the least specialized of the
legs in the bee, and have the simplest form in the drone (fig. 22 A).
In each caste the end of the tibia is armed ventrally with a long spine,
which has been said to be used by the worker for removing wax
from the wax pockets of the abdomen, but the evidence of such use
of the spine has been questioned. The basal tarsomere is long and
cylindrical in the drone (A), in the worker and queen it is wide and
flattened, in the worker (B) its under surface is thickly covered with
long stiff hairs projecting distally, and serves as a pollen brush.
The hind legs—vThe large hind legs are the most specialized in
form of all the legs by reason of the great size and lateral flattening
of the tibia and basitarsus (fig. 23 A, F, H). Though this feature
is common to the three castes, a functional reason for it is not evident
except as it is elaborated for useful purposes in the worker. The
femur has a coating of small hairs in the drone (F), in the queen and
the worker (A) it is bushy with long feathery hairs. The tibia of the
drone (F), aside from its size and shape, has no special characters ;
in the queen (H) the tibia is relatively wider and thick at its base;
in the worker (A) it is narrowed proximally and expanded distally,
and its smooth, somewhat concave outer (anterior) surface (C) is
bordered with long curved hairs, forming the pollen-carrying basket,
or corbicula. The basitarsus is equally large in each caste (A, F, H),
but there is no apparent reason for its size and shape in the queen and
the drone; the cavity of the basitarsus is traversed by the tendon of
the pretarsal claws and is otherwise occupied only by a loose mass of
fat tissue. In the worker the inner (posterior) surface of the basi-
tarsus (A, Btar) is armed with about 10 transverse rows of long,
slender spines directed mesally and distally at an angle of 45°. The
basitarsus of the worker thus resembles a flat brush or currycomb,
and has been termed the strigilis (Betts, 1923), though the same
S40
NO. 2 HONEY BEE—SNODGRASS 67
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ies
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Fic. 22—-The mesothoracic leg and its muscles.
A, right middle leg of drone, posterior view. B, same of worker, posterior view. C, endo-
sternum of pterothorax and mesothoracic coxa, with thoracic muscles of leg, left side (coxa
turned outward). D, base of right middle leg of drone, posterior view, showing trochanteral
and femoral muscles. E, tibio-tarsal joint of right middle leg of drone, dorsal view. F, same,
ventral view. G, trochanter and base of femur of left middle leg of drone, with muscles,
mesal view. H, diagram of dicondylic femoro-tibial joint. I, diagram of monocondylic tibio-
tarsal joint (lateral muscles not shown). J, femoro-tibial joint of right middle leg of drone,
ventral view. K, end of femur with tibia removed, ventral view. L, base of tibia, ventral
view, showing levator muscle (90) and chordotonal ligament (0). M, tarsus and pretarsus of
right middle leg of drone, dorsal view.
d, pleural articulation of coxa; e, sternal articulation of coxa; f, anterior coxal articular
process of trochanter; h, dorsal articulation of trochantero-femoral joint; j, anterior femoro-
tibial articulation; k, posterior femoro-tibial articulation; m, tibial articular process of tibio-
tarsal joint; m, tarsal articular knob of tibio-tarsal joint; 0, tendon of femoral chordotonal
organ; p, process of endosternum giving attachment to muscle 58; q, process of supraneural
bridge of endosternum giving attachment to muscle 52,
68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
name has been used for the antenna cleaner; the brush serves for
the collection and retention of pollen to be stored in the pollen basket
of the tibia. The deep notch in the dorsal margin of the hind leg
between the tibia and the basitarsus is converted in the worker into a
pollen press for the transfer of pollen from the basitarsal brush of one
leg into the tibial basket of the other.
The pollen press of the worker bee is a simple adaptation of the
pincerlike tibio-tarsal notch for pushing pollen from one side of the
leg to the other and for compressing it in the pollen basket. The
proximal end-surface of the basitarsus opposed to the distal end of
the tibia (fig. 23 B) is beveled outward and upward toward the
tibia and is expanded in a small lobe, known as the auricle (au),
bordered by a fringe of long hairs. The surface of the auricle is
covered with small spicules, and is limited mesally by a strong ridge.
The opposing free end of the tibia bears mesally a row of short,
closely set, sinuous spines, forming a little rake, or rastellwm (ras),
which projects down over the base of the auricle (D, E). Laterad
of the rastellum the end-surface of the tibia is concave and is sepa-
rated by a low ridge from the floor of the pollen basket above it on
the outer (anterior) surface of the tibia (C, cbl). When the basi-
tarsus is bent upward at the tibio-tarsal joint (D, E), the fringed lip
of the auricle, guarded mesally by the rastellum, passes within the
semicircle of long curved hairs on the lower end of the tibia and over-
laps the floor of the pollen basket. As this gadget is used by the
bee for loading the pollen basket, a small mass of pollen is first
raked off from the basitarsal brush of the opposite leg by the rastellum
of the tibia; the detached pollen falls on the rough surface of the
auricle and adheres to it. Then, by an upward flexure of the tarsus
on the tibia, the mass of pollen on the auricle, held in place by the
rastelium, is pressed up into the tibial basket. With repeated scrapings
and operations of the press the basket is finally loaded by successive
increments of pollen added at the bottom of the mass.
It is interesting to note that both the antenna cleaner and the pollen
press are developed at the tibio-tarsal joint. This joint differs from
the other leg joints in that the tarsus has a monocondylic hinge on the
tibia, and three muscles. Since this structure pertains to the cor-
responding joint in all the legs of the three castes of the bee, it is
clear that it is not an adaptation to either the antenna cleaner or the
pollen press. On the other hand, it is possible that the development
of these instruments has depended on the peculiar nature of the tibio-
tarsal joint and the unusual mobility of the tarsus.
NO. 2 HONEY BEE—SNODGRASS 69
The coxae and their muscles—The movements of an insect’s leg as
a whole depend on the nature of the coxal articulation with the body.
In the bee the coxae of all the legs swing forward and backward on
“ALA MIWOUW
QOS
Pan
aN
Fic, 23.—The metathoracic leg, external structure.
A, right hind leg of worker, posterior (inner) surface. B, opposing ends
of tibia and basitarsus of right hind leg of worker, posterior view. C, anterior
(outer) face of hind tibia of worker. D, pollen press of worker between tibia
and basitarsus, dorsal view. E, same, better shown by removal of tibial hairs.
F, right hind leg of drone, posterior (inner) surface. G, right hind tarsus
and pretarsus of drone, posterior view. H, right hind tibia, tarsus, and
pretarsus of queen, posterior surface.
transverse axes, and the muscles of each coxa are inserted anterior
and posterior to the coxal axis of movement. The coxal muscles,
therefore, are promotors and remotors. Morison (1927) designates
7O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I03
the coxal muscles “extensors” and “flexors,” the “extensors” being
the lateral muscles, the “flexors” the mesal muscles, but this nomen-
clature ignores the relation of the muscles to the natural movements
of the coxae. Morison adds, however, that a further study “would
doubtlessly result in a better nomenclature” for the muscles and a
“more accurate statement of their functions.”
The fore coxae are attached by articular membranes to the pleuro-
sternal suspensorium of the prothorax with somewhat obliquely trans-
verse axes between weak pleural articulations and the furcasternum
(fig. 11 G) ; sternal articulations are absent, but the mesal angles of
the coxae overlap the furcasternal margins. Though the fore coxae
are thus freely suspended, their principal movements are forward and
backward.
Each fore coxa has six muscles (fig. 13 C, D), two of which are
promotors, three remotors, and one an intersegmental retractor of
the propectus. Of the promotors, one (53) arises on the episternum
and is inserted laterally on the anterior margin of the coxa, the other
(54) takes its origin on the side of the prothoracic endosternum and
is inserted mesally on the anterior coxal margin. The remotors in-
clude two lateral muscles and one mesal muscle. The first lateral re-
motor is a long muscle (C, D, G, 55) arising on the side of the pro-
notum at the base of the spiracular lobe (fig. 17 C), the second (56)
a broad, fan-shaped muscle taking its origin on the ventral surface
of the supraneural bridge of the endosternum (figs. 11 C, 12C);
both muscles are inserted close together on the coxal rim shortly be-
hind the pleural articulation (fig. 13 D, 55, 56). The mesal remotor
(57) is a slender muscle with its origin on the pleural apophysis
(fig. 11 C) and its insertion on the posterior mesal angle of the coxal
base (fig. 13 C, D).
The sixth muscle of the fore coxa (fig. 13 C, D, G, 58) comes from
the mesothorax, where it arises on the anterior part of the median
sternal ridge that supports the superstructure of the pterothoracic
endosternum (fig. 22 C) ; it extends horizontally forward and laterally
(fig. 12 C, 58) to its insertion on a small point of the coxal base
mesad of the pleural articulation (fig. 13 D). The pull of this muscle
can have little specific motor action on the coxa, but the two muscles
diverging from the mesosternal ridge to the fore coxae are evidently
retractors not only of the coxae but of the entire propectus, being thus
adjuncts of the intersegmental ventral thoracic muscles (fig. 12 C, 52).
The intersegmental coxal muscle is classed as an “extensor” of the
coxa by Morison (1927), but the corresponding muscle in Vespula
NO. 2 HONEY BEE—SNODGRASS FM
is termed by Duncan (1939) the “mesosternal retractor of the
propectus.”’
The coxae of the mesothorax are elongate and semicylindrical in
form; each is attached by the entire length of one side to the under
surface of the thorax behind the mesosternum, and the trochanter
arises from its mesal end (figs. 15, 22 A, B). The mesocoxa is
definitely hinged to the body between a lateral pleural articulation
and a mesal sternal articulation (fig. 16 F) ; its axis of movement is
obliquely transverse, with the mesal end somewhat in advance of the
lateral end, and its movement is rotary on the axis.
The musculature of each middle coxa comprises two promotor
muscles inserted at opposite ends of the coxa anterior to the axis of
rotation, and two remotors with the same distribution behind the
axis. The lateral promotor (fig. 22 C, 8o) is a long muscle arising
dorsally on the small pleural apophysis (PIA) of the mesothorax, and
inserted on the anterior coxal rim a short distance from the pleural
articulation (d). The small mesal promotor (S17) arises ventrally on
the median plate of the pterothoracic endosternum, and is inserted on
the anterior coxal margin near the sternal articulation (e). The
lateral remotor of the mesocoxa (82) is the muscle of the subalar
epipleurite (Sa) ; it arises by a broad base on the posterior margin of
the coxa just behind the pleural articulation, and is inserted by a long
tendon on the subalare. This muscle probably has little effect as a
motor of the coxa, but it would appear to be primarily a pleurocoxal
muscle. The mesal remotor of the coxa (83) is the largest of the
coxal muscles; it arises on the median plate of the endosternum and
is strongly attached mesally on the posterior coxal margin.
The metathoracic coxae (fig. 15) are hinged obliquely transverse
between pleural and sternal articulations (fig. 16F), as are the
mesocoxae, but their bases are more nearly horizontal than are those
of the latter, and the coxae themselves project posteriorly (fig. 24 A),
with the trochanters arising from their distal ends.
The musculature of the metacoxae is the same as that of the
mesocoxae, there being inserted on the base of each hind coxa four
muscles (fig. 24B), two of which, attached before the coxal axis
(d-e), are, the one a lateral promotor (103), the other a mesal pro-
motor (104), while a third inserted mesally behind the axis is a mesal
remotor (106). The fourth muscle (705), as in the mesothorax, is
the coxosubalar muscle of the wing; it is attached on the coxa so
close to the pleural articulation (d) and so nearby on the line of
the coxal axis that it can have little effect as a motor of the coxa.
72 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
The mesal promotor and the mesal remotor take their origins on
the metathoracic basal components of the pterothoracic endosternum
(fig. 24 A, 104, 106) ; the lateral promotor (1703) arises on the ridge
between the metapleuron and the propodeum.
The coxo-trochanteral joint—The union between the coxa and the
trochanter in each leg of the bee is a dicondylic hinge with an ap-
proximately horizontal anteroposterior axis. The movements of the
trochanter on the coxa, therefore, are up and down, and are best
expressed as levation and depression. In the bee the trochanter
of each leg is provided with two levator muscles, one anterior, the
other posterior, arising within the coxa, and with two or three de-
pressors having a common point of insertion, but arising both in the
coxa and within the thorax. The two levators are inserted directly
on the trochanter or close to it; the fibers of the depressor muscles
have no direct connection with the trochanter, but are inserted on a
large depressor apodeme arising from a small plate in the ventral
articular membrane at the base of the trochanter. The trochanteral
muscles are the motors of the entire part of the leg (telopodite) be-
yond the coxa, the movements of which are in a plane at right angles
to the coxal plane of movement.
In the fore leg the relatively small anterior and posterior levators
of the trochanter (fig. 13 E, H, 50, 60) arising in the coxa are
inserted on the trochanteral base just beyond the coxo-trochanteral
articulations (E, f, g). The depressors include a long thoracic
muscle (E, H, 61) arising on the prothoracic pleural apophysis (figs.
11 C, 12 C, 61), which is inserted on the end of the depressor apodeme
of the trochanter (fig. 13 E, H), and a large group of fibers (62),
arising on the mesal wall of the coxa, inserted on the sides of the
apodeme and on the apodemal plate. Since the articular surfaces of
the trochanter (E, f, g) are much closer to the insertions of the
levator muscles than to the base of the depressor apodeme, the
effectiveness of the large depressor muscles is greatly increased by
their advantageous leverage.
In the mesothoracic legs the two levator muscles of each trochanter
arise on the lower (lateral) wall of the coxa (fig. 22 D, 84, 85), and
are inserted anteriorly and posteriorly on the base of the trochanter.
The thoracic muscle of the depressor group (D, G, 86) takes its
origin on the side of the endosternum (C); the coxal fibers are
distinctly divided into two muscles (D, G, 87, 88), one anterior, the
other posterior, separated by the pedal nerve (G, Nv) entering
between them into the trochanter.
NO. 2 HONEY BEE—SNODGRASS We
The trochanter of each hind leg has two large levator muscles
(fig. 24 C, D, 107, 108), a long thoracic depressor (C, FE, rog) arising
on the metathoracic arm of the endosternum (A, 100), and a single,
broad coxal depressor (C, E, rzo0). The levators (107, 108) are
inserted in the dorsal articular membrane (D), the depressors (109,
II0) on a long ventral apodeme (Ap).
The trochantero-femoral joint——The hinge between the trochanter
and the femur differs from all the other leg joints in that its axis
traverses the vertical plane of the leg, and is, moreover, strongly
oblique, the dorsal end being distal to the ventral end (figs. 13 H,
24 C). The movements of the femur on the trochanter, therefore, are
those of production and reduction, but, owing to the obliquity of the
hinge, the femur acquires also an upward slant as it turns backward.
A single reductor muscle activates the femur.
The obliquity of the trochantero-femoral joint in the vertical plane
of the leg gives the trochanter a firm pressure or hold on the femur in
responding to the depressor and levator action of the trochanteral
muscles, the trochanter and the femur being thus moved as a unit on
the coxo-trochanteral articulation.
The character of the trochantero-femoral joint differs but little in
the three legs of the bee. In each leg there is anteriorly a narrow
hinge line between the two adjoining segments (figs. 13 H, 24C),
with an articular point (/) at the upper end; only in the hind leg is
there a definite ventral articulation (fig. 24 C, F, 7). Posteriorly the
trochanter and femur are separated by a wide, thick, strongly elastic
conjunctiva (fig. 13 B, cnj). The femoral reductor muscle (fig. 13 H,
Ge fic. 22D, G, So; fig. 24 C, 117) arises anteriorly by a broad
base in the trochanter and its convergent fibers are inserted on a
shelflike apodemal plate projecting from the posterior rim of the basal
foramen of the femur (figs. 13 F, 24 F, Ap). The contraction of
the muscle, therefore, turns the femur posteriorly on the anterior
hinge with the trochanter ; the muscle is opposed by the resistence of
the elastic posterior conjunctiva, which restores the femur to its
original position on relaxation of the reductor muscle.
The femoro-tibial joint —The “knee” joint of each leg has strictly
a hinge motion on a horizontal axis between anterior and posterior
articulations. The movements of the tibia, therefore, are those of
extension and flexion on the end of the femur; they are produced
by dorsal and ventral muscles lying in the femur. The head of the
tibia is somewhat bent toward the femur, and the hard wall of the
femur is deeply emarginate below the joint, allowing the tibia to be
flexed close against the femur.
74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Fic. 24.—The metathoracic leg, muscles and joints (drone).
A, endosternum and ventral wall of posterior part of pterothorax, showing muscles of meta-
thoracic coxae, dorsal view. B, base of right hind coxa and muscle insertions, dorsal view.
C, right trochanter and base of femur, with trochanteral and femoral muscles, anterior (outer)
view. D, proximal view of right trochanter and muscle insertions. E, depressor muscles of
trochanter. FF, base of right femur, anterior view. G, femoro-tibial joint of right hind leg,
ventral view. H, right hind femur with muscles, anterior view. I, distal end of right tibia.
J, proximal end of right basitarsus. K, right hind tibia and muscles, posterior view. L, tibio-
tarsal joint of right hind leg, dorsal view.
d-e, pleurosternal axis of coxa; f, g, anterior and posterior coxal articular processes of
trochanter; h, i, dorsal and ventral trochantero-femoral articulations; j, k, anterior and
posterior femoro-tibial articulations; m, tibial articular process of tibio-tarsal joint; mn, tarsal
articular process of tibio-tarsal joint; 0, ligament of femoral chordotonal organ.
NO. 2 HONEY BEE—SNODGRASS 75
The femoro-tibial joint differs but little in the three legs of the
bee. Its mechanism is well exemplified in the middle leg, which, seen
from below (fig. 22 J), shows the two articular points (j, k) of the
joint and the deep membranous ventral emargination of the wall of
the femur. The articulations consist of two strong basal lobes of the
tibia (L), which fit against corresponding processes concealed within
the end of the femur (K, j, k). The conjunctival membrane below
the joint (H, J) contains a small, partly invaginated genuflexor plate
(gf), to the apex of which is attached the tendon of the broad, pin-
nate flexor muscle of the tibia (97) lying ventrally in the femur.
The fibers of the extensor muscle arise along the dorsal wall of the
femur (H, 90), and are inserted on an axial tendon attached to the
recurved dorsal head of the tibia (H, L). In the hind leg the joint
structures are essentially the same as in the middle leg, though some-
what different in shape (fig. 24 G), and the tibial muscles are larger
BH. 772; 113).
The femur of each leg contains in addition to the tibial muscles a
branch of the flexor muscle of the pretarsal claws (fig. 24 H, rz7a),
which is inserted on the end of a long tendon (117Ap) extending
through the tibia and tarsus. Finally, there is to be found in each
femur a slender, tendinous thread (0) attached on the head of the
tibia just anterior to the extensor tendon (fig. 22 L), which extends
proximally in the femur, and probably belongs to a chordotonal organ
such as occurs in the femur of various other insects.
The tibio-tarsal joint—The joint between the tibia and the tarsus
in each leg of the bee is quite different from that between any of
the other leg segments ; it is monocondylic, and the tarsus is movable
on the tibia by three muscles. The single point of articulation is
median and dorsal, but is concealed within the distal end of the tibia;
two of the muscles are inserted respectively anterior and posterior to
the articulation, the third muscle is ventral.
The essential features of the tibio-tarsal joint are the same in all
three legs, but structural details and the mechanism are somewhat
different in the hind legs. In the first and the second leg the proximal
end of the long basitarsus presents dorsally three knobs (fig. 22 E).
The middle knob (7) is the articular condyle; it is received on the
concave surface of a large, hook-shaped articular process (mm) that
descends from the dorsal wall inside the end of the tibia (I, m). The
lateral knobs give attachment to the anterior and posterior tibial
muscles (E, 92, 93), which arise on the corresponding walls of the
tibia. In the fore leg and the middle leg the attachments of these
muscles on the tibia are so nearly on a level with the articular condyle
76 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I03
that the anterior muscle (92) acts as a productor of the tarsus, and
the posterior muscle (93) as a reductor when tension is exerted on
the muscles in a dead specimen. Morison (1927) calls these two
muscles anterior and posterior “extensors” of the tarsus, but if they
have an extensor effect on the tarsus, the action is weak. The third
tarsal muscle arises on the lower wall of the tibia (fig. 22 F, I, 94)
and is attached by an axial tendon in the ventral conjunctival mem-
brane of the joint. This muscle unquestionably is a depressor of the
tarsus. The opposite movement of the tarsus, if not produced by the
lateral muscles, must result from the elasticity of the articular con-
junctiva. In the fore leg the tibio-tarsal joint mechanism operates the
antenna cleaner.
The tibio-tarsal joint of the hind leg, as above mentioned, differs in
some ways from that of the other legs. The articular condyle of the
basitarsus is a long, curved arm descending from the dorsal rim of
the proximal tarsal foramen (fig. 24 J, ») ; the corresponding articular
process of the tibia (I, m) is a strong sclerotization of the anterior
wall of the tarsal socket in the end of the tibia (L, m), which curves
inward over the tendon of the anterior muscle (114), dips downward,
and turns distally to articulate ventrally with the lower end of the
tarsal condyle (n). The articulation of the two segments is thus
almost at the center of the tarsal foramen. As a consequence, the
attachment of the posterior tarsal muscle (J, K, L, 175) is so far
above the level of the articulation that this muscle, representing the
tarsal reductor in the other legs, becomes in the hind leg a strong
levator of the tarsus. The anterior muscle (714), being inserted nearly
opposite the articular point, is evidently a productor; the ventral
muscle (176) is the tarsal depressor.
The structure of the tibio-tarsal joint of the hind leg is the same in
both the drone and the worker, and hence has no direct relation to the
pollen press of the latter. The mechanism of the joint in the hind
leg, however, especially the combined levator-productor action of the
two lateral muscles, is unquestionably conducive to the effective action
of the press.
The tarsal joints——The narrowed base of each of the small tarsal
subsegments, or tarsomeres, beyond the large basitarsus fits into a
receptive cavity in the end of the preceding tarsomere (fig. 23 G),
and each of these joints is monocondylic, as is the tibio-tarsal joint
itself. The dorsal lip of the minute basal foramen of each small
tarsomere is drawn out into a long articular process, which fits into a
socket formed by an inward extension of the lip of the distal foramen
of the tarsomere preceding. There are no intratarsal muscles giving
NO. 2 HONEY BEE—SNODGRASS TT.
individual movement to the tarsomeres, but the entire tarsus is
traversed by the flexor-muscle tendon (117Ap) of the pretarsal
claws, and a pull on this tendon causes a deflexion of the slender part
of the tarsus beyond the basitarsus.
The pretarsus——The pretarsus of the honey bee has a complex and
highly specialized structure (fig. 25 A, B). The median part of the
segment is differentiated into a conical, stalklike basal part (E) from
which the claws arise laterally, and an oval distal lobe, which is the
arolium (Ar). In the ordinary position of the foot (E) the arolium
stands upward between the claws on the end of the basal support.
Dorsally the arolium presents a deep concavity (A) between its up-
turned lateral parts, and in its outer wall is a dark, elastic, U-shaped
band, the arcus (B, E, arc), which embraces the base of the arolium
ventrally with its arms extending distally in the lateral walls. The
conical base of the pretarsus contains dorsally an elongate median
sclerite (A, E, mn) armed with five or six long, thick, curved spines.
By its widened base the sclerite is articulated to the end of the tarsus
between the bases of the claws, and its narrowed distal end is at-
tached like a handle to the base of the scoop-shaped arolium (H),
for which reason the sclerite may be termed the aroliar manubrium.
In the ventral wall of the pretarsal base is a broad, weakly sclerotized
plate, the planta (B, E, pln), thickly beset with strong spines diverging
distally.
The claws are hollow, strongly sclerotized lateral outgrowths of
the membranous lateral walls of the pretarsal base. Each claw has
two points of unequal length, and in size and shape the claws differ
considerably as between the drone (fig. 25 A, B) and the worker (C,
D). Dorsally the claws are articulated individually to two small
knobs on the end of the tarsus (A, 0, p), corresponding with the ar-
ticular condyles at other dicondylic joints of the leg, but the claws
themselves are devoid of muscles.
The motor apparatus both of the claws and of the arolium consists
of the depressor muscle of the pretarsus, and of accessory sclerites in
the ventral articular membrane at the base of the segment. The prin-
cipal one of the basal sclerites is a large median plate, known as the
unguitractor plate (fig. 25 B, Utr), which is comparable with the
genuflexor plate below the knee joint. In the membrane at the
proximal end of the unguitractor plate is attached the tendon of the
pretarsal muscle (954p), and at the distal angles are two small
auxiliary sclerites (aux) intermediate between the unguitractor plate
and the bases of the claws. The pretarsal tendon extends forward
through the tarsus (fig. 23 G) into the tibia (fig. 24K), where it
78 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103 |
divides into two branches, one branch giving insertion to a group
of fibers arising posteriorly in the tibia (117b), the other, going on
into the femur, to a second group (H, r1r7a) arising posteriorly in
Fic. 25.—The pretarsus.
A, pretarsus of middle leg of drone, dorsal surface, flattened, inner sides of
claws uppermost, arolium turned upward. B, same, ventral surface, arolium
depressed and spread out. C, posterior claw of middle leg of worker, outer
surface. D, anterior claw of hind leg of worker, inner surface. FE, pretarsus
and end of tarsus, claws removed, arolium turned upward, semidiagrammatic.
F, same, arolium depressed. G, ventral surface of fully depressed and flattened
arolium. H, arolium and manubrium, dorsal view. I, paper model of arolium
and manubrium in nonfunctional condition. J, same, illustrating flattening of
arolium by downward pressure on manubrium.
95Ap, tendonlike apodeme of muscle of pretarsus (retractor of the claws) ;
Ar, arolium; arc, arcus; aux, auxilia; mn, manubrium; o, », anterior and
posterior tarso-pretarsal articulations; pln, planta; Tar, end of fifth tarsomere;
Un, claw (unguis) ; Utr, unguitractor plate.
the base of the femur. The tension of the pretarsal muscle on the
unguitractor plate is distributed to the claws through the auxiliary
sclerites, and thus flexes the claws, for which reason the muscle is
NO. 2 HONEY BEE—SNODGRASS 79
known as the flexor of the claws. The direct effect of the muscle pull,
however, is exerted on the base of the pretarsus, and, unless the
claws interfere, it flexes the pretarsus on the end of the tarsus (fig.
25 F) and brings the arolium into a horizontal position.
The arolium is an organ that enables the bee to cling to smooth
surfaces on which the claws cannot hold. When the arolium is in use
as an adhesive organ the claws are turned proximally with their points
directed outward, and the arolium is horizontally extended, its lateral
walls spread out flat, its under surface (fig. 25 G) pressed against
the support. The extension and flattening of the arolium has been
explained by Arnhart (1923; see Snodgrass, 1925) as caused by a
secretion liquid forced into the foot from a gland in the end of the
tarsus; but a study of the mechanism of the pretarsus shows that
the action of the arolium including its changes in shape can be
produced entirely by mechanical means.
With a live bee it may be observed that when the flexing claws
encounter on a smooth surface nothing to restrain them or to give
them a hold, they turn helplessly upward beneath the end of the
tarsus with their points sprawling outward. The tension of the
contracting pretarsal muscle is then exerted on the base of the pre-
tarsus, which turns abruptly downward bringing the arolium into a
horizontal position, after which the arolium extends, spreads out
flat, and clings to the support. If the foot of a dead bee is in the
position shown at F of figure 25, an upward pressure on the retracted
planta (fln) extends the pretarsus until the planta comes into the
same horizontal plane as the arolium (Ar), and now, with continued
pressure, the arolium itself automatically unfolds and spreads out
with its flattened under surface downward. This action of the arolium
can be induced by manipulation of a detached pretarsus, in which
there can be no possibility of liquid pressure from within the tarsus.
The same kind of action, moreover, can be illustrated with a piece of
paper cut and folded into the form of a scoop (I); a vertical com-
pression of the base of the scoop spreads the sides widely apart (J),
giving thus a close imitation of the partly expanded arolium (H).
The similar response of the arolium to pressure against the planta is
evidently caused in the living bee by the downward pressure of the
tarsus, which flattens the pretarsus and compresses the arolium
against the support beneath it. The manubrial sclerite of the pretarsus
(fig. 25 E, H, mn), however, representing the handle of the paper
scoop (I, J), plays an important part in the expansion of the arolium.
With the extension of the foot, the manubrium turns backward on the
end of the tarsus and hence pushes on the upper edge of the aroliar
6
80 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I03
base, thus augmenting the effect of the pressure on the planta;
finally, when fully extended, the manubrium clamps down on the
arolium and holds it securely in the spread condition. There can be
little doubt, therefore, that the entire action of the arolium is a
mechanical result of the pull of the pretarsal muscle in conjunction
with pressure of the foot against the support when the claws fail to
take a hold on the latter. It has been said that the arolium adheres
to smooth surfaces by means of a sticky exudation from the spines
of the planta, but observation fails to reveal the presence of any such
adhesive. The upper surface of the arolium is covered with minute
hairs, but its under surface is almost entirely bare.
VI. THE ABDOMINAL PETIOLE
The outstanding feature in the external organization of the clisto-
gastrous Hymenoptera is the secondary consolidation of the fourth
body segment with the thorax, and the division of the body by a deep
constriction between the fourth and fifth segments, accompanied by
a narrowing and usually an elongation of the anterior end of the
fifth segment, or even of the entire segment, to form a petiole. Two
important mechanical advantages result from these alterations, which
take place during the pupal stage of development. First is the greatly
increased length that the dorsal wing muscles of the thorax are able
to acquire by a posterior extension of the mesothoracic postphragma
into the fourth segment (fig. 16 C); second is the high degree of
potential mobility conferred on the shortened abdomen. The first
advantage is self-evident, the second needs elucidation.
In the more generalized insects the abdomen is broadly attached
to the thorax by its first segment, and the muscles that move the
abdomen as a whole are dorsal and ventral intersegmental muscles
between the metathorax and the abdomen. In winged insects, how-
ever, the dorsal intersegmental muscles of the metathorax become
functionally a part of the flight mechanism (fig. 26 A, d;) by a
transfer of the ridge or phragma of their abdominal attachment to
the thorax. As a consequence, the dorsal metathoracic muscles of
winged insects have no motor effect on the abdomen. Ventral muscles
between the thorax and the abdomen are seldom strongly developed,
and they may be absent; when present they comprise generally one
pair or two pairs of slender muscles arising anteriorly on the meta-
sternal apophyses and attached posteriorly sometimes on the first ab-
dominal sternum (A), but more commonly for greater effectiveness
on the second (C) or even the third, fourth, or fifth sternal plate.
NO. 2 HONEY BEE—SNODGRASS SI
Such insects have little mobility of the abdomen on the thorax, even
though the two parts of the body may be separated by a constriction.
A comparison of the structural relation between the thorax and
the abdomen in clistogastrous Hymenoptera (fig. 26 B) with that in
Bee aise js B
Fic. 26.—Relation between the propodeum and the abdominal petiole; lateral
muscles of an abdominal segment.
A, diagram of thoracico-abdominal relation in a generalized winged insect:
the dorsal intersegmental muscles of metathorax (d;) are wing motors and
do not move the abdomen. B, specialized condition in clistogastrous Hymenop-
tera: dorsal and ventral muscles (d, v) of first abdominal segment (pro-
podeum) become effective motors of rest of abdomen. C, usual structure in
higher winged insects: abdomen but little movable on thorax. D, anterior end-
view of abdominal segment J” of drone, showing mechanism of dorsoventral
expansion by contraction of reversed anterior lateral muscles. E, right half
of abdominal sternum JV of drone, with sternal and lateral muscles.
a, anterior tergal apodeme; b, anterior sternal apodeme; c, lateral sternal
apodeme; d, dorsal intersegmental muscles; v, ventral intersegmental muscles.
either a generalized winged insect (A) or a more specialized form (C)
will show clearly the advantage derived by the Clistogastra from hav-
ing the first abdominal segment solidly joined to the thorax, and a
constricted point of movement established between the first and
82 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103 _
second segments of the abdomen. The modification allows the muscles
of the transposed first abdominal segment (B, d, v) to become effec-
tive motors of the free part of the abdomen. The aculeate Hymenop-
tera thereby acquire a much greater abdominal mobility for the use
of the ovipositor or sting or for mating activities than is possessed by
any other insects.
The mechanism of the clistogastrous petiole is well illustrated in
the honey bee, though the petiole itself in the bee is relatively short.
Particular attention to the petiole as an important part of the or-
ganization of the bee was first given by Betts (1923), who fully
describes its structure and its principal muscles. The propodeum and
the first segment of the functional abdomen are joined by a mem-
branous conjunctiva (fig. 27 A, C), which permits freedom of move-
ment of the abdomen on the propodeum, but the two parts are hinged
dorsally by a pair of articular points (C, f, f). Between the articular
points on the dorsal surface of the petiole is a small elevation of the
conjunctival membrane (e) connecting anteriorly with a notch in
the end of the propodeum (ft). Within this swelling of the con-
junctiva is a dorsal pocket of the anterior end of the petiole (D, F, G)
having a strongly sclerotic floor supporting a median ridge (D, F, r).
Ventrally (A, E) the conjunctiva is a wide membrane between the
weak sternal plate of the propodeum (JS) and the sternal plate of
the petiole (J7S). The muscles that move the abdomen as a whole
are attached dorsally, laterally, and ventrally on the narrowed end
of the petiole (G); they represent dorsal intersegmental muscles
between the primarily first and second abdominal segments (fig. 26 A,
B, C, d), and ventral intersegmental muscles of the metathorax (B,
C, v) that pass over the first abdominal sternum to attach on the
second.
The petiole muscles of the honey bee include two pairs of dorsal
muscles and two pairs of ventral muscles, but on account of the
positions of their attachment on the petiole, one pair of the dorsal
muscles are levators of the abdomen, one pair of ventrals are de-
pressors, and the other two pairs evidently produce sidewise move-
ments. The levator muscles arise by broad, spreading bases on the
lateral walls of the propodeum (figs. 27 B, C, 28 A, 120), and are
inserted by strong tendons that converge to their attachments on the
posterior end of the ridge on the floor of the petiole pocket (fig. 27 D,
F, G). These muscles thus acquire a short leverage on the propodeal
fulcra (f). The depressors of the abdomen are a pair of long
ventral muscles arising on the metathoracic components of the ptero-
thoracic endosternum (fig. 27 E, rz8), and inserted medially on the
NO. 2 HONEY BEE—SNODGRASS 83
anterior margin of the sternal plate of the petiole (E, G). The lateral
muscles of the petiole include a pair of two-branched dorsal muscles
arising anteriorly on the sides of the propodeum (figs. 27 B, 28 A,
121) and inserted on the lateral angles of the tergum of the petiole
/
—
12 —SS—_ __ 4
Ip f
/
Fic. 27—The propodeum and petiole. (Figures, except E, from Snodgrass,
1933.)
A, abdomen of worker, including propodeum. B, muscles in right side of
propodeum, mesal view. C, propodeum and petiole, dorsal wall of propodeum
removed, exposing muscles. D, levator mechanism of abdomen exposed by
removal of left wall of propodeum and petiole. E, ventral muscles of petiole,
dorsal view. F, petiole and end of propodeum, dorsal view, upper wall of
petiolar pocket (e) removed to show insertion of levator muscles (120).
E anterior end of petiole, showing muscle attachments on tergal and sternal
plates.
e, membranous roof of dorsal pocket of petiole; f, f, fulcra of movement of
abdomen on propodeum; r, median ridge on floor of petiolar pocket; t, posterior
area of propodeum adjoining petiole.
(fig. 27 G), and a pair of ventral muscles arising on the metathoracic
sternum (FE, 179), which are inserted laterally on the end of the
petiole sternum (E, G). The cavity of the neck of the petiole is
divided transversely by the ventral diaphragm (G, V Dph), which
84 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
begins in the metathorax. The larger dorsal channel gives passage
to the tracheal trunks, the oesophagus, and the aorta; the ventral
channel contains the nerve cords.
VII. THE ABDOMEN
Structural changes in an animal that bring about conditions in
conflict with standardized nomenclature create embarrassing situa-
tions for the descriptive anatomist. The transfer of the fourth body
segment to the thorax in the Clistogastra, for example, leaves us
without a suitable term for the third region of the body in this
group of Hymenoptera, since the body region called the “abdomen”
in all other insects includes the transposed segment of the Clisto-
gastra. The clistogastrous abdomen has been termed the postabdomen,
and the gaster, but neither name is satisfactory; the first is used also
for the last four or five segments in certain insects in which the
posterior genital region of the abdomen is structurally differentiated
from the anterior visceral region, and the word “gaster’’ ordinarily
refers to the stomach. The most practical solution of the difficulty
for hymenopterists is to use the term abdomen in a functional rather
than a morphological sense for the third body region of the insect.
The abdomen in the Clistogastra, therefore, is not entirely homologous
with that of the Chalastogastra, and to keep in view the identity of
the segments in the two groups the primary abdominal segments may
be designated J to X regardless of their associations. Segment J in
the Clistogastra is the thoracic propodeum, segment JJ is the segment
of the abdominal petiole, segment X is the anus-bearing lobe or tube
known as the proctiger. In many Hymenoptera the true anus lies
within a terminal invagination of the proctiger, which possibly
represents segment XJ.
General structure of the abdomen.—The segments of the pregenital
region of the abdomen in the honey bee have well-developed tergal
and sternal plates, but there are no lateral, or “pleural,” sclerites, and
the spiracles lie in the sides of the terga (fig. 27 A). The tergal
plates, except the first, are provided each with a large anterior tergal
apodeme on each side (figs. 26 D, 28 A, 29 A, a); the sternal plates
have each a pair of anterior sternal apodemes (figs. 26 D, 28 A, B,
29 A, b), and on each side a long lateral sternal apodeme (c). The
sternal plates of segments J/J to VJ widely underlap each the sternum
of the following segment, forming pockets which in the worker con-
ceal the areas of the wax glands on sterna JV to VJ.
NO. 2 HONEY BEE—SNODGRASS 85
In the male bee the dorsum of the exposed part of the abdomen
ends with the declivous tergal plate of segment VIII (fig. 29 A),
but the sternum of this segment (VJIIS) is concealed within that of
segment VJ/, and the visible part of the venter terminates with
sternum JX. The tergum of segment /X, however, is represented
only by a pair of small dorsal sclerites lying at the sides of the
proctiger (figs. 29 A, 30 A, XT), each of which supports internally
a large apodeme (fig. 29 A, ap) giving attachment to the dorsal
muscles of its segment (B). The sternum of segment JX (figs. 29 A,
30 A, B, XS) is a relatively large crescentic plate exposed beyond
sternum JI, and bears on its posterior margin the two pairs of phallic
plates (Jp, pv). The proctiger is a membranous lobe (fig. 30 A, Ptgr)
with a small subanal plate (>) in its ventral wall between the two
sclerites of the ninth tergum.
In the queen and the worker bee the exposed part of the abdomen
ends with the tergum and the sternum of segment VJI (fig. 27 A).
The margins of the apical plates are ordinarily closed upon each
- other, concealing a large invagination cavity that contains segments
VIII and IX, the proctiger, and the sting. Segment VJ/I is a
complete annulus at the base of the cavity, but its walls are mem-
branous except for two lateral remnants of the tergum, which contain
the spiracles and are associated with the base of the sting. The
tergites of segment VJII, therefore, are known as the spiracular
plates of the sting (fig. 28 C, VIJIT). In the posterior wall of the
ventral part of the eighth segment is a depression containing the
female genital openings (D). Segment JX supports the sting, but
it is hardly to be recognized as a segment; its only sclerotized parts
are two lateral tergal plates that form virtually a part of the stinging
apparatus (fig. 31 A, Qd), and are hence called, because of their
position and shape, the quadrate plates of the sting. The venter of
the ninth segment (/X.V) is entirely membranous and, when the
sting is retracted, is mostly concealed between the oblong plates (Ob)
of the basal apparatus of the sting.
The abdominal musculature —The musculature of the abdomen
follows a definite plan, the pattern of which is repeated alike in each
of the segments, except for omissions in the first and the last three
segments, and the addition of a few special muscles in connection
with the genital organs and the sting. The typical segmental mus-
culature is complete in segments J/J to VJ in both sexes, but the
individual muscles of the male (fig. 29 B) are much larger than those
of the female (fig. 28 A). The somatic abdominal muscles, exclusive
86 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
of the muscles of the dorsal and ventral diaphragms, may be classed
as intertergal dorsal muscles, tergosternal lateral muscles, intersternal
ventral muscles, and spiracular muscles. The dorsals and ventrals
again include internal and external groups of fibers, and the internal
dorsal and ventral muscles comprise median and lateral muscles.
‘For a study of the abdomnial musculature segment V of the
worker may be taken as a representative segment (fig. 28 A, V), the
larger muscles of the drone (fig. 29 B) being less isolated. On each
side of the tergum of segment V two bands of muscle fibers diverge
posteriorly to attach on the antecosta (Ac) of tergum VJ. These
muscles are the internal dorsals, one median (755), the other lateral
(156). The single short external dorsal (157) is a reversed muscle,
since it arises posteriorly on tergum VY and extends forward to its
insertion on the anterior apodeme (a) of tergum VJ. The spiracular
muscles include a small occlusor (not seen in the figure) attached
at both ends on the spiracle itself, as shown in segment V JI (C, 180),
and a long, slender dilator (A, 159) arising on the lateral apodeme
of the sternum (c). Of the three tergosternal lateral muscles, the
first (160) is a reversed muscle arising ventrally on the lower part
of the tergum and inserted dorsally on the long lateral apodeme of
the sternum (fig. 26D, 149). The other two lateral muscles cross
each other obliquely in the female (fig. 28 A) from the tergum to the
sternum, the one which is external (161) being attached on the sternal
margin, the other (162) on the posterolateral upper surface of the
sternum. In the male (figs. 26E, 29B) the second and third
lateral muscles are parallel. The external muscle is a broad band of
fibers (fig. 26 FE, 150) attached on the sternal margin; the internal
lateral muscle includes two distinct groups of fibers (75ra, 151b), the
second distinguished by its pale, whitish color, both attached on the
upper surface of the posterior part of the sternum. The sternal
musculature duplicates the pattern of the tergal musculature, there
being a pair of posteriorly divergent internal ventrals on each side
(fig. 28 A, B, 163, 164), and a single reversed external ventral (A,
165) between the overlapping parts of the consecutive sterna.
Functionally the internal dorsal and ventral muscles are contractors
of the abdomen, inasmuch as they retract the individual tergal and
sternal plates. The external dorsal and ventral muscles of the more
posterior segments, on the other hand, are extensors of the abdomen,
since their contraction protracts the segmental plates by reducing the
overlap between them. Similarly, the reversed muscles of the lateral
series are dilators of the abdomen (fig. 26D, 140), because their
contraction separates the tergum and the sternum of each segment,
NO. 2 HONEY BEE—SNODGRASS 87
193
Fic. 28.—The abdominal musculature of the worker and queen. (From Snod-
grass, 1933, with muscles renumbered. )
A, muscles in right half of anterior part of abdomen of worker, including
propodeum, mesal view. B, ventral muscles of abdominal segments V, VI, and
VII of worker, dorsal view. C, muscles of right half of segment VJJ and of
spiracular plate of segment VJII of worker, mesal view. D, genital pouches
and oviducts of queen, with attached muscles, posterodorsal view. FE, right
spiracular plate of segment VJ/J of queen, and its muscles, mesal view.
ad, anterior tetgal apodeme; b, anterior sternal apodeme; c, lateral sternal
apodeme; e, dorsal pocket of petiole; f, edge of shallow external part of genital
chamber; g, vaginal aperture (opening of median genital pouch); h, h, aper-
tures.of lateral genital pouches; 7, median pouch (vagina) of genital chamber;
_j, j, lateral pouches of genital chamber.
88 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
and thus expands the abdomen dorsoventrally. The other lateral
muscles pull the tergal and sternal plates together and are therefore 7
compressors of the abdomen. The entire musculature of the pre-
genital segments is operative in respiration, since the abdominal
breathing movements of the bee are both dorsoventral and longi- |
tudinal. The same muscles, however, produce various other move-
ments of the abdomen, and it would appear that the dorsal and ventral
muscles, or the lateral muscles of opposite sides may act either in
unison or as antagonists. Finally, it should be noted that the external
dorsal and ventral muscles in the more anterior segments of the
abdomen (fig. 28 A, 126, 132, 135) run obliquely or transversely
between the adjoining segmental plates, and thus evidently produce
a partial rotary or torsion movement of the anterior segments on
each other.
The musculature of the petiole segment (fig. 28 A, //) is simplified
by the absence of the first and second lateral muscles and, in the
worker, of the lateral internal ventral, though this muscle is present
in the drone (fig. 29 B, 131).
In segments VII, VIII, and 1X the musculature is progressively
reduced and altered as these segments become involved in modifications
adaptive to the reproductive functions. In the drone (fig. 29 B) seg-
ment VJI retains the typical dorsal musculature, but the second pair
of lateral muscles is absent, and the sternum has on each side only a
small lateral internal muscle (B, D, 186) attached posteriorly on the
long anterior apodemal arm (0b) of sternum VJ//. In segment VIII
of the drone there is on each side a single, short dorsal muscle (B,
187) attached posteriorly on the apodeme of the ninth tergal sclerite
(IXT), one lateral muscle (797), and a short lateral internal ventral '
(B, D, 195) between the eighth sternum and the small parameral
plate (Jp) of the genital apparatus. Segment /X has only one pair
of short dorsal muscles (B, 202) from the apodemes of the ninth
tergal plates to the base of the proctiger at the sides of the subanal
plate, but there are two small phallic muscles on each side (D, 203,
204) arising ventrally on the parameral plate (/p) and attached
dorsally on the arm of the penis valve (pv), while finally a long.
slender muscle (205) goes from the arm of the penis valve to the
mucus gland, to which it is attached just laterad of the termination
of the vas deferens.
In the female the musculature as well as the structure of the pos-
terior abdominal segments is very different from that of the male.
The tergal musculature of segment VJJ in the worker includes on
each side an internal dorsal (fig. 28 C, 178) attached on the upper |
NO. 2 HONEY BEE—SNODGRASS 89
margin of the spiracuiar plate of segment V/J/I (VJIIT), and a re-
versed lateral lying external to the spiracular plate and attached on
the upper angle of the latter (779). In the queen there are two
internal dorsal muscles of segment VJI (FE, 177, 178) attached on
the spiracular plate. The lateral musculature consists of two tergo-
sternal muscles on each side (C, 182, 183), and a very small reversed
muscle, not shown in the figures. The sternum of segment VJI in
the worker (C) has none of the usual ventral muscles, but in the
queen a pair of median internal ventrals is present (D, 185) attached
posteriorly on the copulatory pouch. In both the worker and the
queen, however, there is a large, intersegmental sternotergal muscle
(C, 184) arising anteriorly on sternum VJ/ and attached posteriorly
on the lower anterior angle of the spiracular plate (C, E).
The muscles of segment /J/I in the female all take their origins
on the spiracular plates, which are lateral remnants of the eighth
tergum. On each side two dorsal muscles (fig. 28 C, E, 187, 188) arise
in the upper angle of the spiracular plate and go to the corresponding
quadrate plate of the sting, which represents the tergum of segment
IX. Each spiracle has the usual occlusor (189) and dilator (190)
muscles, but the dilator arises on the spiracular plate, there being no
sternal plate in the eighth segment. From the lower edge of each
spiracular plate a long posterior muscle (192) goes forward to the
triangular plate of the sting (fig. 31 B, D, 792), and in the queen two
slender anterior ventral muscles (fig. 28 E, 193, 194) go mesally to
the dorsal wall of the vagina (D).
The muscles of segment /X in the female pertain to the sting,
except two small muscles from the quadrate plate to the ventral wall
of the proctiger (fig. 31 B, 200, 201). The other muscles will be
described in connection with the sting.
The male genital organ.—The phallic organ of the Apidae differs
from that of all other Hymenoptera in a reduction of the outer parts
and a great development of the endophallic sack. In Apis mellifera
the ectophallic parts consist of two pairs of small plates lying behind
the last abdominal sternum (fig. 30 A, B, C, D, lp, pv), the small outer
pair of which (/p) may be identified with the parameral plates, and
the larger inner pair (pv) with the aedeagal plates, or penis valves,
of other families (see Snodgrass, 1941). Between the penis valves
and thick membranous folds uniting them dorsally and ventrally is
a widely distensible opening, the phallotreme (Phtr), which leads into
the cavity of a huge endophallic invagination (F, Enph) that extends
forward as far as the segment of the petiole.
/ \
/ / \
VS VIS VIIS VIIIS IXS_ pv
Fic. 29—The abdominal musculature of the drone, not including the pro-
podeum. (From Snodgrass, 1941, with muscles renumbered.)
A, right half of abdominal wall, mesal view. B, muscles of right half of abdomen, mesal
view. C, right halves of sternal plates of abdominal segments VJJI and IX, and right phallic
plates, mesal view. D, same, with muscles.
a, anterior tergal apodeme; b, anterior sternal apodeme; c, lateral sternal apodeme.
(90)
NO. 2 HONEY BEE—SNODGRASS gI
The endophallus is the intromittent organ of the bee; during mating
it is everted, in part or entirely, and inserted into the genital tract
of the queen. Structurally the endophallus is differentiated into three
major sections: first, opening directly from the phallotreme, is a
large sack, or bursa (fig. 30, F, Brs); the second, middle part is a
narrowed neck, or cervix (Cer), connecting the bursa with the third,
innermost part, which is a large, thick-walled bulb (Blb). The
ejaculatory duct (Dej) opens into the anterior end of the bulb.
The endophallic bursa (fig. 30 F, Brs) has tough membranous
walls, which are more or less collapsed and folded in the retracted
condition. From the sides of the bursa project a pair of large, taper-
ing, thin-walled, crumpled cornua, or pneumapophyses (bc). In the
ventral wall are two somewhat elevated dark areas, the first (c)
quadrate in form, the second (d) shield-shaped. Internally these
areas are seen to be flat pockets of the bursal wall covered with a
dense growth of small, dark spines or thick curved hairs. Elsewhere
the inner walls of the bursa are closely dotted with minute spicules,
but the linings of the cornua are bare, though wrinkled and finely
alveolate in appearance.
The cervix of the endophallus is ordinarily compressed into an
irregular shape (fig. 30 F, Cer), but when stretched out straight it is
seen to be a wide tube with tough membranous walls. Along the length
of its lower wall runs a rounded fold (e) that forms internally a
ventral gutterlike channel. Externally the fold is crossed by a series of
V-shaped elevations conspicuous by their dark color, which is due to
an internal covering of small spines. Similarly, an internal spine-
covered patch at the posterior end of the dorsal wall appears externally
as a triangular or cordate dark area (f). Near its anterior end the
cervix bears dorsally a large diverticulum (flb), known as the pinnate,
or fumbriate, lobe because its margins are deeply cut into flat, over-
lapping lobules.
The large endophallic bulb (fig. 30 F, Blb) has a complex structure,
which is not fully understood since it has never been thoroughly
studied by histological methods. In the distal half of its dorsal wall
are two parallel, elongate, strongly sclerotic plates (E, F, g), which
are usually shown in illustrations as external structures, but they are
covered by a thin outer tunic that forms a fold in the groove between
them and is continued over the rest of the bulb. The plates, therefore,
would appear to be sclerotizations of the inner wall of a flat dorsal
pouch of the bulb lumen; when the covering tunic is removed the
smooth outer surfaces of the plates are exposed (EF) and the tapering
posterior ends are seen to project freely into the posterior part of
Q2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
the cavity of the bulb. The plates are not separable from a large
spongy body beneath them that forms the dorsal wall of the bulb
cavity and thins out on the sides, where it is continued into a chitinous
intima covering the rest of the lumen. From the inner wall of the
spongy body two small, posterior triangular plates (E, h) project
laterad of the dorsal plates as free flaps into the cavity of the bulb.
In most drones taken in late summer the endophallic bulb is found
to be filled with a dense mass of spermatozoa and fine granular
matter, which, in specimens preserved in alcohol, forms a compact
body molded into the shape of the interior of the bulb (fig. 30 H).
This body is termed by Zander (1911) a Samenpatrone (‘seminal
cartridge,” cartouche of Leuenberger, 1929). It is not, however, a
true spermatophore, or sperm-containing capsule, since it has no
retaining wall; its outer surface in preserved specimens is dense and
regular, but is minutely pilose because of the great number of sperm
tails projecting from it.
The discharge of the spermatozoa is accomplished by eversion of
the endophallus. The endophallic walls being entirely nonmuscular,
eversion of the organ is brought about naturally by a strong muscular
compression of the abdomen, and this function of the abdomen in
the male probably accounts for the great size of the abdominal muscles
of the drone (fig. 29 B) as compared with those of the queen or
worker (fig. 28 A). Artificially a partial evagination of the endo-
phallus is easily produced in a fresh specimen by pressure on the
abdomen, and is often seen in killed specimens (fig. 30 G). Bishop
(1920), however, describes methods by which a complete eversion
can be induced, and Leuenberger (1929, fig. 17) gives a photograph
of the fully everted organ.
In the first stage of the endophallic eversion, or the condition
usually produced by artificial compression of the abdomen, the bursa
is turned entirely inside out (fig. 30 G, Brs) with the everted cornua
(bc) projecting upward and outward from its sides. The stretched
cervix of the organ now traverses the lumen of the bursa and opens
between the bases of the cornua. It is generally supposed (though
apparently never observed) that in the copulatory act the cornua, or
pneumapophyses, enter the lateral pouches of the female genital cham-
ber (fig. 28 D, 7) ; probably the cervical aperture on the distended bursa
is held close to the opening of the median vaginal pouch (g), and
the endophallic bulb then everted through the cervix into the vagina.
Zander (1911) contended that the bulb is not everted, since in newly
mated queens he found it in the uneverted condition grasped between
the base of the ovipositor and the seventh abdominal sternum. On
NO. 2 HONEY BEE—SNODGRASS 93
J is
Tia
|
WES HW
VIS IxS PV
Fic. 30—The male genitalia. (Figures A to F from Snodgrass, 1941.)
A, segments VJ/JI, IX, and X of male abdomen, end view, showing phallo-
treme and genital plates between sternum /X and proctiger. B, end of male
abdomen, ventral view. CC, ectophallus, dorsal view. D, same, phallotreme
distended by separation of penis valves (pv). E, dorsal and inner plates of endo-
phallic bulb. F, ectophallus and invaginated endophallus, dorsal view. G, end of
abdomen with endophallus partly everted. HH, mass of spermatozoa and agglu-
tinating material in bulb of endophallus.
a, membranous dorsal lip of phallotreme; An, anus; ap, anterior apodeme
of sternum V JIT; b, subanal plate of proctiger; bc, bursal cornua; B/b, bulb of
endophallus; Brs, bursa of endophallus; c, spiny area of inner bursal wall;
Cer, neck of endophallus; d, setigerous area of inner bursal wall; Dej, ductus
ejaculatorius; e, corrugations of endophallic neck; Enph, endophallus; f, setig-
erous area of inner wall of endophallic neck; fbl, fimbriate lobe; g, dorsal
plate of endophallic bulb; h, inner plate of bulb; /p, lamina parameralis;
Phir, phallotreme; Pmr, paramere; Ptgr, proctiger; pv, penis valve.
94 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
the other hand, Shafer (1917) describes four cases in which he ©
observed the bulb everted and contained within the vagina. Both —
writers agree that the bulb has an inverted position in the queen, and
Shafer notes that the reversed points of the dorsal plates of the bulb
are pressed upward against the vaginal wall, thereby preventing an
extraction of the bulb. Only the bulb is said to be found adhering
to a mated queen, the other parts of the male organ, the cervix and
the bursa, probably being torn away when the queen detaches herself
from the drone. Both Shafer and Bishop believed that the spermatozoa
are discharged into the vagina directly from the exposed end of the
ejaculatory duct, but according to Zander the sperm are already stored
in the bulb at the time of mating. The observations recorded above
on the presence of a spermatic mass in the bulb of unmated drones
suggests that the condition described by Zander is true probably in
most cases. Differences in the age of drones examined may account
for the observed presence or absence of spermatozoa in the bulb.
The mating of the honey bee is a subject that still needs investi-
gation. We know nothing definitely concerning the function of the
various structural parts of the complex endophallic organ of the
drone, and, notwithstanding evidence that the bulb is everted into the
vagina, it is difficult to imagine how such a thick-walled structure can
be turned inside out.
The sting.—The sting of the bee has been so many times described
that only a summarized account of its structure and mechanism need
be given here; the writer has elsewhere (1933) given a full discussion
of both phases of the subject. Though the hymenopterous sting is
a modified ovipositor, its parts have long been known to students
of the honey bee under special names, and these names will be used
in the following description.
The act of stinging involves movements of the entire abdomen that
accomplish the initial thrust of the shaft of the sting into the skin of
the victim. The farther penetration of the shaft into the wound and
the injection of the poison are produced by movements of the sting
itself. The stinging organ (fig. 31 A) comprises two distinct parts:
one is a large basal part by which the sting is attached within the
sting chamber at the end of the abdomen, and which constitutes the
motor apparatus; the other is the long, tapering shaft, which is the
effective part of the organ, and is alone protracted from the abdomen
during the act of stinging. The two parts of the sting are connected
anteriorly by a pair of curved, flexible arms, and in the position of
rest the shaft lies beneath the basal part and is mostly concealed within
the latter.
NO. 2 HONEY BEE—SNODGRASS 95
Fic. 31.—Structure of the sting. (From Snodgrass, 1933, with muscles re-
numbered. )
A, sting of worker, lateral view. B, sting and its muscles, with ventral wall of proctiger,
dorsal view. C, bases of oblong plate and bulb of stylet connected by second ramus (27).
D, triangular plate and base of lancet connected by first ramus (sr). E, proximal bulbous
part of stylet, showing basal articulation on second ramus, attachment of furcula, and entrance
a duct of poison sack into base of bulb. F, furcula. G, cross section of distal part of shaft
of sting.
a, angle of attachment of quadrate plate to spiracular plate of eighth segment; Ap, apodeme
of quadrate plate; b, apex of triangular plate; BGld, “‘alkaline’’ gland of sting; blb, bulb of
stylet; c, d, basal angles of triangular plate; e, fulcrum of triangular plate on oblong plate;
f, articular lobe of second ramus; Frc, furcula; g, articular lobe of bulb; h, hinge between
second ramus and bulb; i, ridge of second ramus; j, groove of first ramus; Lect, lancet;
Ob, oblong plate; pc, poison canal; PsnSc, poison wack: Ptgr, proctiger; Od, quadrate plate;
Ir, first ramus (basal arm of lancet); 27, second ramus (basal arm of stylet); Sh, sheath lobe;
Stl, stylet; Tri, triangular plate.
7
gO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I03
The basal apparatus of the sting presents on each side three plates —
(fig. 31 A). Uppermost is the large quadrate plate (Qd), which, —
as already explained, is a lateral remnant of the ninth abdominal —
tergum. Below the quadrate plate, and partly overlapped by its —
lower margin, is a horizontally elongate oblong plate (Ob). Above }
the anterior end of the oblong plate and before the quadrate plate
is a small triangular plate (Tri), articulated by the dorsal angle of
its base (c) with the quadrate plate, and by the lower angle (d) with
the upper edge of the oblong plate. The triangular plate and the
oblong plate represent, respectively, the first and the second valvifer
of the ovipositor of other insects. Projecting from the distal end of
each oblong plate is a long, soft appendage (Sh); the pair of ap-
pendages clasp the shaft of the retracted sting and thus form a sheath
for the latter. Between the lower edges of the oblong plates is a
deeply concave membranous fold, which is the unsclerotized ventral
wall of segment JX; it projects posteriorly as a free lobe (JX.V)
when the shaft is depressed.
The shaft of the sting consists of three closely appressed parts
mutually tapering to a point (fig. 31 A). The unpaired upper com-
ponent is the stylet (Stl), the paired lower elements the Jancets (Let).
The lancets are the so-called first valvulae of an ovipositor, the
stylet represents the united second valvulae. Proximally the stylet
is expanded in a bulblike enlargement (b/b), which contains a cavity
open below and continued as a ventral groove to the end of the
stylet. The curved arms that unite the shaft with the supporting base
of the sting are composed each of two parts. The first (zr) is a
proximal ramus of the lancet, connecting the lancet with the apical
angle (b) of the triangular plate (Tri); the second (27) is a cor-
responding ramus of the stylet connecting the base of the stylet bulb
with the anterior end of the oblong plate (Ob). The slender lancets
are held close to the under surface of the stylet by grooves that fit
over tracklike ridges of the latter (G), and the same interlocking
apparatus is continued upon the basal rami, the first ramus being
grooved to its upper end (D, 7), the second having a ridge on its
convex margin (C, 7). The lancets are thus able to slide freely back
and forth on the under side of the stylet. Between the stylet and the
lancets is the poison canal of the sting (G, pc), which expands
proximally into the cavity of the bulb, and here receives the poison
liquid from the great poison sack (A, PsnSc), or reservoir of the
poison glands, which opens into the base of the bulb. Each lancet
bears on its proximal part a valvelike lobe (D, Viv), the two lobes
projecting side by side into the cavity of the bulb, where they serve
feNO. 2 HONEY BEE—SNODGRASS 97
to propel the poison through the poison canal when the sting is in
action. Finally we must note a small forked rod, the furcula (E, Fre,
F), that projects into the body cavity at the base of the bulb above
the neck of the poison sack. The furcula is an apodemal structure
giving attachment to important muscles of the sting (B, Frc).
When the sting is not in use it is concealed within the abdomen,
where its base is supported in the membranous wall of the sting
chamber, and the upper angles of the quadrate plates are loosely
articulated on the spiracular plates of the eighth segment (fig. 32 A).
The shaft is turned upward on the rami so that the bulb is encased
in the membranous fold between the oblong plates, and the distal part
is clasped by the sheath lobes. When the sting goes into action, the
whole structure is swung downward and posteriorly on the dorsal
angles of the quadrate plates (a) until the basal apparatus takes a
vetrical position with the sheath lobes directed upward (fig. 32 B) ;
simultaneously the shaft is depressed and, by the backward swing of
the sting base, is protracted from the tip of the abdomen. The lancets
now begin a rapid, alternate back-and-forth movement on the stylet.
The action of the sting thus involves three separate movements: first
the backward swing of the entire organ, second the depression of the
shaft, third the movement of the lancets. On retraction of the sting,
the shaft is replaced in the sheath, and the base returns to the position
of repose.
The backward swing of the entire organ in the sting chamber was
attributed by the writer in a former paper (1933) to pressure sup-
posedly engendered within the abdomen by contraction of the ab-
dominal segments. More recently, however, Rietschel (1937) has
shown that specifically it is the depression of the seventh abdominal
sternum that projects the sting. Manipulation of a dead specimen, in
fact, will demonstrate that when the seventh sternum is turned down-
ward, its broad anterior part, turning upward and backward, is
pressed deeply into the abdomen against the base of the sting and
forces the latter to swing posteriorly. The shaft, of course, is not
protracted by this action, since it must at the same time be turned at
right angles to the supporting apparatus. The depression of the shaft
is effected by a pair of large, flat muscles arising on the inner surfaces
of the oblong plates (fig. 31 B, 197) that curve anteriorly and mesally
over the base of the bulb and are inserted on the furcula (Frc). The
tension of these muscles on the furcula pulls upward on the base of
the bulb (fig. 32 C), and the shaft is turned downward on a pair of
fulcral points (7) at the lower angles of the bulb that abut against
small lobes on the lower ends of the second rami (2r). The fulcral
98 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
apparatus is shown in greater detail at C and E of figure 31, and the
action of the depressor muscles (197) on the shaft is diagram-
matically expressed at D and E of figure 32. Mesad of the fulcrum —
each ramus of the stylet is continuous with the bulb by a slender, y
flexible connection. The opposite movement of the shaft, by which —
it is again turned upward and ensheathed between the oblong plates —
and their distal appendages, is effected by a pair of slender muscles —
stretched from the second rami to the fulcral lobes of the bulb (figs.
31 B, C, 32C, D, 106), the insertion points of the muscles being —
sufficiently beyond the fulcral points to afford a short leverage on the
shaft.
The movements of the lancets accomplish the penetration of the
tip of the shaft into the victim, and the injection of the poison into
the wound. They are produced by two pairs of large muscles in the
basal apparatus that affect primarily the quadrate plates, and
secondarily the triangular plates and the attached lancets. The muscles
of one pair consist each of two large bundles of fibers (figs. 31 B,
32 C, 198) attached posteriorly, one laterally the other mesally, on
the upper apodemal part of the quadrate plate, and anteriorly on the
anterior end of the oblong plate. The muscles of the other pair (799)
are attached anteriorly by broad, spreading bases on the inner surfaces
of the quadrate plates, and posteriorly on the posterior ends of the
oblong plates. The alternate contraction of these two muscles on each
side pulls the corresponding quadrate plate first forward and then
backward, relative to the oblong plate; the movements of the quadrate
plate are transmitted to the triangular plate (fig. 31 A, T7vi) by the
connection of the former with the upper basal angle of the latter (c) ;
the triangular plate, in turn, rotates on its ventral articulation (d)
with the oblong plate, and the final result is that the alternate up-and-
down movement of the apical angle (b) of the triangular plate is
transmitted to the attached ramus of the lancet (zr), which gives the
lancet itself a back-and-forth movement on the stylet. The mechanism
is illustrated diagrammatically at F and G of figure 32. At F muscle
199, the retractor of the stylet, is contracted, the quadrate plate con-
sequently pulled backward, the apex of the triangular plate lifted, and
the lancet (Let) retracted by the pull on its basal ramus. At G muscle
198, the protractor of the lancet, is in contraction, the quadrate plate
is pulled forward, the triangular plate depressed, and the lancet pro-
tracted. In a living bee, as may be seen in a freshly extracted sting,
the movements of the quadrate plates appear as a rapid vibration, and
the two lancets have alternately opposite motions.
NO. 2 HONEY BEE—SNODGRASS 99
When a bee stings, the initial insertion of the tip of the out-thrust
shaft into the skin of the victim is probably effected by a quick
movement of the abdomen. The subsequent deeper penetration is the
result of the successive alternating movements of the lancets, which
Fic. 32—Mechanism of the sting.
A, sting in position of retraction within sting chamber of abdomen. B, sting
protracted. C, right half of basal parts of sting, mesal view, shaft partly de-
pressed, showing muscles of right side. D, shaft of sting, held in usual position
by contraction of ramus muscle (706). E, same in protracted position, shaft
depressed on basal hinge with ramus (/) by contraction of furcular muscle
(197). F, diagram of sting and muscles of oblong plate: lancet retracted by
contraction of muscle 199. G, same, lancet protracted by contraction of
muscle 108.
Lettering as on figure 31.
at each deeper insertion hold their gain by means of recurved barbs
on their distal ends. The stylet, after the first jab, passively follows
the lancets. The point of the shaft thus automatically goes deeper and
deeper into the wound by the action of the lancets, and continues to
100 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
do so even when the sting is detached from the body of the bee. At
the same time the contents of the poison sack are poured into the bulb
of the sting and driven through the poison canal by the action of the
valves on the bases of the lancets. The poison escapes through a
ventral cleft near the tips of the lancets.
VIII. ANNOTATED LIST OF MUSCLES
The terms origin and insertion are conventional in descriptive
myology for denoting muscle attachments respectively at the fixed
and the movable ends of the muscle. Literally there is no “insertion”
of insect muscles, the muscle fibers being attached directly on the
integument by means of cuticular tonofibrillae. Insect tendons, more-
over, are merely slender apodemal ingrowths of the integument. The
incongruous expression “inserted on’ has thus become current in
entomological writing, referring in some cases to the muscle attach-
ment proper by tonofibrillae, in others to the attachment of the
apodemal tendon on the integument.
In the following list of the bee’s muscles reference is made prin-
cipally to the work of Morison (1927) on the muscles of the honey
bee, and to that of Duncan (1939) on the wasp. Citations of the more
fragmentary work of earlier writers are given by Morison, in part by
the writer (1933, 1941), and need not be repeated. In general there
will be no difficulty in identifying the muscles listed here with those de-
scribed by Morison, notwithstanding differences in functional names;
a few discrepancies are noted.
I. Adductor of the labrum.—A slender muscle arising on the frontal
region between bases of antennae, inserted by a long tendon laterally
on posterior margin of base of labrum. (Levator labri Morison.)
2, 3, 4, 5. Extrinsic muscles of the antenna (fig. 2 A, C).—Four
muscles arising on anterior arm of tentorium, inserted on base of
antenna.
6. Levator of the antennal flagellum (fig. 2H).—\A slender muscle
arising on dorsal wall of scape, inserted dorsally on base of pedicel. |
7. Depressor of the antennal flagellum (fig. 2H).—Origin on
ventral wall of scape, insertion ventrally on base of pedicel.
8. Abductor of the mandible (fig. 31).—A flat muscle arising by
spreading fibers on genal and postgenal walls of cranium behind
lower part of compound eye, inserted by strong apodeme in membrane
between mandibular base and pleurostomal margin of cranium (fig.
3E, 8Ap).
NO. 2 HONEY BEE—SNODGRASS IOI
g. Adductor of the mandible (fig. 3 H)—A larger muscle than
the last, fibers arising in several groups behind upper part of com-
pound eye, on postgena below level of foramen magnum, and one
small group (9a) on top of head, inserted by strong apodeme in
articular membrane mesad of mandibular base (fig. 1 C, 9Ap).
10. Promotor of the maxilla; cardinal protractor of the proboscis
(fig. 4 A).—Origin posteriorly on inner face of postgenal inflection
forming lateral wall of proboscis fossa, inserted on short arm of
cardo beyond articular process.
II, 12, 13. Adductors of the maxilla; stipital protractors of the
proboscis (fig. 4A).—Three muscles arising on extreme anterior
end of anterior tentorial arm, rz inserted on proximal end of stipes,
12 and 13 on mesal margin of stipes. These muscles are primarily
adductors of the maxilla, but because of the angulation between cardo
and stipes they become protractors, and in the bee protract the entire
proboscis.
14. Muscle of the maxillary palpus (fig. 4 B, F).—A large muscle
arising in stipes (F), inserted on base of palpus. Morison describes
this muscle as an extensor of the galea inserted on the base of the
latter near the palpus. In Vespula, according to Duncan, there are
two muscles with a common insertion on the palpus.
15. Flexor of the galea (fig. 4 F).—A large pinnate muscle arising
posteriorly in stipes, inserted by long tendon laterally on base of galea.
The galea of the bee, as in insects generally, has no extensor muscle.
16. Flexor of the lacinia; tensor of the lacinial lobe (fig. 4 F, G).—
Origin in base of stipes, insertion on lever in base of lacinia; tension
of this muscle on the lever raises the lacinial lobe (/cl) and holds it
taut. (Spanner des Segelhalters Wolff ; flexor of the lacinia Duncan ;
not given by Morison.)
17. Anterior adductor of the labium; retractor of the proboscis
(fig. 7A, D).—A long muscle arising on dorsal wall of cranium,
tapering downward to strong tendon inserted on distal end of ligular
bar of prementum (/). It retracts the entire proboscis, also the ligula
into the prementum, and recurves the glossal tongue. (Retractor
linguae longus Wolff; retractor ligulae superior Morison; flexor of
the paraglossa Duncan.)
18. Posterior adductor of the labium; protractor of the labium
(fig. 7D).—Origin on extreme anterior end of anterior tentorial
arm, inserted with muscle of opposite side by common tendon at-
tached on base of prementum. Serves to adduct the labium, and
probably also protracts it by flattening the angle between prementum
102 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
and postmentum. (Protractor labii Morison; posterior labial adductor
Duncan. )
19. Premental retractor of the ligula (fig. 7 A).—Arises laterally
in base of prementum, tapers to tendinous insertion on ligular arm
(h) of prementum just laterad of 177. This muscle is probably the
paraglossal muscle of the labium; functionally it is accessory to 17
in retraction of the ligula’ (Retractor linguae brevis Wolff ; anterior
flexor of the ligula Duncan; wrongly described by Morison as flexor
palpi maxillaris.)
20. Flexor of the glossa, retractor of the tongue (fig. 7A, B).—
Origin posteriorly in base of prementum, insertion by long tendon
close to side of base of glossal rod. The muscles of this pair serve
to shorten the tongue or to roll it from side to side. (Smaller branch
of retractor linguae biceps Wolff ; retractor ligulae inferior Morison;
posterior flexor of the ligula Duncan.)
21. Flexor of the labial palpus (fig. 8 A).—A flat pinniform muscle
arising on side of prementum, inserted by long tendon on rod in
posterior wall of palpiger. It depresses the palpus and folds the latter
back behind the stipes. (Extensor palpi maxillaris Morison ; depressor
of the labial palpus Duncan. Morison mistakenly assigns two muscles
to the labial palpus in the bee. Duncan finds but one in Vespula.)
22. Muscle of the second segment of the labial palpus (fig. 8 A).—
Origin in base of first palpus segment, insertion on base of second
segment ; flexes the second segment mesally, opposed by elasticity of
the joint.
23. Dilator of the salivary syringe (fig. 7A, C).—A flat muscle
arising on anterior lateral margin of prementum, inserted medially on
anterior wall of salivary syringe. (Protractor liguae Wolff; dilator
ampullae superior Morison; dilator of the salivarium Duncan.)
24. Compressor of the salivary syringe (fig. 7A, C).—Largest
muscle in prementum, arising proximally, inserted distally on lateral
margins of salivary syringe. Probably compresses the syringe by
lateral tension. (Principal branch of retractor linguae biceps Wolfe;
dilator ampullae salivae posterior Morison; posterior muscle of
salivarium Duncan.)
25. Retractor, or levator, of the epipharynx (fig. 10 C).—A group
of fibers arising on distal part of clypeus, diverging to insertions
within the epipharynx. (Levator veli palatini Wolff.)
26, 27, 28, 29, 30. Dilators of the suctorium (fig. 10 ©} .—Five
large, paired bundles of short fibers arising on clypeus and inserted
on anterior wall of cibarial region of sucking pump.
NO. 2 HONEY BEE—SNODGRASS 103
31. Compressors of the suctorium (fig. 10 C).—Five thick bundles
of fibers on anterior wall of cibarial region of sucking pump, the
first transverse over the mouth, the others oblique between the dilators,
attached laterally on oral plate (op/).
32. Protractor of the oral plate (fig. 10 B, C).—Origin ventrally
on a process of frontal aspect of base of anterior tentorial arm, in-
sertion dorsally on upper part of pharyngeal arm (y) of oral plate
above frontal ganglion connective. (Lateral pharyngeal muscle
Duncan. )
33. Retractor of the oral plate (fig. 10 B, C)—A smaller muscle
than the last arising medially on frons below median ocellus, inserted
ventrally opposite 32 on arm of oral plate. (No corresponding muscle
given by Duncan in the wasp.)
34, 35. Precerebral dilators of the pharynx (fig. 10 C).—Two
small muscles arising on frons, inserted on pharyngeal region of suck-
ing pump just above frontal ganglion. (Frontal dilators of the anterior
pharynx Duncan.)
36. Postcerebral muscle of the pharynx (fig. 10 C).—Long slender
muscle arising on vertex, inserted laterally on upper extremity of
pharynx, probably mainly suspensory in function. (Dorsal dilator of
the posterior pharynx Duncan.)
37. Posterior contractor of the pharynx (fig. 10C).—A long,
median, fusiform muscle lying against posterior wall of pharynx, aris-
ing by slender tendon on median process of tentorial bridge, inserted
ventrally on inner edge of oral plate. (Posterior dilator of the anterior
pharynx Duncan.)
38. Parietal contractor of the pharynx (fig. 10 C).—A broad sheet
of fibers spreading over lateral and posterior walls of pharynx from
two points of origin laterally on tentorial bridge, fibers going beneath
circular muscles on lower half of pharynx to attachments on arms
of oral plate. This muscle and the last are termed “dilators” of the
pharynx in the wasp by Duncan, but their lengthwise distribution
would suggest that they are contractors of the pharynx, though at the
same time they may dilate the cibarium.
39. Constrictors of the pharynx (fig. 10 C).—The layer of circular
fibers investing the entire pharyngeal region of the sucking pump,
continued on the oesophagus.
40. Superior phragmatic levator of the head (fig. 12 A).—One of
a pair of submedian, parallel muscles arising on prephragma of meso-
notum, inserted laterally on dorsal margin of foramen magnum.
(Levator capitis horizontalis Morison.)
t
104 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
41. Inferior phragmatic levator of the head (fig. 12 A).—One of
a pair of long muscles arising laterally on prephragma of mesonotum,
converging beneath 40 to insertion in dorsal angle of foramen
magnum. (Rotator capitis superior Morison.)
42. Pleural levator, or rotator, of the head (fig. 12 B, D).—A large,
three-branched muscle arising on episternum, inserted by common
tendon attached in laterodorsal notch of foramen magnum; none of
its fibers observed to arise on endosternum as stated by Morison.
(Rotator capitis inferior Morison; pleural elevator of the head
Duncan. )
43. Endosternal levator, or rotator, of the head (fig. 12 B, C).—
A large muscle with wide base on dorsal surface of prothoracic endo-
sternum, fibers convergent on long thick tendon attached in latero-
dorsal notch of foramen magnum above 42. (Levator capitis obliquus
Morison; furcal elevator of the head Duncan.)
44. Depressor of the head (fig. 12B).—A large muscle of two
parts arising on dorsal surface of prothoracic endosternum anterior
to 43, inserted by strong tendon on ventral margin of foramen
magnum. (Depressor capitis horizontalis Morison; first and second
depressors of the head Duncan.)
45. Intersegmental dorsal muscle of the prothorax (figs. 11 F,
12 A).—A thick muscle arising on middle of pronotum, attached
posteriorly on prephragma of mesonotum; probably depresses the
pronotum. (Depressor prothoracis Morison; inner pronotal retractor
of the mesonotum Duncan. )
46. Phragmatopleural muscle of the prothorax (fig. 12 A).—
Arises farthest laterad on prephragma of mesonotum, attached an-
teriorly on cervical apodeme of pleuron. (Lateral branch of levator
prothoracis internus Morison; anterior notal elevator of the pro-
pleuron and head Duncan.)
47. A small prothoracic muscle attached close to 46 on cervical
apodeme of pleuron (fig. 12 A, B) may be a branch of 48, though its
fibers appear to run along the under side of the latter.
48. Tergoepisternal muscle of the prothorax (fig. 12 A)—A
large, straplike muscle arising mesally on posterior inflection of pro-
notum (not on phragma), attached anteriorly on horizontal apodeme
of episternum (A, B). (Levator prothoracis externus Morison.)
49, 50. Protractors of the propectus (fig. 12B, C)—Two fan-
shaped muscles arising anteriorly on side of pronotum, inserted on
apex of pleural apophysis. (Depressor episterni prothoracis anterior
and posterior Morison; first and second posterior notal protractors of
the propleuron Duncan.)
NO. 2 HONEY BEE—SNODGRASS I05
51. Adductor of the propleuron (fig. 12 B).— A large muscle aris-
ing mesally on straplike apodeme of prothoracic endosternum (fig.
11 E, 1), inserted anteriorly on cervical apodeme of pleuron. (Re-
tractor episterni prothoracis Morison; prothoracic furco-pleural
muscle Duncan.) The muscles of this pair cannot be retractors of
the episterna because of the endosternal attachments of the pleura.
52. Intersegmental ventral muscle of the prothorax; endosternal
retractor of the propectus (fig. 12 C).—A slender muscle attached
anteriorly on base of proendosternum, posteriorly on supraneural
bridge of mesoendosternum (fig. 22C). (Retractor prothoracts
Morison; first furcal retractor of the propectus Duncan, the second
retractor of Vespula absent in Apis.)
53. Lateral promotor of the fore coxa (fig. 13 C, D, G).—A broad
muscle arising on episternum, inserted laterally on coxa anterior to
pleural articulation.
54. Mesal promotor of the fore coxa (fig. 13 C, D)—A small
muscle arising on side of endosternum, inserted on anterior mesal
angle of coxal base.
55. First lateral remotor of the fore coxa (figs. 12 C, 13 C, D, G).—
Longest of the coxal muscles, arises laterally on pronotum at base
of spiracular lobe (fig. 17 C), inserted on outer rim of coxa behind
pleural articulation.
56. Second lateral remotor of the fore coxa (figs. 11 C, 12C,
13 C, D, G).—A large muscle arising on under surface of supra-
neural bridge of endosternum (figs. 11 C, 12 C), inserted on margin
of coxal base close behind 55 (fig. 13 D, G).
57. Mesal remotor of the fore coxa (figs. 11 C, 13 C, D).—Smallest
of the coxal muscles, arising on upper end of pleural apophysis (fig.
11 C), inserted mesally on posterior edge of coxal base (fig. 13 C, D).
58. Coxal retractor of the propectus (figs. 11 C, 12 C, 13 C, D, G).
—A large muscle arising posteriorly on anterior part of median
ridge of pterothoracic endosternum (fig. 22 C), the pair diverging
forward to insertions on fore coxae mesad of pleural articulations
(fig. 13 C, D). (Extensor coxae prothoracis Morison; mesosternal
retractor of the propectus Duncan.)
59. Anterior levator of the fore trochanter (fig. 13 E, H).—Origin
in coxa, inserted anteriorly on base of trochanter beyond coxal
articulation.
60. Posterior levator of the fore trochanter (fig. 13 E, H).—Origin
in coxa, inserted posteriorly on base of trochanter beyond coxal ar-
ticulation. Morison mentions only one trochanteral levator, but two
are present in both the drone and the worker.
106 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
61. Thoracic depressor of the fore trochanter (figs. 11 C, 12C,
13 D, E, H).—A long muscle with origin on pleural apophysis of
prothorax (not on endosternum as stated by Morison), going down-
ward into coxa where fibers inserted on depressor apodeme of
trochanter.
62. Coxal depressor of the fore trochanter (fig. 13 E, H).—A
broad muscle occupying entire mesal wall of coxa, fibers inserted on
depressor apodeme and apodemal plate of trochanter (fig. 13 E).
63. Reductor of the fore femur (fig. 13 H).—A short broad
muscle arising on anterior wall of trochanter, inserted somewhat
ventrally on posterior margin of base of femur.
64. Extensor of the fore tibia—Origin dorsally in femur, inserted
dorsally on base of tibia.
65. Flexor of the fore tibia—Origin ventrally in femur, inserted
on ventral genuflexor plate at base of tibia.
66. Productor of the fore tarsus (fig. 14 E).—A flat muscle arising
~ on anterior wall of tibia, inserted anteriorly on base of tarsus.
67. Reductor of the fore tarsus (fig. 14 E).—A flat muscle arising
on posterior wall of tibia, inserted posteriorly on base of tarsus.
68. Depressor of the fore tarsus (fig. 14 D, E).—A longer muscle
than the last two, arising ventrally in tibia, inserted ventrally on base
of tarsus.
69. Flexor of the pretarsal claws of the fore leg—Fibers in two
groups arising one in femur, the other in tibia, inserted on long
slender tendon traversing tarsus, tibia, and femur from attachment
at base of unguitractor plate of pretarsus.
70. External longitudinal dorsal muscle of the mesothorax (fig.
16 D).—A small submedian muscle traversing the mesocutellum.
71. Internal longitudinal dorsal muscle of the mesothorax ; indirect
depressor of the wing (fig. 16 A, C).—A huge muscle attached an-
teriorly on prephragma of mesonotum and on median area (fig. 16 B)
of scutum, attached posteriorly on postphragma of mesonotum.
72. Tergosternal muscle of the mesothorax ; indirect.elevator of the
wing (fig. 16 A, C).—A thick column of fibers attached dorsally on
lateral area of mesoscutum (fig. 16 B), attached ventrally on sternal
area of pleurosternal region.
73. Occlusor of the first spiracle (fig. 17 C, F)—A slender muscle
arising on anterior margin of mesepisternum (C), inserted in spiracu-
lar membrane in front of spiracular aperture (F).
74. Pleuroalar muscle of the mesothorax (fig. 19 C).—A small,
two-branched muscle arising on anterior margin of mesepisternum
NO. 2 HONEY BEE—SNODGRASS 107
(fig. 17 C), inserted on ventral side of humeral complex of first wing
pae) 5O'C ).
75. Pleurotergal muscle of the mesothorax (fig. 17 E).—A large,
fan-shaped muscle arising on episternum, the fibers convergent to
short tendon attached on lateral margin of posterolateral plate of
scutum (fig. 19 F). (Retractor scutelli mesothoracis Morison.)
76. Pleuroaxillary muscles of the mesothorax; flexors of the fore
wing (figs. 17 E, 19 H).—Three small muscles arising on meso-
pleuron, inserted by separate tendons on third axillary of wing base.
77. Basalar muscle of the mesothorax (figs. 17 E, 19 G).—A broad
muscle with spreading fibers arising ventrally on anterior part of epi-
sternum, inserted dorsally by strong tendon on basalar sclerite.
78. Muscle of the axillary lever of the mesothorax (fig. 20 A, B).—
Origin by broad base ventrally on mesothoracic arm of pterothoracic
endosternum, insertion dorsally on end of lever sclerite of fourth
axillary. (Musculus furco-lateralis mesothoracis Morison.) In non-
apoid Hymenoptera this muscle is a muscle of the postphragma (fig.
20°C to.H).
79. Pleurosternal muscle of the mesothorax (fig. 17 G).—A mass
of short fibers arising on posterior epimeral area and on pleural
apophysis of mesothorax, attached on end of mesothoracic arm of
pterothoracic endosternum.
80. Lateral promotor of the middle coxa (figs. 17 G, 22C)—A
slender muscle arising on the small pleural apophysis of mesopleuron,
inserted laterally on coxal base anterior to pleural articulation.
81. Mesal promotor of the middle coxa (fig. 22 C).—Smallest of
the mesocoxal muscles, arising on basal plate of endosternum, in-
serted mesally on anterior margin of coxa.
82, Lateral remotor of the middle coxa; coxosubalar muscle of the
mesothorax (figs. 17 G, 22 C).—A long muscle attached ventrally on
base of coxa just posterior to pleural articulation, attached dorsally
by long tendon on subalar sclerite. Probably has little motor effect
on the coxa; deflects the wing posteriorly by indirect tension on the
third axillary. (Flexor coxae mesothoracis Morison; no correspond-
ing muscle given by Duncan in mesothorax of the wasp.)
83. Mesal remotor of the middle coxa (fig. 22 C).—Largest of the
coxal muscles, origin on basal plate of endosternum, insertion mesally
on posterior margin of base of coxa.
84. Anterior levator of the middle trochanter (fig. 22 D) —Origin
in lower part of coxa, insertion anteriorly on base of trochanter beyond
coxal articulation.
8
108 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
85. Posterior levator of the middle trochanter (fig. 22 D).—Origin
posteriorly in coxa, insertion posteriorly on base of trochanter beyond
coxal articulation.
86. Thoracic depressor of the middle trochanter (fig. 22 C, G).—
Origin dorsally on side of endosternum (C), insertion on end of
depressor apodeme of trochanter (G).
8&7. Anterior coxal depressor of the middle trochanter (fig. 22 D,
G).—Origin anteriorly in base of coxa (D), insertion on base of
depressor apodeme of trochanter (G).
88. Posterior coxal depressor of the middle trochanter (fig. 22 D,
G).—Origin posteriorly in base of coxa (D), insertion on base of
depressor apodeme of trochanter (G).
89. Reductor of the middle femur (fig. 22 D, G).—Origin antero-
dorsally in trochanter, insertion posteroventrally on base of femur.
90. Extensor of the middle tibia (fig. 22L)—A long pinnate
muscle arising on dorsal wall of femur, inserted by axial tendon on
dorsal process of base of tibia.
o1. Flexor of the middle tibia (fig. 22 J) —A muscle similar to but
larger than the extensor, arising on ventral wall of femur, inserted by
its tendon on ventral genuflexor plate (gf) at base of tibia.
92. Productor of the middle tarsus (fig. 22 E).—A slender pinnate
muscle arising anteriorly in tibia, inserted by tendon anteriorly on
base of tarsus.
93. Reductor of the middle tarsus (fig. 22 E).—Similar to the pro-
ductor, arising posteriorly in tibia, inserted posteriorly on base of
tarsus.
94. Depressor of the middle tarsus (fig. 22 F).—A broad pinnate
muscle arising on ventral wall of tibia, inserted ventrally on base of
tarsus.
95. Flexor of the pretarsal claws of the middle leg—Same as in fore
leg and hind leg, one group of fibers arising in base of femur, another
in tibia, inserted by long tendon on unguitractor plate at base of
pretarsus.
96. Retractor of the mesothoracic postphragma (figs. 16 C, 27 C).—
A small fan-shaped muscle arising by narrow end on posterior median
process of propodeum, fibers spreading anteriorly and laterally to
attachments on side of postphragma of mesothorax. (Musculus re-
tractor mesophragmati Morison, mesophragma retractor muscle
Betts.) The muscles of this pair probably are the longitudinal dorsal
muscles of the metathorax with their propodeal attachments shifted
posteriorly.
