Y^f-t?
HARVARD UNIVERSITY
Library of the
Museum of
Comparative Zoology
m'h.^
BuLLQtln OF THE
Museum of
6 m para five
ogy
Volume 137
1968-1969
'*
HARVARD UNIVERSITY
CAMBRIDGE, MASSACHUSETTS 02138 U.S.A.
CONTENTS
No. 1. The Carabid Beetles of New Guinea. Part III. Harpalinae (Con-
tinued): Perigonini to Pseudomorphini. By Philip J. Darlington.
July, 1968 1
No. 2. Geographic Variation in Anolis distichus Cope ( Lacertilia, Iguan-
idae ) in the Bahama Islands and Hispaniola. By Albert Schwartz.
September, 1968 255
No. 3. Ammonoids of the Late Scythian (Lower Triassic). By Bernhard
Kummel. April, 1969 311
/^K«J
Museum of
Comparative
Zoology
The Carabid Beetles of New Guinea
Part III. Harpalinae (Continued):
Perigonini to Pseudomorphini
p. J. DARLINGTON, JR.
HARVARD UNIVERSITY VOLUME 137, NUMBER 1
CAMBRIDGE, MASSACHUSETTS, U.S.A. JULY 30, 1968
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THE CARABID BEETLES OF NEW GUINEA PART III. HARPALINAE
(CONTINUED): PERIGONINI TO PSEUDOMORPHINI
p. J. DARLINGTON, JR.^
•■ CONTENTS
Introduction to Part III.
Purpose; other parts; acknowledgments _.„ 2
Sources and disposition of material 2
Policies and methods; type examinations;
measurements; drawings 3
Localities 4
Findings 5
Taxonomic section
Tribe Perigonini 5
Perigona 6
Tribe Licinini 14
Badister 15
Physolaesthus 15
Omestcs 16
Dicrochile 16
Microferonia 18
(Tribe Amblystomini) 19
( Aniblystoinits) 19
Tribe Chlaeniini 20
Chlaeniits 1 20
Tribe Oodini 30
Anairichis 31
Oodes 32
Tribe Harpalini 38
Gnathaphanus 4 1
Diuphoromerus 42
Hijphurpax 1 44
Lecanomerus 45
Chydaeiis 47
Platyrnciopus 48
TricJiotichnus 48
Hcirpaloxenus _*._: 59
Lyter ; 63
Coleolissus 64
Hyphacreon 66
Anoplogenius 68
Egadroma 69
Stenolophus 71
Acupalpiis 72
Tribe Anaulacini 76
^ Work supported by National Science Founda-
tion Grant GB-93; see also p. 2.
* Bull. Mus. C
Odoutomasoreus 76
Amuilactis 77
AeplinkUus 77
Cuphoia 78
Tribe Cyclosomini 78
Sarotlirocrepis 78
Tribe Lebiini 80
{ Soniotrichus) 82
Aristolehia 83
Lehia 85
Lach noderma 89
Sinurits QQ
Stenotelus 90
Misceliis 91
Holcoderti.s 94
Minuthodcs 95
Catuscopus 100
Pericalus no
Coptfldera 1 10
Minuphloetis 117
Agonochilu 1I8
Oxyodotitus 122
Moc}ithcni.s 122
( Mochtheroides) 123
DolicJtoctL^ 124
Stricklandia 132
Peliocypas 134
Celacnephes 135
Syntom us 135
Microlestes 135
Apristus 137
( Plochionus) 138
Parena 133
Anch i.sta 139
Eiidynomena 140
Denietrida 140
Phlococarahus 183
Trigunotlwps 184
Nototarus ....'. 185
Anomotarus 186
Tribe Pentagonicini 191
Pentagonica 192
Parascopodes 195
Scopodes 197
Tribe Hexagoniini 202
omp. Zool., 137(1): 1-253, July, 1968 1
Bulletin Museum of Comparative Zoology, Vol. 137, Xo. 1
CONTENTS
Hexagonia 202
Tribe Odacanthini 203
Collium 204
Casnoidca - 207
Bcisi.stich us 208
Clurcncia 209
Dicraspeda 210
Lachnotlionix 214
Etidalia 214
Dobodura 215
Tribe Dryptini 216
Drtjpta - .' 216
Desera 218
Tribe Zuphiini 218
Zuphitim 219
Planctes 220
Tribe Helluodini 222
Pofiona^Iossus- 222
Tribe Ilclluonini 228
Creagris 229
IlcUuoiiidius 229
Ih'lbiopapua 232
HelUiosoma 233
HcUuodcma _. „ 233
Gigadcina 233
Tribe Brathinini 234
Phcro})so])}\its 234
Bracliinus 239
Tribe Pseudomorphini 239
Adelatopus 240
Cn/pt(>cc))halomorpha 242
S))haU(>nH)rpli(i 242
INTRODUCTION TO PART III
Purpose: other parts: acknoidcdiimcnts.
Tliis is the third part of a taxonomic survey
of the beetles of the family Carabidae
(predaeeoiis ground beetles) of the island
of New Guinea.- The present part covers
the tribes of the subfamily Ilarpalinae from
Perigonini through Pseudomorphini. and
thus completes coverage of the famiK
Carabidae in the approximate order of the
Junk-Schenkling Catalogue (Csiki 1932-
f933). I'art IV, which is now being pre-
pared, will be primarily a summary, analy-
- Part I, covering the Cieiiidelinae, Carabinae,
and Flarpalinac llnontili Ptiiostichini (in tlic ordc-r
of the Junk-Schenkhn!4 Catalogue) and Part II,
covering the Agonini, are in the Bulletin nt the
Museum of Comparative Zoology: Part I, in \'ol.
126, No. .3, 1962, pp. 321-564, 4 plates; and Part
II, in Vol. 107, No. 3, 1952, pp. 87-252, 4 plates.
(Because of my special interest in the Agonini,
Part II was written and published lalore Part I.)
sis, and discussion of the New Guinean
carabid fauna as a whole. Among subjects
to be considered are the general nature of
the fauna, its geographic relationships and
origins, its ccologic composition, and its
evolution including specific evolutionary
trends (toward wing atrophy, etc.) and
evolutionary radiations on New Guinea.
However, Part IV will include also a taxo-
nomic supplement to list important new
records of previously recorded species and
to describe a number of additional species
received recently, especially new Agonini
from high altitudes.
I have already acknowledged, in Parts I
and II, aid received from the Guggenheim
Foundation. I have now to acknowledge
also aid received from the National Science
Foundation ( Grant GB-93 ) . which has sup-
ported my work on Carabidae of Ne^^
Guinea in many ways, including publica-
tion of the results.
For meticulous editing and typing of the
manuscript of Part III, I am indebted to
Mrs. Judith Koivumaki, and for the accurate
outline drawings and realistic w atercolors,
to Mrs. Mary Catron.
Sources (iiul disposition of material. Prin-
cipal initial sources of material used in my
\\'ork on New Guinean C^arabidae ha\e been
acknowledged in Part I, page 323, and Part
II, pages 90-91. However, notable addi-
tional material has been received recently.
Most important are thousands of specimens
collected for the Bishop Muscnmi by several
entomologists und(>r the direction of Dr.
J. L. (>ressitt; Mr. Josei Sedlacck and his
wife and son have obtained an especially
large number of Carabidae for the Bishop
Museum. An important collection has hcvu
submitted for stud\ also b\- the Department
ol Agriculture, I'ort Mor(\sb\-. through the
kindness of Mr. J. j. II. S/cnl-I\an\'; this
collection includc\s much matcMial from the
Port Moresby area, and also specimens
from other localities including some Irom
high altitudes. Sent with this collection,
but belonging to him personalK', is a fine-
lot of Carabidae collected bv Dr. U. W.
The Carabid Beetles of New Guinea • Darlington
Hornabrook; this too includes material from
high altitudes. A collection submitted by
the Australian Commonwealth Scientific
and Industrial Research Organization, at
Canlierra, includes specimens from the
Morehead River, on the south coast of
Papua almost opposite the tip of Cape
York; several Australian species not known
elsewhere in New Guinea were found at
this locality. And an interesting collection
has been submitted by the South Australian
Museum, including much material from
Mt. Lamington, Papua.
Because different collections have been
received at different times, and because
different portions of my manuscript have
been finished at different times, I have not
set a single deadline for material included
in the present part of my work. I have
simply used in each case the specimens
available when a given group was studied,
with only a few especially important addi-
tional records interpolated later. Additional
noteworthy records will be included in the
supplement in Part IV, referred to above.
Several of the most productive New
Guinean carabid collectors, whose names
appear many times in the following pages,
are associated with single museums. In
order to save space, I shall cite these col-
lectors without repeating the names of their
museums. The persons in question, and the
museums with which they are associated
and to which their specimens belong, are:
(Ludwig) Biro: Hungarian National
Museum, Budapest
(Miss L. Evelyn) Cheesman: British
Museum
(P. J.) Darlington .(Jr. ): Museum of
Comparative Zoology, Cambridge, Mas-
sachusetts, abbreviated M.C.Z.
(J. L. ) Gressitt: Bishop Museum, Plono-
lulu
Sedlacek(s): Bishop Museum, Honolulu
( Citation of this name in the singular
indicates Mr. Josef Sedlacek; in the
plural, additional or different members
of the Sedlacek family: Marie and/or
J. H. Sedlacek)
(L. J.) Toxopeus: Leiden Museum
Other museums and collections of which
the names are abbreviated are:
American Museum of Natural Ilistorv
(New York): A.M.N.H.
California Academy of Sciences, San
Francisco: Cal. Acad.
Commonwealth Scientific and Industrial
Research Organization, Canberra, Aus-
tralia: C.S.I.R.O.
United States National Museum, Wash-
ington, D. C: U.S.N.M.
Policies and methods; type examinations;
measurements; drawings. My work is
second-stage taxonomy ( see Part I, pp. 328-
330). My methods have been described in
Part I, page 330, and Part II, pages 91ff.
However, I should repeat and stress certain
things. I have tried to be reasonably con-
sistent in preparing descriptions but have
not followed a single model exactly. I
have treated some tribes and some genera
in much more detail than others, the rule
being to give the information that has
seemed worth giving in each case. My
descriptions do follow a basic form but are
flexible in detail. I do not like check-list
taxonomy, in which descriptions are (in
effect) drawn by inserting adjectives in
blank spaces in a standard form. This kind
of taxonomy is easy, but it is likely to be
poor taxonomy. I think it is better to
describe each species individually, following
of course some sort of basic pattern, and if I
state under one species that a character is
striking, I see no reason to state (say)
twenty times under other species that it is
not striking.
Although the present part is consistent
with Parts I and II in general, I have made
a few slight changes of usage to conform
to two publications that have appeared
recently. One is the "Style Manual for
Biological Journals," pulilished in 1960 bv
the American Institute of Biological Sci-
ences, 2000 P St., NW, Washington, D. C,
20016. The other is the revised edition
( 1964 ) of the International Code of Zoologi-
cal Nomenclature. I have in general adopted
BuUetin Museum of Comparative Zoology, Vol 137, No. 1
the details of style suggested by the former,
and ha\'e tried to follow the rules and most
recommendations of the latter. However,
although I have followed the Style Manual
in most ways including most abbreviations,
I have occasionally preferred to follow
Webster's Collegiate Dictionary on points
of general style where I see no reason why
biologists should be different.
References listed under tribes, genera,
and species are limited to items directly
concerned with New Guinea plus selected
items likely to be specially useful to workers
on New Guinean Carabidae.
Type examinations: In the present part of
my work I have indicated what types of
previously described species have been
seen and not seen. I have borrowed for
study a few types in especially difficult
genera in which my work has been in
effect revisionary ( in Perigona, for ex-
ample), but I have not attempted to see or
to borrow types in most cases. There are
two reasons for this. First, I do not think
types should be loaned merely to confirm
identifications in faunal work, especially
when the types come from outside the area
under study, in the present case often from
other islands or from Australia rather than
Irom New Guinea. And second, H. E.
Andrewes saw many of the types in question
and made comparisons with them (see my
Part I, p. 325), and my study of the An-
drewes Collection has enabled me to place
not only his own but also most of the older
Oriental species with reasonable confidence.
I do, however, plan to see many of the
older types, including those in Paris, before
completing Part IV, so that I should be
able to correct errors of identification then.
Measurements. Statements of j^roportions
have been calculated (with a slide rule)
Irom actual measurements made with a
ruled disc in the ocular of a stereoscopic
microscope. Proportions eannol be esti-
mated satisfactorily by eye. When possible,
the proportions are based on measurements
of an average-looking 6 9 . The specimens
thus measured are usually specified in a
paragraph headed Measured specimens, but
this paragraph is omitted under species of
which only one or two indi\iduals are
known. Measurements of length and width
are extremes of all available specimens.