NO. 2 HONEY BEE—SNODGRASS IO0Q
07. First tergosternal muscle of the metathorax (fig. 21 F, G).—
Origin by broad base on metathoracic arm of endosternum, insertion
on anterior angle of lateral plate of metanotum (E).
98. Second tergosternal muscle of the metathorax (fig. 21 F, G).—
Origin on metathoracic endosternal arm posterior to 97, insertion on
posterior angle of lateral notal plate (E).
99. Third tergosternal muscle of the metathorax (fig. 21 F).—At-
tached ventrally by narrowed end on small tongue (c) of metathoracic
arm of endosternum, fibers spreading dorsally to triangular lateral
area of metanotum.
z00. Pleuroaxillary muscle of the metathorax; flexor of the hind
wing (fig. 21 B, H).—A small, three-part, fan-shaped muscle arising
on anterior marginal inflection of metapleuron, fibers converging to
small nodule in membrane of wing base (a) close to third axillary.
Jor. Basalar muscle of the metathorax (fig. 21 H).—A large muscle
arising by broad base on apodemal lobe of metathorax behind meso-
coxal cavity (fig. 16H, r), inserted dorsally by a strong tendon on
basalar sclerite (fig. 21 B).
102. Pleurosubalar muscle of the metathorax (fig. 21C, H)—A
broad, thick, fan-shaped muscle arising on metapleuron, fibers con-
vergent to narrow insertion on subalar sclerite. (Not given by Mori-
son; two corresponding muscles in Vespula, Duncan.)
103. Lateral promotor of the hind coxa (fig. 24 A, B).—A fan-
shaped muscle arising on ridge between mesopleuron and propodeum,
the convergent fibers inserted laterally on anterior margin of coxa.
104. Mesal promotor of the hind coxa (fig. 24 A, B).—Origin on
metasternal component of pterothoracic endosternum, insertion
mesally on anterior coxal margin.
105. Lateral remotor of the hind coxa; coxosubalar muscle of the
metathorax (fig. 24 A, B).—A long muscle attached below on coxa
immediately behind pleural articulation, inserted dorsally by long
tendon on apodeme of subalar sclerite (fig. 21 B, C). Deflects the
wing posteriorly. (Wrongly said by Morison to arise on endo-
sternum. )
106. Mesal remotor of the hind coxa (fig. 24 A, B)—Origin by
wide base on metasternal component of pterothoracic endosternum,
inserted by broad tendon in articular membrane close to posterior
mesal angle of base of coxa.
107. Anterior levator of the hind trochanter (fig. 24 C, D).—Origin
anteriorly in coxa, insertion by tendon in dorsal membrane of coxo-
trochanteral joint beyond anterior coxal articulation (D).
Ilo SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
108. Posterior levator of the hind trochanter, (fig. 24C, D).—
Origin posteriorly in coxa, insertion in dorsal membrane of coxo-
trochanteral joint beyond posterior coxal articulation (D).
109. Thoracic depressor of the hind trochanter (fig. 24 A, C).—
Origin on metathoracic arm of pterothoracic endosternum (A),
fibers convergent into coxa to insertion on long depressor apodeme
of trochanter (D, Ap).
110. Coxal depressor of the hind trochanter (fig. 24 C, E).—A
broad muscle arising ventrally in coxa, the convergent fibers inserted
on base of depressor apodeme of trochanter (E).
111. Reductor of the hind femur (fig. 24C).—A short thick
muscle arising anterodorsally in trochanter, inserted ventroposteriorly
on apodemal inflection of base of trochanter (F, Ap).
112. Extensor of the hind tibia (fig. 24H)—A long pinnate
muscle arising along dorsal wall of femur, inserted by tendon on
dorsal knob of base of tibia.
113. Flexor of the hind tibia (fig. 24 H).—A large pinnate muscle
lying in ventral half of tibia, fibers attached anteriorly and posteriorly,
inserted by strong axial tendon on ventral genuflexor plate at base
of tibia, (G, H, gf).
114. Promotor of the hind tarsus (fig. 24 J, K, L).—Origin ven-
trally on anterior wall of wide distal part of tibia (K, L), inserted
anteriorly on base of tarsus (J).
115. Levator of the hind tarsus (fig. 24 J, K, L).—A fan-shaped
muscle arising on posterior wall of distal part of tibia (K), inserted
posteriorly by narrow stalk on base of tarsus but above level of tibial
articulation (J). Corresponds with the reductor of the other legs.
116. Depressor of the hind tarsus (fig. 24 J, K, L)—Much larger
than the other two tarsal muscles, arises on anterior wall of tibia (K),
inserted ventrally on base of tarsus (J, L).
117. Flexor of the pretarsal claws of the hind leg (fig. 24 H, K).—
As in the other legs, a two-branched muscle, one branch arising
posteriorly in base of femur (H, 117a@), the other posteriorly in base
of tibia (K, 117b), both branches inserted on long apodemal tendon
extending through tarsus and tibia into femur from its origin in
articular membrane at base of unguitractor plate of pretarsus, as in
middle leg (fig. 25 B, 95Ap).
118. Median intersegmental ventral muscle of the metathorax;
median depressor of the abdomen (fig. 27 E).—A long muscle aris-
ing anteriorly on metasternal component of pterothoracic endosternum,
inserted posteriorly by broad tendon medially on sternum of petiole
segment (J/S).
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NO. 2 HONEY BEE—SNODGRASS er
11g. Lateral intersegmental ventral muscle of the metathorax;
ventral lateromotor of the abdomen (fig. 27 E).—A smaller muscle
arising on median ridge of metasternum (in drone), inserted by long
tendon laterally on sternum of petiole segment (J/S).
120. Outer dorsal muscle of the propodeum; median levator of the
abdomen (fig. 27 B).—A large muscle with spreading fibers arising
on lateral wall of propodeum, inserted by strong tendon on median
ridge of pocket on dorsal side of petiole (D, F, G). (Petiole muscle
Betts; musculus propodei retractor dorsalis postabdominis Morison.)
121. Inner dorsal muscle of the propodeum; dorsal lateromotor of
the abdomen (fig. 27 B).—A two-branched muscle arising laterally
on propodeum at base of 120, inserted by long thick tendon on lateral
angle of tergal plate of petiole (G).
122. Occlusor of the propodeal spiracle (fig. 27 B).—A _ short
intraspiracular muscle.
123. Dilator of the propodeal spiracle (fig. 27 B).—A long slender
muscle arising from small process on dorsal margin of hind coxal
cavity.
124. Median internal dorsal muscle of abdominal segment IT (fig.
28 A).
125. Lateral internal dorsal muscle of abdominal segment II (fig.
28 A).
1206. External dorsal muscle of abdominal segment IT (fig. 28 A) .—
Runs transversely in fold between tergum // and tergum JJ/I.
127. Occlusor of the spiracle of abdominal segment II—A short
intraspiracular muscle.
128. Dilator of the spiracle of abdominal segment IT (fig. 28 A).—
A long slender muscle arising ventrally on lateral margin of sternum
5
129. Lateral muscle of abdonunal segment II (fig. 28 A).—The
only tergosternal muscle of segment II, being attached on dorsal
surface of sternum it is probably the third lateral of other segments.
130. Median internal ventral muscle of abdominal segment IT (fig.
28 A).—The only internal ventral muscle in segment // of the female.
131. Lateral internal ventral muscle of abdonunal segment II (fig.
29 B).—Present in the male only.
132. External ventral muscle of abdominal segment II (fig. 28 A).
—Runs transversely from posterior part of sternum // to anterior
apodeme of sternum J/I.
The muscles of the next four abdominal segments are repeated
alike in each of these segments, and are present in both the male and
the female; they are numbered in the female (fig. 28 A), and may
I1I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I03
easily be identified by comparison in the male (fig. 29 B), though the
muscles of the male are much larger and the second and third laterals
are not crossed.
133, 144, 155, 166. Median internal dorsal muscles of abdominal
segments III to VI (fig. 28 A).
134, 145, 150, 167. Lateral internal dorsal muscles of abdominal
segments III to VI (fig. 28 A).
135, 146, 157, 168. External dorsal muscles of abdominal segments —
III to VI (fig. 28 A).—Oblique in anterior segments, reversed in
posterior segments; origins posteriorly on anterior tergum, insertion
anterior on anterior apodeme of posterior tergum.
136, 147, 158, 169. Occlusors of the spiracles of abdominal seg-
ments III to VI (not shown in figures).
137, 148, 159, 170. Dilators of the spiracles of abdominal segments
III to VI (fig. 28 A). —Origins on long lateral apodemes of the sterna.
138, 149, 160, 171. First lateral mscles of abdominal segments III
to VI (figs. 26D, 28 A).—Reversed tergosternal muscles arising
ventrally on lower parts of terga, inserted dorsally on lateral apodemes
of corresponding sterna.
139, 150, 161, 172. Second lateral muscles of abdominal segments
IlI to VI (fig. 28 A).—External tergosternal muscles attached
ventrally on lateral margins of the sterna; dorsoventral in the male
(figs. 26 E, 29 B), oblique in the female (fig. 28 A).
140, 151, 162, 173. Third lateral muscles of abdominal segments III
to VI (fig. 28 A).—Internal tergosternal muscles attached ventrally
in posterolateral angles of dorsal surfaces of sternal plates (fig. 26 E),
parallel with second laterals in the male (figs. 26 E, 29 B), crossing
the latter obliquely in the female (fig. 28 A).
I4I, 152, 163, 174. Median internal ventral muscles of abdominal
segments III to VI (fig. 28 A, B).—In each segment the median
ventrals form a V with the convergent ends posterior (B) ; in the male —
these muscles are wide bands of fibers.
142, 153, 164, 175. Lateral internal ventral muscles of abdominal
segments III to VI (fig. 28 A, B).—The lateral ventrals are oblique
in an opposite direction from the median ventrals, the two sets form-
ing in each segment an M.
143, 154, 165, 176. External ventral muscles of abdominal segments
III to VI (fig. 28 A).—Short reversed muscles from posterior parts
of sterna to under surfaces of anterior apodemes of preceding sterna.
The musculature of abdominal segments VII to IX is unlike in the
two sexes, in some cases also between queen and worker, and differs
from that of the preceding segments by the absence of certain muscles
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NO. 2 HONEY BEE—SNODGRASS TLS
and the presence of special muscles in connection with the ovipositor
and the male genital organ.
177. Lateral internal dorsal muscle of abdonunal segment VII.—
In the drone (fig. 29 B) as in preceding segments; in the queen (fig.
28 E) attached on anterior part of spiracular plate of segment VJII;
absent in the worker (C).
178. Median internal dorsal muscle of abdominal segment VII.—
In the drone (fig. 29 B) as in preceding segments; in the queen (fig.
28 E) a small muscle attached on spiracular plate of segment VJ//
near spiracle ; in the worker (fig. 28 C) a much larger muscle similarly
attached.
179. External dorsal muscle of abdominal segment VII.—As in
preceding segments in the drone; in the female a reversed dorso-
ventral muscle from lower edge of tergum VJ/ to upper angle of
spiracular plate of segment VJII (fig. 28 C, E).
180, 181. Spiracular muscles of abdominal segment VII (fig. 28 C).
—As in preceding segments.
182, First and second lateral muscles of abdominal segment VII.—
First lateral present as a reversed muscle inserted dorsally on lateral
apodeme of sternum JI in both the drone and the worker, but very
small in the latter (No. 5b of Rietschel, 1937). The muscle designated
182 in the worker (fig. 28 C) also is attached on the lateral apodeme
of sternum JJ, but arises posteriorly on the tergum and is probably
the second lateral of segment V//.
183. Third lateral muscle of abdominal segment VII (fig. 28 C).—
Present in both the male and the female as a dorsoventral muscle
between tergum and sternum.
1&4. Intersegmental sternotergal muscle of abdominal segment VII
(fig. 28 B, C).—Present in the female only; origin anteriorly on
sternum JJ in line with lateral ventral muscles of preceding seg-
ments (B), inserted posteriorly on anterior ventral angle of spiracular
plate of segment VJ/J.
185. Median internal ventral muscle of abdominal segment VII
(fig. 28 D).—Present in the queen only, attached posteriorly on lateral
genital pouch.
186. Lateral internal ventral muscle of abdominal segment VII
(fig. 29 B, D).—Present in the male only; a short muscle arising on
sternum JJ at base of anterior apodeme (B), inserted posteriorly
on long lateral arm of sternum VJ/I.
187. Median internal dorsal muscle of abdominal segment VIII.—
A short muscle in the male (fig. 29 B) attached posteriorly on apodeme
of small tergite of segment [X ; represented in the worker (fig. 28 C)
IIl4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
and the queen (E) by a muscle (787) from spiracular plate of seg-
ment V J// to quadrate tergal plate of segment /X.
188, Lateral internal dorsal muscle of abdominal segment VIII.—
Absent in the male, perhaps represented in the worker (fig. 28 C)
and the queen (E) by a muscle (788) from upper angle of spiracular
plate to posterior dorsal angle of quadrate plate of segment /X (fig.
37, By),
189. Occlusor of the spiracle of abdominal segment VIII (fig.
28 C, E).—A short intraspiracular muscle.
190. Dilator of the spiracle of abdominal segment VIII (fig. 28 C,
E).—In the worker and the queen the dilator of the spiracle of
segment ’//J arises in the ventral anterior angle of the tergal spiracu-
lar plate, not on the sternum as in preceding segments.
ror. Lateral muscle of abdominal segment VIII (fig. 29 B, D).—
Present in the male only, apparently represents the second tergo-
sternal muscle of preceding segments.
192. Tergovalvifer muscle of abdominal segment VIII —Present
in the female only, arising on posterior ventral angle of spiracular
plate (fig. 28 C, E), goes anteroventrally to insertion on dorsal margin
of triangular plate (first valvifer) of sting (fig. 31 D).
193, 194. Tergovaginal muscles of abdominal segment VIII—Two
slender muscles present in the queen only, arising on anteroventral
part of spiracular plate (fig. 28 E), inserted on dorsal wall of vagina
(BD):
195. Lateral internal ventral muscle of abdominal segment VIII
(fig. 29 B, D).—A short muscle present in the male only, arising
laterally on sternum V’///, inserted on lateral angle of parameral plate
(Ip).
196. Levator of the shaft of the sting (fig. 31 B, C)—A small
muscle arising on middle of ramus of stylet, inserted on small process
(C, g) of base of stylet bulb a short distance beyond hinge (h) be-
tween ramus and bulb.
197. Depressor of the shaft of the sting (fig. 31 B).—A large
muscle arising posteriorly on oblong plate of sting, curving anteriorly
and dorsally over base of stylet bulb to insertion on furcula (Frc).
A small branch of this muscle is described by Rietschel (1937, No.
19b) as arising laterally on the oblong plate.
198. Protractor of the lancet of the sting (figs. 31 B, 32 C).—A
large muscle of two parts attached posteriorly on outer and inner
surfaces of upper, apodemal part of quadrate plate ; attached anteriorly
on anterior end of oblong plate at base of second ramus (2r). Pulls
the quadrate plate forward, depresses the triangular plate, and pro-
NO. 2 HONEY BEE—SNODGRASS II5
tracts the lancet (fig. 32 G). The two parts of this muscle are anatom-
ically two muscles, but functionally a single muscle. The upper part
has mistakenly been said (Betts, Morison, and others) to be attached
on the triangular plate.
199. Retractor of the lancet of the sting (figs. 31 B, 32C)—A
broad fan-shaped muscle arising anteriorly on quadrate plate, inserted
posteriorly on upper edge of oblong plate. Pulls the quadrate plate
backward, elevates the triangular plate, and retracts the lancet (fig.
B2F),
200, 201. Muscles of the proctiger in the female (fig. 31 B).—Two
long slender muscles arising on posterior dorsal angle of the quadrate
plate (tergite of segment /X), inserted laterally and ventrally on
lower wall of proctiger.
202. Median dorsal muscle of abdominal segment IX in the male
(fig. 29 B).—A short muscle from apodeme of small tergite of seg-
ment JX to side of proctiger. This muscle falls in line with the median
dorsals of the preceding segments, and probably is represented by
muscles 200 and 201 in the female.
203, 204. Phallic muscles of the male (fig. 29 D)—Two small
muscles, or divergent parts of one muscle, arising ventrally on para-
meral plate (Jp), inserted dorsally on end of apodeme of penis valve
(pv).
205. Muscle of the mucus gland of the male (fig. 29 D).—A slender
muscle from apodeme of penis valve to mucus gland just laterad of
end of vas deferens. '
ABBREVIATIONS USED ON THE FIGURES
A, anal vein.
Ac, antecosta.
af, antennafer.
An, anus.
ANP, anterior notal wing process.
Ant, antenna.
Ap or ap, apodeme.
apa, apodeme of aedeagus.
Ar, arolium.
arc, arcus
at, anterior tentorial pit.
AT, anterior tentorial arm.
au, auricle.
aux, auxilia.
Ax, axillary sclerite of wing base (144%, 2Ax, 3Ax, 4Ax, first, second, third,
and fourth axillaries).
AxC, axillary cord.
axlv, axillary lever.
i ne ee
116 SMITHSONIAN MISCELLANEOUS COLLECTIONS
Ba, basalare.
bc, bursal cornua, pneumapophysis.
BGld, “alkaline” gland of sting.
bib, biblike suboral flap.
bib, bulb of sting.
Blb, bulb of endophallus.
Br, brain.
Brs, bursa of endophallus.
Bs, basisternum.
Btar, basitarsus.
C, costa, marginal vein of wing.
Ca, corpus allatum.
Cb, cibarium (suctorium of cibario-pharyngeal pump).
cbl, pollen basket (corbicula) of hind tibia.
Ce, corpus cardiacum.
Cer, neck of endophallus.
Cd, cardo.
Clp, clypeus.
cnj, conjunctiva.
Cvx, neck, cervix.
Cz coxa,
CxC, coxal cavity (where coxa removed from body).
CxP, pleural coxal process.
Dej, ductus ejaculatorius.
dicb, dilator muscles of cibarium.
dlphy, dilator muscles of pharynx.
DT, dorsal arm of tentorium.
E, compound eye.
Endst, endosternum.
Enph, endophallus.
Ephy, epipharynx.
Epm, epimeron (epm, subdivision of epimeron).
Eps, episternum.
ER, epistomal ridge.
es, sulcus of epistomal ridge (epistomal “suture”).
Fbl, flabellum.
fc, food canal of proboscis.
FC, food channel on base of proboscis.
Fl, flagellum.
flb, fimbriate lobe of endophallus.
Fm, femur.
For, foramen magnum.
Fr, frons.
Frc, furcula.
FrGng, frontal ganglion.
Fs, furcasternum,
VOL. 103
NO. 2 HONEY BEE—SNODGRASS TT]
Ga, galea.
Ge, gena.
gf, genuflexor plate.
Gls, labial tongue (glossae).
H, head.
HB, hypostomal bridge.
Hphy, hypopharynx.
IbS, infrabuccal sac.
IS, sternal plate of propodeum.
IT, tergal plate of propodeum.
IX.V, venter of ninth abdominal segment.
L, leg.
Lb, labium.
LbPlp, labial palpus.
Lc, lacinia.
Icl, lacinial lobe.
Let, lancet (first valvula).
Lm, labrum.
10, lateral ocellus.
lp, lamina parameralis.
Lr, lorum.
Lsp, lamina spiracularis (tergite of eighth abdominal segment).
lvr, lever of lacinia.
Mb, intersegmental membrane.
MCu, mediocubital vein.
Md, mandible.
MdC, cavity where mandible removed from head.
MdGld, mandibular gland.
mn, manubrium of arolium.
mO, median ocellus.
Mth, mouth.
Mv, maxilla.
M-xPlp, maxillary palpus.
N, notum (thoracic tergum).
nf, neural foramen.
NR, notal ridge.
ns, sulcus of notal ridge (scutoscutellar “suture’’).
Nv, nerve.
O, ocellus.
Ob, oblong plate of sting (second valvifer).
Oc, occiput.
occ, occipital condyle.
Odc, oviductus communis.
Odl, oviductus lateralis.
118 SMITHSONIAN MISCELLANEOUS COLLECTIONS
Oe, oesophagus.
Op, operculum of first spiracle.
opl, oral plate.
pc, poison canal of sting.
Pdc, pedicel of antennal flagellum.
PF, proboscis fossa.
Pge, postgena.
Pgl, paraglossa.
PgL, postgenal lobe.
Ph, phragma (1Ph, first phragma, 2Ph, second phragma).
Phtr, phallotreme.
Phy, pharynx.
PI, pleuron (fl, subdivision of pleuron).
pla, pit of pleural apophysis.
PIA, pleural apophysis.
PIf, palpifer.
Plg, palpiger.
pln, planta.
Plp, palpus.
PIR, pleural ridge.
pls, sulcus of pleural ridge (pleural “suture’’).
Pmr, paramere.
Pmt, postmentum.
PN, postnotum.
Poc, postocciput.
pos, postoccipital suture.
pr, pollen press.
Prb, proboscis.
Prmt, prementum.
PsnS‘c, poison sack.
pt, posterior tentorial pit.
Ptar, pretarsus.
Ptgr, proctiger.
pv, penis valve (lamina aedeagalis).
Qd, quadrate plate of sting (tergite of ninth abdominal segment).
r, ramus of sting (zr, ramus of lancet; 2r, ramus of stylet).
R, radius.
ras, rastellum.
rd, glossal rod.
Rdp, reduplication of posterior margin of notum.
RNv, recurrent nerve.
S, sternum.
Sa, subalare.
SA, sternal apophysis.
sc, salivary canal of tongue.
Scl, scutellum.
VOL. 103
NO. 2 HONEY BEE—SNODGRASS IIQ
Scp, scape.
Sct, scutum (sct, subdivision of scutum).
sf, scutal fissure (trans-scutal “suture’”).
Sga, subgalea.
Sh, sheath lobe of sting.
SID, salivary duct.
SIO, salivary orifice.
Slv, salivarium (salivary syringe).
Sp, spiracle.
spl, spiracular lobe of pronotum.
spmb, spiracular membrane.
SptD, spermathecal duct.
St, stipes.
Stl, stylet (united second valvulae).
Syr, salivary syringe.
T, tergum.
Tar, tarsus.
Tb, tibia.
TB, tentorial bridge.
Tg, tegula.
Tr, trochanter.
Tra, trachea.
Tri, triangular plate of sting (first valvifer).
Un, unguis (lateral claw of pretarsus).
Utr, unguitractor plate.
V, venter.
V Dph, ventral diaphragm.
Vlv, valve on lancet.
W, wing.
WP, pleural wing process.
REFERENCES
ARNHART, L.
1923. Das Krallenglied der Honigbiene. Archiv Bienenkunde, vol. 5,
pp. 37-86, 1 pl.
Betts, ANNIE D.
1923. Practical bee anatomy, 88 pp., 12 pls. The Apis Club, Benson,
England.
BisuHop, G. H.
1920. Fertilization in the honeybee. Journ. Exp. Zool., vol. 31, pp. 225-265,
3 pls.
Duncay, C. D.
1939. A contribution to the biology of North American vespine wasps.
Stanford Univ. Publ., Biol. Sci., vol. 8, No. 1, 272 pp., 54 pls.
120 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Ferris, G. F.
1940. The myth of the thoracic sclerites of insects. Microentomology,
vol. 5, pp. 87-90.
LEUENBERGER, F.
1929. Les abeilles, anatomie et physiologie, 195 pp., 104 figs. Paris.
Morison, G. D.
1927. The muscles of the adult honey-bee (Apis mellifera L.). Quart.
Journ. Micr. Sci., vol. 71, pp. 395-463, 12 figs.
Park, W.
1925. The storing and ripening of honey by honeybees. Journ. Econ. Ent.,
vol. 18, pp. 405-410, 3 pls.
RIETSCHEL, P.
1937. Bau und Funktion des Wehrstachels der staatenbildenden Bienen und
Wespen. Zeitschr. Morph. Okol. Tiere, vol. 33, pp. 313-357, 20 figs.
SHarFer, G. D.
1917. A study of the factors which govern mating in the honey bee. Michi-
gan Agr. Coll. Div. Ent., Techn. Bull., No. 34, 19 pp., 3 pls.
Snoperass, R. E.
1925. Anatomy and physiology of the honey bee, 327 pp., 108 figs. New
York.
1933. Morphology of the insect abdomen. Part II. The genital ducts and
the ovipositor. Smithsonian Misc. Coll., vol. 89, No. 8, 148 pp.,
48 figs.
1941. The male genitalia of Hymenoptera. Smithsonian Misc. Coll.
vol. 99, No. 14, 86 pp., 33 pls., 6 text figs.
Wotrr, O. J. B.
1875. Das Riechorgan der Biene. Nova Acta. Ks]. Leop.-Carol. Deutschen
Acad. Naturf., vol. 38, No. 1, 251 pp., 8 pls.
ZANDER, E.
1911. Der Bau der Biene, 182 pp., 20 pls., 149 text figs. Stuttgart.
Rh @ i : ; ; it BY p Stes
H. G. DEIGNAN -
Associate Curator, Division of Birds
U.S. National Museum -
2 PUBLICATION 3689)
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PUBLISHED BY THE SMITHSONIAN INSTITUTION
SEPTEMBER 4, 1942.
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 103, NUMBER 3
A REVISION OF THE INDO-CHINESE
FORMS OF THE AVIAN GENUS PRINIA
BY
H. G. DEIGNAN
Associate Curator, Division of Birds
U. S. National Museum
(PUBLICATION 3689)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
SEPTEMBER 1, 1942
TBe Bord Gaftimore Press
BALTIMORE, MD., U. 8. A.
A REVISION OF THE INDO-CHINESE FORMS OF
THE AVIAN GENUS PRINIA
By H. G. DEIGNAN
Associate Curator, Division of Birds, U. S. National Museum
”?
One using the “Fauna of British India, Birds,’ 2d edition, can
scarcely fail to be astonished by the unnatural classification employed
in certain families in consequence of Stuart Baker’s use in a generic,
or even subfamilial, sense of characters which, in themselves, are, at
most, of not more than subspecific importance. The pitfalls into which
such false evaluation may lead are nowhere more evident than in his
treatment of the Sylviidae, where, merely on a basis of whether the
species have I0 or 12 rectrices, we find Phyllergates separated from
Orthotomus by 21 genera, while Prinia is divided from Franklinia by
23 and from Cisticola by 24 genera. In the case of Franklinia, one
can find not a single character of generic importance by which the
putative genus may be upheld as distinct from Prinia and, once the
principle is accepted that the number of tail feathers may be related
to geographical variation within the species (as it is, indeed, generally
accepted in the cases of certain African warbler species), one hesitates
to accept even the specific distinctness of the Indian Prinia socialis
from the Indo-Chinese Franklinia rufescens. As for the so-called
genus Suya, which has not one character by which it may be distin-
guished from Prinia, it is hardly necessary to say more than that the.
form called Swya crinigera cooki is only with difficulty differentiated
subspecifically from the Javanese bird accepted by all modern authors
as Prinia polychroa. The seasonal change in tail length, assumed by
Baker to have generic importance in Prinia, Suya, Franklinia, and
Laticilla, is reduced to its proper insignificance by the realization that
such variation gradually disappears, from north to south, even within
the limits of the several species as conceived by Baker himself. In
short, the genus Prinia, in this paper, is considered to be a poorly
defined aggregation of grass warblers, nearly allied (through /ncana)
to those placed by Lynes in Cisticola and, in Asia, embracing all the
forms distributed by Baker among Prinia, Suya, Laticilla, and
Franklinia.
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 103, No. 3
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
The present study has been inspired and facilitated by the valuable
contributions to an understanding of the Indian Priniae made by
Whistler and Kinnear * and by Ticehurst and Whistler.’
For the loan of long series of specimens to be added to those in the
collection of the United States National Museum (U.S.N.M.) and
without which the work would. not have been possible, I am indebted
to the authorities of the Academy of Natural Sciences of Philadelphia
(A.N.S.), of the American Museum of Natural History (A.M.N.H.),
of the Museum of Comparative Zoology (M.C.Z.) and of the Prince-
ton Museum of Zoology (P.M.Z.).
I
The Indo-Chinese populations of Prinia rufescens, hitherto sub-
merged in Franklinia rufescens (Blyth), prove, upon examination of
series in comparable plumages, to fall into no fewer than five well-
characterized races, localized in just those areas where subspecific
differentiation normally appears in numerous other passerine species.
1, PRINIA RUFESCENS RUFESCENS Blyth
Pr{inia] rufescens Brytu, Journ. Asiat. Soc. Bengal, vol. 16, p. 456, 1847
(Arakan).
Front, crown, and nape brownish slate (summer) ; rufous brown,
tinged with slate (winter).
Mantle dark rufous brown (summer); bright rufous brown
(winter).
Well-marked supraloral streak (often continued beyond the eye as
a short supercilium) and the tiny feathers of both eyelids white
(summer and winter).
Under parts white, washed with buff, most strongly along the flanks
and on the thighs and under tail-coverts (summer) ; white, washed
with rufous buff, most strongly along the flanks, over the greater part
of the abdomen, and on the thighs and under tail-coverts (winter).
Tail short in summer, slightly longer in winter.
Range.—In Indo-China (excepting Burma): the northern half of
Indochine; East Siam, south to the river Me Mu’n (U:bon Prov-
ince) ; all North Siam; West Siam, south to lat. 14° N.
Walden has named Prinia beavani from two examples collected at
Shwegun, Salwin River, Tenasserim (Proc. Zool. Soc. London, —
* Journ. Hones Nat. Hist. Soc., vol. 36, pp. 564-566, 573-582, 1933.
* Ibis, 1030, pp. 761-763.
NO. 3 AVIAN GENUS PRINIA—DEIGNAN 3
[ =1867], p. 551); 1 find no points of difference between the birds of
northern Tenasserim and those of other parts of southern Burma and
unqualifiedly pronounce beavani synonymous with rufescens.
2. PRINIA RUFESCENS PENINSULARIS, new subspecies
Type—Adult female, U.S.N.M. No. 160572, collected at Trang,
Peninsular Siam, January 20, 1897, by W. L. Abbott.
- Front, crown, and nape brownish slate (summer) ; brownish slate
or slaty brown (winter).
Mantle dark rufous brown (summer and winter).
Well-marked supraloral streak (often continued beyond the eye as
a short supercilium) and the tiny feathers of both eyelids white
(summer and winter).
Under parts white, washed with buff, most strongly along the flanks
and on the thighs and under tail-coverts (summer and winter).
Tail short (summer and winter).
Range.—Peninsular Siam, from the Isthmus of Kra: south to (but
not including) Pattani Province.
3. PRINIA RUFESCENS EXTREMA, new subspecies
Type.—Adult male, U.S.N.M. No. 308245, collected at Bangna: ra,
Peninsular Siam, July 13, 1926, by H. M. Smith.
Front, crown, and nape dark brownish slate (summer and winter).
Mantle dark rufescent olivaceous brown (summer and winter).
Well-marked supraloral streak white (summer and winter).
Tiny feathers of upper eyelid white, those of lower eyelid mixed
white and slaty (summer and winter).
Under parts white, washed with pale buff, most strongly along the
flanks and on the thighs and under tail-coverts (summer and winter).
Tail short (summer and winter).
Range.—Peninsular Siam (Pattani Province) and the Malay States,
south to Negri Sembilan and Pahang.
4. PRINIA RUFESCENS OBJURGANS, new subspecies
Type.—Adult male, U.S.N.M. No. 334623, collected at Siracha,
southeastern Siam, April 19, 1934, by H. M. Smith.
Front, crown, and nape brownish slate (summer and winter).
Mantle dark rufous brown (summer and winter).
Well-marked supraloral streak white (summer and winter).
Tiny feathers of upper eyelid mixed white and slaty, those of lower
eyelid slaty (summer and winter).
4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Under parts white, washed with buffy cream, most strongly along
the flanks and on the thighs and under tail-coverts (summer and
winter ).
Tail short (summer and winter).
Range.—Southeastern Siam.
5. PRINIA RUFESCENS DALATENSIS (Riley)
Franklinia rufescens dalatensis Ritey, Proc. Biol. Soc. Washington, vol. 53,
p. 79, 1940 (Fimnon, South Annam).
Front, crown, and nape dark brownish slate (summer); dark
rufous brown, tinged with slaty (winter).
Mantle dark rufous brown (summer and winter).
Indistinct supraloral streak slaty or mixed white and slaty (summer
and winter ).
Tiny feathers of both eyelids slaty (summer and winter).
Under parts white, washed with buffy rufous, most strongly along
the flanks and on the thighs and under tail-coverts (summer and
winter ).
Tail short in summer, slightly longer in winter.
Range.—South Annam.
I am tempted to consider Prinia rufescens and its races merely the
more eastern forms of the species named Pr. socialis by Sykes.
Judging by the literature, however, there seems to be considerable
overlapping of ranges in the eastern Himalayas and Assam and, as-
suming that such overlapping is real, rather than merely apparent, I
keep them, at least for the present, distinct specific entities.
II
With the inclusion of the genus Franklinia Jerdon 1863 in Prinia
Horsfield 1821, Prinia gracilis Franklin 1831 is preoccupied by Sylvia
| =Prinia| gracilis Lichtenstein 1823 and Blyth’s Pr[inia| Hodgson
again comes into use for the bird of Central India, thus becoming
untenable for the Himalayan race, which may be called Prinia hodg-
soni rufula Godwin-Austen 1874.
Of Prinia hodgsonii, I find at least four forms in the Indo-Chinese
countries.
1. PRINIA HODGSONII HODGSONII Blyth
Pr{inia] Hodgsonii Brytu, Journ. Asiat. Soc. Bengal, vol. 13, p. 376, 1844.
New name for Prinia gracilis Franklin 1831 (Mirzapur District, by restric-
tion, Ticehurst and Whistler, Ibis, 1930, p. 763), not Sylvia gracilis Lichten-
stein 1823.
NO. 3 AVIAN GENUS PRINIA—DEIGNAN 5
Upper parts nearest hair brown (Ridgway), this color almost pure
on front, crown, and nape, faintly tinged rufescent on mantle, rump,
and upper tail-coverts (summer) ; light olivaceous brown, suffused
with rufescent, faintly on front, crown, and nape, more strongly on
mantle, conspicuously on rump and upper tail-coverts (winter).
Supraloral streak ashy white (summer and winter).
The swollen edges of the eyelids orange and the tiny feathers of
both eyelids white (summer) ; the edges of the eyelids neither swollen
nor brightly colored, the tiny feathers of both eyelids white (winter).
Under parts white, with a broad pectoral band and flanks deep ashy
gray, the throat and center of abdomen very faintly tinged with cream
(summer) ; white (more or less sullied with ashy gray on breast and
upper flanks), washed with cream, which posteriorly changes to light
rufous cream (winter). *
Tail short in summer, longer in winter.
Range.—In Indo-China: Burma, south of Manipur and west of the
Irriwadi River.
2. PRINIA HODGSONII RUFULA Godwin-Austen
Prinia rufula Gopw1n-AusTEN, Proc. Zool. Soc. London, 1874, p. 47 (Naga
Hills and Manipur).
Upper parts between hair brown (Ridgway) and chetura drab
(Ridgway ), this color almost pure on front, crown, and nape, faintly
tinged rufescent on mantle, rump, and upper tail-coverts (summer) ;
dark olivaceous brown, suffused with rufescent, faintly on front,
crown, and nape, more strongly on mantle, conspicuously on rump and
upper tail-coverts (winter).
Supraloral region concolorous with crown (summer) ; supraloral
streak ashy white (winter).
The swollen edges of the eyelids orange and the tiny feathers of
both eyelids slaty (summer) ; the edges of the eyelids neither swollen
nor brightly colored, the tiny feathers of the upper eyelid mixed ashy
gray and white, those of the lower eyelid white (winter).
Under parts white, with a broad pectoral band and flanks deep ashy
gray, the throat and center of abdomen washed with cream (summer) ;
white (more or less sullied with ashy gray on breast and upper flanks),
washed with cream, which posteriorly changes to light rufous cream
(winter ).
Tail short in summer, longer in winter.
Range—lIn Indo-China: Upper Burma, except for those districts
inhabited by hodgsonit.
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I03
3. PRINIA HODGSONII CONFUSA, new subspecies
Type.—Adult male, M.C.Z. No. 129216, collected at Mengtze, SE.
Yunnan, December 5, 1920, by J. D. La Touche.
Upper parts chetura drab (Ridgway), this color almost pure on
front, crown, and nape, faintly tinged rufescent on mantle, rump, and
upper tail-coverts (summer) ; dark olivaceous brown, suffused with
bright rufous, faintly on front, crown, and nape, strongly on the
mantle, very conspicuously on the rump and upper tail-coverts
(winter ).
Supraloral region concolorous with crown (summer) ; supraloral
streak ashy white (winter).
The swollen edges of the eyelids orange and the tiny feathers of
both eyelids slaty (summer) ; edges of eyelids neither swollen nor
brightly colored, the tiny feathers of the upper eyelid mixed ashy gray
and white, those of the lower eyelid ashy white (winter).
Under parts white, with a broad pectoral band and flanks deep ashy
gray, the throat and center of abdomen faintly washed with cream
(summer) ; white (very slightly sullied with ashy gray on breast and
upper flanks), washed with creamy rufous, faintly on throat and
breast but strongly posteriorly (winter).
Tail short in summer, longer in winter.
Range.—Yunnan, Tongking, and adjacent parts of Laos.
In confusa, the northeasternmost form of Prima hodgsonu, we find
the greatest differentiation between summer and winter plumages and,
in the latter, the strongest resemblance to the corresponding dress of
a race of Prinia rufescens inhabiting the same area. It is evidently
this approximation which puzzled Whistler and Kinnear * and which,
at one time, misled Delacour into believing hodgsonii and rufescens
mere color phases of a single species. Despite their similarity, how-
ever, confusa may readily be known from rufescens by a character
which serves equally well to separate all other cohabitant forms of the
two species: in the former, the under side of the rectrices is brownish
ashy, with black subterminal spot and ashy white tip; in the latter, the
under side of the rectrices is ashy brown, with black subterminal spot
and rufous gray tip.
4. PRINIA HODGSONII ERRO, new subspecies
Type.—Adult female, U.S.N.M. No. 350283, collected at Chieng-
mai, 1,000 feet, North Siam, January 24, 1936, by H. G. Deignan.
* Journ. Bombay Nat. Hist. Soc., vol. 36, p. 565, 1933.
NO. 3 AVIAN GENUS PRINIA—DEIGNAN Fi
Upper parts chetura black (Ridgway), this color almost pure on
front, crown, and nape, faintly tinged rufescent on mantle, rump, and
upper tail-coverts (summer) ; front, crown, and nape slaty brown or
brownish slate, changing gradually to dark olivaceous brown, suffused
with rufescent on the lower back, rump, and upper tail-coverts
(winter ).
Supraloral region concolorous with crown (summer) ; supraloral
streak dark ashy, very rarely ashy white (winter).
The swollen edges of the eyelids orange and the tiny feathers of
both eyelids slaty (summer) ; the edges of the eyelids neither swollen
nor brightly colored, the tiny feathers of the upper eyelid ashy gray,
those of the lower eyelid ashy gray or mixed ashy gray and white
(winter).
Under parts white, with a broad pectoral band and flanks deep ashy
gray, the throat and center of abdomen faintly tinged with cream
(summer) ; white (more or less sullied with ashy gray on breast and
upper flanks), washed with cream, which posteriorly changes to
rufous cream (winter).
Tail short (summer and winter, but occasionally longer in winter).
Range.—Kengtung State, North Siam, and Middle Laos.
Prinia hodgsonii occurs also in East and Southeast Siam and in
South Annam but the material from those areas available to me is too
scant to warrant expression of opinion as to subspecies.
In summer dress, all the races of Pr. hodgsonii are very distinct
from those of Pr. rufescens but variation within the species appears
only in the most subtle characters; in winter dress, they are easily
distinguishable from each other but then show a strong, though
superficial, resemblance to those of the other species.
feet
Prinia inornata (with which I consider the African Drymoica
superciliosa Swainson and its races conspecific) is represented in the
Indo-Chinese countries (excluding Burma) by three forms.
1. PRINIA INORNATA BLANFORDI (Walden)
Drymeeca blanfordi Wa vEN, in Blyth, Cat. Mamm. Birds Burma, Journ. Asiat.
Soc. Bengal, vol. 43, pt. 2, extra No., p. 118, 1875 (Toungoo, Burma).
Range.—In Indo-China (excepting Burma): North Siam (Chieng-
mai Province).
This species has been taken in northern Siam only at Chiengmai,
Mu’ang Fang, and Nan. Fine-plumaged January birds from Chieng-
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
mai may quite safely be called blanfordi; a couple of worn summer
specimens from Mu’ang Fang are subspecifically indeterminable ; the
sole example yet known from Nan is discussed below.
2. PRINIA INORNATA EXTENSICAUDA (Swinhoe)
Drymoica extensicauda SwInuHOok, Ibis, 1860, pp. 50-51 (Amoy).
Range.—In Indo-China: Tongking, North Annam, High Laos.
The single worn and discolored specimen from Nan, placed by
Riley * with “exter” because it “matches . . . perfectly’ summer
birds from western Szechuan, in fact agrees equally well with similar
examples from Amoy (extensicauda) and from Mu’ang Fang (pre-
sumably blanfordi). There is a possibility that the birds of Nan (and
perhaps of Mu’ang Fang as well) are indeed extensicauda (from
which, in my opinion, exter is inseparable) ; until, however, speci-
mens in good winter dress are known from those areas, we have not
the least evidence that they differ from the population of Chiengmai,
which is certainly blanfordi.
3. PRINIA INORNATA HERBERTI Stuart Baker
Prinia inornata herberti Stuart BaKker, Bull. Brit. Orn. Club, vol. 38, p. 39,
1918 (Bangkok, Siam).
Range.—East Siam and all Indochine, south of the territory occu-
pied by Pr. 1. extensicauda.
Riley ° says: “In my opinion [herberti] is not a form of inornata
at all, but of Prima blythi of Java, which should not be in the same
form group.” He goes on to list differences between blythi and
imornata, not one of which, from my point of view, is of more then
subspecific value. Actually, the relationship of blythi and herberti to
other members of the imornata group is much like that of polychroa
and cooki to other members of the polychroa group (vide infra).
IV
Prinia flaviventris, in one form or another, is now known to occur,
however rarely, at suitable localities in all parts of Siam and Indo-
chine. The greater part of the subregion is inhabited by a bird which
has always been called Prima fi. flaviventris, but which, while agreeing
perfectly with that race in color, differs from it in its rather heavier
bill and its acquisition in winter of a tail only slightly longer than that
*U. S. Nat. Mus. Bull. 172, p. 443, 1938.
°Idem, pp. 442-443.
NO. 3 AVIAN GENUS PRINIA—DEIGNAN 9
of summer, thus approaching the very strong-billed Pr. fl. rafflesi,
which shows no seasonal variation at all in length of tail. I propose
to name this form
PRINIA FLAVIVENTRIS DELACOURI, new subspecies
Type—Adult male, P.M.Z. No. 10301, collected at Chiengmai,
1,000 feet, North Siam, January 30, 1932, by H. G. Deignan.
The races now known from the Indo-Chinese countries, exclusive
of Burma, are:
1. PRINIA FLAVIVENTRIS SONITANS Swinhoe
Prinia sonitans Sw1nuok, Ibis, 1860, p. 50 (Amoy).
Range.—In Indo-China: NE. Tongking.
2. PRINIA FLAVIVENTRIS DELACOURI Deignan
Range.—Pegu, Siam, and all Indochine, except those areas in-
habited by sonitans and rafflesi.
3. PRINIA FLAVIVENTRIS RAFFLESI Tweeddale
Prinia rafflesi TWEEDDALE, Ibis, 1877, p. 311, pl. 6, fig. 1 (Lampongs, South
Sumatra).
Range.—In Indo-China: Peninsular Siam, south of the Isthmus
of Kra.
V
Since, as mentioned in my introduction, the southernmost race of
Suya crinigera auctorum (type of the genus Swya) is clearly only
subspecifically distinct from Malurus polychrous Temminck 1828
(Java), polychroa must take precedence over crinigera 1836 as the
oldest name for the species.
The recognizable races of Prinia polychroa are seven or eight, de-
pending on whether Suya parumstriata David and Oustalet (Fukhien)
is distinct from the Formosan Prinia striata Swinhoe, which I have
not seen.
1. PRINIA POLYCHROA STRIATULA (Hume)
Blanfordius striatula Hume, Stray Feathers, vol. 1, p. 195, 1872 [nomen nudum] ;
ibid., vol. 2, p. 300, 1873 (Karachi).
2. PRINIA POLYCHROA CRINIGERA (Hodgson)
Suya crinigera Hovcson, Asiat. Res., vol. 19, p. 183, 1836 (Nepal).
10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
3. PRINIA POLYCHROA ASSAMICA (Stuart Baker)
Suya crinigera assamica Stuart Baker, Bull. Brit. Orn. Club, vol. 44, p. 20,
April 26, 1924 (Shillong, Assam). °
Suya crinigera assamica StTuART BAKER, Fauna Brit. India, Birds, 2d ed., vol. 2,
p. 520, May 7, 1924 (no locality given).
4. PRINIA POLYCHROA CATHARIA Reichenow
Prinia catharia REICHENOw, Ornith. Monatsber., vol. 16, p. 13, 1908 (‘‘Ta-tsieng-
lu-ting in Setschuan’’).
Range.—Szechuan and NW. Yunnan.
Feathers of front, crown, nape, and mantle blackish brown, suf-
fused with chestnut and margined brownish ashy to give a boldly
streaked appearance (summer); equally boldly streaked, but the
feathers overlaid with a rich dark rufous wash which masks the light
margins and gives the bird a generally darker appearance (winter).
Two examples from Kiangyin, Kiangsu Province, agree perfectly
with a series of catharia and not at all with a series of parumstriata.
If these birds are not abnormal, it would seem that catharia follows
the Yangtze from Szechuan to its mouth.
5. PRINIA POLYCHROA PARUMSTRIATA (David and Oustalet)
Suya parumstriata Davin and OusTALetT, Oiseaux de la Chine, p. 259, 1877
(Fukhien).
Range.
East Yunnan, Kwangsi, Kwangtung, and Fukhien.
Olivaceous brown above, lightly washed with rufescent or ochra-
ceous, the feathers of front, crown, nape, and mantle with faintly
darker centers which scarcely appear as broad streaks (summer) ;
much like catharia in winter but without the strong dark rufous wash
and with the dark centers of the feathers much less clearly defined,
especially on nape and mantle (winter).
Two specimens from Hunan Province are intermediate between
catharia and parumstriata, having the obsolescent streaking of the
latter race combined with the rufous suffusion of the former.
Parumstriata possesses considerable interest in that the summer
dress shows its close relationship to the following more southern races,
while the winter dress shows its equally close affinity with the pre-
ceding forms.
6. PRINIA POLYCHROA STRIATA Swinhoe
Prinia striata Swinwor, Journ. North-China Branch Roy. Asiat. Soc., vol. 1,
p. 227, 1859 (Hongsan, NW. Formosa).
NO. 3! AVIAN GENUS PRINIA—DEIGNAN ieu
7. PRINIA POLYCHROA COOKI (Harington)
Suya crinigera cooki Hartncton, Bull. Brit. Orn. Club, vol. 31, p. 109, 1913
(Thayetmyo, Burma).
Upper parts dark olivaceous brown, almost free of rufescent or
ochraceous wash, the feathers with faintly darker centers which
scarcely appear as broad streaks (summer) ; similar, but with a faint
rufescent wash and the obsolescent darker centers of the feathers a
little more distinct, although less so than in the corresponding dress
of parumsiriata (winter).
It is remarkable that this species is known from the Indo-Chinese
countries (exclusive of Burma) only in East Siam, Low Laos,
Cambodia, and South Annam. It is still more remarkable that the
isolated population inhabiting those districts should be identical with
that of Thayetmyo but I have been unable to examine topotypical
material of cooki and Dr. Delacour assures me that the two popula-
tions are indeed inseparable.
8. PRINIA POLYCHROA POLYCHROA (Temminck)
Malurus polychrous Temminck, Nouv. Rec. Pl. Col. Ois., vol. 3, [livr. 78],
pl. 466, fig. 3, 1828 (Java).
Polychroa closely resembles cooki of Annam at any season but
differs in having the upper parts with an ashy, rather than a rufescent,
wash. Worn specimens of the two races tend to approximate each
other but the distinction “here given is readily seen in fresh-plumaged
examples.
The relationship between Prinia polychroa and Pr. sylvatica of
India and Ceylon should be investigated by someone with better
material than is available to me.
VI
Chasen’ has already, in my opinion correctly, made the Malaysian
races of “Suya” superciliaris Anderson 1871 conspecific with “Suya”
atrogularis Moore 1854, with the result that atrogularis becomes the
proper name for the species. With the union of the genera Suya and
Prinia, however, the Chinese subspecies becomes Prima atrogularis
superciliaris Anderson 1871 and the Bornean bird hitherto called
Prinia flaviventris superciliaris Salvadori* is left without a valid
name; for it I propose
© Handlist Malaysian Birds, p. 254, 1035.
7 Ann. Mus. Civ. St. Nat. [Genova], ser. 1, vol. 5, p. 240, 1874.
12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
PRINIA FLAVIVENTRIS CHASENI, new name
Three well-defined forms of Prinia atrogularis occur within Indo-
Chinese limits.
1. PRINIA ATROGULARIS SUPERCILIARIS (Anderson)
Saya [sic] superciliaris ANDERSON, Proc. Zool. Soc. London, 1871, pp. 212-213
(Momien [= Tengyueh], Yunnan).
Range.—Northern Burma, Yunnan, Kwangtung, Fukhien, Tong-
king.
2. PRINIA ATROGULARIS ERYTHROPLEURA (Walden)
Suya erythropleura WALDEN, in Blyth, Cat. Mamm. Birds Burma, Journ. Asiat.
Soc. Bengal, vol. 43, pt. 2, extra No., p. 116, 1875 (Toungoo, Burma).
A series of five summer and eight winter birds from northern Siam
and the adjacent parts of Laos differ from superciliaris in having the
upper parts, throughout the year, more olivaceous brown, much less
rufescent brown; the under parts, in winter, with less rufescence and
that of a paler hue; the tail shorter than that of superciliaris at the
same season and showing less seasonal variation in length than appears
in the more northern race.
No topotype of erythropleura has been seen and the name is here
applied to the Siamese population on the assumption that all birds of
the same latitudes are likely, in this species, to be the same. If
erythropleura be synonymous with superciliaris, the series before me
is without an available name.
3. PRINIA ATROGULARIS KLOSSI (Hachisuka)
Suya superciliaris klossi HAcHtsuKA, Bull. Brit. Orn. Club, vol. 47, p. 53, 1926
(Dalat, South Annam).
Range.—Cochinchine, South Annam, Low Laos _ (Bolovens
plateau).
In the Malaysian subregion appear also the two following forms:
1. PRINIA ATROGULARIS WATERSTRADTI (Hartert)
Suya waterstradti Hartert, Noy. Zool., vol. 9, p. 568, 1902 (Gunong Tahan,
Malay Peninsula).
2. PRINIA ATROGULARIS ALBOGULARIS (Hume)
Suya albogularis Hume, Stray Feathers, vol. 1, p. 459, 1873 (Acheen, Sumatra).
The type of Burnesia dysancrita Oberholser, which lies before me,
shows that this name is a complete synonym of Hume’s albogularis.
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VOLUME 103, NUMBER a
ARCHEOLOGICAL AND GEOLOGICAL
ce _ INVESTIGATIONS IN THE SAN |
JON DISTRICT, EASTERN. |
ey MEXICO
(WITH 9 PLATES)
a ee POBYe (bE Gc
uf “FRANK H.. H. ROBERTS, OR. y
Archeologist, Bareau of American Ethnology,
1
ETH
a (PusLicaTion 3692)
“ei talaapaged OTe “ary OF WASHINGTON.
Bee ahs Post ~ PUBLISHED BY THE SMITHSONIAN INSTITUTION.
BS; ae Se 3 _ OCTOBER 12; 1942
SMITHSONIAN MISCELLANEOUS. COLLECTIONS
VOLUME 103, NUMBER 4
ARCHEOLOGICAL AND GEOLOGICAL
INVESTIGATIONS IN THE SAN
TONEDISTRIC 1 EAS TERN
NEW MEXICO
(WITH 9 PLATES)
BY
FRANK H. H. ROBERTS, JR.
Archeologist, Bureau of American Ethnology
(PUBLICATION 3692)
GITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
OCTOBER 12, 1942
The Lord Baltimore Press
BALTIMORE, MD., U. 8. A,
ARCHEOLOGICAL AND GEOLOGICAL INVESTIGATIONS
IN THE SAN JON DISTRICT, EASTERN NEW MEXICO
By FRANK H. H. ROBERTS, Jr.
Archeologist, Bureau of American Ethnology
(Wi1tTH 9 PLATES)
INTRODUCTION
In continuance of a study of early horizons in North American
archeology a Bureau of American Ethnology-Smithsonian Institution
expedition spent the field season, June 20 to September 6, 1941,
conducting investigations in eastern New Mexico. Members of the
party, in addition to the writer, were: Dr. Kirk Bryan, Department
of Geology and Geography, Harvard University ; Herbert J. Dick, Jr.,
University of New Mexico; Robert Easterday, Colorado State
Teachers’ College; Walter B. Greenwood, Bureau of American Eth-
nology; S. Sheldon Judson, Harvard University; Bert E. Lohr and
Edison P. Lohr, Loveland, Colo.; Donald D. McPhail, Michigan
State College; Robert H. Merrill, engineer, Grand Rapids, Mich. ;
Robert M. White, Harvard University; Eugene C. Worman, Jr.,
Harvard University; Beulah J. Lohr, Linda B. Roberts, and Mrs.
Merrill. Numerous interested scientists and amateurs from nearby
southwestern and other institutions visited the camp during the prog-
ress of the work.
Most of the excavations centered around a site on the north rim
of the Staked Plains 10$ miles (16.9 km.) south of the town of
San Jon and 20 miles (32.2 km.) south of the Canadian River.
At this location there is a shallow basin that appears to be a remnant
of an old filled-in lake bed or series of ponds that had formed in
the fill resulting when an original valley in the escarpment was
blocked by sand deposits. The bottom of the basin is traversed
by a series of deep ravines and broad arroyos (pl. 1, fig. 2) that
come together and cut through the rim to join one of the intermittent
tributaries of the Canadian heading in the brakes below. The local
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 103, No. 4
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
name of the formation is Sand Canyon. Heavy erosion of the gully
banks in recent years exposed deposits of alluvium and sporadic
concentrations of animal bones including mammoth, bison, and deer
as well as other smaller mammals, mainly of the Rodentia. Stone
implements occurring near some of these outcroppings indicated that
aboriginal hunters had been active in the area and suggested that
both former camping places and associations between man-made
objects and extinct species of animals could be found there.
The site was discovered by Keith Martin, a local ranchman, who
reported the presence of the bones and showed the artifacts that he
had found there to various people at the Museum of New Mexico
and the Laboratory of Anthropology at Santa Fe, and at the Uni-
versity of New Mexico in Albuquerque. In the spring of 1940 Dr.
Frank C. Hibben and a group of students from the Department of
Anthropology at the University did some preliminary prospecting
there and obtained a collection of animal bones and a few stone
artifacts. In August of the same year the writer visited the site in
company with Dr. Hibben. Because of the amount of work involved
in a thorough investigation and the fact that its efforts were occupied
with other archeological researches, the University of New Mexico
offered to turn the site over to the Smithsonian Institution. The
combination of features at the site and in the adjacent district was
so promising that the offer was accepted. Permission to carry on the
work was obtained from Mrs. George Wilburn, lessee, and Mrs.
H. Bonem, owner, of Tucumcari, N. Mex.
Several factors were considered in making the decision to investi-
gate the site. In addition to the occurrence of artifacts with fossil
bones, there was the matter of the projectile points picked up from
the weathered surfaces by Martin and found in situ by the University
group. They are of the so-called Yuma type, a form purported to be
of some antiquity but about which further data are needed to deter-
mine its true status. Furthermore, the location, some 130 miles
(209.2 km.) south of the original Folsom bison quarry (Figgins,
1927; Roberts, 1935, 1940), and some 60 miles (96.6 km.) north
of the important deposits along Blackwater Draw between Clovis and
Portales where Folsom and Yuma materials occur (Howard, 1935;
Roberts, 1940), as well as its proximity to some of the west Texas
places where reputedly old finds have been made, seemed significant.
In addition the area was the scene of considerable activity during
late protohistoric and early historic times. It was hunting territory
for numerous bands of the Apache—the Querechos of Coronado and
NO. 4 SAN JON DISTRICT, NEW MEXICO—ROBERTS 3
the Vaqueros of Benavides; possibly for some of the Jumanos—the
Patarabueyes of Espejo and the Rayados of Ofiate (Mera, 1935;
Scholes and Mera, 1940); for occasional groups from the eastern
Pueblos; perhaps for some of the Wichita roving that far west;
for the later Comanche and Kiowa; and even some of the Pawnee.
Coronado’s party probably followed the Canadian north of the site
and other Spanish explorers visited the region. One of the main
Comanche trails, the northern, skirted the base of the bluffs.
Numerous survey parties from the eastern States passed that way
during and immediately after the Mexican War; the California
emigrant trail from Fort Smith, Ark., followed the Canadian in
1849; and still later one of the famous trails of the cattle barons
traversed the valley. All contribute a share of interest and enhance
the possibility of adding to the fund of knowledge pertaining not
only to the “early man” occupation of the region, but to later Indian
groups and historic events as well.
The escarpment of the Staked Plains rising some 800 feet
(243.8 m.) above the broad valley of the Canadian River is one of
the striking topographic features of the district (pl. 1, fig. 1). The
brakes, from the rim to the plain below, are covered with juniper,
pifion, and scrub oak, and there are sporadic clusters of cottonwood
trees along the stream beds that start in the brakes and wind across
the lower plain from the base of the bluffs. On top a sparse scattering
of juniper and pifion extends back a short distance from the rim
and occurs in protected places in gullies and ravines, but for the
most part the upper level quickly merges into the vast, rolling, tree-
less terrain that so bewildered both the Spanish and early American
explorers. Prior to the recent heavy cultivation of large areas for
wheat and grain sorghums it was covered with a thick, heavy carpet
of grass that furnished ample forage for the great herds of buffalo
and antelope, as well as for the wild horses of the Canadian in historic
times, that roamed its stretches. Several varieties of cacti, including
the bushlike cholla or coyote candles, grow on the slopes of the escarp-
ment and along the ravines, and yucca is fairly abundant. Wild
grapes can be found in places along the base of the bluffs. There
are several kinds of bushes with edible berries and numerous plants
whose roots, tops, and seeds contributed in no small measure to
indian subsistence.
The winter and spring of 1941 were unusually wet in this district
with an attendant abundance of plant forms. Among those noted
were a number that are known to have been used by Indians in other
4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
parts of New Mexico and it is quite possible that they may have
served the peoples living here. Although it is not definitely known
that all of the forms were present in earlier times, they are so
widespread that it may be taken for granted that they were available.
Included in this list are: The sunflower (Helianthus annuus) that
furnished seeds for parching and for grinding into a paste used in
thickening stews and making cakes ; the four-o’clock (Mirabilis multi-
fiora) the roots of which were used in tanning; purslane (Portulaca
oleracea) that furnished greens; amole or yucca (Yucca glauca)
with seed pods for food and roots for soap; datil (Yucca baccata)
the fruit of which was eaten raw and cooked in various ways; rabbit-
brush (Chrysothamnus latisquameus) the buds of which were eaten
and the flowers of which were the base for a yellow dye; pigweed
(Amaranthus retroflexus) that supplied fresh greens or was dried for
winter use; lambsquarter (Chenopodium leptophyllum) the leaves of
which were used as greens and the seeds of which were ground into
meal; panic grass (Panicum obtusum) with seeds that were ground
into a kind of flour; wild peas (Lathyrus leucanthus) that contributed
the whole pod as well as seeds; wild potatoes (Solanum fendleri and
Solanum jamesi) the small tubers of which were boiled with the skins
on or were eaten raw with a white clay or talc to counteract certain
unpleasant effects they otherwise might have.t Then, of course, there
were the nuts from the pinons (Pinus edulis) and the prickly pears
(Opuntia spp.) that were in high favor with the Indians.
The buffalo has disappeared, but a few antelope (Antilocapra
americana americana) and white-tailed deer (Odocoileus virginianus
texanus) can still be seen. Some of the local residents report that
on occasions they have observed mule deer (Odocoileus hemionus
macrotis) in the brakes. This is a little south and east of the normal
range for that particular deer, but the animals would not need to
stray far to wander into the district. Jack rabbits (Lepus californicus
melanotis) and cottontail rabbits (Sylvilagus audubonu neome.xt-
canus) are plentiful, the prairie dog (Cynomys ludovicianus ludo-
vicianus) and pocket gopher (Geomys breviceps llanensis) are com-
mon. Coyotes (Canis latrans) were seen by members of the party,
lynx or bobcat (Lynx rufus baileyi), badgers (Taxidea taxus ber-
landiert), a buff-gray fox (probably Vulpes velox velox, but this
1 These plants were identified by Linda B. Roberts. For further information
on native plants as sources of food in the Southwest see Castetter, 1935; Castetter
and Opler, 1936.
NO. 4 SAN JON DISTRICT, NEW MEXICO—ROBERTS 5
identification is based on only fleeting glimpses of the animal), skunks
(Mephitis mesomelas varians), rock squirrels (Citellus variegatus
grammurus), striped and spotted ground squirrels (Citellus tridecem-
lineatus arenicola and Citellus spilosoma major), the hoary wood rat
(Neotoma micropus canescens), and many kinds of small mice
abound.
Birds are numerous in the locality and among those observed were:
Lark bunting (Calamospiza melanocorys), the cowbird (Molothrus
ater ater), mourning doves (Zenaidura macroura marginella), western
meadowlark (Sturnella neglecta), horned lark (Otocoris alpestris
leucolaema), western mockingbird (Mimus polyglottos leucopterus),
kingbird (Tyrannus tyrannus), cactus woodpecker (Dryobates sca-
laris cactophilus), pifion jay (Cyanocephalus cyanocephalus), cliff
swallow (Petrochelidon albifrons albifrons), western lark sparrow
(Chondestes grammacus strigatus), Arizona scaled quail (Callipepla
squamata pallida), road runner (Geococcyx californianus), western
burrowing owl (Speotyto cunicularia hypogaea), sparrow hawk
(Falco sparverius sparverius), western redtail hawk (Buteo jamai-
censis calurus), nighthawks (Chordeiles minor henryi and Chordeiles
minor howelli), Bullock’s oriole (Icterus bullocki bullocki), and the
Cassin sparrow (Peucaea cassin:). Wild turkeys formerly were abun-
dant along the Canadian near the Texas border but have not been
reported since the middle of the nineteenth century. They probably
were the eastern form (silvestris) and had worked their way up the
river and into eastern New Mexico.
There are several kinds of snakes, including the diamondback and
prairie rattlers, as well as a few lizards and the horned toad. From
the standpoint of the flora and fauna the district by and large is still
one that would hold considerable attraction for aboriginal peoples.
The general problem was attacked from several angles. One per-
tained to the locating of archeological materials and discovering
what the associations were, if any, between different types of artifacts
and animal remains. In this connection there also was the matter of
determining to what extent the various kinds of stone tools and the
bones could be correlated with the several strata in the fill. Then
there was the question of a sequence in implement types and the
possibility of obtaining important information on the relationship
between the Yuma and points from already established cultural
horizons. Another phase consisted of a careful study of the material
in the fill in the basin at the main site, of deposits in the general
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
area, and of possible correlations of both with geologic phenomena
of known age. To facilitate the recording of archeological informa-
tion and to aid in the geologic studies a detailed 5-foot (1.524 m.)
contour map of the basin and its gullies was prepared by Mr. Merrill.
Results were more gratifying in some cases than in others. This
is in part attributable to an insufficient quantity of archeological
evidence rather than to its quality, and to the fact that certain phases
require further work. On the whole, however, an interesting outline
of developments in the area is apparent from present information.
ARCHEOLOGICAL WORK
The archeological activities, under the writer’s direction, were
carried on by the students in the group. The grid system was followed
in the digging. Areas to be investigated were staked off in 5-foot
(1.524 m.) squares, the lines running east-west and north-south, and
exploratory trenches were laid out in 5-foot widths with their lengths
continuing as required on the basis of 5-foot units to provide con-
sistency in the records and in order that the sections would be uniform
in case it was found necessary to expand the trenches into larger
areas. The men were given separate sections, each working his own
from the surface to the bottom of the excavation (pl. 2, fig. 1). The
material was removed layer by layer, following the natural strata,
and the vertical and horizontal positions and the nature of the earth
surrounding all objects found were carefully recorded. On the com-
pletion of a section a scale drawing was made of the face of the next
section before work was started on it. In this way profiles showing
the nature of the deposits as they occurred at 5-foot intervals through-
out the digging were made available for study and as a part of the
data pertaining to the excavations. The archeological material pre-
sented no handling difficulties, but in the case of the animal bones it
was often necessary to harden them with preservatives and to apply
plaster jackets before removal.
Excavations in three sizeable areas and four trenches in and around
the basin (fig. 1), digging in four places on the level top of a terrace
in the brakes, midway between the rim and the plain below, and in
buried hearth levels exposed by gullies in the lower plain, produced
an interesting series of implement types with geologic evidence for
a sequence in some of the forms. This was also true for some of
the animal bones, although the number of variations involved was
not as great. Four types of projectile points came from the basin
and its environs and from the terrace site. They occurred in four
' NO. 4 SAN JON DISTRICT, NEW MEXICO—ROBERTS 7:
stratigraphic horizons and while actual superposition of specimens
was lacking, the geologic nature of the layers is such that there seems
little reason to doubt that the sequence indicated is correct. A fifth
type was found in a buried occupation level in the lower plain. Its
position in the sequence is somewhat doubtful at present writing,
yet it seems to be approximately contemporaneous with the third or
next to most recent in the series from the basin.
TRENCH I
= TRENCH 2
= TRENCH 2
Fic. 1—Map showing outline of main arroyos in the basin and the break
through the escarpment. Location of work areas, trenches, and places where
bones were found without accompanying artifacts are also indicated.
The earliest type of point, a rather roughly chipped, thick-bodied
blade with a square base, parallel sides and rounding tip (fig. 2, a),
occurred in a layer of dark earth corresponding to the top level of
the silt in one of the old ponds or lake beds. The point in general
might be considered as an example of the Indeterminate Yuma
(Wormington, 1939), but the lack of many of the characteristic
minor details makes its assignment to that classification debatable.
This doubtful status is accentuated by the fact that many obviously
unrelated forms have been lumped together under the name Indeter-
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I03
minate, and it is very much a question whether any of them should
be considered as belonging in the Yuma category. The uncertain
nature of the class was emphasized by the discussions at a conference
held at the Laboratory of Anthropology in Santa Fe, N. Mex., in
September 1941 (L. L. Ray, 1942; C. T. Hurst (The Editor), 1942),
when it was proposed that the Indeterminate group be dropped from
the Yuma series. Consensus was that as each of the forms became
established it could be named on the basis of its place of origin and
proper relationships. In view of this it probably is best, for the time
being, to refer to the present type as the San Jon point, adding the
proper qualifier when its affinities have been determined.
The San Jon point was associated with the remains of a large bison,
probably the extinct taylori.?, The bones from this level are completely
fossilized, and in many cases articulated members were so firmly
mineralized that they were lifted from the earth as a unit. Numerous
examples of articulated feet and lower leg bones were found in
upright positions in the upper part of the silt or alluvium (pl. 3, fig. 1).
They definitely suggested an animal wading out to drink, becoming
inextricably mired in ooze and perishing there. Perhaps the Indians,
certainly some of the carnivores in the area, preyed on the creatures
trapped in this fashion because in many cases bones from the upper
parts of the legs and the body were either entirely missing or else
were considerably scattered.
The geologic age of this deposit is still to be established. At
another part of the site, however, a portion of a true Folsom point
was found weathering out of the same layer in association with frag-
ments of similarly fossilized bone. This, coupled with the fact that
a large bison is involved, points toward possible contemporaneity with
the Folsom horizon, one of the oldest thus far recognized in North
America and dated in the closing stage of the Pleistocene (Antevs,
1935; Bryan and Ray, 1940). But because the Folsom specimen was
not wholly in situ and represents only a single occurrence it can be
regarded merely as an indication. On the other hand the presence
of man-made objects with extinct bison remains and the occurrence
of mammoth bones, teeth, and tusk fragments in the same stratum
a few feet away, although no direct associations have thus far been
* This material was identified by Dr. C. Lewis Gazin, assistant curator,
division of vertebrate paleontology, U. S. National Museum, who stated that
it obviously was from an extinct form closely approximating taylori in size,
but because only lower jaws and leg bones are available for study the species
cannot be indicated with certainty.
NO. 4 SAN JON DISTRICT, NEW MEXICO—ROBERTS 9
found between the latter animal and artifacts at this location, implies
some antiquity. That contemporaneity of Indians and the mammoth
in this region was within the bounds of possibility is indicated by the
finds near Clovis (Howard, 1935; Cotter, 1937), at Sandia Cave
(Hibben, 1941), and in various Texas sites (Sellards, 1940).
The second type of point, a characteristic Yuma of the form desig-
nated the Collateral (Wormington, 1939) and since the 1941 Santa Fe
a b Cc d
Fic. 2—Some projectile point types. a, San Jon; c, Collateral or Eden Valley
Yuma; b, and d, unnamed forms. (Actual size.)
conference tentatively called the Eden Valley Yuma, comes from a
mixed layer of reddish clay and sand and is found in association
with bones from an essentially modern species of bison, a much
smaller animal than that of the lower level. The bones show some
fossilization, although it is in much less degree than that of the first
group. Geologic evidence is that this horizon was separated from
the first by an appreciable period of erosion followed by marked
deposition. This argues for considerable time lapse and a much
younger age for the cultural material. The Collateral or Eden Valley
IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Yuma at San Jon has an elongated subtriangular shape with slightly
convex edges, a rudimentary tang formed by a slight inset on the
edges near the base, and a slightly convex base (fig. 2, c). There is
a suggestion of the median ridge on the faces and the inset edges near
the base are smoothed. All the Yuma points reported from the site
have not been seen by the writer as some of the earlier finds have
been scattered, but those that were studied conform to the above
description. It is possible that some of the Oblique Yuma (Worming-
ton, 1939) may be there, although the presence of one form does not
necessarily imply that the other also will be found. There are
numerous cases where only one of the types occurs at a site.
The third type of point, one with a long, leaf-shaped blade, round-
ing shoulders, and a roughly straight tang with a slight bevel to the
left, and a concave base (fig. 2, b), is found at a little higher level
in deposits essentially the same as in the case of the preceding form.
This point suggests some of Ray’s Clear Fork types (C. N. Ray,
1938), belongs in the same category as one from the Taos Plateau
reported by Bryan and Butler (1940), possibly has some relationship
to Sayles’ Brazos River forms (Sayles, 1935), and also bears some
similarity to points from the Big Bend region identified as forms
frequently found in Pecos River sites* (Kelley, Campbell, and
Lehmer, 1940, pl. 5, fig. 1, b, upper left; fig. 2, b, left); yet it is
sufficiently different to make correlation with the latter questionable.
The only animal remains associated with these points are those of
modern bison, the bones showing no traces of fossilization. The
chief distinction between the remains of this and the Yuma horizon
appears to be in the fossilization of the bones. In general the degree
or absence of mineralization has little value as a criterion because
of the varying rates at which it takes place under diverse conditions
and in different localities. When the material comes from a small
area at a single site where there is no difference in conditions, how-
ever, it probably does have some significance and can be considered
as giving at least an indication of relative age. Hence on the basis
of the nature of the bones and of the earth from which they and the
points were dug it seems evident that they are somewhat, although
not markedly, later than the Yuma.
The fourth type in the series is actually a group of several subtypes
of the small, notched and tanged arrowheads common at recent Indian
sites. Sporadic examples are found on the surface around the basin,
on the Slopes of the brakes, and on the plain below. In the excava-
8 Not fet type of point named the Pecos River point, however.
NO. 4 SAN JON DISTRICT, NEW MEXICO—ROBERTS Il
tions, however, they occur just below the present sod line or, as in
the case of those from the terrace in the brakes where there is no
grass, are from 2 inches (5.1 cm.) to 4 inches (10.2 cm.) beneath
the surface in the upper part of an ash- and charcoal-stained deposit
underlying a layer of sterile sand. Curiously enough none of these
seems to belong to any of the types identified as characteristic of the
Apache, Wichita, Jumano, or Panhandle groups (Sayles, 1935) that
might be expected to have left a few points in the region.
The triangular form with broad, expanding tang with convex base
(pl. 4, a) appears sporadically over most of the southern plains, is
occasionally seen in collections from the northern plains, occurs in
some eastern Pueblo ruins, and may be found in sites east of the
Mississippi River. It apparently does not occur in large numbers
in any one locality and has not been correlated with any particular
horizon or specific cultural group, but it undoubtedly is a relatively
late type.
The same is true for the group with a longer, expanding tang and
more rounding base (pl. 4, c). A comparable point is reported from
a rock shelter in the southern escarpment of the Staked Plains in
Winkler County, Tex., some 220 miles (354.1 km.) southeast from
the San Jon site (Holden, 1938, pl. 28, No. 10). There the asso-
ciation included potsherds, some of Pueblo origin, other types of
projectile points, knives, scrapers, drills, bone and shell artifacts.
The culture represented is considered to be non-Puebloan, many of
the points and artifacts seem to be Sayles’ Wichita and Jumano types
although other groups are also included, and evidently does not belong
in an early category.
The triangular point with slightly convex and serrated edges,
medium tang and concave base (pl. 4, b) suggests some of the Texas
forms from the Edwards Plateau section, might possibly be derived
from some of the bifurcated-base types found there, and also has
a resemblance to examples occurring sporadically in the eastern
periphery of the Pueblo area, yet it does not appear to have definite
affinities with established cultural phases. Perhaps, like the others,
it represents a minor type that had a rather widespread distribution
and as far as present knowledge goes is without any particular
significance.
The last in the group (pl. 4, d) is a type in which the basal portion
constitutes a much larger proportion of the blade than is usually the
case. The tip end is triangular in shape, while the section below the
side notches is roughly rectangular with a deeply concave base.
[2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Slightly variant examples of the type have been reported from
numerous sites, mainly of the Classic and later periods, in the Pueblo
area where they generally are considered as “unusual” forms (Cos-
grove, 1932, pl. 51, a; Kidder, 1932, fig. 6). They probably bear
some relationship to the more highly developed points found in Okla-
homa, some of the southern States, and even in parts of Mexico,
that are characterized by such deep basal concavities that they appear
to have wings. Beyond their late horizon, however, they do not seem
to have any special local significance and certainly cannot be regarded
as a criterion for the presence of any specific group in the area. All
four of these types of points are in association with bones from
modern bison, deer or antelope, and jack rabbits.
The fifth type, found in a buried occupation level in the lower plain
(pl. 3, fig. 2), is a rather stubby form with roughly parallel sides,
flat base, and sharply tapering tip (fig. 2, d). First thought was that
the implement was a drill made from a broken point, but careful
reconsideration and study of several examples indicates that it prob-
ably was a form of projectile. In a general way it bears resemblance
to some of the points from the Edwards Plateau district in Texas
and also some of those from the Big Bend region, yet it is sufficiently
distinct to be considered as representing a separate class. Possible
relationships may be determined by future work in the region. Asso-
ciations in this case consisted of modern bison, deer, and bones from
small mammals. The type, as previously mentioned, appears to be
contemporaneous with the third main group, yet eventually may
prove to be somewhat older.
Projectile points were the only artifacts obtained from the oldest
and subsequent Yuma horizons, unless a few nondescript flakes
capable of utilization as knives or scrapers be considered as tools,
and there is no information regarding other kinds of implements
comprising the complexes to which they belonged. Further data
along this line are essential to a proper understanding of the subject
as a whole. Several kinds of scrapers, knives, choppers, hand ham-
mers, and grinding stones were associated with the later type points.
In the case of the fourth series there was the addition of potsherds.
The ubiquitous snub-nosed or end scraper (pl. 4, e-k) is one of
the more prevalent forms in the group of scrapers. As is usually the
case, most of the examples are of the uniface, planoconvex type with
the flat side showing little or no chipping and the curved side having
had just enough flakes removed to give the implement its characteristic
shape. The convex, broad end and two edges were retouched by the
NO. 4 SAN JON DISTRICT, NEW MEXICO—ROBERTS 5
pressure method, the fine flaking producing sharp scraping surfaces.
The smaller end, which on most examples still shows the bulb of
percussion produced when the flake was struck from the original
nodule, rarely has any flaking. An exceptional specimen is shown
in plate 4, k, where the smaller end was given a concave edge of the
form that was used for scraping and smoothing arrow shafts. No
examples were found with a small, sharp point for perforating or
egraving purposes. In some localities these are a fairly common feature
on snub-nosed scrapers and may occur at various places on the
implement. In some cases they are at one of the corners formed by
the convex end and one edge, occasionally they may be noted at both
corners, or they may be along one of the edges or at the small end.
Whether the absence of the feature from the San Jon examples is
purely coincidence in that specimens bearing it just weren’t found
or is a matter of significance is not known at present. Taken as a
group or singly, there is nothing to distinguish the scrapers in this
series from similar forms found elsewhere in either Plains or Pueblo
sites and they cannot be considered as a criterion for cultural or
period identification. They apparently were concomitant with hunting
among most Indian groups.
Side scrapers of simple form were common (pls. 5, 6). They
were made from flakes of various sizes and shapes and in most cases
exhibit a minimum of flaking, except along the functioning edge
which was given a low-angled bevel. A few specimens have a fine,
secondary flaking or retouch along the working edge, but for the
most part the regular flaking seems to have been sufficient and there
are examples where it occurs on only a portion of the edge. As a
matter of fact numerous sharp flakes undoubtedly were used as
scrapers without any attempt being made to better the edges through
the flaking process. This is particularly true of various quartzite
forms occurring in abundance at the places where the other types
were found. Such flakes break away from a core or nucleus with
a fine, straight edge that will stand considerable abuse before being
dulled to the extent that it is no longer efficient. While such imple-
jents are in use, small flakes are generally knocked off the edge.
This is a good criterion for identifying utilized flakes. Numerous
examples in the collection show this feature. A large majority of
the working edges are convex, only a few are straight, concave forms
are rare, and there is no example of a double-edged tool in this series.
4Flakes found in association with the San Jon and Yuma points belong in
this category.
I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Closely associated with the foregoing implements is a similar group
that is arbitrarily considered to consist of rough flake knives. The
main distinction is that of thickness and the nature of the functioning
edge. In the case of the knives the flakes are much thinner and the
edge has an acute, longer-tapering bevel. Both scrapers and knives
of this type are commonly regarded as fortuitous flakes and little
attention is paid to them. They seem to have been an important part
of the implement complex in this district, however. The closest
similarities between them and artifacts from other localities are found
in a series from caves along the Cimarron Valley in northeastern New
Mexico and western Oklahoma (Renaud, 1930). The latter were part
of a complex that has been considered as a primitive form of Basket
Maker culture and although, because of peripheral lag, it may be later
chronologically than the material from the main Basket Maker centers
farther west, it is, if correctly identified, older than the San Jon speci-
mens. In view of this situation, perhaps the main significance in the
similarity of forms is their indication that in this general region the
hunting peoples over a long period of time tended to rely on implements
produced with a minimum of chipping.
Better knives of the more conventional type consist of bifaced
blades of a general broad-leaf shape with rather blunt points and
broad, convex base (pl. 6, 4, 71). The workmanship on these imple-
ments is not so good as that on the projectile points, but they are
definitely made tools. There is nothing, however, to distinguish them
from similar artifacts scattered abundantly and widely over most
of the Plains area. Fragments from a diamond-shaped, double-ended
or four-edged, beveled type of knife that is common in districts
farther east in Texas and Oklahoma (Poteet, 1938), and extending
northward along the western edge of the Plains, were found, but
no complete specimen was obtained. This type of knife has been
attributed to the Wichita (Sayles, 1935), yet it has been found under
conditions suggesting that it may derive from other, and in some
cases possibly older, groups. Whether the form actually was made
in the San Jon district or was an importation is an unsettled problem.
On the basis of its small representation and the lack of complete
specimens it would seem to belong in the trade category. Sayles,
however, notes that it was a common Panhandle artifact and the
present fragments appear to be of Poteet’s type B which is believed
to center in the Panhandle. Hence it may be at home in the San Jon
collection. The kinds of stone used in this instance are not much
help as they occur locally as well as farther east. The implement
NO. 4 SAN JON DISTRICT, NEW MEXICO—ROBERTS 15
unquestionably belongs to the general bison-hunting complex of the
Plains and where found in Pueblo sites of the eastern periphery
is an introduced form and good evidence for trade relations. (Kidder,
1932, Pp. 30-34).
Choppers consist of large pebbles or fragments of stone with broad
flaking along one side or an end (pl. 7). In rare cases there was a
little secondary chipping but they mainly were fashioned, like many
of the scrapers, with a minimum of effort. Frequently the removal
of one or two flakes sufficed to form a satisfactory working edge.
The remaining smooth surfaces would serve to protect the hand of
the user. From their occurrence in the excavations it was evident
that these objects had been employed in cracking and splitting bones.
This type of tool is commonly found at sites attributable to hunting
peoples or those who depended on game for a large part of their
sustenance.
Hand hammers are simple round or ovoid-shaped pebbles that
generally exhibit a slight amount of pecking or pitting, probably the
fortuitous result of use, along one side or on each end (pl. 7). They
are of a size that can be gripped in the hand, 3 inches (7.6 cm.) to
5 inches (12.7 cm.) in diameter, and no doubt served for knocking
flakes from nodules in the manufacture of implements and for general
pounding purposes under conditions where such a tool was needed.
Some are suggestive of what has been termed the Plains Maul
(Sayles, 1935), although in no case is there any indication of polish
around the central portion of the body nor the slightest trace of an
encircling groove, features common to that implement. The Plains
Maul usually is associated with the complex of agricultural and
seed-gathering peoples. One of its purposes may have been that
of pecking out depressions in stones intended for grinding and in
shaping the smaller stones used with them, in which case it would
be most efficient as a hand tool. That it had other functions, however,
is indicated by the polished central portion or shallow groove evincing
hafting in some sort of handle. There was not much indication that
the inhabitants had practiced agriculture in the vicinity of the San Jon
site, but that they took advantage of the abundance of natural products
is suggested by the presence of hand stones or manos and milling
stones or metates. Both of which may have been shaped by artisans
employing hand hammers or mauls of the type found here.
The hand stone or manos for grinding are rather nondescript and
all are of the single hand size (fig. 3). Some are oval in outline
with one flat and one convex grinding face. Others are roughly
16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
circular with one flat and one convex face. Still others are circular
with two flat faces and a wedge-shaped cross section. There are
examples that are little more than natural pebbles with a single
flat or only slightly convex face. In some cases the edge between
the two grinding surfaces was pecked, in others it was not. There
Fic. 3—Manos or hand grinding stones. (One-third actual size.)
apparently is no correlation between outline, grinding surface forms,
and the pecking of the edges. The latter may or may not be present
on any of the two-faced forms. No one type of mano seems to have
prevailed over the others or to be characteristically associated with
one type of point. In general they are similar to those found at most
sites throughout the region occupied by users of native food plants.
In contrast to many locations, none of the San Jon examples has a
os
oe ee
eit i a a
NO. 4 SAN JON DISTRICT, NEW MEXICO—ROBERTS Wi;
pit or depression in the face. The latter implements possibly were
a combination type that could be used either as a grinding stone or
as a hammer, the pits or depressions making it easy to hold the stone
between the thumb and fingers and strike with the edge. Hard nuts
and seeds could be cracked in this fashion and then be ground by
rubbing with the flat surfaces without the necessity of changing tools
in the process.
The metates or nether milling stones are of two types. One is a
thin slab of stone with a shallow, oval or circular bowl and the other
is a deep, narrow, troughlike stone that actually might be considered
as boat-shaped. Little effort seems to have been made to give form
or shape to the thin examples. A roughly rectangular or polygonal
slab was obtained, and a shallow basin was pecked in one face.
Through use the basin would be enlarged and deepened. The trough
or boat-shaped type, fashioned from large tabular blocks, was given
some external dressing, the rounding of sharp edges and the removal
of projecting points, but most of the work was on the interior and
pertained to the pecking of the groove or trough. The latter extended
a greater part of the length and width of the block. The deepest
portion was along the center with the bottom curving up toward the
ends, the sides being fairly vertical. The ends of the trough were
closed, not pecked open as in the case of the Pueblo milling stones.
Both of these metate forms are reported for the Jumano and Pan-
handle groups, while the first is also attributed to the Wichita
(Sayles, 1935) and extends northward into other portions of the
Plains area. The difference in metates no doubt accounts for the
difference in manos. The circular forms would be used for the most
part with the shallow bowl type, the grinding being done with a
circular motion of the hand and arm, while the longer oval examples
would serve in the trough form, the grinding being accomplished with
a backward and forward movement of the hand stone.
Materials used in the manufacture of implements consisted of
chalcedony, chert, jasper, quartzite, dolomite, sandstone, and granite.
The projectile points of the San Jon and Yuma types were made
from gray and mottled chert. The points in the third group are all
of quartzite, gray, brown, red, or yellowish white. There is greater
variety in the fourth group, chalcedony, jasper, chert, and quartzite.
Fifth type points were made from gray or white quartzite. The
fragment of Folsom point is of dolomite. The snub-nosed scrapers
are of jasper, chert, dolomite, and quartzite. Other types of scrapers
are mainly quartzite, with a few of jasper. The flake knives are
18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
mostly quartzite, although there are sporadic examples of jasper
and chert. Jasper and dolomite were used in the four-edged knives.
Choppers were made of quartzite and from granite pebbles. Hand
hammers are quartzite, granite, and sandstone. Manos and metates
are sandstone. Most of these materials could be obtained locally and
there are several places on the slopes of the brakes where nodules
have weathered out from the formation. At each of these locations
there is definite evidence of workshop activities in the form of
numerous chips and flakes, fragments from partially completed
implements and hammer stones used in the breaking of nodules.
Identical scraps and flakes were found in the digging around the
basin and on the terrace site, showing that some of the material was
carried back to camp and worked on there. Some of the chert prob-
ably came from deposits farther east and was mostly imported in
the form of completed implements as there is little of that kind of
stone in the chipper’s debris around the basin. The dolomite is the
silicified form generally known as Amarillo dolomite which occurs
along the Canadian River and some of its tributaries and which was
obtained by the Indians from extensive quarries near Amarillo, Tex.
Some of this material undoubtedly was taken to San Jon, possibly
in the form of blanks to be fashioned into completed tools at the
maker’s leisure, as small bits and flakes of it are found in the chips
from the occupation areas. Sandstone and granite pebbles are avail-
able at numerous places in the vicinity of the basin and other sites.
One kind of stone used in snub-nosed and other types of scrapers
seems to be a silicified sandstone that has actually become a quartzite.
It has a characteristically laminated appearance suggestive of shale
(pl. 5, center right), but in contrast to that material is very hard and
compact. It is of particular interest because flakes, nodules, and
implements made from it are abundant at a site lying between Big
and Little Tucumcari Mountains a few miles south of the town of
Tucumcari. Present evidence is that it was not extensively used
elsewhere in the region which suggests a possible relationship between
the occupants of the basin area at San Jon and the site near Tucum-
cari. This is also indicated by other factors and if contemporaneity
should be established it would be of significance in the matter of
certain geologic correlations still to be demonstrated. The source of
this material has not been definitely located as yet, but it unques-
tionably is in the immediate district.
The potsherds found in association with the artifacts and fourth
type points occur only in small fragments and are not numerous.
NO. 4 SAN JON DISTRICT, NEW MEXICO—ROBERTS IQ
While they indicate that the people had a knowledge of and possessed
some pottery, they also suggest that it was not a highly important
trait in the complex as far as activities around the basin were con-
cerned. Most of the sherds are from a buff or brownish-colored
ware with smooth surface and hard, compact paste. The tempering
material, as studied with a hand glass, appears to be sand. In a few
cases there is a slight admixture of mica. None of the fragments
show any form of decoration. Both bowls and jars seem to be
represented. As a group the sherds fit the description for Sayles’
Panhandle wares, although no paddle-marked examples were found.