Drawings. My drawings are designed
primarily to show gross form, which is very
difficult to describe in words. Mouthparts,
antennae, and legs are sketched in semi-
diagrammatically. The drawings have been
outlined by Mrs. Mary Catron (usually
with use of a crosslined disc in the ocular
of a stereoscopic microscope), checked by
me ( the checking including measuring and
calculating of proportions ) , and then inked
by Mrs. Catron. I have not tried to figure
all species or even all new ones, but have
tried to show unusual ones and also new
species that are based on only one or two
specimens. I expect to deposit representa-
tive sets of specimens in museums in Lon-
don, Honolulu, Canberra, and elsewhere, as
well as in the continental United States,
and persons working on New Guinean
Carabidae in the future should use m\-
specimens rather than figures of them \\'hich
( like all figures ) are sure to be inadequate.
I have usually not used and therefore not
illustrated genitallic characters. I expect to
discuss this matter — when and how to use
genitallic characters in carabid taxonomy —
in Part IV.
Localities. I plan to include in Part IV
a map showing, as far as possible, all
localities at which C>arabidae ha\e been
obtained in New Guinea. In the meantime
the preliminary map published in Part II,
page 93, shows my own localities, most ol
Miss Cheesman's, and some others, and the
sketch map in Part I, page 326 shows the
route of my collecting on the Bismarck
Range. Also, the Rishoi) Museum has
issued a 19-page "List of New (hiinea
Localities" (to 1966) which gi\es approxi-
mate latitud(\s, longitudes, and altitudes ol
the localities ol Hishop Museum collectors
and ol some other persons. This list is, I
suppose, a\ailal)le on recjuest. I have used
it as a standard h)r spt-Uing ol place names.
The Carabid Beetles of New Guinea • Darlington 5
Certain localities have become especially
important in the course of my work.
Dobodura, Papua, where I collected from
March to July 1944 (see Part I, pp. 325-
326), is by far the best known lowland
locality in New Guinea, for Carabidae.
Wau, in the Morobe District, N-E. N. G., is
by far the best known middle-altitude
locality, thanks principally to the efforts of
the Sedlaceks. And Mt. Wilhelm on the
Bismarck Range, N-E. N. G. (where I
collected), and the Snow Mts., West N. G.
(where Toxopeus collected during the
Netherlands Indian-American (Third Arch-
bold) Expedition of 1938-1939), are the
only venj-higJi-altitude localities well known
for Carabidae. Comprehensive collections
from other localities, especially at high
altitudes, are much needed to show at
what intervals localized species replace
each other on New Guinea. Until this is
known, the total number of species of
Carabidae on the island cannot even be
guessed at closely.
Additional evidence that the label
"Dor(e)y" has been wrongly placed on
many of Wallace's Carabidae that prob-
ably really came from Celebes or the Moluc-
cas is given in the present part of my work:
see, for example, under AmbJystonius (p.
20). For general discussion of this locality
see Part I, pages 330-331. Although many
specimens so labeled evidently did not
come from Dorey, Wallace did go there.
Some of his field notes from there are
quoted under Catascopus in the present
part of my work (p. 102).
Findings. Although analysis and discus-
sion of the New Guinean carabid favma as
a whole will be postponed to Part IV, a
few special points are worth noting now.
Several genera that are chiefly Australian
have been found at high altitudes on New
Guinea, Java, and sometimes other islands
in the Malay Archipelago. These genera
include Mecyclothorax (Part I, pp. 498,
505); Microferonia (present part, p. 18);
and Scopodes (present part, p. 197). One
genus, Chydaeus (p. 47), has been fomid
with an opposite pattern of occurrence, on
the mainland of Asia and at high altitudes
on mountains in the Malay Archipelago
east to New Guinea. However, Bemhidion
and Trechiis have not been found on moun-
tains in New Guinea, although Asiatic stocks
of these genera have reached high moun-
tains farther west in the Malay Archipelago
(Darlington, 1959, Pacific Insects, Vol. 1,
pp. 331-345).
Important evolutionary patterns, of not-
able radiations of Carabidae on New
Guinea, have been found in the Agonini
(Part I) and are described and discussed
for several genera treated in the following
pages. The most striking, in fact exciting,
case is in the lebiine genus Demetrida,
which seems to be in the midst of an
evolutionary explosion. The situation among
these diversely colored carabid beetles in
the mountain rain forests of New Guinea
parallels in some ways the situation among
the birds of paradise in the same forests.
I have seen about 1250 specimens of
Demetrida from New Guinea, representing
apparently 56 species, all new! See discus-
sion under the genus (pp. 142-143) for
further details. Less striking, but neverthe-
less important, radiations of species chiefly
within the confines of New Guinea are
described in Trichotichnus (pp. 48-59),
Catascopus (pp. 101-110), Dolichoctis (pp.
124-132), Anomotarus (pp. 186-191),
Scopodes (pp. 197-202), Dicraspeda (pp.
210-214), and IleUuonidius (pp. 229-232).
TAXONOMIC SECTION
Tribe PERIGONINI
Plattjnini group Perigonae G. H. Horn 1881, Trans.
American Ent. Soc. 9, p. 143.
Perigonini Csiki 1931, Coleop. Cat., Carabidae,
Harpalinae 5, p. 894 ( see for synonymy and
additional references ) .
Jedlicka 1964, Reichenbachia 2, No. 61, pp. 267-
274 ( Oriental forms ) .
Perigonitae Jeannel 1941, Rev. frangaise d'Ent. 8,
p. 137.
Perigonidae Jeannel 1948, Coleop. Carabiques de
la Region Malgache, Part 2, p. 733.
The taxonomic limits of this tribe and its
6 Bulletin Museum of Comparaiive Zoology, Vol. 137, No. 1
relation to other tribes of Carabidae are
doubtful but need not be discussed here.
The only genus of the tribe in New Guinea
is Perigona itself (sensu lato).
Genus PERIGONA Castelnau
Castelnau 1835, Etude Ent., p. 151.
Sloane 1903, Proc. Linn. Soc. New South Wales
28, p. 635.
Andievves 1929, Tijdschrift voor Ent. 72, p. 326
( Suniatran species ) .
Csiki 1931, Coleop. Cat., Carabidae, Haipalinae
5, p. 895 (see for synonymy and additional
references ) .
Jedlicka 1935, Neue Carabiden aus Ostasien, Part
10, pp. 17-19 (Philippine species).
1964, Reichenbachia 2, No. 61, pp. 267-
274 ( Oriental species ) .
Jeannel, see references under tribe, above.
Euriipcri<iona Jeannel 1941, Rev. francaise d'Ent.
8, pp. 138, 149 (new synonymy).
Subgenus Trechicus Leconte 1853, Trans. Ameri-
can Philosophical Soc. 10, p. 386.
Dkigno.sis. Small Tachijs- or Trcchiis-\ike
Carabidae; with usually 2 setae over each
eye; apical segments of palpi rather long,
usually subconical; other technical char-
acters given by Jeannel.
Description. None needed here, except
note that all known New Guinean species
are fully winged.
Tiij)e species. Of Ferigona, P. pallida
Castelnau of Africa; of Eun/perifi,ona, P.
j)rocera Fauvel of Java; of Trechicus, T.
umhripennis Leconte (= Perigona nigriceps,
below ) .
Generic distribution. World-wide in
tropical and warm temperate regions. S(>e
also 4th paragraph of following Notes.
Notes. Ennjperigona Jeannel is based on
Perigona procera Fauvel, a very large
species with maxillary palpi long, slender,
with penultimate segments relatively long.
Perigona rex (below) would go in Euryperi-
goruu if this genus were recognized. How-
ever, "Euri/])crigonu' nitida Jeannel 1941
{= Perigona grandis Jedlicka 1935) of die
Philippines has maxillary palpi relatively
shorter and with pcMiultimate segments
shorter than in procera, and tends to con-
nect the latter with more typical Perigona,
and I do not think generic separation is
advisable.
Jeannel divides Perigona into 2 subgenera
which seem natural and useful. They differ
in arrangement of submarginal elytral
punctures and they differ also in habits:
Perigona sensti stricto occurs (in my ex-
perience) only or mostly on or in logs or
rotting wood; subgenus Trechicus, among
dead leaves or debris on the ground, usuallv
in forest. Perigona (Trechicus) nigriceps
( Dejean ) has extended its ecological range
to include fermenting vegetation and vari-
ous plant materials carried by man, and
has been spread over all the warmer parts
of the world.
Variation of supraocular and lateral
prothoracic setae in this genus is note-
worthy. In rev (below) all these setae are
absent. In P. lata Andrewes of Sumatra
the anterior supraocular and median-lateral
prothoracic setae are absent in both type
and "cotype" in the British Museum. And
in P. astrolahica Csiki the posterior-lateral
prothoracic setae are present or absent, as
described inider this species below.
Species of Perigona are numerous in
tropical Asia and the Malay Archipelago.
Fourteen occur in New Guinea. However,
only 5 (nigriceps and 4 endemic species)
occur in Australia (Darlington, 1964, Psxche
71, pp. 125-129). The New Guinean
Perigona fauna is therefore Oriental in
general naturt> and diversity, and it is Ori-
ental also in relationships of most species,
so far as relationships can be determined.
Farlier ke\s to specic\s of Perigona of the
Malay Archipelago (Andrewes 1929; Jed-
licka 1935, 1964) and Australia (Sloane
1903) have been based principalh' on size
and color, but most ol the 14 New Guinean
species haxc diagnostic structural char-
acters, as the follow ing Key shows. I am
indebted to Dr. Z. Kaszab h)r an opportu-
uit\ to examine the t> pes ot Csikis New
( iuiiicaii species.
""JMic I ol low lug species leeorded Ironi New
Guinea are still unknown to ine, and are
not included in the Key.
The Carabid Beetles of New Guinea
Darlington
SPECIES OF PERIGONA PREVIOUSLY
RECORDED FROM NEW GUINEA BUT
NOT RECOGNIZED FROM DESCRIPTION
Perigona litura (Perroud & Montrousier)
Perroud & Montrousier 1864, Ann. Soc. Linneenne
Lyon 11, p. 72 (Trcchiis).
And'rewes 1929, Tijdschrift voor Ent. 72, p. 372
(in key).
1933, Tijdschrift voor Ent. 76, p. 3C3.
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae
5, p. 897 (see for additional references).
This species was described from New
Caledonia. It is listed by Csiki from sev-
eral islands in the Malay Archipelago, in-
cluding New Guinea, but I cannot find the
source of the New Guinean record. An-
drewes did not know the species. Details
given in the original description, and the
fact that the type(s) occurred in detritus,
suggest that it may be a color form of
nig,nceps.
Perigona subcordata Putzeys
Putzeys 1875, Ann. Mus. Civ. Genoa 7, p. 730.
This species was described from the
Kei Islands and is likely to occur in New
Guinea. The size and other details suggest
that it may be an earlier name for astrolabica
Csiki.
Perigona suturaiis Putzeys
Putzeys 1875, Ann. Mus. Civ. Genoa 7, p. 728.
The type was collected at Sorong, West
New Guinea, by Beccari and D'Albertis,
and is now in the Genoa Museum. Putzeys'
description does not permit an exact deter-
mination but suggests a small specimen of
astrolabica Csiki or a large one of sub-
cyanescens Putzeys.
Key to Species of Perigona of New Guinea
1. Group of 3 punctures in outer submarsinal
channel of elytron (at % or % of elytral
length) forming a straight line {Perigona
scnsii stricto) 2
- These 3 punctures forming a triangle ( sub-
genus Trechiciis) 9
2. Supraocular and lateral prothoracic setae
absent; very large (9.4-12.4 mm) (p.
8 ) rex
- Two pairs supraocular and usually 2 pairs
lateral prothoracic setae present; size
smaller 3
3. Frontal foveae weak, subobsolete; elytra
each with 2 dorsal punctures, no subapical
puncture above marginal channel; length
c. 2.0-2.5 mm (p. 8) pygmaea
- Frontal foveae short but distinct, margined
externally by weak elevations; elytra with
3 punctures, the 3rd either posteriorly on
disc or subapically above marginal channel;
usually larger 4
4. Elytra with 3rd (posterior) dorsal puncture
on disc, separated from marginal channel
by more than width of latter; if in doubt,
refer here specimens over 4 mm long „ 5
- Elytra with 3rd puncture farther back, just
above edge of marginal channel 7
5. Posterior dorsal elytral punctures less than
Yw of elytral length from apex; length c.