The Panhandle wares are associated with house ruins concentrated
along the Canadian River (Sayles, 1935, p. 84) and on the basis of
associated sherds from the Rio Grande have been dated as about
middle fourteenth century. The affinities of the wares have not been
definitely established. They probably belong to the broad basic form
represented in the pottery of the Wichita and the Jumano which in
turn points toward the Caddo. It is possible that the presence of
such sherds near the basin at San Jon is to be attributed to hunting
parties from the Canadian settlements to the northeast and that the
small number is due to the fact that people on the move rarely bur-
dened themselves with many pottery vessels.
There are a few black-on-white sherds from vessels that probably
belonged to a type of pottery from an adjacent area to the west.
This form has been called Chupadero Black on White (Mera, 1931)
and is one that is commonly found in association with brown wares
such as those represented by the sherds in the present collection.
Chupadero Black on White is a Pueblo ware that was taken over
bodily and adopted by the makers of the brown wares (Scholes and
Mera, 1940, p. 293). Traces of it have been found as far east as
the sand-dune camp sites in western Texas (Mera, 1935, p. 30).
It was a late development out of an earlier type of Pueblo ware and
seems to have been absorbed into the ceramic complex of the brown-
ware region during the thirteenth century. It apparently persisted
into early historic times and its presence in an assemblage of ma-
terials dating from the late fourteenth or early fifteenth century in
the San Jon district is wholly in keeping with the general picture.
Although both bowls and jars were made in this ware, jar fragments
only were found during the San Jon investigations.
The occurrence of actual hearths was noted only in the horizons
represented by the fourth and fifth point types. However, scattered
pieces of charcoal and sporadic ashes were found in all of the archeo-
20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
logical levels. In no case was the hearth an actual fire pit lined with
stones or stone slabs. They were simple basins in former surfaces
of occupation, were roughly circular or oval in outline, and were
comparatively shallow. They ranged from diameters of 1 foot 4
inches (40.6 cm.) and 1 foot 6 inches (45.7 cm.) to 2 feet (61.0 cm.)
and 3 feet (91.4 cm.), and from depths of 5 inches (12.7 cm.) to
8 inches (20.3 cm.). All of those examined contained ashes, char-
coal, pieces of fire-burned rock, and sporadic chips and spalls from
implement making. In a few cases charred fragments and splinters
of bone, even complete small bones, were found mixed with the other
debris. No traces of a habitation, tent circle, or shelter occurred
in the vicinity of any of these hearths. As a matter of fact no such
indications were found in any of the excavations. The probabilities
are that any dwellings used by these people were of a temporary
nature, tents made from buffalo skins being the most likely form,
and as a consequence little evidence remains of their former placement.
GEOLOGICAL STUDIES 5
Considerable progress was made in the study of the deposits and
their possible age. This phase of the investigations was carried on
by Dr. Bryan and Mr. Judson. Two lines of approach were followed.
The first pertained to the belt of plains between the Staked Plains
escarpment and the Canadian River, and the second to the archeo-
logical site at the basin. In the plain three stages of alluviation fol-
lowed by periods of erosion, arroyo cutting, and the formation of
sand dunes were established. This sequence has many similarities
to deposits in the Hopi country in Arizona (Hack, 1942) and the
Big Bend region in Texas (Albritton and Bryan, 1939) but because
of insufficient field work is subject to revision.
Interesting items appear in the sequence, even though the results
are regarded as still tentative. The last cycle of erosion, continuing
at the present time, is characterized by arroyos and sand dunes. The
beginning and heavy cutting of the arroyos representing this stage
appears to belong to the decade from about Ig00 to 1910. The forma-
tion of the first dunes in the present series started about 1910. The
5 The present summary is mainly for the purpose of rounding out the account
of the expedition’s activities and to indicate the archeological concomitants of
the results. It is based on notes furnished by Bryan and Judson and on state-
ments made during discussions of the problems in the field camp. The authori-
tative and detailed geologic report will be issued by Bryan and Judson when
their studies are completed.
eS ee, eee Te
ent ae i en Ol nn ee
NO. 4 SAN JON DISTRICT, NEW MEXICO—ROBERTS 21
third alluviation is indicated by loose and friable alluvium. Fire pits,
animal bones, and stone artifacts, late in type, occur in this layer.
The preceding stage of arroyo cutting and wind-blown sand, the
second erosion interval, is correlated with Pueblo type potsherds that
date in the period 1300 to 1540. The second alluviation is charac-
terized by an alluvium, humic in places, with irregular, hollow, twig-
like limestone concretions (calcareous tubules) in the finer portions.
Fire pits, artifacts, and bones from modern bison are found in this
level. The first erosion period was marked by arroyos and sand-
dune formation, but thus far is not correlated with any evidence of
kuman activity or animal remains. The deposit of the first alluviation
contains large limestone concretions and compact humic alluvium
in small bodies. Mammoth, sloth, and horse bones have been found
in this horizon. Projectile points purportedly of Folsom and Yuma
types are supposed to have come from this layer, but the authenticity
of this occurrence and the kind of Yuma involved remains to be
established. If substantiated, however, and the Yuma proved to be
of the Indeterminate type the combination of features would be an
important link in correlating the plain deposits and those in the
basin on top.
The deposits in the basin form a complex but definite sequence,
one that is in part reproduced in the next gulch to the east. There
are II stages in all and, as previously suggested, the sequence is
related to the filling of an original valley in the escarpment with
layers of sand, the “basal sand” of the section. The last or present
stage is characterized by the deep ravines and the broad arroyos in
the basin and the continuing erosion of today. Preceding this was
an interval of alluviation characterized by coarse alluvium that re-
mains as terraces. The small barbed arrowheads, potsherds, and
grinding stones probably belong to this horizon. The disconformity
between this layer and the next lower alluvium is marked by deep,
narrow arroyos. The latter were cut into alluvial fill consisting of
reddish clay and sand. It is in this horizon that the Yuma (Collateral
or Eden Valley) and third type points belong. Broad, shallow
arroyos characterize the disconformity between this layer and the
underlying alluvium composed of a sequence of clay, red sandier
alluvium, and clay. The San Jon point and extinct bison and mam-
moth come from this deposit, the lowest level for either animal or
human evidence. Below this is a layer of water-laid ash, volcanic
in origin, resting on an iron cap and associated clays resulting from
the deposition of limonite and clays under wet conditions. The iron
22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
cap and clays were laid down in the hollows in the top of the white
and buff cross-bedded sand filling the basinlike valley that had been
formed in the top of the Tertiary beds underlying the area.
The question of the relationship and the correlation between the
two sequences is an important one. Although there are certain indi-
cations that are suggestively significant, more work is needed to
establish them as facts. On the basis of the similar circumstance
of the alluvium of the first alluviation in the plain sequence being
subsequent to a deposition of volcanic ash and the lowest alluvium
of the basin sequence resting on such material, that both contain
similar faunal remains, and that Folsom and possible Yuma points
are reported from them, it would seem that they are approximately
the same age. Because of the uncertain archeological evidence for
the presence of Folsom and Yuma material in the first alluviation
and of Folsom at the basin site such a tie can be regarded only as
highly tentative at best. There is an additional complication in the
nature of the Yuma points involved. If those from the first alluvia-
tion are the Indeterminate, even perhaps the San Jon type, the case
is much stronger than it would be if they are the Collateral or Eden
Valley form, as the latter definitely occurs in a much later horizon
at the basin. No doubt much of the difficulty will disappear when
there is more complete geologic and archeologic information from
which to draw conclusions. At present it is not possible to estimate
how long a period of time is represented by the first arroyo-cutting
cycle and the second period of alluviation; hence it cannot be indi-
cated how many years before the 1300-1540 second arroyo stage
the first alluviation ended.
While studying the deposits in the valley floor some miles west
of the main site Mr. Judson found an interesting cache of 44 stone
implements. The artifacts, consisting of five large blades (pl. 8),
end scrapers, side scrapers, and flake knives (pl. 9), had been placed
in a hole beside a fire pit. The blades were lying side by side in a
line extending in a general east-west direction. The other specimens
were grouped around and above the blades in no particular order.
The fire and cache pits were exposed by the caving of a gully bank.
Their tops were 2 feet (61.0 cm.) below the present surface. The
overlying deposits consisted of a layer of structureless sand con-
taining numerous charcoal flecks, a layer of red-clayey sand, and the
layer of modern buff-white blow sand. The pits were dug into a
layer of water=laid buff-colored sand to a maximum depth of 8 inches
(20.3 cm.). This layer corresponds to the third alluviation period
NO. 4 SAN JON DISTRICT, NEW MEXICO—ROBERTS 23
in the sequence for the valley plain. On the basis of the evidence
from Pueblo type potsherds, previously mentioned, Judson concluded
that the earliest possible date for the third alluviation was in the
late 1400’s. Hence at most the specimens could only just antedate
the early historic period, and it is more likely that they actually
belong in that stage.
Neither in the type of the implements nor in the material from
which they were made, a gray chert, is there any suggestion of rela-
tionship to the artifacts excavated in and around the basin at the
main site. The absence of projectile points adds to the difficulty
of attempting to correlate the material with any definite group or
cultural horizon. The writer has seen blades and scrapers of this
type, made from similar stone, that came from Lipscomb County,
Tex., and adjacent Ellis County, Okla. Similar blades and scrapers
are also found in southern Kansas, north of this Texas-Oklahoma
district in what has been called the Province of Quivira (Brower,
1898, 1899), in fact large blades of this type have been called the
Quivira Blade. Throughout this general area are large deposits of
gray chert where much material was quarried by the Indians and
the present specimens may have come from there. That is old Wichita
country and on the basis of a fifteenth-century date it is possible
that the implements may indicate the presence of hunting parties
from that tribe in this New Mexican area. In his listing of typical
Wichita artifacts, however, Sayles (1935) does not include such
blades and scrapers; hence they may have belonged to some other
group. Such caches are not uncommon in areas farther east and
north but, as far as evidence in the literature is concerned, seem to
be rare in this district. A similar series of blades is reported to have
been found about to miles (16.1 km.) south of the present location.
They were not seen by any of the field party and the occurrence
has not been verified. That these blades were actual implements and
not blanks intended for further refinement into more specific tools
is demonstrated by the secondary chipping along some of the edges
and a certain amount of polish, in some areas on their surfaces,
such as is acquired through use. Implements of this type were
employed in the preparation and dressing of skins, particularly those
from the buffalo. The snub-nosed and side scrapers in the series
are of the types usually found in a hunting complex, although as a
group the sizes are larger than similar tools from the basin and buried
sites in the plain.
24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103 }
The fire pit, measuring 2 feet (61.0 cm.) by 3 feet (91.4 cm.),
had no lining. It contained charcoal, ashes, splinters and fragments
of charred bone, several vertebrae, one of which showed the effects
of fire, and a calcaneum. The vertebrae and calcaneum are from
the American antelope or pronghorn (Antilocapra).6 There was
an additional fire pit nearby, and a few bones from modern bison *
were found in the vicinity, but no other cultural material was
recovered.
MOLLUSKS
Numbers of mollusk shells were found in the digging, and in order
that there would be sufficient material for comparative purposes,
specimens were collected from typical wet-weather lakes in the area
surrounding the site. Others were gathered from shallow depressions
where sand and surface soil had been blown out by wind action.
These shells were sent to Dr. Frank C. Baker, University of Illinois,
who reported that the collection contained a number of land and
fresh-water species. Commenting on the collections Dr. . Baker
writes: ®
The species of Helisoma from the pits and fossil deposits are the same as those
living in the Recent fauna; none are extinct, as is the case among some of the
vertebrates. The other species of land and fresh-water mollusks are also the
same as typical members.of the Recent fauna. This difference in geological
faunal contents has been observed in many other places and is to be noted
especially in some Pleistocene deposits where the mollusks show little change
in form while the vertebrates, principally the mammals, contain several species
now extinct.
It is interesting to note, although there may be no particular sig-
nificance in the fact, that all the shells from the excavations around
the basin are from a single, fresh-water species (Helisoma tenue
sinuosum Bonnet) common in New Mexico, Arizona, Texas, and
northern Mexico. Dr. Baker states that the form is “Usually named
tenuis but it is a distinct race of the typical tenwis which is confined
to Mexico, the types being described from the vicinity of the City
of Mexico.” The fact that the material from the basin represents
a fresh-water species is additional corroboration for its having been
a lake or series of ponds.
® Identified by Dr. David H. Johnson, assistant curator, division of mammals,
U. S. National Museum.
7 Bones identified by Dr. C. L. Gazin, assistant curator, division of vertebrate
paleontology, U. S. National Museum.
8 Letter to Dr. Paul Bartsch, curator, division of mollusks, U. S. National
Museum.
;
;
NO. 4 SAN JON DISTRICT, NEW MEXICO—ROBERTS 25
Samples from the deposits representing the bottoms of the various
filled-in ponds were examined for fossil diatoms by Paul S. Conger,
custodian, section of diatoms, United States National Museum.
None were found, however, and no help toward determining the
age of the deposits was obtained from this possible source of
information.
SUMMARY AND DISCUSSION
From excavations on the northern rim of the Staked Plains, the
brakes below, and the plain of the Canadian River valley south of
the town of San Jon, N. Mex., came an interesting sequence of
projectile point and other artifact types that sheds some light on
the aboriginal occupation of that portion of the Southwest. Faunal
associations and geologic horizons give good indication of relative
age, but more evidence is needed before an actual chronology can
be suggested. The investigations to date tend to corroborate impli-
cations observed in finds made elsewhere, particularly with respect
to the relationship between certain kinds of projectile points, yet
do not furnish the complete and detailed evidence essential to a
thorough understanding of developments in the earlier stages of the
lithic industries in the western Plains. Later types of stone tools
and implements probably were made by Indian groups known to have
hunted over that area. On the basis of present knowledge, however,
it is not possible to assign specific forms to definite peoples. There
unquestionably was a mixing of materials in late times when various
bands swept back and forth across the region, following the great
herds of bison, camping at the same water holes, and otherwise
making use of the same territory. Until sites attributable to occu-
pancy by single groups have been worked and the character of the
artifacts made and used by them is established to the extent that
they are readily recognized the identification of different objects in
a mixed series as the product of a definite tribe is not feasible. The
best that can be done is to suggest certain probabilities.
The oldest horizon in the district is represented by one type of
projectile point occurring in association with the bones of an extinct
species of bison and coming from a stratum in which mammoth
bones and teeth also are present. The point is of a type that might
be called Indeterminate Yuma, but in order to clarify a confusing
situation with respect to that classification it is designated the San Jon
point. Indications are that it may be contemporaneous with the
Folsom horizon, one of the oldest thus far established, but more
26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
satisfactory evidence is necessary to establish that fact. In some
ways this type of point is quite like others found under somewhat
similar circumstances, points also identified as Indeterminate Yuma,
that apparently have a late contemporaneity with Folsom types and
that persisted after the latter were no longer made (Roberts, 1940,
pp. 64-65). Hence it is possible that this early San Jon level dates
from the closing stages of the Folsom horizon, some 10,000 to 15,000
years ago. Geologic studies tend to substantiate such a conclusion,
although it must be regarded as tentative until more work has been
done on that particular phase of the problem.
The next oldest remains comprise an assemblage of modern bison
and deer bones and points of the Collateral or Eden Valley Yuma
type. The significant factor in connection with this horizon is that
geologic evidence demonstrates that between it and the preceding
occupation there was a lengthy period of erosion followed by another
during which there was considerable deposition of new material.
The time lapse indicated is a relatively long one and during the
interval the large bison were replaced by a smaller species, the modern
buffalo. The cultural material from this level is unquestionably much
younger than that from the underlying one, a fact that has important
implications with respect to the status of different types of projectile
points in the Yuma series. There has been a general tendency to
regard all forms of the Yuma as more or less contemporaneous and
indicative of the same cultural age. If the San Jon point from the
oldest level is considered as an example of the Indeterminate Yuma,
a questionable classification discussed in the detailed consideration
in preceding pages, it is obvious that at least two of the Yuma forms
occurred in sequence and are characteristic of chronologically dif-
ferent stages. The present evidence tends to support a previous
suggestion to the effect that some of the generalized points regarded
as Indeterminate Yuma may represent the form out of which the
true Yuma developed. The definite break in occupation between the
two levels here, however, shows that the transition did not take place
in the San Jon district but if it actually did occur was accomplished
elsewhere.
The break in occupancy evinced by the nature of the deposits is
of interest because it is in accord with indications noted in other
areas. For some as yet unexplained reason at most sites where traces
of the earliest hunting cultures now recognized are found there is a
definite hiatus between them and the following stages. Whether there
actually was an interval during which the regions where these mani-
NO. 4 SAN JON DISTRICT, NEW MEXICO—ROBERTS 27
festations occur were uninhabited or whether it was only the particular
sites that were not lived on, the people camping elsewhere and leaving
materials not yet properly identified, is a problem still to be solved.
The consensus of many is that there was an actual interruption in
the stream of migration flowing from the Old to the New World
and that for a time vast stretches of the western Plains and the
Great Basin were uninhabited; that the first peoples had pushed on
southward and into Middle America, although traces of them have
not yet been found there, and those who were following had not yet
arrived. Others maintain that once man had reached the North
American continent there was no break in the continuity of occupa-
tion, that the evidence just has not been found or else has not been
recognized. There are places where an unbroken sequence from early
to modern times is suggested (Sayles and Antevs, 1941), but there
still is a question as to whether or not there is a gap between the
beginning of that series and some of the other older forms. Perhaps
the correct conception is that in some districts remnants of the early
migration persisted and ultimately were joined by incoming groups
of a later movement, while in other sections the dispersal of peoples
into the more southerly regions left large areas unoccupied sufficiently
long for natural agents to cover their former camping places before
others drifted in and settled at the same locations. Such certainly
seems to have been the case at San Jon.
From the appearance of the Yuma type points down to late proto-
historic times there was no break in the occupation of the area investi-
gated. Projectile points progress through forms similar to some of
those found in the Texas area to the east to the small, notched types
associated with late sites in many parts of the country. Accompanying
these are stone implements of the kinds that normally occur in the
complex of hunting or hunting—seed-gathering peoples. In addition
the material from the latest level contained potsherds. Among the
stone objects are forms similar to those that have been attributed
to the Wichita and the Jumano Indians, others that are like artifacts
from sites in the Panhandle district along the Canadian, and still
others that can be duplicated in material coming from eastern Pueblo
ruins. The potsherds are of two types, one a buff or brownish-colored
ware similar to that occurring in house ruins along the Canadian
River that probably belongs to the broad, basic type represented in
the wares of the Jumano and the Wichita, that have some relationship
to those of the Caddo, and the other a black-on-white ware that was
derived from a Pueblo type. The latter, called Chupadero Black
28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
on White, is commonly found with brown wares such as those in the
San Jon collection, and is a form that was taken over bodily, becoming
a part of the ceramic complex of the brown-ware-making peoples.
Considered as a whole, the series of implements from the late horizon
definitely indicates that several different Indian groups used that area
as hunting territory, a fact that is borne out by historical documents
of later times.
On the basis of geologic studies and the presence of certain types
of Pueblo potsherds in deposits at other locations in the district,
supplemented by the evidence of the Chupadero Black on White at
the basin site, the period of the last horizon is judged to be in the late
fourteenth or early fifteenth century. Even with full allowance for
an appreciable interval for the intervening point types, the period
of the Collateral or Eden Valley Yuma is much later than has
generally been supposed. If the San Jon point horizon is approxi-
mately of the age suggested, the gap between the two is indeed a
broad one. Subsequent work on the geology may show that the
deposits containing the San Jon point and extinct species of animals
is more recent than indicated at present, but the break in the sequence
would still be of sufficient proportions to cast doubt on the idea of
a continuity of peoples in the area.
In general it may be said that the remains in the San Jon district
are those of a hunting and hunting—seed-gathering peoples whose
closest affinities were with the Plains cultures. Such traits as are
suggestive of the Pueblo pattern to the west were either borrowed’
or are present because of trade relations. Pueblo peoples on occasion
did get that far east, as is shown by sporadic finds along the Canadian
and in later times by historical records, but apparently did not linger
long enough to have any marked effect on the archeological picture.
The earliest occupants of the region depended on the large bison for
subsistence, while later groups hunted buffalo, deer, and antelope,
and gathered the native food plants.
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Bull. Geol. Soc. Amer., vol. 51, pp. 373-432, March.
Wormincton, H. M.
1939. Ancient man in North America. Colorado Mus. Nat. Hist., Pop.
Ser., No. 4.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOE. 103, NOo 4, PE
%
1. VIEW OF THE NORTH RIM OF THE STAKED PLAINS
Photograph by Edison P. Lohr.
2. MAIN ARROYO
Line of old lake bottom can be seen along top of light-colored earth.
Expedition camp at left.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLES 10S iNOS a pees
1. STARTING EXCAVATION IN ONE OF THE ARCHEOLOGICAL AREAS
2. MASS OF BISON BONES AT BOTTOM OF ONE OF SECTIONS
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 4, PL. 3
1. BISON FOOT AND LOWER LEG BONES IN SILT LAYER AT BOTTOM OF
FORMER POND
2. OCCUPATION LEVEL EXPOSED IN ARROYO BANK IN LOWER PLAIN
Workman pointing to hearth.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 4, PL. 4
k / mM
ARROWHEADS, SNUB-NOSED SCRAPERS, AND POINTS FROM KNIFE BLADES
(Actual size.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS WOES WOS5 IN@a ain eo &
SCRAPERS AND ROUGH FLAKE KNIVES
(Actual size.)
SMITHSONIAN MISCELLANEOUS
COLLECTIONS
SCRAPERS AND KNIVES
(Actual size.)
VOL. 103,
NO; 4, (PEs6
|
;MITHSONIAN MISCELLANEOUS COLLECTIONS
CHOPPERS AND HAND HAMMERS
(One-fourth actual size.)
VOL.
103, NO. 4, PL.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 4, PL. 8
LARGE BLADES FROM CACHE IN LOWER PLAIN
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOE> 1035, NO! 4, PES 9
SCRAPERS AND FLAKE KNIVES FROM CACHE IN LOWER PLAIN
(One-fourth actual size.)
on
——
_ SMITHSONIAN MISCELLANEOUS COLLECTIONS
ea gee VOLUME 103, NUMBERS
A BY
pe eae ds) KSHARLES: E.RESSER <<
Sate Curator, Division of Invertebrate Paleontology and Paleobotany
ey U.S. National Museum ~
are - CITY OF WASHINGTON
_- PUBLISHED BY THE SMITHSONIAN INSTITUTION
pea OCTOBER 21, 1942
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 103, NUMBER 5
NEW UPPER CAMBRIAN TRILOBITES
(WiTH 21 PLATES)
BY
CHARLES E. RESSER
Curator, Division of Invertebrate Paleontology and Paleobotany
U. S. National Museum
(PUBLICATION 3693)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
OCTOBER 21, 1942
The Lord Baltimore Press
BALTIMORE, MD., U. 8. A.
CONTENTS
MUEToch te ELOT MM N rere ners meter ty errs tens eee a roau teva arouses: eins ih ats my easel inmie 5a
DESCEIpLOnrot genera, and! SPECS a0 «.4 jersiie co cideics clode wceais sae ces cen sigasle
Orta s pid demo Daya Slitenna verse voreie vero reren isis erersicseie ls Gk cyaher oles sie ecsisierover:
Ghaniocepnalismntvall le ster ecyervere) ci ticle arse. s srese ayers oicclele clot aarereia’s
Irvingella Ulrich and PRESCOT Ae eeh eee ER eae yc Sie Siar oS a
ORG UINGEL GeO DAV AS Ii Me lolae ieee etiam Serrieleeia csie ae sie cincre area
Dy UMLES US MUTE WANE CllUStaerare cites lenin ierols a ote oie wae ele eretoare caret
(Oirclacsiinedwsoenet amen erry teeiachate clemre cds stolacets crele tore le evenrers eversuela cyeneie
GhevlocepialusmperdceyeaNarascrern crc tencia eicie cle eke eaetyo enews eolerelcraretae la
(COTO pa SmmN cy TO 11 MMR ge ea een a apes, tele atotei sravorcusteverehero, cra eheiericle. dere
GCN Prom a Ie War C CMULS eis chol sie elope Syslershe oles e/crov nes, 4 slavele love elm otereisvsverwis
UL CCHUIGe Semlclialll ae one nctererepecverenste (oratctorsiav sive ciavcusr athe arate hare wxereierarorace
UGLY GO UPS mlRcy til O 10d ateyaseste ie onesencxefeicesi or cis, e009) 4m. 1s. <e Sues niles ties [nlchsi eveh ey ee
LOMO WLC MMLC WAC Steere tsreierefotereueveiato' s/o) <t.5 156/510 Seo Go. tre cohen seb ue
Vi ace Tomi CimNeSSe rater tase ree ee oie a etailss Suehet otal oy Save IS Oho oe ei ede Reese
VAC POH OLmNIA Cte Meese, Scseiter eee east teva) eros aves Suez ars lepe Mra cieret nin, clove
eletwomvero piusmOiliiGly mycin ca cases cict a cias sie eisiclel cisiare sterecctns
LGWOSEONIM Vial COLEM ee reee ot orn aa ie slosh sicjavers ol cie) eeralececsi cise eee orev
BAAN All COLE ise ciner ele he Tinie icin co sie era e eee eve elisuny ene cers aieie.s
EV MAUILE LL Cmte Win CLUS tare eter eieia croichercrerscia ois e: cre: ciara ots cvevogerm a arsvelete
AV MALSIULILCR ETC Wer & CILULS Ey sbctorere hs tev elayaiaie: o. Gre -eystcrs ela sicicu'sile) sveuecwi.ersl @ieterere o1ehate
EE IT TIUS OS AVAMONG Hyapeicraeciaveiaserele'esce.9-¢ <4, < cis otis ste sieisle tiero/eidiaeiniele
EOLA CEN Al GO Leama rere tcte ster aioe hs erate. sya: Stee! etn didyaie) sisUo sms ieie cuele oyetateace
PUOMM VELL MINESS Claa te ricteretaieieiete oisie eve leis cieinie nis: 5.0/0 sydiciele oid evel e isretnshs «
VG PSOGEpHOLOLAESMNNODAYASIIe aes eier-eisveseic oor e lei s)s)sj0 2 s/eleveleyeis ereie\s
VCore ZAI TCMNNIAl CO Ltd ee acol ra etta oad eros ees coke: s/c) ater sre) ois elelieseveiesstove asia) she
VUCEC ORGS PUSHIN ESSER aytiio evel ictosaiele alate ci oiele <iaveie-olejs lols sieve elesais ainisajporets
(GOO SINN FA COE tamer steretel. race onset arene oh cvsrat save evn 2) ainsoh of ciel bhayesaucraza) eiapahere
(Goosellambachtnanr mays c relocates asi oveus orcs Ne ai eiensre sista eens
TALC HOCE DN GUNG INCSS Cline ee cicietes ce cheraie saci eies> slots. etcl che) sraielicleraloterseatarotore
Extiitiees Ae vane waa tatiitliygelercia setae eiea siers)ausie seta «eis « 20) sietole 6 /e)eys ie) Sieusie el seiels
ESALIGIFC LATIMMN NIAC OE DEMERS CT Te ores rote tae oo aroerles Sie aE are ST aL ORiar Sheed Gl her eeoeaayareicee
MecdamGStaMNvial COtt aera eiciae caeleleinis eiic sielornr® toss eleteleicieleetayels\ tevenonoieys
ESE Ia CACM IN ESS Cae eters sy ey acon = lea oterol cca cael vel sree balon'eihersi ee euehsveroneuerele
Wha lasstirecMoetiete eerie) fe oo ovorescrckersictcheve sai otc -Nie sister ars ousieta: «setorene mieten ererare
NCUA CEMA 211 OL EMME) Shere ros chs ate oee oie, as aie teat Shovetciove epmiai si ceaheskelovetencle tye
Paenicephalus Ulrich atid: Resser. 0: 2. ..j0csicce- seed ncce ce eeaes
THD: WEA NHR pepedoGn et OUS OOD HOOD TOnOboOnn cocoon oto doOsc
Bx crta et ORO sO LAL EG ae aies to ie nce i soa less velsi of'sl svete slave) sneterere¥aycici= eretVarcteretctela) secs satel
Index
eecceece este eee eet cece ese es esto eee ese eee eee eee ee seseeeeeseeesaeesece
NEW UPPER CAMBRIAN TRILOBITES
By CHARLES E. RESSER
Curator, Division of Invertebrate Paleontology and Paleobotany,
U.S. National Museum
(WitTH 21 PLaTEs)
INTRODUCTION
Soon after 1878 when Dr. C. D. Walcott centered his main efforts
on the study of the Cambrian system, he conceived the idea of a
monograph series, each of which should contain descriptions of all
Cambrian fossils representing a major animal group. With this in
mind he published new genera from time to time, frequently accom-
panied by descriptions of only one species, because the others were
left for the monograph. Thus, for example, brachiopod genera were
established and in 1912 the monograph of this group appeared. Like-
wise trilobite genera were described with the view of producing such
a monograph in due course. For this reason practically all the genera
then known from the older beds were described in 1891." ©
By 1920 it was clear that a monograph by him on Cambrian trilo-
bites, like that on the brachiopods, was not feasible. Nevertheless he
continued segregating specimens and in 1924 and 1925 published more
generic descriptions.” In the two years remaining before his death in
1927 Dr. Walcott set aside some of the specimens here presented, and
less than 2 weeks before his passing he urged me to carry on and
describe species as rapidly as possible so that they would become
available for use.
Since 1925 there has been increased activity in the study of Cam-
brian geology. Numerous papers describing many new genera have
* Walcott, C. D.:
1891. Fauna of the Lower Cambrian or Olenellus zone. toth Ann. Rep.
U. S. Geol. Surv. 18091, pp. 507-774.
1916. Cambrian trilobites. Smithsonian Misc. Coll., vol. 64, No. 3, pp. 157-
258, pls. 24-38.
1916. Cambrian trilobites. Ibid., vol. 64, No. 5, pp. 303-456, pls. 45-67.
aWwiatcott, C!D.: ;
1924. Cambrian and Lower Oczarkian trilobites. Smithsonian Misc. Coll.,
vol. 75, No. 2, pp. 53-60, pls. 9-14.
1925. Cambrian and Oczarkian trilobites. Ibid., vol. 75, No. 3, pp. 61-146,
pls. 15-24.
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 103, No. 5
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
appeared, but in most cases they also included only one or two species
in addition to the genotype. This situation leaves many of the genera
inadequately portrayed and also fails to make available the species
needed for stratigraphic purposes. Hundreds of undescribed species
contained in the National Museum collections need to be described,
both to improve the concept of genera and to supply species for
stratigraphic work. Many of them cannot appropriately be included
in faunal or regional studies and are therefore best described in
papers of this type.
Only a few of the genera represented by species described in this
paper can be referred to higher taxonomic categories. It would be a
simple matter to assign old family names or apply new ones to genera
that. bear superficial resemblances to one another, but such action
would perpetuate existing errors and introduce new ones. Trilobites
cannot yet be satisfactorily classified, and to attempt to do so merely
adds to the confusion now existing. |
DESCRIPTION OF GENERA AND SPECIES
KOMASPIDAE Kobayashi, 1935
Kobayashi established this family to include Komaspis Kobayashi,
Chariocephalus Hall, Irvingella Ulrich and Resser, Irvingelloides
Kobayashi, and Bathynotus Hall. Komaspidae was a poor selection
as the family name, for both Chariocephalus and Irvingella are far
more characteristic of the family than the obscure Komaspis. More
than 50 species of Chariocephalus and Irvingella now studied show
that the genotype of Chariocephalus is nowhere near the median point
of the generic limits, and that Chariocephalus, Irvingella, and several
other genera intergrade to the extent that arbitrary lines of separation
must be drawn. No question regarding family relationships of these
two genera can be raised, but there is no certainty that they belong
with Komaspis.
Why Kobayashi should have assigned Bathynotus to this family,
even tentatively, is difficult to understand. It was evidently done
because of its large eyes, but there is not the remotest relationship
with the Komaspidae expressed thereby.
In 1938 Kobayashi added Dartonaspis Miller to the family, at the
same time erecting two subgenera, /;vingellina and Parairvingella,
which are discussed on a later page. Also, he added the genus
Komaspidella based on Agraulos (?) thea Walcott. The latter species,
however, is a Kingstonia and therefore belongs in a different family.
The type of Dartonaspis belongs to Chariocephalus, and hence this
name becomes a synonym.
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 3
On the other hand the subgenus Parairvingella, based on Chario-
cephalus (?) tumifrons Walcott (not Hall and Whitfield) deserves
generic rank. Kobayashi’s other subgenus Jrvingellina based on Hall
and Whitfield’s species falls in Jrvingella, as subsequently discussed.
It is doubtful whether Jrvingelloides, which is based on a single
incomplete cranidium, belongs in the family. The new genus, Drum-
aspis, is added to the family Komaspidae.
The family Komaspidae is characterized by a large, more or less
quadrate glabella which extends almost the full length of the
cranidium. Occipital furrow is deep and wide, and three pairs of
glabellar furrows are usually developed. The real pair, generally
deep, may or may not be interrupted in the center, and the next pair,
if visible, consists of short lateral indentations while the anterior pair
is always faint and situated far forward. Fixigenes essentially con-
fined to palpebral lobes. Brim narrow, usually simple, but may have
a narrow rim. Eyes large, in some species longer than the glabella.
Libragenes usually rather narrow. Pygidium has a wide well-seg-
mented axis; pleural lobes fused, pleural furrows deep. Border
flattened or concave.
I recently discussed the relation of Chariocephalus and Irvingella,
but now that many more species are determined, further remarks
will be in order. As the matter now stands this family contains the
following genera.
Komaspis Kobayashi, 1935
Glabella subconical; brim concave with slight rim; eyes moderate
in length; anterior fixigenes developed.
Middle Cambrian, Asia.
Chariocephalus Hall, 1863
Glabella quadrate or expanded slightly forward; brim a simple bar ;
eyes moderately to extremely long ; libragenes large ; anterior fixigenes
reduced or absent.
Upper Cambrian (upper Franconia zones), North America.
Irvingella Ulrich and Resser, 1924
Glabella quadrate to subconical; eyes always large; anterior fixi-
genes present; brim simple or slightly rimmed; libragenes very
narrow ; pygidium like Chariocephalus.
Upper Cambrian (lower and middle Franconia zones), North
America.
4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Parairvingella Kobayashi, 1938
Glabella subconical to quadrate; eyes moderately large; brim con-
cave, with rim and preglabellar area; anterior fixigenes present.
Upper Cambrian (Franconia zones), Nevada and Novaya Zemlya.
Drumaspis, new genus (genotype, D. walcotti, new species)
Glabella quadrate or subconical; eyes smallest in family; brim
always has well-developed rim; anterior fixigenes very narrow, rela-
tively wide rearward.
Upper Cambrian (middle Franconia zones), North America.
CHARIOCEPHALUS HALL, 1863
Chariocephalus HAtt, 16th Ann. Rep. New York State Cab. Nat. Hist., p. 175,
1863; Trans. Albany Inst., vol. 5, p. 165, 1867.—Z1rTEL, Handbook Pal.,
Bd. 2, p. 603, Munich, 1885——Mrrier, N. A. Geol. Pal., p. 538, 1889.—
GraBAU and SuHimer, N. A. Index Foss., vol. 2, p. 279, 1910.—RESSER,
Smithsonian Misc. Coll., vol. 97, No. 10, p. 28, 1938.
Dartonaspis MiLiEr, Journ. Pal., vol. 10, No. 1, p. 20, 1936.
Now that a large number of species is available, the distinction
between Chariocephalus and Irvingella disappears, for the gap between
the genotypes is bridged by a series of species, so that it becomes
necessary to draw an arbitrary line of separation. As the line has
been drawn in this paper, Chariocephalus occupies a slightly higher
stratigraphic position than Jrvingella, and thus far the two genera
have not been found together at any locality. The stratigraphic
position of Chariocephalus seems to be in the beds of upper Fran-
conia age.
In view of the arbitrary separation now necessary between Chario-
cephalus and Irvingella, a new generic diagnosis is presented, taking
into account the features added by study of the many additional
species. It will be noted that the relative size and position of the eye
has become a less diagnostic criterion.
Chariocephalus is characterized by its very large, essentially quad-
rate glabella that occupies nearly the full cranidial length. It may be
slightly expanded forward, and in rare instances constricted near the
midpoint. Anterior angles are usually well rounded, and the front
outline may be gently curved, straight, or even slightly indented. If
present on the outer surface, the glabellar furrows are wide and
shallow, but most exfoliated specimens show three pairs of prominent
furrows. The rear pair may be united but usually is interrupted. If
present, the third pair is situated so far forward that the short depres-
sions arise from the rounded anterior angles. Dorsal furrow usually
deep and wide even on the outer surface. A wide occipital furrow is
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 5
apparently visible on most unexfoliated cranidia. Occipital ring heavy
and of nearly even width throughout. Brim consists of a simple bar
of even width throughout, and never as long as the glabella is wide.
It may be somewhat thickened and has various attitudes with respect
to the horizontal plane. Eyes always very large, strongly bowed, and
set off by a clearly defined palpebral furrow. Fixigenes confined to
the large palpebral lobes between the eyes and dorsal furrow. Because
the eyes usually extend far forward, they continue part way around
the anterior angles. At the rear they may extend behind the ends of
the occipital furrow, but if not, only very narrow and short postero-
lateral limbs are formed.
Libragenes narrow and long. Ocular platform narrow, long, and
curved somewhat less than the eye. Outer rim usually clearly defined,
increasing in width toward the genal angle. Genal spines present.
They vary from short to long slender spines several times the cranidial
length. They are usually directed outward at a considerable angle to
the axis. Some Wisconsin species, however, appear to have long
curved genal spines which cross over the thorax to the extent that
their outer extremities overlap.
Hypostoma nearly circular in outline, large, and characterized by a
large central lobe. The alate portions are upturned toward the edges.
Pygidium ovate, with about half the area of the cranidium. The
well-segmented axis is wide and long but tapers little. From two to
four rings are defined and the terminal lobe may have faint impres-
sions of others. The pleural platforms are convex, usually less than
half the axial width at the same point, and drop off into a narrow
flattened border.
Surface smooth.
Genotype.—C. whitfieldi Hall.
The fragment described by Raymond as Chariocephalus peloris
(Bull. Geol. Soc. Amer., vol. 48, No. 8, p. 1119, pl. 3, fig. 12, 1937)
is excluded from the genus by its small eye, and the direction and
depth of the glabellar furrows.
As stated above, Kobayashi’s Irvingella (Irvingellina) protuberans
(Jap. Journ. Geol. Geogr., vol. 5, Nos. 3-4, p. 176, pl. 15, figs. Ia-c,
2, 1938) falls into Chariocephalus.
CHARIOCEPHALUS TENERUS, new species
PLATE I, FIGURES t-3
This moderately small species, represented by a single almost
entirely exfoliated cranidium, is the only example of the genus found
in this collection.
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
The glabella is slightly expanded and the anterior angles are rounded
off. Because it is exfoliated, the three pairs of furrows show clearly,
the wide posterior pair continuing across the glabella. On the test
this glabellar furrow is very shallow and possibly interrupted in the
middle. The brim is of moderate width and slightly upturned. Exclu-
sive of the wide dorsal furrow the palpebral lobes, which also are
the fixigenes, are nearly half the width of the glabella and maintain
most of their width to the occipital furrow. The eyes’ measured in a
straight line from end to end are almost as long as the glabella and,
of course, measured around their outer edge are longer than that.
The eyes extend from the occipital furrow to a point about one-sixth
the width of the glabella beyond the lateral dorsal furrow. This
leaves a narrow fixigene slightly exceeding the width of the eye band
at the forward end of the eye. The glabella is rather evenly convex
both laterally and longitudinally, with. a moderate curvature. As can
be seen in the front view, the libragenes are rather sharply and evenly
curved, first rising considerably above the bottom of the dorsal furrow.
Honey Creek limestone; (loc. 37t) 4 miles southeast of Hennepin,
Arbuckle Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108631.
CHARIOCEPHALUS BELLUS, new species
PLATE I, FIGURES 4-6
This species, represented by cranidia only, is associated with C.
bulla,
C. bellus is characterized by its quadrate glabella. Although the
front of the glabella and the anterior angles have the aspect of a
quadrate form, the illustration shows that both are definitely rounded.
In addition to the occipital furrow, there are the usual three pairs of
glabellar furrows, the posterior pair being connected across the middle
by a faint depression while the anterior pairs are very shallow
throughout. Brim convex and turned down slightly. The fixigenes
at their widest point are about one-third the width of the glabella.
The exceedingly long eyes extend forward well beyond the latera,
dorsal furrow. The glabella is evenly convex laterally. It is highly
convex longitudinally, the curvature being concentrated in the forward
third. The fixigenes are only slightly curved in cross section and
slope down from the relatively shallow dorsal furrow at about the
same angle as the lateral slopes of the glabella. Surface punctate.
Honey Creek limestone ; (loc. 37v) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108632.
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER ye
CHARIOCEPHALUS BULLA, new species
PLATE I, FIGURES 7-II
Several cranidia and a pygidium are placed in this moderate-sized
species. C. bulla is a large-eyed form and hence resembles C. bellus
and C. tenerus.
The glabella is essentially quadrate, with rather strongly marked
furrows on the exfoliated holotype. The anterior outline is nearly
straight between the broadly rounded anterior corners, and the narrow
brim is only slightly thickened. Fixigenes are confined to the palpebral
lobes and at their widest point are a little less than half the width of
‘the glabella. The long eyes do not reach the posterior furrow, and in
front, they end exactly at the outer edge of the dorsal furrow, and
hence are relatively shorter than in C. bellus or C. tenerus. The
glabella is convex laterally with the greatest curvature near the dorsal
furrow. Longitudinally the greatest curvature is toward the anterior
end. Fixigenes are considerably curved and slope downward from
the moderately deep dorsal furrow. The pygidium is somewhat
narrow and rather convex, with the axis standing wholly above the
pleural lobes. As a whole the pygidium slopes rearward at a moderate
rate, terminating with an abrupt slope. In fact, the rear end of the
axis slightly overhangs the border. The pleural lobes form triangular
platforms that drop off to the slightly flattened border, which becomes
rather narrow at the rear of the axis.
Honey Creek limestone ; (loc. 37v) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108633a; paratype, No. 108633b.
CHARIOCEPHALUS TERES, new species
PLATE I, FIGURES I2, 13
This large-eyed species, resembling C. tenerus, is represented by a
few cranidia, none of which is complete. The illustrated holotype is
partially exfoliated and the anterior portion of the glabella abraded.
The glabella is somewhat ovate, owing more to the rounded front
margin and rounded anterior angles than to the slight constriction
toward the rear. The furrows are developed normally but may be
fainter than the average species from the region. Incomplete preser-
vation prevents an exact description of the brim but it seems to be
wide and slightly upturned, though not thickened. The fixigenes are
nearly half the width of the glabella. Eyes are very long, extending
from the occipital furrow well beyond the lateral course of the facial
suture, and therefore exceed the glabella in length. Surface punctate.
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Honey Creek limestone ; (loc. 37v) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Holotype-—U.S.N.M. No. 108634.
CHARIOCEPHALUS AFFINIS, new species
PLATE I, FIGURES 14-20
This species, represented by a number of examples of all parts, is
perhaps most closely allied to C. bulla.
The glabella is well rounded at the anterior angles, but has parallel
sides and only two pairs of furrows. It is convex, having an even
curvature laterally, but longitudinally its greatest curvature is in the
anterior third. The nearly flat fixigenes, a little more than a third the
width of the glabella at their widest point, slant downward from the
shallow dorsal furrow at a smaller angle than the lateral slope of the
glabella. The eyes are long, extending forward beyond the lateral
dorsal furrow. An associated libragene and a pygidium are assigned
to this species. The pygidium is transverse and the axis does not rise
wholly above the pleural lobes. The pleural lobes form very narrow
platforms, which are elongate rather than triangular, and which slope
into a broad concave border with a thickened, slightly upturned rim.
Honey Creek limestone ; (loc. 370) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Cotypes—U.S.N.M. Nos. 108635a-c.
CHARIOCEPHALUS AGRARIUS, new species
PLATE I, FIGURES 21-28
In present collections this moderately large-eyed form is the most
common Chariocephalus species at the locality. It is represented by
all cranidial parts and the pygidium.
The wide glabella expands only slightly, standing entirely above
the fixigenes. It is very convex laterally, with a slightly flattened
profile. Longitudinally it is also strongly convex, with the greatest
amount of curvature in the front half. The front outline is rounded,
and the dorsal furrow is relatively shallow. The brim is very slightly
thickened and is turned up to a nearly horizontal position. The fixi-
genes at their widest point are nearly half the width of the glabella.
The eyes extend from a short distance forward of the occipital furrow
to the dorsal furrow. Exfoliated specimens show normal depth for
glabellar furrows. The fixigenes are only slightly convex and slope
down from the rather broad and shallow dorsal furrow. Distinct pits
are developed at the anterior angles in the dorsal furrow. The libra-
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 9
gene selected for this species is of ordinary shape and dimensions.
The rarely found hypostoma is here preserved. It is characterized by
a tumid central lobe and a relatively narrow upturned alate portion.
The pygidium assigned to the species has a wide axis with three
well-defined rings, and stands completely above the pleural platforms.
Pleural lobes form an elongate platform which slopes abruptly to the
horizontal, slightly thickened border.
Honey Creek limestone ; (loc. 37v) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108636a; paratypes, Nos. 108636b-e.
CHARIOCEPHALUS ANGUSTATUS, new species
PLATE I, FIGURES 35-40
This is one of the smaller species and in several respects approaches
closer to the genotype than most Oklahoma forms. It is represented
by several cranidia, together with a pygidium assigned to it.
The slightly expanded glabella is rather long and has three pairs
of furrows visible. It is highly arched transversely. Longitudinally
it stands completely above the fixigenes, but the convexity in that
direction is confined almost entirely to the anterior fourth. The
occipital furrow is exceptionally wide and deep. The brim consists
of a simple bar. Pits are present at the anterior angles in the dorsal
furrow, which is wide but not as deep as usual. The fixigenes are
considerably less than half the glabellar width, and are flatly convex,
as a whole sloping only slightly downward from the dorsal furrow.
Rather long eyes are bowed most strongly anterior to the middle of
the glabella, which causes the species to have the aspect of the
genotype.
The associated pygidium assigned to the species has a moderately
high axis and relatively narrow pleural platform. The border is also
relatively narrow with slightly thickened edge.
Honey Creek limestone ; (loc. 37v) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Holotype -—U.S.N.M. No. 108637a; paratype, No. 108637b.
CHARIOCEPHALUS GRACILENS, new species
PLATE I, FIGURES 20-31
This is the smallest species in the Wichita Mountains collections,
and as far as brim structure is concerned conforms rather closely to
the genotype.
IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
The glabella is quadrate, being rounded slightly at the anterior
corners. The dorsal furrow is wide and rather deep and has pits at
the anterior angles. The occipital furrow is wide and deep, but the
glabellar furrows are faint, the rear pair showing only as faint
depressions. The fixigenes are slightly less than half the width of
the glabella. The eyes extend far forward, terminating about a half
millimeter behind the dorsal furrow pits. The brim consists of a
slightly thickened narrow bar.
Honey Creek limestone (Ptychaspis beds) ; (loc. 9y) 2 miles south-
east of Canyon Creek, Wichita Mountains, Oklahoma.
Holotype—U.S.N.M. No. 108638.
CHARIOCEPHALUS MONTIS, new species
PLATE I, FIGURES 32-34
Only a few cranidia of this small species have been located.
The glabella is practically rectangular in outline with the anterior
corners rounded. In the exfoliated holotype the furrows are so
shallow that they presumably did not show on the exterior surface.
The brim consists of a rather stout, slightly thickened bar. The dorsal
furrow is wide and fairly deep. The fixigenes at their widest point are
a little more than one-third the glabellar width, while the anterior
fixigenes are about as wide as the eye band. The eyes are very long,
extending from the occipital furrow to a point beyond the sides of
the glabella, and are highly bowed with a fairly even curvature. The
glabella is rather evenly curved laterally, and longitudinally the rate
of curvature increases from the rear pair of the glabellar furrows
forward. The fixigenes are rather flat, sloping slightly downward
from the dorsal furrow.
Honey Creek limestone (Ptychaspis beds); (loc. 91q) 8 miles
southeast of Mountain View, Wichita Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108639.
CHARIOCEPHALUS WICHITAENSIS, new species
PLATE 2, FicurEs 1-8
This is one of the larger species and is represented by several
cranidia and other parts. The cranidium expands slightly and is
rounded at the anterior corners to the straight front outline. In fact,
the front of the glabella is slightly indented. The occipital furrow is
deep, but only the rear pair of glabellar furrows shows on the exterior,
while the other two pairs are faint on exfoliated specimens. The
glabella stands above the level of the fixigenes and its convexity
laterally is produced by a gently curving top and relatively steeply
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER Tar
sloping sides. Longitudinally the moderate convexity is practically
confined to the anterior lobe. The brim consists of a narrow band in
a slightly upturned position. The fixigenes at their widest point are
about half the width of the glabella and the long eyes have a nearly
circular curvature. They extend forward to the dorsal furrow and
back practically to the occipital furrow. The narrow libragene has a
long, slender, straight genal spine which extends outward at a con-
siderable angle to the axis. The pygidium is characterized by a wide
tapering axis with three rings and terminal segment. Fusion has not
entirely eliminated the pleural grooves. Pleural platforms narrow
because of the long slope to the marginal furrow.
Honey Creek limestone (Ptychaspis beds); (loc. 91q) 8 miles
southeast of Mountain View, Wichita Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108640a; paratypes, Nos. 108640b-e.
CHARIOCEPHALUS ULRICHI, new species
PLATE 2, FIGURES 9-14
This is another large and well-represented species, similar to C.
wichitaensis. It fact the two are so similar that a formal description
of this species is not needed. C. ulrichi differs in several respects, the
most notable of which are the presence of a narrow anterior fixigene
and greater curvature of the eyes so that the rear portion of the
fixigene is relatively more contracted. The increased curvature of
the eye is accompanied by a slightly greater relative width of the
palpebral lobe making it equal to half the glabellar width.
The associated pygidium of C. ulrichi differs considerably from the
others found in Oklahoma. Its axis slopes rearward into the dorsal
furrow, which grows more shallow in the same direction, so that the
rear portion of the axis merges into the pleural lobes. The whole
pygidium slopes in all directions from the median point of the
anterior axial segment.
Honey Creek limestone (Ptychaspis beds) ; (loc. gy) 2 miles south-
east of Canyon Creek, Wichita Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108641a; paratypes, Nos. 108641b-d.
CHARIOCEPHALUS MAGNUS, new species
PLATE 2, FIGURES 15-17; PLATE 12, FIGURE I3
This beautiful, large species is represented by the complete holo-
type and several imperfect cranidia.
The quadrate glabella is slightly constricted in its middle portion.
It stands well above the fixigenes and is convex laterally, with a
circular profile. Longitudinally the considerable convexity is confined
I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
to the anterior third. The brim is not well preserved, but apparently
narrow, with a slightly upturned edge. Fixigenes, confined to the
palpebral lobes, at their widest point are less than half the width of
the glabella. Anterior fixigenes wanting. The eye, with a nearly cir-
cular curvature, extends forward to the dorsal furrow. The palpebral
lobes are flat for most of their width, but at the outer margins turn
down very sharply so that their anterior portions actually bulge over
the eye band. Dorsal furrow wide and deep on exfoliated cranidia,
while the first pair of glabellar furrows are wide but shallow. Test
smooth. Where partially exfoliated, the remaining thin portion of the
test is finely punctate.
Lyell formation; (loc. 20j) Tilted Mountain Brook, 94 miles east
of Lake Louise, Alberta.
Holotype-—U.S.N.M. No. 108642.
CHARIOCEPHALUS BADGERENSIS, new species
PLATE 2, FIGURES 18-20
This species, represented by only a few cranidia, is about normal
in size, and in general appearance approaches close to the genotype.
The glabella is quadrangular, with rounded anterior corners. Two
pairs of glabellar furrows are faintly indicated, besides the excep-
tionally wide occipital furrow. Brim relatively wide with an upturned
margin. Dorsal furrow wide and deep. Fixigenes at their widest
point less than one-third the width of the glabella. Very narrow
anterior fixigenes remain. Since the eye does not reach to the occipital
furrow, fixigenes remain posterior to the eye and are about equal to
one-fourth the glabellar width at that point. The eye extends from
a point opposite the forward ends of the occipital furrow on the
glabella to the dorsal furrow, and is rather sharply bowed. In longi-
tudinal section the cranidium is moderately convex with the greater
portion of the convexity concentrated in the anterior lobe. Laterally
the glabella is rather evenly convex and the palpebral lobes moderately
convex. Exfoliated surface punctate.
Lyell formation; (loc. 19u) Badger Pass, Johnson Canyon, 8 miles
east of Lake Louise, Alberta.
Holotype —U.S.N.M. No. 108643.
CHARIOCEPHALUS BURLINGI, new species
PLATE 2, FIGURE 21
This is the largest species of Chariocephalus found thus far.
Although the material is hardly adequate as the basis of a species, yet
its description is included because the fauna is very important for
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 13
stratigraphic reasons and further collecting is difficult. It is most
closely related to C. badgerensis.
The incomplete holotype cranidium has a large quadrate glabella,
slightly constricted in the middle. The occipital furrow is wide and
deep but does not join the dorsal furrow with the same strength. No
doubt on the outer surface of the test this furrow was either rather
shallow or obscelescent toward its ends. The rear pair of glabellar
furrows is somewhat less recurved than in C. badgerensis. The fixi-
genes at their widest point are equal to nearly half the glabellar width.
In cross section the entire cranidium is somewhat flat, and the
palpebral lobes slope gently from the dorsal furrow. The eyes appear
to have been rather long but their anterior ends are not preserved.
Both the test and the exfoliated portions are smooth.
Lynx formation; (loc. 190) Iyatunga Mountain, Robson Peak
District, British Columbia.
Holotype—U.S.N.M. No. 108644.
IRVINGELLA ULRICH AND RESSER, 1924
Irvingella Uric and Resser, in Walcott, Smithsonian Misc. Coll., vol. 75,
No. 2, p. 58, 1924—WatcorTt, ibid., No. 3, p. 98, 1925——WaLt.cortT and
Resser, Rep. Sci. Results Norwegian Exp. Novaya Zemlya, No. 24,
Videnskp., p. 10, Kristiania, 1925.—ReEssER, Smithsonian Misc. Coll., vol. 97,
No. 10, p. 33, 1938.
Irvingellina KopayasHtl, Jap. Journ. Geol. Geogr., vol. 15, Nos. 3-4, p. 175, 1938.
Relationships between /rvingella and the other genera of the
Komaspidae have been discussed. Kobayashi erected the subgenus
Trvingellina on Chariocephalus tumifrons Hall and Whitfield. Al-
though the brim of that species is not fully typical of IJrvingella,
owing to the upturned edge forming a rim, its essential structure
remains that of a bar and therefore agrees in every respect with
Irvingella. The other species that Kobayashi assigned to Jrvingellina
are referred as follows: Jrvingella arctica Walcott and Resser to
Pararvingella; Irvingella gibba Miller and J. (Irvingellina) pro-
tuberans Kobayashi to [rvingella.
Even though published descriptions of /rvingella are very brief,
they continue to be adequate in spite of the addition of many new
species.
Genotype—I. major Ulrich and Resser, 1924.
IRVINGELLA SILVESTRIS, new species
PLATE 2, FIGURES 22-27
This average-size plump species is represented at several localities
in the Arbuckle Mountains by numerous cranidia.
2
14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
The wide glabella is constricted to a rounded anterior outline. The
occipital and first pair of glabellar furrows are narrow, but clearly
defined, and in exfoliated specimens the anterior pair is visible. The
dorsal furrow is wide and deep. The glabella, slightly flattened on
top, stands wholly above the fixigenes and the dorsal furrow on the
sides while in front it overhangs that furrow. Longitudinally the head
is highly convex, the glabella curving throughout has its greatest
convexity in the anterior half. The brim is narrow, its position con-
tinuing the downward trend of the cranidial curvature. At their
widest point the fixigenes are a little more than one-third the glabellar
width. They contract rearward from their widest point which is
situated anterior to the middle of the head. Their anterior portion is
about half their average width, but because of high convexity in
dorsal view they appear much narrower. Laterally the fixigenes are
considerably curved and like the glabella have the greatest curvature
at the outer margins where they drop off to a wide palpebral furrow
and eye band, which is practically horizontal. Longitudinally the
fixigenes rise abruptly from the depressed posterolateral limbs, and
then slope gradually until near the forward end of the eye, where
they drop off abruptly to meet the depressed brim.
Honey Creek limestone ; (loc. 89w) west side of the West Timbered
Hills; and (loc. 12p) 4 miles east of Alpers, Arbuckle Mountains,
Oklahoma.
Holotype—U.S.N.M. No. 108645a; paratype, No. 108645b.
IRVINGELLA ARBUCKLENSIS, new species
PLATE 2, FIGURES 28-33
This is a very prolific species of average size. The glabella, normal
in size and depth of furrows, is constricted toward the front. The
dorsal furrow is relatively shallow, but the glabella is well differen-
tiated in cross section because it stands completely above the fixigenes.
Laterally the glabella is evenly curved, attaining a height equal to
about one-half its width. In the opposite direction the considerable
convexity is attained by declivity of the forward moiety. The brim,
of normal width, slopes downward from the anterior furrow. The
fixigenes are slightly less than half the width of the glabella; thus
the eyes are set at a considerable angle to the dorsal furrow. They
increase in width from front to back. The anterior portion is flexed
downward into a nearly vertical position and so is much wider than
appears in dorsal view. The eyes extend from about the occipital
furrow forward only to the anterior glabellar furrow, and hence are
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER I
S|
the shortest of any species observed. The anterior fixigenes are
slightly abnormal owing to the presence of a shallow anterior furrow,
which causes them to appear tumid in lateral section. Laterally the
fixigenes are only slightly convex, except at their outer edges, and
as a whole slope downward from the dorsal furrow. The eye band
is upturned a little.
Honey Creek limestone ; (loc. 89w) west side of the West Timbered
Hills, Arbuckle Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108646a; paratypes, Nos. 108646b, c.
IRVINGELLA MESLERI, new species
PLATE 2, FIGURES 34-38
This average-size species is represented by a number of cranidia.
The long glabella decreases in width forward to the rounded anterior
outline. Occipital and first pair of glabellar furrows deep; two other
pairs visible. The dorsal furrow is wide and moderately impressed.
The brim is rather wide and thick in the middle, and has a rimlike
edge owing to the presence of an incipient anterior furrow. Fixigenes,
at their widest point, nearly half the glabellar width. Their anterior
portions, which are short owing to the fact that their greatest width
occurs far forward, are about as wide as the rim and equally convex.
The long eyes extend approximately from the occipital furrow to the
brim. Laterally the fixigenes are only moderately convex and slope
down slightly from the dorsal furrow.
Honey Creek limestone; (loc. 12n) 7 miles north of Springer,
Arbuckle Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108647a; paratype, No. 108647b.
IRVINGELLA RECURVA, new species
PLATE 2, FIGURES 39-41
The glabella is rather wide and well rounded in front. The glabellar
furrows are broad and deep, and the dorsal furrow is particularly
wide and, in exfoliated specimens, also deep. In cross section the
glabella stands completely above the dorsal furrow, and is flattened
‘slightly on top. Longitudinally the rear third has a nearly straight
profile, while the anterior portion bends down sharply to overhang
the anterior dorsal furrow. The narrow brim is slightly thickened,
and when viewed from the front has a sinuate outline. The fixigenes,
which stand vertical, are only about one-fourth the width of the
glabella. They are slightly convex and have considerable width in
their anterior portions,
16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Honey Creek limestone; (loc. 12m) 7 miles north of Springer,
Arbuckle Mountains, Oklahoma.
Holotype.—U.S.N.M. No. 108648.
IRVINGELLA ALIA, new species
PLATE 2, FIGURES 42-47
This species is represented by a few cranidia which are slightly
above average size. The glabella is wide and rounded off at the
anterior angles. All furrows are narrow but only moderately im-
pressed. The glabella stands wholly above the dorsal furrow. Later-
ally it is rather convex with a nearly even curvature, but longitudinally
the curvature extends from the rear margin to the front, increasing
slowly for half the distance and then more rapidly. The brim is
narrow and slightly thickened. Fixigenes narrow, being a little more
than one-fourth the glabellar width. Their anterior angles turn down
into a vertical position. Laterally the fixigenes are slightly convex,
and slope down rather evenly from the dorsal furrow. The eyes are
very long, extending from the occipital furrow to the brim.
Honey Creek limestone ; (loc. 9p) about 15 miles northwest of Fort
Sill, Wichita Mountains, Oklahoma.
Holotype-—U.S.N.M. No. 108649a; paratypes, No. 108649b.
IRVINGELLA DAVISENSIS, new species
PLATE 2, FiGuRE 48
The genus /rvingella is very poorly represented in the Davis forma-
tion of Missouri, this being the only species that can now be described.
The glabella is rather narrow. The occipital furrow is wide and
deep, but the glabellar furrows are shallow. Laterally the glabella is
strongly convex, but not longitudinally. Brim is relatively wide and
convex. The fixigenes at their widest point are more than half the
width of the glabella at the same point. The anterior fixigenes are
somewhat wider than the brim. They rise slightly from the dorsal
furrow toward the eyes, then slope off gradually with increasing
convexity toward the outer edges. The posterolateral limbs are
sharply, and the anterior angles slightly, depressed. The eyes are of
moderate length.
Davis formation ; (loc. 22m) Shaw Branch, Davis Creek, Missouri.
Holotype —U.S.N.M. No. 108650.
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 7,
IRVINGELLA OTTERTAILENSIS, new species
PLATE 2, FIGURE 49
One imperfect cranidium is available, but the species is described
because of the importance of the fauna.
The glabella is rather large, with the occipital and first furrows
sharply impressed, but the anterior pairs do not show. Convexity
cannot be determined because of the partial compression of the fossils
from this locality. The fixigenes are more than half the width of the
glabella and the eyes are very long, extending from the occipital
furrow around the sharply bowed cheeks to the brim.
“Goodsir” formation; Moose Creek, Ottertail Range, British
Columbia.
Holotype —U.S.N.M. No. 108651.
IRVINGELLA OBLONGA, new species
PLATE 3, FIGURES 1-3
Although this locality has yielded numerous specimens of other
Irvingella, only this crandium of a small distinctly marked species
was found.
The glabella is long and subcylindrical, well rounded in front. Its
width equals about three-fourths its length. The occipital furrow and
the complete rear glabellar furrow are both deep and approximately
parallel. The glabella stands completely above the dorsal furrow, and
in cross section has practically vertical sides rounding to the slightly
flattened top surface. Longitudinally the high convexity is confined
largely to the front half so that the anterior lobe overhangs the brim.
Brim narrow and slightly upturned. Fixigenes about one-fourth the
width of the glabella, retaining about that width throughout their
length. The anterior portion decreases to about half the greatest
width. The eyes are long and, because of the rather even width of
the fixigenes, are not greatly bowed.
Wilberns formation; (loc. 70) Baldy Mountain, 8 miles northwest
of Burnet, Texas.
Holotype.—U.S.N.M. No. 108652.
IRVINGELLA AGRESTIS, new species
PLATE 3, FIGURES 4-6
The glabella tapers, owing to the even convergence of the dorsal
furrow from the occipital furrow forward. The occipital and first
18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
glabellar furrows are about equally deep and nearly parallel, and a
faint second pair is present. Glabella stands completely above the
fixigenes and is moderately and evenly curved laterally. Longi-
tudinally there is considerable curvature which increases in amount
from the occipital furrow forward. Brim of moderate width and
development ; only slightly thickened. Anterior fixigenes a little wider
than brim. At their widest point they are about one-fourth the width
of the glabella, decreasing in. width toward the occipital furrow.
Moderately bowed eyes’ extend forward nearly to the front end of
the glabella.
Wilberns formation; (loc. 69) Honey Creek, 8 miles southeast of
Llano, Texas.
Holotype—vU.S.N.M. No. 108653.
IRVINGELLA ARDMORENSIS, new species
PLATE 3, FIGURES 7-12
Only one comparatively large species of Jrvingella occurs at this
locality. An unusually wide variation in size has been allowed, for
some cranidia are larger, while others are less than one-third the
size of the specimens illustrated.
The glabella is nearly quadrangular in outline, tapering slightly
forward to a rounded frontal outline in which there is a slight
flattening medially. In cross section the glabella has considerable
elevation produced by a fairly even curvature. Longitudinally it is
also high, as shown by the illustrations. The glabellar profile shows a
rise from the rear margin in a shallow curve to the front lobe of the
elabella and then a very steep drop to the anterior furrow. Brim
narrow and slightly thickened. Anterior fixigenes taper rapidly to
the narrow brim, at their widest point being about one-third the
elabellar width. The fixigenes have only a slight convexity. The eyes
are a little more than half the length of the glabella and are only
moderately bowed.
Honey Creek limestone ; (loc. 12p) 4 miles east of Alpers, Arbuckle
Mountains, Oklahoma.
Holotype—U.S.N.M. No. 108654a; paratype, No. 108654b.
IRVINGELLA PLENA, new species
PLATE 3, FIGURES 13-15
This species, represented by numerous cranidia, is found in asso-
ciation with several other forms from which it is distinguished by its
plumpness.
*
i
if
4
a nll eae ane
a a
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 19
The cranidium is nearly as wide as long and is well rounded off at
the anterior corners. The second pair of glabellar furrows is fairly
distinct although they fail to register in the photograph. In cross
section the glabella is moderately high with a somewhat flattened top.
Longitudinally its profile is rather straight in the rear, dropping off
steeply in the anterior fourth. Brim of average width, slightly thick-
ened and situated considerably below the dorsal furrow. Fixigenes
very narrow ; about the same width as the brim at the anterior angles,
and at their widest point they equal about a fourth the glabellar width.
They are rather highly arched so that they partly overhang the eye
band.
Wilberns formation; (loc. 14b) Cold Creek, opposite north end
of Sponge Mountain; (loc. 70) Baldy Mountain, 8 miles northwest of
Burnet, Texas.
Holotype —U.S.N.M. No. 108655.
IRVINGELLA ALTA, new species
PLATE 3, FIGURES 16-18
The wide glabella is only about a fifth longer than wide. The entire
cranidium is highly convex in both directions, with the posterolateral
limbs depressed almost to a vertical position. The dorsal furrow is
narrow. Three pairs of glabellar furrows are visible; both the
occipital and the rear pair are relatively narrower and less deep than
usual. Laterally the glabella is almost evenly curved, with a tendency
toward forming a keel in the middle. The longitudinal profile is
circular except in the anterior fourth where the curvature increases
until the anterior glabellar lobe overhangs the brim. Brim narrow
and compressed closely against the glabella. Fixigenes are less than
a third the glabellar width and slope downward at a sharp angle from
the dorsal furrow. The eyes are long and somewhat angulated at the
widest point of the palpebral lobe. Surface of test evidently smooth.
Honey Creek limestone ; (loc. gp) about 15 miles northwest of Fort
Sill, Wichita Mountains, Oklahoma.
FHolotype-—U.S.N.M. No. 108656.
IRVINGELLA DECKERI, new species
PLATE 3, FIGURES 19-27
This species is represented by numerous cranidia. Specimens vary
considerably in size, but their average is about that for the genus.
Taken as a whole the cranidium has a more ovate shape than many
other species. The large glabella is rounded in front. Dorsal furrow
20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
shallow but clearly defined. Occipital furrow narrow. First pair of
glabellar furrows continues across center unchanged in width but the
anterior pairs are not visible. Laterally the glabella is moderately
convex, with concentration of the greatest convexity near the center.
Longitudinally the rear half is only slightly convex, but in the anterior
portion the convexity increases until the slope becomes vertical. The
brim is narrow and slightly thickened, sloping down from the anterior
furrow. Fixigenes are about one-third the width of the glabella, their
anterior portion measuring about one-third their average width. The
eyes are as long as the glabella, extending beyond the line of the
lateral dorsal furrow in front. The fixigenes slope downward rather
evenly from the dorsal furrow.
Honey Creek limestone ; (loc. 89y) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Holotype-—U.S.N.M. No. 108657a; paratypes, Nos. 108657), c.
IRVINGELLA BURNETENSIS, new species
PLATE 3, FIGURES 28-33
‘The cranidia representing this species are of average size and, as
the illustrations show, are rather typical in form. The cranidium is
subcylindrical in shape. The dorsal furrow is wide and relatively deep,
as are the other furrows. The glabella, convex in both directions,
stands completely above the libragenes. In cross section it has a sub-
circular outline and longitudinally it is curved throughout, but has
nearly all of its convexity in the anterior half. The brim is relatively
wide and flat, with rather sharply depressed ends. The fixigenes are
about one-third the width of the glabella and their anterior portions
are about half of their average width. The eyes are long, extending
forward almost the full length of the glabella.
Wilberns formation; (loc. 70) Baldy Mountain, 8 miles northwest
of Burnet, Texas.
Holotype-—U.S.N.M. No. 108658a; paratypes, Nos. 108658b, c.
IRVINGELLA BACCA, new species
PLATE 3, FIGURES 34-36
This small, well-defined species is not as abundantly represented as
the others found at the locality. The glabella contracts slightly to a
rounded front. Occipital and rear glabellar furrows deeply impressed ;
anterior pair faint and the median pair well developed as shown in
side view. The entire cranidium is convex longitudinally, the glabella
having its greatest convexity in the anterior half, where it overhangs
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 21
the anterior furrow. In cross section it has a distinctly flattened out-
line, although it stands completely above the fixigenes. The brim is
of normal width, slightly thickened, and because of the depressed
anterior angles, appears to be sinuate when viewed from the front.
Fixigenes, nearly half the width of the glabella, are convex and slope
downward from the dorsal furrow at an angle less than the lateral
slope of the glabella. They are also arched longitudinally and contract
in the anterior portion to a small fraction of their average width. In
dorsal view the anterior fixigenes appear narrow, owing to the nearly
vertical position of their outer portions.
Wilberns formation ; (loc. 14b) Cold Creek Canyon, opposité north
end of Sponge Mountain, San Saba County, Texas.
Holotype-—U.S.N.M. No. 108659.
IRVINGELLA ACCINCTA, new species
PLATE 3, FIGURES 37-39
The glabella, clearly defined by a rather deep dorsal furrow, con-
tracts rather strongly in the anterior portion to a rounded front. In
the exfoliated holotype the occipital and two pairs of glabellar furrows
are well developed, but the anterior pair seems to be lacking. In cross
section the glabella is distinctly flattened, and longitudinally has most
of its convexity in the anterior third. The brim is rather wide and at
its ends considerably depressed. The anterior fixigenes first slope
downward toward the front, then upward to meet the down-turned
ends of the brim. The fixigenes at their widest point are about half
the width of the glabella. They are convex laterally, rising con-
siderably above the bottom of the dorsal furrow and then sloping
down more gently, so that the eye is only slightly below the dorsal
furrow. The eye band is prominent because of the deep palpebral
furrow and the slight thickening of the band. The posterolateral
limbs are also curved into a vertical position.
Wilberns formation ; (loc. 14b) Cold Creek Canyon, opposite north
end of Sponge Mountain, San Saba County, Texas.
Holotype —U.S.N.M. No. 108660.
IRVINGELLA ABRUPTA, new species
PLATE 3, FIGURES 40-45
Like the other species at the locality, /. abrupta is about average
size for the genus. It is represented by several cranidia. The glabella
is prominent both for its large size and for its convexity, the species
deriving its name from the latter feature which serves to distinguish
22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
it from its associates. In the exfoliated holotype the occipital and the
three glabellar furrows increase in depth toward the rear of the —
cranidium. The dorsal furrow is broad and on the outside of the test
is indicated only by change of curvature. The glabella is constricted
anteriorly by the circular course of the dorsal furrow. Laterally the
glabella is highly arched, being only slightly flattened on top. Longi-
tudinally it is very convex so that the front half slopes downward
sharply from the horizontal line. The thick brim is sinuate in front
view and is slightly up-turned with respect to the anterior furrow.
Fixigenes nearly half the width of the glabella, narrowing rapidly
forward from their midpoint, to one-fourth average width. Eyes
moderately long extending from about the ends of the occipital furrow
forward to the anterior angles of the dorsal furrow. In dorsal view
the eyes curve only slightly for half their length from the occipital
furrow, then more abruptly. Laterally the fixigenes are only slightly
convex, except where they are rolled under at their outer edges, and
as a whole assume a nearly horizontal position with respect to the base
of the cranidium. Longitudinally they conform to the great convexity
of the head, but more by means of sloping forward as a whole than
by curving. In this respect /. abrupta is unique. Test smooth, but
exfoliated specimens have the usual strongly punctate surface.
Wilberns formation ; (loc. 14b) Cold Creek Canyon, opposite north
end of Sponge Mountain, San Saba County, Texas.
Holotype —vU.S.N.M. No. 108661a; paratype, No. 108661b.
IRVINGELLA MEDIA, new species
PLATE 3, FIGURES 46-54
This relatively small, abundant species is wider than the other
Texas species and in this respect finds closer relatives in the Canadian
Rockies.
The glabella is long, narrow, and considerably constricted toward
the front. The occipital furrow and rear glabellar furrows are wide
and fairly deep, while the anterior pairs have normal development.
Viewed vertically, this species seems to be very broad and flat. From
the front it is broad even though the glabella is highly arched. Longi-
tudinally, however, as the side views show, this species is strongly
convex. The brim is rather wide and has a somewhat thickened edge
and a shallow anterior furrow. The fixigenes at their widest are
about half the width of the glabella at the same point. Their rear
moiety is nearly rectangular in shape, but the anterior portion de-
creases rapidly in width, being reduced finally to the width of the
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 23
brim. The eyes are long, extending almost from the occipital furrow
to a point beyond the anterior angle of the dorsal furrow. They first
curve outward from their rear end for about a third of their length,
then bend at a rather sharp angle toward the glabella.
Wilberns formation ; (loc. 70) Baldy Mountain, 8 miles northwest
of Burnet; and (loc. 14b) Cold Creek, opposite north end of Sponge
Mountain, San Saba County, Texas.
Holotype —U.S.N.M. No. 108662a; paratypes, Nos. 108662b,
108663.
IRVINGELLA ALBERTA, new species
PLATE 4, FIGURES I-3
The wide glabella tapers more rapidly than the average for the
genus. The occipital and first pair of glabellar furrows are very deep
and wide. The middle pair consists of short, sharply impressed inden-
tations at the sides of the glabella, and the anterior pair appears only
as slight depressions. Laterally the glabella stands completely above
the dorsal furrow, its outline being slightly flattened on top. Longi-
tudinally it is curved throughout. The brim is evenly convex and of
average width. The fixigenes are less than one-third the glabellar
width anterior to the eyes, narrowing rapidly to match the rim width.
Owing to the sharp depression of the anterior angles and the tapered
glabella the entire cranidium narrows perceptibly forward.
Lyell formation; (loc. 191) Johnson Creek, 8 miles east of Lake
Louise, Alberta.
Holotype —U.S.N.M. No. 108664.
IRVINGELLA RICHMONDENSIS, new species
PLATE 4, FIGURES 4-6
This small species with a wide cranidium has relatively the widest
fixigenes of any Nevada species.
The glabella is subcylindrical, tapering slowly forward. The occi-
pital and first pair of glabellar furrows are narrow and their position
is more nearly parallel to the rear margin of the glabella than is
usually the case. Laterally the glabella has a nearly semicircular
outline. Longitudinally it is rather convex, increasing in convexity
from the rear margin forward. The brim is relatively large and some-
what thickened, sloping down in line with the declivity of the head.
Fixigenes more than half the width of the glabella, decreasing in their
anterior portions to the width of the brim. Longitudinally they are
24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
rather tumid in the rear moiety. Laterally they first rise from the
broad dorsal furrow, then slope gradually to their outer margins.
Secret Canyon shale; (loc. 60) near Richmond Mine, Eureka
District, Nevada.
Holotype-—U.S.N.M. No. 108665.
IRVINGELLA ADAMSENSIS, new species
PLATE 4, FIGURES 7-II
Like the other species at this locality, 7. adamsensis has a relatively
slender glabella and wide cranidium. It is represented by a number
of cranidia.
The glabella is constricted toward the truncated front. The furrows
are developed about as usual, but a peculiar feature is found in the
slight depression in the sides of the ridge between the occipital and
first glabellar furrows. In cross section the glabella has a flattened
curvature, and longitudinally it curves with the cranidial convexity.
The rather wide and thickened brim is striated in the middle. The
fixigenes are about half the width of the glabella, a width they main-
tain with little change in the rear half, but in the anterior portion
gradually contract to the width of the brim. Laterally the fixigenes
are rather convex, rising from a fairly deep glabellar furrow.
Secret Canyon shale; (loc. 60) near Richmond Mine, Eureka
District, Nevada.
Holotype -—U.S.N.M. No. 108666a; paratype, No. 108666b.
IRVINGELLA FLOHRI, new species
PLATE 4, FIGURES 12-14
This well-represented species is large compared to its associates.
The glabella tapers forward to a rather straight front margin, and
the furrows are developed about as usual. Laterally the convexity of
the glabella is greatly reduced, though longitudinally it curves at an
even rate. The brim is of normal width, slightly rolled. The fixigenes
are about half the width of the glabella in their rear half, while the
anterior portion is very convex, overhanging the eyes at the anterior
angles. The eyes are very long, and the eye band is conspicuously
developed.
The specific name is given in recognition of Dr. M. C. Flohr, who
was my assistant at the time when some of the fossils were collected.
Secret Canyon shale; (loc. 60) near Richmond Mine, Eureka
District, Nevada.
Holotype —U.S.N.M. No. 108667.
a EE Gr SE ES
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 25
IRVINGELLA TUMIFRONS (Hall and Whitfield)
PLATE 4, FIGURES 42, 43
Chariocephalus tumifrons Hatt and Wuirrtetp, U. S. Geol. Expl. 4oth Par.,
vol. 4, p. 224, pl. 2, figs. 38, 30, 1877.
Irvingella tumifrons RESSER, Smithsonian Misc. Coll., vol. 97, No. 10, p. 33, 1938.
Irvingella (Irvingellina) tumifrons KopayAsuHti, Jap. Journ. Geol. Geogr., vol.
15, Nos. 3-4, p. 175, 1938.
The present study has led to the conclusion that this species belongs
in Irvingella. Several additional cranidia, distorted in other directions
than in the holotype, allow a reconstruction. It will be observed in
figure 43 that on the left side of the specimen, as mounted, an anterior
furrow seems to separate a rim and preglabellar area, but on the other
side none appear>. From the fact that other specimens also fail to
show an anterior furrow, we may conclude that this seeming furrow
is merely the result of compression. It is, of course, possible that all
species of both Jrvingella and of Chariocephalus, if they have a fairly
wide anterior angle, may possess an incipient anterior furrow.
Secret Canyon shale; Pogonip Mountain, White Pine District,
Nevada.
Holotype.—U.S.N.M. No. 24561.
PARAIRVINGELLA Kobayashi, 1938
Parairvingella KopayASuHI, Jap. Journ. Geol. Geogr., vol. 15, Nos. 3-4, p. 175,
1938.
Kobayashi erected Parairvingella as a subgenus, based on the
Nevada specimen which Walcott erroneously identified as Chario-
cephalus tumifrons Hall and Whitfield, renaming it Jrvingella
(Parairvingella) angustilimbatus. Long ago I recognized the distinct-
ness of this specimen, not only specifically, but generically, and so
labeled it. Kobayashi extracted his data from my partially completed
notes. Even though there is no striking difference between Parair-
vingella and related genera, there seems to be sufficient difference to
raise it to full generic rank. I have referred Jrvingella arctica Walcott
and Resser from Novaya Zemlya to the same genus as Walcott’s
Nevada specimen, although some doubt of its true generic affinity still
remains. At the same time three new species are recognized among
the Nevada material.
Parairvingella has the glabellar and eye structure like /rvingella,
and a brim consisting of a narrow preglabellar area and rim. The
distinctive feature is the preglabellar area in which we find a transi-
20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 1038
tional structure between Drumaspis on the one hand and Elvinia on
the other.
Genotype.—Chariocephalus (2) tumifrons Walcott (not Hall and
\WWhitfield).
PARAIRVINGELLA ANGUSTILIMBATA Kobayashi
PLATE 4, FiGuRES 18-22
Chariocephalus tumifrons Waxvcort (not Hall and Whitfield), U. S. Geol. Surv.
Monogr. 8, p. 61, pl. 10, fig. 16, 1884.
Irvingella (Parairvingella) angustilimbatus JopayAsui, Jap. Journ. Geol.
Geoer., vol. 15, Nos. 3-4, p. 175, 1938.
This species, represented by a number of cranidia, is recognized at
several localities. It is characterized by a wide glabella with furrows
of normal development. The glabella stands well above the fixigenes.
In cross section it is very steep at the sides but is flattened on top.
Longitudinally it has a marked convexity, most of which is in the
anterior half. The narrow brim consists of a clearly defined thickened
rim and a narrower preglabellar area. The slightly convex fixigenes —
are almost exactly half the width of the glabella, and slope down
from the dorsal furrow. Posterolateral limbs depressed to a vertical
position. Anterior fixigenes also sharply depressed. The eyes are
long, extending forward almost to the anterior furrow, and are
angulated at about the midpoint.
Secret Canyon shale; (locs. 62, 63) north of Adams Hill, Eureka
District, Nevada.
Holotype and paratypes ——U.S.N.M. No. 24643; plesiotypes, No.
108672.
PARAIRVINGELLA EUREKENSIS, new species
PLATE 4, FIGURES 15-17
The narrow glabella tapers a little in the anterior third to a rounded
front. The occipital and first pair of glabellar furrows are wide and
deep, and in the exfoliated holotype the other two pairs are visible.
Laterally the glabella is arched above thé dorsal furrow. Longitu-
dinally it also has considerable convexity, most of which is attained
in the anterior fourth. The fixigenes are slightly more than half the
width of the glabella and, since the eyes are not greatly bowed, main-
tain their maximum width in the posterior two-thirds. Anteriorly the
fixigenes contract to about half their greatest width, and are con-
siderably depressed. The eyes are moderately long, and, because of
the wide anterior fixigenes, do not reach the line of the dorsal furrow.
'
&
/
\
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 27
Brim rather wide, with a thickened, slightly upturned rim and a
preglabellar area of about equal width. Viewed laterally the pre-
glabellar area appears to be merely an extraordinarily wide anterior
furrow. That this is not the case is shown in the anterior fixigenes,
on which the anterior furrow is clearly seen in normal position.
Secret Canyon shale; (loc. 61) south of the Hamburg Mine, Eureka
District, Nevada.
Holotype —U.S.N.M. No. 108668.
PARAIRVINGELLA INTERMEDIA, new species
PLATE 4, FIGURES 25-31
This is the most common species in the Eureka District. It is char-
acterized by great width, comparable to typical Jrvingella. The glabella
is wide and has a circular outline in front, with the usual furrows
clearly defined. It stands completely above the shallow dorsal furrow,
and longitudinally attains its convexity in the anterior third. The
fixigenes are about half the width of the glabella, contracting in their
anterior portions to about half their average width. The long eyes
attain their moderate curvature by a sharp bend near their midpoint.
The brim is narrow and consists of a thickened upturned rim and a
narrow preglabellar area of equal width. The posterolateral limbs
are bent down very sharply, while the anterior fixigenes decline into
the anterior angles to a lesser degree.
Secret Canyon shale; (loc. 61) south of Hamburg Mine, ele
District, Nevada.
FHoloty
S.N.M. No. 27018.
PARAIRVINGELLA HAMBURGEWNSIS, new species
PLATE 4, FIGURES 23, 24
Development of a wide, vertically striated preglabellar area gives
this species somewhat the appearance of Elvinia, but for the present
it may remain in Parairvingella.
The subconical glabella shows the usual furrows. Although not
highly convex in cross section it rises above the fixigenes. Longi-
tudinally the rear portion is little curved, but the anterior third curves
sharply downward. The brim width is nearly equal to one-fourth the
cranidial length and consists of a thickened upturned rim and a pre-
glabellar area of about equal width. The fixigenes average less than
half the glabellar width and maintain their width except in the anterior
portion where there is a reduction of about a third. The test is granu-
lated and the preglabellar area marked by vertical lines.
28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Secret Canyon shale; (loc. 61) south of Hamburg Mine, Eureka
District, Nevada.
Holotype —U.S.N.M. No. 108669.
DRUMASPIS, new genus
This beautiful trilobite, represented by numerous species, carries
development of the Chartocephalus-Irvingella line in a logical direction
toward more normal trilobite structure by reducing eye size and en-
larging the posterolateral limbs. The brim always has a narrow pre-
glabellar area, but taken as a whole, the departure from the bar
structure, characteristic of the other komaspid genera, is not great.
The quadrangular glabella is more or less tapered and usually has
rounded anterior angles. Occipital furrow deep. Three pairs of
glabellar furrows normally developed, the rear pair generally, and the
second pair sometimes, connected across the middle. Dorsal furrow
deep throughout. Fixigenes confined very largely to palpebral lobes
and posterolateral limbs. Eyes large, varying from less than half to
more than two-thirds the cranidial length; usually set at an angle to
the cranidial axis. Eye bands wide. Brim narrow, of even width,
and its length equals the width of the rear portion of the glabella.
Thickened rim about as wide as preglabellar area. Anterior fixigenes
narrow. Anterior facial suture usually vertical, but sometimes diver-
gent. Posterolateral limbs fairly large in the smaller-eyed species.
Libragenes small, with thickened rim. Ocular platforms narrow in
front, increasing in width posteriorly.
Pygidium not found. Surface usually granulose.
Genotype.—D. walcotti, new species.
DRUMASPIS WALCOTTI, new species
PLATE 4, FIGURES 37-41
This species has a long glabella which tapers slightly. Three pairs
of glabellar furrows are clearly discernible. The fixigenes vary con-
siderably in width owing to the strong curvature of the eyes. At their
widest point they are a little more than a third of the glabellar width.
The brim consists of a thickened rim and a slightly narrower pre-
glabellar area. The glabella is slightly arched in cross section, and
longitudinally curves considerably more, particularly in the anterior
portion. The conspicuously wide eye bands at once set the species off
from others found in the vicinity.
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 29
St. Charles formation; (locs. 4y, 5e) Two Mile Canyon, Wasatch
Mountains, Idaho.
Holotype —U.S.N.M. No. 108670a ; paratype, No. 108670b.
DRUMASPIS IDAHOENSIS, new species
PLATE 4, FIGURES 32-36
This species is represented by many cranidia. The glabella tapers
slightly. Peculiar structure is shown by the occipital furrow, which
is narrow next to the dorsal furrow, then widens suddenly, main-
taining that width to the similar point on the opposite side. The rear
pair of glabellar furrows is interrupted in the middle both by growing
shallow and by making a reversed forward curve. The anterior pairs
show on exfoliated specimens. In cross section the glabella is rather
flat but stands completely above the dorsal furrow. In longitudinal
section it curves gently for most of its length, then turns down sharply
at the front. The narrow brim consists of a rim in horizontal position.
The fixigenes are a little less than one-third the glabellar width and
in their anterior portion decrease to about the width of the anterior
dorsal furrow. They slope downward rather evenly from the dorsal
furrow, their declivity increasing in the rear to meet the sharply
depressed posterolateral limbs, and anteriorly to join the rim. The
moderately bowed eyes attain a length equal to about half the length
of the glabella.
St. Charles limestone; (loc. 4y) Two Mile Canyon, 2 miles south
of Malad, Wasatch Mountains, Idaho.
Holotype—U.S.N.M. No. 108671a; paratype, No. 108671b.
DRUMASPIS ALBERTA, new species
PLATE 5, FIGURES 1-3
The description of this species is based on several cranidia, the
most complete example being figured as the holotype.
The quadrangular glabella has a slightly indented anterior margin.
Anterior angles rounded. In cross section the glabella is only slightly
arched above the dorsal furrow. Longitudinally there is considerable
elevation, most of which occurs in the anterior fourth. Brim narrow,
apparently with a narrow upturned rim. Fixigenes about half the
width of the glabella; anterior portion reduced to brim width. Eyes
not strongly bowed, set at a wide angle to the dorsal furrow, forming
a fixigene of fairly even width for the rear two-thirds of the head.
3
30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I03
The fixigenes are almost flat and slope down from the dorsal furrow
in line with the lateral slope of the glabella. Surface finely granulose.
Lyell formation; (loc. 64x) Ranger Brook Canyon, Sawback
Range, Alberta.
Holotype-—U.S.N.M. No. 108673.
DRUMASPIS BRISCOENSIS, new species
PLATE 5, Ficures 4-8
This species is represented by several cranidia, of which two ex-
foliated examples are figured.
The glabella is rectangular, with a practically straight front margin
and slightly rounded anterior angles. Four sets of furrows are clearly
defined in exfoliated specimens. In cross section the glabella rises
with a flat curvature above the dorsal furrow, while longitudinally it
is curved throughout, the curvature increasing from the rear forward.