4-6 mm (p. 9) astrolabica
- Posterior dorsal elytral punctures more than
Yxo of elytral length from apex; usually
smaller 6
6. Form normal, moderately broad and de-
pressed; length c. 3.3-4.0 mm (p. 9) _...
sxihcijanescens
- Form narrower, subcylindrical; length 2.6-
3.7 mm (p. 10) papimna
7. Larger, c. 4.5 mm; dark castaneous with
reddish suture and appendages (p. 10 ) - rossi
- Smaller; // approaching rossi in size, form
more depressed and color testaceous ._._ - 8
8. Depressed; nearly uniform testaceous with
head browner but elytra not plagiate; length
slightly over 3 mm (p. 10) livens
- Less depressed; partly testaceous but with
head and much of elytral discs darker;
length under 3 mm (p. 11) plagiata
9. Submarginal channel of elytra behind
puncture-triangle (at % or % of elytral
length ) wide, with bottom flat or convex .. 10
- Submarginal channel behind puncture-tri-
angle very narrow 12
10. Color cither testaceous with dark head and
elytral apices or brownish castaneous with
head slightly darker and suture paler; eyes
large, forming c. right angles with neck;
front and neck with distinct c. isodiametric
reticulation (p. 11) nigriceps
- Color dark castaneous with suture not or
not much paler; eyes variable; microreticu-
lation of head often less distinct, often ( not
always) transverse posteriorly 11
11. Eyes larger; microsculpture of posterior
part of head ( if visible ) not obviously
transverse; length usually c. 3.2—3.6 mm
(rarely smaller) (p. 12) erimae
- Eyes smaller and less prominent; micro-
sculpture of posterior part of head (if
8 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
visible) more trans\'erse; length c. 2.7-
3.3 mm (rarely larger) (p. 12) --_ hidovici
12. Prothorax with sides not strongly sinuate
and posterior angles not denticulate; length
c. 2.8-3.3 nnn (p. 13) lebioides
- Sides of prothorax strongly sinuate or
posterior angles denticulate 13
13. Sides of prothorax strongly sinuate about
% of length before base; (fine) micro-
sculpture present; length c. 3.4 mm (p.
13 ) cordens
- Sides of prothorax nearly straight and con-
verging posteriorly, but posterior angles
aliruptly promini'ntly denticulate; micro-
sculpture absent or nearly so; length c.
3.2-3.4 mm (p. 14) dcntifcr
Perigona is.s.) rex n. sp.
Description. With characters of genus;
form as in Figure 1; very large, broad,
depressed; brownish castaneous, lower sur-
face and legs more reddish; rather shining,
reticulate microsculpture fine, lightly im-
pressed, c. isodiametric on head, slightly
transverse on pronotum and elytra. Head
0.58 and 0.60 width prothorax; mandibles
shorter and more curved than usual in
genus; eyes rather small but prominent,
enclosed behind by genae; antennae with
middle segments c. 1V-' X long as wide;
maxillary palpi slightly shorter than in P.
proccra Fauvel, with apical segments slightly
more conical, and with subapical segments
c. equal length of apical ones; frontal im-
pressions vague; supraocular setae absent;
mentum with a long, triangular tooth.
Prothorax: width length 1.64 and 1.56;
base/apex c. 1.33 and 1.22 (exact measure-
ments imjiossible because basal angles
broadly rounded); lateral setae absent; disc
with fine middle line, other imjiressions
vague. Elytra: width elytra/prothorax 1.20
and 1.24; striae absent or faintly indicated;
each elytron with 2 to 4 dorsal punctures
(variation individual, sometimes xms\in-
metric), anterior puncture larth(>r than
others from suture. Secondary .sexual char-
acters: i front tarsi scarcely dilated but
usually with inconspicuous 2-seriate squa-
mae on first 3 segments below (only near
apex of 1st segment, and soiiietimes missing.
perhaps broken off); S with posterior
femora dentate on upper posterior side near
apex; $ with usually 3, 9 4 or 5 seta-
bearing punctures each side last ventral
segment. Measurements: length 9.4-12.4;
width 3.(S-5.0 mm.
Types. Holotype i (Bishop Mus.) and
1 9 paratype (M.C.Z., Type No. 31,344)
from Sepalakembang, Salawaket Rge., N-E.
N. G., 1920 m, holotype Sept. 11-14 and
paratype Sept. 15, 1956 (E. J. Ford, Jr.);
and the following additional paratvpes.
N-E. N. G.: 2, Wau, Morobe Dist., 1400 m.
Mar. 29, 19^3 (Sedlaceks); 1, same locality,
1650 m, Feb. 23, 1962 (Sedlaceks); 1 L
Feramin, 1200-1500 m, May 2.3-31, 1959
(W. W. Brandt, Bishop Mus.). 1 c? , Okapa
(Busa), [1650-1800 m], Oct. 17, 1964
( Homabrook ) ; 1 c^ , Morae, Kukukuku
[Rge.], E. Highlands, 6000 ft. (c. 1850 m),
Mar. 1, 1964 ( Hornabrook ) . West N. G.:
1 A , Mt. Cyclops, 3500 ft. (1067 m). Mar.
1936 (Cheesman).
Mea.sured specimens. The 6 holotype
and 9 paratype from Sepalakembang.
Notes. This remarkable species would go
in Eiiryperip,ona if the latter were recog-
nized (see discussion under genus). So
far as I know it is unicpie in Feriiiona in loss
of all supraocular and latcMal prothoracic
setae and in the toothed posterior i
femora. It is comparable to P. proccra
Fauvel of Java in size but is broader, and
proccra has the above-mentioned setae and
does not have toothed A femora.
Perigona is.s.) pygmaea Andrewes
Andrewes 1930, Treubia, Supplement 7, pp. 3.34.
345.
Description (for recognition only). A
very small Pcriiiona characterized by wt-ak
frontal sulci and absence of 3rd (subapical)
eKtral punctures; IcMigth c. 2.0-2.5 mm.
Type. Vvo\\\ Bum, collected by Toxo-
peus; now in British Mus. (seen).
Occurrence in New Gtnnea. I*apiia: 4,
Dobodura, Mar.-July lf>44 (Darlington).
N-E. N. G.: 2, lower Busu R., Iluon Pen..
Mav 12 and 17, 1955 (E. O, Wilson. M.C.Z.,
The Carabid Beetles of New Guinea • Darlington 9
1 specimen numbered 1056 ) ; 20, Sattclberg,
1899 ( Biro ) ; 3, Stephansort, Astrolabe Bay,
1898, 1900 (Biro); 5, Aitape, Aug. 1944
(Darlington). West N. G.: 12, Maffin
Bay, Aug. 1944 (Darlington).
Notes. I have this species also from
Leyte and Luzon in the Philippines, and
have examined Andrewes' type from Biiru.
My Dobodura specimens were taken under
bark of rotting logs in rain forest.
Of all New Guinean Peritonei, this
seemed most likely to include short-winged
individuals, but I have examined all speci-
mens listed above, and all are in fact long-
winged.
Perigona is.s.) asfrolabica Csiki
Csiki 1924, Ann. Mus. National Hungary 21, p. 172.
Description. None required here; size,
and number and position of dorsal elytral
punctures are diagnostic, in New Guinea;
length 4.3-6.0 mm except only 3.8 mm in
an apparent dwarf of this species from
Dobodura.
Tijpe(s). From Stephansort, Astrolabe
Bay, N-E. N. G., collected by Biro in 1898;
in Hungarian National Mus. (seen).
Occurrence in New Guinea. Papua: 13,
Dobodura, Mar.-Julv 1944 (Darlington);
1, Kokoda, 1200 ft.' (366 m), June 1933
(Cheesman). N-E. N. G.: holotype + 2,
Stephansort, Astrolabe Bay, 1897 (Biro);
1, Sattelberg, 1899 (Biro); 1, Finschhafen,
Huon Pen., 150 m, Apr. 14, 1964 (Sed-
lacek); 6, Saidor, Gabumi Village, Finis-
terre Rge., June 24-30, July 1-21, 1958
(W. W. Brandt, Bishop Mus.); 1, Wum,
Upper Jimmi Vy., 840 m, July 18, 1955
(Gressitt); 1, Wau, Morobe Dist, 1150 m,
Nov. 7, 1961 (Sedlaceks); 1, same localitv,
1450 m, Feb. 5, 1963 (Sedlacek); 1, same
locality, 1700 m, Feb. 19, 1963 (Sedlacek);
1, Bulolo, "G. Pines," 600 m, Feb. 19, 1962
(Sedlacek). West N. G.: 1, Hollandia,
July-Sept. 1944 (Darlington); 1, same lo-
cality, May 1945 (B. Malkin, U.S.N.M.); 1,
Ifar, Cyclops Mts., 450-500 m, Sept. 9,
1962 (Sedlacek); 1, Maffin Bay, July 8,
1944 (E. S. Ross, California Acad.); 4,
Rattan Camp, 1150 m, Feb.-Mar. 1939
( Toxopeus ) .
Notes. P. astrolahica seems close to but
probably distinct from jacohsoni Andrewes
of Sumatra, in which the suture is red
(usually not red in astrolahica) and the
microreticulation of pronotum and elytra
more transverse. Two more, perhaps re-
lated, apparently undescribed species occur
in Luzon.
All specimens seen from New Guinea
have all usual supraocular and prothoracic
setae (or punctures marking positions of
setae) except that the 4 from Rattan Camp
and the 1 from 1700 m at Wau lack pos-
terior-lateral prothoracic setae. However,
presence or absence of these setae is ap-
parently simple dimorphism, for of 6 speci-
mens from Cape Gloucester, New Britain
(Darlington), 5 lack and 1 has posterior-
lateral setae. Because the distribution of
individuals with and without posterior-
lateral prothoracic setae may be of interest
in the future, I have listed ( above ) all New
Guinean specimens of the species in detail
rather than summarizing the species' oc-
currence.
Perigona is.s.) subcyanescens Putzeys
Putzeys 1875, Ann. Mus. Civ. Genoa 7, p. 732.
Csiki 1924, Ann. Mus. National Hungary 21, p. 172.
Andrewes 1930, Treul^a, Supplement 7, p. 334.
Louwerens 1953, Verhandlungen Naturforschenden
Gesellschaft Basel 64, p. 305.
horni Jedlicka 1935, Neue Carabiden aus Ostasien,
Part 10, pp. 18-19 (new synonymy).
Description. None required here. See
preceding Key to Sjjccies for identification;
length ( in New Guinea ) c. 3.3—4.0 mm.
Types. Of subcyanescens, from Andai,
near Dorey, West N. G., collected by
Beccari and D'Albertis, in Genoa Museum.
Of horni, from Imungan, Luzon, in Jed-
licka's collection. (See 2nd paragraph of
following Notes.)
Occurrence in New Guiiiea. Widely dis-
tributed and common on the island: 48
specimens from 13 localities scattered from
Milne Bay to Sansapor, and including
Dobodura and Wau (to 1100 m).
10 BuUetin Museum of Comparative Zoology, Vol. 137, No. 1
Notes. Outside New Guinea, this species
is recorded from West Sumba (Louwerens);
Borneo; Mindanao, Samar, and Luzon in
the Philippines; and doul:)tfully from Burn
(Andrewes); and I have seen specimens
from New Britain and the Solomons.
My identification of mbcyanescem is
based on specimens borrowed from the
Genoa Museum, one marked as compared
with Putzeys' type presumably by Csiki,
and my identification of liorni is based on
comparison with Philippine "cotypes" in
the British Museum.
Perigona is.s.) papuana Csiki
Csiki 1924, Ann. Mus. National Hungary 21, p. 173.
Description. None required here. See
Key to Species, and note subparallel cy-
lindrical form; length 2.6-3.7 mm.
Types. Lectotype (by present designa-
tion) and paratvpe from Stephansort,
Astrolabe Bay, N-E. N. G., 1898 (Biro);
in Hungarian National Mus. The specimen
( sex not determined ) now designated lecto-
type bears the original "Holotypus" label,
although no holotype was specified.
Occurrence in New Guinea. N-E. N. G.:
4 (in addition to the types), Stephansort,
1898 (Biro); 1, lower Busu R., Huon Pen.
May 17, 1955 (E. O. Wilson #1066, M.C.Z.),
in lowland rain forest; 1, Wau, 1300 m,
July 27 (year and collector not given).
Notes. This distinct species seems to be
confined to a limited area on the north side
of N-E. New Guinea.
Perigona is.s.) rossi n. sp.