Brim of moderate width consists of a thickened rim, widened slightly
in the middle. Fixigenes at their widest point slightly more than a
fourth the glabellar width. Eyes rather sharply bowed. Anterior
fixigenes depressed sharply. Portion of the test preserved appears to
be smooth.
Sabine formation; (loc. 17p) Radium Hot Springs, Brisco Range,
British Columbia.
Holotype-—U.S.N.M. No. 108674a; paratype No. 108674b.
DRUMASPIS GOODSIRENSIS, new species
PLATE 5, FIGURE 9
Only the holotype cranidium has been found at this locality.
The glabella tapers slightly forward to a straight anterior margin.
The usual three pairs of glabellar furrows are developed. Owing to
crushing in the shale matrix, the convexity cannot be determined. The
fixigenes at their widest point are a little more than a third the glabellar
width. The eyes are not greatly bowed and, because of the relatively
great width of the anterior fixigenes, depart from the course of the
dorsal furrow only at a small angle. Brim of average width, con-
sisting of a slightly thickened rim of about the same width as the
anterior furrow.
“Goodsir” formation; Moose Creek, Ottertail Range, British
Columbia.
Holotype —U.S.N.M. No. 108675.
,
i
,
i
;
¥
m
NO. 5 UPPER CAMBRIAN TRILOBITES—-RESSER ou
DRUMASPIS SABINENSIS, new species
PLATE 5, FIGURES I0, IT
This distinctive species is represented only by a few cranidia.
The rectangular glabella is rather long and rounded at the anterior
angles. Both laterally and longitudinally it has average convexity for
the genus. The moderately wide brim is made up about equally of
anterior furrow and rim. The rim has a horizontal position and
widens out in the middle. The fixigenes, at their widest point, are less
than one-third the glabellar width. Their anterior portions are turned
down sharply, leaving a small ridge which extends into the preglabellar
area. The eyes are of moderate size and moderately bowed.
Sabine formation; (loc. 17s) Sabine Mountain, 2 miles northeast
of Canal Flats, Brisco Range, British Columbia.
Holotype —U.S.N.M. No. 108676.
DRUMASPIS MAXWELLI, new species
PLATE 5, FIGURES I2, 13
About a dozen cranidia of this species are available.
The glabella is of normal shape and proportion, rounded at the
anterior corners. The usual glabellar furrows are present but not
deeply impressed. In cross section the glabella stands completely above
the fixigenes and dorsal furrow, rising steeply on the sides, but is
flattened on top. Longitudinally the cranidium is considerably arched,
the convexity increasing from the rear forward. The brim is of
normal width and upturned to a slightly thickened brim, which in the
holotype cranidium is partly broken off. Fixigenes are narrow, being
little more than one-fourth the giabellar width. Their anterior portion
is about as wide as the brim and is turned down at the anterior angles.
The eyes are of moderate size and slope outward at a small angle.
St. Charles limestone; (loc. 66z) St. Charles Canyon, Bear River
Range, Idaho.
Holotype —U.S.N.M. No. 108677.
DRUMASPIS DECKERI, new species
PLATE 5, FIGURES 14-16
Even though the available material is confined to the illustrated
holotype, the species is so distinctive that it merits description. The
quadrangular glabella is rounded at the anterior angles and slightly
indented in front. Glabellar furrows developed as usual, the rear
32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
pair containing pits on each side of the medial line. Laterally the
glabella is only moderately convex, but longitudinally it is considerably
more curved. The brim is of normal width, consisting mainly of a
wide rim, in horizontal position, with a lingual extension in the middle.
The fixigenes at their widest point are a little less than one-third the
glabellar width, and their anterior portions, being sharply depressed,
are narrower than the brim. The large eyes are sharply bowed.
Honey Creek limestone; (loc. gtb) 4 miles southeast of Hennepin,
West Timbered Hills, Arbuckle Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108678.
DRUMASPIS OSELLA, new species
PLATE 5, FIGURES 17-20
Among the numerous specimens of the genus in the Texas collec-
tions, two cranidia present very distinct characteristics and are thus
separated from D. tevana. The glabella tapers forward more than
the average amount for the genus and is rounded at the anterior
angles to a straight front margin. Four pairs of glabellar furrows
are defined. In cross section the entire cranidium is convex, the
glabella standing above the dorsal furrow with a rather flat curvature.
Longitudinally the head is rather convex, the greatest amount being
in the anterior third. Brim rather wide, consisting of a thickened
rim and a slightly narrower preglabellar area. The fixigenes at their
widest point are less than one-third the width of the glabella. Their
anterior portion is slightly wider than the preglabellar area. The eyes
are evidently sharply bowed and not very large. Both the anterior
angles and the posterolateral limbs are depressed rather sharply. The
surface, except in the furrows, is ornamented with anastomosing lines,
which, in places, practically form granules.
Wilberns formation; (loc. 70) Baldy Mountain, 8 miles northwest
of Burnet, Texas.
Holotype—U.S.N.M. No. 108679a; paratypes Nos. 108679b, c.
DRUMASPIS TEXANA, new species
PLATE 5, FIGURES 27-30
This species is represented by numerous specimens from several
localities. The rather wide glabella contracts slightly forward and is
well rounded at the anterior angles. The rear pair of glabellar furrows
is deeply impressed, the next pair faint, and the forward pair very
faint. In cross section the glabella is elevated as usual, longitudinally
it is curved throughout. The brim consists of a narrow preglabellar
NO. 5 UPPER CAMBRIAN TRILOBITES—-RESSER 33
area and a slightly thickened rim expanding in the middle. Fixigenes
at their widest point are a little less than one-third the glabellar width.
Eyes of moderate size, rather sharply bowed. Anterior fixigenes
about as wide as the brim, increasing forward into the down-turned
anterior angles. Surface granulated.
Wilberns formation ; (loc. 70) Baldy Mountain, 8 miles northwest
of Burnet, Texas; and (loc. 670) Potatotop, 7 miles northwest of
Burnet, Texas.
Holotype.—U.S.N.M. No. 108684a; paratype Nos. 108684b, c.
DRUMASPIS WICHITAENSIS, new species
PLATE 5, FIGURES 21, 22
More than 10 cranidia of this species are in the collection. The
quadrate glabella is rounded by even curves at the anterior angles.
Glabellar furrows are developed normally except that the front pair
is almost obsolescent. In cross section the glabella stands completely
above the fixigenes, rising rather steeply at the sides and flattened on
top. Longitudinally the cranidium is high, with the front lobe of the
glabella standing vertical. Brim of moderate width, consisting of a
narrow preglabellar area and a slightly widened rim in horizontal
position. Fixigenes narrow, at their widest point being a little more
than one-fourth the width of the glabella. Anterior fixigenes sharply
down-curved, attaining a width at the anterior angles of slightly more
than the brim. Eyes rather small and highly bowed, with practically
a circular course. Posterolateral limbs curved downward into a
vertical position. Surface faintly marked by anastomosing lines.
Honey Creek limestone; (loc. 91u) 1 mile east of Canyon Creek,
15 miles northwest of Fort Sill, Wichita Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108680.
DRUMASPIS CLARA, new species
PLATE 5, FIGURES 23-26
Several cranidia and a libragene represent this species.
The glabella tapers forward to a nearly circular frontal outline.
Glabellar furrows developed normally. In cross section this is a highly
convex species. The glabella, standing completely above the libra-
genes, rises with a rather even curvature throughout. Longitudinally
the cranidium also has high relief, likewise attained by a rather even
curvature. Brim is of normal width. The slightly thickened rim
widens in the middle, and occurs in a horizontal position. Fixigenes
slightly less than one-third the glabellar width, their anterior portion
34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
rather wide and depressed at the anterior angles. The eyes are of
moderate size and not greatly bowed. The associated libragene has a
heavy, striated rim, so constructed that it shows the suture to be
intermarginal for more than one-third the width of the head. Ocular
platform rather small, and covered with anastomosing lines similar
to the elevated portions of the cranidium.
Honey Creek limestone; (loc. 91s) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Holotype-——U.S.N.M. No. 108681a; paratype No. 108681b.
DRUMASPIS NITIDA, new species
PLATE 5, FIGURES 34, 35
This species is represented by several cranidia. The wide, nearly
quadrate glabella is well rounded in front. All the furrows are
moderately developed. The brim is normal in width and consists of
a narrow preglabellar area, and a slightly thickened rim which
broadens toward the middle. The fixigenes are narrow, at their
widest point being little more than one-fourth the glabellar width. In
their anterior portions they are about the same width as the brim, but
widen toward the depressed anterior angles. The eyes are somewhat
larger than average size and slightly bowed, the bowing attained by
a sharp angular turn near the midpoint. A distinctive feature is the
extraordinary width of the eye band. Surface covered by irregular
raised lines which are equivalent to irregular curved granules.
Honey Creek limestone; (loc. 91s) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108682.
DRUMASPIS UTAHENSIS, new species
PLATE 5, FIGURES 31-33
In the National Museum collections are several cranidia which
when first studied were thought to belong to Chariocephalus. How-
ever, when comparison was made with typical forms of that genus
and of Drumaspis, they appeared to be nearer the latter.
The quadrangular glabella has a straight anterior margin and
rounded anterior angles. The glabellar furrows are well developed,
and even the anterior pair may be traced across the glabella. In cross
section the glabella stands completely, but not high, above the dorsal
furrow and is nearly flat on top. Longitudinally the relief is con-
siderable, but the curvature is nowhere great except in the very front
of the cranidium. The fixigenes are wide, being half the glabellar
NO. 5 UPPER CAMBRIAN TRILOBITES—-RESSER 35
width at their widest point. The very long eyes are curved so sharply
that the two halves are almost at right angles to each other. Anterior
fixigenes practically wanting and on the whole are nearly flat, sloping
down from the dorsal furrow. The posterolateral limbs are very
sharply depressed.
St. Charles limestone; (loc. 54x) Two Mile Canyon, 2 miles south
of Malad, Wasatch Mountains, Idaho.
* Holotype —U.S.N.M. No. 108683.
UNCLASSIFIED GENERA
CHEILOCEPHALUS Berkey, 1898
CHEILOCEPHALUS WICHITAENSIS, new species
PLATE 5, FIGURE 39
Several cranidia of this interesting genus have been found in the
Oklahoma collections. It is a very simple’ trilobite, the cranidium
consisting of a large glabella with no furrows except a shallow
occipital furrow. The fixigenes are simple. The glabella tapers
slightly to a nearly straight anterior margin. In cross section it stands
well above the dorsal furrow, rising at a fairly even rate to approxi-
mately the center. Longitudinally, the head has considerable relief,
attained by even curvature. The brim is narrow and consists of a
simple, slightly concave, and nearly horizontal extension, the outer
edge of which is very slightly thickened but does not form a rim. The
fixigenes have nearly the same width throughout, and are about equal
to one-fourth the glabellar width. Test smooth.
Compared to C. stcroixensis the Oklahoma species is slightly smaller
and differs in the distribution of convexity in both directions.
Honey Creek limestone; (loc. gp) 15 miles northwest of Fort Sill,
Wichita Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108685.
CHEILOCEPHALUS TEXANUS, new species
PLATE 5, FIGURES 36-38
The one small cranidium in the Texas collections has a smooth
rectangular glabella, rounded in front. In cross section it forms a
continuous curve with the fixigenes, which are barely separated by
the dorsal furrow. Longitudinally, the cranidium is convex, with a
rather even curvature, increasing in the anterior portion. The brim,
on the whole, is slightly concave. No anterior furrow is developed,
but a thickened rim turns up slightly, and is thereby separated from
30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
the preglabellar area. The fixigenes, at the rear of the eyes, are about
half the glabellar width and form wide triangular posterolateral limbs.
Anterior to the rather small eye they are somewhat wider than the
brim. As the cranidium has no occipital furrow, it presents a very
smooth appearance.
Hickory sandstone; (loc. 68z) Packsaddle Mountain, 11 miles
southeast of Llano, Texas.
Holotype.-—U.S.N.M. No. 108686.
CHEILOCEPHALUS BUTTSI, new species
PLATE 11, FIGURE 6
Unfortunately many years ago the holotype and other cranidia
representing the species were damaged by poor preparation. This
cranidium is fully typical of the genus, even as to size. The large
glabella extends the full length of the head and has three pairs of
shallow furrows. It tapers only slightly. Though incomplete the
fixigenes are seen to be small anterior to the large posterolateral limbs.
Longitudinally the cranidium is little curved except in the anterior
fourth where the slope becomes vertical. In cross section the entire
head has considerable convexity attained by a rather evenly convex
glabella and continued by the lateral slopes of the fixigenes.
Surface beautifully shagreened.
Ore Hill limestone; (loc. 107v) 43 mile northwest of Drab, Penn-
sylvania.
Holotype.
U.S.N.M. No. 108746.
CHOLOPILUS Raymond, 1924
CHOLOPILUS NEVADENSIS, new species
PLATE 6, FiGcuRES I-2
The main features of this species relate it to Cholopilus. Further-
more its stratigraphic position is approximately the same as other
representatives of the genus. Numerous cranidia are in hand, but
no other parts.
C. nevadensis is characterized by its simplicity. The very large,
smooth glabella extends nearly to the anterior margin. The dorsal
furrow, fairly well impressed in the rear, decreases in depth forward
until beyond the eye it can be observed only as a very faint line
beneath the surface. A weak occipital furrow is developed but no
glabellar furrows show on the outer surface. Exfoliated specimens
have the merest trace of three pairs of glabellar furrows, the first pair
recurved, the second pair turning a bit forward, and the third again
NO. 5 UPPER CAMBRIAN TRILOBITES—-RESSER 37
turned backward, so that the second and third pairs form a faint cross
on the glabella. Laterally the glabella is rather evenly but flatly
arched ; longitudinally it is considerably curved. The brim continues
the rounded curvature of the glabella and consists of a preglabellar
area and a narrow, wirelike rim, but faintly indicated. Fixigenes at
their widest point are about one-fifth of the glabellar width, and they
maintain practically the same width throughout. The eyes are of
normal size, situated about the midpoint of the cranidium. A faint
tubercle which has more the appearance of a median eye than of a
spine occurs on the occipital ring.
Hamburg limestone; (loc. 23d) near Hamburg Mine, Eureka
District, Nevada.
Holotype —U.S.N.M. No. 108687.
CHOLOPILUS (?) ALBERTENSIS, new species
PLATE 10, FIGURE 3
This cranidium of doubtful generic position is described in order to
present the faunal element. Because there are no good illustrations
of the type of Cholopilus extant, one cannot be sure that the published
drawings rightly portray the glabellar proportions. The general ap-
pearance of this trilobite places it nearest to Cholopilus, in which
genus it may be placed for the present.
The glabella is wide, occupying more than half the cranidial area.
It is defined by a dorsal furrow so faint that it disappears in certain
light. This cranidium is convex in both directions, the curvature being
greater longitudinally than laterally. The anterior margin is slightly
indented in the middle and a very faint rim is indicated.
Lyell formation ; (loc. 64b) Glacier Lake Canyon Valley, 48 miles
northwest of Lake Louise, Alberta.
Holotype —U.S.N.M. No. 108733.
CALYPTOMMA, new genus
A small trilobite characterized by simplicity of structure. The large
glabella is quadrate except for constriction at the anterior angles.
Glabellar furrows are very faint even on exfoliated specimens. Brim
a simple bar. Fixigenes very small, confined to the palpebral lobe,
and in the genotype are less than half the width of the eye band. Eyes
so large that they envelop the glabella, extend from the forward
margin of the glabella beyond the occipital ring. Eye band wide.
This genus resembles Cholopilus Raymond in the simplicity and
arrangement of the various cranidial parts. It differs from that genus
38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
in two respects: first, the fixigenes are narrower, and second, the
eyes are very much larger.
Surface marked by elongate granules on the elevated portions.
Name.—«xaAvzrw = envelop ; 6upa=eye.
Genotype.—C. typicale, new species.
CALYPTOMMA TYPICALE, new species
PLATE 12, Ficures 8, 9
The generic description and illustrations present the specific
characters.
Red Lion formation; (loc. 150e) Boulder Creek, 1 mile north of
Princeton, Montana.
Holotype-—U.S.N.M. No. 108753a; paratype, No. 108753b.
ILLAENURUS Hall, 1863
ILLAENURUS PRISCUS, new species
PLATE 6, FIGURES 3-7
This species, though not fully typical of /Waenurus, does not seem
to warrant the establishment of a separate genus. The four cranidia
figured differ slightly in proportions, and therefore it is possible that
two species are included. Departure from essential features of
Illaenurus is found in the great width and faint delimitation of the
glabella. Exfoliated specimens show a completely outlined glabella,
and a considerable width of fixigene. It has generally been assumed
that Jllaenurus has no fixigenes, or in other words, that the dorsal
furrow passes through the two ends of the eye lobe.
In general outline the cranidium is nearly quadrate. J. priscus has
a narrow, wirelike rim, and eyes of moderate size situated about the
midpoint of the cranidium. Anterior to the eye the facial suture
diverges slightly, and the anterior angles are turned down somewhat
by a continuation of the cranidial slope. In cross section the glabella
has a flat curvature, but longitudinally a much greater one.
Lyell formation; (locs. 66j, k) Ranger Canyon, Sawback Range,
Alberta.
Holotype—U.S.N.M. No. 108688a; paratypes Nos. 108688b,
108689a, b.
ILLAENURUS ALBERTENSIS, new species
PLATE 6, Ficures 8-12; PLATE 14, FicurE 18
This species is represented by numerous cranidia, and by a pygidium
and libragene.. It is not greatly unlike several undescribed species
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 39
from the Upper Mississippi Valley. On exfoliated cranidia the dorsal
furrows show faintly for a short distance anterior to the eye, touch-
ing both ends of the eye lobe. Exfoliated specimens have a median
eye situated about one-third the distance forward from the rear
margin. In front of the eyes, which are situated behind the middle
of the cranidium, the facial suture diverges considerably to form wide
anterior angles. The thickened, heavily striated rim is almost vertical
to the horizontal plane of the cranidium. In cross section the cranidium
has a slight even curvature, but longitudinally it has considerably more
relief, also attained by even curvature.
The libragene forms nearly a quarter circle, with its convexity
similar to that of the cranidium. The heavily striated border con-
tinues around the margin, decreasing toward the genal angle, which
evidently bore no spine.
The pygidium is short, very wide, and lacks differentiation of axis
or pleura. At the anterior margin two slight depressions indicate the
presence of a very wide axis. The pygidium evidently came to rather
sharp points at its lateral angles.
Portions of six simple thoracic segments remain articulated on one
rock fragment.
Lyell formation; (loc. 20d) Tilted Mountain Brook, 9 miles east
of Lake Louise, Alberta.
Holotype —U.S.N.M. No. 108690a; paratype, No. to86gob.
ILLAENURUS (?) SINCLAIRENSIS, new species
PLATE 6, FIGURES 13-15
This species, doubtfully referred to Jllaenurus, comes from some-
what older beds than those that contain the more characteristic species.
The cranidium is wider than long and exclusive of the posterolateral
limbs is rectangular in outline. No trace of dorsal furrows has been
observed. In cross section this species is rather flat, but longitudinally
it is considerably convex, with the anterior margin of the cranidium
turned under. A peculiar feature is the development of a rim like
that of Platycolpus. A small occipital tubercle suggests the presence
of a medial eye, and short shallow multiple furrows near it evidently
represent the occipital furrow. The eyes are of moderate size, situ-
ated slightly forward of the median point of the cranidium.
Sabine formation ; (loc. 16t) Sinclair Canyon, Brisco Range, British
Columbia.
Holotype —U.S.N.M. No. 108691a; paratypes, Nos. 108691b, c.
40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
ILLAENURUS ELONGATUS (Walcott)
PLatre 6, Frcures 18, 19
Tsinania elongata Watcott, Smithsonian Misc. Coll., vol. 64, No. 3, p. 228,
pl. 36, figs. 10, 10a, 1916.
The cranidium on which this species is based is not average for
Tilaenurus. It is long and narrow, with little curvature in cross section
but with somewhat more in longitudinal section. No dorsal furrow
or rim seems to be differentiated. The anterior angles are rounded
more than average for the genus. The eyes are about normal size,
situated slightly forward of the glabellar midpoint.
McKay group; (loc. 23z) 2 miles west of Donald, Dogtooth
Mountains, British Columbia.
Holotype-—U.S.N.M. No. 61736.
PLATYCOLPUS Raymond, 1913
PLATYCOLPUS QUINNENSIS, new species
PLATE 6, FIGURES 16, 17
The small collection from this locality has yielded a good cranidium
of Platycolpus, which is described because of its significance in the
fauna. It is normal in all essential characters of the genus. A wide
glabella, faintly indicated by a shallow dorsal furrow, extends a little
in front of the eyes. The rim is wide and heavily striated. Anterior
to the eyes the facial suture diverges, forming large anterior angles.
The eyes are of moderate size, situated well back on the head. A
shallow occipital furrow delimits a wide flat occipital ring.
Mendha limestone; (loc. 7j) Quinn Canyon Range, Nevada.
Holotype—U.S.N.M. No. 108692.
PLATYCOLPUS OKLAHOMENSIS, new species
PLATE 6, FIGURES 20-25
This species is recognized at several localities. As usual only frag-
ments of the cranidium are preserved, a condition characterizing most
species thus far recognized in the genus. The portion of the glabella
available shows that it is wide, and rather evenly but not highly arched
in both directions. It has the usual wide brim, although it lacks the
usual striations.
The pygidium is normal in every respect. Exfoliated specimens
show a well-separated axis. On the outer surface it is marked simply
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER AI
by a change in slope. The pygidium is flatly convex in transverse
section, but as may be observed from the illustration is much steeper
in the opposite direction. The species is of average size and the
surface seems to have been smooth.
Signal Mountain formation; (loc. 186s) 4 miles northwest of
Ravia; (loc. 201j) Royer Ranch, Honey Creek, Arbuckle Mountains,
Oklahoma.
Holotype-—U.S.N.M. No. 108693 ; paratypes, Nos. 108694, b.
PLATYCOLPUS WICHITAENSIS, new species
PLATE 6, FIGURES 26-29
This species is represented by several cranidia and a pygidium, and
even by a fragment of a libragene. P. wichitaensis averages slightly
smaller than P. oklahomensis. The fragmentary cranidium figured
has a faint dorsal furrow behind the eyes, and in both lateral and
longitudinal sections is evenly and considerably arched. The eyes are
of normal size and are situated about the midpoint of the cranidium.
The brim is sharply demarcated, thickened, and heavily striated.
The pygidium on the same piece of rock with a cranidium of
Eurekia, is also highly arched in both directions and shows faint
dorsal and axial furrows.
Signal Mountain formation ; (loc. 12g) 2 miles southwest of Signal
Mountain, Wichita Mountains, Oklahoma.
Holotype-—vU.S.N.M. No. 108695a; paratype, No. 108695b.
PLATYCOLPUS HIGHLANDENSIS, new species
PLATE 6, FIGURES 30, 31
This species constitutes another important element of the fauna
from the upper beds in the Highland Range, and consequently is
described, even though the material is fragmentary. As usual no
cranidia are preserved. The pygidium on which this species is based
is itself incomplete, but has the normal generic features, and it is of
rather large size. As shown in the illustrations the axis is faintly
defined, tapering less rapidly than in other species. It is not highly
arched in either direction, but has normal distribution of convexity.
Mendha limestone; (loc. 88) 7 miles north of Bennett Springs,
Highland Range, Nevada.
Holotype —U.S.N.M. No. 27017.
42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 108
PLATYCOLPUS SINCLAIRENSIS, new species
PLATE 6, FicuRES 32-36
Though there are in hand about Io pygidia and about an equal
number of libragenes, only one fragmentary cranidium, not altogether
typical, has been found in the collection.
The libragene is almost flat in its rear portion but increases greatly
in convexity forward, indicating that the cranidium must have been
considerably arched longitudinally. The ocular platform is approxi-
mately a quarter circle and is surrounded by a wide, thickened, heavily
striated border, with the striations turning toward the margin as the
genal angle is approached.
The pygidium is rather convex, with the axis standing well above
the pleural lobes, but the dorsal furrow is merely a change in slope.
Several pleural furrows are visible on exfoliated specimens. The axis
is rather long, extending beyond the midpoint of the pygidium.
Sabine formation; (loc. 16t) Sinclair Canyon, Brisco Range,
British Columbia.
Holotype.—U.S.N.M. No. 108697a; paratypes Nos. 108697), c.
ENONTIOURA, new genus
Although up to the present time only the holotype pygidium has
been discovered, it is given a new generic name because it represents
such a strange form. On the same piece of rock is a rounded
cranidium similar to that of Camaraspis but it is not well enough
preserved to warrant description.
The specimen assumed to be a pygidium is characterized by a very
large axis, which occupies most of its area. At the front of the wide
axis is a half segment which looks more like a rim than the usual
segmental division. Shallow dorsal furrows separate narrow pleural
lobes, the dorsal furrow continuing very faintly around the rear of
the axis. Beginnning at the anterior angles a thickened, slightly
upturned rim borders the outer edge of the pygidium. Aside from
the peculiarity of shape and convexity, a very unusual feature is found
in the striations which transverse the entire specimen from side to
side, crossing both the axis and the pleural lobes. It is this feature
which has several times raised the question as to whether this specimen
is actually a trilobite pygidium or some oddly formed hypostoma, or
even an undescribed crustacean. In view of these conditions, a descrip-
tion of this specimen is presented primarily to call it to the attention
of geologists with the hope that its real position may be discovered.
Name.—evwvrios = contrary ; ovpa=tail.
Genotype.—E. typicalis, new species.
|
’
.
a
MO 5 UPPER CAMBRIAN TRILOBITES—RESSER 43
ENONTIOURA TYPICALIS, new species
PLATE 6, FIGURES 37, 38
The generic description, together with the illustrations, presents the
characteristics of this peculiar form.
Mendha limestone; (loc. 88) 7 miles north of Bennett Springs
Highland Range, Nevada.
Holotype —U.S.N.M. No. 108696.
MACELLOURA Resser, 1935
MACELLOURA TRANSVERSA, new species
PLATE 7, FIGURE I
When first observed this pygidium was thought to represent M. dia.
Comparison shows that, while typical of the genus in every respect,
the pygidium is wider than that of M. dia.
Wilberns formation; (loc. 14f) Bartlett Hollow, Burnet County,
Texas.
Holotype—U.S.N.M. No. 108608.
ARAPAHOIA Miller, 1936
Arapahoia Miter, Journ. Pal., vol. 10, No. 1, p. 24, 1936.
Hesperaspis StoyANow, Bull. Geol. Soc. Amer., -vol. 47, No. 4, p. 460, 1936.
At the time Arapahoia was originally described, only the cranidium
was known to the author. Specimens from the eastern front of the
Beartooth Mountains identified with the genotype A. typa now add
the libragene and pygidium.
In the original description Miller pointed out the relationship of
Arapahoia to Plethometopus, which occurs in younger Upper Cam-
brian strata. From the present study it becomes clear that Norwood-
ella described later is also related and that Arapahoia, Norwoodella,
and Plethometopus form a single line of development. Thus far
Arapahoia seems to be confined to horizons in the lower part of the
Cedaria zone, and is the oldest genus of this group.
In spite of the abundance of examples, it is difficult to free good
specimens of Arapahoia, because the matrix is refractory.
Exfoliated cranidia have well-defined dorsal and occipital furrows,
and some species have three pairs of short glabellar furrows. A node
on the base of the occipital spine may possibly be a median eye.
The libragene is of moderate size. In some specimens a border is
faintly indicated, but it scarcely breaks the even curvature as the
ocular platform slopes to the margin. The entire libragene extends
into a longer or shorter genal spine.
44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
The pygidium is short and wide, with a large axis, which seems to
be segmented in all species. Some species have well-developed pleural
furrows, and a few show a border.
ARAPAHOIA TYPA Miller
PLATE 7, FIGURES 37, 38
Arapahoia typa MiLier, Journ. Pal., vol. 10, No. 1, p. 25, pl. 8, fig. 6, 1936.
The figured specimens from the Beartooth Mountains are identified
with the type species. However, without the holotype at hand for
comparison, this identification is not without question.
Maurice formation; Butte north of Little Rocky Creek, Beartooth
Mountains, Wyoming.
Holotype——Columbia Univ. No. 12612; plesiotypes, U.S.N.M. No.
108708.
ARAPAHOIA STANTONI, new species
PLATE 7, FIGURES 2-4
This species is similar to A. spatulata Miller in that the glabella is
not separated by relief from the fixigenes. The very shallow dorsal
furrow can be seen very clearly in cross light. The keel shows in
about the same degree as the glabella. Exclusive of the neck spine,
the holotype canidium, measured immediately in front of the eyes, is
13.7 mm. long and 8.8 mm. wide. The anterior margin of the brim
is slightly angulated. Few specimens retain the slender neck spine.
The associated pygidium has two pleural furrows clearly defined.
Pilgrim formation ; (loc. 26b) Mill Creek, Little Rocky Mountains,
Montana.
Holotype-—U.S.N.M. No. 108699a; paratype, No. 108699b.
ARAPAHOIA REESIDEI, new species
PLATE 7, FIGURES 5-9
This species is represented by many specimens, but only a few can
be cleaned reasonably well. A. reesidei, named for the collector, Dr.
John B. Reeside, Jr., is nearly flat in cross section but has more con-
vexity longitudinally. The brim is turned down rather sharply at the
margin. The dorsal furrow is expressed by shallow furrows on the
sides as far forward as the eyes; beyond that point there is only a
faint indication of it. The eyes are in an elevated position. The
anterior facial suture diverges slightly.
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 45
The libragenes have short slender genal spines.
Maurice formation ; (loc. 26c) T. 30 N., R. 96 W., 6 miles south of
Hailey, Wyoming.
Holotype -—U.S.N.M. No. 108700a; paratypes, Nos. 108700b, c.
ARAPAHOIA POLITA, new species
PLATE 7, FIGURES I0, II
The cranidium, exclusive of the large occipital spine and postero-
lateral limbs, forms approximately a rectangle. A weakly defined
dorsal furrow showing on the upper surface, probably is strongly
indicated in exfoliated specimens. In cross section the cranidium is
flatly arched but longitudinally the front third turns down rapidly.
The eyes are of moderate size, situated at about the midpoint of the
cranidium. Facial suture diverges slightly in front of the eye and
rapidly behind the eye to form large posterolateral limbs. A broad,
shallow furrow separates the greatly thickened neck ring which ex-
tends into a long, tapering, up-curved spine, the sides of which are
strongly striated. No other parts have been assigned to the species.
Sullivan formation; (loc. 64h) East Lyell Glacier, 48 miles north-
west of Lake Louise, Alberta.
Holotype.—U.S.N.M. No. 108701.
ARAPAHOIA ELONGATA, new species
PLATE 7, FIGURES I5-19, 32
This is a common species of the group found in this abundant
fauna. It is characterized by a relatively narrow, long, tapering
glabella which in exfoliated specimens is outlined by a shallow dorsal
furrow. Fixigenes narrow. Eyes small. Facial suture diverges in
front of eyes, then becomes intermarginal for a considerable distance,
rounding the anterior angles. In cross section the head is flat, and
longitudinally it has about the same flat curvature, except at the front
edge, where it is rolled under slightly. The neck spine evidently was
long. The libragene assigned to this species is rounded in contour on
all edges, and in exfoliated specimens shows faint lines radiating from
the eye. The pygidium assigned to the species is smooth and evidently
_ has a punctate surface, but lacks a well-differentiated border.
Sullivan formation; (loc. 64h) East Lyell Glacier, 48 miles north-
west of Lake Louise, Alberta.
Holotype —U.S.N.M. No. 108703a; paratypes, Nos. 108703b, c.
4
46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
ARAPAHOIA ALBERTENSIS, new species
PLATE 7, FIGURES 27-31
This is a very abundant species and is normal for the genus. Its
outstanding characteristic is the enormous posterolateral limbs, which
have pushed the rear facial suture forward nearly to the front margin
of the glabella. In exfoliated specimens the glabella is clearly marked
by four pairs of glabellar furrows. The shallow occipital furrow
marks off a triangular neck ring extending into a long occipital spine.
A tubercle near its base seems to be a median eye. Anterior facial
suture diverges very little and evidently is intermarginal nearly to
the center. The libragene assigned to the species is short and stout,
to fit with the contour ef the facial suture. The pygidium assigned to
the species is also short and has a well-defined border.
Sullivan formation; (loc. 64h) East Lyell Glacier, 48 miles north-
west of Lake Louise, Alberta.
Holotype —U.S.N.M. No. 108705a; paratypes, Nos. 108705p, e.
ARAPAHOIA PROLIXA, new species
PLATE 7, FIGURES 20-26
This large species is represented by a number of specimens. In
length it resembles .4. elongata. On exfoliated specimens the glabella
is clearly defined and shows faint traces of glabellar furrows. The
outer test has a keel. At the anterior end of the eye the fixigenes are
narrow, measuring a little more than the width of the dorsal furrow.
Anterior to the eyes the suture diverges, but less than in A. elongata.
In cross section the species has low convexity, attained by an even
curvature. Longitudinally it has greater convexity but this is also
attained with even curvature. Concentric lines appear just behind the
midpoint of the cranidium and possibly surround an eye. On the other
hand, the brim is characterized by vertical anastomosing lines, which
are stronger on exfoliated specimens. Unfortunately no examples are
complete but a rather wide libragene seems to represent the species.
The pygidium assigned to the species is convex, with a fairly well-
defined border. It has definite pleural furrows and grooves.
Sullivan formation; (loc. 64h) East Lyell Glacier, 48 miles north-
west of Lake Louise, Alberta.
Holotype —U.S.N.M. No. 108704a; paratypes, Nos. 108704b-d.
ARAPAHOIA WALCOTTAE, new species
PLATE 7, FIGURES 12-14
This is a short form similar to A. albertensis. In cranidia that
retain the test the outline of the glabella is indicated by a shallow
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 47
dorsal furrow. Exfoliated specimens have the glabella much more
sharply defined, and glabellar furrows are represented by pits. The
eyes are slightly farther back than in A. albertensis. The suture
diverges in front of the eye and forms large posterolateral limbs
behind the eye. The middle portion of the brim projects somewhat.
In cross section the cranidium has considerable elevation, attained by
sharp curvature of the top of the glabella from which the postero-
lateral limbs bend downward with approximately the same slope.
Longitudinally the species is not greatly curved.
Sullivan formation; (loc. 64e) East Lyell Glacier, 48 miles north-
west of Lake Louise, Alberta.
Holotype-—U.S.N.M. No. 108702a; paratype, No. 108702b.
PLETHOMETOPUS Ulrich, 1930
PLETHOMETOPUS ALBERTENSIS, new species
PLATE 12, FIGURE 13
A single cranidium, typical of the genus in all respects is sufficiently
well preserved to warrant naming. The simple large glabella, indi-
cated by a faint dorsal furrow, is about three-fourths the length of
the head, and tapers slightly to a rounded anterior margin. A strong
occipital furrow separates a neck ring that extends into a short blunt
spine. Fixigenes narrow, measuring at the eyes about a fifth the
glabellar width. The posterolateral limbs are small and possibly short.
Eyes small, situated about the midpoint of the glabella. In cross
section the cranidium is rather evenly curved, while longitudinally
the curvature is much greater, particularly in the anterior half. As
a result the simple brim is convex in both directions. A faint broad
anterior furrow is visible in the anterior angles.
Lyell formation; (loc. 20j) Tilted Mountain Brook, 94 miles east
of Lake Louise, Alberta.
Holotype —vU.S.N.M. No. 108755.
KINGSTONIA Walcott, 1924
KINGSTONIA ELECTRA, new species
PLATE 7, FIGURES 33-36
This species is represented by a number of cranidia, but thus far
no pygidium which can be assigned to the species has been located.
On the outer surface the cranidium shows few features. It is highly
arched in both directions, becoming almost hemispherical. Indeed its
outline departs slightly from a semicircle only by a flattening of the
curvature at about the midpoint of the cranidial length. A narrow
brim is present. 'Exfoliated specimens show a clearly defined glabella,
48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
which extends to the anterior margin of the cranidium. A narrow
occipital ring is indicated.
Weeks formation; (locs. 30h, 1) 5 miles south of Marjum Pass,
House Range, Utah.
Holotype —U.S.N.M. No. 108706a ; paratypes, No. 108707.
KINGSTONIA MUCRO, new species
Piate 8, Figures 1-8
This rather large species is represented by some 20 specimens. It
is characterized by enormous posterolateral limbs, which give the
cranidium great width and cause it to depart from the more nearly
semicircular outline of most species. The glabella is faintly indicated,
more particularly in the rear where shallow furrows extend forward
for some distance. There is a narrow rim of the usual type. The
eyes are of normal size and situated somewhat in front of the mid-
point. In cross section the head is strongly arched. The posterolateral
limbs continue the downward slope, with increasing curvature near
their distal ends. Longitudinally the cranidium is highly arched. The
associated pygidium, when viewed from the dorsal surface, has a
rounded triangular shape. In profile it is very convex. At the anterior
angles the sides stand vertical, but posteriorly the curvature increases
until at the rear of the axis the pygidium is folded under. The axis
is faintly outlined and in exfoliated specimens shows six rings.
Pleural grooves are visible.
Sherbrooke limestone; (loc. 58f) ridge west of Mount Bosworth,
British Columbia.
Holotype-—U.S.N.M. No. 108709a; paratypes, Nos. 108709a-d.
KINGSTONIA BOSWORTHENSIS, new species
PLATE 8, FIGURES 9-13
This species has normal size and shape. The cranidium is quite
globular with only short posterolateral limbs, and narrow brim around
the front. In cross section the cranidium stands very high with its
greatest curvature toward the rear. Longitudinally it is curved rather
evenly throughout, with a slight flattening in the front fourth. The
associated pygidium is short and very convex.
Sherbrooke limestone; (loc. 57p) ridge west of Mount Bosworth,
3ritish Columbia.
Holotype —U.S.N.M. No. 108710a; paratypes, Nos. 108710b, c.
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 49
KINGSTONIA ROBSONENSIS, new species
PLateE 8, FIGuRES 14, I5
A single cranidium, fortunately rather well preserved, was found
in the collection, showing that the Sullivan formation should be recog-
nized in the Robson District. This cranidium is of average size and
shape. The glabella is defined by a complete dorsal furrow, which
shows as a darker line through the test but is not expressed as a
furrow. A narrow striated brim is present and the posterolateral
limbs are wide and bluntly rounded at the ends.
Sullivan formation ; (loc. 61r) Moose River, to miles northeast of
Robson Peak, British Columbia.
Holotype —vU.S.N.M. No. 108711.
KINGSTONIA SULLIVANENSIS, new species
PLate 8, FIGURES 16-19
This globular species with rather large posterolateral limbs has a
faintly defined glabella. A distinctly thickened and striated rim char-
acterizes the holotype. The posterolateral limbs are rounded off
abruptly.
Sullivan formation; (loc. 64m) Sullivan Peak, 48 miles northwest
of Lake Louise, Alberta.
Holotype —U.S.N.M. No. 108712a; paratypes, No. 108712b.
KINGSTONIA PROMISSA, new species
PLATE 8, FIGURES 20-23
Numerous cranidia and a few pygidia represent this species, which
is slightly smaller than average. It is a short form with relatively
wide fixigenes and large posterolateral limbs. The glabella is outlined
only in the rear. In cross section the cranidium is very convex; the
convexity being attained by the steeply inclined posterolateral limbs
and a very sharp bending at the top of the glabella. Longitudinally
the convexity is that of a quarter sphere. The pygidium is very
plump, swelling greatly toward the rim.
Sullivan formation; (locs. 64c, j) Glacier Lake Canyon Valley,
48 miles northwest of Lake Louise, Alberta.
Holotype —U.S.N.M. No. 1087134; paratype, No. 108713b.
50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
KINGSTONIA VULGATA, new species
PLATE 8, FIGURES 26-30
This species is similar to K. promissa, differing in being less convex
in every way, and having wider posterolateral limbs. It has normal
convexity in both directions, in both the head and tail.
Sullivan formation; (loc. 64b) Glacier Lake Canyon Valley, 48
miles northwest of Lake Louise, Alberta.
Holotype —U.S.N.M. No. 108715a; paratypes, Nos. 108715b, c.
KINGSTONIA LOPERI, new species
PLATE 8, FIGURES 24, 25
This species is represented by pygidia from several localities, but
a sufficiently well-preserved cranidium to permit unquestioned deter-
mination has not been found. These fossils occur in very hard arkosic
sandstone and quartzite; consequently not much can be done in the
way of preparation. The illustrated specimen shows the quartz grains
scattered through the siliceous matrix. Owing to its great length, this
pygidium agrees rather well with that of Bynumua, but until positive
evidence is available, it is placed in Kingstonia, the genus blonging to
the horizon of the Sawatch fauna.
This pygidium has an elongate triangular shape, with a long, narrow
axis clearly defined. Some of the exfoliated specimens, when
weathered in a certain way, show the axial rings and the pleural
furrows. On the exterior, however, only the anterior furrow and a
faintly defined brim are visible. Further segmentation is shown but
very faintly by reflection through the test.
Sawatch formation; (loc. 6a) Taylor Peak, 4 miles south of Ash-
croft; (loc. 6) Italian Mountain, 15 miles northeast of Crested Butte,
Colorado.
Holotype —U.S.N.M. No. 108714.
KINGSTONIA (?) PLENA, new species
PLATE 8, FIGURES 31, 32
A single pygidium represents this large and tumid species. Since
the pygidium is exfoliated the lateral portions of the dorsal furrow
are fairly deep, but it is not traceable around the rear of the axis.
The axis rises slightly above the pleural lobes, which slope gently
from the dorsal furrow for about half their width, then turn down
very rapidly. At the rear the entire pygidium has such great convexity
NOES UPPER CAMBRIAN TRILOBITES—RESSER 51
that the margin is turned under. At the rear the test is irregularly
striated.
Sullivan formation ; (loc. 64h) East Lyell Glacier, 48 miles north-
west of Lake Louise, Alberta. ,
Holotype-—U.S.N.M. No. 108716.
KINGSTONIA ARA (Walcott)
PLATE 8, FicuRES 42-46
Ucebia ara Watcott, Smithsonian Misc. Coll., vol. 75, No. 2, p. 60, pl. 14,
fig, 1024; ibid., No: 3) p. 118) pl. 17, figs. 7, 8, 102s.
Kingstonia ara REesseEr, ibid., vol. 95, No. 4, p. 24, 1936.
In addition to the holotype and paratype cranidia that are refigured,
another specimen from the type locality is added to show the changed
aspect resulting from exfoliation. It will be remembered that the
genus Ucebia was erected on this species, because the glabella was
well defined. However, as the illustrations show both exfoliated
specimens and one cranidium retaining part of the test, it is clear
that Ucebia is not a valid genus. When the shell is removed, any
species of Kingstonia becomes Ucebia.
The locality erroneously stated in the original description, is given
correctly below. |
‘Warrior limestone; (loc. 24f) 1 mile southeast of Warriors Mark,
Pennsylvania.
Holotype —U.S.N.M. No. 70257; paratype, No. 70258; plesiotype,
108720.
KINGSTONIA KINDLEI, new species
PLATE 9, FIGURES I-4
This species is named for Dr. Cecil Kindle, who collected the
cranidium and several pygidia from the belt of Warrior limestone
exposed in the southwestern portion of the Bellefonte quadrangle.
K. kindlei differs somewhat from K. ara (Walcott), which occurs
in the same limestone belt to the southwest in the Tyrone quadrangle.
It averages larger and is more convex than K. ara. Also the anterior
outline is more circular. The holotype cranidium retains the test, and
consequently the dorsal furrow can be seen only when the specimen
is held so that strong cross light falls parallel to the slope of the
glabella and fixigene. A narrow striated rim is set in a nearly vertical
position. In cross section this cranidium is very high with a nearly
even curvature. When viewed from the side, the crandium appears
to be a quarter sphere. The associated pygidium has a nearly cylin-
bo
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
on
drical axis, which extends almost to the rear margin. Posteriorly it
coalesces with the pleural lobes, owing to shallowing of the dorsal
furrow.
Warrior limestone; (loc. 38e) 1 mile northwest of Benore, and
5 miles west of State College, Pennsylvania.
Holotype —U.S.N.M. No. 108721a; paratypes, Nos. 108721b, c.
BYNUMIA Walcott, 1924
Bynumia Wa.cort, Smithsonian Misc. Coll., vol. 75, No. 2, p. 54, 1924; ibid.,
No. 3, p. 78, 1025.
The additional species here described add proof that Bynumia is
a valid genus. Examples of the libragene and pygidium are also added
to the genus. The illustrations show that the glabella of Bynumia,
like that of AKingstonia, is clearly marked in exfoliated cranidia, but
in unexfoliated examples it is seldom indicated by more than a sug-
gestion of the dorsal furrow.
Genotype.—B. eumus Walcott (restricted).
BYNUMIA EUMUS Walcott
PLATE 9, FIGURES 5-7
Bynumia eumus Watcott, Smithsonian Misc. Coll., vol. 75, No. 2, p. 54,
pl. 10, fig. 2, 1924; ibid:, (part) No. 3, p. 78; pl. 17, fig. 4, 1925.
A pygidium is figured in addition to the holotype cranidium. Be-
cause the number, poorly written on the type cranidium, was misread,
its locality was erroneously given in the Sawback Range.
Sullivan formation; (loc. 64b) head of Glacier Lake Canyon
Valley, 48 miles northwest of Lake Louise, Alberta.
Holotype —U.S.N.M. No. 70255; plesiotype, No. 108722.
BYNUMIA ELEGANS, new species
PLATE 9, Ficures 8-12
This species, represented by a large number of specimens, is close
to B. eumus, and hence typical of the genus in all respects. In B.
elegans the glabella is rather slender and more truncate than usual.
Its distinguishing characteristics, as in other species, reside chiefly in
the proportions of the brim. The brim width is equal to half the
glabellar length and comes to a sharper point than in Bb. eumus.
Another difference from that species is found in the occipital furrow,
which extends on to the posterolateral limbs. These limbs are further
NO ey UPPER CAMBRIAN TRILOBITES—-RESSER 53
distinguished by their convexity. The associated pygidium is similar
to that of B. eumus, differing chiefly in somewhat greater relief of
the axis and the portions near the margin, particularly toward the
rear.
Sullivan formation; (loc. 64r) Ranger Brook Canyon, Sawback
Range, Alberta.
Holotype —U.S.N.M. No. 108723a; paratypes, Nos. 108723b-d.
BYNUMIA ARGUTA, new species
PLATE 9, FIGURES 13-15
This species is represented by several examples. Its fairly wide
glabella is square in front and rounded at the anterior angles. As a
whole the outline is nearly that of an equilateral triangle. A shallow
dorsal furrow, visible on the exterior, separates the brim from the
glabella. The brim is considerably swollen in front. From the front
the cranidial cross section is little curved, except at the outer edges.
Since the posterolateral limbs are greatly depressed in the rear, the
cranidium is highly arched. Longitudinally the curvature is moderate,
except at the very front, where the swollen rim drops steeply. The
associated pygidium is flat and so fused that the axis is scarcely
traceable.
Sullivan formation; (loc. 66s) Badger Pass, Sawback Range,
Alberta.
Holotype—U.S.N.M. No. 108724a; paratype, No. 108724b.
BYNUMIA WALCOTTI, new species
PLATE 9, FIGURES 16-20
Bynumia ewmus Watcorr (part), Smithsonian Misc. Coll., vol. 75, No. 3,
plet7, figs) 5,6, 1925.
This species was included by Walcott in B. ewmus, which it re-
sembles. It is abundantly represented at several localities by many
specimens. Compared with B. ewmus, B. walcotti appears to be a
stockier form, particularly when not exfoliated. This appearance is
due to the more convex and slightly narrower brim. The pygidium
assigned to the species is flat, with the poorly defined axis raised
slightly above the flat pleural lobes.
Sullivan formation; (loc. 66m) 5 miles northwest of Banff, and
(loc. 66s) Badger Pass, Sawback Range, Alberta.
Holotype—U.S.N.M. No. 108731a; paratypes, Nos. 70256,
10873 1b.
54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
BYNUMIA VENUSTA, new species
PLATE 9, FicureEs 21-28
This abundantly represented species is characterized chiefly by its
narrow cranidial shape, which forms an isosceles triangle. Longi-
tudinally the curvature is not great, and the relatively wide brim
continues the gradual slope of the anterior portion of the glabella.
The libragene figured, the first for the genus, constitutes almost a
quarter circle, the outer margin increasing its rate of curvature toward
the genal angle. Several pygidia are figured, one with a complete test
and two partially exfoliated examples. The latter show how the axial
rings and pleural furrows and grooves are developed on the under
side of the test.
Sullivan formation; (loc. 64s) Ranger Brook Canyon, Sawback
Range, Alberta.
Holotype —U.S.N.M. No. 108725a; paratypes, Nos. 108725b-f.
BYNUMIA ROBSONENSIS, new species
PLATE 9, FIGURES 30-34
One species of Bynumia occurs in the Robson District, far north
of the localities of the other species. This species is characterized by
its triangular outline, with the base being longer than the sides. The
photographs fail to bring out the sharpness of the brim, owing to the
high convexity developed near the tip.
Lynx formation (Sullivan equivalent); (locs. 19m, L) Iyatunga
Mountain, Mount Robson, British Columbia.
Holotype-—U.S.N.M. No. 108726a; paratypes, Nos. 108726b, c.
BYNUMIA SULCATA, new species
PLATE 9, FIGURES 35, 36
A single small distinctive cranidium has the usual triangular outline
and average shape of other species in the Sawback Range. Because
of the depth of the dorsal furrow, B. sulcata is closest to B. arguta,
but carries this development much further. Not only is the dorsal
furrow deeply impressed both on the sides and in front, but in addi-
tion, the brim, set in a nearly horizontal position, does not continue
the glabellar slope. This is a remarkable development in a specimen
that is not completely exfoliated. The side view shows the distribution
of convexity.
Sullivan formation; (loc. 64s) Ranger Brook Canyon, Sawback
Range, Alberta.
Holotype —U.S.N.M. No. 108727.
NO. 5 UPPER CAMBRIAN TRILOBITES—-RESSER 55
BYNUMIA RANGERENSIS, new species
PLATE 9, FIGURES 37-41
This well-represented species is allied to B. ewimus in size and pro-
portions of the glabella. In cross section the brim, and to a lesser
degree the glabella and posterolateral limbs, are highly convex. Longi-
tudinally the convexity is not great, except in the swollen brim. The
brim width is less than half the glabellar length. The rather large
associated pygidium is nearly flat in cross section, but in the opposite
direction has increasing curvature in the posterior moiety.
Sullivan formation; (loc. 66L) Ranger Brook Canyon, Sawback
Range, Alberta. ’
Holotype —U.S.N.M. No. 108728a; paratypes, Nos. 108728b, c.
BYNUMIA SAWBACKENSIS, new species
PLATE 9, FIGURES 42-44
This large, well-represented species is characterized particularly by
its wide brim, which exceeds half the glabellar length. Not only is
the britn wide, but when viewed from the front comes to a blunt,
nearly cylindrical point. The pygidium has a blunt rear margin,
reached by the faintly outlined axis.
Sullivan formation; (loc. 64w) Ranger Brook Canyon, Sawback
Range, Alberta.
Holotype —U.S.N.M. No. 108729a; paratype, No. 108729b.
BYNUMIA (?) MODESTA, new species
PLATE 8, FIGURES 33, 34
This species lies between Bynumia and Kingstonia and should
possibly be referred to the latter genus.
The glabella is large and relatively wide, so that it occupies by far
the greater portion of the cranidium. The front outline is strongly
curved but does not come to such a sharp point as in other species.
Longitudinally the cranidium is convex with an even curvature. It is
also convex laterally, but this convexity is attained by a sharp bend
along the median line causing the glabella and posterolateral limbs,
as well as the palpebral lobes, to slope down sharply on each side. The
steeply inclined halves of the cranidium are only a little curved in
themselves. The eyes are small and are situated well forward. A
narrow rim is visible in front.
Sullivan formation; (loc. 64h) Glacier Lake Canyon Valley, 48
miles northwest of Lake Louise, Alberta.
Holotype —U.S.N.M. No. 108717.
50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
BYNUMIA (?) PRODUCTA, new species
PuaTe 8, FIGURES 35, 36
Although this species is represented by many specimens, only one |
cranidium is figured. In its relatively flat anterior outline and some-
what upturned, thickened rim, and in the lack of divergence of the
anterior facial suture, this species resembles Blowntia. However, since
the fixigenes are almost triangular in shape, it is placed in Bynumua.
Owing to the truncate anterior outline, the glabella is more quadrate
than usual. The very shallow dorsal furrow converges slightly for-
ward and rounds off the anterior angles. A shallow occipital furrow
outlines a narrow ring. A deep depression crosses the preglabellar
area in front of the glabella, and the rim is thickened, so that it is
rather prominent in side view.
Numerous pygidia are in hand, but because the specific association
is rather uncertain they have not been definitely assigned to the
species. These pygidia are characterized by the usual triangular shape,
axial and other features, differing from B. mollis chiefly in the abrupt
slopes along the outer margin, which are so steep that the edges are
rolled over.
Sullivan formation; (loc. 641) Glacier Lake Canyon Valley, 48
miles northwest of Lake Louise, Alberta.
Holotype —U.S.N.M. No. 108718.
BYNUMIA MOLLIS, new species
PLATE 8, FIGURES 37-41
A few specimens characterized by a pronounced triangular shape
represent this species. The glabella is faintly outlined on the sides
by the dorsal furrow and in front by a change in slope. The thickened
rim is slightly upturned. Viewed from the rear, the glabella stands
above the posterolateral limbs, owing to deep notching of the posterior
margin at the ends of the dorsal furrow. Viewed from the front, the
lateral curvature is rather even, with the greatest amount along the
median line. Longitudinally the nearly even curvature of the glabella
is interrupted by the more horizontal position of the brim. The
pygidial axis is long, slender, and tapers gradually, reaching nearly
to the rear margin. A faint border is visible, and on exfoliated
specimens the usual segmentation is clearly defined both in the axis
and the pleural lobes.
Sullivan formation ; (loc. 64h) East Lyell Glacier, 48 miles north-
west of Lake Louise, Alberta.
Holotype —U.S.N.M. No. 108719a; paratypes, Nos. 108719b, c.
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 57
BYNUMIELLA, new genus
It appears that the Bynumia stock continued into Franconia time,
taking on a somewhat changed form. The alterations in structure
may be summed up by stating that they are modifications of Bynumia
in the direction of average trilobite structure. For instance, the brim
structure is retained, but with modification in the direction of reduction
to the more normal band form. Likewise the glabella develops con-
siderable taper, the dorsal furrow becomes deeper and an occipital
furrow separates a neck ring, which is expanded backward in the
middle. Only cranidia are known.
Diagnosis.—Small trilobites with a tapering glabella without fur-
rows. Dorsal and occipital furrows are well defined. The simple
brim varies in width, but tends to extend forward in the middle,
causing the anterior outline of the cranidium to project.
Genotype.—B. typicalis, new species.
BYNUMIELLA TYPICALIS, new species
PLATE 10, FIGURES I, 2
Several small cranidia have been located in the collection, charac-
terized by a glabella which tapers to a rounded point, and which is
demarcated by the well-defined dorsal furrow. An equally deep occip-
ital furrow separates a swollen neck ring. The fixigenes average
about the same width as the brim, and the latter has a width equal to
one-third the glabellar length. The eyes are small, situated well
forward.
Lyell formation; (loc. 66j) Northeast branch Ranger Brook
Canyon, 10 miles northwest of Banff, Sawback Range, Alberta.
Holotype-—U.S.N.M. No. 108732a; paratype No. 108732b.
BYNUMIELLA BRISCOENSIS, new species
PLATE 9, FIGURE 20
Several cranidia from the Sabine formation represent Bynumiella.
In the holotype cranidium the glabella tapers to a rounded front less
sharp than in B. typicalis. The brim width is somewhat less than
one-third the glabellar length. Owing to depression of the anterior
angles, the brim is convex. The cranidium is moderately arched in
both directions.
Sabine formation ; (loc. 16t) Sinclair Canyon, Brisco Range, British
Columbia.
Holotype —U.S.N.M. No. 108730.
58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
BYNUMIELLA (?) OKLAHOMENSIS, new species
PLATE I0, FIGURES 27, 28
Pending the availability of better material several .cranidia that are
not fully typical are referred to the genus. This species differs from
typical forms in that the glabella tapers less rapidly and in the size
and position of the eyes, which are far larger and situated much
farther back than they should be for a Bynumiella. Brim, occipital
ring, and fixigenes are all typical.
The illustrations show the characteristics of the holotype cranidium.
Faint glabellar furrows are present. A thickening of the neck ring
produces a short blunt spine. Longitudinal curvature is shown in
figure 27, and that in the opposite direction is about the same.
Signal Mountain formation; (loc. 2013) I mile south of Royer
Ranch, Arbuckle Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108743.
BYNUMINA, new genus
A second genus Bynumina is erected for the Bynumia stock that
continued into Franconia time. As in Bynumiella this genus represents
development of Bynumia in the direction of more average trilobite
structure. Although glabellar taper was an essential change in this
instance, the head as a whole is less constricted anteriorly. Glabellar
furrows also appear, but the most pronounced change lies in the brim
structure, which brings that part more nearly to form a transverse
band.
Diagnosis —Small trilobites characterized by large, somewhat ta-
pered glabella. Dorsal and occipital furrows well defined. Glabellar
furrows faint. Neck ring narrow and of even width throughout. Eyes
small, situated well forward. Brim simple, of nearly even width
throughout. Faint eye lines present. Pygidium simple; axis long,
tapered, poorly defined, and with faint axial rings. Pleural lobes
completely fused.
Genotype.—B. caclata, new species.
BYNUMINA CAELATA, new species
PLATE 10, FicurEs 18-22
Numerous cranidia and one pygidium represent this species. The
illustrations show both the generic and specific characteristics.
The glabella tapers gradually to a truncated front margin. The
anterior corners are rounded, and along the sides the dorsal furrow
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 59
bows outward slightly. Several pairs of recurved glabellar furrows
are visible. The occipital furrow, both on the glabella and fixigenes,
is deeply impressed. In cross section the cranidium is convex, reach-
ing the greatest curvature as the median line is approached. Longi-
tudinally the convexity is less, and the rate of curvature even through-
out. The brim width is a little more than one-fourth the glabellar
length, and is a simple convex band surrounding the front of the
head. The associated pygidium is also convex, with a broad, faintly
outlined axis and fused pleural lobes.
Davis formation ; (loc. g2d) Federal Lead Mine No. 4, Flat River,
Missouri.
Holotype-—U.S.N.M. No. 108740a; paratypes, Nos. 108740b-d.
BYNUMINA MISSOURIENSIS, new species
PLATE 10, FIGURES 23-26
This is the more common Missouri species, and it differs from
B. caelata mainly in its narrower rim.
B. missouriensis has a tapering glabella, but the curvature of the
lateral dorsal furrows is not even. It expands rapidly as far forward
as the first glabellar furrow, then suddenly begins to contract, main-
taining a nearly straight course until it approaches the anterior angles,
where the rate of taper becomes less again. Besides this peculiarity,
the species is characterized by a rim width of a little more than one-
fifth the glabellar length.
Davis formation; (locs. 11k, 91y) Flat River, Missouri.
Holotype —U.S.N.M. No. 108741 ; paratypes, Nos. 1087422, b.
STENOPILUS Raymond, 1924
STENOPILUS BACCA, new species
PLATE 10, Ficures 8-13
Such a featureless hemispherical trilobite as Stenopilus is difficult
to describe. Several species are here presented, chiefly to show the
differences between small and large examples, and the variation in
degree of sphericity. There seems to be little doubt but that these
trilobites developed from a Kingstonia ancestor.
Large cranidia of S. bacca are nearly hemispherical in shape, but,
as may be seen in figures 10 and .11, small heads show a distinct
Kingstomia shape. Of course, the indentations of the dorsal furrow
in the rear remain, but unless the very convex specimen is specially
posed, they are obscured by the bulging cranidium.
60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Signal Mountain formation; (locs. 121, j) 2 miles southwest of
Signal Mountain, Wichita Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108737; paratypes Nos. 108738a-c.
STENOPILUS ADUNCUS, new species
PLATE 10, FIGURES 14-17
A second species is chosen for description to show the elongate
form taken by species of Stenopilus. This development recalls the
Bynumuia elongation of the Kingstoma stock, but in Stenopilus this
attends obesity and elimination of all furrows. As a result a smooth
trilobite is developed, so convex that the glabella overhangs the
posterior margin. This high convexity is reduced forward, but the
curvature is not eliminated in any portion of the test.
S. aduncus is longer than wide and has its greatest convexity at
the rear. Otherwise it is practically featureless.
Signal Mountain formation; (loc. 12L) Pickens Ranch, Arbuckle
Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108739a; paratype, No. 108739b.
BLOUNTIA Walcott, 1916
BLOUNTIA PLANA, new species
PLATE 14, FIGURES 12-14
Only pygidia have been assigned to the species. While the long
axis is defined, and the usual interrupted axial furrows are visible,
it does not stand above the pleural lobes. The chief characteristic of
the species is found in the flatness of the pygidium. When viewed
from the rear, the pleural platforms, with the axis, form only a
slightly convex shield. The border is slightly concave, thereby in-
creasing the curvature near the lateral margins.
Sullivan formation; (loc. 64c) Glacier Lake Canyon Valley, 48
miles northwest of Lake Louise, Alberta.
Holotype—U.S.N.M. No. 108779a; paratype, No. 108799b.
BLOUNTIA KINDLEI, new species
PLATE 14, FIGURES 27-31
Two small pygidia and a portion of the cranidium are figured. A
somewhat broken pygidia is included to show appearance of the test ;
one much larger pygidium has not been figured. This species, which
is named in honor of the collector, Dr. Cecil Kindle, is typical of the
genus in every respect.
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 61
The large glabella tapers at a normal rate to the rounded front. It
lacks furrows and even the occipital furrow is very faint. The fixi-
genes average less than half the glabellar width, expanding to the
moderate anterior angles, and behind the eye into stout posterolateral
limbs.. Eyes are small, situated about the midpoint of the cranidium.
The rather heavy, somewhat upturned rim is about the same width as
the preglabellar area. When exfoliated, the pygidium shows a long
axis on which at least six rings are faintly indicated. As a whole, the
pygidium is triangular in outline. In cross section the curvature is
gentle, except near the margins where the slope increases so much
that the outer edges are slightly turned under.
Warrior limestone; (loc. 38e) 1 mile northwest of Benore, 5 miles
west of State College, Pennsylvania.
Holotype.—U.S.N.M. No. 108788a ; paratypes, Nos. 108788b, c.
BLOUNTIA (?) DISPARILIS, new species
PLATE 14, FIGURES I0, II
This narrow cranidium is not typical of Blountia. The glabella
tapers to a rounded anterior outtine and lacks glabellar furrows. A
shallow occipital furrow separates a narrow neck ring. The fixigene
is narrow, averaging about one-third the average glabellar width. The
wide brim is subdivided into a preglabellar area and a wider rim. The
rim is thickened and increases in the middle to about twice the width
of the preglabellar area. Eyes small. Longitudinally the cranidium
attains considerable convexity by rather strong curvature in the rear
portion. In cross section the curvature, including the posterolateral
limbs, is more even.
Sullivan formation; (loc. 64h) Glacier Lake Canyon Valley, 48
miles northwest of Lake Louise, Alberta.
Holotype —U. S. N.M. No. 108775.
BLOUNTIELLA Resser, 1938
BLOUNTIELLA ALBERTA, new species
PLATE 14, FIGURES 19-26
Although this species is represented by numerous cranidia, the
pygidium has not been located. Taken as a whole, the cranidium is
rather short, and the glabella occupies most of its cranidial area.
Glabellar furrows are lacking even in exfoliated specimens. The fixi-
genes average less than half the glabellar width. The eyes are small,
situated about the midpoint of the cranidium. A narrow preglabellar
5
62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
area separates the thickened, striated rim from the dorsal furrow.
Laterally the head is moderately and evenly convex, but longitudinally
the curvature is much greater. A shallow occipital furrow separates
a narrow ring.
Sullivan formation; (locs. 64c, b) Glacier Lake Canyon Valley,
48 miles northwest of Lake Louise, Alberta. :
Holotype-—U.S.N.M. No. 108776a; paratypes, Nos. 108776b,
1087772, b.
ELLIPSOCEPHALOIDES Kobayashi, 1935
Ellipsocephaloides Kopayasut, Journ. Fac. Sci. Imp. Univ. Tokyo, sect. 2, '
vol. 4, pt. 2, p. 196, 1935. t
This generic name was assigned without proper study of the
specimens, for it perpetuates the false idea of relationship between
these trilobites and Ellipsocephalus. The species of Ellipsocepha-
loides, moreover, are unique in several respects, and could not belong
to the I/laenuxidae, which should have been apparent even before the
pygidium was known. Under present circumstances the best pro-
cedure is to place the genus in a new family, leaving the determination
of the family’s position in trilobite classification to future study.
Considerable variation in width of the anterior fixigene and the
more advanced course of the facial suture due to a more forward
position of the eye may be observed among the species here described.
Though these variations fall almost entirely on one side of the geno-
type, E. curtus (Whitfield), the narrow forms and those in which
the eye is farther back are not segregated as a separate genus since
sufficient gradation seems to exist to tie all together. Ellipsocepha-
loides is apparently characteristic of Franconia horizons, for it is
associated with Idahoia, Pseudagnostus, Briscoia, and other genera
of Franconia age.
Species of Ellipsocephaloides vary considerably in shape. The
glabella is large and clearly defined, both by the dorsal furrow and
its elevation above the fixigenes. Some species have three pairs of
short glabellar furrows. Occipital furrow always developed, marking
off a neck ring, usually of even width throughout. Brim a simple
band, except in the wider species where a narrow rim is demarcated
by a shallow anterior furrow. The brim continues as the anterior
fixigene around the anterior angles and therefore curves much more
sharply in the narrow than in the wide forms. At their widest point
the fixigenes, exclusive of the eye band, vary from less than half to
more than the glabellar width at the same point. Fixigenes between
the dorsal furrow and the eye are usually nearly flat, though they may
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 63
be concave. Since the very heavy eye bands are elevated, most of the
fixigene may lie below the level both of the dorsal furrow and of the
eye. The moderate-size eyes vary considerably in position. Libragene
unknown.
Pygidium characterized by a stout axis in which two or more seg-
ments are indicated. The axis is from a half to two-thirds the length
of the pygidium and stands above the dorsal furrow. Pleural lobes
flat, extending fanwise into marginal spines. Pleural furrows strong
and pleural grooves usually visible. Marginal spines usually rather
blunt, forming a serrate edge. As the genus is now constituted they
number from one to five on a side, depending on the degree of fusion.
ELLIPSOCEPHALOIDES ARGUTUS, new species
PLATE I0, FIGURES 5, 6
Several cranidia from the Ram Creek area retain sufficient of their
characters to warrant description. The cranidium is broad. The
almost parallel-sided glabella extends nearly the full length of the
cranidium. In cross section it stands entirely above the shallow dorsal
furrow and the fixigenes which slope gently down from it. The
occipital furrow is clearly defined both on the glabella and fixigenes,
and three pairs of glabellar furrows are traceable. The fixigenes
between’ the eyes and the dorsal furrow are about as wide as the
glabella at the same point. Unfortunately the eyes are not completely
preserved, but appear to have been of normal size, shape and position.
Sabine formation; (loc. 12s) Ram Creek, 15 miles south of Canal
Flats, British Columbia.
Holotype-—U.S.N.M. No. 108735a; paratype, No. 108735b.
ELLIPSOCEPHALOIDES BRISCOENSIS, new species
PLATE I0, FIGURE 7
One of a dozen or more cranidia is figured. E. briscoensts is
narrower than E. argutus from the same region. Because of this
feature it bridges the gap between the genotype and the narrow forms.
The glabella is rectangular with rounded anterior angles and has two
or three pairs of glabellar furrows represented by elongate pits. In
width the brim measures less than a fourth the glabellar length. It
has a peculiar narrow elevation in the middle, and also has a faintly
defined narrow rim. As a whole the brim is slightly convex and is
set in a horizontal position. On the sides the brim passes into the
anterior fixigenes which maintain its level. In fact the outer margins
of the fixigenes are curved up and back to the genal angles so that
64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
the ocular platforms are depressed, leaving the eye lobes in a promi-
nent position. In this species the eyes are long, and the eye bands
wide and heavy. Faint eye lines extend straight across the fixigenes
slightly anterior to the second pair of glabellar furrows.
Sabine formation; (loc. 17s) Sabine Mountain, Brisco Range,
British Columbia.
Holotype —U.S.N.M. No. 108736.
ELLIPSOCEPHALOIDES SILVESTRIS, new species
PLATE 11, FiGuRES I-3; PLATE 12, FIGURE 7
This form, intermediate between the broad and narrow species, is
well represented in the collections. The nearly quadrate glabella is a
little longer than wide and is well rounded in front. The occipital
ring is of even width. The fixigenes are less than half the glabellar
width and maintain their width throughout. The simple brim, only
a little narrower than the fixigenes, has an elongate boss in the middle.
A narrow rim is faintly indicated for a short distance in the middle
of the head. Wide eye lines connect the wide eye lobes with the dorsal
furrow.
The associated pygidium has a short wide axis on which three or
four rings are demarcated. The pleural lobes and the rear border,
which is as wide as the axis is long, have both pleural grooves and
furrows. Both extend to the margin, producing sharp ridges. Five
spines are developed on each side.
Honey Creek limestone ; (loc. g1b) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108744a; paratypes, Nos. 108744b-d.
ELLIPSOCEPHALOIDES SAWBACKENSIS, new species
PLATE II, FIGURES 4, 5
Only the two cranidia illustrated represent this distinctive species.
The glabella is longer than wide and has three pairs of glabellar
furrows indicated by pits. The fixigenes are about half the glabellar
width and vary little from front to back. Heavy eye lines connect
the large elevated eye bands with the dorsal furrow at a point in front
of the anterior pair of glabellar furrows. The brim, about half the
width of the fixigenes, is simple in structure but is raised as a boss
in the center.
Lyell formation; (loc. 64x) Ranger Brook Canyon, Sawback
ange, Alberta.
Holotype.—U.S.N.M. No. 108745a; paratype, No. 108745b.
es
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 65
ELLIPSOCEPHALOIDES CARUS, new species
PLATE 11, Ficures 7, 8
This is a small form associated with EF. argutus.
The subcylindrical glabella is rounded in front and has three pairs
of glabellar furrows. The fixigenes are convex in contrast to most
other species; consequently the rather thin eye lines appear to be
more curved than usual. The eyes are on slightly upturned heavy
bands.
Sabine formation; (loc. 12s) Ram Creek, 15 miles south of Canal
Flats, British Columbia.
Holotype —U.S.N.M. No. 1087474; paratype, No. 108747b.
ELLIPSOCEPHALOIDES MONTIS, new species
PLATE II, FIGURES 9-II
The third species from the locality is an intermediate form, ap-
proaching the narrow rather than the broad species. Width may be
an aspect as much as a reality, since the narrowness in dorsal view
is sometimes due to the sloping position of the anterior angles, whereas
other species appear wider because the anterior flanges of the same
size are more nearly in a horizontal position. The wide glabella of
E. montis is well rounded in front, and has a slight indentation at
the center. Glabellar furrows are reduced to faint pits. The most
distinctive feature is the fusing of the broad eye line with the brim,
which slopes back to the eyes, the latter being in a posterior position.
This gives the cranidium the appearance of being made up of the
large glabella about which is draped a wide band that slopes toward
the rear. Anterior fixigenes are eliminated by this juncture of the
eye ridge and brim. Between the eyes and glabella the fixigenes have
just a little more than one-third the glabellar width. In this species
the neck ring is rather wide.
Sabine formation; (loc. 12s) Ram Creek, 15 miles south of Canal
Flats, British Columbia.
Holotype —U.S.N.M. No. 108748a; paratype, No. 108748b.
ELLIPSOCEPHALOIDES BEARENSIS, new species
PLATE II, FIGURE I2
An imperfect cranidium represents the genus in the St. Charles
formation of Idaho, showing that this element is also present in that
fauna. This species is much more like the genotype in general aspect,
but differs distinctly owing to the more posterior position of the eyes.
Glabellar furrows seem to be lacking. The eye lines are very much
66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
reduced, consisting only of a slightly elevated portion of the fixigenes.
The eye band is turned up but does not have a distinct palpebral
furrow.
St. Charles limestone ; (loc. 66z) 5 miles west of St. Charles, Bear
River Range, Idaho.
Holotype.—U.S.N.M. No. 108749.
ELLIPSOCEPHALOIDES BUTLERI, new species
PLATE II, FIGURES 14, 15 +
This species, represented by numerous cranidia, is preserved in
sandstone, and is associated with Briscoia. It has a typical develop-
ment for the genus but is not proportionally as wide as the genotype.
Glabellar furrows are present but may be lacking on the test. The
eyes are strongly bowed and situated about the middle of the
cranidium.
Sawatch formation; (loc. 37x) near Gilman, Mosquito Range,
Colorado.
Holotype-—U.S.N.M. No. 108751a; paratype, No. 108751b.
ELLIPSOCEPHALOIDES NITELA, new species
PLATE II, FIGURE 13; PLATE 12, FIGURES I-3
This narrow form, represented by many examples, most resembles
E. silvestris. Compared with that species FE. nitela is somewhat
broader and in the pygidium the pleura are wider. The nearly quad-
rate glabella has rounded anterior angles. The fixigenes are less than
half the glabellar width, and are a little wider than the brim. The
eye lines are weak and the broad eye bands are clearly defined by the
palpebral furrows. The simple brim has a slight elevation in the
middle. The associated pygidium has a broad axis on which the fur-
rows are deep. In cross section the axis stands above the pleural
lobes, which retain both the pleural grooves and furrows. The latter
bisect the pleura obliquely, decreasing in strength posteriorly. Four
marginal spines, decreasing in size from front to back, terminate the
four pleural segments.
Honey Creek limestone; (loc. 12m) 7 miles north of Springer,
Arbuckle Mountains, Oklahoma.
Holotype-—U.S.N.M. No. 108750a; paratypes, Nos. 108750b-d.
ELLIPSOCEPHALOIDES MONSENSIS, new species
PLATE 12, FIGURES 4-6
This narrow species similar to E. montis is represented by a num-
ber of cranidia and the pygidium. The quadrangular glabella is about
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 67
a third longer than wide, and has several pairs of glabellar furrows.
The brim is narrower than the neck ring, a characteristic common to
several of the Canadian species. The brim joins the eye ridges, which
also form a raised band. The rather large eyes are situated well
forward. The fixigenes, less than half the glabellar width, contract
forward to meet the relatively narrow brim.
A beautiful pygidium with a wide flaring border bears two spines
on each side. Its short, wide axis extends into a long postaxial ridge.
Only two axial rings and pleura are defined. The entire pygidium is
covered by prominent anastomosing lines.
Lyell formation; (loc. 64£) Mons Glacier, 50 miles northwest of
Lake Louise, Alberta.
Holotype —U.S.N.M. No. 108752a; paratype, No. 108752b.
ELLIPSOCEPHALOIDES DECLIVIS, new species
PLATE 12, FIGURES I0-13
This species is represented by several cranidia. It is similar to
E. monensis except for the more posterior position of the eyes. The
long, parallel-sided glabella is rounded in front, and in cross section
stands completely above the palpebral lobe. The rear pair of glabellar
furrows is continuous across the glabella and one shallow pair is
visible anterior to it. The fixigenes, measuring less than half the
glabellar width, slope down steeply from the dorsal furrow, though
at a lesser angle than the sides of the glabella. The eyes are of normal
size and are connected by a thickened eye ridge with the narrow
convex brim.
Lyell formation; (loc. 20j) Tilted Mountain Brook, 94 miles east
of Lake Louise, Alberta.
Holotype —U.S.N.M. No. 108754a; paratype, No. 108754b.
MARYVILLIA Walcott, 1916
MARYVILLIA WYOMINGENSIS (Resser)
PLATE 12, FIGURES 14-20; PLATE 14, FIGURE I
Bathyuriscus sp., Watcott, U. S. Geol. Surv. Monogr. 32, p. 466, pl. 64, fig. 6,
1899.
Coosia wyomingensis RESSER, Smithsonian Misc. Coll., vol. 95, No. 22, p. 7,
1937.
Heads and tails of a species of Maryvillia occur throughout the
Yellowstone Park region ranging from Clark Fork, Wyoming, to the
Castle Mountains, Montana. A good cranidium on the hand specimen
with the holotype of Tricrepicephalus tripunctatus (Whitfield) comes
from Moss Agate Springs in the Castle Mountains, Montana.
68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
The glabella tapers slightly to a rounded front. At the eye the
fixigenes are exactly half the glabellar width. The facial suture forms
rather broad posterolateral limbs. Eyes small and strongly bowed.
Brim concave, with a rim differentiated by a change in slope. Eye
lines traceable on exfoliated specimens.
Pygidium convex, sloping down rather sharply at the anterior
angles. Outer test smooth, and dorsal furrow shallow. On exfoliated
specimens both the pleural furrows and grooves are visible.
Pilgrim formation; Dead Indian Creek; and (loc. 151g) mouth of
Clark Fork River, Absaroka Range, Wyoming ; (loc. 4r) Suce Creek,
Mount Delano, Snowy Range; and near Moss Agate Springs, Castle
Mountains (with type of Tricrepicephalus tripunctatus), Montana.
Holotype—_U.S.N.M. No. 94343; plesiotypes, Nos. 108756, 108757,
108772.
MARYVILLIA VIOLAENSIS, new species
PLATE 12, FIGURES 21-26
About a dozen cranidia and half as many pygidia have been located
in the collections. Only two small cranidia are illustrated, as all the
larger specimens are incomplete.
Two partially preserved cranidia show that the cranidium of the
species attained a length of about 15 mm. and a width of about 25 mm.
The glabella is poorly defined because the dorsal furrow is very
shallow. Eyes, eye lines, and fixigenes are average in development.
The brim on the other hand is excessively concave, and consequently
the rim seems to be more distinct than usual. As a whole the cranid-
ium is gently convex in a transverse direction and much more sharply
convex longitudinally.
In the pygidium the axis is clearly outlined by a shallow dorsal
furrow. Axial rings and the anterior pleural furrows are faintly
indicated. The pleural lobes, nearly flat in their inner half, become
convex in their outer portion. Longitudinally the pygidium slopes
gently rearward.
Maurice formation; (loc. 151k) Lebarge Creek, near Viola, Uinta
County, Wyoming.
Holotype —U.S.N.M. No. 108758a; paratype, No. 108758b.
MARYVILLIA UTAHENSIS, new species
PLATE 12, FIGURES 27-32
Oolitic and soft granular limestone beds of the Weeks formation
have yielded species of Maryvillia. M. utahensis is represented by
NO. 5 UPPER'CAMBRIAN TRILOBITES
RESSER 69
fairly good material which includes all the various parts. In the
striation and structure of the brim, the larger size of the eye, and the
wider border of the pygidium, this species approaches Coosia. How-
ever, it seems better to place the species on the Maryvillia side of the
boundary and recognize these discrepancies.
The glabella tapers to a rounded front, and in exfoliated specimens
is clearly outlined by a deep dorsal furrow. The fixigenes, convex in
cross section, are a little less than half the glabellar width. Rather
heavy eye lines are accentuated in appearance by the abrupt slope
adjacent to them. The width of the brim is equal to half the length
of the glabella exclusive of the occipital ring. The rim is demarcated
by a change in slope. Heavy striations occur on the front edge of
the rim. Eyes rather large, being nearly as long as the brim is wide.
The libragene shows that the suture is intramarginal for a short
distance. Its rather wide and strongly striated rim increases in width
slightly from the front to the rounded genal angle. The ocular plat-
form is very narrow opposite the anterior portion of the eye. At its
widest point, at the rear of the eye, it is not much broader than the
rim. The associated hypostoma is nearly circular in outline with the
central convex portion occupying less than half of its area. The outer
upturned flange is striated in a manner similar to that of the rims on
the cranidium and libragene.
The associated pygidium is strongly convex in lateral direction
and slopes very steeply longitudinally. The axis, occupying only a
little more than half the length of the tail, is not strongly differentiated
from the pleural lobes, although it stands completely above them.
Axial rings and the anterior pleura are faintly indicated.
Weeks formation; (loc. 32w) Fandango Spring Canyon, Dugway
Range, Utah.
Holotype-—U.S.N.M. No. 108759a; paratypes, Nos. 108759b-e.
MARYVILLIA MARJUMENSIS, new species
PLATE I2, FIGURES 33-35
This species is well represented, and on several pieces of rock
cranidia and pygidia lie close together. One fairly large cranidium is
figured, but even larger ones are present in the collection. M. marjum-
ensis is fully typical of Maryvillia in that the pygidial axis extends
nearly the full length of the pygidium. As usual the glabella tapers
to a rounded front and in exfoliated specimens is clearly defined by
a rather deep dorsal furrow, which on the outer test is shallow. Fixi-
genes average just about half the glabellar width. Eyes are of normal
7O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
size. The brim width equals about half the length of the glabella,
including the occipital ring. It is gently concave with a rather wide
rim indicated by a slight change in curvature. The cranidium is
slightly convex in both directions.
The associated hypostoma is almost circular in outline, and is
striated on the wide flange which surrounds the convex but depressed
central elevation.
The plump pygidium with an axis extending nearly the full length
terminates in a postaxial ridge.
Weeks formation ; (loc. 301) 5 miles south of Marjum Pass, House
Range, Utah.
Holotype-—U.S.N.M. No. 108760a ; paratypes, Nos. 108760b-e.
MARYVILLIA LOPERI, new species
PLATE 12, FIGURES 36-38
This species is found at two localities in arkosic, calcareous, and
quartzitic sandstone. It averages rather large in size compared to
other western species. The glabella tapers forward to the rounded
front in the usual manner, and is a little longer than in the other
species here described. At the eyes the fixigenes are slightly more
than half the glabellar width. The eyes are normal in size and position.
The brim width is about one-third the glabellar length. A shallow
anterior furrow separates a slightly thickened rim. In cross section
the cranidium is moderately and evenly convex, but longitudinally it
is rather flat.
The associated pygidium is likewise rather convex in cross section,
but longitudinally proportionately has greater convexity than the
cranidium. The prominent axis occupies nearly three-fourths the
length of the pygidium. In exfoliated specimens three pleural grooves
are visible on each side.
Sawatch formation; (loc. 6a) Taylor Peak, 4 miles south of Ash-
croft; (loc. 6) Italian Mountain, 15 miles northeast of Crested Butte,
Colorado.
Holotype—U.S.N.M. No. 108761a; paratype, No. 108761b.
MARYVILLIA ALBERTA, new species
PLATE 13, FIGURES I-4
This species is founded on a single exfoliated cranidium, ‘and a
pygidium from a nearby locality is referred to it. The cranidium,
typical in all respects, has a large glabella that tapers to a rounded
frontal outline. Exfoliated specimens have a strongly developed keel.
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER ae
At the eyes the fixigenes are half the width of the glabella. The width
of the simple, gently concave brim is a little less than the glabellar
length. The eyes are slightly larger than usual. In both directions
the convexity of the cranidium amounts to little.
The pygidium from a nearby locality, assigned to the species, has
a prominent axis which tapers somewhat more than usual. The rear
border occupies about one-third the pygidial length, which with the
rather large eye, indicates the approach to Coosia.
Sullivan formation; (locs. 641, 64c) Glacier Lake Canyon Valley,
48 miles northwest of Lake Louise, Alberta.
Holotype—U.S.N.M. No. 108762; paratype, No. 108770.
MARYVILLIA MONTIS, new species
PLATE 13, FIGURES II-13
Several cranidia and pygidia are in hand. The glabella which tapers
to a rounded front, is almost without trace of furrows but does have
a faint keel. In this species the brim is rather wide, equaling one-
third the cranidial length. A slightly thickened rim, separated by a
wide anterior furrow, can be seen when the specimen is lighted from
the front, but when the direction of lighting is changed the brim
appears to be concave without a rim. At the eye the fixigenes are
about half the glabellar width. Longitudinally the cranidium is nearly
flat. However, the glabella is gently convex and the preglabellar area
depressed, while the brim rises until its anterior half extends above
the level of the glabella. In cross section there is little convexity.
The prominent pygidial axis occupies three-fourths the length of
the pygidium and has a postaxial ridge that reaches practically to the
rear margin. Axial and pleural furrows shallow. Moderate convexity
is developed in both directions.
Deadwood formation; (loc. 17L) 4 mile west of Deadwood, Black
Hills, South Dakota.
Holotype -—U.S.N.M. No. 108763a; paratype, No. 108763b.
MARYVILLIA HYBRIDA, new species
PLATE 13, FIGURES 14-17
Although this species is not fully typical of the genus, the cranidial
and pygidial characters appear to be closer to Maryvillia than to any
other described genus. Therefore 1. hybrida is placed in this genus
for the present. The exfoliated cranidium has a glabella that tapers
to a rounded front. The occipital ring is narrow. No trace of glabellar
furrows remains. The brim width equals nearly one-third the
72 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
glabellar length. It is concave without differentiation of a rim either
by furrow or change in contour.
At the eyes the fixigenes are about half the glabellar width, increas-
ing posteriorly to expand in rather large posterolateral limbs. In
cross section the cranidium is rather convex with an even curvature,
while longitudinally the convexity is greater. The pygidium has a
conspicuous axis rising above the pleural lobes. Axial and pleural
furrows obliterated. In cross section the profile is rather flat, but
longitudinally the axis is convex and the rear portion of the pygidium
slopes down steeply.
Cap Mountain formation; (loc. 14d) Bartlett Hollow, Burnet
County, Texas.
Holotype-——U.S.N.M. No. 108765a; paratype, No. 10876sb.
MARYVILLIA MOOSENSIS, new species
PLATE 14, FIGURES 15-17
This species is represented by several examples of cranidia and
pygidia from a single boulder found in the Moose River, and there-
fore the association of head and tail is certain. This is the first
identification of the genus in the lower beds of the Lynx formation
in the Robson District. The matrix is a dark crystalline limestone
containing vaugnite pebbles.
The illustrated cranidium is partially exfoliated, and therefore has
a deep dorsal furrow and faint glabellar furrows. The slightly
thickened rim is a little wider than the preglabellar area. Fixigenes
are narrow, averaging about half the glabellar width at its anterior
end. An exfoliated pygidium is illustrated, showing the nodes in the
axial furrows, and also shallow pleural furrows. The axis tapers very
little and slopes down in the rear rather steeply.
Lynx formation (Sullivan equivalent) ; (loc. 6rr) drift in Moose
River, 10 miles northeast of Robson Pass, British Columbia.
Holotype-—U.S.N.M. No. 108771a; paratype No. 108771b.
METEORASPIS Resser, 1935
METEORASPIS BANFFENSIS, new species
PLATE 13, FIGURES 5-10
Meteoraspis is not yet well understood. Reference of these cranidia
and possibly the pygidium to the genus is warranted according to our
present knowledge.
The large tumid glabella occupies about three-fourths of the cranid-
ial area. Furrows are only faintly indicated even on the exfoliated
cranidium. Occipital furrow and ring are well developed. Brim width
NO. 5 UPPER CAMBRIAN TRILOBITES—-RESSER 73
about one-fourth the glabellar length. Thickened rim wider than the
nearly flat preglabellar area. Fixigenes narrower than the rim, convex
adjacent to the eye. The small, strongly bowed eyes are set parallel
to the dorsal furrow. Suture diverges normally anterior to the eye.
Relief of the cranidium is considerable as shown in the illustrations.
Anterior angles and posterolateral limbs are moderately depressed.
Surface closely granulose.
The associated pygidium is similar to certain forms assigned to
Maryvillia or Coosella. Its axis extends about three-fourths the
pygidial Jength. Pleural furrows weakly developed. A rather wide,
flat, border is underlain by a striated doublure.
Sullivan formation ; (loc. 66m) 4% miles northwest of Banff, Saw-
back Range, Alberta.
Holotype —U.S.N.M. No. 108764a ; paratypes, Nos. 108764b, c.
COOSIA Walcctt, 1911
COOSIA CANADENSIS, new species
PLATE 13, FicuRES 18-20
Several pygidia and one cranidium have been segregated. It is a
rather small species for the genus but typical in most other respects.
The glabella, tapering to a rounded front, is without furrows. The
concave brim width is about one-third the glabellar length. A flat
rim demarcated by a shallow anterior furrow about equals the pre-
glabellar area in width. Fixigenes narrow, their width being less than
one-fourth the glabellar width at both ends of the eye lobe. The
fixigenes rise from the dorsal furrow, but the palpebral lobes have a
nearly horizontal position. Laterally the glabella is slightly convex,
and longitudinally the cranidium attains moderate convexity by de-
pression of the brim and the steepness of the slope in the anterior part
of the glabella. The pygidium has a prominent axis occupying a little
more than half its length. In dorsal view a postaxial ridge makes the
axis appear much longer than it is. Axial rings are faintly shown,
but the pleural furrows and grooves are so completely fused that only
the anterior pair remains.
Sullivan formation; (loc. 641) Glacier Lake Canyon Valley, 48
miles northwest of Lake Louise, Alberta.
Holotype —U.S.N.M. No. 108766a; paratype, No. 108766b.
COOSIA ALBERTENSIS, new species
PLATE 13, FIGURES 25-28
Although apparently an abundant species, only a few specimens
have been segregated in the collection. This is not a large species,
74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
but is typical of the genus. The glabella tapers to a rounded front.
A keel is clearly defined and faint shadows indicate glabellar furrows.
Brim, one-third the length of the glabella, has a more clearly defined
rim than usual. The fixigenes are narrow and the palpebral lobes
rather large.
In the pygidium the axis, with fairly deep furrows, occupies about
half the pygidial length. The pleural lobes slope down rather gently
to the wide border which assumes a horizontal position.
Sullivan formation; (loc. 651) Glacier Lake Canyon Valley, 48
miles northwest of Lake Louise, Alberta.
Holotype —U.S.N.M. No. 108768a ; paratype, No. 108768b.
COOSIA DAKOTENSIS, new species
PLATE 13, FIGURES 29-31
Although this large species is abundantly represented, only a few
unbroken examples of each part have been freed from the matrix. It
occurs in thin limestone lenses composed chiefly of trilobite fragments.
Some of the fragments show the heavy doublure characteristic of
the genus.
The cranidium is fully typical of Coosia in its tapering unfurrowed
glabella, wide concave brim, and narrow fixigenes. The brim width
is considerably more than half the glabellar length. It is without an
anterior furrow, but the heavy doublure causes a thickening in the
anterior portion. Fixigenes and eyes are normal in development.
Libragene large, with an elongate ocular platform and a heavy rim
that extends into a long, heavy, nearly straight genal ae Brim on
both the cranidium and libragene striated.
Pygidium normal, with the axis occupying about half ie pygidial
length. Axial rings well defined, and on exfoliated specimens the
pleural furrows are visible.
Deadwood formation ; (loc. 17j) Galena, Black Hills, South Dakota.
Holotype —U.S.N.M. No. 108769a ; paratypes, Nos. 108769b, c.
COOSIA TRIDENTENSIS, new species
PLATE 15, FIGURES 12-17
This species is abundantly represented in the small collection from
this locality. Unfortunately the rock is shattered with injection of
calcite veins, which with the crowding of the fossils accounts for the
difficulty in getting complete specimens.
The glabella tapers rather rapidly to a rounded front. Lighted
from certain directions it appears to be sharply truncate because of
the contour of the anterior lobe. Another peculiar feature is the
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 7.
ou
outward curvature of the dorsal furrow slightly anterior of the mid-
point of the eye, which creates a small side lobe on the glabella. The
concave brim is a little more than one-third the glabellar length. A
sudden increase of curvature develops a broad, shallow anterior
furrow near the middle of the brim. Between the eyes, with their
broad bands, the fixigenes are confined to the palpebral lobes, which
are not as wide as the eye band. Anterior to the eye the facial suture
diverges rapidly, developing large anterior angles. Since the eyes
extend almost to the occipital furrow, there is almost no fixigene
behind the eye, and the posterolateral limbs are therefore very narrow.
The pygidium has the typical oval shape, with a tapering axis that
occupies about two-thirds its length. A light postaxial ridge is
traceable to the margin. Longitudinally the axis is slightly convex,
dropping to the nearly flat border with a smooth concave curve.
Furrows, weak on the axis, become clearly defined on the pleural lobes
when the test is exfoliated.
Pilgrim formation; (loc. 20y) 3 miles north of Trident, north of
Logan, Montana.
Holotype —vU.S.N.M. No. 108786a; paratypes, Nos. 108786b-g.
COOSELLA Lochman, 1936
COOSELLA TEXANA, new species
PLATE 13, FIGURES 21-24; PLATE 14, FIGURES 2-5
The first species of Coosella, typical in all respects, to be recog-
nized in Texas occurs at two localities. Only the holotype cranidium
and the nearly complete pygidium were collected at one locality, but
several of each of the parts occur at the other place.
The large glabella tapers rapidly to the rounded front. Faint broad
depressions shadow the glabellar furrows. The brim width equals a
little less than one-third the glabellar length. In cross light the brim
is simple and concave, but when the lighting is from the front a broad,
slightly thickened rim is demarcated by a shallow anterior furrow.
The diagonal set of the eyes and the nondivergence of the anterior
facial suture cause the brim to be much shorter than the rear of the
cranidium, producing the pinched effect in the anterior part of the
cranidium characteristic of the genus. At the anterior end of the eye
lobe the fixigene equals half the glabellar width at the same point,
while at the rear of the eye lobe the same relationship is less than
one-third. This shows that the fixigene retains its width but little
changed, whereas the glabella expands rapidly. The eyes are of
normal size, situated opposite the anterior portion of the glabella, and
are set parallel to the dorsal furrow.
76 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
The axis is about half as long as the pygidium and although it rises
above the pleural lobes, its convexity is not great. On the exfoliated
portion axial furrows and the pleural grooves are clearly indicated by
shallow furrows.
Cap Mountain formation; (loc. 14d) Bartlett Hollow, and (loc.
67a) Potatotop, 7 miles northwest of Burnet, Texas.
Holotype —U.S.N.M. No. 108767a; paratypes, Nos. 108767b,
10877 3<a, b.
COOSELLA BREVIS, new species
PLATE 14, FicuREs 6-9
Only the illustrated incomplete cranidium and pygidium have been
found in this small collection. This species is fully characteristic of
Coosella in features of both the head and tail. The glabella, which
tapers to a rounded front, lacks glabellar furrows. On the other hand
the occipital furrow and ring are clearly defined. At the anterior end
of the eye the fixigene is about half the glabellar width. The fixigene
maintains the same width with little change. Owing to the moderate
divergence of the anterior facial suture, there is a slight expansion in
front of the eye. The brim width equals nearly half the glabellar
length. It has a heavily striated swollen rim, defined by the anterior
furrow. The preglabellar area, which is wider than the rim, is de-
pressed below both the rim and glabella.
The eye lobes, about half as long as the glabella, are not greatly
bowed and are set nearly parallel to the converging dorsal furrow.
The almost semicircular pygidium is slightly wider than long. The
axis stands above the pleural platforms, and extends back for more
than two-thirds the pygidial length. It slopes sharply downward
near the posterior end to a slight postaxial ridge. Axial and pleural
furrows are shallow, but clearly defined in the exfoliated type. Since
the test 1s thick it is probable they are not to be seen on the outer
surface. The doublure has a width equal to about one-fourth the
pygidial length.
Warrior limestone; (loc. 38e) 1 mile northwest of Benore, and
5 miles west of State College, Pennsylvania.
Holotype —U.S.N.M. No. 108774a; paratype, No. 108774b.
PTEROCEPHALINA Resser, 1938
PTEROCEPHALINA TEXANA, new species
PLATE 14, FIGURES 32, 33
Only the holotype pygidium has been found in this collection. The
wide axis with at least four rings tapers to a rather sharp point, back
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 77
of which a postaxial ridge extends to the indented median margin.
The pleural platforms are shaped as small triangles, considerably
curved to meet the wide concave border. This border develops sizeable
flanges at the posterior angles, giving the rear a rather straight
margin and the entire pygidium a quadrate shape.
Wilberns formation; (loc. 69) Honey Creek, 8 miles southeast of
Llano, Texas.
Holotype —U.S.N.M. No. 108780.
PTEROCEPHALINA NOTHA, new species
PLATE 14, FicuRES 34-38
The pygidium is typical of the genus. The associated cranidium
assigned to the species differs from the head supposed to represent
P. bilobata, but is like those assigned to the other species of the genus.
The glabella which has two pairs of short furrows, tapers to a
poorly defined front, as shown in the side views. This characteristic
is due not only to the shallowness of the anterior dorsal furrow but
also to the fact that the preglabellar area and the anterior fourth of
‘the glabella form a slope of uniform curvature to the depths of the
concavity in the brim. The wide brim rises sharply and the outer
portion of the rim is somewhat thickened. A slight change in slope
marks the inner edge of the rim which in itself is very concave.
Since it is so poorly marked, light must strike this angle properly in
order to make it visible. It will be noted that the rim is scarcely
perceptible in the dorsal view and in one of the side views. The
fixigenes rise rather steeply to the palpebral lobes, on which a strongly
curved eye occupies a prominent position because it has a wide
swollen eye band. Preglabellar area and part of rim marked by
vertical anastomosing lines. In the pygidium the axis is long and
rather slender, a postaxial ridge connecting it with the indented
median margin. A wide border is produced by the concave portion
underlain by the doublure.
Mendha limestone; (loc. 7j) 1 mile north of Italian Ranch foot-
hills, north end of Quinn Canyon Range, Nevada. ;
Holotype -—U.S.N.M. No. 108781a; paratypes, Nos. 108781b-d.
PTEROCEPHALINA BILOBATA (Hall and Whitfield)
PLATE 14, FIGURES 30-43
Dikellocephalus (Pterocephalus) bilobatus Hatt and Wuirtrietp, U. S. Geol.
Expl. 40th Par. vol. 4, p. 226, pl. 2, fig. 36, 1877.
Dicellocephalus bilobatus Watcott, U. S. Geol. Surv. Monogr. 8, p. 40, 1884.
6
78 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Platycolpus bilobatus Wa.cott, Smithsonian Misc. Coll., vol. 57, No. 13, p. 349,
1014.
ee piiiae bilobata ReEssER, ibid., vol. 97, No. 10, p. 42, 1938.
Additional specimens of this species are figured in order that it
may be better understood. A small tail is figtired to show the features
at that stage. The large cranidium, from which a rim of unknown
but evidently considerable width has been broken away, is tentatively
assigned to the species.
Secret Canyon shale; (loc. 65) east side of Sierra Canyon, opposite
Pinnacle Peak; and (loc. 61) south of Hamburg Mine, Eureka
District, Nevada.
Holotype —U.S.N.M. No. 24568; plesiotypes, Nos. 108782a-c.
PTEROCEPHALINA POGONIPENSIS, new species
PLATE I5, FIGURES I-2
Several small pieces of very fossiliferous limestone contain a
pygidium and a cranidium regarded as belonging to the same species.
Originally it was identified as P. flabellifer.
The cranidium is small for the genus. The glabella occupies more
than half the cranidial length and since it is exfoliated, three pairs of
recurved furrows are faintly visible in cross light. The concave brim
is divided into a slightly convex preglabellar area and a wider, flat,
somewhat upturned rim. The convex fixigenes are less than half the
glabellar width.
As may be observed in the illustrations the axis occupies nearly
the full length of the pygidium. It is almost straight on top, sloping
back at an even rate to the short postaxial ridge. Axial and pleural
furrows are well developed, the latter extending to the margins.
Though the lateral margins are somewhat extended, the border is less
flared than in more typical species.
Secret Canyon shale; west side of Pogonip Mountain, White Pine
District, Nevada.
Holotype —U.S.N.M. No. 108783a; paratype, No. 108783b.
PTEROCEPHALINA GRATA, new species
PLATE 15, FIGURES 3-6
This small distinctive species is represented by several cranidia and
pygidia. The narrow cranidium has a large slightly tapered glabella
and a concave brim about one-third as wide as the glabellar length.
The demarcation of a wide rim is so faint that only in a certain light
does the brim appear to be anything but a simple concave plate.
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 79
The axis occupies only about half the pygidial length. It is slightly
concave in a longitudinal direction and slopes abruptly to the wide
border. Furrows are visible on the axis and in the anterior portions
of the pleural platforms. A slight postaxial ridge extends to the rear
margin.
Secret Canyon shale; (loc. 23b) ridge east of Hamburg Ridge,
Eureka District, Nevada.
Holotype:—U.S.N.M. No. 108784a; paratype, No. 108784b.
PTEROCEPHALINA UTAHENSIS, new species
PLATE 15, FIGURES 7-II
A very fine species of Pterocephalina has been found in western
Utah. This genus should be represented in other formations of the
same age throughout the cordilleran region.
The glabella tapers to a truncated front, where the dorsal furrow
is almost obliterated. Three pairs of glabellar furrows are faintly
defined, the posterior pair having a very irregular course. The wide,
gently concave brim occupies nearly half the glabellar length. Fixi-
genes are narrow, being only one-third the glabellar width between
the eyes and the dorsal furrow. Eye bands are heavy. Anterior to
the eyes the facial suture diverges moderately to meet the wide brim.
The axis occupies less than half the length of the pygidium and stands
above the small pleural platforms. Three rings are clearly defined.
Three pleural furrows, decreasing in depth from posteriorly, are
visible. The wide border is nearly flat, and has a wide medial
indentation.
Orr formation ; (loc. 33d) east side of Fish Spring Range, Utah.
Holotype—U.S.N.M. No. 108785a; paratype, No. 108785b.
BURNETIDAE, new family
Kobayashi established the Dokimocephalinae as a subfamily in the
Solenopleuridae Angelin, to include the genera Burnetia, Dokimo-
cephalus, Iddingsia, Elkia, and Acrocephalites. The last-named genus
superficially resembles the others, but since it is a Middle Cambrian
Atlantic Province genus and therefore far removed from the others
in time and space, its relationship may be questioned. In my opinion
Acrocephalites is related to Solenopleura and should remain in that
family. The Burnetidae are not from the Solenopleura line but have
descended from trilobites found in the cordilleran region. To the
named genera must be added Berkeia, and several others still
undescribed.
80 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Thus constituted, the new family is found to play a rather promi-
nent role in the Upper Cambrian, beginning in Crepicephalus time and
extending to about the middle of the Uppet Cambrian. Wide variation
appears to exist between the ordinary swollen rim of Berkeia and the
enormous snout of Dokimocephalus. However, since the glabella,
fixigenes, eyes, and furrows are much alike in these genera, the varia-
tions in rim shape are not to be looked upon as a separating but rather
as a uniting feature of the family. Variability of the rim is found
between species of the same genus. Consequently, with such a char-
acteristic, wide variation must be expected in the family, and as such
becomes a family characteristic.
BURNETIA Walcott, 1924
BURNETIA INTERMEDIA, new species
PLATE 17, FIGURES I0, II
B. intermedia is one of the linguloid forms. Its glabella tapers
moderately to a rounded front. Two pairs of glabellar furrows are
clearly defined. The enlarged neck ring extends into a long, erect
spine. At the anterior end of the eyes the fixigenes are less than one-
third the glabellar width. The palpebral lobes are sharply angulated.
The palpebral furrow is wide and eye band heavy. The brim attains
a width equal to the length of the glabella exclusive of the neck ring, .
and the preglabellar area is less than one-fifth this width. Viewed
from the rear the glabella is roundly convex and the fixigenes rise to
the eyes. Viewed from the front, however, the relative convexity of
the several parts is much less, though the anterior fixigenes drop
steeply to the depressed anterior angles. Greater convexity of the
cranidium is found in the longitudinal profile, attained by a rather
even curvature of both glabella and brim, which together form a
smooth sigmoid curve. Surface granulose.
Honey Creek limestone; (loc. 12p) 4 miles southeast of Hennepin,
Arbuckle Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108805.
BURNETIA ALTA, new species
PLATE 17, FIGURES 12-14
The long glabella has two pairs of furrows. In dorsal view it
appears to be narrow and tapered more rapidly than in most species.
If, however, the excessive convexity in both directions is taken into
consideration the area of the glabella is really large. The brim is not
wide, being less than half the glabellar length exclusive of the neck
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 81
ting. The preglabellar area is less than one-fourth the rim width.
Longitudinally the convexity of the cranidium is so great that the
front fourth has taken the position of a go° angle with the posterior
portion of the glabella. From that position the brim takes a less steep
angle and is in itself nearly flat. In cross section the glabella is also
very convex, attaining a height equal to about one-third its length.
Because the anterior angles are also greatly depressed this species
becomes one of the most convex Cambrian trilobites.
The associated libragene is large. The wide and long ocular plat-
form is only slightly convex and lacks a furrow. Toward the long
genal spine the libragene becomes convex forming a ridge that extends
into the genal spine. Elevated portions of surface of head and cheek
coarsely granulated.
Davis formation ; (loc. 11k) Federal Lead Mine No. 4, Flat River,
Missouri.
Holotype-——U.S.N.M. No. 108801a; paratype, No. ro88o1b.
BURNETIA EXTENSA, new species
PLATE 17, FIGURES 15-22
This species is abundantly represented by numerous cranidia, several
libragenes, and possibly also by a pygidium. It is the extreme of the
linguloid forms thus far found. It will be observed that a slight
variation has been allowed among the specimens figured.
The glabella, rounded in front, has two pairs of furrows. The
occipital furrow is almost interrupted at the center, and the neck ring
carries a large spine. The brim, which comes to a rather sharp point,
exceeds the length of the glabella exclusive of the neck ring. A
narrow preglabellar area is present. Eyes are strongly bowed and have
heavy eye bands. The libragenes illustrated show that the suture is
intramarginal for a considerable distance and that the wide doublure
apparently extends all the way across, with an increase in width
toward the middle. The associated pygidium has a long, stout axis
and pleural lobes that are slightly convex, sloping to a concave border.
Honey Creek limestone ; (loc. 89v) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108807a ; paratypes, Nos. 108807b-g.
BURNETIA EXILIS, new species
PLATE 17, FIGURES 23-27
This species also is well represented by cranidia. Because of its
nearly circular anterior outline it is closer to the normal form. In
82 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
fact, taken as a whole, the cranidium gives the impression of circu-
larity. At the same time 5. evilis is a convex form, the glabella
standing high above the fixigenes and the brim continuing the down-
ward slope of the anterior portion of the glabella. Eyes, fixigenes,
glabellar furrows, and eye bands are similar to corresponding parts in
species of average structure for Burnetia. The brim width is about
three-fourths the glabellar length and has only a narrow preglabellar
area. Viewed from the front, the brim has an even curvature between
the depressed anterior angles. In cross light several shallow secondary
furrows parallel to the front margin are visible, but they may repre-
sent doublure impressions. Surface of elevated portions granulated.
Honey Creek limestone ; (loc. 89v) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108808a ; paratypes, Nos. 108808b, c.
BURNETIA ECTYPA, new species
PLATE 17, FIGURES 30, 31
Only a small fragment in addition to the beautiful holotype cranid-
ium has been found. The glabella, well rounded in front, has the
rear pair of glabellar furrows of normal development and very shallow
identations indicating the second pair. No nuchal spine and practically
no preglabellar area remain. The brim is about two-thirds the length
of the glabella including the neck ring. Longitudinally the glabella is
rather highly and evenly convex. The concave brim is set at such an
angle that it makes the cranidium as a whole high. In cross section
the glabella rises steeply above the dorsal furrow to a somewhat
angulated median portion. The rim is evenly convex between the
moderately depressed anterior angles.
Honey Creek limestone ; (loc. 89v) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108809.
BURNETIA LINGULA, new species
PLATE 17, FIGURES 32, 33
A single cranidium of another linguloid form, much like B. extensa,
has greater convexity and a brim of the same width as the glabellar
length. A narrow preglabellar area remains. Viewed from the side
the glabella is gently convex at the rear, increasing its rate of curva-
ture until the anterior third is turned sharply downward. The pro-
clivity of the glabella is continued by the brim at a decreasing rate
for nearly half its width, beyond which point the rate of slope de-
INO! 5 UPPER CAMBRIAN TRILOBITES—RESSER 83
creases until the anterior part is nearly horizontal. In cross section
the glabella is moderately convex, the anterior angles not greatly
depressed and the anterior portion of the rim nearly flat. Surface not
preserved.
Honey Creek limestone; (loc. 89v) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Holotype.—U.S.N.M. No. 108810.
BURNETIA CURTA, new species
PLATE 17, FIGURES 28, 29
A cranidium has a peculiar large glabella with a nearly circular
front outline. Three pairs of furrows are visible. Fixigenes and eyes
are normal. The brim width is not much more than half the glabellar
length and has a narrow preglabellar area. The neck ring extends
into a long elevated spine that nearly equals the length of the cranid-
ium. Convexity is not great in cross section, but longitudinally it
is considerably more.
Honey Creek limestone; (loc. gq) Blue Creek Canyon, Wichita
Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108811.
BURNETIA CAVA, new species
PLATE 20, FIGURES I-3
A cranidium and associated libragene impression represent this
species. Glabella normal in size and shape although it possibly is a
little wider than average. Two pairs of furrows are well developed
and a third is faintly indicated. The neck ring may have borne a
spine, but if so, it was no more than a node. The brim width is
considerably greater than half the glabellar length. Most of it consists
of a concave rim with only a narrow preglabellar area demarcated by
a shallow furrow. Fixigenes average only a little more than a fourth
of the glabellar width, and the divergence of the suture anterior to the
eye produces large anterior angles into which the wide rim extends.
The sharply bowed eyes are set at a somewhat more divergent angle
than the dorsal furrow and are situated slightly in advance of the
occipital furrow, leaving a considerable posterolateral limb. The broad
fixigene assigned to the species has a rather large genal spine. The
suture evidently is intramarginal to the center, but since it is broken
away in front, that structure is not readily observable.
Surface covered with evenly spaced granules except in furrows.
They are arranged in radiating lines on the libragene.
84 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Ore Hill formation; (loc. 107v) 4 mile northwest of Drab, Penn-
sylvania.
Holotype—U.S.N.M. No. 108827a; paratype, No. 108827b.
BURNETIA EDWARDSI, new species
PLATE 20, FIGURES 20, 21
A large cranidium associated with species of Taenicephalus and
Orygmaspis occurs on a hand specimen of soft sandstone. Although
the brim is narrow, the entire structure is typical of the genus. The
glabella, tapering at a normal rate, has several sets of broad, shallow
furrows faintly indicated. The occipital furrow also is broad and
shallow. The concave brim is subdivided almost equally between a
rim and a preglabellar area. Its width is just a little more than a
fourth of the cranidial length. The fixigenes are a little less than half
the glabellar width. The sharply bowed eyes are of normal size but
situated a little farther forward than the ordinary position for the
genus. In front of the eyes the sutures diverge considerably to form
large anterior angles. Longitudinally the head forms a sigmoid curve
with the rear part of the glabella nearly horizontal and the anterior
part arched down sharply to the concave brim. Viewed laterally the
glabella stands completely above the fixigenes and has a fairly even
curvature.
Franconia sandstone; 2 miles west of Clifton, Monroe County,
Wisconsin.
Holotype.—U.S.N.M. No. 108832.
BURNETIA PENNSYLVANICA, new species
PLATE 21, FIGURES 29-31
Several cranidia and a libragene from the type locality have been
assigned to this species. From the other locality there are other less
complete cranidia and fragments of other parts. This species is typical
of the genus, belonging to the group in which the brim is not ex-
panded. Restoration of the front of the glabella from the counterpart
allows the cranidium to be reproduced in its entirety. The glabella
tapers forward at the usual rate for cranidia of this type. The rear
pair of furrows is deeply impressed, while pits next to the dorsal
furrow indicate the position of the other two pairs. The width of the
moderately concave brim is just half the glabellar length exclusive of
the occipital ring. The rim is much wider than the preglabellar area
and is not evenly concave because it has a furrowlike depression
paralleling the anterior furrow. The eyes of normal size are so
sharply bowed that the palpebral lobe is angulated. At the anterior
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 85
end of the eye the fixigene is less than a third the glabellar width.
Anterior angles and narrow posterolateral limbs are strongly de-
pressed. Surface of exterior granulose, evidently with two sets of
granules. Exfoliated surfaces are even more closely covered with
coarser granules varying considerably in size.
Ore Hill formation; (loc. to7e) I mile south of Ore Hill; and
(loc. 107v) 4 mile west of Drab, Pennsylvania.
Holotype —U.S.N.M. No. 108842a; paratype, No. 108842b.
IDDINGSIA Walcott, 1924
IDDINGSIA NEVADENSIS, new species
PLATE 16, FIGURES 15-17
A cranidium and libragene which were previously identified as
I. robusta, differ from that species in several respects. J. nevadensis
is characterized by the usual large glabella on which the rear pair of
furrows are deep and the next pair short and shallow. A wide occip-
ital furrow separates a prominent neck ring. At the anterior end
of the eye the fixigenes are about one-third the glabellar width. In
width the brim is just half the length of the glabella including the
neck ring. It is divided about equally into a somewhat convex pre-
glabellar area and a slightly thickened rim. Longitudinally the cranid-
ium is convex, the glabella having a nearly even curvature except
toward the front, where there is a slight decrease. On the other hand
the brim as a whole is moderately concave. In cross section the
glabella is convex, and the libragenes rise from the dorsal furrow
rather steeply. The anterior angles are sharply depressed. Likewise
the posterolateral limbs appear to have been considerably depressed.
The associated libragene has a large ocular platform and a rim to
match the brim width. A heavy genal spine of unknown length is
built from the heavy rim and the thickened rear portion of the cheek.
This species differs from J. robusta in its longer glabella and rela-
tively narrower rim. In J. robusta the brim as a whole continues the
downward slope of the anterior half of the glabella, thus giving the
cranidium much greater convexity.
Secret Canyon shale; (loc. 61) South of Hamburg Mine, Eureka
District, Nevada.
Holotype-—U.S.N.M. No. 108796a; paratype, No. 108796b.
IDDINGSIA UTAHENSIS, new species
PLATE 16, FIGURES 18-20
Several cranidia from the Dugway Range are the basis for a species,
which in general appearance resembles /. nevadensis. I. utahensis is
86 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
characterized by a normal glabella on which two pairs of furrows are
clearly impressed. At a point immediately in front of the eye the
fixigene is about one-third the glabellar width. Divergence of the
facial suture in front of the eye develops moderately large anterior
angles. The brim width is about two-thirds the glabellar length in-
cluding the occipital ring. The slightly convex preglabellar area is
about two-thirds the width of the rim. Little curvature exists in the
rear half of the glabella, but the forward part turns down sharply.
The preglabellar area continues that slope, but the rim is turned
toward a horizontal position. Surface granulose, as usual.
Orr formation; (loc. 32t) Fandango Spring Canyon, Dugway
Range, Utah.
Holotype —U.S.N.M. No. 108797a; paratype, No. 108797b.
IDDINGSIA MISSOURIENSIS, new species
PLATE 16, FIGURES 21-26
This is a rather prolific species, associated with Berkeia, Ptero-
cephalia, and other genera. Besides the numerous cranidia several
libragenes also represent the species. The glabella, tapering to a some-
what truncated front, has three pairs of well-developed furrows. The
width of the flaring brim equals about two-thirds of the glabellar
length. In the middle the flat rim is wider than the preglabellar area.
The slightly convex preglabellar area continues the downward slope
of the anterior portion of the glabella, but the nearly flat rim has a
less steep slope. At the anterior end of the eye the fixigene is one-
third as wide as the glabella. It rises sharply from the dorsal furrow
but is in itself not very convex. The strongly bowed eyes are situated
far back, practically in contact with the occipital furrow. An eye line
extends forward from the eye to the anterior glabellar lobe. Longi-
tudinally the cranidium is strongly convex, the relief being attained
by a slight curvature of the posterior half of the glabella and a strong
down-curving of the anterior half, continued by the slope of the brim.
Laterally the glabella rises little above the level of the palpebral lobes,
but the posterolateral limbs are sharply depressed. At the eye lines
the fixigenes slope abruptly downward to the anterior angles. A small
node occurs on the well-defined occipital ring.
The rather large libragene has a long stout genal spine. At the eye
the fixigene rises steeply to the ocular platform which has a globular
convexity. The rim is flat and the suture intramarginal at least for
half and possibly all the distance to the center. None of the specimens
preserves the doublure which probably maintains its width across the
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 87
cranidium. A peculiar feature of the libragene is the convexity that
begins about halfway back in the rim and extends into the round,
elevated genal spine.
Davis formation ; (loc. 11k) Federal Lead Mine No. 4, Flat River,
Missouri.
Holotype.—U.S.N.M. No. 108798a; paratypes, Nos. 108798b-d.
IDDINGSIA BICINCTA, new species
PLATE 16, FIGURES 27-209
Several plump cranidia from Blue Creek Canyon constitute another
species. Compared with others in the region the distinctive feature
seems to be the subdivision of the brim into two approximately equal
portions. The rather short glabella is rounded in front and rises above
the dorsal furrow with an even curvature. Furrows are so faint that
they are scarcely traceable. The occipital furrow on the other hand is
wide and deep, and the ring carries a long slender spine. The brim
width nearly equals the length of the glabella exclusive of the neck
ring. At the anterior end of the eye, the fixigenes are only one-third
the glabellar width. Divergence of the facial suture forms rather
large anterior angles. Longitudinally the cranidium is convex, the
rear half of the glabella being nearly level and the anterior third
sharply curved, and the slightly convex preglabellar area continues
its downward trend, while the rim turns to a nearly horizontal position.
In cross section the glabella appears elevated with a nearly circular
curvature. In like manner the fixigenes near the eyes are also convex.
There is but a slight depression of the anterior angles so that the
slope of the preglabellar area is not much more at the sides than in
the middle. Surface of test nearly smooth. However, the exfoliated
preglabellar area is marked by rather strong vertical anastomosing
lines.
Honey Creek limestone; (loc. 9q) Blue Creek Canyon, Wichita
Mountains, Oklahoma.
Flolotype —U.S.N.M. No. 108799.
IDDINGSIA CRASSIMARGINATA, new species
PLATE 16, FIGURES 30-32
This cranidium, associated with /. missouriensis, is very distinct,
owing to its wide, flaring brim and relatively small glabella. The
glabella tapers to a slightly rounded front and has two pairs of
furrows. The brim is nearly as wide as the length of the glabella
exclusive of the occipital ring. It has a wide, slightly concave rim
88 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
nearly one and one-half times as wide as the preglabellar area. Eyes
are strongly bowed and situated just in front of the occipital furrow.
At the anterior end of the eye the fixigenes are less than half the
glabellar width. In cross section this species has little relief. The
glabella rises to a rounded median angle and the fixigenes rise without
curvature to the palpebral lobes. Longitudinally the glabella is gently
curved, increasing its declivity near the front, while the slightly con-
cave brim slopes less. The anterior angles are moderately depressed,
but the elevated palpebral lobes accentuate the steepness of the slopes
of the anterior fixigenes immediately forward of the eye lines.
Davis formation; (loc. 11k) Federal Lead Mine No. 4, Flat River,
Missouri.
Holotype —U.S.N.M. No. 108800.
IDDINGSIA ALPERSENSIS, new species
PLATE 16, FIGURES 33-38
The glabella is rounded in front and practically without furrows.
The brim width is about half the length of the glabella including the
thickened neck ring, which carries a long spine. Because of the
circular anterior outline of the cranidium and the comparatively much
less curved anterior furrow, the rim widens appreciably toward the
center, where its width is about equal to the preglabellar area. At
the anterior end of the eye the fixigene is about one-third the glabellar
width. In this species the eyes are not so strongly bowed, nor do they
extend back to the occipital furrow. As a whole the cranidium has
considerable convexity. In the rear the glabella stands high above the
dorsal furrow, but this relationship decreases anteriorly. Thus a
forward slope of the whole is created, which is continued by the
convex preglabellar area. The wide flat rim rises somewhat above a
horizontal position. A slight rise brings the fixigenes up to the
palpebral lobes. Anterior angles depressed. Preglabellar area marked
by strong anastomosing lines.
Honey Creek limestone ; (loc. 12p) 4 miles east of Alpers, Arbuckle
Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108801a; paratypes, Nos. 1o881Tb, c.
IDDINGSIA (?) QUINNENSIS, new species
PLATE 16, FIGURES 39-41
At first glance this cranidium seems to be far removed from
Iddingsia. However, when its features are examined individually,
each feature fits Jddingsia more closely than any other genus. In other
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 89
words, while the ensemble does not look so much like Jddingsia, the
separate features fall readily within its generic limits.
The glabella is large, nearly rectangular, slightly rounded in front,
and has three pairs of furrows. The neck ring carries a small spine.
Since the anterior facial suture diverges at a small angle and the
anterior margin is rather straight, the cranidium as a whole has a
quadrangular shape. The brim, divided almost equally between rim
and preglabellar area, equals about half the glabellar length exclusive
of the neck ring. At the anterior end of the eye the fixigene is less
than half the glabellar width. The eyes situated far back against the
occipital furrow are rather strongly bowed and have particularly
heavy palpebral furrows and eyebands.
Mendha limestone; (loc. 7j) 1 mile northwest of Italian Ranch
foothills, Quinn Canyon Range, Nevada.
Holotype—vU.S.N.M. No. 108802a; paratype, No. 108802b.
IDDINGSIA ANATINA, new species
PLATE 17, FIGURES I, 2
This neat species is characterized by its smooth, rounded glabella.
The occipital furrow is narrower than usual, but the neck.ring expands
in the center and has a long spine. The brim width equals the length
of the glabella exclusive of the occipital ring. Owing to the forward
projection of the anterior margin, the rim at its center exceeds the
width of the preglabellar area. Fixigenes are of average size and
shape, being peculiarly marked by the swellings in the inner rear
portion of the palpebral lobe. The evenly and sharply bowed eyes
have heavy bands. In cross section the evenly convex glabella rises
above the dorsal furrow. The fixigenes opposite the eyes are also
convex. Longitudinally the glabella is gently convex. The cranidium
as a whole slopes forward, its declivity being continued by the pre-
glabellar area as far as the anterior furrow, beyond which the rim is
turned up somewhat. Brim covered with vertical anastomosing lines.
Honey Creek limestone; (loc. 9q) Blue Creek Canyon, Wichita
Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108803.
IDDINGSIA SIMPLICITAS, new species
PLATE 17, FIGURES 3-9
This prolific species, represented by many cranidia and several
libragenes was first regarded as representing a new genus. However,
go SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
upon close examination it appears to be merely a simplification of
the regular ddingsia features.
The fairly large glabella has two pairs of faint furrows. The
expanded neck ring, separated from the glabella by a strong furrow,
carries a large spine in an elevated position. Although in the posterior
position of the genus /ddingsia, the eyes are sufficiently removed from
the occipital furrow to leave a small gap. Measuring along the con-
tours of the brim, its width equals the length of the glabella exclusive
of the neck ring. A median boss is developed in the convex pre-
glabellar area, thus accentuating the variant aspect of the species. A
change in contour alone marks the position of the anterior furrow,
In different light directions this concavity shifts its position, causing
the rim to appear relatively wider or narrower than the preglabellar
area. The fairly large libragene has a moderately convex ocular plat-
form. The suture is intramarginal for a considerable distance. A
swelling beginning in the rear portion of the rim and the outer corner
of the ocular platform, continues into the long genal spine which is
in a horizontal position, thus having a conisderable upward and out-
ward course from the ocular platform.
Honey Creek limestone ; (loc. 89y) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108804a; paratypes, Nos. 108804b-d.
BERKEIA Resser, 1937
BERKEIA COMES, new species
PLATE 15, Ficures 18-21
Although several cranidia are available, the pygidium has not been
determined.
All of the cranidia are exfoliated, and therefore the furrows appear
much deeper than on the exterior of the test. The large glabella is
well rounded in front. Because of the enlargement of the dorsal
furrow the glabella appears to be restricted posterior to the rear pair
of glabellar furrows. Three pairs of recurved furrows are clearly
impressed. The swollen occipital ring is elevated and carries a short
spine. The brim width is about one-fourth the glabellar length. In
dorsal view it appears to be divided almost equally between the pre-
glabellar area and the rim, but in side view the rim seems to be
relatively narrower. This subdivision is attained by a change in slope
between the somewhat convex preglabellar area and the slightly
thickened rim, turned up a little above the horizontal position. Oppo-
site the eyes the fixigenes equal one-third the glabellar width. Owing
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER gI
to the sharp depression of the anterior angles, the divergence of the
anterior facial suture is not apparent. Back of the eyes the postero-
lateral limbs, also sharply depressed, are about three times the width
of the occipital ring. Laterally the glabella stands completely above
the fixigenes with very steep sides. Longitudinally the entire cranid-
ium is very convex.
Deadwood formation; (loc. 88a) Deadwood, Black Hills, South
Dakota.
Holotype —U.S.N.M. No. 108787a; paratype, No. 108787b.
BERKEIA SARATOGENSIS, new species
PLATE 15, FIGURES 22-25
Although this species is not fully typical of the genus in that the
rim is narrower than the preglabellar area, the difference in appear-
ance may be due to the fact that on these specimens the test is pre-
served. However, since this form does not fit in any other existing
genus and because its departure from the characteristics of Berkeia
is confined to the narrowness of rim, the species is placed in that genus.
As usual the large glabella tapers to a rounded front and has three
pairs of recurved furrows. The brim width is somewhat less than
a third the glabellar length. The preglabellar area is more than twice
the width of the slightly upturned rim. Opposite and behind the eye
the fixigenes average about half the glabellar width. Divergence of
the suture creates fairly large anterior angles. In cross section the
total convexity across the eyes is moderate, although both the glabella
and fixigenes are individually convex. By depression of the anterior
angles and of the posterolateral limbs, both the front and back portions
of the cranidium attain considerable convexity. Longitudinally the
cranidium is strongly convex, attained by a rather even curvature
throughout.
Potsdam sandstone ; Greenfield, northwest of Saratoga, New York.
Holotype and paratypes—New York S. M.; casts U.S.N.M. Nos.
108792a-c.
BERKEIA NEVADENSIS, new species
PLATE I5, FIGURES 26, 27
B. nevadensis, represented at several localities, differs from B.
affinis chiefly in its lesser convexity in longitudinal direction. This
difference is not merely a matter of preservation, for B. nevadensis
is represented by nearly a score of specimens.
g2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Features which make this an average species of the genus include
similar glabellar proportions and furrows, narrow convex fixigenes,
and brim subdivided into convex preglabellar area and rim of equal
width. In dorsal view this species looks like B. affinis, but in side
view, although nearly equally convex in the glabella, it has a brim
less convex in its individual parts and as a whole much less down-
turned.
Secret Canyon shale; (loc. 61) south of the Hamburg Mine, and
(loc. 62) north of Adams Hill, Eureka District, Nevada.
Holotype —U.S.N.M. No. 108788a; paratype, No. 108788b.
BERKEIA RETUSA, new species
PLATE 15, FicuREs 28-30
This species acquires a peculiar aspect for two reasons. First, the
preglabellar area is wider than the rim, and second, the chief distinc-
tion lies in the slight swelling of the middle portion of the pre-
glabellar area. Glabellar furrows are well developed. Eyes and fixi-
genes are of normal size and shape. As stated, the preglabellar area
is wide and convex. Longitudinally the cranidium is rather evenly
convex, turning down more rapidly in the anterior half. In cross
section the glabella and fixigenes are each convex, but taken together
are rather flat in profile. The anterior angles and posterolateral limbs
are moderately depressed.
Honey Creek limestone; (loc. 12p) 4 miles southeast of Hennepin,
Arbuckle Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108789a; paratype, No. 108789b.
BERKEIA WICHITAENSIS, new species
PLATE 15, FIGURES 31-33
The rather long cranidium has the usual large glabella, in which
two pairs of furrows are well defined. The fixigenes average less
than a third the glabellar width. The preglabellar area is wider than
the rim. In this species the rim expands somewhat to the middle,
which together with the more angular anterior margin exaggerates
the actual length of the cranidium.
The small pygidium, tentatively assigned to the species, has a wide
axis occupying more than half its length, and flat pleural lobes. As
a whole the pygidium is oval in outline.
Honey Creek limestone ; (loc. 91L) Big Baldy, Wichita Mountains,
Oklahoma.
Holotype —U.S.N.M. No. 108790a; paratype, No. 108790b.
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 93
BERKEIA ANGUSTATA, new species
' PLATE 15, FIGURES 34-39
B. angustata is a prolific species which appears to be narrower than
others from the same region. Its aspect is due to the relatively long
glabella and the angulation of the anterior margin. The glabella,
rounded in front, has two well-developed pairs of furrows. The neck
ring extends into a short blunt spine. In this species the preglabellar
area is considerably wider than the brim.
Honey Creek limestone; (loc. 89y) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Holotype —vU.S.N.M. No. 108791a; paratypes, Nos. 108791b, c.
BERKEIA SCULPTILIS, new species
PLATE 16, FIGURES 1-4
This small species is not fully typical of the genus because of the
relative depression and constriction of the middle portion of its pre-
glabellar area. The prominent glabella stands well above the fixigenes.
It is rounded in front and has three pairs of furrows on exfoliated
specimens. The fixigenes average nearly half the glabellar width and
the brim width equals about one-third the glabellar length including
the neck ring. Both the anterior angles and the posterolateral limbs
are sharply downturned so that in dorsal view the strong curve of the
brim is exaggerated. The rim is thickened in the center, where it is
about as wide as the preglabellar area.
Honey Creek limestone; (loc. 12k) Honey Creek, 7 miles north of
Springer, Wichita Mountains, Oklahoma.
Holotype-—U.S.N.M. No. 108793a; paratype No. 108793b.
BERKEIA JUCUNDA, new species
PLATE 16, FIGURES 5-I0
This is the smaller of the two species in the Davis formation. It
is typical of the genus in every respect, agreeing rather closely with
those Oklahoma species which are characterized by the wide pre-
glabellar area. The moderately convex glabella tapers at the normal
rate to a rounded front and has three pairs of furrows on exfoliated
specimens. The fixigenes are narrower than the brim, averaging about
one-third the glabellar width. The eyes are rather large, and the
palpebral lobe is angulated near its center. The brim width equals
half the glabellar length exclusive of the neck ring. It has a nearly
even brim about half as wide as the preglabellar area. The front
margin is slightly angulated in the middle.
Zz
94 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Davis formation; (loc. 11k) Federal Lead Mine No. 4, Flat River,
Missouri.
Holotype -—U.S.N.M. No. 108794a; paratypes, Nos. 108794b, c.
BERKEIA MISSOURIENSIS, new species
PLATE 16, FIGURES II-I4
This abundant species associated with Pterocephalia and Burnetia
is one of the largest found. The slightly truncate glabella has three
pairs of furrows. The very narrow fixigenes at the anterior end of
the eye are only about one-fourth as wide as the glabella. On the
other hand the brim is wide, equaling more than a third the glabellar
length. In this species the eye lobes are only slightly curved. A small
node occurs on the neck ring.
Davis formation ; (loc. 11k) Federal Lead Mine No. 4, Flat River,
Missouri.
Holotype-—U.S.N.M. No. 108795a; paratypes, Nos. 108795b, c.
UNCLASSIFIED GENERA
ELVINIA Walcott, 1924
ELVINIA MONTIS, new species
PLaTE 18, FIGURES I-4
FE. montis is characterized by-a normally tapered, truncate glabella.
Two pairs of furrows are present, the rear pair taking a straight
course in the middle third. The brim width is about half the length
of the glabella exclusive of the occipital ring. It has a narrow rim,
which at its widest point equals about half the preglabellar area. At
the front end of the eye the fixigenes are half the glabellar width.
Back of the rather strongly bowed eyes, which slope outward at an
angle somewhat greater than the course of the dorsal furrow, the
fixigenes expand into wide posterolateral limbs. On the outer surface
none of the furrows are deeply impressed, and the surface apparently
is smooth.
It was hoped that this form would turn out to be E. tetonensis, the
species previously known from western Wyoming. Unfortunately it
proved impossible to place it in that species owing to the more truncate
glabella, less evenly curved glabellar furrows, and more strongly
bowed eyes.
Snowy Range formation; (loc. 37r) Warm Spring Creek, Wind
River Range, Wyoming.
Holotype-—U.S.N.M. No. 108812a; paratype, No. 1o8812b.
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 95
ELVINIA UTAHENSIS, new species
PLATE 18, FicuREs 5, 6
Except for the fact that this is such an important element of the
fauna, this peculiarly weathered cranidium would have awaited the
finding of additional material before its description. Strangely enough,
this is the only Elvinia specimen found in large collections from the
locality. In spite of its poor preservation this cranidium allows the
specific characteristics to be ascertained.
The glabella tapers forward at a normal rate to a truncate front.
Only a very shallow rear pair of furrows is visible, and the wide
occipital furrow is also shallow. The brim, occupying nearly a third
of the cranidial length, is separated into a thickened rim and a slightly
wider convex preglabellar area. Eyes normal in size and position,
and are not strongly bowed. The posterolateral limbs are long.
St. Charles limestone; (loc. 54e) Blacksmith Fork, Bear River
Range, Utah.
Holotype.—U.S.N.M. No. 108813.
ELVINIA RUEDEMANNI, new species
PLATE 18, FIGURES 7-10
Ruedemann identified this form as FE. matheri. However, very
considerable differences in brim structure and the granulated surface
exist. It is not certain that the larger and smaller cranidia here
illustrated belong to the same species, for none of the smaller heads
is well preserved, although it is clear that they are granulated. E.
rucdemanni belongs to a group which may eventually have to be
separated from Elvinia, but at present the altered appearance resulting
from depression of the preglabellar area does not seem to warrant
such action. .
E. ruedemanni is characterized by a rather rapidly tapering glabella.
Two pairs of furrows are developed, the anterior pair being short and
shallow while the deeper rear pair is connected as usual. The rather
large eyes are so far forward that their anterior ends are on a line
with the front margin of the glabella. At this point the fixigenes are
more than half the glabellar width. Rearward the fixigenes expand
less rapidly than the glabella, and extend into large posterolateral
limbs. The brim occupies about one-fourth the cranidial length and
consists of a thickened rim nearly circular in cross section. The rim
is less than half the width of the depressed preglabellar area. The
surface is covered with granules except on the preglabellar area where
vertical anastomosing lines take their place.
96 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I03
Potsdam sandstone (Theresa member) ; Greenfield, west of Sara-
toga Springs, New York.
Holotype and paratypes——New York S. M.; casts, U.S.N.M. Nos.
10881 4a-c.
ELVINIA GRANULATA, new species
PLate 18, Ficures II, 12
This cranidium, lying next to a good example of Berketa affinis, is
the only one found.
The large glabella tapers slightly to a rounded front. Only the rear
pair of furrows is developed. The eyes are situated well forward. At
the anterior end of the eyes the fixigenes are about one-third the
glabellar width. Posterolateral limbs large and long. The brim,
divided about equally between the preglabellar area and the thickened,
somewhat upturned rim, occupies about a fourth the cranidial length.
The surface is covered closely by evenly spaced and evenly sized
granules which in the preglabellar area overlie a system of vertical
anastomosing lines.
Secret Canyon shale; (loc. 63) northeast of Adams Hill, Eureka
District, Nevada.
Holotype —U.S.N.M. No. 108815.
ELVINIA MISSOURIENSIS Resser
PLATE .18, FIGURES 13-17
Elvinia roemeri Bripce (part), U. S. Geol. Surv. Prof. Pap. 186-M, p. 251,
pl. 69, figs. 12, 13, 1937.
Elvinia missouriensis RESSER, Smithsonian Misc. Coll., vol. 97, No. Io, p. 31,
1938.
A well-preserved cranidium, the pygidium, and the libragene are
figured to illustrate specific characteristics not shown in the holotype.
It will be observed that the doublure evidently extends completely
across the head.
Davis formation; (loc. 11k) Federal Lead Mine No. 4, Flat River,
Missouri.
Holotype-—U.S.N.M. No. 93011; plesiotypes, No. 108819.
ELVINIA GREGALIS, new species
PLaTE 18, Figures 18-23
This is one of the smaller species of the genus but seems to be rather
widespread in the Wichita Mountains.
The glabella tapers at a normal rate to a truncate front margin. A
shallow rear glabellar furrow is developed. The fixigenes average
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 97
about half the glabellar width. In front of the eye the suture diverges
little, but back of the eye it forms posterolateral limbs of normal shape.
The brim, almost exactly a third of the cranidial length, has a
somewhat thickened rim considerably narrower than the preglabellar
area. Longitudinally the considerable convexity is attained by a sharp
bend about the midpoint. In cross section the relief is not so great,
as the nearly flat fixigenes slope less steeply, and the glabella is gently
arched. The anterior angles are not greatly depressed. The associated
pygidium is characterized by the usual large, stout axis standing well
above the pleural platforms and terminating very abruptly in the rear.
Deep pleural furrows rib the slightly convex pleural platforms. The
border turns up sharply all around.
Honey Creek limestone; (loc. 9q) Blue Creek Canyon, and other
localities in the Wichita Miguntaing. Oklahoma.
Holotype —U.S.N.M. No. 108816a; paratypes, Nos. 108816b, c.
ELVINIA LONGA, new species
PLaTE 18, FIGURES 24-27
This species resembles E. gregalis but must be regarded as distinct
owing to greater relative and absolute relief and its longer glabella.
The long glabella appears to be slender. It tapers at a normal rate
to a rounded front margin. In spite of considerable relative relief
the glabellar furrows are very shallow. At the anterior end of the
eye the fixigenes are considerably more than half the glabellar width.
Their width is not greatly increased either forward or backward and
the posterolateral limbs are not so long. The brim occupies about
one-fourth the cranidial length, which makes it relatively narrower
than in A. gregalis. Both the rim and the preglabellar area are strongly
convex, the rim occupying a little more than a third the brim width.
Longitudinally the cranidium is decidedly convex, being slightly
curved in the rear half and sharply curved in the front portion. Since
the glabella and fixigenes are both convex, and the anterior angles
sharply depressed, the lateral convexity is great.
Honey Creek limestone; (loc. 9p) Blue Creek Canyon, Wichita
Mountains, Oklahoma.
Holotype-—vU.S.N.M. No. 108817a; paratype, No. 108817b.
ELVINIA BRIDGEI Resser
PLate 18, FicurEs 28-31; PLATE 19, FIGURES I-5
Elvinia roemeri BripcE (part), U. S. Geol. Surv. Prof. Pap. 186-M, p. 251,
pl. 60, figs. 19-21, 1937.
Elvinia bridgei Resser, Smithsonian Misc. Coll., vol. 97, No. 10, p. 31, 1938.
98 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Additional specimens of this species are figured to illustrate char-
acteristics of pygidium and the peculiar flatness of the head.
Honey Creek limestone ; (loc. 12p) 4 miles southeast of Hennepin,
and other localities in the Arbuckle Mountains, Oklahoma.
Cotypes—U.S.N.M. Nos. 93025a, b; plesiotypes, Nos. 108818a, b,
108820a, b.
ELVINIA VAGANS, new species
PLATE 19, FicuRES 6-9
Cranidium, exclusive of posterolateral limbs, nearly quadrate. Gla-
bella, tapered to a slightly rounded frontal outline, is a little more
than two-thirds the glabellar length. Brim divided about equally
between a well-defined, slightly upturned rim, and preglabellar area.
At their narrowest point the fixigenes are about one-third the glabellar
width. Anterior to the eyes the facial suture diverges moderately to
form rather square anterior angles. Posterolateral limbs long and
stout. The moderately bowed eyes are a little less than one-third the
cranidial length and are set nearly parallel to the lateral dorsal furrow.
Occipital furrow deep both on glabella and posterolateral limbs. Rear
glabellar furrow shallow, others not developed. Laterally the glabella
stands above the fixigenes with gentle curvature. Fixigenes nearly
flat. Longitudinally the head has considerable convexity attained by
an angulation near the midpoint. Each half individually is nearly in
a plane, except for the furrows and raised rim. Associate pygidium
normal in all respects. The wide axis reaches nearly to the rim. Three
axial furrows are well developed. The nearly flat pleural platforms
have two wide shallow furrows and a thickened and raised border.
Honey Creek limestone; (loc. 9q) 15 miles northwest of Fort Sill,
Wichita Mountains, Oklahoma.
Holotype-—U.S.N.M. No. 108821a; paratype, No. 1to8821b.
ELVINIA BREVIFRONS, new species
PLATE 109, FIGURES IO-14
This species is so constricted in brim width and length that it is no
longer fully characteristic of Elvinia. Cranidium as a whole forms a
truncated isosceles triangle. The long glabella which tapers to a
truncated front, with slightly rounded anterior angles, is demarcated
by a shallow dorsal furrow. Brim width equals about one-fourth the
glabellar length and has a nearly straight front outline. The occipital
and rear glabellar furrows are deep. Measured at the same point the
fixigenes are about half the width of the glabella. In front of the
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 99
eyes the facial suture diverges slightly. Eyes of moderate size, situ-
ated rather far forward, so that their anterior ends are about opposite
the front of the glabella. They are only slightly bowed, but the palpe-
bral lobes lie entirely outside the margin of the fixigene and are set
at an angle slightly greater than the course of lateral dorsal furrow.
Heavy but low eye ridges are developed. Viewed from the front the
cranidium is evenly convex, with glabella and fixigenes forming one
curved slope. Longitudinally the convexity is greater but is also even,
except for the slight flattening of the brim toward a horizontal posi-
tion. The associated libragene has a large occular platform, a raised
thickened rim, and a slender genal spine.
Honey Creek limestone ; (loc. 12p) 4 miles east of Alpers, Arbuckle
Mountains, Oklahoma.
Holotype —U.S.N.M. No. 108822a ; paratypes, Nos. 108822b, c.
TAENICEPHALUS Ulrich and Resser, 1924
TAENICEPHALUS PEALEI, new species
PLATE 10, FIGURES 15-23
Limestone layers in the Dry Creek formation cropping out from
Logan eastward to the Bridger Range, and thence southeastward
beyond Livingston, contain a species of Taenitcephalus, often occur-
ring in abundance. Most of the rock is a chowder, but here and there
cranidia and other parts escaped being broken before entombment.
This species is characterized by normal features throughout. The
glabella tapers at a normal rate. On the outside two pairs of shallow
furrows may be traced, but on exfoliated examples three pairs are
clearly visible. Where the test remains, the dorsal furrow is rather
deep but narrow, and the occipital furrow is prominent both across
the glabella and on the large posterolateral limbs. The fixigenes are
half the glabellar width at the anterior end of the eyes. Brim width
equals half the glabellar length. Preglabellar area rather wide and
strongly convex. Rim expanded forward in middle, giving the anterior
outline an angulated appearance. Eyes of moderate size and in normal
position.
The pygidium is of normal size and oval shape. The almost com-
pletely exfoliated example figured has deeper furrows and greater
relief than the exterior would show. The wide axis extends nearly
the full length of the tail.
Although the outside of test seems to be smooth, the preglabellar
area and other elevated portions of exfoliated specimens exhibit
100 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
scattered granules. Beneath the anterior furrow a somewhat irregular
row of granules is present.
Dry Creek shale; (loc. 153) north of Gallatin River, east of Logan,
Horseshoe Hills; (locs. 147a, 151, 152) Churn and Cottonwood
Canyons, Bridger Range; Davis Creek, south of Livingston, Snowy
Range, Montana.
Holotype.—U.S.N.M. No. 108823a ; paratypes, Nos. 108823b-g.
TAENICEPHALUS HOLMESI, new species
PLATE 19, FIGURES 24-27
The glabella which tapers to a rounded front, has three pairs of
fairly deeply impressed furrows. The brim width exceeds more than
half the glabellar length. At the center the brim is divided about
equally between rim and preglabellar area, both of which have about
the same convexity. At the eye the fixigenes are almost exactly equal
to half the glabellar width. The small eyes are situated about the
midpoint of the cranidium. They are very strongly bowed in a longi-
tudinal direction, which, together with the distinctness of the palpebral
lobe, projects the eyes considerably above the remainder of the cranid-
ium. In cross section the glabella is moderately, and the fixigenes
greatly, curved. Although individual parts have high relief the longi-
tudinal curvature is not great and is rather evenly distributed. The
pygidium, typical in size and shape, has considerable convexity.
The specific name is given in recognition of the geologic work done
by the collector, Dr. W. H. Holmes.
Dry Creek shale; (loc. 66x) Gallatin Valley, northwestern Yellow-
stone National Park, Wyoming.
Holotype and paratypes —U.S.N.M. No. 9595.
TAENICEPHALUS SPECIOSUS, new species
PLATE 20, FIGURE I9
This rather large species found in the northeastern part of the
Yellowstone National Park differs from T. holmesi in that all
furrows are shallower, and the brim is wider. Also, the glabella and
fixigenes are less highly arched in both directions. The surface is
smooth.
Dry Creek shale; (loc. 62r) Abiathar Mountain, northeastern
Yellowstone National Park, Wyoming.
Holotype.—U.S.N.M. No. 108831a; paratype, No. 108831b.
=e
ee ee
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER IOI
TAENICEPHALUS LIBERTYENSIS, new species
PLATE 10, FiIGuRES 28-32
Several cranidia, a libragene, and several small pygidia are contained
in the collection from this locality. Only the larger of the specimens
have been figured. The species is characterized by normal shape and
development of the several parts, with the possible exception of a
flattening in the preglabellar area. The glabella tapers at a normal
rate. On the outer surface its anterior outline is straighter than on
exfoliated specimens, a condition due to widening of the dorsal furrow
in the latter instance. Two pairs of furrows show on the outside and
three on the inside of the test. The deep occipital furrow is irregular.
The neck ring has a small spine. Fixigenes are about half the glabellar
length exclusive of the occipital ring. Rim slightly elevated and
thickened, expanding forward somewhat in the middle where it is
only slightly less wide than the preglabellar area. Eyes are normal
size and moderately bowed. In cross section the curvature is not
great in the central portion of the cranidium. In spite of the depressed
anterior angles the front margin is nearly straight but has considerable
relief because of the greatly depressed posterolateral limbs. Libragene
normal in size and shape, with a rather wide and possibly short genal
spine. The associated pygidium is normal in all respects.
St. Charles limestone; (loc. 56g) 6 miles west of Liberty, Bear
River Range, Idaho. ,
Holotype —U.S.N.M. No. 108824a; paratype, No. 108824b.
TAENICEPHALUS CORDILLERENSIS Miller
PLATE 10, FIGURE 33
Taenicephalus cordillerensis MiLLer, Journ. Pal., vol. 10, No. 1, p. 33, pl. 8,
figs. 40, 41, 1936.
Many of the fossils of this zone are found on weathered surfaces.
The illustrated example shows several cranidia in various states of
preservation and part of a libragene.
Snowy Range formation; (loc. 37r) Warm Springs Creek, Wind
River Range, Wyoming.
Holotype-—Columbia Univ. No. 12622; plesiotype, U.S.N.M. No.
108825.
TAENICEPHALUS GRANULOSUS, new species
PLATE 20, FIGURES 4-7
This moderately large species is covered with rather large, evenly
distributed granules.
102 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
The glabella tapers considerably to a narrow truncate front. Three
pairs of glabellar furrows are deeply impressed, and the occipital
furrow and ring are strongly developed. The brim width exceeds a
third of the cranidial length, and the preglabellar area is somewhat
wider than the rim at the center. At the eye the fixigene is about
one-third the glabellar width. Longitudinally the head is only slightly
convex, except that the fixigenes are flexed at the palpebral lobe. In
cross section the glabella is moderately convex, with a slight indication
of a keel, while the fixigenes are very convex. Both the rim and pre-
glabellar area are highly convex owing to the depth of both dorsal
and anterior furrows. Posterolateral limbs bent down rather sharply.
St. Charles limestone ; (loc. 4y) Two Mile Canyon, south of Malad,
Wasatch Mountains, Idaho.
Holotype—U.S.N.M. No. 108828a ; paratypes, No. 108828b.
TAENICEPHALUS MALADENSIS, new species
PLATE 20, FicurEs 8-10
Associated with T. granulosus is a smaller form with shallower
furrows. Its surface is not granulose but ornamented by raised
anastomosing lines, which form a more or less reticulate network.
Neither the fixigenes nor the preglabellar area is as convex, which
gives T. maladensis a smoother appearance than T. granulosus.
St. Charles limestone; (locs. 54x, 4y) Two Mile Canyon, south of
Malad, Wasatch Mountains, Idaho.
Holotype-—U.S.N.M. No. 108826a ; paratype, No. 108826b.
TAENICEPHALUS ORNATUS, new species
PLATE 20, FIGURES II, I2
Several species of Taenicephalus have been collected in Blacksmith
Fork canyon. They were not found at the same spot and therefore
may occur at somewhat different levels. Several cranidia, character-
ized by an ornate appearance due to the conspicuous granulation and
deep furrows as well as to the relatively high relief, may appropriately
be named T. ornatus. The rather wide glabella tapers at the average
rate of a rounded front and has three pairs of furrows rather deeply
impressed on the outside of the test. The slightly convex fixigenes
average nearly half the glabellar width. The brim is about one-third
the cranidial length. A deep anterior furrow separates the somewhat
thickened and considerably elevated rim which expands forward in
the center from agpreglabellar area of nearly equal width. The rather
small and very sharply bowed eyes are prominent because the palpebral
lobe projects outward and the fixigenes are arched at the eyes.
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 103
Longitudinally the relief of the cranidium is not great but because
both the dorsal and anterior furrows are deep the relative relief
between the several parts is accentuated. In cross section the con-
vexity of the glabella and sharp rise of the fixigenes to the eyes as
well as sharp depression of anterior angles and posterolateral limbs,
give both the front and rear margins of the head considerable curva-
ture. The surface is covered by evenly spaced granules.
St. Charles limestone; (loc. 54e) Blacksmith Fork, Bear River
Range, Utah.
Holotype —U.S.N.M. No. 108829.
TAENICEPHALUS STRIATIFRONS, new species
PLATE 21, FIGURES I-3
The second species from the locality is characterized by a rather
strongly tapering glabella and a vertically striated brim. The fixigenes
are rather evenly covered by granules but only a few are found on
the more elevated parts of the glabella. Brim width is considerably
less than one-third the cranidial length. The longitudinal convexity
is not great nor does the cross section attain high relief, even though
the glabella and the fixigenes are convex.
St. Charles limestone; (loc. 54e) Blacksmith Fork, Bear River
Range, Utah.
Holotype —vU.S.N.M. No. 108833a; paratype, No. 108833b.
TAENICEPHALUS UTAHENSIS, new species
PLATE 21, FIGURES 4-6
This third species from the locality is represented by several
cranidia. It is most like 7. striatifrons in shape and general appear-
ance but differs in having considerably less surface ornamentation.
The average size glabella tapers at a normal rate and has the usual
sets of furrows faintly outlined. The brim width is just about one-
third the cranidial length. The rim is about two-thirds the width of
the preglabellar area in the center. Both are convex. The fixigenes
are narrow and the eyes of normal size and shape. Longitudinally the
head is not very convex, most of it being attained in the anterior
portion. In cross section the fixigenes rise rather abruptly from the
dorsal furrow. The anterior angles are considerably depressed but
the posterolateral limbs are not.
St. Charles limestone; (loc. 54e) Blacksmith Fork, Bear River,
Range, Utah.
Holotype—U.S.N.M. No. 1088342; paratype, No. 108834b.
104 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
TAENICEPHALUS HYRUMEWNSIS, new species
PLATE 20, Ficures 13-18
This species is represented by a number of cranidia, a hypostoma,
and several pygidia. It is characterized by the smoothness of the test.
Slight indications of granulation are found on parts of the glabella
and rim. They are more clearly developed on the preglabellar area.
The rather long, fairly,slender glabella tapers to a rounded front.
Faint furrows are present on the test, but in exfoliated specimens
three well-developed sets can be seen. Brim width less than half the
glabellar length, divided by the anterior furrow into about equal parts.
The fixigenes average less than half the glabellar width. Postero-
lateral limbs fairly large. Eyes of moderate size, moderately bowed.
The associated pygidium is rather large and flat, approaching that of
Wilbernia in several respects, but it lacks the pronounced doublure
of that genus. It is possible that this pygidium should be referred to
T. modestus from the same locality.
St. Charles limestone; (loc. 55) Blacksmith Fork, Bear River
Range, Utah.
Holotype —U.S.N.M. No. aes paratypes, Nos. 108830b-e.
TAENICEPHALUS MODESTUS, new species
PLATE 21, FIGURE 7
A single cranidium is sharply demarcated from the other species at
the locality. Its reference to Taenicephalus may be questioned because
of the width of the cranidium across the brim and the flatness of the
whole. However, it is closer to this genus than to Elvinia, which it
most resembles. The glabella tapers at the normal rate and has
average proportions. Several sets of furrows are faintly indicated,
while the occipital furrow is rather deep. The brim width is half the
glabellar length exclusive of the occipital ring. Both the preglabellar
area and rim are convex and of about equal width. Fixigenes, eyes,
and course of the suture are approximately normal, except that the
anterior branch of the suture diverges slightly more than average.
St. Charles limestone; (loc. 55h) Blacksmith Fork, Bear River
Range, Utah.
Holotype —U.S.N.M. No. 108835.
TAENICEPHALUS TEXANUS, new species
PLATE 21, Ficures 8-12
All of the 10 or more cranidia are normal in size and shape with a
projecting front due to the forwardly expanded rim. The glabella
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 105
is about average in size and shape and has the usual three sets of
furrows. The brim width is half of the glabellar length exclusive of
the neck ring. Fixigenes are rather narrow, averaging a little more
than a third of the glabellar width. Surface finely granulose.
Wilberns formation; (loc. 68) Packsaddle Mountain, 11 miles
southeast of Llano; and (loc. 70) Baldy Mountain, 8 miles northwest
of Burnet, Texas.
Holotype-—U.S.N.M. No. 108836a; paratypes, Nos. 108836b-d.
TAENICEPHALUS WICHITAENSIS, new species
PLATE 21, FIGURES 13-17
A number of cranidia occur in a coarsely granular limestone full
of trilobite fragments. Much of the rock is nearly white, but it
contains many particles of glauconite and most of the trilobites are
stained red, which gives the rock a peculiar appearance. This species
is normal in all respects. The fairly wide glabella is truncated in the
front. All specimens are exfoliated, and on them three sets of furrows
are visible. The neck ring expands, possibly bearing a short spine.
The brim width exceeds the length of the glabella including the neck
ring. It consists of a rim that expands rapidly forward in the middle
from a nearly straight anterior furrow to give the front of the head
a considerable projection. The rim shrinks to extinction at the
anterior angles from the same width as the preglabellar area at the
center while the latter maintains its width into the anterior angles. At
the front end of the eye the fixigenes are less than half the glabellar
width, an average which they maintain throughout. The fairly large
eyes are in normal position for the genus. Triangular posterolateral
limbs are of considerable size. Surface of available exfoliated speci-
mens smooth, as are the small fragments of test remaining.
Honey Creek limestone; (loc. 9q) Blue Creek Canyon, 15 miles
northwest of Fort Sill, Wichita Mountains, Oklahoma.
Holotype-—U.S.N.M. No. 108837a; paratypes, Nos. 108837b-d.
TAENICEPHALUS QUINNENSIS, new species
PLATE 21, Ficures 18-23
This species is represented by cranidia, libragenes, and a pygidium.
Owing to greater width across the anterior part of the cranidium and
the flatter relief in the brim, the species approaches Elwvinia in
appearance.
The fairly wide glabella, well rounded in front, has the usual sets
of well-defined furrows, the rear pair being connected across the
106 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
middle by a wide bar. The brim width is just about half the glabellar
length including the neck ring. The rim, not greatly expanded beyond
the anterior angles, is much narrower than the preglabellar area.
Fixigenes rather wide, averaging over three-fourths the glabellar
width. Behind the eyes the sutures do not diverge rapidly, thus form-
ing wide and rather short posterolateral limbs. Anterior to the eyes
the divergence of the suture is not great, which leaves only moderate
anterior angles. The eyes are fairly sharply bowed. The palpebral
furrow in line with the facial suture has little curvature and a wide
eye band; hence the eyes are prominent. Longitudinally the head
does not have great convexity. Laterally, although both the glabella
and fixigenes are convex, the total relief is not great. Anterior angles
only moderately depressed and the posterolateral limbs even less.
Mendha limestone; (loc. 7}) 1 mile north of Italian Ranch foot-
hills, Quinn Canyon Range, Nevada.
Holotype-—U.S.N.M. No. 108838a; paratypes, Nos. 108838b-d.
TAENICEPHALUS CASTLENSIS, new species
PLATE 21, FIGURES 24, 25
Several cranidia are in the small collection from this locality, which
is situated between the Yellowstone Park and the occurrences of
Taenicephalus species north of the Gallatin River. It is typical in
most respects. The moderately long glabella tapers to a truncated
front. Three pairs of furrows are sharply impressed on the exfoliated
holotype, and the occipital furrow is wide. In the center the brim
width is a little less than half of the glabellar length including the
occipital ring. It consists of approximately equal subdivisions. The
rim tapers rapidly to extinction at the anterior angles which are
somewhat rounded. Both rim and preglabellar area have about the
same degree of convexity. Anastomosing lines cover the preglabellar
area, and a few scattered granules appear on the rim. Eyes, fixigenes,
and posterolateral limbs are of normal size and proportions.
Dry Creek formation; (loc. 62s) Castle Peak, north of Squaw
Creek, Gallatin Range, Montana.
Holotype-—U.S.N.M. No. 108839.
TAENICEPHALUS WYOMINGENSIS, new species
PLATE 21, FIGURE 32
A small collection from the Big Horn Mountains contains a good
species of Taenicephalus. It is close to T. cordillerensis Miller from
the Wind River Range but has a wider preglabellar area. Compared
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 107
with T. holmesi, the palpebral lobes are less prominent because the
fixigenes are less sharply bowed at that point.
Snowy Range formation; head of Buffalo Fork, west side of Big
Horn Mountains, Wyoming.
Holotype and paratypes —U.S.N.M. No. 11593.
TALBOTINA Lochman, 1938
TALBOTINA CANDIDA, new species
PLATE 21, FIGURES 27, 28
The glabella covers somewhat less than half the cranidial area and
is sculptured by three sets of recurved furrows and a deep, wide
occipital furrow. The fixigenes average about half the glabellar width.
Posterolateral limbs are fairly long and of normal width. The brim,
about equal in width to the fixigenes, is separated into rim and pre-
glabellar area. The somewhat swollen rim widens to the center, chiefly
by rather sudden expansion rearward. The eyes, of average size, are
situated behind the midpoint of the glabella. Eye lines present. Con-
vexity is about average in amount, while the relief is pronounced. The
surface is abundantly granulose. Libragene, thorax, and pygidium
unknown.
Wilberns formation; (loc. 14b) Cold Creek, opposite north end of
Sponge Mountain, Texas.
Holotype—U.S.N.M. No. 108841.
TALBOTINA ULRICHI, new species
PLATE 21, FIGURE 26
A second species from Texas is characterized by being shorter and
by the glabella in consequence occupying a relatively greater pro-
portion of the cranidium. The anterior furrow also has a more even
course near the center, although the rim remains proportionally about
the same as in T. candida. The surface is nearly smooth.
Wilberns formation; (loc. 14b) Cold Creek, opposite north end of
Sponge Mountain, Texas.
Holotype —U.S.N.M. No. 108840.
TALBOTINA CAELATA, new species
PLATE 10, FIGURE 4
When the fauna from the locality was described, this cranidium was
overlooked. T. caelata is a beautifully sculptured trilobite of medium
size. It differs essentially from the genotype in its narrowness.
108 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
The large, well-demarcated glabella tapers at a normal rate to a
rounded front. Two pairs of shallow recurved furrows are visible.
The fixigenes at the anterior end of the eye are about half the glabellar
width. The eye lines are made conspicuous by a furrow which
parallels them on their front side. Brim about half the glabellar length
exclusive of the occipital ring. A conspicuous, somewhat thickened
rim widens backward in the center, greatly reducing the width of the
preglabellar area. The moderate-sized eyes are not strongly bowed
and are situated posterior to the middle of the cranidium. The cranid-
ium has considerable relief in each separate part, as well as in its
entirety. Longitudinally the cranidium stands high, with a fairly even
curvature, slightly accentuated in the glabella. In cross section the
glabella stands completely above the fixigenes. The fixigenes are
rather strongly convex in lateral direction, having greatly depressed
posterolateral limbs, with convexity increasing anterior to the eye lines.
Nolichucky shale; (loc. 27d) U.S. 11, 7 miles northeast of Rogers-
ville, Tennessee.
Holotype-—U.S.N.M. No. 108734.
EX PEANATION OF PLATES
Figures are natural size unless otherwise indicated. Since all the
species described belong to the Upper Cambrian, only the formational
names are given with the locality descriptions.
Ilo SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
PLATE
Chariocephalus tenerus, MEW SPECIES... 650620 e eee e ence ween sewers cesacss
1-3. Dorsal, front, and side views of the holotype cranidium (X 5).
Honey Creek limestone : (loc. 37t) 4 miles southeast of Hennepin,
Arbuckle Mountains, Oklahoma.
Chariocephalus bellus, ‘new SPECIES: a!iee sis sae so ae eens sree ens eee 6
4-6. Dorsal, front, and side views of the holotype cranidium ( X 3).
Honey Creek limestone; (loc. 37v) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Chariocephalus bulla, new SPeCieS.....5...00 eee cece e Pee e eect eee eceees 7
7, 8. a and rear views of the pygidium nected to the species
(X 2)
Q-II. Dorsal, side, and front views of the holotype cranidium ( 3).
Honey Creek limestone; (loc. 37v) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Chartocephalus teres, NeW SPECIES........ 0. eee cece cence eee e nese eee nenes 7
12, 13. Dorsal and front views of holotype cranidium (X 5).
Honey Creek limestone; (loc. 37v) West Timbered Hilly, Arbuckle
Mountains, Oklahoma.
Ghartocephalus. affinis, new SpeCleS= asm y)- cise ee eee aia ieee tae 8
14. Associated libragene (XX 2).
15-17. Side, dorsal, and front views of the cranidium ( X 3).
18-20. Dorsal, side, and rear views of the pygidium (X 3).
Honey Creek limestone; (loc. 37u) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Chartocephalus agrarius, Tew. SPECIES. ccs -ccmices «snes eos oie eye ete eee 8
21. Libragene (X 2).
22. Hypostoma (X 1.5).
23. Large cranidium (X 2).
24-26. Dorsal, front, and side views of holotype cranidium (X 2).
27, 28. Dorsal and side views of pygidium (X 2).
Honey Creek limestone; (loc. 37v) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Gharsocephalus gracitlens; newaSpecies«... sania ae ee eee eee eee 9
29-31. Dorsal, front, and side views of the holotype cranidium (XX 3).
Honey Creek limestone; (loc. 9y) 2 miles southeast of Canyon Creek,
Wichita Mountains, Oklahoma.
Chariocephalis montis, New Species. .-ieeee eae eine ener eee ere 10
32-34. Side, front, and dorsal views of holotype cranidium (X 5).
Honey Creek limestone; (loc. 91q) 8 miles southeast of Mountain View,
Wichita Mountains, Oklahoma.
Chariocephalus angustatus, new Species... 4.05. oe ve eos ces nesses eee 9
35-37. Dorsal, side, and rear views of the associated pygidium (X 2).
38-40. Holotype cranidium ( 6).
Honey Creek limestone; (loc. 37v) West Timbered Hills, Arbuckle
Mountains, Oklahoma,
VOL. 103, NO. 5, PL. 1
UPPER CAMBRIAN TRILOBITES
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 5, PL. 2
UPPER CAMBRIAN TRILOBITES
IN KO) 5 IS UPPER CAMBRIAN TRILOBITES—RESSER II!
PLATE. 2
Chariocephalus wichitaensis, NCW SPECIES... 16.6. e cece cece eee tee eee eee 10
1-3. Dorsal, side, and front views of holotype cranidium (X 1.5).
4. A larger, less perfect cranidium.
5, 6. Libragenes.
7, 8. Associated pygidium (X 1.5).
Honey Creek limestone; (loc. 91q) 8 miles southeast of Mountain View,
Wichita Mountains, Oklahoma.
C@haviocepialus wirichin MEW, SPECIES: + «ec )ei0 22s cle eieiceisieiaieieles sie or oc II
9, 10. Side and dorsal views of holotype cranidium (X 3).
11. Libragene.
Ts Front. and oblique views of cranidia (1.5).
13, 14. Side and dorsal views of the pygidium ( 1.5
Honey Creek limestone; (loc. 9y) 2 miles southeast 5} ‘Canyon Creek,
Wichita Mountains, Oklahoma.
(GHOPIOCEPNOLUSNGGNUS, NEW, SPECIES. wales 0 scene «leon cclewc dlewlicee secemse II
15-17. Holotype cranidium. (See also pl. 12, fig. 13.)
Lyell formation; (loc. 20j) Tilted Mountain Brook, 95 miles east of
Lake Louise, Alberta.
Chariocephalus Vadgerensis, MEW SPECIES. ... 2.0.2. c eee e cece ence ceresenes 12
18-20. Exfoliated holotype cranidium (X 2.5).
Lyell formation; (loc. 191) Badger Pass, Johnson Canyon, 8 miles east
of Lake Louise, Alberta.
iarorepnolus Durlingl, MEW SPECIES... ccc cee os cee eee eens eee neees 12
21. Holotype cranidium ( 1.5).
Lynx formation; (loc. 190) Iyatunga Mountain, Robson Peak District,
British Columbia.
na SIDVESETUS MEW. SDECICS\. 6.0/2 4a) eels dea ialtle sisie.c b.9 0 sacs no's noel wears 13
22-24. Side, front, and dorsal views of holotype Se (S<a))-
25-27. A larger, partially exfoliated cranidium (XX 1.5
Honey Creek limestone; (loc. 89w) west side of the West Timbered
Hills, Arbuckle Mountains, Oklahoma.
Irvingella arbucklensis, new SpeCi€S......-.. eres eee e eee e ese e cece ce eeees 14
28-30. Front, dorsal, and side views of the holotype cranidium (> 1.5).
Bin 2: Similar views of another cranidium CeZr
33. An incomplete cranidium (X 1.5).
Honey Creek limestone; (loc. 89w) west side of the West Timbered
Hills, Arbuckle Mountains, Oklahoma.
er Cacia MesIeri NeW SPECIES. 25) U ees Rial tis oid bie wie oye in 468 ate love etal ole's's 15
34-306. Dorsal, side, and front views of the holotype cranidium C<3a)F
BW Gish Waal incomplete cranidium (X 3).
Honey Creek limestone; (loc. 12n) 7 miles north of Springer, Arbuckle
Mountains, Oklahoma.
MILE Le VE GUNUC. TICW, SPECIES: c/s eee ers di Giane, 5 sie. +isie1 esspe a) sto e) s/o epee eusie eye) ole sie 15
30-41. Dorsal, front, and side views of the holotype cranidium (X 2).
Honey Creek limestone; (loc. 12m) 7 miles north of Springer, Arbuckle
Mountains, Oklahoma.
MCMTUCEN Ua ChIAC TE WisSPCGIES specie cits «ever aaj leie! aye octal eroietisy ol else onel sneusyeseavetehersjetete 16
42-44. Dorsal, front, and side views of the holotype cranidium (X 2).
45-47. A larger cranidium (> 1.5).
Honey Creek limestone; (loc. 9p) about 15 miles northwest of Fort
Sill, Wichita Mountains, Oklahoma.
TCT AR IESCMISIC RTE Ws SPECIES <\. . daib.sc ve. s sat ea Seen cee aise ene res 16
48. Holotype cranidium (X 4).
Davis formation; (loc. 22m) Shaw Branch, Davis Creek, Missouri.
PHN OCIIOLOLLCKIGILCNSIS, TIEW! SPECIES). ..c\-:015,< e516) sclsvehe o/d0e ae) seis elatel wiles sknya 2oele ere 17
49. Impression of holotype cranidium (X 2.5). ;
“Goodsir” formation; Moose Creek, Ottertail Range, British Columbia.
II2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
PLATE 3
Iroingella oblanga;. new Species): «i).i0 wie eames sae ne le teehee 17
1-3. Dorsal, side, and front views of the holotype cranidium (X 2.5).
Wilberns formation; (loc. 70) Baldy Mountain, 8 miles northwest of
Burnet, Texas.
Irvingella agrestis, NeW SPECIES jcsa.c.: iain airy ay ales adidas" vs saa 0s iohoa ol we 17
4-6. Dorsal, front, and side views of the holotype cranidium (> 2.5).
Wilberns formation; (loc. 69) Honey Creek, 8 miles southeast of Llano,
Texas.
Iyvingella ardmorensis: NEW SPECIES a .tio scien cee cisic sree icine aetna 18
7-9. Dorsal, side, and front views of the holotype cranidium (> 1.5).
10-12. Another partially exfoliated cranidium (> 1.5).
Honey Creek limestone; (loc. 12p) 4 miles east of Alpers, Arbuckle
Mountains, Oklahoma. °
Irvingella plena; lew: Speciesc— + «eee an eee Soe See een eee 18
13-15. Dorsal, side, and front views of the holotype cranidium (X 1.5).
Wilberns formation; (loc. 14b) Cold Creek, opposite north end of
Sponge Mountain, San Saba County, Texas.
Irvimgella-alia, new specieS: s..0. cere Ree eee CCC Ree 19
16-18. Dorsal, side, and front views of the holotype cranidium (X 1.5).
Honey Creek limestone ; (loc. 9p) 15 miles northwest of Fort Sill,
Wichita Mountains, Oklahoma.
Trumgelia decker, new Species. 4.4 44sec eee eee 19
19-21. Views of a small cranidium ( 2.5).
22-24. A larger cranidium (1.5).
25-27. Front, side, and dorsal views of the holotype cranidium (x 3).
Honey Creek limestone; (loc. 89y) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Irvingella burnetensis, new <Speciese..uc . 5 acisenighle's aren eek ae eee 20
28-30. Front, dorsal, and side views of the holotype cranidium ( 1.5).
31-33. Dorsal and side views of two cranidia on one base, and front
view of the upper specimen ( 2).
Wilberns formation; (loc. 70) Baldy Mountain, 8 miles northwest of
Burnet, Texas.
Irvingella: bacca; new species?) 4.004 jak cee ee eee ee 20
34-30. Dorsal, side, and front views of the holotype cranidium ( 2.5).
Wilberns formation; (loc. 14b) Cold Creek Canyon, opposite north
end of Sponge Mountain, San Saba County, Texas.
Iruimngella atcimcta. new SRECIESxs..,.,.¢ 5.5 6 Coe ee eee ee 21
37-39. Front, side, and dorsal views of the holotype cranidium (XX 2.5).
Wilberns formation; (loc. 14b) Cold Creek Canyon, opposite north end
of Sponge Mountain, San Saba County, Texas.
Irvingella abrupta. new: Species: ».< asitow tenes os see he eee eee eee 21
40-42. Dorsal, side, and front rN of the holotype cranidium (X 1.5).
43-45. A smaller cranidium a2
Wilberns formation; (loc. 14b) Cold ‘Creek Canyon, opposite north end
of Sponge Mountain, San Saba County, Texas.
lrumgella media, new: speciess iid ig ae eee 22
46-48. Side, front, and dorsal views of the holotype cranidium (> 5).
49-51. Another excellent specimen (5).
Wilberns formation; (loc. 70) Baldy WOwanaE 8 miles northwest of
Burnet, Texas.
52-54. Dorsal, front, and profile views of a larger cranidium (> 3).
W ilberns formation : (loc. 14b) Cold Creek, opposite north end of
Sponge Mountain, San Saba County, Texas.
SMITHSONIAN MISCELLANEOUS COLLECTIONS
UPPER CAMBRIAN TRILOBITES
VOL. 103, NO. 5, PL. 3
SMITHSONIAN: MISCELLANEOUS COLLECTIONS VOL. 103, NO. 5, PL. 4
UPPER CAMBRIAN TRILOBITES
Tin PAS setts
a Peo ee
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 113
PLATE 4
PAGE
TRPMEO CLG LOLUCKIO: NEW SPECIES....:0i:0 soc). tiele vs wc dee ae ey Se eee s mwle els nid ae ae 23
1-3. Dorsal, front, and side views of the holotype cranidium (X 2).
Lyell formation; (loc. 19u) Johnson Creek, 8 miles east of Lake
Louise, Alberta.
Une lGarichmondenstsy ew SPECIES. s1..) > 1s. 2 ss cs yaeiewl cine be ees cee oe 23
4-6. Dorsal, front, and side views of the holotype cranidium ( 6).
Secret Canyon shale; (loc. 60) near Richmond Mine, Eureka District,
Nevada.
UGAIUGLUG AAAMSCNSIS, TIEW SPECIES’. os... cc. cei gis se dsc els boss due ge abe teleeled 24
7-9. Side, dorsal, and front views of the holotype cranidium ( 2.5).
10, 11. Front and dorsal views of a larger cranidium (> 1.5).
nore Canyon shale; (loc. 60) near Richmond Mine, Eureka District,
evada. ;
are ERO LRU TIEW “SDECIES(S 215. fia ay.) k c'a.01» (oh 'ase-ayesetwloia ga, ole) alu o lielejaiabaraseaesniedeld 24
12-14. Front, side, and dorsal views of the holotype cranidium (X 3).
Secret Canyon shale; (loc. 60) near Richmond Mine, Eureka District,
Nevada.
Rorarvingella €urekensis, NEW. SPeCieS....0. 45... ssccceccevececceacceence 26
15-17. Dorsal, front, and side views of the holotype cranidium (> 3).
Secret Canyon shale; (loc. 61) south of Hamburg Mine, Eureka Dis-
trict, Nevada.
Parurvingella angustilimbata Kobayashi..............0..ccccecceccercas 26
18-20. Two dorsal, front, and profile views of a complete cranidium
Sa)
21, 22. Side and dorsal views of the holotype cranidium (> 1.5).
Secret Canyon shale; (locs. 62, 63) north of Adams Hill, Eureka
District, Nevada.
Pararvingella hamburgensis, new SPeCieS...........0.cceccccescceesceeee 27
23, 24. Dorsal and side views of the holotype cranidium (X 2).
Secret Canyon shale; (loc. 61) south of Hamburg Mine, Eureka Dis-
trict, Nevada.
Parairvingella intermedia, new species
SI ee hrel spar fares ara ee Ree A eG 27
25-27. Dorsal, front, and side views of the holotype cranidium (> 3).
28, 29. A larger cranidium (X 2).
30, 31. Large incomplete cranidium (1.5).
Secret Canyon shale; (loc. 61) south of Hamburg Mine, Eureka Dis-
trict, Nevada.
EASES EPS TUGHOCNSTS. TIEW SPECIES. .'o:. 6. cje-wejewalon's slots Sade ooo ane oe ciate as 20
32-34. Dorsal, front, and side views of the holotype cranidium ( 2).
35, 36. Another, less complete cranidium ( 2).
St. Charles limestone; (loc. 4y) Two Mile Canyon, 2 miles south of
Malad, Wasatch Mountains, Idaho.
TE MLES PIS RUCICOLEDS: MEW; SPECIES snore a (oie sree are lato al aah ooh ieee nose eee 28
37-39. Dorsal, side, and front views of the holotype cranidium ( 5).
40, 41. A larger cranidium (X 3).
St. Charles formation; (loc. 4y) Two Mile Canyon, 2 miles south of
Malad, Wasatch Mountains, Idaho.
manngela tumirons (Hall and Whitfield) .........0..3 000002 ssccccecuseces 25
42, 43. Dorsal and front views of the holotype cranidium (X 1.5).
Secret Canyon shale; Pogonip Mountain, White Pine District, Nevada.
IIl4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
PLATES
Drumaspis alberta, new. SPECIES)... io cu' ns. eouys ceinvizce= ens shee oetentenees 29
1-3. Dorsal, side, and front views of the holotype cranidium (X 6).
Lyell formation; (loc. 64x) Ranger Brook Canyon, Sawback Range,
Alberta.
Drumaspis briscoensts, NEW SPECiES.... </cco.civie sie» am Fi sles ele eels tie © Sole ele 30
4, 5. Front and dorsal views of an incomplete cranidium (X 2.5).
6-8. Three views of the holotype cranidium (X 2.5).
Sabine formation; (loc. 17p) Radium Hot Springs, Brisco Range,
British Columbia.
Drumaspis goodswensts, NEW SPECIES. cis om, cia<a os ol nin cininieiw oiete'wie ieee 30
9. Impression of holotype cranidium (X 3).
“Goodsir” formation; Moose Creek, Ottertail Range, British Columbia.
Drumaspis sabimensis, NEW SPECIES. 5% ./2 oe alee eis vie 2a ne somite elelele oie terete hi
10, 11. Dorsal and side views of the holotype cranidium (X 6).
Sabine formation; (loc. 17s) Sabine Mountain, 2 miles northeast of
Canal Flats, Brisco Range, British Columbia.
Drumaspis maxwelle; MEW SPECIES << a5 << tint ee whe me shot sible ye ieee 31
12, 13. Side and dorsal views of the holotype cranidium ( X 3).
St. Charles limestone; (loc. 66z) St. Charles Canyon, Bear River
Range, Idaho.
Drumaspis deckert, New :SpeCies: . 32 ).07 Sian: erie ieieiienre am ee eee pe ees 31
14-16. Dorsal, front, and side views of the holotype cranidium (X 6).