Description. With characters of genus;
form as in Figure 2, c. as in astrolabica but
slightly more slender and convex; dark
castaneous, suture reddish, appendages red-
dish testaceous; rather shining, reticulate
microsculpture isodiametric on head, in part
transverse on pronotum, more transverse
on elytra. Head 0.70 width prothorax; man-
dibles pointed and slightly curved but not
notably elongate; eyes moderate, narrowly
enclosed behind by genae; antennae monili-
form; palpi with apical segment much
longer than subapical, narrowed and almost
pointed apically; frontal impressions short,
shallow, diverging posteriorly; 2 setae over
each eye. Prothorax: width length 1.43;
base apex 0.90; apex broadly emarginate,
with angles well defined but not advanced
beyond arc of emargination; base emargin-
ate-truncate, with basal angles distinct but
obtuse, slightly blunted; sides broadly
rounded, each with usual 2 setae; disc with
middle line distinct, baso-lateral impressions
weak. Elytra: width elytra prothorax 1.25;
humeri rounded-prominent; apices broadly
but irregularly rounded to obtuse but well
defined sutural angles; striae vaguely in-
dicated; intervals punctulate, 3rd with punc-
tures at c. Vs and % of length and at apex
just above submarginal channel. Secondary
sexual characters: $ unknown; 9 vvith
several setae each side apex last ventral
segment. Measurements: length c. 4.5;
\\'idth 1.7 mm.
Type. Holotype 9 (California Acad.)
from Maffin Bay, West N. G., June 1944
(E. S. Ross); the type is unique.
Notes. This species resembles astrolabica
but differs in details of shape especially of
prothorax, and in position ( nearer apex ) of
posterior elytral punctures.
Perigona is.s.) livens Putzeys
PutzL-ys 1873, Ann. Mus. Ci\ . (ienoa 4, p. 225.
Andrewes 1926, Cat. Philippine Caraliidae, p. 354.
1930, Cat. Indian Insects, Part 18, Ca-
raliidae, p. 265.
Tedlicka 1964, Heielienbacliia 2, No. 61, pp. 268,
270.
Description (lor recognition onh'). A
depressed, pale Teri'^ona s.s. with technical
characters indicated in the preceding Key
to Species; length (in New Guinea) c. 3.3
mm.
Ty])r. DoublfulK from Coromandt>l,
India; via C^haudoir and then Oberthiir
Golls. to Paris Nhis. (not seen).
Occurrence in New Guinea. Papua: 2,
Dobodura, Mar.-July 1944 (Darlington).
Notes. P. liv('i}s is listed b\' Andrewes
The Carabid Beetles of New Guinea
Darlington
11
( 1926 ) from Luzon and Mindanao in the
Philippines, and Andrewes ( 1930 ) indi-
cates that he saw Putzeys' type. I have a
PhiHppine (SE. Bataan) specimen identi-
fied as livens by comparison with An-
drewes' collection. The New Guinean speci-
mens do not match Philippine ones exactly,
but my material is too limited to justify
separating the New Guinean form even as
a subspecies.
Perigona (s.s.) plagiata Putzeys
Putzeys 1875, Ann. Mus. Civ. Genoa 7, p. 734.
Andrewes 1930, Cat. Indian Insects, Part 18,
Carabidae, p. 266.
Csiki 1924, Ann. Mus. National Hungary 21, p.
172.
1931, Coleop. Cat., Carabidae, Harpalinae
5, p. 898 ( see for additional synonymy and
references ) .
Jedlicka 1935, Neue Carabiden aus Ostasien, Part
10, p. 18 (in key).
1964, Reichenbachia 2, No. 61, pp. 268,
271.
Van Emden 1937, Stettiner Ent. Zeituns 98, p. .35.
annamita Fauvel 1907, Revue d'Ent. 26, p. 104.
Andrewes 1933, Ann. Mag. Nat. Hist. (10) 11,
p. 110.
Description. A small, brownish testaceous
Perigona s.s. with head and much of elytral
discs darker brown, and with technical
characters as indicated in preceding Key
to Species; length c. 2.2-2.8 mm.
Types. Of plagiata, from Aru and Kei
Islands, collected by Beccari, and from
Andai, West N. G., collected by Beccari
and D'Albertis; in Genoa Mus. Of an-
namita, from Ceylon, Annam, Singapore,
and Andai, West N. G., the specimen(s)
from New Guinea collected by Raffray;
Andrewes (1933) found a "type" in the
Maindron Collection, Paris Mus. Lectotypes
for both plagiata and annamita should be
fixed by the next reviser, after examination
of all the original type material. (Types
not seen. )
Occurrence in Neio Guinea. Common
and widely distributed. I have seen 145
specimens from localities scattered over
most of the length of the island, from
Dobodura to Sansapor; most at low altitudes
but single specimens found at 1100 and
1200 m at Wau.
Notes. Andrewes (1930) records plagi-
ata from a wide range, from SE. Asia and
Japan across the Malay Archipelago to
the Philippines and New Guinea, and
Van Emden lists it from the New Hebrides.
Csiki ( 1924 ) records it from Australia on
the basis of specimens ( which I have seen )
in the Hungarian National Mus., but I think
this is an error (see Darlington 1964,
Psyche 71, p. 125). Perigona rufilabris
(Macleay) of eastern Australia is a similar
but larger species,
Perigona {Trechicus) nigriceps (Dejean)
Dejean 1831, Species General Coleop. 5, p. 44
(Bcmbidium).
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae
5, p. 897 ( see for synonymy and additional
references ) .
Jedlicka 1935, Neue Carabiden aus Ostasien, Part
10, p. 18 (in key).
1964, Reichenbachia 2, No. 61, pp. 268,
270, fig. 2.
Jeannel 1941, Rev. francaise d'Ent. 8, p. 141.
lititra Perroud and Montrousier 1864, Ann. Soc.
Linneenne Lyon 11, p. 72 (Trechus) (new
synonymy ) .
bcccaiii Putzeys 1875, Ann. Mus. Civ. Genoa 7,
p. 732 (new synonymy).
J)iwi Csiki 1924, Ann. Mus. National Hungary
21, p. 173 (new synonymy).
klickai JedHcka 1935, Neue Carabiden aus Ostasien,
Part 10, pp. 18, 19 (new synonymy).
Description (for recognition only). See
preceding Key to Species of Perigona of
Netv Guinea; color either testaceous with
head and apices of elytra darker, or brown-
ish castaneous with suture (and of course
appendages) pale, or intermediate with
elytral disc partly but not entirely clouded;
technical characters include eyes relatively
large and prominent, front isodiametrically
microreticulate, and elytra more conspicu-
ously 3-punctate than usual in the genus,
with posterior puncture usually almost in
line with the others; length c. 2.5-3.0 mm.
Types. Of nigriceps, from North Amer-
ica, sent to Dejean by Leconte; now in
Oberthiir Coll., Paris Mus. Of litiira, from
Kanala, New Caledonia; location of type(s)
12 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
unkno\\'n. Of heccarii, from Sarawak, Bor-
neo, collected by Doria and Beccari; now
in the Genoa Mus. (a lectotype should be
designated by next reviser). Of hiroi, I
now designate as lectot\pe a 9 from Ma-
dang (Friedrich-Wilh.-hafen), N-E. N. G.,
1896 (Biro, Hungarian National Mus.);
this specimen is from Csiki's original series
and is labeled "Holotypus," but the desig-
nation has not been published until now.
Of klichai, from Mt. Makiling, Luzon; in
Andrewes Coll., British Mus. (Types of
hiroi and klickai onlv seen.)
Occurrence in Neic Guinea. Common and
widely distributed at low altitudes: more
than 160 specimens from many localities,
from Milne Bay to "Dorey" and Biak Is.,
and including Dobodura and Wau ( to 1.300
m ).
Notes. P. nigriccps is cosmopolitan,
carried by man to all tropical and warm
temperate regions.
P. lifura, described from New Caledonia
but supposedly widely distributed in the
Malay Archi]ielago, was unknown to An-
drewes. The description fits the dark form
of niii,riceps, and the fact that the type(s)
occurred under vegetable detritus also fits
niiiriceps. (The habitat of niiiriccps is noted
under the genus. ) P. heccarii is another
name lor the dark form of this species ( I
do not consider the dark form worth distin-
guishing by name), and hiroi and klickai,
of which I have seen the types, are also
based on dark examples of ni^riceps.
Perigona {Trechicus) erimae Csiki
Csiki 1924, Ann. Mus. National Hungary 21, p. 173.
Description (for recognition only ). With
characters of Perigona, subgenus Trechicus;
broad, moderately convex; black or castane-
ous with suture usually not paler; eyes
forming c. right angles with neck, but some-
what variable; front with or without (lighth'
impressed) isodianietrie reticulations; pro-
thorax with sides not or slightly sinuate
postci ioily, with angles well defined but
obtuse; elytra not or hiintly striate, with
little or no punctulalion, with subinarginal
channel moderately broad behind puncture-
triangle; length c. .3.2-3.6 mm.
Type. From Erima, Astrolabe Bay, N-E.
N. G., 1896 (Biro); in Hungarian National
Mus. (seen).
Occurrence in Neic Guinea. Thirty-four
specimens from numerous localities in
eastern and central New Guinea, from
Milne Bay and Dobodura to Hollandia and
Cyclops Mts.; not yet found farther west in
New Guinea; most from low altitudes but
reaching 1200 m at Wau.
Notes. Specimens of this species vary
considerablv. The eves varv in size and in
development of genae but usually form
nearly right angles with the neck. Reticulate
microsculpture of the head may be distinct
(but light), or partly obliterated, or c.
absent. And there is some \ariation of
other characters. However, the variation
is not primarily geographic, but occurs at
single localities. I think, but cannot be
quite sure, that only one variable species is
involved.
Csiki's type of erimae is large, with eyes
large and genae slight, and with the front
distinctly reticulate. Proportions of the type
are head 0.83 width prothorax; prothoracic
width length 1.50, base/apex 1.03; width
elytra prothorax 1.53.
Although erimae is known onl\- from New-
Guinea, somewhat similar but apparently
distinct species {andreicesi Jedlicka, arroici
Jedlicka) occur in the Philippines.
Perigona (Trechicus) ludovici Csiki
Csiki 1924, Ann. Mus. National Ilun.uan 21, p. 174.
Description ( for recognition onl\). Form
of PcriiiO)\a, subgenus Trechicus; small;
dark, like dark ni'^riceps but suture not or
not conspicuously reddish; head with e\'es
smaller than in niil.rice))s. Iront less dis-
tinctK' reticulat(> and with rtiieulations
more trans\'erse espeeialK posteriorly;
elytra with .3rd (posti'rior) punctures nearer
sutm(>; length c. 2.7-.3.3 mm.
T\i])cs. Lectotype (present designation)
from Mt. llansemann, Astrolabe Bay, N-E.
N. (;., 1901 (Biro, Tbnigarian National
The Carabid Beetles of New Guinea • Darlington
13
Mus. ), and 8 additional original (co) types,
2 with same data as lectotype and 6 from
Madang (Friedrich-Wilh.-hafen), 1900 and
1901 (Biro) (seen).
Occurrence in Neiv Guinea. Seventy-five
specimens from numerous localities over
almost the whole length of New Guinea
(Milne Bay and Dobodura to the Vogel-
kop); at low altitudes, none above 550 m.
Notes. P. ludovici is compared with
nigriccps in the preceding Description. P.
ludovici is in fact closer to erimac but has
the head narrower, eyes relatively smaller,
and microreticulation of posterior part of
head usually more transverse. Also, ludovici
averages smaller than erimae, although
measurements of length overlap: ludovici,
c. 2.7-3.3; erimae, c. 3.2-3.7 mm. (Csiki
gives 2.5-2.8 mm for ludovici and 3.5 mm
for erimae. ) Nevertheless, these species are
very similar and some individuals are dif-
ficult to place. Proportions of the lectotype
of ludovici are head 0.76 width prothorax;
prothoracic width/length 1.49, base/apex
1.08; width elytra/prothorax 1.59.
Both erimae and ludovici live among dead
leaves on the ground in rain forest.
Perigona (Trechicus) lebioides Csiki
Csiki 1924, Ann. Mus. National Hungary 21, p. 174.
Description ( for recognition only ) . Form
of small, very broad, moderately convex
Perii^ona, subgenus Trechicus; castaneous
with suture not or only faintly reddish;
prothorax with sides not strongly sinuate
and not denticulate posteriorly; elytra with
submarginal depressed space very narrow
behind puncture-triangle; elytra faintly or
irregularly striate, not or not much punc-
tulate; length 2.8-3.3 mm.