Honey Creek limestone; (loc. 91b) 4 miles southeast of Hennepin,
West Timbered Hills, Arbuckle Mountains, Oklahoma.
Drumaspis osella, new SPECIES sm oi <o sieisistn,» ole t= apple) ol nde se ae ele ee 32
17-19. Dorsal, side, and front views of the holotype cranidium (X 3.5).
20. Another less complete cranidium ( 3.5).
Wilberns formation; (loc. 70) Baldy Mountain, 8 miles northwest of
Burnet, Texas.
Drumaspis wichttaensis, MEW SPECIES nase. aise « oes aie ea ajar e oes aie 33
21, 22. Dorsal and side views of the holotype cranidium ( X 2).
Honey Creek limestone; (loc. 91u) 1 mile east of Canyon Creek, 15
miles northwest of Fort Sill, Wichita Mountains, Oklahoma.
Drumaspis clara; New: SPECIES... .c000 5 42 2 see bie oe ee ane Ws ee Se 33
23. Libragene (X 3).
24-26. Front, side, and dorsal views of the holotype cranidium (xX 2).
Honey Creek limestone; (loc. 91s) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Drumasprs tex ana, NEW SPECIES W..6c., set eke -eonels wisiretererel ev aertetiastetotie ieee 32
27, 28. Dorsal and side views of a large cranidium (X 6).
29. Holotype cranidium (X 5).
30. A small cranidium (X 5).
Wilberns formation; (loc. 70) Baldy Mountain, 8 miles northwest of
Burnet, Texas.
Drumaspis utahensis, NOW *SPECIES «2's es cis wale slacatsielpe se cosets eee eee 34
31-33. Side, front, and dorsal views of the holotype cranidium (xX 6).
St. Charles limestone; (loc. 54x) Two Mile Canyon, 2 miles south of
Malad, Wasatch Mountains, Idaho.
Drumaspis ‘witida, New: SPECIES «2%. s/s <lnpers <inja 30 vine bl care wisie eine = be eee 34
34-35. Side and dorsal views of the holotype cranidium (X 5).
Honey Creek limestone; (loc. 91s) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Cheilocephalus texanus, New: SPECIES... cn eeiicle is eteem eer eee eee 35
36-38. Front, side, and dorsal views of the holotype cranidium. (36 and
37, X 3;_38, X 6.)
Hickory sandstone; (loc. 68z) Packsaddle Mountain, 11 miles south-
east of Llano, Texas.
Chetlocephalus wichitaensis; new SpeCieS:. «6. cs eee ete ees eae 35
39. Holotype cranidium (x 2).
Honey Creek limestone; (loc. 9p) 15 miles northwest of Fort Sill,
Wichita Mountains, Oklahoma.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 5, PL. 5
UPPER CAMBRIAN TRILOBITES
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOL. 103, NO. 5, PL. 6
UPPER CAMBRIAN TRILOBITES
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER TT5
PLATE. 6
MARES METAACHSES, MEW SPECIES |) << exces 20s a's sialeieig sv a cla vials tiaeisyeuje wa edu ne 36
I, 2. Dorsal and side views of holotype cranidium ( 2.5).
Hamburg limestone; (loc. 23d) near Hamburg Mine, Eureka District,
Nevada.
ROMA URMNRTIFESUNES, TICW SPECIES i550 2c 2 o's =. 2's 0'e 5.4m wrinis «ce erm ia aie nicia Say oie Sm iets 38
3, 4. Cranidia. (3, X 3; 4, X 5.)
5, 6. Small cranidium (x 5).
7. Holotype cranidium (X 3).
Lyell formation; (locs. 66j, k) Ranger Canyon, Sawback Range,
Alberta.
WOES RCIOCrECHSHS:A NEW. SPECIES. .o'..3 «os scine cides cn Ge scence sce ewe 38
8. Libragene ( 2).
9. Pygidium (X 2).
10, 11. Dorsal and side views of a partially exfoliated cranidium (> 2).
12. Holotype cranidium (2). (See also pl. 14, fig. 18.)
Lyell formation; (loc. 20d) Tilted Mountain Brook, 9 miles east of
Lake Louise, Alberta.
Minenures (©), sinclairensts, New SPECIES... ......50.506.c0ccescceeesessess 39
13, 14. Dorsal and side views of the holotype cranidium (X 2.5).
15. Two associated cranidia (5).
Sabine formation; (loc. 16t) Sinclair Canyon, Brisco Range, British
Columbia.
PCO MNUS, IINMOLSIS: TICW SPECIESs co c.ciam oc asniesisasde sc bic ved sew eraseses 40
16, 17. Dorsal and side views of the holotype cranidium (X 1.5).
Mendha limestone; (loc. 7j) Quinn Canyon Range, Nevada.
MECICROMGAIUS (VV AICOEL))... chee eh coke gees s oa oe tal ta 6 ease va gies ace 40
18, 19. Side and dorsal views of the holotype cranidium (> 5).
McKay group; (loc. 23z) 2 miles west of Donald, Dogtooth Mountains,
British Columbia.
PAaivcolpus orlahomensis, NEW, SPECIES... 2. soa. sacs sees sess ese ccs ees 40
20, 21. Incomplete cranidium (X 1.5).
22, 23. Exfoliated pygidium.
Signal Mountain formation; (loc. 201j) Royer Ranch, Honey Creek,
Arbuckle Mountains, Oklahoma.
24, 25. Side and dorsal views of the holotype pygidium.
Signal Mountain formation; (loc. 186s) 4 miles northwest of Ravia,
Arbuckle Mountains, Oklahoma.
PQICOIPUS WHChiLaensis, NEW SPECIES... .. 2.2... scene dene ence nena aeces 41
26, 27. Dorsal and side views of the holotype pygidium.
28, 29. An incomplete cranidium.
Signal Mountain formation; (loc. 12g) 2 miles southwest of Signal
Mountain, Wichita Mountains, Oklahoma.
Remricoipus highlandensis, Tew SPECIES... 665 nc ccc ee ee cane ee cus eeesesa 4I
30, 31. Squeeze of the holotype pygidium.
Mendha limestone; (loc. 88) 7 miles north of Bennett Springs, High-
land Range, Nevada.
RNEVECOUPUSPMINCIOINCNSIS: IEW SPECIES)..)<ci-t1 viele oe de c.s.cle edie sarees so eles ee oe 42
32, 33. Side and dorsal views of a large pygidium.
34, 35. The holotype pygidium.
36. Libragene.
Sabine formation; (loc. 16t) Sinclair Canyon, Brisco Range, British
Columbia.
PUM EOUT ONLY PIC MILS* STEW: SPECIES 1< «15.4 o0e1h eravaieinia one's yanie bkels eles sing s'sleved 43
37, 38. Side and dorsal views of the holotype pygidium.
Mendha limestone; (loc. 88) 7 miles north of Bennett Springs, High-
land Range, Nevada.
110 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
PLATE 7
PAGE
Macelloura transversa, n€w: SPCClES. «cece. snc clasiis cleeleles ocislt siete sitio le tials 43
1. Holotype pygidium (X 1.5).
Wilberns formation; (loc. 14f) Bartlett Hollow, Burnet County, Texas.
Arapahota. stantoni, new SPECIES. % wince sic p= ae euls sete wee nine alo mieten e's eee mae 44
2. Holotype cranidium.
3) 4. Pygidium) (<2):
Pilgrim formation; (loc. 26b) Mill Creek, Little Rocky Mountains,
Montana.
Arapahota veesidcet, NEW SPECIES. =,.ctew sce sacl scie as ola feel eters ale igiatetetieietetae 44
5. Pygidium (X 1.5).
6, 7. Dorsal and side views of the holotype cranidium (X 2).
8, 9. Another cranidium (X 2.5).
Maurice formation; (loc. 26c) 6 miles south of Hailey, Wyoming.
Arapahota polita,. new SPECIES «cect amas sila, asinicierays en «cere eelw Oseoie atelataiee 45
10, 11. Dorsal and profile views of the holotype cranidium ( X 1.5).
Sullivan formation; (loc. 64h) East Lyell Glacier, 48 miles northwest
of Lake Louise, Alberta.
Arapahowa walcotiae, new. SPCCieS «uc. oie. ois 0 one onic aiesele oles etn smn alm nln elaine 46
12, 13. Dorsal and side views of the holotype cranidium (X 1.5).
14. A smaller, exfoliated cranidium (X 2).
Sullivan formation; (loc. 64e) East Lyell Glacier, 48 miles northwest
of Lake Louise, Alberta.
Arapahoia: elongata, new SPECIES. cis .aia 0. oraen'e fa s'« sonia aialers yale ie /feeieler = alerannian 45
15, 16. Dorsal and side views of the holotype cranidium ( X 1.5).
17. Libragene (1.5).
18, 19. Pygidium (1.5). (See also fig. 32.)
Sullivan formation; (loc. 64h) East Lyell Glacier, 48 miles northwest
of Lake Louise, Alberta.
Arapahota prolixa, NeW - SPECIES 2. cis te oietenys olan ppieia wiaineis eine aes eee 46
20, 21. Side and dorsal views of the holotype cranidium (X 1.5).
22, 23.. Pygidium: (< 1.5).
24, 25. A partially exfoliated cranidium (X 2).
26. Libragene (XX a
Sullivan formation; (loc. 64h) East Lyell Glacier, 48 miles northwest
of Lake Louise, Alberta.
Arapahoia albertensis, NEW SPECIES. .......00ccccccccccceccucnccsercnccas 46
27. Cranidium (X 3).
28, 29. Side and dorsal views of the holotype cranidium (X 3).
30. Side view of unexfoliated cranidium (x 3).
31. Libragene (X 1.5).
32. eee and libragene of A. elongata (X1.5). (See also figs.
17-10.
Sullivan formation; (loc. 64h) East Lyell Glacier, 48 miles northwest
of Lake Louise, Alberta.
Kingstoma electra, new Species. . 0.0 4s 6. cists me mms mains eclait Sn clei eerste 47
33, 34. Dorsal and side views of the holotype cranidium (x 5).
35, 36. A partially exfoliated cranidium (X 5).
Weeks formation; (locs. 30h, i) 5 miles south of Marjum Pass, House
Range, Utah.
Avapahota ty pa. Miller 5... doit <.ccs eae ata were ee mena cals s ane ee 44
37. Cranidium and pygidium (1.5).
38. Libragene and pygidium (X 1.5).
Maurice formation; Butte north of Little Rocky Creek, Beartooth
Range, Wyoming.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 5, PL. 7
Me Selene gel aay aes pat Ga STE
oF sea.
1
5
»
:
UPPER CAMBRIAN TRILOBITES
SMITHSONIAN MISCELLANEOUS COLLECTIONS
UPPER CAMBRIAN TRILOBITES
VOL. 103, NO. 5, PL. 8
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 17,
PLATE 8
PIE SEO MAR MUCHO! TIC Wr SPECIES. /<isjace-4 tierojie crotote level ee ers aie eves mete Breve eikae 48
I, 2. Large cranidium (X 2.5).
3, 4. Dorsal and side views of the holotype cranidium ( 4).
5, 6. Pygidium (X 1.5).
7, 8. An exfoliated pygidium ( 1.5).
Sherbrooke limestone; (loc. 58f) ridge west of Mount Bosworth,
British Columbia.
eOSTOMICEVOSWOTLLENSIS. NEW: SPECIES. «5.5042 242 sci as ccc einen econ seals we 48
9, 10. Small cranidium (5).
II, 12. Side and dorsal views of the holotype cranidium (x 5).
13. Pygidium ( 4).
Sherbrooke limestone; (loc. 57p) ridge west of Mount Bosworth, British
Columbia.
aM STOM(GTODSOMENSIS, NEW. SPECIES» «15:00 00,0.6.50.</1+ 4-0) 0 lela caus oles eke bP eee oe 49
14, 15. Dorsal and side views of the holotype cranidium (X 4).
Sullivan formation; (loc. 61r) Moose River, 10 miles northeast of
Robson Peak, British Columbia.
EM RUNCOMUU SUIIDORENSIS, TIEW SPECIES. ...c. sks eessec cea vee ce awosstcece 49
16, 17. An imperfect cranidium (X 4).
18, 19. Dorsal and front views of the holotype cranidium ( 4.5).
Sullivan formation; (loc. 64m) Sullivan Peak, 48 miles northwest of
Lake Louise, Alberta.
LGM OSEONIG: POMISSA, TIEW. SPECIES). «22% c0s coc escc ster ses cee cewesoes wees 49
20, 21. Dorsal and side views of the holotype cranidium ( 4.5).
22, 23. Pygidium (x 4).
Sullivan formation; (locs. 64c, j) Glacier Lake Canyon Valley, 48
miles northwest of Lake Louise, Alberta.
EO SLOMUTMO PCH aNeW) SPCCleSrucesctin sc aie 40 ls oiwe in eeele od ooes. oe ee aioe 50
24, 25. Side and dorsal views of the holotype pygidium (X 3).
Sawatch formation; (loc. 6a) Taylor Peak, 4 miles south of Ashcroft,
Colorado.
Bere SLOMIRUULG GLE. TIEW: SPECIESs ere mc) ais ese <.s:Alctiwl cowie, stale elbow cies s 4 eee 50
26, 27. Pygidium ( 4).
28, 29. Side and dorsal views of the holotype cranidium ( 4).
30. Exfoliated cranidium (X 3).
Sullivan formation; (loc. 64b) Glacier Lake Canyon Valley, 48 miles
northwest of Lake Louise, Alberta.
rena aNE Ge Ds PICMIL, NEW SPECIES 2. <1 ob. Pehitidiiela es wlow s ven celeeews 0 deplete 50
31, 32. Dorsal and side views of the holotype pygidium (X 1.5).
Sullivan formation; (loc. 64h) East Lyell Glacier, 48 miles northwest
of Lake Louise, Alberta.
ERISA Gre) RIOCESEG, MEW, SPECIESi.o.4, sete c's csc, ols o-0is elneneg so Sislaueevece ove 2 55
33, 34. Side and dorsal views of the holotype cranidium (5).
Sullivan formation; (loc. 64h) Glacier Lake Canyon Valley, 48 miles
northwest of Lake Louise, Alberta.
MILLEN (G7) EP OGWCLO. NEWe SPECIES. =/sjjcicieiiclcwiace Sersies ciel sitla « weave meee ale 56
35, 30. Dorsal and side views of the holotype cranidium ( 5).
Sullivan formation; (loc. 641) Glacier Lake Canyon Valley, 48 miles
northwest of Lake Louise, Alberta.
PRO MAPUSL HILO LEDS? TIEW! SPECIES, ove sos) «i sfare 416) = siricigincis-s © 0:5 cleyn e) otia aye wiele miele aerere 56
37, 38. Dorsal and side views of the holotype cranidium (x5).
30, 40. Somewhat distorted cranidium (5).
41. Pygidium (> 3).
Sullivan formation; (loc. 64h) East Lyell Glacier, 48 miles northwest
of Lake Louise, Alberta.
EMO STOMA As ONVialGOLE his cerca caters. oc iee a Sone orev athe neunletald ised Hatolstotees oltre 51
42. Partially exfoliated cranidium (XX 3).
43, 44. The exfoliated holotype cranidium (5).
45, 46. Complete paratype cranidium (> 6).
Warrior limestone; (loc. 24f) 1 mile southeast of Warriors Mark,
Pennsylvania.
118 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
PLATE 9
King stonia. Rindlet, new species... .0c.su5 vatecss osvew halen ox ait emma ene 51
1, 2. Dorsal and side views of the holotype cranidium (X 3).
3, 4. Two pygidia (X 5). } é
Warrior limestone; (loc. 38e) 1 mile northwest of Benore, and 5 miles
west of State College, Pennsylvania.
Bynumia eumus Walcott cc ccc 5.5 howe decals ee oe beeclamemioclee ie neler 52
5, 6. Dorsal and side views of the holotype cranidium (X 3).
7. Pygidium (X 3). :
Sullivan formation; (loc. 64b) head of Glacier Lake Canyon Valley,
48 miles northwest of Lake Louise, Alberta. |
Bynumia elegans: NewisSPeCies'.,.%. wssecseloeieie tne sts eroidinie ele Gleerareee ae eens 52
8, 9. Dorsal and side views of the holotype cranidium (X 6).
10. Smaller cranidium (X 3).
TY, 12) Dworpyridia lOc aye
Sullivan formation; (loc. 64r) Ranger Brook Canyon, Sawback Range,
Iberta.
Bynumtavarguia,, MEW SPeChesaacrets ise vine clerniss eisis eens e cie etter ile 53
13, 14. Side and dorsal views of the holotype cranidium (5).
15. The associated pygidium ( 3).
Sullivan formation; (loc. 66s) Badger Pass, Sawback Range, Alberta.
Bynumia wolcottt, new: SpECleS. avian a3 nditoel sales oapeaeias es hare eee 53
16, 17. Side and dorsal views of the partially exfoliated holotype
cranidium (X 3).
18, 19. An exfoliated cranidium figured by Walcott as B. eumus (XX 3).
20. The associated pygidium ( 2.5).
Sullivan formation; (loc. 66m) 5 miles northwest of Banff, Sawback
Range, Alberta.
BYynumia, VENUSTA, NEW! SPECIES. Wi Mae vad oe eee ae en eee rie oe nee 54
21, 22. Side and dorsal view of the holotype cranidium (X 3).
23. Libragene (X 5).
24-26. Three pygidia (xX 3).
27, 28. A somewhat variant cranidium (X 2.5).
Sullivan formation; (loc. 64s) Ranger Brook Canyon, Sawback Range,
Alberta.
Bymamiella: briscoensts; new SPECIESs. «sass: fess > coset ceeee ous eee 57
29. Holotype cranidium (5).
Sabine formation; (loc. 16t) Sinclair Canyon, Brisco Range, British
Columbia.
IBIS IT HOd AOS A Se SHANE STU oon ddan soadoounodnbadesddahuoasod cour 54
30, 31. Side and dorsal views of the holotype cranidium (> 4).
32, 33. An exfoliated cranidium (X 6).
34. An exfoliated pygidium (> 3.5).
Lynx formation; (loc. 19m) Iyatunga Mountain, Mount Robson,
British Columbia.
Bymunia! sulcatas news Spectese aanaesece eee nee ee ERE 54
35, 30. Side and dorsal views of the holotype cranidium (> 5).
Sullivan formation; (loc. 64s) Ranger Brook Canyon, Sawback Range,
Alberta.
Bynumia ranger énsis; New Species...) ssi. 2. Hees. eae dee eases See 55
37, 38. Side and dorsal views of the holotype cranidium (X 3).
39, 40. An exfoliated cranidium ( 3).
41. Pygidium ( 2).
Sullivan formation; (loc. 66L) Ranger Brook Canyon, Sawback Range,
Alberta.
Bynumia sawbackensis, new Species s.chi..csice se ehese reas eee eee 55
42, 43. Side and dorsal views of the holotype cranidium ( 3).
44. Pygidium ( 2).
ee formation; (loc. 64w) Ranger Brook Canyon, Sawback Range,
erta.
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOL. 103, NO. 5, PL. 9
ee) ee et tk ee
a
PR SO Te
f; UPPER CAMBRIAN TRILOBITES
VOL. 103, NO. 5, PL. 10
SMITHSONIAN MISCELLANEOUS COLLECTIONS
SE eee
- SR
UPPER CAMBRIAN TRILOBITES
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER I19
PLATE 10
PAGE
PaPHCLIC OEY PICGIES, TOW SPECIES»)... 6/50 vice va wisleje cece abs eislnmee wens 57
1. Holotype cranidium (X 4).
2. A larger cranidium (5).
Lyell formation; (loc. 66j) Northeast branch Ranger Brook Canyon,
10 miles northwest of Banff, Sawback Range, Alberta.
Giplonilusa(? valbertensis, NEW SPECIES Ss 5... + cjenie che cele elec ese eieas 37
3. Holotype cranidium (X 2).
Lyell formation; (loc. 64b) Glacier Lake Canyon Valley, 48 miles
northwest of Lake Louise, Alberta.
MCD OUMOMEGEL ALG NEW, SPECIES co. é.6:0,6 s1a,¢ one(seo 46 le ac sie a e onate eae steele slaenie a6 107 |
4. Holotype cranidium (XX 2.5).
Nolichucky shale; (loc. er U. S. 11, 7 miles northeast of Rogersville,
Tennessee.
Ellipsocephaloides argutus, MEW SPECIES... ......cc cc ccecc cess cescccecsese 63
5. Imperfect cranidium (> 3).
6. Holotype cranidium (X 4).
Sabine formation; (loc. 12s) Ram Creek, 15 miles south of Canal Flats,
British Columbia.
Ellipsocephaloides briscoensts, n€W SPeCi€S........ccccecccccccceccccceess 63
7. Holotype cranidium (X 2.5).
Sabine formation; (loc. 17s) Sabine Mountain, Brisco Range, British
Columbia.
RI EHO MISH AGC MNEW: SPECIES craetn « casio clea ee eisie's as deel h ue owls cs slere ote vele 50
8, 9. A large cranidium (X 2).
10, 11. Dorsal and side views of a young cranidium (X 6).
12. Partially exfoliated cranidium (X 4).
13. Holotype cranidium and a smaller example ( 3).
Signal Mountain formation; (locs. 12i, j) 2 miles southwest of Signal
Mountain, Wichita Mountains, Oklahoma.
EHOPIMUSHOMUNCUS, NEW SPECIES... . o2% occiscc neve ves dele s dass cieles oles 60
14, 15. Dorsal and side views of the holotype cranidium ( 1.5).
16, 17. Partially exfoliated cranidium (X 1.5).
Signal Mountain formation; (loc. 12L) Pickens Ranch, Arbuckle
Mountains, Oklahoma.
Ey PUMNG CACIGIG, MEW SPECIES. 0. 5... ie eee ce sees ste feet cbt estes 58
18-21. Dorsal and side views of two cranidia ( 4).
22. Holotype cranidium and associated pygidium (X 4).
Davis formation; eee 92d) Federal Lead Mine No. 4, Flat River,
Missouri.
MUU ILSS OUNICISIS STEW, SPECIES sa. 0./ac1)s ciccicte cioicis eee see ee wei veces 50
23, 24. Dorsal and side views of the holotype cranidium (5).
25. Another cranidia (xX 4).
26. Surface with several cranidia (x 6).
Davis formation; (loc. 11k) Flat River, Missouri.
Bynumiella (?) oklahomensis, new SpecieS.............ceececccceeccces 58
27, 28. Side and dorsal views of the holotype cranidium (5).
Signal Mountain formation; (loc. a01j) 1 mile south of Royer Ranch,
Arbuckle Mountains, Oklahoma.
I20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
PLATE ii
Eltipsocephaloides, silvestris: new SpecieSa: .cisct isles isietinie ieiele el nel siete aitorereate 64
I. Cranidium (xX 4).
2. Holotype cranidium (> 4).
3. Two pygidia with which are associated the base of a cranidium and
a libragene of Ptychaspis (* 4). (See also pl. 12, fig. 7.)
Honey Creek limestone; (loc. 91b) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Ellipsocephaloides sawbackensts, NeW SPECIES... .....0.2cceescceccescsoss 64
4. Holotype cranidium (> 5).
5. Large incomplete cranidium (5).
Lyell formation; (loc. 64x) Ranger Brook Canyon, Sawback Range,
Alberta.
Cheilocephaluss buttsi,. mew? SpeGleseaaashehiasee eee eee eee 36
6. Holotype cranidium (X 1.5).
Ore Hill limestone; (loc. 107v) 4 mile northwest of Drab, Pennsylvania.
Ellipsocephaloides carus; New Species. . 2... 0.6 .s2k esas ee eae 65
7. Cranidium (5).
8. Holotype cranidium (X 5).
Sabine formation; (loc. 12s) Ram Creek, 15 miles south of Canal Flats,
British Columbia.
Ellipsocephaloides montis, new Species... clan «ise ssle este a oes ee 65
9, 10. Dorsal and side views of the holotype cranidium (X 2.5).
11. A small cranidium (> 5). ~
Both are associated with Pseudagnostus.
Sabine formation; (loc. 12s) Ram Creek, 15 miles south of Canal Flats,
British Columbia.
Ellipsocephalowdes, bearensis, new Species... a -eeeee aces cee cee enien cee 65
12. Holotype cranidium (X 2).
St. Charles limestone; (loc. 66z) 5 miles west of St. Charles, Bear
River Range, Idaho.
Elhpsocephaloides nttela, new Species... 0s. .stns ches cases cteic ae eee eee 66
13. Several cranidia (xX 4). (See also pl. 12, figs. 1-3.)
Honey Creek limestone; (loc. 12m) 7 miles north of Springer, Arbuckle
Mountains, Oklahoma.
Ellipsocephaloides butlert, tlew SPECleS: <). Us sme ence + dese ee eee ee 66
14, 15. Cranidia (XX 2.5). Upper right cranidium of fig. 14 is the
holotype.
Sawatch formation; (loc. 37x) near Gilman, Mosquito Range,
Colorado.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 5, PL. 11
2%,
UPPER CAMBRIAN TRILOBITES
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 5, PL. 12
UPPER CAMBRIAN TRILOBITES
PRATIC 12
ALE PAGES MITCIG MEW. SPECIES; oc ix/eie'nm swirls an ewisc coe eine pe vee siecle as
1. Holotype cranidium (5).
2. Pygidium (x 6).
3. Pygidium with full complement of spines, and cranidium (X 3).
(See also pl. 11, fig. 13.)
Honey Creek limestone ; (loc. 12m) 7 miles north of Springer, Arbuckle
Mountains, Oklahoma.
Ellipsocephaloides monsensis, ME WaiS DECIESH eT are ei ieiaicrs Oise oateictoee Soe
4. Holotype cranidium (X 5).
5, 6. The pygidium (X 3
Lyell formation; (loc. Bat) Mons Glacier, 50 miles northwest of Lake
Louise, Alberta.
Piesocepnaloides Silvestris, NEW SPeClES: 2... 0.05.0 siee ccs ew enw ae be acess
7. Well-preserved pygidia (5). (See also pl. 11, fig. 3.)
Honey Creek limestone; (loc. 91b) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Gala pian LV PICGle NEW SPECIES «2 =o /\a( adie e/eie) «0 a o)sie susie oo ele elm elec sierereters
8. Cranidium (> 4).
9. Holotype cranidium (X 4).
Red Lion formation; (loc. 150e) Boulder Creek, 1 mile north of Prince-
ton, Montana.
Ener niu Ones AEChiUES, NEW SPECIES. x .. s.o.aiee ocd 0's ceisinnsels oie eeieiva dois
to. Cranidium (X 4).
11, 12. Dorsal and side views of the holotype cranidium (5). (See
also fig. 13.)
ICEMOMELO PUS GIOETIENSIS® MEW. SPECIES. ..2...0,0 cle ssc hs siee oed eels elec nen
13. Holotype cranidium, with cranidia of Ellipsocephaloides dechvis
(see figs. 10-12) and Chariocephalus magnus (see pl. 2, fig. 15).
ae)
Lyell formation; (loc. 20j) Tilted Mountain Brook, 94 miles east of
Lake Louise, Alberta.
Meg T OME UYOMANRGERSIS: (AIRESSET i .2g csole css cce ss ae citsemsche das sm@enane
14, 15. Dorsal and side views of the holotype pygidium (1.5).
Pilgrim formation; Dead Indian Creek, Clark Fork, Absaroka Range,
Wyoming.
16, 17. Side and dorsal views of an exfoliated pygidium (1.5).
18. Partially exfoliated cranidium ( X 1.5
Pilgrim formation; (loc. 4r) Suce Creek, ee Delano, Snowy Range,
Montana.
19. Cranidium (XX 1.5).
20. Pygidium (X 1.5).
Pilgrim formation; (loc. 151g) mouth of Clark Fork River, Absaroka
Range, Wyoming. (See also pl. 14, fig. 1.)
VAN PAU THO IQENSIS MCW: SPECIES vais yaetel<\eieieai io eysichs 4, sis sic ols are, clele sweustelaie ls
21, 22. Dorsal and side views of a cranidium. (21, * 1.5; 22, X 3.)
23. Partially exfoliated cranidium (XX 2.5).
24. Partially exfoliated pygidium (X 1.5).
25, 26. The holotype pygidium (X 1.5).
Maurice formation; (loc. 151k) pees Creek, near Viola, Uinta
County, Wyoming. .
MEGPYONMG WLANENSIS, NEW SPECIES... ose eee ne hence teen c en em neces
27, 28. Dorsal and side views of the cranidium ( 2).
29. Libragene (X 2).
30. Hypostoma (X 5).
31. Holotype pygidium (X 1.5).
32. Incomplete pygidium (X 1.5).
Weeks formation; (loc. 32w) Fandango Spring Canyon, Dugway
Range, Utah.
Maryvillia marjumensts, NEW SPECIES... 2.00.00 cnc cnccscsccsccdcesccnce
33. Hypostoma (X 3).
34. oes pygidium associated with an exfoliated cranidium
125 )))-
35. Associated cranidium and pygidium (> 1.5).
Weeks formation; (loc. 301) 5 miles south of Marjum Pass, House
Range, Utah.
MCN LO DErU ANE We SPECIES» e:ctayaycssieia «onze sien oieke ae skal dierel sé sleloldsieie eae e Simic ie
36. Cranidium (X 1.5
37, 38. Side and aoe views of the holotype pygidium.
Sawatch formation; (loc. 6a) Taylor Peak, 4 miles south of Ashcroft,
Colorado.
121
66
64
38
68
68
70
I22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
PEATE 13
: PAGE
Maryuillia alberia, new SpeCi€s): 2 \. 2-65 ynice once k = echo ne waste eine eee 70
1, 2. Dorsal and side views of the holotype cranidium (X 2).
Sullivan formation; (loc. 641) Glacier Lake Canyon Valley, 48 miles
northwest of Lake Louise, Alberta.
3, 4. A pygidium referred to species (X 1.5).
Sullivan formation; (loc. 64c) Glacier Lake Canyon Valley, 48 miles
northwest of Lake Louise, Alberta.
Meteoraspis ban ffensts, New SPECIES... <si.5 00.2 ctewcnass eran oes cece cemenies
5, 6. Dorsal and side views of the holotype cranidium (X 2).
z, 8. An exfoliated cranidium (XX 2).
9, 10. Pygidium (x 4).
Sullivan formation; (loc. 66m) 4} miles northwest of Banff, Sawback
Range, Alberta.
Maryvillia montis; news SPeCleSi scien loxersie oherever tala ncciere sie iorssctokeiaistoler rier inetenets
11. Cranidium (X 2).
12, 13. Side and dorsal views of the holotype pygidium.
Deadwood formation; (loc. 17L) 4 mile west of Deadwood, Black
Hills, South Dakota.
Maryuillia: hybrida, new SPeCieS: os.c3 6. saz tin wcisiev eek Ente eee ae ees
14, 15. Cranidium (X 1.5).
16, 17. Dorsal and side views of the holotype pygidium (1.5).
Cap Mountain formation; (loc. 14d) Bartlett Hollow, Burnet County,
‘Nexas:
Goosta’ canadensis “new, SPeCleSaece ena Aerie ee eee eee eee
18, 19. Dorsal and side views of the holotype cranidium ( 1.5).
20. Pygidium (X 2).
Sullivan formation; (loc. 641) Glacier Lake Canyon Valley, 48 miles
northwest of Lake Louise, Alberta.
Coosellatexand, New, SPECIES) /< Naveore ciarote cioiteniels tre nie Dae te rnie ReEEeeone
21, 22, Pyeidium, (1-5).
23, 24. Dorsal and side views of the holotype cranidium (1.5). (See
also pl. 14, figs. 2-5.)
Cap Mountain formation; (loc. 14d) Bartlett Hollow, Burnet County,
Texas.
Coosiavalbertensis; new; SpeCleSi'=- nse: ese meee ee ee One eee
25,26, Pyeidium:, (25; 5623/20) os.)
27, 28. Side and dorsal views of the holotype cranidium (1.5).
Sullivan formation; (loc. 65i) Glacier Lake Canyon Valley, 48 miles
northwest of Lake Louise, Alberta.
Coosta dakotensis, New. SpeCieS. .c0 nance orice ce coat telstoere oe erie Recents
29, 30. Side and dorsal views of the holotype cranidium (X 1.5).
31. Pygidium, impression of the cranidium, and libragenes showing
the wide doublure (1.5).
Deadwood formation; (loc. 17j)) Galena, Black Hills, South Dakota.
72
71
Fu
73
73
74
SMITHSONIAN MISCELLANEOUS COLLECTIONS
UPPER CAMBRIAN TRILOBITES
VOL.
103, NO: '5, PL, 13
SMITHSONIAN MISCELLANEOUS COLLECTIONS
40
UPPER CAMBRIAN TRILOBITES
VOL. 103, NO. 5, PL. 14
PLATE 4
mined ieyomingencsts (CIRESSEL ).... csc deine seems sie clencs Secs eee cess 67
1. Partially exfoliated cranidium associated with the holotype of
Tricrepicephalus tripunctatus (Whitfield). (See also pl. 12, figs.
14-20. )
Pilgrim formation; near Moss Agate Springs, Castle Mountains,
Montana.
Oe LEO IEW. SPECIES ee ayels ts iooia ls orcesiale wre nls! Cicine Die eee Mae Sore niece 75
2, 3. Dorsal and side views of a nearly complete cranidium.
4, 5. Portion of pygidium. (See also pl. 13, figs. 21-24.)
Cap Mountain formation; (loc. 67a) Potatotop, 7 miles northwest of
Burnet, Texas.
MENG D LADIS ATIGW SPECIES)» o> shiveic ss ais: carers ti caters ora nie in erecnee ae elon ea 76
6, 7. Dorsal and side views of the holotype cranidium ( 3).
8, 9. The pygidium (X 2).
Warrior limestone; (loc. 38e) 1 mile northwest of Benore, and 5 miles
west of State College, Pennsylvania.
Patennee OG) eesparplis, NEW SPECIES 25.6626... oe cane cee en eaewdde eee 61
10, 11. Side and dorsal views of the holotype cranidium. (10, * 2.5;
ig)
Sullivan formation; (loc. 64h) Glacier Lake Canyon Valley, 48 miles
northwest of Lake Louise, Alberta.
aN COIPADE RCM LLC Wat SP ECIESt or. Siaicuce, o:d-S oare-oiy ood Sota Sake le sublets ete Se mae 60
12. Somewhat distorted pygidium (X 1.5).
13, 14. Dorsal and side views of the holotype pygidium ( 1).
Sullivan formation; (loc. 64c) Glacier Lake Canyon Valley, 48 miles
northwest of Lake Louise, Alberta.
MV CIELO O SENSISE MEW) SPECIES <i ja)2t2 eis ccicve «sets @ oppo lees elele solves noe cc 72
15. Holotype cranidium (X 2.5).
16, 17. Side and dorsal views of a pygidium (X 1.5).
Lynx formation; (loc. 61r) drift on Moose River, 10 miles northeast of
Robson Pass, British Columbia.
MGe nue sealbertensis; MEW SPECIES... <liaccecmie ce vses svavscvecscescescedds 38
18. Exfoliated cranidium (X 2.5). (See also pl. 6, figs. 8-12.)
Lyell formation; (loc. 20d) Tilted Mountain Brook, 9 miles east of
Lake Louise, Alberta.
NRE LO MGLDERIGS NEW) SPECIES sacle f csesiccc ois a aisle oom s+ Gisid o cie iela due cate eis ele 61
19, 20. Partially exfoliated cranidium (X 2.5).
21. Longitudinally shortened cranidium (> 3).
22-24. Dorsal, side, and front views of the holotype cranidium ( 3.5).
25, 26. Partially exfoliated cranidium ( 3).
Sullivan formation; (locs. 64b, c) Glacier Lake Canyon Valley, 48
miles northwest of Lake Louise, Alberta.
EME CERO LE Le Me We SPECIES: orice Lows caieind ceiceloesis © ons neite soo sca@eeba ec 60
27. Pygidium retaining most of test (X 3.5).
28, 29. Large pygidium (X 2).
30, 31. Dorsal and side views of the holotype cranidium (> 5).
Warrior limestone; (loc. 38e) 1 mile northwest of Benore, 5 miles west
of State College, Pennsylvania.
EVEROCEPRALING TExONG, NEW, SPECIESs. c+ aca cicud iw se clesieeulo deco eere secs 76
32, 33. Side and dorsal views of the holotype pygidium (X 2).
Wilberns formation; (loc. 69) Honey Creek, 8 miles southeast of
Llano, Texas.
ERCROGEIMULMIMOLNOENG) NEWSPECCLES Cs 20 eise esis bce oassles aa clos se dae sine alee eteels 77
34. Cranidium and holotype pygidium.
35, 36. Small cranidium (x 2.5).
37, 38. Partially exfoliated cranidium.
Mendha limestone; (loc. 7j) 1 mile north of Italian Ranch foothills,
north end of Quinn Canyon Range, Nevada.
izterocepnaina vilobata (Halland! Whitheld)). >.0.55.0-0-4.2.-5-.52-00-- 77
39, 40. Incomplete cranidium.
41, 42. Small pygidium ( 2).
43. Pygidium of normal size.
Secret Canyon shale; (loc. 61) south of Hamburg Mine, Eureka District,
Nevada.
123
124 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
PLATE 15
Pterocephalina pogonipensts, NEW SPECIES. ......020sccccvercscsccevucaceee 78
1. Cranidium and holotype pygidium ( 2).
2. Side view of the holotype (X 2).
Secret Canyon shale; west side of Pogonip Mountain, White Pine
District, Nevada. .
Pterocephalina grata, new SPECIES... 05 sea 2 oe poe udu a niw orsie es ee 78
2. 4. Cranidium (<3):
5, 6. Side and dorsal views of the holotype pygidium ( 2).
Secret Canyon shale; (loc. 23b) ridge east of Hamburg Ridge, Eureka
District, Nevada.
Pterocepholina: utahensis, new: Species.)... n-ne sine eee ss eens 70
7. Large pygidium (X 1.5).
8, 9. Cranidium (X 2).
10, 11. Dorsal and side views of the holotype pygidium (> 2).
Orr formation; (loc. 33d) east side Fish Spring Range, Utah.
Coosia tridentensis: new. Species... canes oe ace oe eee Eee hse
12. Small pygidium (X 1.5).
13. Cranidia and pygidium (> 1.5).
14, 15. Holotype cranidium (X 1.5).
16. Incomplete cranidium (X I aa
17. Impression of pygidium ( 1.5).
Pilgrim formation; (loc. 20y) 3 miles north of Trident, Montana.
Berketa comes, new. Species 25s). acre was 2 oe id ao) cretute alas eee ata 90
18, 19. Incomplete cranidium (X 3).
20, 21. Dorsal and side views of the holotype cranidium ( 3).
Deadwood formation; (loc. 88a) Deadwood, Black Hills, ear Dakota.
Berketa ‘saratogensis, MEW SPeCieS ies «aon ec. edaae cine see eens eee OI
22, 23. Side and dorsal views of the holotype cranidium. (22, X 1.5;
23, X 3.)
24, 25. Two cranidia (1.5).
Potsdam sandstone; Greenfield, northwest of Saratoga, New York.
Berkeia*nevadensts. new, Species. sen a-indica eee eee eee OI
26. Holotype cranidium ( 2.5).
27. Cranidium (> 2.5).
Secret Canyon shale; (loc. 61) south of the Hamburg Mine, Eureka
District, Nevada.
Berkéta retusa, new, iSpecieSiccucit dangers oien oe tO ee ee 92
28, 29. Side and dorsal views of the holotype cranidium (> 3).
30. Cranidium (XX 3).
Honey Creek limestone; (loc. 12p) 4 miles southeast of Hennepin,
Arbuckle Mountains, Oklahoma.
Berkeia wichitaensis, new species aigi'-a 9530 (6 Space ‘Syeaepade Poh ate rasan stetaterensteiasioreee ae teete 92
31, 32. Side and dorsal views of the holotype cranidium ( 3).
33- pyaiitts ($3).
Honey Creek limestone; (loc. 91L) Big Baldy, Wichita Mountains,
Oklahoma.
Berkeia angustata, new SPECIES 55.4 fol ejasratomedeon meets athe sae eile eee 93
34, 35. Side and dorsal views of the holotype cranidium (X 3).
36-39. Two cranidia (3
Honey Creek limestone; (loc. Soy) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 5, PL. 15
UPPER CAMBRIAN TRILOBITES
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 5, PL. 16
UPPER CAMBRIAN TRILOBITES
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 125
PLATE 16
EMEC EH EMTS Y REM. “SPECIES. 2°54 sic. Neate cinta cb.a/tisg ta sides ca-wldiesieo cence cus 93
I, 2. Incomplete cranidium (X 3).
3, 4. Side and dorsal views of the holotype cranidium ( 3).
Honey Creek limestone; (loc. 12k) Honey Creek, 7 miles north of
Springer, Wichita Mountains, Oklahoma.
er CMMeT CUB OTIC We S PCCICSereieyaiciers srsis/oe' sere = orale © isieie ola se aleie’s sie wierd on Ss 93
5, 6. Side and dorsal views of the holotype cranidium (X 3).
7-10. Similar views of two other cranidia (X 3).
Davis formation; (loc. 11k) Federal Lead Mine No. 4, Flat River,
Missouri.
ERE MIE MIESSOUTICHSES, TIEW SPECICS. 2. icc 530 os cnc enc dened sewneasonws cade 04
II, 12. Dorsal and side views of the holotype cranidium ( 3).
13, 14. Two larger cranidia (X 2).
Davis formation; (loc. 11k) Federal Lead Mine No. 4, Flat River,
Missouri.
MaMREISHE ME DAUENSIS, NEW SPECIES. . 2... 6. ..c ccc cress ccsenveeesesesvves 85
15. The associated libragene.
16, 17. Dorsal and side views of the holotype cranidium ( 1.5).
Secret Canyon shale; (loc. 61) south of Hamburg Mine, Eureka Dis-
trict, Nevada.
MOLTO IOMIALCILCISUS SIIGW? SPCCIESticieicpeverets ieis save oe ovels crore’ aie) vi sivielevele is siete ele eth 85
18, 19. Side and dorsal views of the holotype cranidium (X 1.5).
20. Incomplete cranidium (> 1.5).
Orr formation; (loc. 32t) Fandango Spring Canyon, Dugway Range,
Utah.
WANG ORMAISSOUTIENSIS: TIEW. SPECIES. cincc's elsies.o.c'sieis © aims ais Be aicie'eye sie cise ares s 86
21, 22. Dorsal and side views of the holotype cranidium ( 2).
23. Cranidium ( 1.5).
24, 25. An exfoliated cranidium (X 1.5).
26. Libragene.
Davis formation; (loc. 11k) Federal Lead Mine No. 4, Flat River,
Missouri.
DLL SACMUT GU CEO ATI CW. SPECIES sr. cca oie vaie lel eves siisis\ays ous Sleig)e lenatejal's oraidln vi giehalalate 87
27-29. Squeeze, side and dorsal views of holotype cranidium ( 1.5).
Honey Creek limestone; (loc. 9q) Blue Creek Canyon, Wichita Moun-
tains, Oklahoma.
RANUSIA CLASSUNGPOINGLG, NEW SPECIES... 2... ceca csecesesctcccsvdeneds 87
30-32. Squeeze, dorsal, and side views of holotype cranidium (X 1.5).
Davis formation; (loc. 11k) Federal Lead Mine No. 4, Flat River
Missouri.
MEP II SAE E) POY SCUSIS, TIEW SPECIES 2 2.6. 5 ola ogc siete cies wae oieiie eieeiena sinus des ciate 88
Bane4e Cranidium: (>< 2.5);
35, 30. Dorsal and side views of the holotype cranidium (X 1.5).
37, 38. Small cranidium with neck spine (X 3).
Honey Creek limestone; (loc. 12p) 4 miles east of Alpers, Arbuckle
Mountains, Oklahoma.
naar Ce ) HUANENSIS, TIEW SPECIES. «|<. . oie aice oaieteesiss <6 cpeielnces ace es 88
39. Holotype cranidium (X 1.5).
40, 41. Small cranidium (X 3).
Mendha limestone; (loc. 7j) 1 mile northwest of Italian Ranch foothills,
Quinn Canyon Range, Nevada.
10
126 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
PEATE 17
PAGE
Iddingsta anatina, new SPGCieS 40s ssk fase as Sew ea oe es ee ee 890
1, 2. Dorsal and side views of the holotype cranidium ( 2).
Honey Creek limestone; (loc. 9q) Blue Creek Canyon, Wichita Moun-
tains, Oklahoma.
Iddwngsta simplicttas, new SPECiES)..< 5 sos s'ac smn es cen tombe ets 890
3, 4. Side and dorsal views of the holotype cranidium.
5-8. Two other cranidia (1.5).
go. Libragene (X 1.5).
Honey Creek Cea (loc. 89y) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Burnetia intermedia, new species Me ceteelatetscs siatel a arelete ie laherosert omer aate bit eT eS
10, 11. Dorsal and side views of the holotype cranidium ( 1.5).
Honey Creek limestone; (loc. 12p) 4 miles southeast of Hennepin,
Arbuckle Mountains, Oklahoma.
Burnetia alia, new: Species sa. descren o toiee ee ee
12, 13. Dorsal and side views of the holotype cranidium.
14. Libragene.
Davis formation; (loc. 11k) Federal Lead Mine No. 4, Flat River,
Missouri.
Burnetia extensa, New SPECIES: kai. 4. cee esl teenie eee ee eee
15, 16. Partially exfoliated cranidium ( X 2).
17, 18. Side and dorsal views of the holotype cranidium (> 1.5).
19. Small cranidium (xX 4).
20. Another cranidium (X 3).
21. Libragene (X 2).
22. Libragene and possible pygidium ( 2).
Honey Creek limestone; (loc. 890v) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Burnetia -extlis Mew: SPECIES. ).4..0 enone oaeinlce ne hese ee
23. Small cranidium (X 1.5).
24, 25. More complete cranidium (X 2).
26, 27. Side and dorsal views of the holotype cranidium (X 1.5).
Honey Creek limestone; (loc. 89v) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Burnetia | Curta, NEW, SPECIES xcs -\iciaee core eee ee Oe ee
28, 29. Side and dorsal views of the holotype cranidium ( 2).
Honey Creek limestone; (loc. 9q) Blue Creek Canyon, Wichita Moun-
tains, Oklahoma.
Burnetia. ectypa, New. SPeCleS ane sia «:.-0 «o/c. cweroctai de eee on ne Tee eae
30, 31. Dorsal and side views of the holotype cranidium.
Honey Creek limestone; (loc. 89v) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
Burnetia lingula, new species; cs). st hace ta cee cee eae ee eee
32, 33. Side and dorsal views of the holotype cranidium (> 1.5).
Honey Creek limestone; (loc. 89v) West Timbered Hills, Arbuckle
Mountains, Oklahoma.
80
81
81
83
82
3 Fe
SMITHSONIAN
MISCELLANEOUS COLLECTIONS
UPPER CAMBRIAN TRILOBITES
VOL. 103, NO. 5, PL. 17
VOL. 103, NO, 5, PL. 18
SMITHSONIAN MISCELLANEOUS COLLECTIONS
UPPER CAMBRIAN TRILOBITES
NO. 5 UPPER CAMBRIAN TRILOBITES—RESSER 127
PLATE 18
: i : PAGE
WL CAMEMAONTLS., TIGW., SPECIES a12 ois sia.6 s.«, 4 500s sie w/s-d'elansve ose elaie 406 sperelecicleieiocis.s 04
1, 2. Dorsal and side views of the holotype cranidium (> 1.5).
3, 4. Large exfoliated cranidium.
Snowy Range formation; (loc. 37r) Warm Spring Creek, Wind River
Range, Wyoming.
PA IAMULO LENSES! MEW, SPECIES <6) 5(0s\4<1s)s, cocisisidic« ss ¢ bible wis ge hcla cise elo Oe es 95
5, 6. Dorsal and side views of the holotype cranidium.
St. Charles limestone; (loc. 54e) Blacksmith Fork, Bear River Range,
Utah.
iA CUA MEW SPECIES o...60% 6 ds oa cas sek ciiewaccsesei ee uesccdees 95
7. Libragene (X 1.5).
8, 9. Dorsal and side views of the holotype cranidium (> 1.5).
10. Small cranidium with Berkeia saratogensis (X 1.5).
Potsdam sandstone (Theresa member); Greenfield, west of Saratoga
Springs, New York.
EN aM CIAL Me Wr SPECIOS ria spe lera{orsscisinlorase 0) © sxe. s.«, vias alale'e oe ores clei ale susteee 06
II, 12. Side and dorsal views of the holotype cranidium (X 2.5).
Secret Canyon shale; (loc. 63) northeast of Adams Hill, Eureka Dis-
trict, Nevada.
ee MEET ACMSUCOIRCSSCIY 5. 0.5/4.5 2 Soi x a's. din oes kote e Shad owned bared Seaa ne 96
13, 14. Nearly complete cranidium.
15, 16. Pygidium.
17. Libragene.
Davis formation; (loc. 11k) Federal Lead Mine No. 4, Flat River,
Missouri.
PE AE COILEG HCW, SPECICS ~ 5+) 4, afsys 24.5) vei aie, sobs seats dis dia a has svi Sle a aale lala ea 96
18, 19. Dorsal and side views of the holotype cranidium (X 2).
20, 21. Smaller cranidium (> 1.5).
225238 Pyeidium (>< 1.5):
Honey Creek limestone; (loc. 9q) Blue Creek Canyon, Wichita Moun-
tains, Oklahoma.
PRIN PAOD EW SPECIES. <-.5. 4 s.0.c7010 s/h) «cides Aves w MDa ae aka cies vlae Dog ead 07
24, 25. Side and dorsal views of the holotype cranidium.
26, 27. Small cranidium.
Honey Creek limestone; (loc. 9p) Blue Creek Canyon, Wichita Moun-
tains, Oklahoma.
MME ALITA ESSE TY sio)oc cite aise Sow ok cis lse woke oss e Lag ce wlasisidaiiadieansbes 97
28, 29. Squeeze of a pygidium.
30, 31. A large cranidium, with injury at left anterior angle. (See also
pl. 10, figs. 1-5.)
Honey Creek limestone; (loc. 12p) 4 miles southeast of Hennepin,
Arbuckle Mountains, Oklahoma.
128 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
PLATE 19
Bluamia bridge: Resser ..\c3568. 2. See cas esate eee aire se ee ee ee ee 07
1, 2. Partially exfoliated cranidium.
3, 4. Well-preserved cranidium.
5. Libragene. (See also pl. 18, figs. 28-31.)
Honey Creek limestone; (loc. 89v) West Timbered Hills, Arbuckle
Mountains, Oklahoma. (This is the type locality.)
Elumia vagans; NEW Species: 5 aaare eto s sities decade «oem Lee nana ee 08
6, 7. Dorsal and side views of the holotype cranidium.
8, 9. Pygidium (X 1.5).
Honey Creek limestone; (loc. 9q) 15 miles northwest of Fort Sill,
Wichita Mountains, Oklahoma.
Elana. brevifrons, new. Species. 1d Gus : Fee eee ea ee eee 08
10, Ir. Cranidium (xX 1.5).
12, 13. Side and dorsal views of the holotype cranidium.
14. Libragene.
Honey Creek limestone; (loc. 12p) 4 miles east of Alpers, Arbuckle
Mountains, Oklahoma.
Taentcephalus pealet, new Spectess os acs .«2- vas dtieancnye «ae See ee ee 99
15, 16. Small cranidium retaining the test (xX 4).
17. Large exfoliated cranidium showing granules under anterior furrow.
18. Partly exfoliated head and libragene (X 3).
19, 20. Pygidium (x 2.5).
21. Exfoliated cranidium showing scattered granules (XX 3).
22, 23. The holotype cranidium ( 3).
Dry Creek shale; (loc. 153) Horsehoe Hills, east of Logan, Montana.
Laencephalus holmest; Mew: SPpeCes iaje'savesaies cle aaicie sales eine sere en 100
24. Exfoliated cranidium (> 2).
25, 26. Side and dorsal views of the holotype cranidium ( 3).
27. Pygidium (X 3).
Dry Creek shale; (loc. 66x) Gallatin Valley, northwestern Yellowstone
National Park, Wyoming.
Toaenmcephalus libertyensis, new’ SpecieS..\c....0.2440<)oe es cee eee eee 101
28, 29. Holotype cranidium (X 2.5).
30, 31. Small cranidium (X 4).
32. Libragene (X 2).
St. Charles limestone; (loc. 56g) 6 miles west of Liberty, Bear River
Range, Idaho.
Taenicephalus.cordillerensis. Miller.) vcc8s tee eee IOI
33. Surface with cranidia and libragene (X 3).
Snowy Range formation; (loc. 37r) Warm Springs Creek, Wind River
Range, Wyoming.
UPPER CAMBRIAN TRILOBITES
VOL. 103, NO. 5, PL. 19
SMITHSONIAN MISCELLANEOUS COLLECTIONS
UPPER CAMBRIAN TRILOBITES
NO. 5 UPPER CAMBRIAN TRILOBITES—-RESSER 129
PLATE 20
a NMEA ME TI SPVEN OS oat cag halt o o/s, 5) <'ala = iatalate nuclei ata = -iatape erence aides 83
1, 2. Dorsal and side views of the holotype cranidium (X 2.5).
3. Libragene (XX 3).
Ore Hill formation; (loc. 107v) 4 mile northwest of Drab, Pennsylvania.
MaCMECPNAIUS Granulosus, NEW SPECIES... 2. 5..6060s0000ce0ccecesecdoinecs IOI
4, 5. Side and dorsal views of the holotype cranidium (X 2).
6, 7. A less complete cranidium ( 2.5).
St. ae limestone; (loc. 4y) Two Mile Canyon, Wasatch Mountains,
aho.
UREPIACEPGIUS INGIGGENSIS, NEW SPECIES. ......6cce ccna ssaeesceacnacevsens 102
8, 9. Side and dorsal views of an exfoliated cranidium (X 3).
10. The holotype cranidium and libragene (X 2.5).
St. Charles limestone; (loc. 54x) Two Mile Canyon, 2 miles south of
Malad, Wasatch Mountains, Idaho.
EP RERCTIUTUSVOTNGILS, NEW SPECIES... ooo lien csleis oe cnies vee den ee ele sels le 102
II, 12. Dorsal and side views of the holotype cranidium ( 3).
St ees limestone; (loc. 54e) Blacksmith Fork, Bear River Range,
tah.
Taemcephalus hyrumensis, new SpeCieS.......0..ccecc cece cccescccetecaees 104
13, 14. Dorsal and side views of the holotype cranidium retaining
most of test (X 2).
15. Exfoliated cranidium and hypostoma (X 2).
16. Side view of the cranidium in fig. 15 (X 2).
17, 18. Two different views of a cranidium and associated pygidium
(X 2.5).
St. Charles limestone; (loc. 55h) Blacksmith Fork, Bear River Range,
Utah.
MP MIEE PLS SPECIOSUS, NEW SPECIES... ic cc.c cas escacseeessicsacece dense 100
19. Holotype cranidium and libragene, associated with Biullingsella
exasperata Bell. Portion of another cranidium lies under the
libragene. (XX 1.5.)
Dry Creek shale; (loc. 62r) Abiathar Mountain, northeastern Yellow-
stone National Park, Wyoming.
ERE UMACTOULU St ATLEW) SPECIES as sie sicle tia djaie hems save sisielole hale sels avarehs bela 84
20, 21. Side and dorsal views of the holotype cranidium and examples
of Taenicephalus shumardi and Orygmaspis eryon.
Franconia sandstone; 2 miles west of Clifton, Monroe County,
Wisconsin.
130 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
PLATE 21
Taemchephalus striati{rons, Mew Species ...«. 21+ cei onesie cco eee en
Holotype cranidium (XX 4.5).
2, 3. A larger cranidium (> 2).
St. Charles limestone; (loc. 54e) Blacksmith Fork, Bear River Range,
Utah.
Maenicephalus utahensis, Mew Species.1.). 20s dee o he ae els ieee eee
4, 5. Exfoliated cranidium (X 2.5).
6. Holotype cranidium (X 2.5
St. Charles limestone; (loc. ae Blacksmith Fork, Bear River Range,
Utah.
Taemcephalus modestus, new SpeCies sae aust aoe ous eee cee oe
7. Holotype cranidium ( 3).
St. Charles limestone; (loc. 55h) Blacksmith Fork, Bear River Range,
Utah.
Taencephalus texanus, mew Species... .:4 on). va ote usdeene eos poe
8, 9. Side and dorsal views of the holotype cranidium ( 2.5).
10-12. Three other cranidia in various states of preservation (x 2.5).
Wilberns formation; (loc. 68) Packsaddle Mountain, 11 miles south-
east of Llano, Texas.
Taenicephalus wichitaensis, mew SpeCieS......%.. 0.20 is. ceeses odsscecdebeeuess
13. Exfoliated cranidium (X 2).
14, 15. Dorsal and side views of the holotype cranidium (X 2)
16, 17. Two other cranidia (X 2.5).
Honey Creek limestone; (loc. 9q) Blue Creek Canyon, 15 miles north-
west of Fort Sill, Wichita Mountains, Oklahoma.
Taentcephalus quinnensts, Tew SPECIES. us. oaece- ae. «cide doe eeuiee eee
18, 19. Incomplete cranidium ( 1.5).
20, 21. A pygidium (> 2).
22. Holotype cranidium (X 2.5).
23. Libragene (X 2).
Mendha limestone; (loc. 7j) 1 mile north of Italian Ranch foothills,
Quinn Canyon Range, Nevada.
Taemcephalus castlensis; new Speeies:, «4... 5..¢.acvans Came denen one ween
24, 25. Dorsal and side views of the holotype cranidium ( 2).
Dry Creek formation; (loc. 62s) Castle Peak, north of Squaw Creek,
Gallatin Range, Montana.
Talbotina ulricht, lew: Species: 4% suco« ote ine ine to a ne Oe eee
26. Holotype cranidium (X 2).
Wilberns formation; (loc. 14b) Cold Creek, opposite north end of
Sponge Mountain, Texas.
Talbotina. candida, new~ Species... mvs tse soc ces te oe ee eee
27, 28. Dorsal and side views of the.holotype cranidium (x 2).
Wilberns formation; (loc. 14b) Cold Creek, opposite north end of
Sponge Mountain, Texas.
Burnetia pennsylvanica, new SPECIES. We. cihu tee akie ates ee eae 2 el
29, 30. Side and dorsal views of the holotype cranidium.
31. Smaller cranidium ( 2).
Ore Hill formation; (loc. 107e) 1 mile south of Ore Hill, Pennsylvania.
Faentcephalus wyomingensis, new Species;..,..-.suee des eeen eee eens
32. Holotype and other cranidia, and libragene (X 1.5).
Snowy Range formation; head of Buffalo Fork, west side of Big Horn
Mountains, Wyoming.
103
104
104
105
105
106
107
107
84
106
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 5, PL. 21
UPPER CAMBRIAN TRILOBITES
INDEX
abrupta, Irvingella, 21
accincta, Irvingella, 21
Acrocephalites, 79
adamsensis, Irvingella, 24
aduncus, Stenopilus, 60
affinis, Chariocephalus, 8
Berkeia, 96
agrarius, Chariocephalus, 8
Agraulos (?) thea, 2
agrestis, Irvingella, 17
alberta, Blountiella, 61
Drumaspis, 29
Irvingella, 23
Maryvillia, 70
albertensis, Arapahoia, 46
Cholopilus (?), 37
Coosia, 73
Illaenurus, 38
Plethometopus, 47
alia, Irvingella, 16
alpersensis, Iddingsia, 88
alta, Burnetia, 80
Irvingella, 19
anatina, Iddingsia, 89
angustata, Berkeia, 93
angustatus, Chariocephalus, 9
angustilimbata, Parairvingella, 25, 26
ara, Kingstonia, 51
Ucebia, 51
Arapahoia, 43
albertensis, 46
elongata, 45
polita, 45
prolixa, 46
reesidei, 44
stantoni, 44
typa, 44
walcottae, 46
arbucklensis, Irvingella, 14
arctica, Irvingella, 13
ardmorensis, Irvingella, 18
arguta, Bynumia, 53
argutus, Ellipsocephaloides, 63
bacca, Irvingella, 20
Stenopilus, 59
badgerensis, Chariocephalus, 12
banffensis, Meteoraspis, 72
Bathynotus, 2
bearensis, Ellipsocephaloides, 65
bellus, Chariocephalus, 6
Berkeia, 79, 90
affinis, 96
angustata, 93
comes, 90
jucunda, 93
missouriensis, 94
nevadensis, QI
retusa, 92
saratogensis, QI
sculptilis, 93 .
wichitaensis, 92
bicincta, Iddingsia, 87
bilobata, Pterocephalina, 77, 78
bilobatus, Dicellocephalus, 77
Dikellocephalus (Pterocephalus), 77
Platycolpus, 78
Blountia, 60
(?) disparilis, 61
kindlei, 60
plana, 60
Blountiella, 61
alberta, 61
bosworthensis, Kingstonia, 48
brevifrons, Elvinia, 98
brevis, Coosella, 76
bridgei, Elvinia, 97
briscoensis, Bynumiella, 57
Drumaspis, 30
Ellipsocephaloides, 63
bulla, Chariocephalus, 7
burlingi, Chariocephalus, 12
burnetensis, Irvingella, 20
Burnetia, 79, 80
alta, 80
cava, 83
curta, 83
ectypa, 82
edwardsi, 84
exilis, 81
extensa, 81
intermedia, 80
131
132
lingula, 82
pennsylvanica, 84
Burnetidae, 79
butleri, Ellipsocephaloides, 66
buttsi, Cheilocephalus, 36
Bynumia, 50, 52, 57, 58, 60
arguta, 53
elegans, 52
eumus, 52
(?) modesta, 55
mollis, 56
(?) producta, 56
rangerensis, 55
robsonensis, 54
sawbackensis, 55
sulcata, 54
venusta, 54
walcotti, 53
Bynumiella, 57
briscoensis, 57
(?) oklahomensis, 58
typicalis, 57
Bynumina, 58
caelata, 58
missouriensis, 59
caelata, Bynumina, 58
Talbotina, 107
Calyptomma, 37
typicale, 38
Camaraspis, 42
canadensis, Coosia, 73
candida, Talbotina, 107
carus, Ellipsocephaloides, 65
castlensis, Taenicephalus, 106
cava, Burnetia, 83
Chariocephalus, 2, 3, 4
affinis, 8
agrarius, 8
angustatus, 9
badgerensis, 12
bellus, 6
bulla, 7
burlingi, 12
gracilens, 9
magnus, II
montis, 10
peloris, 5
tenerus, 5
LELeS; 7
INDEX
tumifrons, 3, 25
ulrichi, 11
wichitaensis, 10
Cheilocephalus, 35
buttsi, 36
stcroixensis, 35
texanus, 35
wichitaensis, 35
Cholopilus, 36
(?) albertensis, 37
nevadensis, 36
clara, Drumaspis, 33
comes, Berkeia, 90
Coosella, 73, 75
brevis, 76
texana, 75
Coosia, 73
albertensis, 73
canadensis, 73
dakotensis, 74
tridentensis, 74
wyomingensis, 67
cordillerensis, Taenicephalus, ror
crassimarginata, Iddingsia, 87
curta, Burnetia, 83
curtus, Ellipsocephaloides, 62
dakotensis, Coosia, 74
Dartonaspis, 2, 4
davisensis, Irvingella, 16
deckeri, Drumaspis, 31
Irvingella, 19
declivis, Ellipsocephaloides, 67
Dicellocephalus bilobatus, 77
Dikellocephalus (Pterocephalus) bilo-
batus, 77
disparilis, Blountia (?), 61
Dokimocephalus, 70
Drumaspis, 3, 4, 28
alberta, 2
briscoensis, 30
clara, 33
deckeri, 31
goodsirensis, 30
idahoensis, 29
maxwelli, 31
nitida, 34
osella, 32
sabinensis, 31
texana, 32
utahensis, 34
walcotti, 28
wichitaensis, 33
ectypa, Burnetia, 82
edwardsi, Burnetia, 84
electra, Kingstonia, 47
elegans, Bynumia, 52
Elkia, 79
Ellipsocephaloides, 62
argutus, 63
bearensis, 65
briscoensis, 63
butleri, 66
carus, 65
curtus, 62
declivis, 67
monsensis, 66
montis, 65
nitela, 66
sawbackensis, 64
silvestris, 64
elongata, Arapahoia, 45
Tsinania, 40
elongatus, Illaenurus, 40
Elvinia, 26, 94
brevifrons, 98
bridgei, 97
granulata, 96
gregalis, 96
longa, 97
matheri, 95 '
missouriensis, 96
montis, 94
roemeri, 96, 97
ruedemanni, 95
tetonensis, 94
utahensis, 95
vagans, 98
Enontioura, 42
typicalis, 43
eumus, Bynumia, 52
eurekensis, Parairvingella, 26
Eurekia, 41
exilis, Burnetia, 81
extensa, Burnetia, 81
flohri, Irvingella, 24
gibba, Irvingella, 13
goodsirensis, Drumaspis, 30
INDEX 133
gracilens, Chariocephalus, 9
granulata, Elvinia, 96
granulosus, Taenicephalus, Io1
grata, Pterocephalina, 78
gregalis, Elvinia, 96
hamburgensis, Parairvingella, 27
Hesperaspis, 43
highlandensis, Platycolpus, 41
holmesi, Taenicephalus, 100
hybrida, Maryvillia, 71
hyrumensis, Taenicephalus, 104
idahoensis, Drumaspis, 29
Iddingsia, 79, 85
alpersensis, 88
anatina, 89
bicincta, 87
crassimarginata, 87
missouriensis, 86
nevadensis, 85
(?) quinnensis, 88
simplicitas, 89
utahensis, 85
Illaenurus, 38
albertensis, 38
elongatus, 40
priscus, 38
(?) sinclairensis, 39
intermedia, Burnetia, 80
Parairvingella, 27
Irvingella, 2, 3, 13
abrupta, 21
accincta, 21
adamsensis, 24
agrestis, 17
alberta, 23
alia, 16
alta, 19
(Parairvingella) angustilimbatus, 26
arbucklensis, 14
arctica 13
ardmorensis, 18
bacca, 20
burnetensis, 20
davisensis, 16
deckeri, 19
flohri, 24
gibba, 13
media, 22
134
mesleri, 15
oblonga, 17
ottertailensis, 17
plena, 18
(Irvingellina) protuberans, 5, 13
recurva, I5
richmondensis, 23
silvestris, 13
tumifrons, 2
Irvingellina, 3, 13
Irvingelloides, 2, 3
jucunda, Berkeia, 93
kindlei, Blountia, 60
Kingstonia, 51
Kingstonia, 47, 60
ara, 51 :
bosworthensis, 48
electra, 47
kindlei, 51
loperi, 50
mucro, 48
(?) plena, 50
promissa, 49
robsonensis, 49
sullivanensis, 49
thea, 2
vulgata, 50
Komaspidae, 2
Komaspidella, 2
Komaspis, 2, 3
libertyensis, Taenicephalus, 101
lingula, Burnetia, 82
longa, Elvinia, 97
loperi, Kingstonia, 50
Maryvillia, 70
Macelloura, 43
transversa, 43
magnus, Chariocephalus, 11
maladensis, Taenicephalus, 102
marjumensis, Maryvillia, 69
Maryvillia, 67
alberta, 70
hybrida, 71
loperi, 70
marjumensis, 69
montis, 7I
INDEX
moosensis, 72
utahensis, 68
violaensis, 68
wyomingensis, 67
matheri, Elvinia, 95
maxwelli, Drumaspis, 31
media, Irvingella, 22
mesleri, Irvingella, 15
Meteoraspis, 72
banffensis, 72
missouriensis, Berkeia, 94
Bynumina, 59
Elvinia, 96
Iddingsia, 86
modesta, Bynumia (?), 55
modestus, Taenicephalus, 104
mollis, Bynumia, 56
monsensis, Ellipsocephaloides, 66
montis, Chariocephalus, 10
Ellipsocephaloides, 65
Elvinia, 94
Maryvillia, 71
moosensis, Maryvillia, 72
mucro, Kingstonia, 48
nevadensis, Berkeia, 91
Cholopilus, 36
Iddingsia, 85
nitela, Ellipsocephaloides, 66
nitida, Drumaspis, 34
Norwoodella, 43
notha, Pterocephalina, 77
oblonga, Irvingella, 17
oklahomensis, Bynumiella (?), 58
Platycolpus, 40
ornatus, Taenicephalus, 102
osella, Drumaspis, 32
ottertailensis, Irvingella, 17
Parairvingella, 4, 13, 25
angustilimbata, 26
eurekensis, 26
hamburgensis, 27
intermedia, 27
pealei, Taenicephalus, 99
peloris, Chariocephalus, 5
pennsylvanica, Burnetia, 84
plana, Blountia, 60
Ce +92 Dea
INDEX
Platycolpus, 39, 40
highlandensis, 41
oklahomensis, 40
quinnensis, 40
sinclairensis, 42
wichitaensis, 41
plena, Irvingella, 18
Kingstonia (?), 50
Plethometopus, 43, 47
albertensis, 47
pogonipensis, Pterocephalina, 78
polita, Arapahoia, 45
priscus, Illaenurus, 38
producta, Bynumia (?), 56
prolixa, Arapahoia, 46
promissa, Kingstonia, 49
protuberans, Irvingella (Irvingellina),
SES
Pterocephalina, 76
bilobata, 77
grata, 78
notha, 77
pogonipensis, 78
texana, 76
utahensis, 79
quinnensis, Iddingsia (?), 88
Platycolpus, 40
Taenicephalus, 105
rangerensis, Bynumia, 55
recurva, Irvingella, 15
reesidei, Arapahoia, 44
retusa, Berkeia, 92
richmondensis, Irvingella, 23
robsonensis, Bynumia, 54
Kingstonia, 49
roemeri, Elvinia, 96, 97
ruedemanni, Elvinia, 95
sabinensis, Drumaspis, 31
saratogensis, Berkeia, 91
sawbackensis, Bynumia, 55
-Ellipsocephaloides, 64
sculptilis, Berkeia, 93
silvestris, Ellipsocephaloides, 64
Irvingella, 13
simplicitas, Iddingsia, 89
sinclairensis, Illaenurus (?), 39
Platycolpus, 42
135
Solenopleura, 79
Solenopleuridae Angelin, 79
speciosus, Taenicephalus, 100
stantoni, Araphahoia, 44
stcroixensis, Cheilocephalus, 35
Stenopilus, 50
aduncus, 60
bacca, 59
striatifrons, Taenicephalus, 103.
sulcata, Bynumia, 54
sullivanensis, Kingstonia, 49
Taenicephalus, 99
castlensis, 106
cordillerensis, 101
granulosus, IOI
holmesi, 100
hyrumensis, 104
libertyensis, Ior
maladensis, 102
modestus, 104
ornatus, 102
peali, 99
quinnensis, 105
speciosus, I00
striatifrons, 103
texanus, 104
utahensis, 103
wichitaensis, 105
wyomingensis, 106
Talbotina, 107
caelata, 107
candida, 107
ulrichi, 107
tenerus, Chariocephalus, 7
teres, Chariocephalus, 5
tetonensis, Elvinia, 94
texana, Coosella, 75
Drumaspis, 32
Pterocephalina, 76
texanus, Cheilocephalus, 35
Taenicephalus, 104
thea, Kingstonia, 2
transversa, Macelloura, 43
Tricrepicephalus tripunctatus, 67
tridentensis, Coosia, 74
tripunctatus, Tricrepicephalus, 67
Tsinania elongata, 40
tumifrons, Chariocephalus, 3, 26
Irvingella, 13, 25
136
typa, Arapahoia, 44
typicale, Calyptomma, 38
typicalis, Bynumiella, 57
Enontioura, 43
Ucebia ara, 51
ulrichi, Chariocephalus, 11
Talbotina, 107
utahensis, Drumaspis, 34
Elvinia, 95
Iddingsia, 85
Maryvillia, 68
Pterocephalina, 79
Taenicephalus, 103
vagans, Elvinia, 98
INDEX
venusta, Bynumia, 54
violaensis, Maryvillia, 68
vulgata, Kingstonia, 50
walcottae, Arapahoia, 46
walcotti, Bynumia, 53
Drumaspis, 28
wichitaensis, Berkeia, 92
Chariocephalus, 10
Cheilocephalus, 35
Drumaspis, 33
Platycolpus, 41
Taenicephalus, 105
wyomingensis, Coosia, 67
Maryvillia, 67
Taenicephalus, 106
+.
SA ENS kG
J, MANSON VALENTINE
ea! : , Bureau of Entomology and Plant Quarantine,
U, S. Department of Agriculture |
2
(PuBLICATION 3696)
Puree “arty OF. WASHINGTON
ie ‘PUBLISHED BY THE SMITHSONIAN INSTITUTION
. NOVEMBER at igee 75
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 103, NUMBER 6
ON THE PREPARATION AND PRESERVATION OF
INSECTS, WITH PARTICULAR REFERENCE
TO COLEOPTERA
BY
J. MANSON VALENTINE
Bureau of Entomology and Plant Quarantine
U. S. Department of Agriculture
(PUBLICATION 3696)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
NOVEMBER 21, 1942
|
The Lord Baltimore Press
BALTIMORE, MD., U. 8. A,
ON THE PREPARATION AND PRESERVATION OF
INSECTS, WITH PARTICULAR REFERENCE
LO, COLEOPTERA
By J. MANSON VALENTINE
Bureau of Entomology and Plant Quarantine
U. S. Department of Agriculture
It is beyond the scope of the present paper even to enumerate the
multiplicity of methods employed and customs adhered to in the prepa-
ration of insect specimens for the cabinet. Rather, what is intended
is solely a presentation of certain procedures which have proved most
useful to the writer after a period of considerable experimentation.
Broadly classified, there are two schools of technique in the mount-
ing of Coleoptera. The European entomologist habitually displays his
smaller specimens by gluing each of them, ventral surface down, to
a standard rectangular card which is then pinned. The American
prefers to mount his at the apices of small cardboard triangles whose
bases hold the pins. In the first method, the appendages are protected
but only the dorsal aspect of the insect is visible, an examination of
its ventral characters necessitating the removal of the specimen from
the card. This is a tedious and dangerous routine which the average
collector is reluctant to undertake ; its avoidance, however, cannot fail
to result in identifications based entirely on dorsal anatomy. The
American system, on the other hand, while providing opportunity to
study lateral and ventral characters (though only those not obscured
by the legs in their flexed positions) fails to afford proper protection
to the specimen. Neither technique ordinarily includes any degreas-
ing treatment with the result that too often, during the passage of
time, escaping oils render the specimen unfit for study.
The following is a description of a third procedure which attempts
to combine the merits of both schools and, at the same time, to elimi-
nate the more undesirable features of each. This technique was
developed specifically for the purpose of preparing good research
material in the Cicindelidae and Carabidae. However, it has proved
equally useful in other groups of beetles; and it has been adapted
with success to insects of various additional orders, especially to
those whose membranous wings, if present, are folded and concealed.
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 103, No. 6
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
I. KILLING
Since the final results are, in large measure, dependent upon the
killing agent used, and the manner in which killing is accomplished,
the choice of a suitable lethal chemical and its proper application be-
come matters of the highest importance. In general it may be said
that killing by means of the vapors from volatile anesthetics (lipoid
solvents) is to be preferred to direct submergence in any fluid. Two
such substances are recommended :
a. Carbon tetrachloride—lI{ specimens do not come into direct
contact with the fluid, they will remain relaxed after death, especially
if the catch is large and is allowed to remain in the killing bottle at
least 24 hours. A full bottle, however, should not be neglected longer,
as disintegration of soft parts will soon set in with consequent loss
of setae and abdominal segments. A convenient vehicle for carbon
tetrachloride is chopped elastic bands ; rubber imbibes the fluid readily
and retains it a long time. Crumpled paper toweling is also satisfac-
tory, especially when used in connection with light traps requiring
the efficient operation of large killing jars at least partly open to the
outside air. The heavy fumes arising from a paper towel saturated
with carbon tetrachloride will seek the bottom of the jar, which will
remain lethal throughout the night.
Advantages :
1. Produces fair to good relaxation. In this respect it is better
than either cyanide or alcohol.
2. Assists in the extraction of oils and fats.
3. Prepares the specimen for genitalic examination by causing ex-
cessive swelling when subsequently dropped into ether, an event usu-
ally resulting in the extrusion of the genital apparatus.
b. Ethyl acetate (acetic ether).—For all general purposes, this is
an ideal killing agent. Its advantages when used as such were first
pointed out to the author by Prof. Candido Bolivar, whose technique
was to half fill a collecting tube with coarsely ground cork moistened
(not wet to the point of adhesion) with the ether. Strips of paper or
pieces of cotton tape moistened with ethyl acetate and placed in a
vial are equally effective and are better for very small specimens.
A still more efficient method is to introduce a half inch or so of wet,
mixed plaster of paris in the bottom of a tube or vial. Allow the
plaster to set; dry it thoroughly in an oven; then saturate it with
ethyl acetate, pouring off any excess fluid after complete impregna-
tion. A collecting bottle of this sort may “stand up” under months
of use, if not left uncorked. When exhausted, it can be dried again
in the oven and recharged with ethyl acetate. It is advisable to allow
No. 6 PREPARATION OF COLEOPTERA—VALENTINE 3
the day’s catch to remain in the killing bottle at least overnight, in
order to insure the maximum relaxing effect of the vapor-filled atmos-
phere. Insects may thus be preserved, while awaiting mounting, for
an indefinite period, especially if they receive an occasional wetting of
ethyl acetate. It is better, however, to remove them for drying, de-
greasing (II), or preserving in fluid (X) before many weeks have
passed.
Advantages :
1. Specimens killed by the fumes of ethyl acetate are completely
relaxed and retain their flexibility when subsequently degreased in
ether (I1) or preserved in Barber’s fluid (X). Success in mounting
such material on a flat, smooth surface, or in arranging appendages
after direct pinning, or in relaxing specimens that have dried is far
greater than when either cyanide or alcohol has been used as the
killing agent.
2. Unlike the average cyanide jar, an ethyl acetate-charged killing
bottle acts promptly, permanently, and uniformly over a considerable
period of time. Insects show no tendency to revive if they are allowed
to remain in the lethal atmosphere a few minutes after all motion has
ceased, and delicate Lepidoptera as well as powerfully jawed beetles
usually succumb before they can injure themselves or other specimens
in the same bottle.
3. No fading or discoloration has so far been observed by the
author as a result of killing beetles and their larvae with ethyl acetate.
However, the green pigment of certain moths may turn yellow if the
specimens are not removed from the killing bottle as soon as dead.
4. The use of ethyl acetate presents no such hazard as does cyanide,
which is far more toxic to human beings.
5. Ethyl acetate is an ingredient of the relaxing fluid described
below (XI) and a solvent for cellulose cement ; hence its use as a kill-
ing agent simplifies the field technique by reducing the number of
necessary fluids to be carried on a trip of long duration.
II. DEGREASING
The use of some lipoid solvent in the preparation of Coleoptera,
especially carabid, cicindelid, and scarabaeid material, is of the great-
est importance. Ordinary sulfuric ether? (the commercial product)
suffices very well. Specimens should be soaked in ether until the
1 Ether can be conveniently and economically stored in 1-pound cans fitted
with small screw caps seated with cork. While in use as a grease solvent, it
will keep well in tightly corked homeopathic vials.
4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
fluid ceases to grow yellow owing to dissolved oils, one or two changes
of the bath aiding the process when the bulk of material is great.:
The duration of treatment varies from about 12 hours to a week,
depending upon the size and number of specimens per bottle, the
volume of ether in proportion to material, and the fat content of
the particular insects being degreased. To protect the specimens from
the clinging, watery exudate which escapes from them and tends to
work its way to the bottom of the container, a small wad of loosely
folded absorbent tissue or filter paper should first be placed in the
bottle. This will serve to capture such waste.
Ether will preserve ethyl acetate-killed material in a perfectly re-
laxed condition for an indefinite period. However, a certain degree
of surface etching will take place in very oily ether where specimens
should never be left very long. Also, very small specimens isolated
in a volume of ether too large to be discolored by them may become
temporarily brittle if given this treatment for more than a few hours.
Should such an event occur, a drop of Barber’s fluid (XI) on the
dry insect will serve to free its articulations instantly.
sulky specimens which have swelled in the ether bath should be
removed to a pad of absorbent paper where all turgidity can be elimi-
nated by gentle pressure after pricking through the dorsal abdominal
and nuchal membranes with a needle. Such material should always be
returned to clean ether for further degreasing before mounting.
Very hairy insects * and those possessing easily detached scales or
farinose areas should be subjected to the ether bath only when the
oily nature of the species at hand demands such treatment. Rela-
tively few Coleoptera (certain groups of farinigerous Buprestidae,
Curculionidae, etc.) fall under this category. These cannot be ether-
treated for long without the loss of at least some of their powdery
secretion.
2 Degreasing of pilose insects, such as some bees, flies, etc., must be thorough
if attempted at all. When body fats have been completely removed, the pile
may rather easily be restored to its natural fluffiness by lifting it with a
camel’s-hair brush while applying a stream of air through a blowpipe. The
detachable pile which covers the bodies of moths cannot, of course, withstand
rubbing or brushing. Gravid females of large-bodied species, however, often
require degreasing because of the high lipoid content of their egg masses;
since immersion is to be avoided in such cases, it is therefore desirable, before
spreading these, to remove the viscera through an incision in the ventral
abdominal wall. By clamping the pin (head end) in a horizontal position, it is
possible to perform this operation with the abdomen hanging free. The ab-
dominal cavity should be dusted with cotton dipped in dry plaster, blown
clean, and loosely stuffed with fresh cotton.
NO. 6 PREPARATION OF COLEOPTERA—VALENTINE 5
Advantages :
1. Ether effects the extraction of body fluids as well as complete
degreasing. After the ether bath, piceous, castaneous, and fulvous
specimens will acquire a remarkable freshness of color, lightly pig-
mented areas appearing in vivid contrast to dark as soon as evapora-
tion has taken place. Likewise, the true texture of the chitin, a
useful habitus character dependent upon clean microsculpture, will
be preserved as in life.
2. Never soiled with a sticky layer of grease to which dust and
fine litter adhere, the ether-treated specimen can always be cleaned
with a dry camel’s-hair brush with minimum danger to setae.
3. The greaseless insect can be firmly cemented to the mounting
support ; should it become dislodged, its lightness and flexibility will
insure a good chance of survival intact. Specimens exuding grease
after they have been mounted on points or cards invariably discolor
the paper and not infrequently work loose; when pinned directly,
they usually corrode their pins.
4. Ether-treated specimens acquire atmospheric moisture rapidly
and never become brittle under ordinary climatic conditions. In this
respect they contrast very favorably with grease-soaked specimens
whose ligaments and musculature eventually harden.
5. Swelling is accomplished by means of the ether bath, specimens
killed with the fumes of carbon tetrachloride becoming so turgid in
ether that usually the aedeagus and frequently its internal sac are
extruded (III). This may be a great advantage, insofar at least as
small carabids are concerned, since it eliminates the difficult process
of dissecting such material. The more moderate distention of ethyl
acetate-killed specimens in ether is useful not only in the extraction
of genitalia, but also in the preparation of soft-bodied larvae (IX).
Soaking in ether, however, will not cause swelling or extrusion of
genitalia in previously dried material.
There are, of course, other satisfactory, though perhaps not equally
efficient, lipoid solvents which may be used in place of ether. Chloro-
form, benzol (benzene), and diethyl carbonate all yield good results
but tend neither to be imbibed nor to expel body fluids so readily.
Xylol (xylene) is pleasant and convenient to use, but its tendency to
stiffen articulations and to deposit a whitish film are disadvantages.
Chloroform is the only one of these solvents heavy enough to float
both the specimens and their extracted body fluids, a fact which
makes it desirable to separate the two with a wire screen. Ether,
on the other hand, being lightest of all, permits both to sink, while
diethyl carbonate, xylol, and benzol effect a more or less temporary
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
separation, the water alone, at first, falling to the bottom. Of the
three last named, diethyl carbonate is the most efficient solvent. It is
less volatile than ether and therefore safer to handle.
III. STEPS PREPARATORY TO MOUNTING
When thoroughly degreased and ready for mounting, specimens
should be removed from the ether bath to a pad of absorbent tissue,
where, if desired, jaws may be separated and genital extrusions com-
pleted. Carabid jaws may best be opened by springing them apart
by means of fine forceps applied ventrally. Provided the killing tech-
nique has been followed as indicated, the jaws will yield readily and
the disclosed mouth parts may then be easily cleaned with a soft
camel’s-hair brush dipped in ether. Partial extrusion of the genital
apparatus of males may be successfully completed by slight pressure
on the abdomen ; should this not produce the desired result, the insect
is placed ventral surface uppermost and held while a sharp needle is
inserted in a membranous portion of the median lobe and the latter
gently extracted. The laterally curved aedeagi of carabids require
urging counterclockwise in the direction of their curvature. Any
fluids escaping from the body during the process of handling should
be washed off in a bath of clean ether. If the specimen is minute,
the operation should be performed in fluid (ether or Barber’s fluid)
under a binocular, a special tool of fine pin wire or drawn glass hold-
ing the beetle firmly against the bottom of the dissecting dish (fig. 1).
It is desirable to transfer drying specimens to a smooth, clean sur-
face (preferably glass) for the final arrangement of their appendages.
In preparation for the slip method of mounting, the legs are oriented
close to and on a plane with the body, and the antennae are directed
backward along the sides. Until cleared mounts of genitalia are de-
sired, these organs had best be left attached to the abdomen; in
male Carabidae and Cicindelidae, they tend to orient pointing clock-
wise as seen from above and in this position they are most convenient
for study.
IV. MOUNTING ON TRANSPARENT SLIPS
A sufficient series of each species collected at one time and under
the same ecological conditions are assembled and are ready for
mounting as soon as their surfaces are dry. Mounting should be
done before the insects have become stiff, since the legs in drying
tend to elevate the body slightly and this necessitates relaxing the
specimens again before they can be properly cemented to a plane
No. 6 PREPARATION OF COLEOPTERA—VALENTINE 7
surface. Should this eventuality arise, the dried insects can be com-
pletely and instantaneously relaxed merely by dipping them in
Barber’s fluid (XI).
The foregoing technique is preliminary to mounting on a trans-
parent supporting surface, though it does not preclude pinning in the
usual manner. The former, or slip system, applicable to large speci-
mens as well as small, has been developed in two ways:
a. Cellulose acetate mounting.—Only the best quality, heavy (.015,
020, .025 inch=15, 20, 25 gauge) acetate sheeting * can be used to
Wass
cov
a SONS
tN0\ £0
KN wee,
Fig. Z
Fic. 1—Preparing a minute male carabid for mounting.
Fic. 2.—Preferred arrangement of parts in a pinned carabid.
advantage. This material, unlike celluloid, will not curl when pinned
nor, apparently, will it discolor or lose any of its transparency when
exposed to the light. Up to 25 gauge, it can be cut easily with ordi-
nary scissors, can be punctured with a No. 3 pin, and will grip the
latter firmly without need of reinforcement.
On a piece of acetate sheeting of convenient size and gauge, each
species-time-locality series is cemented separately in a compact group,
usually of one row. Adequate space is left between the groups so
3 Acetate sheeting ordinarily tends to fog by collecting droplets of an oily
fluid when in contact with the fumes of pest and mold repellents such as
’ naphthalene, paradichlorbenzene, phenol, creosote, etc. Carbon tetrachloride,
however, has no such effect on it. Sheeting which will mot fog under the above
conditions can be obtained from Eastman Kodak Company, Rochester, N. Y.
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
that they may be cut apart without danger to appendages; and an
ample margin of free sheeting should remain in back of each group
in order that the pin, which is placed there, may not interfere with
hand-lens observations of the specimens nearest it (fig. 3). Standard
strips of 15-gauge sheeting cut in $- and 3-inch (13-mm. and 10-mm.)
widths will greatly facilitate the mounting of small to minute speci-
mens and will lend a neat appearance to the collection. The beetles
should face and be close to a long edge; the slips bearing series or
single specimens can then be speedily cut apart in one operation yield-
ing mounts of uniform dimension from back to front with the speci-
mens thereon occupying similar relative positions. If sufficiently
heavy sheeting is used for large beetles, it is seldom necessary to
i
sl
Fig 3 Fig. 4
Fic. 3—The cellulose acetate slip» method of mounting in series.
Fic. 4——The cover glass slip method of mounting in series.
place the pin in any position other than the standard one, in the
center of the rear of the mount. Should, however, an extra large
mount require reinforcement, a drop of cement placed at the point
of exit of the pin will suffice.