Types. I now designate as lectotype the
specimen marked "Holotypus" by Csiki. It
is from Erima, Astrolabe Bay, N-E. N. G.,
1896 (Biro) in Hungarian National Mus.
(seen). Seven paratvpes are from Sattel-
berg, N-E. N. G., 1899 (Biro). (Two addi-
tional specimens labeled as paratypes of
lebioides, from Simbang, Huon Gulf, Biro,
in Plungarian National Museum, are not
lebioides but erimae. )
Occurrence in Neiv Guinea. Sixty-one
specimens (including 44 from Dobodura)
from localities in all 3 political divisions of
New Guinea; most at low altitudes but 1
from Sibil, Star Rge., at 1260 m ( Leiden
Mus.).
Notes. This, like the other small Perigona
of subgenus Trechicus that occur in New
Guinea, lives among dead leaves on the
floor of rain forest. Biro presumably col-
lected the types by sifting. I took mine
by throwing raked-up leaves and leaf mold
into still water, and picking up the beetles
as they came to the surface.
Perigona {Trechicus) cordens n. sp.
Description. With characters of Perigona,
subgenus Trechicus; form broad, rather
convex; black or castaneous, suture not or
not much paler, elytra subiridescent, ap-
pendages reddish testaceous; reticulate
microsculpture faint, not clearly visible at
c. 100 X l)ut apparently isodiametric on
front, somewhat transverse posteriorly on
head, fine and strongly transverse on
pronotum and elytra. Head 0.80 and 0.79
width prothorax; eyes rather large, forming
c. right angles with neck, mandibles average
for genus; antennae with middle segments c.
IVi X long as wide; front with impressions
irregular but distinct, margined externally
by short elevations. Prothorax cordate;
width length 1.47 and 1.47; base apex 0.98
and 0.97; sides strongly sinuate about Vs
from base; posterior angles nearly right but
blunted; disc with fine middle line, shallow
poorly defined baso-lateral impressions.
Elytra short, wide; width elytra prothorax
1.64 and 1.66; submarginal impressed space
very narrow^ behind puncture-triangle; each
elytron with parts of at least 6 striae, inner
ones moderately impressed and irregular
or vaguely punctate; intervals not distinctly
punctulate, 3rd 3-punctate. Secondary sex-
ual characters normal: c5 front tarsi with
3 segments (only apex of 1st) narrowly 2-
seriately squamulose; 6 with 2, 9 c. 4
14 BuUetin Museum of Comparative Zoology, Vol. 137, No. 1
setae at apex last ventral segment. Mea-
surements: length c. 3.4; width 1.5-1.6 mm.
Types. Holotype $ (M.C.Z., Type No.
31,345) and 3 paratypes (broken <^ , 9 $ )
all from Dobodura, Papua, Mar.-July 1944
( Darlington ) .
Measured speeimens. The 6 holotype
and 1 9 paratype.
Notes. This species occurred in leal
litter in rain forest, with erimae, ludoviei,
and lehioides, from all of which cordens is
immediately distinguishable by its strongly
cordate prothorax.
Perigono (Trechicus) denfifer n. sp.
Description. With characters of Perigona,
subgenus Trechicus; form as in Figure 3;
broad, moderately convex; reddish castane-
ous with suture slightly paler, appendages
reddish testaceous; shining, not iridescent,
microsculptme absent or nearly so. Head
0.78 and 0.78 width prothorax; mandibles
slender, pointed, weakly arcuate near apex;
eyes moderately large but less prominent
than usual, forming obtuse angles with
neck; antennae with middle segments c.
Vri X long as wide; front with slight median
puncture and distinct short anterior frontal
impressions. Prothorax broadly subcordate,
very wide anteriorly; width/length 1.40
and 1.45; base/apex 0.95 and 0.88; sides
weakly rounded, strongly converging pos-
teriorly almost to base, then abruptly sinu-
ate with basal angl(\s right-denticulate; disc
with usual middle line and transverse im-
pressions, basal transverse impression sub-
foveate at middle and running into slightly
dccjier but poorly defined baso-lateral im-
pressions. FJijtra wide; width elytra pro-
thorax r. 1.61 and 1.60 (exact measurement
impossible because elytra spread in both
specimens); submarginal impressed space
very narrow behind puncture-triangle; 6
abbrc-viated striae on each elytron, inner on(\s
impressed, all plainU' punctate; int(>r\als
not punclulate, 3rd 3-puuctate. Secondary
.sexual characters as in jireceding species
(cordens). Measuremenl.s: length r. 3.2-
3.4; width r. 1.4-1.5 mm.
Types. Holotype S (M.C.Z., Type No.
31,346) and 1 9 paratype both from Milne
Bay, Papua, Dec. 194.3 (Darlington).
Notes. The form of prothorax, absence of
microsculpture, and impressed punctate
elytral striae are diagnostic of this species.
Tribe LiCININI
Sloane 1898, Proc. Linnean Soc. New South Wales
23, pp. 487 ff . ( Australian genera ) .
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae
.5, p. 899.
Ball 1959, Mem. American Ent. Soc, No. 19, p.
5 ( see for synonymy and additional references ) .
Most Licinini, including all those known
from New Guinea, have the labrum and
usually also the clypeus deeply emarginate,
the labrum often so deeply so as to appear
2-lobed. This alone is almost a sufficient
recognition character of the tribe, in New
Guinea. Other diagnostic characters are
discussed by Ball ( 1959, pp. 5-8).
Licinines are nearly world-wide in distri-
bution but are relatively few in Central
and South America and relatively numerous
in Australia: about 10 genera, including
some that are probably primitive or relict,
occur in Australia. Five genera occur in
New Guinea: Badister, which is \\idel>
distributed in other parts of the world;
Omestes, a monotypic genus confined to the
eastern part of the Mala\^ Archipelago;
and Physolaesthus, Dicrochilc. and Micro-
feronia, which are primarih Australian.
Thrc^e species of Dicrochile and one of each
of the other genera are known in N(^w
Guinea. All the New Guinean species are
winged, except Microferonia haro.
The following Key is based on Balks
(1959, p. 11) key to Oriental licininc^
genera.
Kky to Gfnfha ok Licinini ok Xknv Ciinka
1. One mandible deeply notched alioxe, with a
prominent boss beliind tiir iiotrh 2
Neither mandible notched as described 4
2. I.cil mandible notched; only basal seginent
oi antenna glabrous (p. 15) ._. Badister
- \\\si}\{ mandible notchecb each antciuKi w itii
3 segments glabrous 3
3. Smaller (c. 5 mm); elytra not spined (p.
15) Phtjsolacsthus
The Carabid Beetles of New Guinea • Darlington
15
- Larger (c. 11-15 mm); elytra with short
apical spines (p. 16) Omcstc.s
4. Form Agonu)n-\\\.e; mandibles blunt at apex
(p. 16) — Dicrochile
- Form elongate-oval with very small head;
mandibles ( at least the right one ) con-
spicuously 2-dentate at apex, with upper
tooth large, acute (p. 18) Miciofeiouia
Genus BADISTER Clairville
Anonymous [Clairville] 1806, Entomologie Hel-
vetique 2, p. 90.
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae
5, p. 901 (see for synonymy and additional
references ) .
Jeannel 1942, Faune de France, Coleop. Carabiques,
Part 2, p. 1000.
Louwerens 1956, Treubia 23, p. 236 (key to
Indonesian species ) .
Ball 1959, Mem. American Ent. Soc, No. 16, pp.
189-191.
Diagnosis. See preceding Key.
Description. None required here.
Type species. Carabus bipustidatus Fab-
ricius, of Europe, etc.
Generic distribution. Temperate and
tropical Eurasia, the Malay Archipelago,
and eastern Australia; Africa and Mada-
gascar; North America and some West
Indies, but not South America.
Notes. See Jeannel (1942) and Ball
( 1959 ) for further information on this
widely distributed genus.
Badisfer (Baudia) sundaicus Andrewes
Andrewes 1926, Ann. Mag. Nat. Hist. (9) IS, p.
275.
Louwerens 1956, Treubia 23, p. 236.
Description. See Andrewes (1926); length
c. 4.0-4.5 mm.
Type. From Soekaboemi, Java; in
Andrewes Coll., British Mus. (seen).
Occurrence in New Guinea. Papua: 4,
Dobodura, Mar.-July 1944 (Darlington).
N-E. N. G.: 1, Maprik, Sepik Dist., 150 m,
Dec. 29, 1959-Jan. 17, 1960 (T. C. Maa,
Bishop Mus.). West N. G.: 2, Hollandia,
July-Sept. 1944 (Darlington).
Notes. I tentatively identify as sundaicus
specimens from Siani and the Malay Pen.
(British Mus.); Sumatra; Java; Luzon and
Leyte in the Philippines; Morotai Is. in
the Moluccas; New Guinea (listed above);
New Britain; and widely scattered locali-
ties in eastern Australia. Specimens from
all these places have the mandibular and
antennal characters indicated in the pre-
ceding Key to Genera. However, variation
is obvious, and further study may show
that more than one species is involved.
Specimens of this and related species
that I have collected were usually among
dead leaves and vegetation on the ground
in very wet places by standing (not running)
water.
Genus PHYSOLAESTHUS Chaudoir
Chaudoir 1850, Bull. Soc. Nat. Moscow 23, Part
1, No. 2, p. 411.
Diagnosis. See preceding Key to Genera.
Description. See Chaudoir (1850), and
following Notes.
Type species. P. australis Chaudoir, of
Australia.
Generic distribution. Primarily Austra-
lia; one species described from New
Zealand; and the following species (if
correctly assigned) on New Guinea, Java,
and the Philippines.
Notes. I have not been able to identify
australis in the Australian material before
me. Chaudoir does not describe its antennal
pubescence but states that the right man-
dible is tuberculate, and this character is
always associated with 3 antennal segments
glabrous, among Australian licinines known
to me. Whether the following species is
really a Fhysolaesthus and how this genus
is related to Badistcr will have to be decided
by future revisers.
Fhysolaesthus caviceps (Andrewes)
Badistcr caviceps Andrewes 1936, Ann. Mag. Nat.
Hist. (10) 17, p. 312
Louwerens 1956, Treubia 23, p. 236.
Description. See Andrewes, and my
Figure 4; length c. 5 mm.
Type. A $ from Toeloengagoeng, Java;
in Andrewes Coll., British Mus. (seen).
Occurrence in New Guinea. West N. G.:
4, all from Wissel Lakes area, as follows:
16 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
1, Itouda, Kamo Vy., 1500 m, Aug. 12,
1955 (Gressitt), in light trap; 1, Lake
Paniai, 1570 m, Aug. 28, 1939 (H. Boschma,
Leiden Mus.); 2, Enarotadi. LSOO m, Aug.
1, 1962 (Sedlacek).
Notes. I have seen specimens with the
characters of caviceps from Java and
Luzon as well as New Guinea but am not
sure whether they represent one species or
two or more related species. Except for
the different mandibles and antennae, this
species is remarkably similar to Badisfcr
sundaicus (above), and I think the habitats
of the two species are similar, judging
from what I have seen of them in the
Philippines.
Genus OMESTES Andrewes
Andrewes 1933, Treubia 14, p. 276.
Diafinosis. See preceding Key to Genera.
Description. See Andrewes.
Type species. Omestes torta Andrewes,
below.
Generic distribution. Same as that of O.
torta, below.
Notes. I suspect that Omestes torta may
prove to be only a large, specialized
(spined) Physokiestlius, but I shall leave a
decision about this to future revisers,
Omestes torta Andrewes
Andrewes 1933, Treuljia 14, p. 277.
Louvverens 195(-), Treul)ia 23, p. 224.
Description. See Andrewes, and m>
Figure 5; length 11-14 mm.
Type. A £ from Sangi Is.; in Andrewes
Goll., British Mus. (seen).
Occurrence in New Gttinea. Papua: 1,
Dobodura, Mar.-July 1944 (Darlington); 1,
Milne Bay, Dec. 1943 (Darlington); 2,
Kiunga, Fly R., Aug. 14-17, 1957 (W. W.
Brandt, Bishop Mus.); 1, Darn Is., Mar. 16-
31, 1936 (Archbold Expedition, A.M.N.H.);
1, Woodlark Is. (Murua), Kuhunadau ITill,
Apr. 16-22, 1957 (W. W. lirandl. Bishop
Mus.). West N. C;.: 19, Mollandia, Julv-
Sept. 1944 (Darlington); 1, Maffen, Tor R.
(mouth), 4 km E. of Hollandia, JuK 2.
1959 (T. G. Maa, Bishop Mus.), at light;
1, Bernhard Gamp, 50 m, Apr. 12, 1939
( Toxopeus ) .
Notes. Omestes torta is now known from
New Guinea, the Moluccas ( Ilalmahera
and Morotai), Celebes, the Sangi and
Talaud Islands, and the Philippines
(Leyte). My material is not sufficient to
show details of geographic variation. The
insect lives among dead leaves and vege-
tation on the ground in deep swamps.
Genus DICROCHILE Guerin
Guerin 1<S46, Ann. Soc. Ent. France (2) 4, Bnll.
p. cm.
Sloane 1923, Proc. Linnean Soc. New South Wales
48, pp. 35-36 (key to Australian species).
Csiki 1931, Colcop. Cat., Carabidae, Harpalinae 5,
p. 921 ( see for synonym\- and additional refer-
ences ) .
Diagnosis. See preceding Key to Genera.
Description. None required here.
Ty})c species. Presumabh- Dicrochilc
fabrii Guerin, of New Zealand. (I do not
wish to designate a type species. If no
formal designation has been made, it should
be left to the next reviser. )
Generic distribution. New Zealand, Aus-
tralia, New (Guinea, Moluccas (Obi Is.),
Solomons ( a probably undescribed species
near alternans from Bougainville), New
(Caledonia.
Notes. All species of this genus that I
know, in Australia as well as New Ciuinea,
arc \\'inged. Most of them live in swamps
or other wet places, but alternans (de-
scribed below) is a nicsophile.
Kkv k) Species oi- Dickociiilk ok Xeav Guinea
1. I']l\tra w'itli atiitc tcctli or sliort spini's at
sutnral and outer-apical ant^lcs; dorsal elytral
intcr\als ccjnal or ncarK so (p. 10) acuta
- Elytra not toothed or spined; dorsal ehtral
inter\als uncciual _ ._ 2
2. I'^ront of head normalK' con\e\; smaller,
length 11.5-12.5 nun (p. 17) altcntans
— I'roiit ol licad sliyhtK depressed; hir^cr,
length 13. .5-14. 5 nnn ( [i. IS) tiro
Dicrochile acuta n. sp.
Description. Form (Fig. 6) ol ALi,()num-
likc Dicrochilc; piccous black, lateral mar-
gins ol proiiotiiin ;iiul elytra slightK trans-
The Carabid Beetles of New Guinea • Darlinfiton
17
lucent, elytra iridescent; microsculpture
fine and isodiametric on front, indis-
tinct (at 100 X ) but probably strongly
transverse on pronotum and elytra. Head
0.74 and 0.72 width prothorax; eyes large;
front slightly convex, weakly impressed at
sides anteriorly. Frothorax quadrate-sub-
cordate; width length 1.35 and 1.39; base/
apex 1.13 and 1.09 (base measured across
posterior-lateral setae); base slightly emar-
ginate, not margined; apex broadly emar-
ginate, with impressed marginal line; sides
rounded except c. straight toward base;
margins rather broad, moderately explanate,
each with usual 2 setae ( at base and before
middle); basal angles very obtuse, almost
rounded; pronotum with usual impressions,
impunctate at middle, closely punctate at
base and sides. Elytra subparallel, slightly
narrowed toward base; width elytra/pro-
thorax 1.46 and 1.40; outer-apical and
sutural angles each with an acute tooth or
very short spine; striae shallow, faintly
punctulate; intervals c. flat, subequal on
disc, 3rd with 2 punctures attached to 2nd
stria. Legs: middle and hind tarsi broadly
grooved each side above; 5th segment hind
tarsi with e. 6 strong setae each side below.
Secondary sexual charaeters: i front tarsi
somewhat obliquely dilated, with 3 seg-
ments squamulose below; 6 with 1, 9 with
2 setae before apex each side last ventral
segment. Measurements: length c. 12.5-
15.5; width c. 5.0-6.4 mm.
Types. Holotype c^ (A.M.N.H.) and 1 9
paratype (M.C.Z., Type No. 31,347) from
Lake Daviumbu, Fly R., Papua, Sept. 1-10
(holotype) and Aug. 19-30 (paratype),
1936 (Archbold Exp.), evidently taken in
a light trap; 1 9 paratype (Bishop Mus.),
Oriomo R., Papua, 6 m, Feb. 13, 1964,
"H. C", in light trap; 1 9 paratype, "Highl.
Agr. Exp. Sta./Aiyura, E. Highl./D", N-E.
N. G., 5600 ft. (c. 1700 m), May 26, 1960
(J. J. H. Szent-Ivany, Dept. Agr. Port
Moresby), at light; 1 S paratype (Bishop
Mus.), Nabire, S. Geelvink Bay, West
N. G., 10-40 m, Oct. 7, 1962 (H. Holtmann),
m light trap in jungle.
Measured specimens. The 6 holotype
and 9 paratype from Lake Daviumbu.
Notes. This species is closely comparable
only with D. gigas Castelnau, of Australia.
It resembles gigas in most technical char-
acters including the denticulate-spinose
elytra, but differs from gigas in being much
smaller (Australian gigas measure 20 mm
and over) and in having a relatively nar-
rower prothorax and less impressed front.
Louwerens (Treubia 24, 1958, p. 250)
records D. gigas from Obi Is. in the Moluc-
cas, on the basis of 2 specimens 18 mm long,
which differ in some details from the single
Australian specimen of gigas with which
they were compared. Whether the Obi Is.
specimens are gigas or a related species
remains to be decided, as Louwerens hints.
Dicrochile alternans n. sp.
Description. Form (Fig. 7) of rather
broad Australian Dicrochile (e.g., goryi
Boisduval); black, appendages blackish ex-
cept outer segments of antennae brown;
both sexes moderately shining but not
iridescent, with reticulate microsculpture
faint and c. isodiametric ( where detectable)
on front, vague or irregular but apparently
transverse on pronotum and elytra. Head
0.74 and 0.73 width prothorax; eyes mod-
erate; front convex at middle, irregularly
longitudinally impressed each side anteri-
orly. Frothorax slightly transverse, width/
length 1.36 and 1.38; base/apex 1.15 and
1.15; base and apex broadly emarginate,
apex strongly and base less strongly or in-
distinctly margined; sides broadly rounded;
margins broadly flattened and moderately
reflexed posteriorly, each with usual 2
setae, at base and before middle; basal
angles broadly rounded; disc convex, with
usual impressions, punctate at base, sides,
and apex, impunctate at middle. Elytra
elongate-subquadrate; width elytra/pro-
thorax 1.48 and 1.54; apices sinuate but not
denticulate; striae deep, punctulate; in-
tervals convex, unequal on disc (3rd, 5th,
7th nearly 2x as wide as others at % of
elytral length), 3rd usually 2-punctate with
18 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
punctures near or behind ^.-i and 7;5 of
elytral length, but anterior puncture some-
times duphcated on one or both elytra.
Legs: middle and hind tarsi sulcate each
side above; 5th segments hind tarsi with c.
6 strong setae each side below. Sccondanj
sexual characters: 6 front tarsi dilated and
squamulose as usual in genus; 6 with
1, $ 2 setae each side last ventral segment.
Measurements: length c. 11.5-12.5; width
4.5-5.0 mm.
Types. Holotype $ (M.C.Z., Type No.
31,34(S) and 12 paratvpes from Chimbu Vv.,
Bismarck Rge., N-E'. N. G., 5000-7500 ft.
(c. 1500-2300 m), Oct. 1944 (Darlington);
1 paratype, Feramin, N-E. N. G., 1200-1500
m, June 15-18, 1959 (W. W. Brandt, Bishop
Mus. ); 1 paratvpe, Minj, W. Highlands,
N-E. N. G., 5200 ft. (c. 1600 m), May 20,
1960 (J. 11. Barrett, Dept. Agr. Port
Moresby), by mercury vapor lamp.
Additional material. Papua: 1 c5 , S.
Highlands, Aiyuro nr. Mendi, 1530 m, Oct.
7, "l958 (Gressitt), in light trap. West
N. G.: 16, Wissel Lakes, Urapura-Itouda,
Kamo Vy., 1500 m, Aug. 12, 1955 (Gressitt).
Measured specimens. The i holotype
and 1 9 paratype from Ghimbu Vy.
Notes. The usually 2-punctate 3rd elytral
intervals and the deep, punctulate striae
suggest that this new species is allied to
the common Australian Dicrochilc goryi
Boisduval, but the elytral intervals of goryi
do not alternate in width, and there arc
other smaller differences.
I found th(> Ghimbu specimens under
cover on the ground in fairly open places.
Dicrochile tiro n. sp.
Descriptio)\. Similar to the preceding
(aUerruitis) but larger, with flatter front
and relatively wider prothorax. Head 0.71
and 0.69 width prothorax, lormed as in
alternans except Hatter anteriorly. Vro-
thorax: width length 1.39 and 1.45; base
apex 1.16 and 1.16; otherwise as in alternans.
Elytra: width clytra/prothorax 1.47 and
1.40; most details including alternation of
elvtral intervals c. as in (illcr)unis: 3rd in-
terval 2- to 4-punctate, the number of punc-
tures often different on the 2 elytra of 1
individual (actual punctures on the left
and right elytra of 6 individuals are 2-3,
2-2, 4-2, 2-2, 2-2, 2-3). Measurements:
length 13.5-14.5; width 5.5-6.1 mm.
Types. Holotype 9 (Leiden Mus.) and
7 paratypes (some in M.G.Z., Type No.
31,349) all from Wissel Lakes, West N. G.,
as follows: holotype and 4 paratypes. Lake
Paniai, 1750 m, and 1 paratype, Arabu
Camp, ISOO m, various dates in Sept., Oct.,
Nov. 1939 (H. Boschma, Leiden Mus.); 2
paratypes, Enarotadi, 1800-1900 m, July 31,
Aug. 9, 1962 (Sedlacek).
Measured specimens. One 6 paratype
from Lake Paniai and the 9 holotype, in
this order.
Notes. Sufficiently compared with al-
ternans in the preceding Description and in
the Key to S})ecies of Dicrochile of New
Guinea. Whether tiro is a separate species
or a local form of alternans is uncertain.
The matter is complicated by the occur-
rence of a specimen of alternans in the
Wissel Lakes area.
Genus MICROFERONIA Blackburn
Blackburn 1(S90, Proc. l-iiuicaii Sor. New Sdutli
Wales (2) 4, p. 738.
Sloane 1898, Proc. Linncaii Soc. Xcw South Wales
23, pp. 490-491 (Australian species).
Csiki 1931, Coleop. Cat., Caiabidae, Harpalinae
5, p. 920 (see lor additional rcferenc(^s ).
Ck'nyccnis Andrewes 1933, Trenliia 14. \i. 277
( new s\non\ luy).
Diagnosis. See Key to (Uiura of Licinini.
Description. None re(inirc'd here; see
Notes, below.
Ty})e s))ccies. Oi M icrofcronicL M. adc-
laidae Blackburn, Australia; of Genycerus,
Cx. hicanoides Andrewes, of Ja\a.
Generic distribution. Aii.«^tralia, N<mv
(Fiiiiioa. Ja>a. and presumably inlciAcniiig
islands.
Notes. W'lu'n Anchx'wcs described Gouj-
cerus, he thought the mandiblc\s uniciue
among Licinini, but he was not lamiliar
with the Australian members ol the tribe. 1
l)a\(' seen the t\ pe ol (U'nyccnis lucanoidcs
The Carabid Beetles of New Guinea • Darlington
19
and have a photograph of it, and it seems
to me that the mandibles are eomparable
to those of Microfcronia. The diseovery of
another comparable species in New Guinea
links the Australian and Javan forms geo-
graphically. I therefore tentatively suggest
the synonymy cited above.
Microferonia baro n. sp.
Description. Form as in Figure 8, elon-
gate-oval with very small head; brownish
piceous, legs and antennae slightly reddish;
moderately shining, reticulate microsculp-
ture c. isodiametric on front, transverse on
pronotum, more transverse on elytra. Head
0.51 width prothorax; eyes large, genae
short; 2 setae over each eye; antennae with
3 basal segments glabrous; right mandible
2-dentate, with inner tooth strong and
acute ( left mandible probably c. similar but
partly hidden ) ; front almost evenly convex
except with slight frontal impressions an-
teriorly; clypeus subtruncate, with narrow
transverse membrane; labrum emarginate to
c. middle of length, with lobes equal;
mentum without tooth; ligula and paraglos-
sae apparently subequal, ligula apparently
2-setose; palpi slender except apical seg-
ments of both pairs slightly thickened.