A good-quality, clear cement of cellulose base * should be used. It
should not be applied straight from the container, however, but
should first be diluted with sufficient solvent to allow small drops to-
form slowly on the head of a pin after the mixture has been thor-
oughly stirred. At least two lots should be made up, one diluted
with ethyl acetate and the other with amyl acetate. The former is
relatively quick-drying and is used in cementing firmly all but the
smallest speciinens to their acetate mounts; the latter is slower to
dry and therefore more useful in attaching minute specimens and
in mending broken appendages. A convenient applicator is the
head of a long, fine insect pin stuck into the cork of the vial in which
4 Duco “household” cement has been found very serviceable.
4
%
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;
4
na
No. 6 PREPARATION OF COLEOPTERA—VALENTINE 9
the cement is mixed. Good adhesion with a minimum droplet of
cement depends, of course, upon the consistency of the latter, which
should not be so thick as to provide poor capillarity, nor so thin as to
spread beyond the original confines of the droplet.
Advantages :
1. An assemblage of specimens illustrative of a circumscribed popu-
lation or of an ecological sample of a species provides far more
valuable data than isolated specimens and should be kept intact under
the same label. The slip method of mounting makes this possible for
large specimens as well as small, and promotes comparison of
individuals.
2. Too often the customary procedure of mounting small, flexed
insects on cardboard points results in specimens whose appendages
and ventral anatomy are seriously obscured by the adhesive. Another
handicap to study frequently arises from undue arching of the body
and depressing of the head. The transparent-slip technique allows
maximum visibility of all parts, which, in all but the most convex
forms (some weevils, mordellids, etc.), can easily be arranged so as
to appear nearly on one plane. Under these conditions, buccal and
appendicular anatomy and comparative dimensions of legs, etc., can
be observed with a maximum degree of ease.
3. One of the commonest causes of damage in a collection is the
inadvertent contact between protruding labels and specimens deli-
cately mounted on points. The slip method obviates this hazard.
4. Mutilation of specimens and interference with hand-lens exami-
nation by pins thrust through elytra are eliminated.
5. Labels under acetate mounts are readable from above; usually
_ they can be larger than the minimum-sized label customarily attached
to specimens mounted individually.
6. A great saving of time is effected when specimens are mounted
in series by reducing the number of labels necessary and speeding
up transfer.
7. Space is conserved through the close approximation of specimens
on the mounting slip.
8. Far fewer pins are used.
g. The problem of which method of mounting—pin or point—
should be applied to a rather small specimen is eliminated ; this makes
for uniformity in the collection.
to. Any specimen can instantly be removed from the acetate mount
merely by touching it with a brush containing ethyl acetate.
b. Cover-glass mounting.—This is a refinement of the above tech-
nique, micro cover glass (No. 1 or No. 2) being used in place of
Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I03
acetate sheeting. Cover glass is best cut with a diamond point; a
small chip cemented to the end of a drawn-glass tube makes a perfect
instrument for the purpose. The slip may be cut to the required size
after the specimen or series has been attached to it. It is imperative
to work on a perfectly smooth, level surface, preferably plate glass;
and to use a rule which will not slip, such as a microscope slide faced
on one side with adhesive tape. The best adhesive for mounting on
glass is acetate cement thinned with ethyl acetate. A generous appli-
cation of a slightly thicker mixture should be used to attach two
narrow strips of bond paper or single-ply bristol board (about 4 inch
wide for small slips), one on each side of the glass along the entire
rear margin of the mount. Approximately half the width of these
strips should be occupied by the glass between them, the other half
being left for direct contact of the strips and subsequent perforation
by the mounting pin. The strips should be gently pressed together
and carefully aligned. When working with a quantity of material
of fairly uniform size, a number of slips can be attached at intervals
to a long bottom strip, and a top strip of equal width can then be
cemented over the whole. After 15 minutes or so of drying, before
the cement has become thoroughly hardened, the mounts should be
cut apart, the paper trimmed close to the glass with fine scissors, and
each mount carefully pinned on a flat-topped, gauged block, prefer-
ably slotted (VI) (fig. 4). It is advisable to pin the mount tilted
slightly upward, and to immobilize it with a drop of cement deposited
on the bottom strip around the pin.
A well-constructed cover-glass mount is neat, strong, and optically
ideal. However, unless its specimens have been exceedingly thor-
oughly degreased, it may, in time, collect fine droplets of oil emanat-
ing from them. Cover glass is, therefore, best employed in mount-
ing small specimens (10 mm. or less) which can easily be cleaned of
all grease.
The writer has experimented with a variety of materials for the
supporting strips of these mounts. None of the transparent “plastics”
and celluloids used has proved so efficient as heavy paper. Besides
its stability, another advantage in the paper base is that it can effec-
tively serve to display data, numbers, etc., written thereon.
V. DIRECT PINNING
Provided specimens are of sufficient size to justify impaling them
on insect pins, they can always be effectively mounted in accordance
with this standard technique. It should be pointed out, however, that
substituting direct pinning in the larger forms for the transparent-
No. 6 PREPARATION OF COLEOPTERA—VALENTINE II
slip method entails some sacrifice of utility, safety, and uniformity.
On the other hand, an obvious advantage in pinning lies in the result-
ing mobility of individual specimens. Whichever method is chosen,
the quality of the finished product depends, to a large extent, upon
two factors: proper killing (1) and adequate degreasing (II).
In mounting Coleoptera, the location of the pin is internationally
agreed upon: It should enter the anterior discal portion of the right
elytron and emerge on the right side between the mid and hind legs,
passing through the metasternum laterad of the mid line; it should
be so oriented that both longitudinal and transverse axes of the beetle
are at right angles to it. The writer recommends the European sys-
tem of appendage arrangement as combining neatness and compact-
ness with maximum visibility. The head and pronotum are extended,
with antennae close to the dorsal surface and directed straight back;
the legs are directed downward and toward the pin, but with femora
more or less horizontal and close to the body, the anterior femora
pointing forward, the mid and hind pairs pointing backward. A
fresh specimen naturally assumes this attitude when pinched laterally
between the fingers. If the insect has been killed with ethyl acetate,
its appendages will usually dry in position without the necessity of
guard pins. However, drooping of the head and pronotum may oc-
cur. This can be remedied easily by resting the jaws (open if possi-
ble) of the drying specimen on an outwardly inclined pin placed
before it (fig. 2).
VI. GAUGING
Mounting specimens at an approximately uniform height not only
makes for neatness in the collection but facilitates microscopic com-
parisons. Two or three standard heights should be decided upon as
proper for corresponding categories of beetles of different body
depths, and a’ gauge constructed which will automatically elevate the
mounting support to the desired distance on the pin.
A glass-topped, slotted pinning block greatly facilitates both pin-
ning and gauging. It can easily be constructed of plaster in a mold
slightly larger (about 4 inch on each side) than the future glass
working surface. The latter is composed of two pieces of plate glass,
each 3 inches square, aligned along one edge but held slightly apart
by two narrow strips cut from the ends of a microscope slide, in-
serted in the slot between the plates at the corners, and cemented
there in an upright position. The thickness of these lateral pieces
should gauge the width of the slot just to accommodate the shaft of
a No. 5 insect pin. Two microscope slides are then placed at right
I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
angles with the top plates, flush with and on each side of the slot
and contiguous with the lateral pieces to which they are cemented.
The space between the slides from the ends of the lateral pieces to
the longitudinal free margins of the slides (a distance equal to the
thickness of the top plates) is filled by a strip of appropriate width
cut the full length of a slide and cemented in place to form the floor
of the slot. If preferred, a strip of hard wood, to cushion the points
of the pins, may be substituted here for glass. The slot depth, from
the free surface of the top plates to the inner edge of the bottom
strip, now measures the width of a microscope slide (about 1 inch)
BB
Y
J
highest mounting gauge \ <
S| \ el:
slot “tf
label gauges LW | |
SN r micro. slides
ee ee ae Sear e —)
plate-glass top side of mold clay
Fig. 5
A slotted pinning block in process of construction.
and gauges the correct position for slips bearing medium to small
beetles on the standard 34-mm. (12-inch) pin. Additional gauges for
lower levels of the mounting slip, as well as for pin-label heights,
may be had simply by inserting small removable glass rectangles of
the proper widths at the ends of the slots, an operation which had
best be performed after the block is otherwise completed. The glass
assemblage is now placed, top downward, in the center of the mold
and mixed plaster of paris poured over it to a depth greater than the
walls of the slot (fig. 5).
A slotted pinning block makes it possible to pin mounting slips
squarely with reference to the transverse axis, yet with the longitudi-
nal axis at a slight angle to the horizontal. It is advisable, when
pinning heavy mounts, to tilt them slightly upward in front in order
to compensate for gravity, and to strengthen them by means of a
drop of cement placed so as to embrace the pin at its point of exit.
NO. 6 PREPARATION OF COLEOPTERA—VALENTINE 13
VII. LABELING
There exists an established precedent of long standing that pin
labels, when oriented parallel to the specimen, should face to the
right. This custom originated at a time when hand-lens observa-
tions were the rule; its function was to insure legibility of labels
when specimens were examined in the left hand, the right remain-
ing free for manipulation of the lens. Since the binocular dissecting
microscope has come into almost universal usage in entomology and
the lifting of specimens with the right hand is, ordinarily, the safer
procedure, it follows that a system which would permit labels on
specimens held in the right hand to be read without rotating them
is to be preferred. Consequently, the writer recommends pinning
labels for slip mounts through the right end so that the printing will
face left. Additional reasons for this procedure are these: There is
usually a pinning space in the uneven right-hand margin of a label;
and also, any identification label thus oriented may, when desired, be
swung into the standard, left-hand position without becoming in-
verted. Labels on specimens pinned individually, without mounting
support, had best be pierced through the center and oriented facing
left, parallel to the specimen.
Double-ply Reynolds bristol board or single-ply Strathmore makes
an admirable label paper of the proper thickness to grip the pin effec-
tively. Should a label work loose, a drop of acetate cement on the
under surface around the pin will serve to fasten it securely.
VIII. MOUNTING DISSECTIONS
Preparations of mouth parts, legs, genitalia, etc., may be attached
dry to the mount bearing the specimens which yielded them. Should
a transparent preparation be required, the subject, partly dried on
absorbent paper, can be both cleared and dehydrated in xylol, and
from this transferred directly to a drop of balsam on the glass or
acetate mounting slip close to the specimen to which it belongs. After
proper orientation of the dissection by means of a needle, a tiny
square of cover glass, or thin acetate sheeting (7.5 gauge), is placed
over it. Another method, useful in attaching cleared preparations to
specimens mounted individually, is to prepare a small balsam mount
on one end of a rectangle of acetate sheeting and run the pin through
the opposite end. When not being examined, such a mount may be
swung out of the way under the specimen.
Bulkier genitalia, which require examination in more than one
plane, may be conveniently prepared for study as cleared objects
14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
by placing them (after dehydration in strong alcohol and clearing
in xylol) in small sections of glass tubing drawn to an appropriate
gauge and filled with balsam or, better still, with a color-stable gum
damar. One end of the tube is left open for addition of more mount-
ing medium as contraction takes place; the other end is closed with
a tiny cork plug, or by embedding it in a small cork block in which
a suitable recess has been cut with a cork borer. Through the free
end of the cork is thrust the pin of the specimen which yielded the
dissection. The tube should eventually be sealed with thick acetate
cement. Examination of its contents should be made in xylol or
cedar oil, where virtually all optical interference due to the curved
glass will be eliminated.
The presence of small air pockets trapped within or adjacent to a
dissection in a freshly made balsam mount need not be viewed with
concern ; after a few hours, these will disappear.
IX. MOUNTING LARVAE
Soft-bodied forms, especially the larvae of Coleoptera, have been
successfully treated exactly as the adults and mounted with them
on the same slips. There is but one departure in technique: While
drying, after having been thoroughly soaked in ether, the specimen
is warmed under an electric lamp sufficiently to cause expansion of
the gas within its body and consequent distention. The heat must
be carefully applied and maintained for a few minutes until the speci-
men has dried in an inflated condition. Too sudden an increase in
heat will cause rupture, too gentle an application will cause collapse
of the body.®
X. CARE OF ACCESSORY MATERIAL
a. Dry storage-—The flat tin boxes in which 50 cigarettes are
sold provide admirable storage facilities for Coleoptera. They may
be made to do double duty if both top and bottom are utilized. After
the metal is first scored with a sharp instrument, two pieces of heavy,
smooth cardboard are attached, one to each of the inner surfaces,
by means of a suitable cement (such as “Metallic X”). To these
cards the ethyl acetate-killed, ether-treated specimens are cemented
in close array, with legs at the sides and antennae pointing backward.
This should be done while the beetles are still relaxed, and the ad-
5 This method has yielded excellent results in the preparation of degreased
and dehydrated lepidopterous larvae of both smooth and hairy species.
NO. 6 PREPARATION OF COLEOPTERA—VALENTINE I5
hesive employed should be acetate cement, somewhat diluted with
ethyl acetate. Each lot, representing a day’s catch or an ecological
aggregate, is circumscribed with an ink line and labeled with the
date and locality, or with the date plus some symbol referring to a
category in the chronologically arranged notes. The boxes are stored
chronologically in cardboard filing cases (12 by 6} by 5 inches),
where they stand vertically, hinge uppermost, each bearing an adhe-
sive-tape label along its upper edge. If the cementing has been care-
fully done and the specimens are clean, the chances of their breaking
loose, even in shipping, are extremely remote. Space for naphthalene
on the bottom of the filing case is provided by the recess in which
the filing mechanism operates. However, should dermestids succeed
in entering the tin boxes, they can be destroyed without disturbing
the contents simply by dousing specimens and all with carbon tetra-
chloride and shutting the lids for a few hours.
The chief advantages gained by the above technique lie in the
visibility of the specimens and in the compactness of their arrange-
ment. If at any time a beetle is required for mounting, it may be
detached immediately from the cardboard after an application, by
brush or pipette, of ethyl acetate. It should then be immersed in the
relaxing bath (XI) where any residual cement will be dissolved.
b. Preservation in fluid —Adequate preservation of soft parts with
minimum hardening of tissues and stiffening of joints are the criteria
for a good entomological preservative. In the writer’s opinion, Bar-
ber’s relaxing mixture (XI) meets these requirements better than
any preservative in common use. Ethyl acetate-killed material, stored
in this fluid either before or after ether treatment, has emerged years
later in an excellent state of preservation and perfectly relaxed. It
has proved particularly useful as a preservative or softener for
carabid beetles captured in molasses traps, the specimens being soaked
in the fluid after having been thoroughly washed and partly dried.
In using Barber’s fluid as a preservative over a long period of time,
it is important to remember to change the supply as often as it be-
comes darkened by dissolved oils. When the bulk of material is great,
several changes may be required.
Because of their hardening properties, neither alcohol nor formalin
should be employed alone as a preservative when flexibility of articu-
lations is prerequisite to the mounting technique adopted. Further-
more, the use of these fluids in killing and preserving seems to in-
hibit, somewhat, the action of degreasing agents. However, the ease
with which specimens can be collected and preserved in alcohol fre-
quently justifies its use, especially when time, simplicity of method,
16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
and quantity of material are considerations, or when a hard fixation
of perishable internal organ systems is desired for purposes of
dissection. If alcohol is used for preserving Coleoptera, strengths
neither exceeding 70 percent nor less than 50 percent are
recommended.
XI. RELAXING
The writer has found the relaxing mixture developed at the United
States National Museum by Herbert S. Barber to be extremely effi-
cient and versatile. In this fluid, ethyl acetate-killed specimens be-
come plastic almost instantly and genitalic dissections may be made
a very short time after immersion. It is invaluable for rejuvenating
old, greasy specimens, and will dissolve every mounting adhesive now
in common use. The formula is quoted below:
Barber's fluid
Ethyilalcohols(osupercent) mene eee enone 265 parts
Wratten uoaicseccsccteseiee ae lee Ce eee 245 parts
Ethyltacetater (aceticrether) pace eee eee 95 parts
Benzolr( benzene) Ms.-nieac tne eee alae 35 parts
Should the benzol separate out, a little alcohol, added slowly wit
shaking, will serve to bring it back into the mixture.
Relaxing alcohol-killed material preparatory to slip mounting pre-
sents a difficult problem. However, if such specimens are completely
dried under a lamp, then slightly moistened with Barber’s fluid and
quickly blotted, recalcitrant appendages can usually be made to re-
main in the desired positions. A simpler and perhaps more efficient
method consists in a thorough soaking (several hours at least) in
chloroform. Articulations then become fairly easy to manage, espe-
cially in the smaller specimens whose appendages, while they are
still moist, readily yield to manipulation with needles.
ACKNOWLEDGMENTS
The author wishes to express his gratitude for the helpful sugges-
tions and encouragement received throughout the course of this
work from Edward A. Chapin, of the United States National
Museum, and from Herbert S. Barber and L. L. Buchanan, of the
Bureau of Entomology and Plant Quarantine, United States Depart-
ment of Agriculture.
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dibs VOLUME 103, NUMBER 7
ie OF ADULT AND. IMMATURE _
COLEOPTERA
(wire oe Puares)
fas DB
ce GARL KESTER DORSEY
ty Bureau of Entomology and Plant Quarantine
| U.S. Department of Agriculture
t
INcee
sea IN
2 CRO VIA
if ee ese Ania |
(PUBLICATION 3697)
CITY OF WASHINGTON
_ PUBLISHED BY THE SMITHSONIAN INSTITUTION
s JANUARY 20, 1943
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 103, NUMBER 7
Sees MUSCULATURE OF THE LABRUM,
LABIUM, AND PHARYNGEAL REGION
OF ADULT AND IMMATURE
COLEOPTERA
(WITH 24 PLATES)
BY
CARL KESTER DORSEY
Bureau of Entomology and Plant Quarantine
U. S. Department of Agriculture
(PUBLICATION 3697)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
JANUARY 20, 1943
The Lord Battimore Press
BALTIMORE, MD., U. 8 A.
THE MUSCULATURE OF THE LABRUM, LABIUM, AND
PHARYNGEAL REGION OF ADULT AND
IMMATURE COLEOPTERA
By CARL KESTER DORSEY
Bureau of Entomology and Plant Quarantine, U. S. Department of Agriculture
(WITH 24 PLATEs)
CONTENTS
PAGE
LISTYIE® GST ECGS 7 RRR RE RN PRR A a 8 2
ienGencral discussion of the musculature: . 6... 0.6 0cc ounce nese ce we 4
Hee NGusclesanrancement im) Coleopterasssssssce oe nee neers 9
MME TAC ee ert rou cite din icles mn mete, | isheterea dpa neater ee Tae END 10
AO ACa ae dec loeye ena ac Oke AGS ears Pe Ce ate 10
Gicindelidae BAS hansen os ce Baie orc ern hate op tee Aen 10
DB yhISCi ela e aire ae tinvanneiey euny sca ckeriy sre) net bai nlctwovaye Reve paceman 10
AMES ETOH ELAN rte 3 Si ctags sens sensvarany & bacca Sons chars vs ios Siete keno ena he Baie II
METI AGT “aise See ae ae Bence Aer eats Mare nana ate es eRe II
Semen S THA IEMO TC Grin, aa echo crn isis, =o enue eis eae aie le a Roch Pear ce ate eae 12
SHUI nel Re eters eeta ein ore ave Oe cece eed Hare Ee age le RP en ee 12
Stes HMA TAECIUG | eaten sie ter ohaeeeae entre’ nuke) cedecncrerkio Mite neato ci eeae 12
oT, TRIE ale eT EL (o) fa Fee Ue ape eee ay Lee eke eae Mee TE get eA hes a a 13
ly cro platlidae ses ese ter tye att cicts croatian aso orci Ee ROLE ene 16
MIE SE TN OCA ence 5-5/5 ce awk clals) 4d comneie ane iets Sop bls, Hey aren ee te wee ae oe 14
SMETICHEIOUIGAG ote MM ank Bed ove sie ou ee le eee eee ER oe ae 14
ROR EIMENITGAE, i evectesaitae Gis cameakrantel UR: oA Grate Ne eae acta ie 14
SITCOM ot ac eetee lerortsier Sarees csisvacte eye ee cerormecobacnse ta ote nes 14
Sepa G EOE) cw anisteec acsoestienisis macpiinyerie dar a's oi eike eines ac aig a aiwe 15
PSV at ME xe ear here aes ee ok se vise FS Saag eens bone ne ee ee 15
PX ascill oid eat ree ses cyshs cranes tetova ie ee ralar elaine Gia Behe ears Oe ree on Ra lonerene 16
INosodenmdridaey oY aatotscees vite ere ee Le cde iss Seen Ramee eTe 16
PELELOCEDIC AG; aces cout oI eRe cia Be Sic te em eet 16
Beam NIE CANS 1s, voc veo. aise eins wicbttea rade treluate © aves Ade ale ake elute atons 16
HBS! Gasset ehh be ieee Aird ota FR ah 5 ehd nie of taieta ale ici gece cane te 16
Mod Achy LG aed sc: eis save ons coos sas lahe are bh a iehe & oem eran eee 17
PROM CGani Harold Cage ex cts weeps eee eae CE SER ER enemas: 17
GEACENATIGAC (cia fiu,eoe.s hike a Seine esis 6 owes s aha ero rene ame orn rae 17
Bree a A a 8h Ss bc tusis cod Deane, sides Pier aot ol peserat chs reiake tony eee entrees 17
IB aIME VEE! pose niches Wan es 2 ete gs eho CON eKIeY cote ee ME tem ater 18
BAT IMES FAL GTi L CL Get Wisi csscia ois vie Sraraye ovens © aeckore satus [asd ates wie wis cle Nabe aaNet Bae ie gene 18
PET AIRE TIGA Ss ee eects te ache Cee eo het At ated et alin Bid hase RNY Oe een ecee 18
THE CS EGA Gy wiiareece asia St eset ela ton kciar tic iatstepoamuae op eate atatae 18
MBE EIONIC ACh fo2 no its, See oe a aes eter eitece See, Cece Ae ate 19
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 103, No. 7
XIII.
XIV.
XVI.
XVII.
XVIII.
XIX.
XX.
XXI.
XXII.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
PAGE
carabacOldeal ic 8 ie. % treiscs «a's nisin’ ea Mught a eave ciate ee ener eee 19
SCarabacidae™ 7,254 haiewalesmiae Se oo Sous oat oe Oe oe 19
Ejucanidae: 4. adda fegaediee ed eal ean Sea ea ee 19
Clerojded ss ins acs, sda ate tan enue eet eee ae eee 21
Dierimestidae’ sca tia nw causes satire ers ei teeta ete ev 21
Cleridae:s co cig oc stores os ae seach oO eae ine te 21
Ostomatidaes joc se.dh-aiein aye thera ntacceten ee, fe reelae al ae ne 22
Melotdea yaa % sm ais tees ois tas ores a nalts ee ene 22
Meloidae™ 3c Sade eh Ge ee Je 22
Mordelloidéass 3; pada uinasc Sc ctaciac oh mehig watts Cite anna 23
Mordellidae. 5. i:.2 av montane earnest Bake See ee eee 23
Ceranbycoided (25 vias dkcars) acauet date acm ae eta eh tent mea 23
Cerambycidae =. n0c Naa secon chek cok aot eid aes ae ee 23
Ceraniby cinae: (ls adamacder ai tte te ela oe cs Coe pe ee eee 23
Priomintae 267k eihtain a een 6s ee teeters ean eae ele 23
Bostrichoideay hyo eee. aera tins save dates aucinle ante ae ee eee 25
Bostrichidaes ¥s)5 2 2cheirae suacnsh cleteetliant ane oes Mn ee 25
Ghrysomieloidess 200 aisatd sole, acces nreas ae ee ape ae la React gear at ee 25
Chrysomelidaes = ).is ao cck y's seine ctare odin Pan Oe Mena Ha ORI 25
Monneudae 3) C/N Lan tava ae ane eta tate eae ROE ee 25
Galérucidae: ide Pete a daldoee ete ae ee ctonaS een eee 25
Platystoinoidea. i Site ass esc oe k race ee Ae eee eee 2
Platystomidae \\7eocr tone ise eee. suate eiatlc Gece a elaiake eel mare 27
Cuteulighoideas 2: 35 ue etek tats tor ARC ais eae ere ste en el a ee 27
Curculionidae, 60k foetal OR ere ne aes 27
Scolytiddet-do 0 sitvewrsneueoe aes cis ate mein ans aoe 27
Brentidae: te seein sta elven tee een the Sane n er: ee 28
Bymexyloideds .s..e orucm tate iene eee neve ater ene ec ce LE eC eee 29
Tey mex VTC ies. ieeccoyh arate cick cots ices tv aeanter tani te ieee oy ogee nae 29
SUmM anys tials hes okey bce arenes cree Cee MEIC ee eee 30
Muscles:considered im: this: study.1.\s4 <cwsciescten kien eee Dee 34
Literattire sconsulted: 20). 03 te2 28 cai eretnwsieie eee eyeeieas ae eee 35
Abbreviations: used:on the: figures), <2 "sca. seein eee ve See ee ee 36
Explanation: of the splatesin,./de soe osauk ec wans men ere Gee eae ar
INTRODUCTION
This paper presents the results of a comparative study of the muscle
arrangement in certain regions of the insect head and stomodaeum.
A preliminary study was made in Orthoptera, Dermaptera, Hymen-
optera and Neuroptera, and was followed by a similar but more in-
clusive study of representatives of adult and immature stages of the
order Coleoptera.
Several species of orthopteroid insects were dissected to determine
the muscular arrangement in a generalized form of insect. The other
orders are included to observe similarities with and variations from
the type of muscle system in Orthoptera, of which Periplaneta ameri-
cana L. is chosen as the most generalized form.
NO. 7 MUSCULATURE OF COLEOPTERA—DORSEY 3
In order to observe carefully the places of origin and insertion of
the labral, labial, hypopharyngeal, and pharyngeal muscles it is neces-
sary to remove the maxillae and mandibles and their muscles. The
maxillary, mandibular, and antennal muscles are not included in this
study.
This particular work evolved from an interest as to how chewing
insects actually accomplish the feat of getting the food into the true
mouth, which, of course, is located in the posterior region of the
cibarium. Much is known as to how these insects procure and chew
their food, but the hypopharynx though tonguelike in position could
not be very efficient in accomplishing the work done by the tongue of
a vertebrate. In many insects the hypopharynx, for all practical pur-
poses, can be considered to be absent; such insects also ingest food
and thrive, so the hypopharynx alone cannot be responsible for the
act of placing the food in the mouth from which point it can be
swallowed.
It is difficult and often impossible to identify corresponding muscles
on a functional basis. A muscle, for example, as found in one species
may be an adductor while its homolog in another species is perhaps
a retractor because of some slight change in the position of the point
of insertion. For this reason the general application of functional
names to insect muscles is limited; a careful study of each species
individually is necessary to ascertain muscle functions.
It is hoped that the facts of the arrangement of muscles, and cer-
tain anatomical observations herein presented will contribute to a
better understanding of the means of ingestion of food by chewing
insects, and also of the homologies of muscles between orders of
insects, or between families within an order.
The Leng system of classification as modified by Boving and
Craighead (1930) is followed, and at least one family from each
superfamily selected from which a genus is chosen for study. When
possible the immature stage of the same genus is used. The species
of Coleoptera included in this study are not considered as representa-
tive, necessarily, of their respective families.
This work is presented in partial fulfillment of the requirements
for the degree of Doctor of Philosophy in the Graduate School,
Entomology Department, University of Maryland. The study was
carried on under the direct supervision of R. E. Snodgrass of the
United States Department of Agriculture, Bureau of Entomology
and Plant Quarantine, who is also a member of the staff of the Uni-
versity of Maryland, Entomology Department. The writer is sin-
cerely grateful to Mr. Snodgrass for his guidance and encourage-
4 SMITHSONIAN MISCELLANEOUS COLLECTIONS | VOL. 103
ment throughout this entire study, and also to Dr. Ernest N. Cory,
head of the Entomology Department, University of Maryland, for
his helpful criticisms. The assistance of the United States National
Museum in the loan of certain immature stages of Coleoptera for
study is appreciated as is also that of several specialists in the Divi-
sion of Insect Identification, Bureau of Entomology and Plant
Quarantine, for their help in the identification of certain species.
I. GENERAL DISCUSSION OF THE MUSCULATURE
Among the orthopteroid insects there is apparently, with minor
variations, a general plan of muscle arrangement. The American
cockroach, Periplaneta americana L., is selected as a representative
of Orthoptera exhibiting the generalized plan of musculature. The
muscles included in this work are numbered for convenience of study
and their homologies in other species and orders are considered.
The muscles are considered as occurring in definite pairs unless
indicated as otherwise. The muscles in the dorsal region of the head
inserted on the stomodaeum posterior to the paired nerve (frontal
connective) that connects the frontal ganglion with the brain are con-
sidered as belonging properly to the frons; those muscles that are
anterior to this nerve belong to the clypeal region proper.
1. Compressor muscle of the labrum (fig. 1)—An asymmetrical
muscle arising in the posterior region of the dorsal wall of the labrum;
inserted along the lateral margin of the ventral wall of the labrum.
2. Dilator muscle group of the labrum (figs. I, 2).—A transverse
band of fibers extending across the ventral wall of the labrum.
3. Ventral (posterior) muscle of the labrum (fig. 2).—Arises in
the posterior region of the frons adjacent to the inner margin of the
antennal rim and is inserted laterally at the base of the labrum on the
torma.
4. Dorsal (anterior) muscle of the labrum (fig. 2)—Arises near
the center in the posterior region of the frons and is inserted me-
dially on the posterior dorsal margin of the labrum.
5, 6, and 7. Dorsal dilator muscles of the cibarium (figs. 1, 2).—
Arise on the clypeal region slightly laterally and, with the exception
of muscle 7, are inserted in the anterior region of the dorsal wall of
the cibarium. Muscle 7 is inserted on the dorsal wall of the cibarium,
midregion, between the transverse muscles, 12, of this area.
8. Dorsal dilator muscle of the cibarium (figs. 1, 1A, 2).—Arises
centrally in the anterior clypeal region ; it surrounds muscle 4 (fig. 1A)
and extends ventrad to be inserted in the anterior region of the dorsal
surface of the cibarium. This muscle was found only in Periplaneta
americana L.
NO. 7 MUSCULATURE OF COLEOPTERA—DORSEY 5
g. Dorsal dilator muscle of the anterior region of the pharynx
(figs. 1, 2).—Arises on the frons anterior to muscle rz and is inserted
on the dorsal wall of the pharynx immediately behind the frontal
ganglion connective.
10. Muscle of hypopharyngeal bar Y (figs. I, 2).—A branched
muscle: branch a arises on the frons immediately laterad of muscle rz
(fig. 1) ; branch 0 arises laterally in the anterior region of the frons.
Both branches are inserted on the dorsal end of the sclerotized bar Y
(fig. 1), branch 0 laterally.
11. Dorsal dilator muscle of the pharynx (figs. 1, 2).—Arises on the
frons immediately laterad of the dorsal muscle of the labrum, 4, and
is inserted slightly laterally on the dorsal wall of the pharynx.
12. Compressor muscle group of the cibarium (fig. 2).—Arises on
one side of the dorsal wall of the cibarium, central region, and ex-
tends transversely across the surface to be attached on the opposite
side.
13. Compressor muscles of the anterior region of the cibarium
(fig. 2) —Arise laterally on the dorsal wall of the cibarium in the
anterior region of muscle group 12; they extend diagonally anteriorly
to be inserted on the dorsal surface of the cibarium mesad of one of
the dorsal dilator muscles of the cibarium, 5.
14. Hypopharyngeal muscle of the mandible (fig. 1).—Arises on
the lateral wall of the mandible and is inserted on the distal end of
sclerite X (fig. 1). This muscle was found only in Orthoptera; it
is, of course, common in apterygote insects and other arthropods.
15. Anterior dorsal dilator muscle of the salivarium (fig. 1).—
Arises on sclerite X adjacent to muscle 76 and is inserted laterally
on the ventral wall of the hypopharynx.
16. Dorsal salivary cup muscle (fig. 1).—Arises on the inner sur-
face of sclerite X and is inserted, near the center, on the dorsal sur-
face of the salivary cup.
17. Anterior salivary cup muscle (fig. 1).—Arises laterally on the
ventral wall of the labium adjacent to the palpal base and is inserted
laterally in the anterior region of the salivary cup anterior to muscle 78.
18. Lateroventral salivary cup muscle (fig. 1).—Arises laterally at
the base of the prementum near the dorsal surface and is inserted
laterally in the anterior region of the salivary cup.
19. Muscle of the hypopharynx (fig. 1).—Arises on the tentorial
bridge immediately dorsad of muscle 20 and is inserted laterally on
the base of the hypopharynx.
20. Dorsal (anterior) muscle of the labium (figs. 1, 3).—Arises
on the tentorial bridge immediately dorsad of muscle 27 and is in-
serted laterally in the anterior region of the dorsal wall of the labium.
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I03
21. Ventral (posterior) muscle of the labium (figs. 1, 3).—Arises
laterally on the tentorial bridge at the posterior end of the anterior
tentorial arm and is inserted laterally on the base of the prementum.
22. Muscle of the prementum (figs. I, 3).—Arises medially in the
posterior region of the submentum and is inserted slightly laterally
on the base of the prementum.
23. Depressor muscle of the labial palpus (figs. I, 3).—Arises
centrally on the dorsal wall of the labium at the base of the glossa
and is inserted dorsally on the base of the palpus.
24. Levator muscle of the labial palpus (figs. 1, 3).—Arises later-
ally at the base of the prementum and is inserted posteriorly on the
base of the labial palpus.
25. Muscle of the labial palpus (figs. 1, 3A).—Arises laterally on
the prementum and is inserted basally and somewhat dorsally on the
second palpal segment.
26. Muscle of the labial palpus (figs. 3, 3A).—Arises in the middle
region of the ventral (posterior) wall of the first palpal segment and
is inserted ventrally (posteriorly) in the middle region of the base
of the third segment.
27. Muscle of the labial palpus (figs. 3, 3A).—Arises anteriorly
at the base of the first palpal segment, inserted on the third segment
basally on the anterior margin.
28. Flexor muscle of the glossa (fig. 3).—Arises medially at the
base of the prementum and is inserted on the ventral wall of the
labium at the base of the glossa.
29. Flexor muscle of the paraglossa (fig. 3).—Arises on the ven-
tral wall of the labium at the base of the prementum and is inserted
basally on the ventral wall of the paraglossa.
30. Ventral dilator muscle of the pharynx (fig. 1).—Arises on the
tentorium and is inserted medially on the pharyngeal region just
within the mouth.
31. Lateral dilator muscles of the pharynx (fig. 2).—Arise later-
ally on the dorsal head region adjacent to the inner margin of the
compound eye posterior to the muscle of hypopharyngeal bar Y, 10,
branch A, and are inserted on the side of the pharynx ventrad of
the dosal dilator muscle of the pharynx, 11 (fig. 1).
32. Dorsal appressor muscle group of the cibarium (fig. 2).—
Arises on the dorsal pharyngeal surface posterior to the frontal
ganglion and extends anteriorly beneath the compressor muscle group
of the cibarium, 12, to be inserted on the dorsal wall of the cibarium
adjacent to the anterior limits of muscle 72. These muscles are found
in a well-developed state in the specimens used in this work only in
Orthoptera, Dermaptera, and Neuroptera.
INO: 7 MUSCULATURE OF COLEOPTERA—DORSEY 7
Two other Orthoptera were studied, the cricket Gryllus assimilis
Fab. and the grasshopper Melanoplus differentialis (Thos.). In the
latter there are no essential differences in the musculature. In Gryllus
the muscle arrangement is practically the same as that found in
Periplaneta except for a few minor variations; the dilator muscle
group of the labrum, 2, is absent, the ventral muscle of the labrum, 3,
is branched, and the muscle of hypopharyngeal bar Y, Io, is not
branched.
As a representative of Dermaptera, Anisolabis maritima Gene was
chosen (figs. 4,5,6). The dorsal (anterior) muscle, 20, of the labium
(fig. 5), the ventral (posterior) muscle, 21, of the labium (fig. 5),
and the muscle of the prementum, 22 (figs. 5, 6), are essentially the
same as those in Periplaneta except that muscle 20 arises posterior
to muscle 22. The muscle of the hypopharynx, 19 (figs. 5, 6), arises
centrally in the middle region of the mentum instead of on the ten-
torial structure as it does in the cockroach; this is an unusual origin
for this muscle, as it usually arises at some point on the posterior
tentorial structure. The insertion of this muscle, however, does not
vary. The flexor muscle of the glossa, 28 (fig. 1), and the depressor
muscle of the labial palpus, 23 (fig. 1), are absent. The muscles of
the labrum, cibarium, and hypopharynx are similar to those of
Periplaneta.
In Neuroptera, Corydalus cornutus L. adult and larval stages were
studied. The salivary muscles, in the adult form (figs. 8. 9), are
similar to those of Periplaneta; the anterior dorsal dilator muscle of
the salivarium, 15 (fig: 1), is absent. The full complement of labial
muscles is present. The muscle of the hypopharynx, 10 (figs. 8, 9),
arises laterally in the posterior region of the gula and is inserted at
the base of the hypopharynx, but not on the sides of the salivary cup
as is the case in Periplaneta. In addition to the usual compressor
muscle of the labrum, 1 (fig. 8), there is another muscle that arises
immediately posterior to muscle 7 and is inserted in the anterior re-
gion of the cibarium on the dorsal wall; this muscle is probably only
a division of muscle 7. The ventral muscle of the labrum, 3, and the
dorsal muscle of the labrum, 4 (fig. 8), are of the general form and
arrangement. Of the dorsal dilator muscles of the cibarium, 5, 6, 7
(fig. 8), muscle 5 is represented by a single pair of muscles ; muscle 6
consists of two pairs, and muscle 7 is also made up of two groups of
muscles on each side of the dorsum of the cibarium. The dorsal dila-
tor muscle of the anterior region of the pharynx, 9 (fig. 7), arises
laterad of the muscle of hypopharyngeal bar Y, ro, instead of mesad
which is the usual way.
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
In the larval stage of Corydalus (figs. I0, 11, 12) the muscles
vary little from those of the adult, with a few exceptions and varia-
tions. Of the salivary muscles, 17 is absent. The dorsal dilator
muscles, 5 and 7 (figs. 10, I2), are represented by a single pair of
muscle groups. The ventral wall of the cibarium is furnished with
well-developed transverse, longitudinal, and diagonal muscle groups
(fig. 11, 33). These muscles were observed only in Neuroptera.
In Hymenoptera, Sphecius speciosus (Dru.) adults were selected
for study (figs. 13, 14, 15, 16). This species possesses muscles that
are massive and somewhat complicated in arrangement, and which
are not easy to homologize with muscles of Periplaneta, though cer-
tain of them can be considered homologous. The flexor muscle of
the paraglossa, 29 (fig. 13), arises. basally on the ventral wall of the
prementum and is inserted at the base of the paraglossa. There is
present only one labial-palpus muscle, 24 (fig. 16); it is similar to
that found in Periplaneta. The dorsal salivary cup muscle, 16 (fig.
13), arises on the inner surface of sclerite X (fig. 14) and is in-
serted laterally on the salivary cup. The anterior salivary cup muscle,
18 (figs. 13, 16), which arises immediately posterior to the levator
muscle of the labial palpus, 24, on the ventral wall of the prementum,
is inserted laterally in the anterior region of the salivary cup. The
muscle of the hypopharynx, 19 (fig. 13), arises in the posterior re-
gion of the head and is inserted in the posterior region of the salivary
cup immediately ventrad and caudad of the dorsal salivary cup
muscle, 16. The ventral muscle of the labium, 21 (fig. 13), takes its
origin on the inner side of the base of the anterior tentorial arm
and is inserted centrally at the base of the prementum. The apparent
dorsal muscle of the labium, 20A (fig. 13), arises on the anterior
tentorial arm slightly above the ventral muscle of the labium, 21, and
is inserted on the base of the cardo proximally, so it probably be-
longs to the maxilla and not to the labium. The ventral muscle of the
labrum, 3 (figs. 14, 15), arises and is inserted in the usual manner.
The dorsal dilator muscles of the cibarium, 5, 6, 7 (figs. 14, 15), are
powerful and specialized. Muscle 5 arises centrally in the anterior
region of the clypeus and is inserted centrally in the anterior region
of the dorsal wall of the cibarium. Muscle 6 arises laterally in the
anterior region of the clypeus adjacent to the clypeal suture and is
inserted on the dorsal wall of the mid-region of the cibarium.
Muscle 7 is posterior to muscle 6 in origin and insertion. The com-
pressor muscle group of the cibarium, 12 (figs. 14, 15), is similar to
that of Periplaneta. The specialized muscle 34 (figs. 14, 15) arises
centrally on the dorsal surface of the modified bar Y (fig. 14) and
NO. 7 MUSCULATURE OF COLEOPTERA—DORSEY 9
extends slightly diagonally and anteriorly to become inserted on the
dorsal wall of the cibarium, anterior region; this is a compressor
muscle of this region that may be homologous with the compressor
muscles of the anterior region of the cibarium, 13 (fig. 2), in the cock-
roach. The two modified sclerites, bars Y (fig. 14), support the sides
of the cibarium and stomodaeum; they are united by a continuously
sclerotized area that strengthens the ventral wall of the stomodaeum.
Muscle 34 (fig. 15) is inserted laterally on the distal end of the bar
Y ; this muscle extends laterally and anteriorly to its point of origin
in the posterior region of the frons adjacent to the inner margin of
the compound eye. The dorsal dilator muscle of the pharynx, 11
(figs. 14, 15), consists of a compact group of fibers and probably is
a homolog of muscle zz in orthopteroid forms. The ventral dilator
muscle of the pharynx, 30 (fig. 14), is especially large and powerful;
it arises on a sclerotized central projection from the anterior (dorsal)
surface of the tentorial bridge by means of a tendon; it is inserted
medially on the ventral surface of the continuous sclerotized area
that supports the stomodaeum. This is a dilator muscle of the
stomodaeum. It was thought on first examination that muscle 35
(figs. 14, 15) was homologous with the muscle of hypopharyngeal
bar Y, ro (fig. 1), in Periplaneta; this is not the case because the
small frontal ganglion with its connectives lies posterior to this muscle,
the function of which is not certain. In the honeybee, Apis mellifera
L., there is a muscle which undoubtedly is homologous with the
muscle of the hypopharyngeal bar Y, ro, found in Periplaneta;, it
consists,of an anterior and a posterior branch inserted on the dis-
tal end of bar Y, and is posterior to the frontal connective nerve.
II. MUSCLE ARRANGEMENT IN COLEOPTERA
The muscles of adult and immature Coleoptera included in this
work are numbered as in Periplaneta and are considered to be
homologous with those of the same designation in the orthopteroid
forms unless indicated as being otherwise.
Certain muscles appear generally and consistently throughout the
adult and immature stages in the order, though there are specialized
muscles found in certain species and also generalized muscles that
are greatly modified in size, shape, origin, and insertion. By con-
sidering the muscles of each superfamily separately it is easier to
interpret homologies and the probable function of the various muscles.
IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
III. CARABOIDEA
In this group adult Harpalus caliginosus F., Carabidae (figs. 17,
18, 19), Tetracha carolina (L.), Cicindelidae, and Lacoophilus sp.,
Dytiscidae, were selected for study. The labral muscles are absent in
the three genera studied. In Harpalus the muscles of the cibarium 5,
6, and 7 (figs. 17, 19) and those of the dorsal and lateral pharyngeal
regions are similar to those found in Periplanecta. Bar Y (fig. 19) is
attached to a subpharyngeal brace. The lateral dilator muscle of the
pharynx, 31 (fig. 19), is not considered, necessarily, as a homolog of
31 in Periplaneta or in any of the other species of Coleoptera since
it varies greatly in size, form, and position; but the function being
the same, it is given the same number. The place of origin of the
labial and hypopharyngeal muscles is unusual. In the anterior region
of the gula, caudad of the posterior tentorial pits there is a central
invagination, or apodeme (fig. 19, PT.N), which is finlike and of
considerable size. The muscle of the hypopharynx, 10, the dorsal and
ventral muscles of the labium, 20 and 21 respectively, and muscle
2IA (fig. 19), arise on this apodeme and not on the tentorial struc-
ture as is the usual manner. It is possible that this apodeme represents
a greatly modified tentorial bridge and tentorial arms. The dorsal
muscle of the labium, 20 (figs. 18, 19), is inserted centrally on a pro-
jection that extends inward from the ventral labial wall in the basal
region; it possibly performs the same function as muscle 20 in Peri-
planeta. Muscle 21A (figs. 18, 19) was found only in this species ;
it probably in some way supplements the action of the other labial
muscles. ;
In Tetracha only the dorsal and ventral muscles of the labium, 20
and 21, respectively, and the muscle of the prementum, 22, are pres-
ent; they arise centrally in the posterior gular region. The dorsal
dilator muscles of the cibarium, 5, 6, and 7, are similar to those of
Harpalus; the compressor muscle group of the cibarium, 12, is more
strongly developed. The dorsal dilator muscle of the pharynx, 11,
is powerfully developed. The tormae of the labrum extend posteriorly
and then bend sharply ventrad to form a rigid lateral support on
each side of the mouth; this rigid brace is continuous with the ventral
head structure. Bar Y projects from the posterior dorsal region of
this torma modification; the muscle of hypopharyngeal bar Y, Io, is
inserted on its distal end.
In Laccophilus the ventral muscle of the labium, 21, originates in
the posterior gular area. The muscle of the hypopharynx, 19, is single,
broad, thin, and arises by means of a bifurcate tendon high on each
anterior tentorial arm. The compressor muscle group of the cibarium,
ENOon 7 MUSCULATURE OF COLEOPTERA—DORSEY ne
I2, is well developed. The dorsal dilator muscles of the cibarium, 5,
6, and 7, are essentially the same as those in Harpalus.
For the larval forms in this superfamily, Amara sp. and Galerita
sp. were studied. In Amara (figs. 20, 21, 22) the levator muscle of
the labial palpus, 24 (fig. 22), is strong and the dorsal and ventral
muscles of the labium, 20 and 21 respectively, are large. The dorsal
dilator muscles of the cibarium, 5, 6, and 7 (figs. 21, 22), are well
developed. The pharynx is small; the dorsal dilator muscle of the
pharynx, II, is absent. Bar Y extends posteriorly along the sides of
the cibarium and is connected with the one on the opposite side by a
long, subpharyngeal, sclerotized area. In Galerita the dorsal and
ventral muscles of the labium, 20 and 21 respectively, are similar to
those in Amara. The dorsal dilator muscles of the cibarium, 5, 6, and
7, are grouped together to form a massive unit of muscle fibers. The
muscle of hypopharyngeal bar Y, ro, is similar to that of Amara.
IV. GYRINOIDEA
The adults studied in this superfamily are Dineutes discolor Aubé
and D. vittatus (Germ.), Gyrinidae. In D. discolor (figs. 23, 24, 25)
the labral muscles are absent. The dorsal dilator muscles of the
cibarium, 5, 6, and 7, the dorsal dilator muscle of the anterior region
of the pharynx, 9 (fig. 24), the muscle of hypopharyngeal bar Y, 10
(fig. 23), and the compressor muscle group of the cibarium, 12 (fig.
24), are all similar to those in Harpalus. The dorsal dilator muscle
of the pharynx, I1, is absent. Bar Y is also continuous with a sub-
pharyngeal support. There is an apodeme originating in the anterior
gular region essentially the same as the one in Harpalus; however,
only the dorsal and ventral muscles of the labium, 20 and 21 respec-
tively (fig. 23), arise on it. The muscle of the hypopharynx, 19
(figs. 23, 25), arises by means of a T-shaped tendon, the lateral arms
of which originate on the walls of the gular suture. The musculature
of D. vittatus is practically the same as that in D. discolor. ;
The larvae used for study are Dineutes sp. (figs. 26, 27, 28). The
dorsal dilator muscles of the cibarium, 5, 6, and 7, and the muscle
of hypopharyngeal bar Y, to (figs. 26, 28), are similar to those of
the larva of Amara sp.; they are proportionately much more massive.
The dorsal dilator muscle of the pharynx, II, is present and the com-
pressor muscle group of the cibarium, 12, is either absent or so poorly
developed that it is not noticeable. The labial muscles are essentially
the same as those of Amara with some variation as to shape. The
muscle of the prementum, 22 (figs. 27, 28), is present in Dineutes.
I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
V. STAPHYLINOIDEA
The adults studied in this superfamily are Silpha americana L.,
Silphidae (figs. 29, 30, 31), and Creophilus villosus Grav., Staphy-
linidae. In Silpha the labral muscles are present. The compressor
muscle of the labrum, 1, is well developed (fig. 31), and the ventral
muscle of the labrum, 3 (fig. 31), is inserted on the labrum by means
of a long, thin tendon. The dorsal dilator muscles of the cibarium,
5, 6, and 7 (figs. 29, 31), the compressor muscle group of the cibarium,
12 (figs. 29, 31), the dorsal dilator muscles of the pharynx, 9 and
Ir (figs. 29, 31), and the muscle of hypopharyngeal bar Y, ro (figs.
29, 31), are all massive and undoubtedly furnish the power for a
strong and efficient sucking mechanism. The musculature of the
labium is essentially the same as that found in Periplaneta, excepting
the salivary muscles, which are not found in Coleoptera. The muscle
of the hypopharynx, 19 (figs. 30, 31), the dorsal and ventral muscles
of the labium, 20 and 21 (figs. 30, 31), the muscle of the prementum,
22 (figs. 30, 31), and the levator muscle of the labial palpus, 24 (fig.
30), are present. The preoral cavity is provided with a sizeable, nar-
row lobe (fig. 31) that extends downward from the cibarial wall and
also is equipped with a suboral grooved shelf (fig. 31) that originates
in the posterior dorsal region of the labium, and which receives the
cibarial lobe in somewhat of a semi-piston-and-cylinder relationship,
possibly to facilitate sucking operations.
In Creophilus the labial muscles are the same in number and ar-
rangement as in Silpha; they arise slightly more posteriorly on the
gular sutures. The labral muscles are similar also except that the
ventral muscle of the labrum, 3, arises on the anterior surface of the
distal end of the anterior tentorial arm at the junction of the arm
and the head wall. The muscles of the cibarial and pharyngeal regions
are practically the same as those found in Silpha.
Silpha americana L., Silphidae (figs. 32, 33, 34), and a specimen
of Staphylinae of the Staphylinidae are the two larval forms used
for study. The muscles in Silpha are very well developed. The com-
pressor muscle of the labrum, 1, is absent and the ventral muscle of
the labrum, 3 (figs. 32, 34), is strong. The dorsal dilator muscles
of the cibarium, 5, 6, and 7 (figs. 32, 34), have become grouped to-
gether in a large mass. The compressor muscle group of the cibarium,
12 (figs. 32, 34), is well developed as are the dorsal dilator muscles
of the pharynx, 9 and 11 (figs. 32, 34). The muscle of hypo-
pharyngeal bar Y, ro (fig. 34), is strong and multibranched. Muscle
IIA (fig. 34), is an additional dorsal dilator of the pharynx which
possibly belongs to the dorsal dilator muscle group of the pharynx,
NO 7 MUSCULATURE OF COLEOPTERA—DORSEY 13
11. The labium is a good example of the prementum with two divi-
' sions I and 2 (fig. 33). The muscle of the hypopharynx, 19 (figs. 33,
34), the dorsal and ventral muscles of the labium, 20 and ar (figs.
33, 34), and the muscle of the prementum, 22 (fig. 34), are all present,
well developed, and take their origin on the tentorial structure. In
the specimen of Staphylinae the muscle of the prementum, 22, is in-
serted on the prementum at the base in the same plane as the dorsal
and ventral muscles of the labium, 20 and 21. The prementum is not
divided into two parts as in Silpha. The cibarial, pharyngeal, and
hypopharyngeal muscles are much the same as those in Silpha, with
a few variations. The labral muscles are absent; a tendon arises on
the anterior tentorial arm, dorsal region, and extends anteriorly to
be inserted basally and laterally on the nasale in a manner very simi-
lar to the insertion of the ventral muscle of the labrum, 3, in the adult
Creophilus villosus Grav.
VI. HYDROPHILOIDEA
Sphaeridium scaraboides Linn., Hydrophilidae (figs. 35, 36, 37),
is the adult form studied. The muscles, though well developed, are
reduced in number. Muscle 1 (fig. 35) possibly corresponds with
the compressor muscle of the labium, 1, in Periplaneta; the insertion
has become changed from the ventral wall of the labrum to a sclero-
tized mesal extension of the torma which is continuous with the dor-
sal wall of the anterior region of the cibarium. The ventral muscle
of the labrum, 3 (figs. 35, 37), arises on the distal end of the dorsal
branch of the anterior tentorial arm and is inserted in the usual way.
The dorsal dilator muscles of the cibarium, 5, 6, and 7, are absent.
The anterior tentorial arms are joined by a sclerotized suboesophageal
brace, SB (fig. 37); a lateral dilator muscle of the pharynx arises
on this brace. The dorsal and ventral muscles of the labium, 20 and
2I (figs. 36, 37), both take their origin in the anterior area of the
postmental region.
Sphaeridium bipustulatum Fabr. (figs. 38, 39, 40) and Hydrous
triangularis (Say) are the larvae studied in the family Hydrophilidae.
In Sphaeridium the labral muscles are absent. The dorsal dilator
muscles of the cibarium, 6 and 7 (figs. 38, 39), are present ; muscle 6
is long, 7 compact and linear. The dorsal dilator muscles, 9 and 11
(fig. 39), are well developed as is also the muscle of hypopharyngeal
bar Y, ro (fig. 39). Bar Y is continuous with the sclerotized sub-
pharyngeal area. The three muscles 36, 37, and 38 (fig. 39) are
probably maxillary muscles. Muscle 36 arises laterally from the pos-
terior region of this sclerotized subpharyngeal area and 37 and 38
I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
arise on a median apodeme in the posterior gular region; all these
muscles are inserted at the base of the maxilla. There is a small labial
muscle, probably the ventral muscle of the labium, 21 (figs. 39, 40),
that arises at the base of the postmentum adjacent to the posterior
tentorial pits; it is inserted medially and basally on the prementum.
In Hydrous there are five pairs of powerful muscles that arise
on the anterior tentorial arm and are inserted at the base of the
maxilla. There is a muscle present, probably the ventral muscle of
the labium, 21, and another that is inserted on the dorsal region of
the labium which may be the dorsal muscle of the labiwm, 20. The
dorsal dilator muscles of the cibarium, 6 and 7, the dorsal dilator
muscles of the pharynx, 9 and 11, the muscle of hypopharyngeal bar
Y, ro, and the compressor muscle group of the cibarium are all similar
to those in Sphaeridium though they are much more massive. The
ventral muscles of the pharynx are large and complicated in
arrangement.
Vit CUCUIOIDEX
In this group Scotobates calcaratus (Fab.), Tenebrionidae, and
Epilachna corrupta Muls., Hippodamia convergens Guer., Coccinel-
lidae, adults were used for study. In Scotobates (figs. 41, 42, 43)
the labral, cibarial, pharyngeal, and hypopharyngeal muscles are pres-
ent and well developed. Bar Y (fig. 41) is connected with the anterior,
distal edge of the anterior tentorial arm by means of a flat, sclerotized
band. The ventral muscle of the labium, 21 (figs. 41, 42), may be
homologous with that of Periplaneta, but its origin is on the anterior
region of the postmentum and not on the tentorium. The dorsal
muscle of the labium, 20 (figs. 41, 42), arises in the usual manner,
but it is inserted medially on the inner side of the anterior labial wall
and might easily be interpreted as the muscle of the hypopharynx, 10.
In Epilachna there is only one labial muscle; it corresponds with
the ventral muscle of the labium, 21, in Scotobates. The muscles of
the labrum, cibarium, and pharynx are essentially the same as those
in Scotobates, with a slight variation in arrangement. The muscle of
hypopharyngeal bar Y, 10, lies laterad of the ventral muscle of the
labrum, 3. The dorsal dilator muscles of the pharynx are very
powerful.
In Hippodanua convergens Guer. the musculature is practically the
same as in Epilachna. The compressor muscle group of the cibarium,
12, is well developed.
Scotobates calcaratus (Fab.), Tenebrionidae, Hippodamia con-
vergens Guer., Coccinellidae, and Synchroa puncta Newn., Syn-
NO. 7 MUSCULATURE OF COLEOPTERA—DORSEY 15
chroidae, larvae were studied. In Scotobates (figs. 44, 45, 46) the
muscles are well developed. The compressor muscle of the labrum,
1, the dorsal dilator muscle of the pharynx, 11, and the compressor
muscle group of the cibarium, 12, are absent. The dorsal dilator
muscles of the cibarium, 5, 6, and 7 (figs. 44, 46), have become
grouped together and are indicated only by 7. The muscle of hypo-
pharyngeal bar Y, ro (fig. 44), has numerous subdivisions. There
is a large, heavily sclerotized median cuspidate sclerite (CPS) on
the dorsal surface of the hypopharynx (fig. 46). An arm extends
posteriorly and ventrally from each side of the posterior region of
this sclerite to become joined to a heavily sclerotized intralabial brace
the lateral limits of which are the sides of the posterior hypo-
pharyngeal region. The ventral muscle of the labium, 21 (figs. 45,
46), arises and is inserted in the usual manner; the dorsal muscle of
the labium, 20 (fig. 46), arises in the posterior lateral region of the
submentum just anterior to the posterior end of the anterior tentorial
arm. The muscle of the prementum, 22 (fig. 45), is a broad, powerful
muscle.
In Hippodamia only the ventral muscle of the labium, 21, is present.
The ventral muscle of the labrum, 3, is absent; the tormae are slender
and rodlike and extend posteriorly to become united with hypopharyn-
geal bar Y. The dorsal dilator muscles of the pharynx, 9 and I1, are
especially powerful.
In Synchroa the musculature is similar to that of Scotobates though
the compressor muscle group of the cibarium, 12, is present and the
muscle, of hypopharyngeal bar Y, 10, is not so profusely branched.
VIII. BYRRHOIDEA
Only larval forms were available for study in the family Byrrhidae.
In Byrrhus (figs. 47, 48, 49) the labral muscles are absent. The dor-
sal dilator muscles of the cibarium, 5, 6, and 7 (figs. 47, 48), are re-
duced in size and are grouped together. The dorsal dilator muscle of
the anterior region of the pharynx, 9 (figs. 47, 48), is small; the
muscle of hypopharyngeal bar Y, 10 (figs. 47, 48), consists of two
large branches, ro and 10d. The labial muscles are somewhat con-
fusing because of an additional labial muscle, 224 (figs. 48, 49).
The regular labial muscles, 20 and 27, occur as usual and the muscle
of the prementum, 22 (figs. 48, 49), is consistent. The additional
labial muscle, 224A, may be a second muscle of the prementum inserted
at the base of prementum 1 (figs. 48, 49).
16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
IX. DASCILLOIDEA
From this superfamily adult and immature stages of Nosodendron
californicum Horn, Nosodendridae, and Heterocerus sp., Hetero-
ceridae, larvae were studied. In the adult Nosodendron (figs. 50,
51, 52), of the labral muscles only the ventral muscle of the labrum, 3
(figs. 51, 52), is present. The cibarial, pharyngeal, and hypopharyn-
geal muscles are well developed. The muscle of hypopharyngeal bar
Y, ro (figs. 51, 52), is especially massive. Bar Y (fig. 52) is greatly
modified ; it forms a strong, lateral support to the mouth and is con-
tinuous with a broad, thin, sclerotized structure that extends ven-
trally and posteriorly to become united. with the ventral head wall.
The dorsal and ventral muscles of the labium, 20 and 21 (figs. 50, 52),
are present; they originate in the gular area posterior to the submen-
tum. The mentum completely covers the prementum in this species
(fig. 52).
In the larval Nosodendron (figs. 53, 54, 55) the labral muscles are
absent as is also the compressor muscle group of the cibarium, 12
(fig. 54). The dorsal dilator muscles of the cibarium, 5, 6, and 7,
the muscle of hypopharyngeal bar Y, ro, and the dorsal dilator muscles
of the pharynx, 9 and II, are all massive and are collected more or
less in the region of the posterior area of the cibarium (figs. 53, 54).
Bar Y (fig. 54) is large and forms a strong support on the sides of
the mouth; it extends ventrally on the sides of the posterior hypo-
pharyngeal region. The full complement of labial muscles is present.
In Heterocerus the labral, cibarial, pharyngeal, and hypopharyngeal
muscles are similar to those of Nosodendron; the muscle of hypo-
pharyngeal bar Y is trifurcate. The dorsal muscle of the labium, 20,
arises at the posterior end of the anterior tentorial arms and is
inserted on the dorsal wall of the labium at what is probably the base
of the hypopharynx. The ventral muscle of the labium, 21, takes its
origin medially in the posterior region of the postmentum and is
inserted medially at the base of prementum z. Muscle 22 (probably
the muscle of the prementum) originates on a transverse bar that
connects the anterior tentorial arms; it is inserted laterally in the
posterior region of prementum 2.
X. DRYOPOIDEA
Helichus fastigiatus (Say), Dryopidae (figs. 56, 57, 58), is the
adult form selected for study from this superfamily. The muscula-
ture in this species is, with some modification, of the generalized
plan. The compressor muscle group of the cibarium, 12, is absent
NO. 7 MUSCULATURE OF COLEOPTERA—DORSEY 17
as also is the anterior dorsal dilator muscle of the pharynx, 9, and the
dorsal dilator muscle of the cibarium, 5. Hypopharyngeal bar Y
(fig. 57) is very weakly sclerotized. The labial muscles consist of
two pairs, the dorsal and ventral muscles of the labium, 20 and 21
(figs. 57, 58).
The larvae considered from the family Ptilodactylidae are Ptilo-
dactyla serricollis (Say) (figs. 59, 60, 61). The ventral muscle of
the labrum, 3, two dorsal dilator muscles of the cibarium, 6 and 7,
and the muscle of hypopharyngeal bar Y, ro (figs. 59, 61), are present
and well developed. The dorsal and ventral muscles of the labium,
20 and 21 (figs. 60, 61), are present. Muscle 20 originates medially
at the inner base of the anterior tentorial arms; muscle 27 arises on
the base of the submentum between the posterior ends of the anterior
tentorial arms.
XI. CANTHAROIDEA
Adults from two families in this group were studied, Chauliogna-
thus pennsylvanicus Deg., Cantharidae, and Plateros timidus Lec.,
Lycidae.
In Chauliognathus (figs. 62, 63, 64) the muscles are very well de-
veloped; those of the labrum, cibarium, pharynx, and hypopharynx
are massive (figs. 62, 63). The transverse compressor muscle group
of the cibarium, 12, is absent. Hypopharyngeal bar Y (fig. 62) is
continuous with the greatly modified tentorial structure that forms
a strong support to withstand the action of the powerful muscles of
this region; it unites the dorsal and ventral head walls rigidly. One
side of the structure is joined with the other by means of a sub-
pharyngeal sclerotized area. The posterior (ventral) ends are con-
tinuous with the ventral head wall. The epipharyngeal region of the
labrum and the posterior dorsal labial region are equipped with a
lobe and a grooved receptacle respectively, similar to that already
described in the adult Silpha americana L., but in Chauliognathus it
is more definitely associated with the epipharyngeal area. The labial
muscles are much the same as those of the generalized form and are
not nearly so powerful proportionately as are those of the dorsal
head regions. Each lateral posterior margin of the labium is fur-
nished with a slender, sclerotized resilient bar, RB (fig. 62), which
abuts against the anterior external face of the rigid tentorial structure.
This resilient bar probably assists the labium in returning to its nor-
mal resting position after contractions of the labial muscles have
disturbed its position.
18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
In Plateros there is only one pair of labial muscles, the ventral
pair, 27; they arise on the posterior ends of the anterior tentorial
arms and are inserted medially on the base of the prementum. The
levator muscle of the labial palpus, 24, is present. The rigid tentorial
structure that was found in Chauliognathus is absent in Plateros. The
labral, cibarial, pharyngeal and hypopharyngeal muscles are all essen-
tially the same as in Chauliognathus.
Chauliognathus pennsylvanicus Deg., Cantharidae (figs. 65, 66,
67), and Photuris sp., Lampyridae, are the larval forms considered.
In Chauliognathus (figs. 65, 66) muscle 5 does the same work as
the dorsal dilator muscle of the cibarium, 5 (figs. I, 2), in the gen-
eralized form; it is considered homologous though it is greatly
modified in origin and somewhat in its insertion. The other dorsal
dilator muscles of the cibarium, 6 and 7 (figs. 65, 67), form a com-
pact, linear group as do also the dorsal dilator muscles of the pharynx,
11 (fig. 67). There is a heavy tentorial modification present, but it
is less extensive than that of the adult; it is also continuous with
the subpharyngeal plate. Hypopharyngeal bar Y (fig. 67) extends
posteriorly from the dorsal region of the structure. There is a
heavily sclerotized postoral subpharyngeal support that joins the head
on each side laterad of the mouth. The labial muscles are similar to
those of the adult and the posterior lateral sclerotized resilient bar,
RB (fig. 67), that was described in the adult form is also present in
the larvae.
In Photuris sp. the musculature is similar to that of Chauliogna-
thus; the muscles, however, are less well developed.
XII. ELATEROIDEA
Neotrichophorus carolinensis (Schfr.), Elateridae, and Chryso-
bothris femorata (Oliver), Buprestidae, are the adult forms from
this group used for study.
In Neotrichophorus (figs. 68, 69, 70) the full complement of labral
muscles is present. Upon first examination it was thought that only
the ventral muscle of the labrum, 3 (figs. 68, 70), was present, but
it was later determined that muscle 3 and the dorsal muscle of the
labrum, 4, are so closely appressed as to appear as one. Muscle ¥ is
inserted laterally on the posterior dorsal rim of the labrum while
muscle 3 is inserted ventrally and laterally as is the usual manner.
This is the only species in the order Coleoptera, examined in this
work, that exhibited any indication of an anterior (dorsal) labral
muscle. Another elaterid adult, Heteroderes nicholsi Notman, was
NO. 7 MUSCULATURE OF COLEOPTERA—DORSEY IQ
examined, but only the ventral muscle of the labrum, 3, was found
to be present; muscle 3 in this species is unusual in that it originates
laterally in the anterior clypeal region (fig. 74). This condition has
been observed only in this species.
In Neotrichophorus the cibarial, pharyngeal, and hypopharyngeal
muscles are of the usual form (figs. 68, 70). Hypopharyngeal bar
Y more or less parallels the lateral margins of the cibarium and is
joined to the bar Y on the opposite side by a subpharyngeal support-
ing area on which are inserted strong ventral dilator muscles of the
pharynx, 30 (fig. 68). The labial and hypopharyngeal muscles are
of generalized form, though the ventral muscle of the labium, 21, is
absent unless the muscle of the prementum, 22 (figs. 68, 69), has
been misinterpreted as 21, in which case, of course, muscle 22 would
be absent.
In Chrysobothris the labial and palpal muscles are similar to those
in Neotrichophorus; the muscle of the prementum, 22, is absent. The
labral muscles are also similar except that the dorsal muscle of the
labrum, 4, is absent; the ventral muscle of the labrum, 3, arises in
the midfrontal region on the inner (mesal) surface of the suture that
separates the inner margin of the compound eye from the frons. The
cibarial, pharyngeal, and hypopharyngeal muscles are essentially the
same, but are all poorly developed.
Parallelostethus attenuatus (Say) (figs. 71, 72, 73), Alaus sp.,
Elateridae, and Cebrio antennatus Schfr., Cebrionidae, are the larvae
studied in this superfamily.
In Parallelostethus the labral muscles are absent. The cibarial,
pharyngeal, and hypopharyngeal muscles are powerful, but other-
wise similar to those found in the adult Neotrichophorus; hypo-
pharyngeal bar Y is also similar. The labial muscles, except for being
long and thin (figs. 72, 73), are of the generalized form. The extra
length is made possible by a posterior, spurlike extension of the
anterior tentorial arm (fig. 73) ; the dorsal and ventral muscles of the
labium, 20 and 21, arise on the inner surface on the posterior tip
of this spurlike extension.
In Alaus sp. the musculature is essentially the same as that in Paral-
lelostethus, and in the cebrionid larva, Cebrio, the muscles were found
to be practically the same as those in the two elaterid larvae studied.
XIII. SCARABAEOIDEA
The adults of Popillia japonica Newman and Macrodactylus sub-
spinosus (Fab.), Scarabaeidaé, and Lucanus placidus Say, Lucanidae,
were selected for study in this superfamily.
2
20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
In Popillia (figs. 75, 76, 77), of the labral muscles only the ventral
muscle of the labrum, 3 (figs. 75, 76), is present. The cibarial,
pharyngeal, and hypopharyngeal muscles are similar to those of the
generalized plan except that the dorsal dilator muscle of the pharynx,
II, is absent. Muscle group 13 (figs. 75, 76) may or may not be
homologous with the compressor muscles of the anterior region of
the cibarium, 13 (fig. 2), in Periplaneta; they are well-developed
diagonal muscles inserted on the anterior dorsal wall of the cibarium
and they apparently have the same function. The compressor muscle
group of the cibarium, 12 (fig. 75), is wide, but not long and it is
confined to the posterior dorsal region of the cibarium. There is
only one labial muscle present, 39 (figs. 76, 77); it is a single unit
of fibers arising centrally in the middle region of the labium and
extending slightly anteriorly and dorsally to become inserted me-
dially on the dorsal labial wall. The function, probably, is to assist
in moving the hypopharyngeal region in some manner. The levator
muscle of the labial palpus, 24 (fig. 77), arises on an intralabral
tormalike process extending mesad from the lateroventral base of
the labium; this structure extends in a hoop form dorsally and is
joined to the process on the opposite side of the labium. The ten-
donous insertion end of one of the maxillary muscles enters the in-
tralabial region (fig. 77) and extends through the modified, hoop-
like structure to be inserted on the maxillary base; it has no effect
on the movement of the labium.
In Macrodactylus the median labial muscle, 39, is absent; a muscle
probably corresponding to the ventral muscle of the labium, 21, of
the generalized form arises rather medially in the anterior region of
the postmentum and is inserted ventrally on the base of the pre-
mentum. The cibarial, pharyngeal, and hypopharyngeal muscles are
similar to those in Popillia; the compressor muscles of the anterior
region of the cibarium, 13, are absent as is also the ventral muscle of
the labrum, 3.
In Lucanus the median labial muscle, 39, and the ventral muscle
of the labrum, 3, are absent; the ventral muscle of the labium, 21, is
similar to that in Macrodactylus. The muscle of hypopharyngeal
bar Y, 10, is large and branched because of the fact that bar Y has
a wide, thin, multibranched distal end, and muscle so is, of course,
inserted on these distal branches. There is a muscle group present
that possibly is homologous with the compressor muscles of the an-
terior region of the cibarium, 13, in Popillia; the fibers are arranged
on the dorsal wall of the cibarium more longitudinally than diago-
nally, which is the case in Popillia. The cibarial muscles are all
NO. 7 MUSCULATURE OF COLEOPTERA—DORSEY 21
rather slender and the oesophagus is small considering the size of the
insect.
The larval forms of Scarabaeoidea studied pertain to Popillia
japonica Newman and Cotinis mitida (L.), Scarabaeidae.
In Popillia (figs. 78, 79, 80) the ventral muscle of the labrum, 3
(figs. 78, 79), is strong; the muscle of hypopharyngeal bar Y, ro
(figs. 78, 79), is massive. All of the cibarial muscles are absent.
The dorsal dilator muscle of the anterior region of the pharynx, 9
(fig. 78), is so weak as to be almost threadlike. The compressor
muscle group of the cibarium, 12 (fig. 78), is poorly developed.
There is a large, heavily sclerotized cuspidate sclerite on the posterior
dorsal surface of the labium. The dorsal and ventral muscles of the
labium, 20 and 21 (figs. 78, 80), and the muscle of the prementum,
22 (figs. 78, 80), are well developed.
In Cotinis the labral and labial muscles are similar to those in
Popillia; the muscle of hypopharyngeal bar Y, 10, is also similar.
There is only one pair of weak muscles inserted on the dorsum of
the cibarium just anterior to the frontal connective nerves; it is
probably one of the dorsal dilator muscles of the cibarium, 5, 6, or 7.
A pair of weak dorsal pharyngeal muscles, probably 9, is present.
XIV. CLEROIDEA
Dermestes caninus Germ., Dermestidae, and Enoclerus spinolae
Lec., Cleridae, are the adult forms studied in this superfamily.
In Dermestes (figs. 81, 82, 83) the labral, cibarial, pharyngeal, and
hypopharyngeal muscles are all well developed. The compressor
muscle of the labrum, 1, and 5 of the dorsal dilator muscles of the
cibarium are absent. The muscle of hypopharyngeal bar Y, ro (figs.
81, 82), is branched as in the generalized form. Bar Y is rather
heavy and is continuous ventrally with a sclerotized subpharyngeal
rod that connects the bar Y of one side with that of the other. In
the labium a muscle is present that might be considered either as the
muscle of the prementum, 22, or as the ventral muscle of the labium,
2I (figs. 81, 83). It arises medially in the anterior region of the
submentum and is inserted medially on the base of the prementum.
The muscle 20 (figs. 81, 83) may in reality be homologous with the
muscle of the hypopharynx, 19; it arises on the inner surface of the
posterior end of the anterior tentorial arm and is inserted medially
on the dorsal labial wall in the anterior region by means of a tendon.
In Enoclerus the musculature is similar to that in Dermestes, with
a few variations. One additional pair of ventral muscles of the labium
is present.
22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
For the larval stages, Dermestes caninus Germ., Dermestidae, and
Tenebriodes sp., Ostomatidae, were chosen for study.
In Dermestes (figs. 84, 85, 86) the labral muscles are absent; the
dorsal dilator muscle of the cibarium, 5, the dorsal dilator muscle of
the pharynx, 11, and the compressor muscle group of the cibarium,
12, are also absent. The other cibarial, pharyngeal, and hypopharyn-
geal muscles are present and well developed. The muscle of hypo-
pharyngeal bar Y, ro (figs. 84, 85), is branched and especially mas-
sive. Bar Y (fig. 84) is heavily sclerotized and forms a rigid lateral
support to the dorsal cibarial region and to the mouth. As is the
case in the adult Dermestes, the labial muscle, 21 (figs. 84, 86), is
difficult to homologize with those of the generalized form; the selec-
tion of their numerical designations (names), in this case, is influ-
enced entirely by their points of insertion.
In Tenebriodes sp. the ventral muscle of the labrum, 3, is present.
The cibarial, pharyngeal, hypopharyngeal, and labial muscles are simi-
lar to those of Dermestes; the ventral muscle of the labium, 21, arises
on the tentorial structure and not on the submentum as it does in
Dermestes.
XV. MELOIDEA
Macrobasis immaculata (Say), Meloidae (figs. 87, 88, 89), is the
adult studied in this superfamily. The labral muscles, 1 and 3 (figs.
87, 88), are present and well developed. The dorsal dilator of the
cibarium, 5, is absent; the compressor muscle group of the cibarium,
12, is weak; the other cibarial and pharyngeal muscles are strong.
The proximal (ventral) region of bar Y (fig. 88) supports the ven-
tral pharyngeal wall and the sides of the mouth. In the labium the
levator muscle of the labial palpus, 24 (figs. 88, 89), takes its origin
on the intralabial sclerotized bar, JR (fig. 88), that terminates later-
ally in a broad lateral hypopharyngeal sclerite. The muscle of the
hypopharynx, 19 (figs. 88, 89), arises in the usual area and is in-
serted medially on a short apodeme process by means of a tendon.
The larva studied is Zonabris phalerata (Pall.) (figs. 90, 91, 92).
The muscles are well developed in this species, though the compressor
muscle of the labrum, 1, and the dorsal dilator muscle of the cibarium,
5, are absent. The dorsal muscle of the labium, 20 (figs. 91, 92),
arises in the posterior region of the submentum and the ventral muscle
of the labium, 21 (figs. 91, 92), originates medially in the anterior
region. Hypopharyngeal bar Y (fig. 91) is continuous with a long
sclerotized subpharyngeal structure that joins the two bars together
in the posterior region of the ventral wall of the cibarium.
NO. 7 MUSCULATURE OF COLEOPTERA—DORSEY 23
XVI. MORDELLOIDEA
Mordella quadripunctata (Say), Mordellidae (figs. 93, 94, 95), is
the adult form used for study in this superfamily. The muscles are
powerful and well developed. The cibarial, pharyngeal, and hypo-
pharyngeal muscles are of the generalized form. Hypopharyngeal
bar Y (fig. 93) is weakly sclerotized. The muscle of the prementum,
22 (fig. 95), might be the ventral muscle of the labium, 21, with the
origin moved anteriorly from the tentorial structure rather than
muscle 22, though it is difficult to interpret a condition like this. The
ventral dilator muscle of the pharynx, 30 (fig. 93), and the lateral
dilator muscle of the pharynx, 31 (fig. 93), are present, the latter
being very large. There is a sclerotized subpharyngeal rod (fig.
93) that lies immediately under the mouth opening; it terminates
laterally in the integument of the posterior dorsal hypopharyngeal
area.
The larvae studied are of the same species as the adult (figs. 96,
97, 98). The musculature is, as the figures indicate, well developed
and unusual. There is a large, thin, sclerotized apodeme that extends
ventrally from the center of the middle region of the frons on which
the ventral muscle of the labrum, 3 (fig. 97), the muscle of hypo-
pharyngeal bar Y, 10, 10A, 1oB (fig. 97), and the dorsal dilator
muscle of the pharynx, 11 (fig. 97), all take their origin. The ventral
muscle of the labrum, 3 (figs. 96, 97), is inserted on the distal end
of a modified torma that originates on the ventral labral wall and
which extends laterally and dorsally. The dorsal dilator muscles of
the cibarium, 6 and 7 (fig. 97), are thin and very long; the muscle
of hypopharyngeal bar Y, 10, 10A, 10B (fig. 97), is divided into
three branches and the dorsal dilator muscle of the pharynx, 11 (fig.
97), originates on the ventral margin of the posterior region of the
apodeme. The labial muscles, 20 and 22 (figs. 97, 98), do not re-
quire special comment as the figures are self-explanatory. There is
present a rigid, heavily sclerotized, subpharyngeal intralabial structure,
IR (fig. 97). The crosspiece that lies beneath the anterior region
of the pharynx terminates in the lateral integumental wall of the
posterior labial or hypopharyngeal region.
XVII. CERAMBYCOIDEA
The adults studied in this superfamily are Cyllene robiniae
(Forst.), Cerambycidae, Cerambycinae, and Prionus pocularis Dalm.,
Prioninae.
In Cyllene (figs. 99, 100, 101) the muscles are well developed and
are of the generalized plan. The muscle of hypopharyngeal bar Y,
24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
to (figs. 99, 100), is especially massive. The dorsal wall of the
cibarium is decidedly dome-shaped and appears to be able to accom-
modate the suboral lobe, SL (fig. 99). The dorsal muscle of the
labium, 20 (figs. 99, 101), is inserted laterally on the prementum
near the dorsal surface. In the posterior lateral region of the sub-
pharyngeal area the integument of one side is joined with that of the
opposite by a sclerotized intralabial rod, JR (fig. 99). The anterior
tentorial arm (fig. 99, AT) is broad but thin. The suboral lobe
(SL) is probably the hypopharynx; a muscle interpreted as the
muscle of the hypopharynx, 19 (figs. 99, 101), is inserted medially
at its base.
In Prionus the compressor muscle of the labrum, 1, is absent; the
muscle of hypopharyngeal bar Y, ro, is branched. The labral, cibarial,
pharyngeal, and hypopharyngeal muscles are similar to those in
Cyllene. The labial muscles are similar also, except that the muscle
of the prementum, 22, is absent and the ventral muscle of the labium,
2I, is present.
In the larval Cyllene robimiae (Forst.), Cerambycinae (figs. 102,
103, 104), the muscles are powerful and in the dorsal head region
are usually multibranched or subdivided. The labral muscles are
absent ; the dorsal dilator muscles of the cibarium, 5, 6, and 7 (figs.
102, 104), occur in groups and the muscle of hypopharyngeal bar Y,
ro and 10A (fig. 104), consists of two main branches each of which
is subdivided into several units. The compressor muscle group of the
cibarium, 12 (figs. 102, 104), is strong; immediately beneath this
group the dorsal wall of the posterior region of the cibarium and the
anterior pharyngeal region invaginates to form a pouch which, when
viewed laterally, is somewhat obovate. This pouch or lobe extends
down into the region of the mouth opening. There is a median muscle,
40, beneath the compressor muscle group of the cibarium, 12 (figs.
102, 104), that arises medially in the posterior region of the pouch
and is inserted centrally in the anterior region of the same. The
dorsal dilator muscle of the pharynx, 11 (fig. 104), is inserted on
the posterior dorsal region of the pouch and the dorsal dilator muscle
of the cibarium, 7, in the anterior dorsal region. By the combined
action of muscles 7, 11, 12, and 4o (fig. 104) it would seem that the
lobe formed by the ventral wall of the pouch could pull food into
the mouth from the shelflike hypopharynx, HY, that lies immedi-
ately anterior to it. In the ventral head region the muscle of the
hypopharynx, 19, the dorsal and ventral labial muscles, 20 and 21,
the ventral and lateral dilator muscles of the pharynx, 30 and 31
a7)
NO. 7 MUSCULATURE OF COLEOPTERA—DORSEY 25
(figs. 103, 104), are all well developed and conform rather closely
to the generalized plan of musculature.
XVIII. BOSTRICHOIDEA
Apatides fortis (Lec.), Bostrichidae, adult and immature forms,
were used for study in this superfamily.
In the adult Apatides (figs. 105, 106, 107) the epipharynx is large
and conelike; the compressor muscle of the labrum, I (fig. 106), is
elongate and is inserted laterally near the tip of the epipharynx. The
epipharynx (fig. 106, EP) is supported laterally by a slender sclero-
tized rod (SF) arising dorsally on a tormalike process that extends
mesad from the side of the labrum anterior to the base of the normal
torma. The ventral muscle of the labrum, 3 (figs. 105, 106), is in-
serted on the distal end of the torma which is more elongate than
usual. The cibarial, pharyngeal, and hypopharyngeal muscles are
strong. The dorsal dilator muscle of the cibarium, 7 (figs. 105, 106),
apparently has a branch, 74 (fig. 105), the point of origin of which
has migrated anteriorly to the posterior surface of the epistomal
suture. The labial muscles are well developed; the ventral muscle of
the labium, 21 (figs. 106, 107), possibly in reality is the muscle of
the prementum, 22, since it takes its origin laterally in the posterior
region of the submentum.
The larval Apatides (figs. 108, 109, 110) exhibits the first labral
compressor muscle, I (figs. 108, 109), observed in immature Coleop-
tera. The powerful ventral muscle of the labrum, 3 (figs. 108, 109),
is inserted on the distal end of an elongate torma; the muscle of hypo-
pharyngeal bar Y, 10 and 10A (fig. 108), is branched and strong.
The ventral and lateral dilator muscles of the pharynx, 30 and 31
(fig. 108), are especially well developed. In the labium the dorsal
and ventral labial muscles, 20 and 21 (figs. 108, 110), lie so close
together that superficially they appear to be one muscle. The unusual
muscle 41 (figs. 108, 110) was observed only in this species; it arises
laterally in the anterior region of the submentum and extends dorsally
almost vertically to become inserted laterally at the base of the
labium near the dorsal surface.
XIX. CHRYSOMELOIDEA
Leptinotarsa decimlineata (Say), Chrysomelidae, Donacia distincta
Lec., Donaciidae, and Galerucella xanthomelaena (Schr.), Galeru-
cidae, are the specimens of adults studied in this superfamily.
In Leptinotarsa (figs. III, 112, 113) the muscles of the dorsal
head region are well developed. The muscle of hypopharyngeal bar Y,
206 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
ro (figs. I1I, 112), is branched; the anterior branch ro/A arises
laterally in the anterior frontal area on the posterior surface of the
epistomal suture. The labial muscles are strong, especially the dorsal
muscle of the labium, 20 (figs. 112, 113). There is an unusual ten-
donous structure (fig. 112, TS) that arises broadly, medially and
basally from the ventral labial wall; it extends anteriorly to be
inserted medially on the dorsal wall of the labium in the anterior
region. A contraction of the dorsal labial muscle, 20 (fig. 112),
would exert a pull on the labium at the point of insertion; this tendon
that joins the anterior dorsal labial surface with that of the posterior
ventral region probably retracts the anterior labial area when muscle
20 contracts. There is a strongly sclerotized intralabial brace, JR
(fig. 112), in the posterior region of the labium that terminates
laterally in the integument.
' In Donacia the compressor muscle of the labrum, 1, is absent;
the anterior branch of the muscle of hypopharyngeal bar Y, 10A,
arises as it does in Leptinotarsa. The cibarial, pharyngeal, hypo-
pharyngeal, labral, and labial muscles are apparently homologous with
those found in Leptinotarsa. .
In Galerucella the compressor muscle of the labrum, I, is absent;
the anterior branch of the muscle of hypopharyngeal bar Y, 10A,
arises as it does in Leptinotarsa. The cibarial, pharyngeal, and labial
muscles are homologous with those found in Leptinotarsa.
For the larvae in this superfamily Leptinotarsa decimlineata (Say)
and Typophorus viridicyaneus (Cr.), Chrysomelidae, were selected
for study.
In Leptinotarsa (figs. 114, 115, 116) the muscles are very well
developed. The compressor muscle of the labrum, 1 (figs. 114, 115),
is present; the cibarial muscles, 6 and 7, and pharyngeal muscles, I1
(fig. 114), are long because of the space between the dorsal wall of
the stomodaeum and the head wall. The muscle of the hypopharyngeal
bar Y, ro (fig. 114), is subdivided into many units. The ventral dila-
tor muscle of the pharynx, 30 (fig. 114), is especially massive; the
dorsal and ventral labial muscles, 20 and 21 (figs. 114, 116), and
the muscle of the prementum, 22 (figs. 114, 116), are all strong.
The latter muscle is inserted on the ventral wall of the labium on an
area in which the division of the prementum is indistinct; the dorsal
and ventral labial muscles, 20 and 21, are inserted on the base of
prementum rz (fig. 116). The muscle of the prementum, 22 (fig.
116), is inserted on the base of the second division of the prementum,
though the dividing suture is indistinct.
The musculature in Typophorus is similar to that of Leptinotarsa
except that the compressor muscle of the labrum, 1, is absent.
NO: 7 MUSCULATURE OF COLEOPTERA—DORSEY 27
XX. PLATYSTOMOIDEA
Euparius marmoreus (Oliv.), Platystomidae (figs. 117, 118, 119),
is the adult form studied in this superfamily. The labral muscles are
absent ; the cibarial, pharyngeal, and hypopharyngeal muscles are not
well developed. The hypopharyngeal bar Y (fig. 118) is modified into
a flat plate. There is a heavily sclerotized subpharyngeal rod, JR
(fig. 118), present that supports the ventral region of the mouth and
also strengthens the posterior lateral labial walls in which region
the ends of the rod terminate. The labial muscles are well de-
veloped. The dorsal muscle of the labium, 20 (figs. 118, 119), is an
unusual muscle in this species; it arises laterally in the posterior gular
area as a thickened, powerful group of muscle fibers that quickly
converge to become inserted on the end of a thin tendon in the anterior
gular region. The tendon extends anteriorly and is inserted some-
what laterally in the dorsal region of the posterior labial area.
In the larva of this species (figs. 120, 121, 122) the ventral muscle
of the labrum, 3 (figs. 120, 121), is present. The muscle of hypo-
pharyngeal bar Y, ro (fig. 10), is massive; the dorsal dilator, muscles
of the pharynx, 9 and 11, are absent in this species. The hypopharyn-
geal bar Y (fig. 121) is large; it supports the mouth opening later-
ally and is continuous with a heavily sclerotized suboral lobe that is
joined on each side by a heavily sclerotized bar, JR (fig. 121), that
terminates in the integumental wall of the posterior dorsal lateral
region of the labium. The ventral muscle of the labium, 21 (figs.
I2I, 122), arises on the tentorium at the base of the tentorial arm;
it is inserted medially on the base of the prementum. There is a divi-
sion of the muscle of the prementum, 22 (figs. 121, 122), into two
parts; each arises laterally in the anterior region of the submentum
and each is inserted laterally at the base of the prementum.
XXI. CURCULIONOIDEA
From this superfamily Pantomorus godmani (Crotch), Curculioni-
dae, and Dendroctonous valens Lec., Scolytidae, were selected for
study of the adults.
In Pantomorus (figs. 123, 124, 125) the muscles are reduced in
number and those present are poorly developed. The labial muscles
are absent and the cibarial muscles, 7 (figs. 123, 124), are repre-
sented by a row of weak fibers inserted through the transverse com-
pressor muscle group of the cibarium, 12 (figs. 123, 124), and, since
they all do insert through 12, are probably divisions of 7. The muscle
of hypopharyngeal bar Y, ro (fig. 123), is also weak and bar Y (fig.
28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
123) has practically disappeared ; it is very lightly sclerotized. There
is a large, soft suboral lobe, SL (fig. 123), present that extends
anteriorly into the cibarium in a tonguelike manner (fig. 123). The
labium is furnished with a single pair of muscles (figs. 123, 125)
that probably correspond with the muscle of the prementum, 22, in
the generalized form.
In Dendroctonous the musculature ‘is similar to that of Pantomorus
with a few minor variations.
Pantomorus leucoloma Boh. and Listroderes obliquus Klug., Cur-
culionidae, and Eupsalis minuta Drury, Brentidae, are the larval
species studied.
In Pantomorus (figs. 126, 127, 128) the ventral muscle of the
labrum, 3 (fig. 127), and the muscle of hypopharyngeal bar Y, ro
(fig. 127), are very well developed; the cibarial and pharyngeal
muscles are rather weak. The pharyngeal muscles, 11 (fig. 127), are
probably homologous with the dorsal dilator muscles, 11, in the gen-
eralized plan; they are inserted dorsally on the wall of the pharynx
posterior to the frontal connective. Hypopharyngeal bar Y (fig. 127)
is continuous with a broad lateral and ventral sclerotized plate that
passes beneath the anterior region of the pharynx to become joined
with the bar Y of the other side. A median, flat, sclerotized posterior
extension arises from the posterior dorsal margin of the labrum (fig.
127, ME); it is a resilient piece. When the ventral muscle of the
labrum, 3 (fig. 127), is retracted, the labrum is depressed and the
sclerotized dorsal extension, WE, abuts against the posterior inner
surface of the epistomal suture (fig. 127, ES). When the ventral
muscle of the labrum, 3, relaxes, the resiliency of the extended proc-
ess, ME (fig. 127), pushes the labrum back into its normal position
thus doing the work of a muscle. There is a heavily sclerotized intra-
labial rod, JR (fig. 127), present that terminates laterally on the
posterior lateral walls of the labium and which is joined to the dorsal
labial wall adjacent to the mouth opening. The labial muscles con-
form with those of the generalized form in origin and insertion.
In Listroderes the muscles are essentially the same as those in
Pantomorus. The posterior dorsal margin of the labrum is furnished
with the median posterior extension, ME, also. The dorsal muscle
of the labium, 20, is inserted laterally on the dorsal wall of the labium.
The muscles in Eupsalis are similar to those of Pantomorus. The
dorsal muscle of the labium, 20, is inserted as in the case of Listro-
deres. The dorsal dilator muscle of the anterior region of the pharynx.
9, is present in this species.
NO. 7 MUSCULATURE OF COLEOPTERA—DORSEY 29
XXII. LYMEXYLOIDEA
Hylecoetus lugubris Say, Lymexylidae, adults and larvae were
selected for study in this superfamily.
The muscles of the labrum in the adult (figs. 129, 130) are repre-
sented only by the ventral muscle of the labrum, 3. The pharyngeal
muscles, 9 and 11 (figs. 129, 130), occur in paired groups. The
muscle of hypopharyngeal bar Y, ro (fig. 129), lies between the two
branches of pharyngeal muscle, 9. A small, longitudinal band of
muscles, 13 (figs. 129, 130), is present; it possibly is homologous with
the compressor muscles of the anterior region of the cibarium, 13
(fig. 2), of the general plan. They arise medially, adjacent to the
anterior margin of compressor muscle group of the cibarium, 12
(figs. 129, 130), and are inserted medially and dorsally on the
cibarium a short distance anterior to the point of origin. Hypopharyn-
geal bar Y (fig. 130) is rather large and furnishes a strong support
to the sides of the mouth and to the posterior lateral cibarial region.
The lateral dilator muscle of the pharynx, 31 (fig. 130), is long and
powerful. The muscle of the prementum, 22 (figs. 130, 131), is the
only muscle present in the labium.
In the larvae of Hylecoetus (figs. 132, 133, 134) the labral muscles
are absent; the muscle of hypopharyngeal bar Y, ro, and its branch
10A (figs. 132, 133), are especially massive; the cibarial and pharyn-
geal muscles are comparatively long and slender. Muscle zo (fig.
133) arises by means of a tendon in the posterior region of the frons.
The posterior lateral labial region is furnished with a heavily sclero-
tized intralabial brace, /R (fig. 132), the ends of which terminate in
the integumental wall. Externally there is a heavily sclerotized U-
shaped area present that extends over the dorsal surface of the pos-
terior labial region and is continuous laterally with the ends of the
intralabial brace, /R (fig. 132). The labial muscles present an unusual
plan of arrangement. The prementum is elongate and possesses a
small median sclerite in the anterior region (fig. 134). The muscle
of the prementum, 22 (figs. 132, 134), which is short and very
broad, arises medially along the posterior margin of the tentorial
bridge; it is inserted along the base of the prementum. The ven-
tral muscle of the labium, 21 (figs. 132, 134), originates medially
on the posterior margin of the tentorial bridge and it is inserted rather
medially on the posterior edge of the median premental sclerite.
The dorsal muscle of the labium, 20 (figs. 132, 134), arises laterally
on the inner surface of the posterior end of the anterior tentorial
arm; it is inserted laterally in the anterior region of the median
sclerite of the prementum.
30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
SUMMARY
A general plan of muscle arrangement can be recognized in
orthopteroid insects, and the same plan is found in representatives
of other orders. It is possible to discover apparent homologies among
muscles on the basis of origin and insertion, though one or both
attachments may vary considerably among the different species.
There are certain muscles that evidently are homologous with those
of the generalized plan, but which because of some variation or modi-
fication in origin or insertion are difficult to identify. On the other
hand, some muscles are aberrant and cannot be reconciled with any
plan.
Some unusual muscles and structures, and relationships between
muscles and skeletal parts, were encountered in this study.
The following muscles occurred only in the species of Orthoptera
included in this work: the hypopharyngeal muscle of the mandible,
14 (fig. 1); the anterior dorsal dilator muscle of the salivarium, 15
(fig. 1); the depressor muscle of the labial palpus, 23 (fig. 1); the
muscles of the labial palpus, 25, 26, and 27 (figs. 1, 3A); and the
flexor muscle of the glossa, 28 (figs. I, 3).
In Periplaneta americana L., Orthoptera, the compressor muscle
of the labrum, 1 (fig. 2), is asymmetrical; the dilator muscle group
of the labrum, 2 (fig. 2), and the dorsal dilator muscle of the cibarium,
§ (figs. 1, 1A, 2), were observed only in this species.
The muscle of the hypopharynx, 19 (figs. 5, 6), in Anisolabis
maritima Gené, Dermaptera, arises centrally in the midregion of the
mentum and not on the tentorial bridge as is the usual manner.
In both the adult and larval forms of Corydalus cornutus L.,
Neuroptera (figs. 9, 11), the prementum is divided into three sclerites.
The muscle group 33 (fig. 11) was observed to be especially well
developed only in the larva of Corydalus.
The two muscles 34 and 35 (figs. 14, 15) were observed only in
the adult Sphecius speciosus (Dru.), Hymenoptera. The muscle of
hypopharyngeal bar Y, 10, of the generalized plan is absent; this is
the only case in which this condition was observed. The dorsal muscle
of the labium, 20A (fig. 13), was found only in this species.
In Harpalus caliginosus F., Carabidae (adult), muscle 2rd (fig. 19)
is an additional labial muscle that is not found in the other insects
included in this work. It is in this species also that the posttentorial
apodeme, PTN (fig. 19), occurs; all of the labial muscles arise on this
structure, as does also the muscle of the hypopharynx, 19 (fig. 19). A
similar apodeme is found in Dineutes discolor Aube., Gyrinidae
NO: 7 MUSCULATURE OF COLEOPTERA—DORSEY 31
(adult) (fig. 23, PTN), but the hypopharyngeal muscle, 19, does
not originate on it in this species.
The prementum is divided into two sclerites in the coleopterous
larvae of: Silpha americana L., Silphidae; Byrrhus sp., Byrrhidae;
Nosodendron californicum Horn., Nosodendridae; Hylecoetus lugu-
bris Say, Lymexylidae ; and Leptinotarsa decimlineata (Say), Chrys-
omelidae. The muscle of the prementum, 22, occurs in all of these
larvae; and the muscle of prementum. 1, 22A (fig. 48), was found
only in the larvae of Byrrhus sp., Byrrhidae.
In Sphaeridium scarabaeoides L., Hydrophilidae (adult), the ven-
tral muscle of the labrum, 3 (figs. 35, 37), arises on the distal end
of the anterior tentorial arm instead of on the dorsal head wall as
is the usual manner. In S. bipustulatum Fabr. (larva) muscle 37
(fig. 39) arises on the posterior lateral margin of the subpharyngeal
supporting area and is inserted on the base of the cardo; this muscle
was not observed in other insects.
It was found that in Scotobates calcaratus (Fab.), Tenebrionidae
(adult), the anterior tentorial arm, AT (fig. 41), is connected with
hypopharyngeal bar Y by means of a thin, flat, sclerotized band.
A heavily sclerotized intralabial brace or rod (JR) that terminates
laterally in the integument of the posterior lateral labial region is to
be found in Coleoptera in: Cyllene robiniae (Forst.), Cerambycidae
(adult) (fig. 99, IR); Macrobasis immaculata (Say), Meloidae
(adult) (fig. 88, JR); Leptinotarsa decimlineata (Say), Chryso-
melidae (adult) (fig. 112, IR); Scotobates calcaratus (Fabr.),
Tenebrionidae (larva) (fig. 46, JR); Mordella quadripunctata (Say),
Mordellidae (larva) (fig. 97, IR); Pantomorus leucoloma Boh.,
Curculionidae (larva) (fig. 127, IR); Hylecoetus lugubris Say,
Lymexylidae (larva) (fig. 132, JR).
In Nosodendron californicum Horn, Nosodendridae (adults and
larvae), the hypopharyngeal bar Y (figs. 52, 54) is greatly modified
and enlarged (more so in the adult) and is continuous with the
tentorial structure. In the adult form the mentum covers the pre-
mentum (figs. 50, 52).
In Chauliognathus pennsylvanicus Deg., Cantharidae (adults and
larvae), the tentorial structure (figs. 62, 67, TN) is greatly modi-
field (more so in the adult) ; a sclerotized resilient bar is present in
the integument of the posterior lateral margin of the labium.
Only in Neotrichophorus carolinensis (Schfr.), Elateridae (adult),
of the Coleoptera studied, were both the dorsal (anterior) and ven-
tral (posterior) labral muscles, 4 and 3 (figs. 68, 70), present. In
32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Heteroderes nicholsi Notman (adult) the ventral muscle of the
labrum, 3 (fig. 74), arises laterally on the anterior region of the
clypeus; this is an unusual point of origin for this muscle. In the
larval Parallelostethus attenuatus (Say) there are posterior exten-
sions of the anterior tentorial arms that provide the points of origin
for the long labial muscles, 20 and ar (figs. 72, 73).
In Popillia japonica Newman, Scarabaeidae (adult) (figs. 75, 76,
77), the labial muscles are absent and an unusual muscle in the
labium, 39 (figs. 76, 77), is to be found only in this species.
The levator muscle of the palpus, 24 (figs. 88, 89), in Macrobasis
immaculata (Say), Meloidae (adult), arises on the intralabial brace
(Tie):
In the larvae of Mordella quadripunctata (Say), Mordellidae, there
is present a thin apodeme (fig. 97, AP) that arises medianly in the
posterior region of the frons and extends deeply into the head cavity.
All the labral, cibarial, dorsal pharyngeal muscles, and the muscles of
the hypopharyngeal bar Y, 10, 10A, 10B (fig. 97), arise on this
apodeme.
In the larva of Cyllene robinae (Forst.), Cerambycidae, there is
an uusual lobe that extends ventrally from the dorsal cibarial wall
into the mouth region (figs. 102, 104); the lobe is furnished with
muscle 40 and probably assists in some manner in food ingestion.
The epipharynx is large and unusual in structure in the adult
Apatides fortis (Lec.), Bostrichidae (fig. 106, EP). In the larva
of this species muscle 4r (fig. 108) is unusual in that it is not found
in the other insects studied in this work.
In the adult of Leptinotarsa decimlineata (Say), Chrysomelidae,
branch 10A of the muscle of hypopharyngeal bar Y, to (figs. 111,
112), arises on the posterior surface of the epistomal suture (fig.
111, ES). In this species the ventral part of the labial wall is con-
nected with the anterior dorsal wall by means of an intralabial
tendonous structure (fig. 112, TS).
The dorsal muscle of the labium, 20 (figs. 118, 119), in Euparius
marmoreus (Oliv.), Platystomidae (adult), arises laterally in the
posterior region of the head, posterior to the tentorium, and is
inserted on the labium by means of a very long tendon.
In the larva of Pantomorus leucoloma Boh., Curculionidae, the
resilient median process (fig. 127, ME) that extends posteriorly from
the dorsal margin of the labrum probably does the work of a muscle
in assisting in the return of the labrum to its resting position after
the ventral muscle of the labrum, 3, relaxes.
NO. 7 MUSCULATURE OF COLEOPTERA—DORSEY 33
Further investigations are necessary to determine the nature of
the complete act of the ingestion of food by chewing insects. The
facts presented in this paper indicate, however, that the cibarial,
pharyngeal, and hypopharyngeal muscles, which are powerfully de-
veloped in many species, play an important part in ingestion. The
action of these muscles is undoubtedly supplemented by that of the
labial muscles to a varied degree depending on the species.
onh WN
©
10.
rr
TIA.
12.
12:
14.
15.
16.
17s
18.
19.
20.
21e
2iA.
22.
22A.
23
24.
25.
20.
a7.
28.
20.
30.
ar
22)
33-
34.
35.
36,
39-
40.
4I.
34
s
MUSCLES CONSIDERED IN THIS STUDY
. Compressor muscle of the labrum.
. Dilator muscle group of the labrum.
. Ventral (posterior) muscle of the labrum.
. Dorsal (anterior) muscle of the labrum.
6, 7. Dorsal dilator muscles of the cibarium.
. Dorsal dilator muscle of the cibarium.
. Dorsal dilator muscle of the anterior region of the pharynx.
Muscle of hypopharyngeal bar Y.
Dorsal dilator muscle of the pharynx.
Dorsal dilator muscle of the pharynx.
Compressor muscle group of the cibarium.
Compressor muscles of the anterior region of the cibarium.
Hypopharyngeal muscle of the mandible.
Anterior dorsal dilator muscle of the salivariwm.
Dorsal salivary cup muscle.
Anterior salivary cup muscle.
Lateroventral salivary cup muscle.
Muscle of the hypopharynx.
Dorsal (anterior) muscle of the labium.
Ventral (posterior) muscle of the labium.
Muscle of the labium.
Muscle of the prementum 2.
Muscle of the prementum 1.
Depressor muscle of the labial palpus.
Levator muscle of the labial palpus.
Muscle of the labial palpus.
Muscle of the labial palpus.
Muscle of the labial palpus.
Flexor muscle of the glossa.
Flexor muscle of the paraglossa.
Ventral dilator muscle of the pharynx.
Lateral dilator muscles of the pharynx.
Dorsal appressor muscle group of the cibarium.
Muscles of the ventral wall of the cibarium.
Laterodorsal muscle of the cibarium.
Lateral muscle of the posterior cibarial region.
37, 38. Maxillary muscles.
Median muscle of the labium.
Retractor muscle of anterior pharyngeal region.
Muscle of the labium.
LITERATURE CONSULTED
ANbERSON, W. H.
1936. A comparative study of the labium of coleopterous larvae. Smith-
sonian Misc. Coll., vol. 95, No. 13, 29 pp., 8 pls.
Bovine, A. G., and CratcHeEap, F. C.
1930. An illustrated synopsis of the principal larval forms of the order
Coleoptera. Entomologica Amer., n. s., vol. II, 351 pp., 125 pls.
CARPENTER, G. H., and MacDoweELt, M. C.
1912. Mouthparts of some beetle larvae (Dascillidae and Scarabaeidae)
with especial reference to the maxillulae and hypopharynx. Quart.
Journ. Micr. Sci., vol. 57, pp. 373-396, pls. 35-37.
CRAMPTON, C. C.
1912. The sclerites of the head and mouthparts of certain immature and
adult insects. Ann. Ent. Soc. Amer., vol. 14, pp. 65-103, pls. 2-8.
1928. The eulabium, mentum, submentum and gular regions of insects.
Journ. Ent. Zool., vol. 21, pp. I-15, 3 pls.
Das, G. M.
1937. The musculature of the mouthparts of insect larvae. Quart. Journ.
Micr. Sci., vol. 80, pp. 39-80, pls. 1-12.
DeEGrysE, J. J.
1915. Some modifications of the hypopharynx in lepidopterous larvae.
Proc. Ent. Soc. Washington, vol. 17, pp. 173-178, pls. 17-109.
ErpMaAnn, H.
1925. Vergleichend-anatomische Studien iiber die Pharynx-musculature der
Insekten. Zool. Anz., vol. 62, pp. 49-64.
JANET, C.
1899. Essai sur la constitution morphologique de la téete de l’insecte. 74 pp..
Zepish baris- ,
I91I. Constitution morphologique de la bouche de l’insecte. 35 pp., 2 pls.
Limoges.
Kanpic, O.
1902. Studien tiber das Labium der Coleopteren. Jenaische Zeitschr.
Naturwiss., vol. 36, pp. 207-228.
Rivey, W. A.
1908. Muscle attachments in insects. Ann. Ent. Soc. Amer., vol. I, pp.
265-269, I pl.
Snoperass, R. E.
1928. Morphology and evolution of the insect head and its appendages.
Smithsonian Misc. Coll., vol. 81, No. 3, 158 pp., 57 figs.
1932. Evolution of the insect head and the organs of feeding. Ann. Rep.
Smithsonian Inst., 1931, pp. 443-489, 25 figs.
1935. Principles of insect morphology. 667 pp., 319 figs. New York.
WALKER, E. M.
1931. On the clypeus and labium of primitive insects. Canadian Ent.,
vol. 63, pp. 75-81.
Wit.iams, I. W.
1938. Comparative morphology of mouthparts of Coleoptera. New York
Ent. Soc., vol. 42, pp. 65-85.
3 35
ABBREVIATIONS USED ON THE FIGURES
AF, antennal fossa.
ANT, antenna.
AT, anterior tentorial arm.
CD, cardo.
CIB, cibarium.
CL, clypeus.
CPS, cuspidate sclerite.
CS, coronal suture.
E, compound eye.
EP, epipharynx.
ES, epistomal suture.
FC, frontal connective nerve.
FGN, frontal ganglion.
FR, frons.
FS, frontal suture.
‘GL, glossa.
GU, gula.
HY, hypopharynx.
IR, intralabial rod or brace.
LB, labium.
LBP, labial palpus.
LIG, ligula.
LM, labium.
M, true mouth opening.
MD, mandible.
ME, median posterior extension.
36
MT, mentum.
MX, maxilla.
MXP, maxillary palpus.
NA, nasale.
O, ocellus.
OE, oesophagus.
PGL, paraglossa.
PH, pharynx.
PMT, prementum.
PSMT, postmentum.
PT, posterior tentorial pits.
PTN, posttentorial structure.
RB, resilient bar.
SB, suboesophageal brace.
SC, salivary cup.
SD, salivary duct.
SL, suboral lobe.
SMT, submentum.
SR, sclerotized rod.
ST, stomodaeum.
i torma.
TB, tentorial bridge.
TN, tentorium.
TS, tendinous structure.
Y, hypopharyngeal bar Y.
1A.
3A.
14.
15.
16.
17.
18.
EXPLANATION OF PLATES
PLATE I
. Periplaneta americana L., Orthoptera, sagittal section of head slightly
to the left of median plane.
Same, diagrammatic detail of the relation of the dorsal dilator muscle
of the cibariwm, 8, to the dorsal muscle of the labrum, 4.
Same, dorsal view of head with part of head wall removed to expose
muscles.
Same, ventral view of labium and posterior head region with part of
integument removed to expose muscles.
Same, diagrammatic detail of relationship of the muscles of the labial
palpus, 25, 26, and 27.
PLATE 2
Anisolabis maritima Gené, Dermaptera, dorsal view of head with part
of head wall removed to expose muscles.
Same, sagittal section of head slightly to the left of median plane.
Same, ventral view of head with part of integument removed to expose
muscles.
Corydalus cornutus L., Neuroptera (adult), dorsal view of part of head
with integument removed to expose muscles. j
Same, sagittal section of head slightly to the left of median plane.
Same, ventral view of head with part of integument removed to expose
muscles.
PLATE 3
Corydalus cornutus L., Neuroptera (larva), dorsal view of head with
part of integument removed to expose muscles.
Same, ventral view of head with part of integument removed to expose
muscles.
Same, sagittal view of head slightly to the left of median plane.
. Sphecius speciosus (Dru.), Hymenoptera (adult), lateral view of labium
with part of integument removed to expose muscles and also portion
of anterior tentorial arm with points of muscle origin.
Same, sagittal section of anterior (dorsal) head region slightly to the
left of the median plane.
Same, dorsal view of anterior region of head with part of integument
removed to expose muscles.
Same, ventral view of labium and tentorium with part of integument
removed to expose muscles.
PLATE 4
Harpalus caliginosus F., Carabidae (adult), dorsal view of head with
part of integument removed to expose muscles.
Same, ventral view of labium with part of integument removed to expose
muscles.
37
21.
22.
23.
24.
25.
26.
ar
28.
20.
30.
3i
32.
33.
34.
35.
30.
37-
38.
39.
40.
4I.
42.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Same, sagittal section of head slightly to the left of median plane.
Amara sp., Carabidae (larva), ventral view of labium with part of
integument removed to expose muscles.
Same, dorsal view of head with part of integument removed to expose
muscles.
Same, sagittal section slightly to the left of median plane.
PLATE 5
Dineutes discolor Aubé, Gyrinidae (adult), sagittal section of head
slightly to the left of median plane.
Same, dorsal view of head with part of integument removed to expose
muscles.
Same, ventral view of labium with part of integument removed to expose
muscles.
Dineutes sp., Gyrinidae (larva), dorsal view of head with part of integu-
ment removed to expose muscles.
Same, ventral view of labium with part of integument removed to expose
muscles.
Same, sagittal section of head slightly to the left of the median plane.
PLATE 6
Silpha americana L., Silphidae (adult), dorsal view of head with part
of integument removed to expose muscles.
Same, ventral view of head with part of integument removed to expose
muscles.
Same, sagittal section of head slightly to the left of median plane.
Siulpha americana L., Silphidae (larva), dorsal view of head with part.
of integument removed to expose muscles.
Same, ventral view of labium with part of integument removed to expose
muscles.
Same, sagittal section of head slightly to the left of median plane.
PLATE 7
Sphaeridium scarabaeoides L., Hydrophilidae (adult), dorsal view of head
with part of integument removed to expose muscles.
Same, ventral view of head with part of integument removed to expose
muscles.
Same, sagittal section of head slightly to the left of median plane.
Sphaeridium bipustulatum Fabr., Hydrophilidae (larva), dorsal view of
head with part of integument removed to expose muscles.
Same, sagittal section of head slightly to the left of median plane.
Same, ventral view of labium with part of integument removed to expose
muscles.
Scotobates calcaratus (Fab.), Tenebrionidae (adult), modified sagittal
section of head slightly to the left of median plane.
PLATE 8
Scotobates calcaratus (Fab.), Tenebrionidae (adult), ventral view of
head with part of integument removed to expose muscles.
45.
46.
47.
48.
40.
50.
Gi.
52.
53-
54-
55-
56.
57-
58.
59.
61.
62.
63.
64.
65.
sia MUSCULATURE OF COLEOPTERA—DORSEY 39
Same, dorsal view of head with part of integument removed to expose
muscles.
. Scotobates calcaratus (Fab.), Tenebrionidae (larva), dorsal view of
head with part of integument removed to expose muscles.
Same, ventral view of anterior portion of head with part of integument
removed to expose muscles.
Same, sagittal section of head slightly to the left of median plane.
PLATE 9
Byrrhus sp., Byrrhidae (larva), anterior portion of head with part of
integument removed to expose muscles.
Same, sagittal section of head slightly to the left of median plane.
Same, ventral view of head with part of integument removed to expose
muscles.
Nosodendron californicum Horn, Nosodendridae (adult), ventral view of
head with part of integument removed to expose muscles.
Same, dorsal view of head with part of integument removed to expose
muscles.
Same, sagittal section of head slightly to the left of median plane.
PLATE I0
Nosodendron californicum Horn, Nosodendridae (larva), dorsal view of
head with part of integument removed to expose muscles.
Same, sagittal section of head slightly to the left of median plane.
Same, ventral view of head with part of integument removed to expose
muscles.
Helichus fastigiatus (Say), Dryopidae (adult), dorsal view of head with
part of integument removed to expose muscles.
Same, sagittal section of head slightly to the left of median plane.
Same, ventral view of head with part of integument removed to expose
muscles.
PLATE II
Ptilodactyla serricollis (Say), Ptilodactylidae (larva), dorsal view of
head with part of integument removed to expose muscles.
Same, ventral view of head with part of integument removed to expose
muscles.
Same, sagittal section of head slightly to the left of median plane.
Chauliognathus pennsylvanicus (Deg.), Cantharidae (adult), sagittal sec-
tion of head slightly to the left of median plane.
Same, dorsal view of head with part of integument removed to expose
muscles.
Same, ventral view of head with part of integument removed to expose
muscles.
PLATE I2
Chauliognathus pennsylvanicus (Deg.), Cantharidae (larva), dorsal view
of head with part of integument removed to expose muscles.
Same, ventral view of head with part of integument removed to expose
muscles.
40
67.
68.
77-
78.
79.
80.
81.
82.
83.
84.
85.
86.
87.
89.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Same, sagittal section of head slightly to the left of median plane.
Neotrichophorus carolinensis (Schfr.), Elateridae (adult), sagittal sec-
tion of head slightly to the left of median plane.
Same, ventral view of head with part of integument removed to expose
muscles.
Same, dorsal view of head with part of integument removed to expose
muscles.
PLATE 13
Parallelostethus attenuatus (Say), Elateridae (larva), dorsal view of
head with part of integument removed to expose muscles.
Same, ventral view of head with part of integument removed to expose
muscles.
Same, sagittal section of head slightly to the left of median plane.
Heteroderes nicholsi Notman, Elateridae (adult), lateral view of labrum
and clypeus with part of head wall removed to expose muscles.
. Popillia japonica Newman, Scarabaeidae (adult), dorsal view of head
with part of integument removed to expose muscles.
Same, sagittal section of head slightly to the left of median plane.
PLATE 14
Popillia japonica Newman, Scarabaeidae (adult), ventral view of head
with part of integument removed to expose muscles.
Same (larva), sagittal section of head slightly to the left of median plane.
Same, dorsal view of head with part of integument removed to expose
muscles.
Same, ventral view of head with part of integument removed to expose
muscles.
Dermestes caninus Germ., Dermestidae (adult), sagittal section of head
slightly to the left of median plane.
Same, dorsal view of head with part of integument removed to expose
muscles.
PLATE 15
Dermestes caninus Germ., Dermestidae (adult), ventral view of head
with part of integument removed to expose muscles.
Same, sagittal section of head slightly to the left of median plane.
Same, dorsal view of anterior portion of head with part of integument
removed to expose muscles.
Same, ventral view of head with part of integument removed to expose
muscles.
Macrobasis immaculata (Say), Meloidae (adult), dorsal view of head
with part of integument removed to expose muscles.
Same, sagittal section of head slightly to the left of median plane.
PLATE 16
Macrobasis immaculata (Say), Meloidae (adult), ventral view of head
with part of integument removed to expose muscles.
Zonabris phalerata (Pall.), Meloidae (larva), dorsal view of head with
part of integument removed to expose muscles.
NO.
gl.
92.
93.
94.
95.
096.
97.
08.
99
100.
Iol.
102.
103.
104.
105.
106.
107.
108.
100.
110.
TITS
112.
113.
14.
7 MUSCULATURE OF COLEOPTERA—DORSEY AI
Same, sagittal section of head slightly to the left of median plane.
Same, ventral view of head with part of integument removed to expose
muscles.
Mordella quadripunctata (Say), Mordellidae (adult), sagittal section of
head slightly to the left of median plane.
Same, dorsal view of head with part of the integument removed to expose
muscles.
PLATE 17
Mordella quadripunctata (Say), Mordellidae (adult), ventral view of
head with part of integument removed to expose muscles.
Same (larva), dorsal view of head with part of integument removed to
expose muscles.
Same, sagittal section of head slightly to the left of median plane.
Same, ventral view of head with part of integument removed to expose
muscles.
Cyllene robiniae (Forst.), Cerambycidae (adult), sagittal section of head
slightly to the left of median plane.
PLATE 18
Cyllene robiniae (Forst.), Cerambycidae (adult), dorsal view of head
with part of integument removed to expose muscles.
Same, ventral view of head with part of integument removed to expose
muscles. .
Same (larva), dorsal view of head with part of integument removed to
expose muscles.
Same, ventral view of head with part of integument removed to expose
muscles.
Same, sagittal section of head slightly to the left of median plane.
PLATE 19
Apatides fortis (Lec.), Bostrichidae (adult), dorsal view of head with
part of integument removed to expose muscles.
Same, sagittal section of head slightly to the left of median plane.
Same, ventral view of head with part of integument removed to expose
muscles.
Same (larva), sagittal section of head slightly to the left of median plane.
Same, dorsal view of head with part of integument removed to expose
muscles.
PLATE 20
Apatides fortis (Lec.), Bostrichidae (larva), ventral view of portion of
head with part of integument removed to expose muscles.
Leptinoiarsa decimlineata (Say), Chrysomelidae (adult), dorsal view of
head with part of integument removed to expose muscles.
Same, sagittal section of head slightly to the left of median plane.
Same, ventral view of portion of head with part of integument removed
to expose muscles.
Same (larva), sagittal section of head slightly to the left of median plane.
42
115.
116.
117.
118.
II9.
120.
720:
122:
123.
124.
125.
120.
127.
128.
129.
130.
131.
122,
133,
134.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
PLATE 21
Leptinotarsa decimlineata (Say), Chrysomelidae (larva), dorsal view of
head with part of integument removed to expose muscles.
Same, ventral view of head with part of integument removed to expose
muscles.
Euparius marmoreus (Oliv.), Platystomidae (adult), dorsal view of head
with part of integument removed to expose muscles.
Same, sagittal section of head slightly to the left of median plane.
Same, ventral view of anterior portion of head with part of integument
removed to expose muscles.
Same (larva), dorsal view of head with part of integument removed to
expose muscles.
PLATE 22
Euparius marmoreus (Oliv.), Platystomidae (larva), sagittal section of
head slightly to the left of median plane.
Same, ventral view of head with part of integument removed to expose
muscles.
Pantomorus godmani (Crotch), Curculionidae (adult), sagittal section
of head slightly to the left of median plane.
Same, dorsal view of head with part of integument removed to expose
muscles.
Same, ventral view of anterior portion of head with part of integument
removed to expose muscles.
PLATE 23
Pantomorus leucoloma Boh., Curculionidae (larva), dorsal view of
anterior region of head with part of integument removed to expose
muscles.
Same, sagittal section of head slightly to the left of median plane.
Same, ventral view of head with part of integument removed to expose
muscles.
Hylecoetus lugubris Say, Lymexylidae (adult), dorsal view of head
with part of integument removed to expose muscles.
PLATE 24
Hylecoetus lugubris Say, Lymexylidae (adult), sagittal section of head
slightly to the left of median plane.
Same, ventral view of head with part of integument removed to expose
muscles.
Same (larva), sagittal section of head slightly to the left of median plane.
Same, dorsal view of anterior portion of head with part of integument
removed to expose muscles.
Same, ventral view of head with part of integument removed to expose
muscles.
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 7, PL. 1
MUSCULATURE OF COLEOPTERA
(For explanation of plate see page 37.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 7, PL. 2
<Y
a
a 4),@
TEA
TH )
z=
Ze
Ses
is
MUSCULATURE OF COLEOPTERA
(For explanation of plate see page 37.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 7, PL. 3
MUSCULATURE OF COLEOPTERA
(For explanation of plate see page 37%)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 7. PL. 4
TD eA /)
|
yD
fA)
By.
MUSCULATURE OF COLEOPTERA
(For explanation of plate see pages 37-38.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 7, PL. 5
MUSCULATURE OF COLEOPTERA
(For explanation of plate see page 38.)
VOL. 103, NO. 7, PL. 6
rh
oa
Ty
rea
ao
Ly
SMITHSONIAN MISCELLANEOUS COLLECTIONS
MUSCULATURE OF COLEOPTERA
(For explanation of plate see page 38.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 7, PL. 7
MUSCULATURE OF COLEOPTERA
(For explanation of plate see page 38.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 7, PL. 8
MUSCULATURE OF COLEOPTERA
(For explanation of plate see pages 38-39.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 7, PL. 9
a ee Jj 6 7
YY
Es (0x A
3 {iW YA fp
Cl | i iI Le
MUSCULATURE OF COLEOPTERA
(For explanation of plate see page 39.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 7, PL. 10
MUSCULATURE OF COLEOPTERA
(For explanation of plate see page 39.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103. NO. 7, PL. 11
eS
bE
iy
MUSCULATURE OF COLEOPTERA
(For explanation of plate see page 30.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 7, PL. 12
MUSCULATURE OF COLEOPTERA
(For explanation of plate see pages 39-40.)
SMITHSONIAN MISCELLANEOUS§COLLECTIONS VOL. 103, NO. 7, PL. 13
MUSCULATURE OF COLEOPTERA
(For explanation of plate see page 40.)
VOL. 103, NO. 7, PL. 14
SMITHSONIAN MISCELLANEOUS COLLECTIONS
MUSCULATURE OF COLEOPTERA
(For explanation of plate see page 40.)
NO. 7, PL. 15
VOL. 103
SMITHSONIAN MISCELLANEOUS COLLECTIONS
MUSCULATURE OF COLEOPTERA
(For explanation of plate see page 40.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL, 103, NO.°7, PL. 16
MUSCULATURE OF COLEOPTERA
(For explanation of plate see pages 40-41.)
J
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 7, PL. 17
99 SS Ss U f
3
MUSCULATURE OF COLEOPTERA
(For explanation of plate see page 41.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 7, PL. 18
MUSCULATURE OF COLEOPTERA
(For explanation of plate see page 41.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 7, PL. 19
MUSCULATURE OF COLEOPTERA
(For explanation of plate see page 41.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS
MUSCULATURE OF COLEOPTERA
(For explanation of plate see page 41.)
VOL. 103, NO. 7, PL. 20
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 7, PL. 21
MUSCULATURE OF COLEOPTERA
(For explanation of plate see page 42.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 7, PL. 22
MUSCULATURE OF COLEOPTERA
(For explanation of plate see page 42.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 7, PL. 23
I28
MUSCULATURE OF COLEOPTERA
(For explanation of plate see page 42.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 7, PL. 24
MUSCULATURE OF COLEOPTERA
(For explanation of plate see page 42.)
? si SMITHSONIAN MISCELLANEOUS COLLECTIONS
4 ‘VOLUME 103, NUMBER 8 -
"THE 101! TESTS OF THE LANGLEY
}
Hee)
- “AERODROME”
BY
G. G. ABBOT
Secretary, Smithsonian Institution
Z anise
(PUBLICATION 3699)
GITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION |
OCTOBER 24, 1942
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 103, NUMBER 8
ae 1914 TESTS OF THE LANGLEY
“AERODROME”
BY
CG. G. ABBOT
Secretary, Smithsonian Institution
(PUBLICATION 3699)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
OCTOBER 24, 1942
The Lord Waltimore (Press
BALTIMORE, MD., U. 8. A.
THE 1914 TESTS OF THE LANGLEY “AERODROME”!
By C..G; ABBOT
Secretary, Smithsonian Institution
Note—This paper has been submitted to Dr. Orville Wright, and under
date of October 8, 1942, he states that the paper as now prepared will be
acceptable to him if given adequate publication.
It is everywhere acknowledged that the Wright brothers were the
first to make sustained flights in a heavier-than-air machine at Kitty
Hawk, North Carolina, on December 17, 1903.
Mainly because of acts and statements of former officers of the
Smithsonian Institution, arising from tests made with the recon-
ditioned Langley plane of 1903 at Hammondsport, New York, in
1914, Dr. Orville Wright feels that the Institution adoped an unfair
and injurious attitude. He therefore sent the original Wright Kitty
Hawk plane to England in 1928. The nature of the acts and state-
ments referred to are as follows:
In March 1914, Secretary Walcott contracted with Glenn H. Curtiss
to attempt a flight with the Langley machine. This action seems
ill considered and open to criticism. For in January 1914, the United
States Court of Appeals, Second Circuit, had handed down a decision
recognizing the Wrights as “pioneers in the practical art of flying
with: heavier-than-air machines” and pronouncing Glenn H. Curtiss
an infringer of their patent. Hence, in view of probable further
litigation, the Wrights stood to lose in fame and revenue and Curtiss
stood to gain pecuniarily, should the experiments at Hammondsport
indicate that Langley’s plane was capable of sustained flight in 1903,
previous to the successful flights made December 17, 1903, by the
Wrights at Kitty Hawk, N. C.
The machine was shipped to Curtiss at Hammondsport, N. Y. in
April. Dr. Zahm, the Recorder of the Langley Aerodynamical
Laboratory and expert witness for Curtiss in the patent litigation,
was at Hammondsport as official representative of the Smithsonian
Institution during the time the machine was being reconstructed and
tested. In the reconstruction the machine was changed from what it
was in 1903 in a number of particulars as given in Dr. Wright’s
1For an account of early Langley and Wright aeronautical investigations,
see Smithsonian Report for 1900 and The Century Magazine of September 1908.
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 103, No. 8
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
list of differences which appears later in this paper. On the 28th of
May and the 2d of June, 1914, attempts to fly were made. After
acquiring speed by running on hydroplane floats on the surface of
Lake Keuka the machine lifted into the air several different times.
The longest time off the water with the Langley motor was approxi-
mately five seconds. Dr. Zahm stated that “it was apparent that
owing to the great weight which had been given to the structure
by adding the floats it was necessary to increase the propeller thrust’.
So no further attempts were made to fly with the Langley 52 HP
engine.
It is to be regretted that the Institution published statements
repeatedly * to the effect that these experiments of 1914 demonstrated
that Langley’s plane of 1903 without essential modification was the
first heavier-than-air machine capable of maintaining sustained human
flight.
As first exhibited in the United States National Museum, January
15, 1918, the restored Langley plane of 1903 bore the following label:
THE ORIGINAL, FULL-SIZE
LANGLEY FLYING MACHINE, 1903
For this simple label others were later substituted containing the
claim that Langley’s machine “was the first man-carrying aeroplane
in the history of the world capable of sustained free flight.’
Though the matter of the label is not now an issue, it seems only
fair to the Institution to say that in September 1928, Secretary Abbot
finally caused the label of the Langley machine to be changed to read
simply as follows:
LANGLEY AERODROME
THE ORIGINAL SAMUEL PIERPONT LANGLEY
FLYING MACHINE OF 1903, RESTORED.
Deposited by
The Smithsonian Institution
301,613
This change has frequently been overlooked by writers on the
controversy.
2 Smithsonian Reports: 1914, pp. 9, 219, 221, 222; I9I5, pp. 14, 121; I9I7,
p. 4; 1918, pp. 3, 28, 114, 166. Report of U. S. National Museum, 1914, pp. 46
and 47.
i 2
No. 8
1914 TESTS OF THE LANGLEY “AERODROME’’—ABBOT 3
In January 1942, Mr. Fred C. Kelly, of Peninsula, Ohio, com-
municated to me a list of differences between the Langley plane as
tested in 1914 and as tested in 1903, which he had received from
Dr. Wright. This list is given verbatim below. The Institution accepts
Dr. Wright’s statement as correct in point of facts. Inferences from
the comparisons are primarily the province of interested experts and
are not discussed here.
COMPARISON OF THE LANGLEY MACHINE OF 1903 WITH THE
HAMMONDSPORT MACHINE OF MAY-JUNE, 1914.
LANGLEY, 1903.
HAMMONDSPORT, IQT4.
WINGS.
e
S1zE: 11/6” x 22'6” (L.M. p. 206)
1040 sq. ft. (L.M. p. 206)
1.96
CAMBER: 1/12 (L.M. p. 205)
Leapinc Epce:. Wire 1/16” di-
ameter (L.M. PI1.66)
AREA:
Aspect RATIO:
an b&b WwW HN
6 CovEeRING: Cotton fabric, not var-
nished.
7 CENTER SPAR: Cylindrical wood-
en spar, measuring 14” dia. for
half its length and tapering to 1”
at its tip. (L.M. p. 204). Located
on upper side of wing.
8 Riss: Hollow box construction.
(L. M. Plates 66,67)
SIZE:
AREA: 988 sq. ft.
2.05
10’ 113” x 22'6”
Aspect RatTIo:
1/18
1”
Leapinc Epce: Cylindrical spar 14
dia. at inner end, tapering to 1” dia. at
outer end.
CAMBER:
Covertnc: Cotton fabric, varnished.
CENTER SPAR: Cylindrical spar about
13” dia. at inner end, tapering to about
1” dia. at outer end. Located on upper
side of wing. This center spar was
reinforced (1) by an extra wooden
member on the under side of the wing,
which measured 1” x 14” and extended
to the 7th rib from the center of the
machine; and (2) by another wooden
reinforcement on the under side ex-
tending out about one-fourth of the
length of the wing.
Riss: Most of the original Langley
box ribs were replaced with others
made at Hammondsport. (Manly let-
ter, 1914). The Hammondsport ribs
were of solid construction and made
of laminated wood. That part of the
rib in front of the forward spar was
entirely omitted.
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOL. 103
Go
10
II
I2
13
14
s % Feet
Lower Guy-Posts: A _ single
round wooden post for each pair
of wings (see Fig. 3), 13” in dia.
63’ long. (L.M. Plate 62, p. 184).
The front wing guy-post was lo-
cated 284” in front of the main
center spar. (L.M. Plate 53).
The rear wing guy-post was lo-
cated 315” in front of the main
center spar. (L.M. Plate 53).
Upper Guy-Posts: For each pair
of wings a single steel tube 3” dia.,
43” long. (L.M. p. 184, pl. 62).
Front wing upper guy-post lo-
cated 284” in front of the main
center spar. (L.M. pl. 53).
The rear wing upper guy-post was
located 314” in front of the main
center spar. (LM. pl 53).
Trussinc: The wing trussing
wires were attached to the spars
at the 5th, 7th and oth ribs out
from the center (L.M. pl. 54).
The angles between these wires
HAMMONDSPORT Winc TRuSSING 19/4.
Lower Guy-Posts: Four for each
pair of wings (see Fig. 4), two of
which were of streamline form mea-
suring 11” x 33”x 54” long; and two
measuring 2” x2” with rounded cor-
ners, 3'9” long.
The front wing guy-posts were lo-
cated directly underneath the main
center spar, 283” further rearward
than in 1903.
The rear wing guy-posts were located
directly under the main center spar,
314” further rearward than in 1903.
Upper Guy-Posts: For each pair of
wings, two streamline wooden posts
each 14” x 33”, 76” long, forming an
inverted V. (See Fig. 4).
Front wing upper guy-posts located
directly over main spar, 284” further
rearward than in 1903.
The rear wing guy-posts were located
directly over the main center spar,
314” further rearward than in 1903.
Trussinc: A different system of wing
trussing was used, and the wing trus-
sing wires were attached to the spars
at the 3rd, 6th and oth ribs from the
center. The angles between these wires
No. 8
16
17
18
19
20
21
22
and the spars to which they were
attached are shown in Fig. 3.
IQ14 TESTS OF THE LANGLEY “AERODROME’’—ABBOT 5
and the spars to which they were at-
tached were all different from those
in the original Langley machine. (See
Fig. 4).
CONTROL SURFACES.
VANE Rupper: A split vane com-
posed of two surfaces united at
their leading edges and separated
15” at their trailing edges, thus
forming a wedge. Each surface
measured 2’3” x 4'6”, with aspect
ratio .5. (L.M. p. 214, pls. 53,54).
Operated by means of a wheel lo-
cated slightly in front of the pilot
at his right side and at the height
of his shoulder (L.M. p. 216, pls.
53,54).
Used for steering only. (L.M.
ih Bite
PENAUD TAIL: This was a dart-
shaped tail having a vertical and
a horizontal surface (Penaud
tail), each measuring 95 sq. ft.
It was located in the rear of the
main frame.
Attached to a bracket extending
below the main frame.
“Normally inactive’, (L. M. p.
216) but adjustable about a trans-
verse horizontal axis by means of
a self-locking wheel located at the
right side of the pilot, even with
his back, and at the height of his
shoulder. (L.M. pls. 51, 53).
Immovable about a vertical axis.
(L.M. p. 214, pl.56, Fig. 1). No
means were provided for adjust-
ing this rudder about a vertical
axis in flight. “Although it was
VERTICAL RuppER: The Langley vane
rudder was replaced by a single plane
vertical rudder which measured 3'6” x
5’, with aspect ratio of .7.
Operated at Hammondsport through
the Curtiss steering wheel in some
tests, (Zahm affidavit pp. 5, 6),
through the Curtiss shoulder yoke in
some others (Manly letter, 1914), and
fixed so as not to be operable at all in
still others, (Zahm affidavit p. 7).
Used “as a vertical aileron to control
the lateral poise of the machine”,
(Zahm affidavit p. 6) as well as for
steering, (Zahm affidavit p. 7).
TatL Rupper: Same size and con-
struction as in 1903.
Attached to same bracket at a point
about 8” higher than in 1903.
Operable about a transverse horizontal
axis and connected to a regular Curtiss
elevator control post directly in front
of the pilot (Zahm affidavit p. 5).
Immovable about a vertical axis on
May 28, 1914, only. Thereafter it was
made movable about a vertical axis
and was connected through cables to
a Curtiss steering wheel mounted on a
23
24
to
on
26
SMITHSONIAN MISCELLANEOUS COLLECTIONS
necessary that the large aero-
drome should be capable of being
steered in a horizontal direction,
it was felt to be unwise to give
the Penaud tail and rudder mo-
tion in the horizontal plane in
order to attain this end’. (L.M.
p. 214).
KeeL_: A fixed vertical surface
underneath the main frame mea-
suring 3/2” in height by 6’ aver-
age length. Area 19 sq. ft. (L.M.
pl. 53).
SYSTEM OF
LATERAL STABILITY: The dihedral
only was used for maintaining
lateral balance. (L.M. p. 45).
LONGITUDINAL STABILITY: Lang-
ley relied upon the Penaud system
of inherent stability for maintain-
ing the longitudinal equilibrium.
“For the preservation of the equi-
librium [longitudinal] of the aero-
drome, though the aviator might
assist by such slight movements
as he was able to make in the
limited space of the aviator’s car,
the main reliance was upon the
Penaud tail.” (L.M. p. 215).
STEERING: Steering in the hori-
zontal plane was done entirely by
the split-vane steering rudder lo-
cated underneath the main frame.
(L.M. p. 214).
VOL. 103
Curtiss control post directly in front
of the pilot.
KeeL_: Entirely omitted.
CONTROL.
LATERAL STABILITY: Three means
were used for securing lateral balance
at Hammondsport: The dihedral
angle as used by Langley, a rudder
which “serves as a vertical aileron”
(Zahm affidavit p. 6), and the Penaud
tail rudder. The last two constituted
a system “identical in principle with
that of Complainant’s [Wright] com-
bined warping of the wings and the
use of the vertical rudder”. (Zahm
affidavit p. 6).
LONGITUDINAL STABILITY: At Ham-
mondsport the Penaud inherent longi-
tudinal stability was supplemented
with an elevator system of control.
STEERING: On one day, May 28, 1914,
steering in the horizontal plane was
done with the vertical rudder which
had been substituted for the original
Langley split-vane steering rudder.
After May 28th the steering was done
by the vertical surface of the tail rud-
der (Zahm affidavit p. 7), which in
1903 was immovable about a vertical
axis, (L.M. p. 214).
No. 8
27
28
29
30
31
32
33
34
35
IQI4 TESTS OF THE LANGLEY “AERODROME’’—ABBOT a
POWER PLANT.
Moror: Langley 5 cylinder radial.
IGNITION: Jump spark with dry
cell batteries. (L.M. p. 262).
CARBURETOR: Balzer carburetor
consisting of a chamber filled with
lumps of porous cellular wood
saturated with gasoline. The air
was drawn through this wood.
There was no float feed. (L.M.
p. 225).
RADIATOR:
fins.
Tubes with radiating
PROPELLERS: Langley propellers
(L.M. pl.53, pp. 178-182).
LAUNCHING
LAUNCHING:
on a houseboat.
Catapult mounted
Fioats: Five cylindrical tin
floats, with conical ends, attached
to underside of main frame at ap-
propriate points, and about six
feet above lowest part of machine.
Motor: Langley motor modified.
IGNITION: Jump spark with magneto.
CARBURETOR: Automobile type with
float feed.
Raptator: Automobile radiator of
honeycomb type.
PROPELLERS: Langley propellers modi-
fied “after fashion of early Wright
blades”.
AND FLOATS.
LauncHING: Hydroplanes, developed
1909-1914, attached to the machine.
Fioats: Two wooden hydroplane
floats, mounted beneath and about 6
feet to either side of the center of the
machine at the lateral extremities of
the Pratt system of trussing used for
bracing the wing spars of the forward
wings; and one (part of the time two)
tin cylindrical floats with conical ends,
similar to but larger than the Lang-
ley floats, mounted at the center of the
Pratt system of trussing used for
bracing the rear wings. All of the
floats were mounted from four to
five feet lower than the floats of the
original Langley, thus keeping the en-
tire machine above the water.
WEIGHT.
Tota, WEIGHT: With pilot 850
pounds (L.M. p. 256).
CENTER GRAVITY: 3/8” above line
of thrust.
ToraL WeicHT: With pilot,
pounds.
1170
CENTER Gravity: About one foot be-
low line of thrust.
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Since I became Secretary, in 1928, I have made many efforts
to compose the Smithsonian-Wright controversy, which I inherited.
I will now, speaking for the Smithsonian Institution, make the follow-
ing statement in an attempt to correct as far as now possible acts
and assertions of former Smithsonian officials that may have been
misleading or are held to be detrimental to the Wrights.
1. I sincerely regret that the Institution employed to make the
tests of 1914 an agent who had been an unsuccessful defendant in
patent litigation brought against him by the Wrights.
2. I sincerely regret that statements were repeatedly made by
officers of the Institution that the Langley machine was flown in
1914 “with certain changes of the machine necessary to use pontoons”,
without mentioning the other changes included in Dr. Wright’s list.
3. I point out that Assistant Secretary Rathbun was misinformed
when he stated that the Langley machine “without modification”
made “successful flights’.
4. I sincerely regret the public statement by officers of the Insti-
tution that “The tests” [of 1914] showed “that the late Secretary
Langley had succeeded in building the first aeroplane capable of
sustained free flight with a man.”
5. Leaving to experts to formulate the conclusions arising from
the 1914 tests as a whole, in view of all the facts, I repeat in substance,
but with amendments, what I have already published in Smithsonian
Scientific Series, Vol. 12, 1932, page 227:
The flights of the Langley aerodrome at Hammondsport in
1914, having been made long after flying had become a common
art, and with changes of the machine indicated by Dr. Wright’s
comparison as given above, did not warrant the statements pub-
lished by the Smithsonian Institution that these tests proved
that the large Langley machine of 1903 was capable of sustained
flight carrying a man.
6. If the publication of this paper should clear the way for Dr.
Wright to bring back to America the Kitty Hawk machine to which
all the world awards first place, it will be a source of profound and
enduring gratification to his countrymen everywhere. Should he
decide to deposit the plane in the United States National Museum,
it would be given the highest place of honor, which is its due.
SMITHSONIAN MISCELLANEOUS COLLECTIONS
* VOLUME 103, NUMBER 9
_ MYSTACOCARIDA A NEW ORDER OF
_ CRUSTACEA FROM INTERTIDAL
ie _ BEACHES IN MASSACHUSETTS.
a AND: CONNECTICUT
(With Two PLATES)
BY
ROBERT W. PENNAK
Biology Department, University
of Colorado
_ AND
DONALD J. ZINN
Osbo:'n Zoological Laboratory,
Yale University
(PUBLICATION 3704)
GITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION |
FEBRUARY 23, 1943
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 103, NUMBER 9
MYSTACOCARIDA, A NEW ORDER OF
CRUSTACEA FROM INTERTIDAL
BEACHES IN MASSACHUSETTS
AND CONNECTICUT
(WitTH Two PLATEs)
BY
ROBERT W. PENNAK
Biology Department, University
of Colorado
AND
DONALD J. ZINN
Osborn Zoological Laboratory,
Yale University
(PUBLICATION 3704)
GITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
FEBRUARY 23, 1943
ay es to) i ee a al oo
Pe Tene he
TBe Lord Baltimore Preas
BALTIMORE, MD., U. S. A.
MYSTACOCARIDA, A NEW ORDER OF CRUSTACEA
FROM INTERTIDAL BEACHES IN MASSACHU-
SETTS AND CONNECTICUT
By ROBERT W. PENNAK
Biology Department, University of Colorado
AND
DONALD J. ZINN
Osborn Zoological Laboratory
Yale University
(WirtH 2 PriatEs)
During the course of recent investigations on the ecology of the
micrometazoa inhabiting the capillary waters of intertidal beaches in
Massachusetts and Connecticut (Pennak, 1942a, 1942b; Zinn, 1942),
more than 65 specimens of a small peculiar entomostracan were found.
When these organisms'were first examined superficially it was thought
that they were copepods. In size, in their basic 16-segmented struc-
ture, in the general organization of the body into head, thorax, and
abdomen, and in the number and arrangement of the appendages (see
pl. 1, fig. 3), they appeared to be an aberrant species of Harpacticoida.
A more detailed study, however, revealed that in certain fundamental
anatomical features they were markedly different from copepods—so
different, in fact, as to warrant the erection of a new order, the order
Mystacocarida. A brief diagnosis of this group is given in the follow-
ing paragraph.
ORDER MYSTACOCARIDA
The Mystacocarida are microscopic Entomostraca inhabiting the
interstitial waters of intertidal beaches. Only females are known.
The body is colorless, distinctly segmented, cylindrical, vermiform,
and elongate ; it is divided into a large head, a single-segmented post-
cephalosome, a 4-segmented thorax, and a 6-segmented abdomen. The
head is elongate, with a cephalic shield, prominent rostrum, and ocelli;
it bears two pairs of antennae, a labrum, a pair of mandibles, and two
pairs of maxillae. The first antennae are large, elongate, uniramous,
and sensory. The second antennae are prominent, biramous, sensory,
and elongate. The large labrum is flat, broad, and movable; it is at-
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 103, No. 9
2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
tached to the head between the bases of the antennae and extends pos-
teriorly as far as the postcephalosome. The mandibles are biramous
and unmodified, while the first and second maxillae are uniramous and
elongate. The postcephalosome bears a pair of flat, broad, irregular
mouthparts (ordinarily corresponding to the maxillipeds of other
groups). Each thoracic segment has a pair of small, unsegmented
lamellar appendages. The terminal (anal) segment bears two large
caudal rami which have prominent setae. None of the appendages
are prehensile. The dense setation of the median margins of the
mouthparts indicates that the Mystacocarida probably feed by strain-
ing particles of food from the interstitial water. They move through
the capillary spaces by wormlike wriggling movements aided by
natatory movements of the second antennae, mandibles, and to some
extent the first and second maxillae. Most of the body segments-
are provided with a pair of lateral, dorsoventral, shallow troughs
which have heavily chitinized edges. The indistinct genital aperture
is on the first thoracic segment, and the anus is on the last abdominal
segment between the bases of the caudal rami. Several larval stages
have been found. Only one species is known.
So far as is known, the Mystacocarida are confined to the intertidal
zones of marine beaches. They are a part of the complex community
of bacteria, protozoa, and microscopic metazoa including harpacticoid
copepods, Tardigrada, Nematoda, Oligochaeta, Acarina, and several
other groups which inhabit the capillary water between the grains
of sand. The Mystacocarida are probably facultative anaerobes since
they may occur in portions of the intertidal sand where oxygen may
be either absent or present in small quantities.
Superficially, the Copepoda and the Mystacocarida are somewhat
similar. The two groups may be most easily distinguished from each
other, however, on the basis of the number of segments comprising the
head, thorax, and abdomen. In copepods the maxilliped segment is
fused with five other anterior segments to form the head, and, in
addition, the first thoracic segment is usually fused with the head,
the whole thus constituting a cephalothorax. In the Mystacocarida,
on the other hand, both the maxilliped segment and the first thoracic
segment are distinct and separate from the head. Furthermore, the
maxilliped segment is clearly separated from the thorax by a ventral
constriction. According to the terminology of Sars (1901, 1903-I91T)
as clarified by Monk (1941), copepods typically have five thoracic
(metasome) segments and five abdominal (urosome) segments. Al-
though there are 10 segments posterior to the maxilliped segment in
the Mystacocarida, there is no specialized movable articulation between
NO. 9 A NEW ORDER OF CRUSTACEA—PENNAK AND ZINN 3
thorax and abdomen. The first four segments following the maxilliped
segment, however, bear appendages which presumably are comparable
with the thoracic appendages of the Copepoda. There are no signs
or rudiments of a pair of appendages which would indicate the fifth
thoracic segment. The maxilliped segment is so distinctive and unique,
and is so sharply set off from the adjacent regions of the body in
the Mystacocarida, that it may be conveniently termed a postcephalo-
some. The body of the Mystacocarida therefore may be divided into
a head, a single-segmented postcephalosome, a 4-segmented thorax,
and a 6-segmented abdomen.
Except for the first and second antennae, the appendages of the
Mystacocarida differ markedly from those of the Copepoda (see
pls. 1 and 2). The mandibles are biramous and elongate, the first
and second maxillae are uniramous and elongate, and the post-
cephalosome appendages (maxillipeds) are irregular, broad, and flat.
The large size and the dense median setation of the relatively un-
specialized and primitive, mouthparts indicate that they function in
straining food particles out of the interstitial water of sandy beaches.
The smaller mouthparts of the Copepoda, on the other hand, are
usually composed of relatively few segments and are more or less
specialized for the manipulation of food; this is especially true of
harpacticoid copepods which inhabit sandy beaches. Not only are
the trophic rami of the Mystacocarida large and unspecialized, but
also the head is proportionately much longer than it is in the
Copepoda. All of the thoracic appendages show a reduction and
simplicity found only in the fifth legs of a few copepods. Each leg
is a simple, unsegmented lamella which is probably useless in swim-
ming or crawling.
Between the bases of the mouthparts of the Mystacocarida there
is an enormous, movable, broad, flat labrum. The labrum of the
Copepoda is an insignificant structure.
On the posterior part of the cephalic shield, on the maxilliped
segment, on each thoracic segment, and on each of the first five
abdominal segments, there is a pair of lateral, dorsoventral, shallow
troughs with heavily chitinized, sculptured edges. Structures of a
somewhat similar nature are known in only one species of Copepoda
(Zinn, 1942). No function can be ascribed to them.
Finally, the Mystacocarida may be differentiated from the Copepoda
on the basis of the location of the genital pore; in the former group
it is on the first thoracic segment, while in the latter it is on the last
thoracic segment.
4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
The peculiar location of the genital pore in the Mystacocarida
represents an unusual condition. In the great majority of Crustacea
the genital pore is at or near the posterior end of the thorax. Indeed,
according to the Calman scheme, the thorax is defined as that region
behind the head and in front of, and including, the segment which
bears the male genital pore; also, with the exception of the hermaph-
roditic Cirripedia, the female genital pore is on the last or third-last
thoracic segment. Such a definition is arbitrary, to be sure, but it
is in striking contrast to the situation in the Mystacocarida.
In spite of the numerous significant morphological differences be-
tween the Copepoda and the Mystacocarida, the similar segmentation,
the general body organization, and the number and arrangement of
the appendages constitute convincing evidence that these two orders
are very closely related. Certainly it does not seem possible to demon-
strate important phylogenetic affinities between the Mystacocarida and
any other order.
Undoubtedly, the Mystacocarida are the most primitive living
Crustacea which have thus far been discovered. In addition to the
simple body organization, this is clearly indicated by the persistence
of larval characters in the head, including the unspecialized primitive
morphology of the large mouthparts, the labrum, and the rostral
shield. The lamellar structure of the abdominal appendages, on the
other hand, is obviously a reduced condition.
DEROCHEILOCARIDAE, new family
Characters as of the type genus, Derocheilocaris
DEROCHEILOCARIS, new genus
Body elongate and cylindrical; all segments clearly marked off
from each other; thorax and abdomen about the same diameter
throughout; head and postcephalosome distinctly narrower than rest
of body. Head covered with a cephalic shield which has a large
rostral plate bearing a prominent, notched rostrum. Anal segment
with a large papilla dorsal to the anus. A pair of lateral, dorsoventral
troughs with heavily chitinized edges on the posterior part of the
cephalic shield and on all other segments except the last.
First antennae 8-segmented, the terminal segment longest. Second
antennae tactile and natatory; both rami elongate. Both rami of
mandibles elongate, but endopod distinctly shorter than exopod;
gnathobase on basipod. Instead of projecting ventrally, the second
antennae and mandibles are flexed so that they extend dorsally and
laterally away from the median plane of the body. Labrum very
NO. 9 A NEW ORDER OF CRUSTACEA—PENNAK AND ZINN 5
large and flat; posterior end, between bases of maxillipeds, forming
a broad, smooth semicircle at its extremity. First maxillae uniramous,
elongate, and composed of seven segments. Second maxillae uniram-
ous, elongate, and 6-segmented ; segments not so broad as those of first
maxillae. Postcephalosome appendages (maxillipeds) irregular, flat,
and broad; not distinctly divisible into exopod and endopod. Each
thoracic appendage reduced to a small, simple, unsegmented lamella
bearing a few terminal setae. Each caudal ramus prominent and
curved dorsally to form a large terminal claw.
Genotype—Derocheilocaris typicus.
DEROCHEILOCARIS TYPICUS, new species
PLATES I AND 2
Head about one-third as long as the rest of the body and separated
from postcephalosome by a@ ventral constriction. Head and post-
cephalosome distinctly narrower than rest of body. Thoracic and
abdominal segments all about the same size. Rostral plate separated
from cephalic shield by a well-defined groove, and with three promi-
nent notches at its anterior edge. The median notch is deep and
ovoid. The lateroventral notches are also deep but are more nearly
circular and are toothed along the ventral margins; it is possible
that these may be remnants of chitinous troughs. Near the posterior
margin of the median notch are several eyespots. Their number and
exact position are variable but typically there are two pairs, one of
which is rather small and close to the median line at the posterior
end of the notch, while the other two ocelli are larger and more
anterior and lateral. The last segment has a prominent median
dorsal papilla which bears a single seta and a minute toothed papillule.
On each side of the dorsal papilla there is a broadly triangular, flat,
short, spinous process. The two caudal rami are slightly longer than
the last segment, broad at the base, and are curved dorsally in the
form of a large claw. Each ramus bears three large plumose setae,
one dorsal and two lateral. The two lateral setae originate at about
midlength of the ramus, one being slightly shorter than the ramus
and the other about twice as long as the ramus. The dorsal seta
originates near the base of the ramus and is slightly longer than the
ramus. There are two small papillae on the dorsal surface of each
ramus, the anterior one bearing a fine seta. Along the median ventral
edge of each ramus are three small equidistant processes each of
which bears a spinule.
The lateral chitinous troughs of the head are situated near the
posterior margin of the cephalic shield; each of these is roughly
6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
elongate but the dorsal and ventral extremities are irregular in out-
line; there are six to eight minute chitinous teeth along both the
anterior and posterior edges. The troughs of the postcephalosome
are irregular in shape but roughly cross-shaped with the vertical axis
longer than the horizontal axis; there are four groups of one to four
teeth along the inner margin of these two troughs. The troughs of
the thorax and abdomen are situated in approximately the middle
of the segments. They are all similar, being long and narrow with
10 to 14 triangular interlocking teeth on each side.
The location of the genital pore on the ventral side of the first
thoracic segment is indistinctly indicated by the presence of a broadly
obtuse chitinous ridge which projects anteriorly and ventrally near
the anterior margin of the segment.
The first antennae are long and 8-segmented; the first three seg-
ments are short and broad, but the others are elongate, the terminal
segment being the longest. The first segment bears one seta; the
second has three, two of which are minute; the third has seven, four
of which are minute ; the fourth through eighth segments bear 3, 4, 4, 5,
and 3 to 5 terminal setae, respectively. The first and second segments
each have a small dorsal hook, and the terminal segment bears a
single delicate aesthetask.
The second antennae are biramous with the 5-segmented endopod
being about half again as long as the 9-segmented exopod. The basal
endopod segment bears two short median setae while the third seg-
ment has a single seta. The terminal endopod segment is long and
fingerlike and bears a fringe of fine hairs at its tip. The exopod
segments are short; segments one, four, five, and six have a single
short seta, while the seventh and eighth have a long plumose seta;
the terminal segment is tipped with three long plumose setae. There
are five aesthetasks, one on the basipod, one on the first exopod
segment, one on the third, and two on the terminal segment. All
the aesthetasks project laterally. Because of their delicacy they are
often broken off in specimens which have been permanently mounted
on slides.
The mandibles are of a generalized larval type. The 2-segmented
endopod is about two-thirds as long as the 7-segmented exopod. The
basal endopod segment bears a single median seta, while the other
has two median and three terminal setae. Exopod segments three,
four, five, and six bear a single terminal seta, the last three of which
are plumose. The last exopod segment has three terminal plumose
setae, two of which are short while the third is slightly more than
twice the length of the entire exopod. The basipod bears a single
NO. 9 A NEW ORDER OF CRUSTACEA—PENNAK AND ZINN 7
distal median seta and a prominent, curved, stout, spinelike gnathobase
which has several small median spinules. All the segments of the
mandibles are beset with small rows of fine hairs.
The seven segments of the uniramous first maxillae are somewhat
broad and flat, the third segment being the largest. The basal segment
is narrowed at its base where it is attached to the trunk; the second
segment is short and is set off from the first and third segments by
poorly defined articulations. Except for one small lateral seta on the
sixth segment, all of the setae are stout and are arranged along the
median margins of the segments (median and terminal margins in
the case of the last segment). Beginning with the basal segment the
setation of the first maxillae is 3-1-4-2-2-2-8. All of the setae except
those of the terminal segment are borne on small projections.
The uniramous 6-segmented second maxillae are narrower and
slightly longer than the first maxillae. The first and second segments
are considerably larger than the other four. Like the first maxillae,
these appendages are densely setose along the median margins;
typically, the setation is 8-8-2-2-2-5. Three of the setae of the basal
segment are borne on a prominent process; two setae of the terminal
segment and one on the fifth segment are geniculate.
The structure of the postcephalosome appendages (maxillipeds)
is unique. They are small, irregular, but somewhat elongate, and
usually lie appressed to the trunk. It is likely that several exopod
and endopod segments have become fused with the basipod to form
the main body of the maxilliped. The remnant of the exopod con-
sists of three small segments, the second and third of which bear two
setae. The remaining endopod segments comprise a small bilobed
outgrowth which bears five setae. The large basal portion of this ap-
pendage has three small lateral and two small median processes all
of which bear one or two broad setae.
In studying the first antennae, first maxillae, second maxillae, and
maxillipeds of about 20 specimens, it was found that there are occa-
sional variations in the number and precise positions of the setae.
The four pairs of thoracic appendages are small, simple, unseg-
mented lamellae which lie rather closely appressed to the trunk. They
are all about the same size and differ only slightly in shape. The first,
second, and fourth appendages are roughly oblong, while the third
is tapered slightly, is more ovate, and has a small median notch at
about half length. The first abdominal appendages have two terminal
setae, while the others have three terminal setae.
From the tip of the rostrum to the posterior end of the caudal
rami the length of the adult ranged from 409 to 491 microns; the
average length was 458 microns.
8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Cotypes, U.S.N.M. No. 80450 from Nobska Beach, Mass., have
been deposited in the United States National Museum.
Larval stages—A few specimens of five different larval stages
were found. For convenience in making comparisons, these larvae
have been arbitrarily numbered I, 2, 3, 4, and 5 in order of their
increasing size, complexity, and maturity. All have two pairs of
antennae, mandibles, labrum, second maxillae, and caudal rami. Only
the two largest stages have first maxillae and maxillipeds, although
all five stages have a maxilliped segment (postcephalosome).
Except for the second antennae and mandibles, all of the larval
appendages are strikingly similar to the corresponding appendages
of the adult. Attached to the basipod of the second antenna, and more
median than the base of the endopod, is an unsegmented biramous
quasi palp in all larval stages. The undivided basal portion of this
quasi palp is very short while the two rami are about as long as the
-adjacent endopod (see pl. 2, fig. 8). The distal half of the outer
ramus and the distal third of the inner ramus consist of two setae
of similar length. In addition, the outer ramus has two small setae
near its base. Also, the endopod of the larval second antenna has
four segments while that of the adult has five.
The presence of the unique quasi palp on the larval second antennae
suggests interesting conjectures. Although the quasi palp is not a
very well-developed or robust structure, it is possible that it may
function, or at one time did function, as a clasping organ. The
possibility then arises as to whether the fifth larval stage may be
(or may have been in the past) a functional male phase as is the
case in certain other protandrous Crustacea, Mollusca, and Nematoda.
In none of the fifth larval stages, however, could any internal repro-
ductive structures be discerned.
The endopod of the larval mandible also differs markedly from
that of the adult. In the former it is composed of three broad seg-
ments, the terminal segment being much smaller than the other two.
In the adult this ramus is composed of two long narrow segments.
The smallest larvae found have a total of only 3 segments posterior
to the postcephalosome; the second smallest stage has 5 such seg-
ments; the next largest has 7; the other two have Io, the same as
the adults. No thoracic appendages are present on the first three
larval stages, but the fourth and fifth stages have 3 and 4 thoracic
appendages, respectively. In all immature stages the posterior part
of the cephalic shield, the postcephalosome, and all posterior seg-
ments except the last segment have chitinized troughs.
elie as
NO. Q A NEW ORDER OF CRUSTACEA—PENNAK AND ZINN 9
From the standpoint of morphological characteristics and size (see
table 1), it is logical to assume that larval stage 4, larval stage 5, and
the adult form a natural sequence. On the other hand, however, it
seems very probable that the three earlier stages are an incomplete
series and that there are still other early stages which were not dis-
covered during the present investigation. It is particularly likely
that there are very early larval stages which are comparable with
copepod nauplii in their simplicity. Because of incomplete informa-
tion concerning the various stages in the development of the Mysta-
cocarida, therefore, no attempt will be made at this time to assign any
descriptive, designative names to the five different types of larvae
that are known.
TABLE 1.—Summarised morphological characteristics of larval stages of
Derocheilocaris typicus
o o =
o Sun 3 = = = z n
s eee Soe % oe = Sd.) Sous
% ne Sa Soa - coi o oes ge GUM
= POEs os < o = 998O 3 OF mS
s Eos *a.e Ss bey o8 mS 2Sce 2485 St, 5
> Saws aang a +B * Botvm 8sAoa D2
a S358 S55 = 3 S gee 385s Fee
Hy Z a a As le ae Zip hexane
RPP ee oso: 8 present absent present absent 3 oO 187
Paes isle 3 3 present absent present absent 5 oO 203
Beery e so 3 present absent present absent o oO 250
Amrartion iets rs < 3 present present present present I0 3 350
eestor yess <) s 3 present present present present I0 4 390
(adult), .;.. 2 absent present present present 10 4 458
Internal anatomy.—Although only a superficial study was made
of the internal anatomy of Derocheilocaris typicus, it might be worth
while to indicate a few of the more obvious structural details. In
sexually mature individuals the most distinctive feature is a large,
elongate, dorsal, granular yolk gland; the posterior end of this yolk
gland extends as far as the fourth or fifth abdominal segment;
anteriorly, it may not extend beyond the first abdominal segment, or
it may extend as far forward as the second thoracic segment. The
ovary consists of several immature ova (customarily three to seven) ;
they are rather compactly arranged in the dorsal portion of the third
and fourth thoracic segments; often the ovary is partially imbedded
in the anterior portion of the yolk gland. A gonoduct could not be
definitely distinguished. The digestive tract is tubular and without any
pronounced constrictions or diverticula. Neither maxillary nor anten-
nary glands could be discerned. The nervous system, as shown by
longitudinal sections, is characterized by a series of extremely large
Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
segmentally arranged ventral ganglia. There is considerably less
fusion of the ganglia of the head than in other groups.
Occurrence—Of five beaches examined in the vicinity of Woods
Hole, Mass., Derocheilocaris typicus was found only in Nobska Beach
near the town of Woods Hole and in a beach 5 km. east of Woods
Hole near the town of Falmouth. About 50 mature and 12 immature
specimens were collected during June and July, 1939. The great
majority of these organisms occurred within a meter of the high-tide
line at a depth of 12 to 16 cm. in the damp sand. Although many
series of intertidal sand samples were collected at Baxter’s Beach,
Pine Orchard, Conn., between November 1937, and May 1942, this
form was found only during April 1940.
ACKNOWLEDGMENTS
The senior author carried on the field work associated with this
study during June and July, 1939, when he was a visiting investigator
at the Woods Hole Oceanographic Institution. He wishes to express
his gratitude to Prof. H. B. Bigelow, then director of the Institution,
for his kindness in providing laboratory facilities. Both authors thank
Prof. G. E. Hutchinson, of Yale University, for his many helpful
suggestions.
LITERATURE CITED
Monk, C. R.
1941. Marine harpacticoid copepods from California. Trans. Amer. Microsc.
Soc., vol. 60, pp. 75-99.
PENNAK, R. W.
1942a. Harpacticoid copepods from some intertidal beaches near Woods Hole,
Massachusetts. Trans. Amer. Microsc. Soc., vol. 61, pp. 274-285.
1942b. Ecology of some copepods inhabiting intertidal beaches near Woods
Hole, Massachusetts. Ecology, vol. 23, pp. 446-456.
Sars, G. O.
1901. An account of the Crustacea of Norway, vol. 4, pp. 1-4. Bergen
Museum.
1903-1911. An account of the Crustacea of Norway, vol. 5, Copepoda Har-
pacticoida. 449 pp. Bergen Museum.
ZINN; De):
1942. An ecological study of the interstitial microfauna of some marine
sandy beaches with special reference to the Copepoda. Dissertation,
Yale Univ.
li,
ee se ee
————————
NO. 9 A NEW ORDER OF CRUSTACEA—-PENNAK AND ZINN TE
EXPLANATION OF PLATES
PLATE I
Derocheilocaris typicus
I. Ventral view of head region of adult showing relationships between labrum
and basal portions of first antennae through maxillipeds.
2. Dorsal view of cephalic shield, showing notches, sutures, and ocelli.
3. Lateral view of adult.
4. Lateral view of abdominal segment showing detailed structure of a chitinous
trough.
5. Lateral view of posterior part of head and postcephalosome showing details
of chitinous troughs.
6. Lateral view of posterior end.
7. Ventral view of first thoracic segment showing region of genital opening.
(Large scale refers to figure 3; small scale refers to all other figures.)
PLATE 2
Derocheilocaris typicus
First antenna of adult.
Second antenna of adult.
Mandible of adult.
First maxilla of adult.
Second maxilla of adult.
Maxilliped of adult.
Right thoracic appendages of adult.
Second antenna of fifth larval stage.
Mandible of fifth larval stage.
OC PIAWKKRY DH
et
a
a
~
so
oOo
=
-
*
o
a
=
oO
>
SMITHSONIAN MISCELLANEOUS COLLECTIONS
|
ES
Bia
Cw WY Z
DOK ye
SN ASS
oo
DEROCHEILOCARIS TYPICUS
(For explanation, see page II.)
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103, NO. 9, PL. 2
Loos SS S
ses 3 IIIT
7,
aes aN
lime.
DEROCHEILOCARIS TYPICUS
(For explanation, see page II.)
——
|| SMITHSONIAN MISCELLANEOUS COLLECTIONS
de eres eS hh VOLUME 103, NUMBER 10
(End of Volume)
Roebling Fund —
| A REMARKABLE REVERSAL IN THE DISTRIBUTION
| OF STORM FREQUENCY IN THE UNITED STATES
|| IN DOUBLE HALE SOLAR CYCLES, OF
4 INTEREST IN LONG-RANGE
FORECASTING
BY
G, J. KULLMER |
(PUBLICATION 3729)
_ CITY OF WASHINGTON |
_ PUBLISHED BY THE SMITHSONIAN INSTITUTION
| APRIL 5, 1943
SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 103, NUMBER 10
(End of Volume)
Roebling Fund
A REMARKABLE REVERSAL IN THE DISTRIBUTION
OF STORM FREQUENCY IN THE UNITED STATES
IN DOUBLE HALE SOLAR CYCLESOF
INTEREST IN LONG-RANGE
FORECASTING
BY
CG. J. KULLMER
(PUBLICATION 3729)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
APRIL 5, 1943
‘The Lord Baltimore Drees
BALTIMORE, MD., U. 8. A.
Roebling Fund
A REMARKABLE REVERSAL IN THE DISTRIBUTION
OF STORM FREQUENCY IN THE UNITED STATES
EN DOUBLE HALE SOLAR’ CYCLES; OF INTEREST
IN LONG-RANGE FORECASTING
By C. J. KULLMER1
We have for the United States the only long fairly uniform series
of maps of tracks of barometric depressions. These have been pub-
lished in the Monthly Weather Review for each month from 1874
to the present. Dunwoody assembled the material for the 10-year
international period, 1878-87, in storm-frequency maps for the whole
Northern Hemisphere. Dunwoody’s method was to divide the maps
into 5° squares and record the number of centers of barometric de-
pressions that crossed each square. In 1911 I remade the maps of
storm frequency in the United States for 1899-1908 according to
Dunwoody’s plan of 5° squares.* In the interval of 21 years a slight
but definite southerly and westerly shift had taken place. But 5° in
latitude, about 345 miles, is evidently unnecessarily large if we wish
to test latitude shifts. I wished to ascertain whether there is any
correspondence between the latitude shifts of sunspots and the lati-
tude of the vortexes in our own atmosphere. Accordingly, I cut
Dunwoody’s square in half, making the unit 5° in longitude and 24°
in latitude, and made in 1913 a series of year maps from 1874 to
1912, furnishing comparison material for three solar cycles. Since
that time three more solar cycles have become available. The
results for the five cycles, with a series of year maps, 1883-1930,
appeared in 1933 in a Smithsonian publication.’ I shall now present
the results for the last solar cycle.
1 Published posthumously; Dr. Kullmer, formerly of Syracuse University,
died in 1942.
2 The shift of the storm track. Chap. 16 11 Huntington, The climatic factor,
Carnegie Inst. Publ. 192, 1914. :
3Kullmer, C. J., The latitude shift of the storm track in the I1-year solar
period, Smithsonian Misc. Coll., vol. 89, No. 2, 1933. Preliminary publication
of parts of the study appeared in Huntington, The solar hypothesis of cli-
matic changes, Bull. Geol. Soc. Amer., vol. 25, pp. 477-590, 1914. See also
Huntington, Earth and Sun, Yale Univ. Press, 1923.
SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 103, No. 10
to
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
For comparison, the most powerful method of attack seemed to be
to add together 3-year maps at solar minimum and 3-year maps at
solar maximum and compare them with each other. This was done
for five solar cycles, with the result that a uniform pattern appeared.
In figure 1 I present one such cycle, perhaps the most typical of the
five. Plus figures indicate the excess in years of solar maximum
over years of solar minimum; minus figures, the reverse. In the
north there was a strong increase along the main storm track, but
also a southerly projection. I have drawn a heavy dotted line through
the main area of excess at times of maximum and also through the
projection; at both sides of the projection there are areas of decrease,
i.e., more storms at years of minimum than at maximum. In figure 2
I have assembled the dotted lines through the area of excess and the
projection. It will be noted that in the fifth period there was an un-
expected move to the south and that the southerly projection had
shifted even farther west than in the second period. Because of a
certain orderly progression in the shifts of the southerly projection
I ventured a hypothetical position for the sixth solar cycle, and was
correct in assuming an easterly motion of the southerly projection.
1 now present in figure 3 the results for the last solar cycle, showing
an unexpected still greater shift to the south of the main area of
excess at solar maximum. There was some doubt as to the year
of maximum, 1937 or 1938, and consequently I made the maps for
both periods, figure 4. Except for slight variation in figures, the
maps are identical.
Now what does the continued southerly shift of the pattern signify,
with the large increases at minimum in western Canada? The in-
terpretation of a relation between sunspot latitudes and latitudes of
earthly storms seemed to be justified by the first four periods, but
contradicted by the fifth and sixth periods. There remained one
way to test the relationship. In figure 5, taken from Clayton’s
“World Weather,’ we have Maunder’s chart of sunspot latitudes.
All these years I have been comparing 3 years at solar minimum
with 3 years at solar maximum, but evidently the latitude effect
would be intensified if I compared 3 years at the beginning of the
cycle with 3 years at the end of the cycle. There is a noticeable
overlapping of the new cycle with the old; in order to avoid this
source of error I would eliminate by inspection the year of maximum
overlapping, as indicated in the chart, and proceed to test the latitude
effect by comparing 3 years at the beginning of a cycle with 3 years
at the end of a cycle. I began with the first cycle available, figure 6,
and the results were a succession of surprises. Gone was all thought
NO. 10 STORM FREQUENCY IN THE UNITED STATES—KULLMER 3
of a relation to sunspot latitudes. Gone entirely was the pattern
with the southerly projection, which I had found consistently for
six solar cycles. This pattern still remains an enigma, the result of
some solar relationship, the difference between years of maximum
solar activity and those of minimum activity. Instead I discovered
a solar relationship of possibly great significance.
The first period, figure 6, shows three horizontal bands. Plus
figures indicate more storms at the beginning of the cycle; minus
figures, more storms at the end of the cycle than at the beginning.
The following period, figure 7, shows a map of an entirely different
character—a plus band in the north, and curving down the Atlantic
coast and across the south a minus area, forming a script T pattern.
The succeeding period, figure 8, returns to the pattern of the first,
three horizontal bands, but in reversed order, with a pronounced
minus area between the plus bands. The succeeding map, figure 9,
was the greatest surprise of all—a return to the almost identical
script T pattern of the second period, but in reverse, with a minus
area in the north and curving down the Atlantic coast, and across the
south a plus area. And now with the-fifth period, figure 10, we re-
turn to the three horizontal bands, but again in reverse, with a plus
area between two still widened minus areas. These complicated re-
lationships will appear more simple in a table, figure 11. Because
of the appearance of three bands in the first and third period and
the strange script T pattern in the second and fourth periods, we
may divide the first four periods into two Hale solar cycles. But
whatever the solar influence may be that determines the location
of storms on earth, that influence was completely reversed in the two
Hale cycles. The evidence of a reversal is further supported by the
return in the fifth period to the conditions found in the first.
A crucial test will be offered by the coming sixth solar cycle, which
will have to show the script T pattern with a plus area in the north
and the long curved minus area in the south. Fortunately we are
able to look somewhat into the future. I have combined the maps for
1935-1937, and they show the highest known figures for the eastern
Canadian region. It might be considered that this shows merely the
increase in the network of northern Canadian recording stations.
But I present the evidence of the two squares in Quebec, north of
the Gaspé. Figure 12 shows the number of barometric depressions
that crossed these two squares from 1883 to 1940. This diagram is
another evidence that tends to confirm my confidence in the early
records of tracks of storms. The early years show approximately
the same high frequency as the later years; the year 1894 with 56
aa SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
barometric depressions almost equals the high of 1937 with 57
storms. We note that in 1895 there was a sudden drop from 56 to
17 and in a few years to 2. Furthermore, we notice that in 1940
there was a sudden drop from a high of 57 to 29 and that the interval
between the two sudden drops was 45 years, exactly four solar cycles.
I think there can be little question that the years of minimum, 1941-
43, will show a decrease in eastern Canada. With the plus area in
eastern Canada assured, we shall have to wait 3 or 4 years for the
possible completion of the lower part of the script T pattern.
How shall we interpret these findings? We naturally first think
of the alternations in solar activity as shown in the accompanying table,
figure 13, and of Clayton’s northerly shift of the centers of action
with increased solar activity, but the figures resist attempts at cor-
relation with the observed reversal. It seems, therefore, necessary
to consider the theory that solar activity may possibly be completed
in four cycles, of which the third and fourth are in some mysterious
way the exact reversal of the first and second. We have become ac-
customed to the idea of a reversal of solar activity through Hale’s
discovery of a reversal of magnetic polarity with each new solar
cycle. But in what way could such a Hale cycle itself be reversed ?
Finally a word concerning the years selected for comparison. It
will be remembered that for the first three and a half periods I
selected by inspection from Maunder’s chart of sunspot latitudes
the years of maximum overlapping and used 3 years on either side
of such a year. Maunder’s chart does not include the minima of
1923 and 1933; consequently I selected these years as boundaries.
Figure 14 summarizes the official years of minimum and maximum
solar activity, together with my periods selected as described. I
would call attention to the fact that the first and third periods begin
2 years after the official minimum; the second period, 1 year after
minimum, and the fourth period begins with the minimum. There
are very slight differences in the Wolfer sunspot numbers which
determine the solar minimum, and it seemed worth while to test the
matter by making a series of maps using the official minimum year
as the boundary; for the fifth test period, which was originally so
bounded, I included the minimum year and have already presented it
as the superior map. In figure 15 I present the first period. The
central plus band is strengthened, but the southern minus band is
weakened. My original period seems to be the best for the three-
banded effect. In the second period, figure 16, the plus band in the
north is weakened, uncovering, as it were, a narrow band of minus
which crosses the map. Also the southern curved minus area is
NO. 10 STORM FREQUENCY IN THE UNITED STATES—KULLMER 5
weakened. Again my original period is the best. In the third period,
figure 17, the northern plus band is weakened; otherwise, there is
little choice. The fourth period, figure 18, shows the greatest dis-
crepancy, with a great weakening of the northern minus band and
the introduction with the year 1916 of a new strong plus band. There
is also a weakening of the southern plus curve. It is evident that for
the solar influence presented in this paper my original period is far
the superior. The last period, figure 19, shows a widening and
strengthening of the northern minus band and I have consequently
chosen it as the best representative of the three-banded effect. Why,
with this one exception, my original periods best bring out the solar
effect here presented I cannot venture an opinion; it is, however,
of interest, that with the exception of the fourth period, a shift of
I year, and in one case, of 2 years does not vitiate the results.
These, then, are my findings, and I offer them to those more com-
petent than I am for interpretation.
I append the year maps of storm frequency for 1931-1940, form-
ing a series of 58 year maps, made with care according to a uniform
technique, to which can be added the years 1874-1882, now in manu-
script.
VOL. 103
SMITHSONIAN MISCELLANEOUS COLLECTIONS
6
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8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
NO. 10 STORM FREQUENCY IN THE UNITED STATES—KULLMER 9
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NO. 10 STORM FREQUENCY IN THE UNITED STATES—KULLMER Tee
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1880-82 1890-92 1903-05 1913-15 1924-26 1935-37
vs. VS. VS. Vs. vs. VS.
1886-88 1899-01 IQO0Q-II 1920-22 1931-33 1941-43?
Two bands Two bands
Three bands ina Three bands ina Three bands ?
“f pattern “f pattern
minus plus plus minus minus plus
plus — minus -- plus ?
minus minus plus plus minus
Engst
SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
Sip ee a Ne.
Fig. 12.
TOGO Mais City ete ace oe Max. 139.1
TOSS ki hia eee ree Max. 63.7
MOOS, ened eu eos nee: Max. 84.9
LOO 5S eharsiars orev cay ne Max 63.5
TOL awe leek ee Max. 103.9
LOZSar ee cs ernie MeN SEN: Max 77.8
LUG er cceaces ee Aan te Max. 137.0
Rireeane)
Three Bands
‘f Pattern
Three Bands
‘{ Pattern
Three Bands
?f Pattern
NO. 10 STORM FREQUENCY IN THE UNITED STATES—KULLMER IS
Years selected Min. Max. Test years
1880-82 vs. 1886-88 1878 1879-81 vs. 1886-88
1883
1890-92 vs. 1899-01 1889 1803 1890-92 vs. 1898-00
1903-05 vs. 1909-11 IQOI 1905 1902-04 vs. IQIO-12
IQ13-I5 vs. 1920-22 1913 1017 1914-16 vs. 1920-22
1924-26 vs. 1930-32 1923 1928 1924-26 VS. 1931-33
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T4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
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160 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 103
1931 and 1932
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KULLMER
NO. 10 STORM FREQUENCY IN THE UNITED STATES
1933 and 1934
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1935 and 1936
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1939 and 1940
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