Prothorax: width/length 1.40; base/apex
1.70; base truncate-emarginate, vaguely mar-
gined at middle; apex broadly emarginate,
with marginal line entire; sides rounded
anteriorly, nearly straight toward base, nar-
rowly margined, each with 2 setae, at base
and before middle; disc broadly convex
except depressed baso-laterally, impunctate,
with middle line distinct but transverse im-
pressions c. obsolete. Elytra long-oval;
width elytra prothorax c. 1.30; margins
entire at base, bluntly (almost rectangularly)
angulate at humeri, not distinctly sinuate
toward apex; sutural angles narrowly
rounded; striae fine, irregular but scarcely
punctulate; intervals nearly flat, somewhat
irregular but scarcely alternating; each 3rd
interval with a conspicuous seta-bearing
puncture about % from base, a less con-
spicuous puncture without seta near or be-
hind middle, apparently no more-posterior
puncture. Inner icin(j,s evidently atrophied.
Loner surface almost impunctate but ex-
tensively alutaceous, not pubescent; met-
episterna less than Vj longer than wide.
Legs: tarsi slender, not sulcate above; 5th
segments hind tarsi with 5 long setae each
side below. Secondary sexual characters:
$ with 3 segments each front tarsus mod-
erately dilated, squamulose below; i with
1 seta each side last ventral segment; i
copulatory organs as in Figure 170; ? un-
known. Measurements: length c. 8; width
3.4 mm.
Type. Holotype $ (M.C.Z., Type No.
31,350) from Mt. Wilhelm, Bismarck Rge.,
N-E. N. G., 7000-10,000 ft. (2135-3050 m),
Oct. 1944 ( Darlington ) ; the type is unique.
It was taken on the ground under cover
in mountain rain forest.
Notes. Microferonia baro is more oval
and smaller-headed than M. (Genycerus)
lucanoides (Andrewes) of Java. I do not
have a specimen of lucanoides, and I do
not want to dissect the mouth parts of the
single type of haro (which should be re-
served for specialists in Licinini), but so
far as I can determine the two species are
similar in generic characters although dif-
ferent in detail. ^L baro is larger, more
oval, and smaller-headed than anv Aus-
tralian Microferonia known to me.
(Tribe AMBLYSTOMINI)
(Genus AMBLYSTOMUS Erichson)
Erichson 1837, Kiifer Mark Brandenburg 1, 1, p.
59.
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae 5,
p. 922 ( see for synonymy and additional ref-
erences ) .
Andrewes 1939, Ann. Mag. Nat. Hist. (11) 3, p.
130.
Diagnosis. Small Carabidae with most
technical characters of large-headed Har-
palini but with labrum usually unsymmetri-
cally emarginate and scutellar striae in first
(not second) intervals; length usually less
than 5 mm.
Description. None required here.
20 BuUetin Museum of Comparative Zoology, Vol. 137, No. 1
Type species. Acupalpus mauritaniciis
Dejean, of the Mediterranean region (An-
drewesl939).
Generic distribution. Most of the wanner
parts of the Old World, inckiding Aus-
tralia but perhaps not New Guinea.
Notes. In the British Museum are 14
specimens labeled as from Dor(e)y, New
Guinea, some marked as collected by Wal-
lace and all probably from his material.
They include 4-macuIate, 2-maculate and
immaculate individuals, probabh represent-
ing different species. However, these speci-
mens may be mislabeled and may really be
from Celebes or the Moluccas (see Part I
of my "Carabid Beetles of New Guinea,"
p. 331 ). I have received no other specimens
from New Guinea and found none there
myself, although I collected series of the
genus in the Philippines, so that my collect-
ing methods are evidently adequate to ob-
tain it, and Amhhjstomus is usually common
where it occurs at all. I therefore doubt
its occurrence in New Guinea. I list the
genus here, in parentheses, but see no rea-
son to name or discuss the "Dor(e)y"
species individually.
Tribe CHLAENIINI
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae
5, \). 927 (see for earlier references and
synoinmy ).
Callistitac Jeannel 1942, Faune de France, Coleop.
Carabiques, Part 2, p. 961.
Jeannel 1949, Coleop. Carabifjues de la Rei^ion
Malgache, Part 3, p. 776.
Calllstinae Basilewsky 1953, Exploration Pare Na-
tional I'Upemba, Fasc. 10, Carabitlae. p. 119.
A single, well known genus of this tribe
is represented in New (Guinea.
Genus CHLAENIUS Bonelli
iioiiclii bSlO, Observations Eiit. 1, Tab. Syiiopt..
Mem. .Acad. Sci. Tnrin 18, pp. 21-7(S.
Chaudoir 1876, Ann. Mus. Civ. Genoa 8, p. 10
(in m()n()L,'rai)li of "Chleniens").
Sloane 1910, Proc. Linnean Soc. New South \\ ales
35, p. 437 (Anstralian species).
Csiki 1931, Coleop. Cat., Carabidae, Haipalinae 5,
p. 934 (see for additional references).
Andrewes 1941, Ann. Maji. Nat. Hist. (11) 7, p.
307 (with key to Javan species),
jeannel 1942; 1949 (see works cited nnder tribe).
Bell 1960, Misc. Pub. Ent. Soc. America 1, pp.
98, 108 (North American species).
Diasi,nosis. See works cited. I use
Chlaenius in a xery broad sense, as noted
below. In this sense it is the only genus of
the tribe in New Guinea and Australia.
Description. None required here. For
discussion of some characters of the New
Guinean species, see Notes below.
Tyj)c species. Chlaenius nmrii,inatus Rossi
{= vclutinus Duftschmidt), of Em-ope.
Generic distribution. Nearly world-wide.
Tlie genus is most diverse in structure and
most numerous in species in Africa and the
Oriental Region, less diverse and less nu-
merous in Eurasia and America north of the
tropics, and still less in South America and
the Australian Region. This suggests that
the genus has evolved primarily in the
Old World tropics and spread from there.
In the Asiatic-Australian area, scores of
species of CJda-cnius are known in tropical
Asia, about 30 in Java (Andrewes 1941),
but only 12 in New Guinea, and only 10
(including Ilololeius) in Australia. Some of
the species in New Guinea and Australia
are undifferentiated Asiatic forms. Others
are endemic to New Guinea or Australia.
And the endemics difft^r in degree of dis-
tinctness. This suggests continual trickling
of species from Asia toward New Guinea
and Australia rather than concerted move-
ments. The fact that all Chlaenius in New
Guinea and Australia are still winged sug-
gests that their dispersals haxc been rela-
tively recent.
Notes. Vov discussion ol the authoi-, date j
of publication, and type species of Chhicnius !|
see Jeannel 1942, page 963, footnote.
Chlaenius is a huge genus of 700 or SOO
or more known species, and the species are
diverse and can be di\ided into man\- well
characterized groups. Ne\-ertheless, the
genus as a whole seems natural, not poK-
phyletic. Under these circumstances, al-
though the genus can and should be
subdivided, the taxononiie lexcl of tlu- sub-
dixisions should be detcMinined b\- utilit)'
and intelligibilit) . Chlaenius is known to
The Carabid Beetles of New Guinea • Darlington 21
many entomologists who are not specialists species. Segment 3 is not strictly glabrous
in Carabidae, and there seems much to lose in any species; a few minute setules are
and little to gain by splitting it into small aWays visible in fresh, clean specimens at
genera with new and unfamiliar generic 50x or lOOx magnification.
names, many of them unfamiliar even to The palpi (lioth pairs, in both sexes) are
me, a specialist in Carabidae! I shall there- usually slender with apices narrowly trun-
fore use ChJacnius in a very broad sense, cate, but are almost acuminate in guttula,
as a matter of considered policy. I expect and more broadly truncate in flaviguttatus
to discuss this policy in more detail in Part (tenninal segments with apical edges Vs or
IV of my "Carabid Beetles of New Guinea." % segments' length ) .
The ChJocniiis of New Guinea are few and The mentum is toothed, and the tooth is
some of them do not fit well in recognized usually variably emarginate.
subgenera, and no one is likely to be misled The pronotum has a basal hair fringe in
if I treat them simply as species of the all species except daer, ceylanicus, and
great genus Chloenius sensu Into. guttula, which lack it. These 3 species are
Almost every author who has worked apparently not related to each other,
extensively on C/?/rtcn/n.5 has used new char- The pronotum always has a pair of pos-
acters to group the species, but the works terior-lateral and in some cases also median-
of different authors have not been well cor- lateral seta-bearing punctures, but they are
related. Chaudoir used a variety of obvious often hard to distinguish in the general
characters, beginning with extent of ab- punctation. The posterior-lateral punctures
dominal punctation. Sloane noted that are Vo or Ve of the prothoracic length before
presence or absence of a basal pronotal the base in Chlaenius pan, daer, and guttula,
hair fringe and presence or absence of inter- but closer to or at the posterior angles in
ruptions of the outer elytral margins are the other species. Median-lateral punctures
promising taxonomic characters within the (just before middle of prothoracic length)
genus. Jeannel and Basilewsky derived new are present in some (all?) individuals of
group characters from the male genitalia, occidtiis and siccus, but apparently absent
And Bell found additional characters in the in the other species.
labial pit organs of both sexes and in the The elytra have the basal margin entire
chaetotaxy of the valvulae of the female, except in Chlaenius pan. The margin is
The following notes on certain characters obtusely angulate at humeri in daer,
apply only to the New Guinean species of rounded or at most vaguely subangulate in
Chlaenius, unless otherwise indicated. the other species.
Tlie mandibles are short in all Chlaenius The outer elytral margins are interrupted
in New Guinea, and are exceptionally before apex except in Chlaenius pan and
strongly semicircularly arcuate in maculiger. daer, in which the interruption is obsolete.
The clypeus and labrum are truncate or The punctation of the elytral intervals
weakly emarginate except in amplipcnnis, is 2-seriate in Chlaenius pan and daer, but
in which the labrum is deeply emarginate. irregular in the other species, in which it
The antennae have segment 3 much (c. varies from sparse {ceylanicus only) to
V2) longer than segment 4 in Chlaenius pan dense.
and daer, slightly longer in guttula and The inner wings are full and probably fit
amplipennis, and subequal in the other for flight in all Chlaenius in New Guinea
species. Segment 3 is pubescent in guttula and also in Australia, although wang atrophy
(although the pubescence differs in quality has occurred in various African, Asiatic,
from that on the outer segments), more and North American stocks of the genus,
sparsely pubescent or setulose in pan and The punctation of the lower surface of
daer, and still more sparsely so in the other the body is more diverse than some authors
22 BiiUetin Museum of Coinparative Zoology, Vol. 137, No. 1
have realized. Almost the whole lower
surface including the abdomen is punctate
or punctulate and setulose in CJilaenitis
guttula and amplipennis and also in dacr,
although the latter belongs to the circiiin-
datiis group in which Andrewes (1941)
considered the middle of the abdomen
glabrous. C. pan, ceylanicus, and muculiiicr
are more or less intermediate in this char-
acter. The other species have the middle
of the abdomen widely glabrous.
The metepisterna are differently mar-
gined in different Chlocnhis in New Guinea,
but I doubt if this character deserves the
importance Andrewes (1941) gives it.
The tarsi are obviously setulose above
in CJihicnius U,uttula, glabrous or nearly so
in the other species, but minute setules are
usually visible on the upper surface of the
tarsi at 50 X or l(X)x magnification, even in
the "glabrous" species.
The hind tarsi have the 5th segments
always with 2 rows of strong setae below.
The number of setae in each row varies
from about 4 to about 8 in different species.
Males of all New Guinean species of
CJihicnius have each front tarsus with 3
segments dilated (least so in ccylaniciis,
see following Key ) and densely squamulose
below. And i 6 have 1, 9 9 2 setae each
side before apex of last ventral segment,
with extra adventitious setae som(>times
present.
The aedeagus is open above for much
ol its length in most species (especially
widely open in iiuttiila) but relatively long
and closed lor almost half its length in pcni.
I have not studied the chaetotaxy of the
9 valvulae.
Six unrelated species of CJihicnius, de-
rived from groups that normally have pale
markings on the elytra, are losing or have
lost the markings in New Guinea (see Notes
under (hier, guttula, flaviguttafus, himac-
uhiius jiongraczi, maculiger, and iunnifer
malclicri). This suggests a local climatic
or other selective factor favoring dark color
and loss of markings in New Guinea.
In habits, all New Guinean Chhienius
are ground-living. C. daer, ceylanicus,
himaculatus pongraczi, and occultus are
found on river banks; occultus especially
may occur only beside rivers. C. hamifer
malclicri and siccus are commonly found
under cover in comparatively dry places.
C. maculiger is, I think, a rain forest species.
The other species live in more or less damp
places, but I cannot give their habitats
exactly. I took specimens of several species
at light or in floods.
Key to Species of Chlaenius of New Guinea
1. Elytra with outer margins not interrupted;
elytra! intervals each with 1 re^iular row
of punctures on each side; antennae with
3rd segments c. ^-j longer than 4th 2
- Elytra with outer margins interrupted be-
fore apex; elytral intervals irregularly punc-
tate; antennae witli 3rd segments not or
not much longer than 4th 3
2. Very large (c. 25 nun); pronotmn with
basal hair fringe (p. 23) )Hin
- Smaller (c. 12-15 mm); pronotum without
basal hair fringe (p. 24 ) .— dacr
3. Elytral intervals very sparsely punctulate;
c^ front tarsi narrower, with 2nd segments
V-t or V-i longer tlian wide (p. 24)
ceylanicus
- Elytral intervals more closely pimctulate;
S front tarsi wider, 2nd segment c. wide
as long - 4
4. Mandibles very short, semicircularly arcuate
(p. 25) macidi^cr
- Mandibles normal, moderately arcuate ^^^ 5
5. Abdomen plainK' punctulate and pubescent
or setulose at nu'ddlc as well as at sides 6
- Abdomen broadh' smooth and glaliious
(or nearly so) at middle 7
6. Labrum subtruncate or weakly emarginate;
pronotum without basal liair iringe; pos-
teiior-lat(Mal selae c. la of i^rothoracic
length before base; size very small ( c.
8 mm) (p. 25) •guttula
- Labniiii deeply emarginate; pronotum with
basal hair Iringe; posterior-lateral setae
near ( slighlK rounded ) posterior angk'S;
larger ( r. 12 nmi ) {p. 26) --- (iiniiliiunnis
7. Male bout leniora each with a small tooth-
like tub('rel<' Ix'low, near base; 5th seg-
mcnls liiiul tarsi with c. 7 or cS setae each
side below S
- Male front b^nora without tubercles; 5th
segments hind taisi with r. 5 setae each
side below 10
The Carabid Beetles of New Guinea • Darlington 23
8. Pionotum closely and coarsely punctate
and head including front punctulate ( this
combination of characters separates both
sexes of this species from all following
ones, the closest approach being siccus,
see couplet 10) (p. 26) flaviguttatus
— Pronotum with only base coarsely punctate;
head not or only sparsely irregularly
punctulate 9
9. Pronotum with anterior margin entire or
only narrowly interrupted at middle; sides
of prothorax usually sinuate; elytra usually
2-niaculate ( p. 27 ) -.- hiimiculatus pongraczi
— Pronotum with anterior margin obsolete,
indicated only toward sides; sides of pro-
thorax not sinuate; elytra not maculate
( p. 27 ) olthofi
10. Pronotum punctate at base and in narrow
zone along midline but much of disc im-
punctate; posterior-lateral pronotal setae
often c. 1/4(1 of pronotal length before angles
(but variable) (p. 28) occultus
- Pronotum more extensively ( but not always
evenly) punctate; posterior-lateral pronotal
setae almost at basal angles 11
11. Pronotum more sparsely punctate near mid-
dle, with punctiues tending to form ir-
regular longitudinal rows; front exten-
sively but irregularly punctate (p. 28) _^_-
Jimnifcr malcheri
- Pronotum coarsely pimctate, with punc-
tm^es somewhat irregular but less so than in
preceding species; front shining, widely
impunctate ( a few punctines posteriorly
and laterally); (this species characterized
also by coarse punctation of proepisterna
and of elytral striae) (p. 29) siccus
Chlaenius pan n. sp.
Description. Form as in Figure 9, large,
rather slender; black, appendages brownish
piceoiis except c. outer halves of femora
reddish testaceous; rather shining; reticulate
microsculpture fine, faint on front, slightly
more distinct on pronotum and elytra, c.
isodiametric except slightly transverse on
part of pronotum. Head 0.81 and 0.84 width
prothorax; eyes rather abruptly prominent;
antennae with 3rd segments about V.>
longer than 4th segments and plainly but
sparsely setulose; mandibles short, mod-
erately arcuate; mentum with deeply emar-
ginate tooth; clypeus subtruncate; labrum
slightly emarginate; palpi narrowly truncate
at apex in both sexes. Fvothorax quadrate;
width/length 1.11 and 1.09; base/apex 1.16
and 1.11; sides weakly arcuate anteriorly,
slightly converging and very broadly weakly
sinuate posteriorly, each with seta c. Vt> of
length before base, without median-lateral
seta; disc with impressed middle line and
rounded basal impressions, wrinkled-punc-
tate at base, nearly smooth (sparsely punc-
tulate) elsewhere; posterior pronotal hair
fringe present. Elytra long, narrowed to-
ward base; width elytra/prothorax 1.67 and
1.75; margins c. obliterated at base (inside
bases of 4th striae), rounded at humeri, not
interrupted posteriorly; intervals rounded-
subcostate, each with an irregular row of
punctures on each side. Lower surface
partly irregularly punctulate, but much of
abdomen smooth at middle; metepisterna
long, with outer edges raised but not
channeled. Inner wings full. Legs slender;
tarsi not pubescent above; 5th segments
hind tarsi with 4 or 5 strong setae each side
below. Secondary sexual characters normal;
2nd segments S front tarsi c. wide as long
(by measurement); d front femur not
dentate; aedeagus long, slender, closed
above for nearly half its length (Fig. 171).
Measurements: length c. 25-26; width c.
9.1 mm.
Types. Holotype $ (Bishop Mus. ) from
Torricelli Mts., Mokai Village, N-E. N. G.,
750 m, Jan. 1-23, 1959 (W. W. Brandt);
and paratypes as follows. N-E. N. G.: 1
$ , Maprik, Sepik Dist., 19^5 ( Dept. Agr.
Port Moresby). West N. G.: 1 9 (M.C.Z.,
Type No. 31,351), Kota Nika, Res. Hol-
landia, Jan. 9, 1958 (R. T. Simon Thomas);
1 9 , Tanahmerah, Res. Boven Digoel, Feb.
1958 (R.T.Simon Thomas).
Measured specimens. The S holotype and
9 paratype from Kota Nika.
Notes. This new species probably repre-
sents Chlaenius femoratus Dejean of Java,
Sumatra, etc. but is narrower (especially
the jjrothorax) and duller than femoratus
and lacks subapical interruptions of the
elytral margins, which are present though
weak in my 5 specimens of femoratus from
Java. I have 1 9 of a related undescribed
s^Decies from Celebes, which partly fills the
24 Bulletin Museum of Comparative Zoologij, Vol. 137, No. 1
geographic gap between femoratus and
pan.
Chloenius doer n. sp.
Description. Form of Chlacnius of cir-
cumdatus group; slender; greenish l)lack,
head green, elytra sometimes with vestige
of very narrow yellowish margin at apex,
appendages testaceous brown; reticulate
microsculpture alisent or faint on head and
pronotum, deep, fine, isodiametric on
elytra. Head 0.82 and 0.(S2 width prothorax;
eyes large, prominent; antennae with 3rd
segments c. ^2 longer than 4th, setulose;
mandibles moderate; clypeus and labrum
subtruncate; surface of head irregularly, not
densely punctate; mentum with ± emargi-
nate tooth. Prothorax narrow, quadrate-sub-
cordate; width length 1.11 and 1.16; base/
apex 1.04 and 1.09; sides arcuate except
broadly usually strongly sinuate posteriorly;
margins narrow, each with seta c. % of
length before base, without median-lateral
seta; disc irregularly punctate, with fine
middle line, linear baso-lateral impressions
nearer sides than middle but shallower than
usual in the group; posterior pronotal hair
fringe absent. Elytra slightly narrowed an-
teriorly; width elytra/prothorax 1.63 and
1.66; margins entire at base, obtusely angu-
late at humeri, not interrupted pcxsteriorly;
intervals weakly convex, each with a row of
punctures on each side. Lower surface in-
cluding middle of abdomen extensively
punctulate and pubescent; metepisterna
long, w(>akly margined externally. Inner
winii-s full. Lcii^s: tarsi nearly glabrous
above; 5th segments hind tarsi with c.
4 short setae each side below. Secondary
sexual characters normal: 2nd segment male
front tarsi c. Vio longer than wide; male
femora not dentate; aedeagus open abov(>
for much of length. Measuremenls: length
c. 12-15.5; width 4.4-6.0 mm.
Types. Holot>T)e i (M.C.Z., Type No.
31,352) and 2 paratypes from Nad/ab, N-E.
N. G., July 1944 (Darlington); and addi-
tional paratypes as follows. Papua: 1<S,
Kiunga, Fly U., dates in July, Aug. 1957
(W. W. Brandt, Bishop Mus.); 1, Lake
Da\iumbu, Fly R., Aug. 19-30, 1936
(Archbold Exp., A.M.N.H.); 1, Palmer R.
at Black R., July 22-31, 1936 (Archbold
Exp., A.M.N.H.); 1, Kokoda, 1200 ft. (366
m), Aug. 1933 (Cheesman). N-E. N. G.: 2,
Aitape, Aug. 1944 (Darlington); 1, Main R.,
Sepik, Feb. 1965 (R. Hornabrook). West
N. G.: 1, Hollandia, July-Sept. 1944
(Darlington); 1, Tanahmerah, Boven Di-
goel Res., 17 m, April 15, 1955 (L. D.
Brongersma, Leiden Mus.); 1, Idenburg R.,
400 m, July 15-Sept. 15, 193S (J. Olthof,
Leiden Mus.); 1, lebele Camp, Snow Mts.,
2250 m, Sept. 1938 (Toxopeus).
Measured specimens. The i holotype and
1 9 paratype from Kiunga.
Notes. C. daer is the only species of the
Chlaenius circumdatus group in New
Guinea. This group is widely distributed
and common in the warmer part of the Old
World including Australia. The present
new species seems nearest acroxanthus
Chaudoir (which ranges from the south-
eastern corner of Asia to the Moluccas —
Louwerens 1956, Treubia 23, p. 223) but
has baso-lateral pronotal impressions shal-
lower and punctation less coarse. I have
used for comparison a series of acroxantJuis
from Java, collected by Thomas Barbour.
Chlaenius ceylanicus Nietner
Nietner 1856, j. Asiatic Soc. Bengal 25, p. 385.
Csiki 1931, Cok'op. Cat., Carabidat', ilarpalinae
5, p. 932 (HoloUns) (.see for additional refer-
ences ).
niliditlus Dejean (not Sehraiik) 182(i, Species
General Coleop. 2, p. 34 1 .
onuitus Tryon 1890, Second Annual Report
Adnn'nistrator British New (Guinea, Appendix 5,
p. 109 (Poecilus).
Csiki 1931, Coleop. ("at., Carahidae, Ilarpalinae 5,
p. 563 (?Focciloi(lca).
Descri])lion. None recjuired here; see
preceding Key to Sj)ccies for recognition
characters. Note 2nd segments 6 front tarsi
'1 (Ja\;ui specimen) or ':i (Australian speci-
men) longer than wide (l)y measurement);
length c. 11-12.5 mm.
Types. Of ceylanicus. from w(\stern and
southern Cevloii; now in I^erlin U. Zool.
The Carabid Beetles of New Guinea • Darlington 25
Mus. and Stettin Town Mus. (t. Andrewes).
Of nitidulus, from "Indes orientales"; in
Oberthiir Coll., Paris Mus. Of omofus. a
6 from St. Joseph (Angabimga) R. District,
Papua, collected by A. C. English; present
location of type unknown (not seen).
Occurrence in Neic Guinea. Papua: the
type of ornafus; 1, Rouku, Morehead R.,
Apr. 1962 (W. W. Brandt, C.S.I.R.O.).
N-E. N. G.: 5, Kamindibit, Main R., Sepik,
Feb. 1965 (R. Hornabrook), on water
weeds in swamp. West N. G. : 1 9 ,
Garian, Lake Jamoer, Dec. 8, 1954 (L. D.
Brongersma, Leiden Mus.).
Notes. ''Hololius" ceyJonicus ranges from
southern Asia to eastern Australia, and
will probably be found on all the inter-
vening islands, although records are still
incomplete. I have found it in Australia
under cover by backwaters of rivers and in
river floods. Nietner says it flies to light
in Ceylon.
Chlaenius maculiger Casteinau
Castelnau 1867, Notes on Australian Colcop., p. 62.
Chandoir 1876, Ann. Mus. Civ. Genoa 8, p. 67.
Sloane 1910, Proc. Linnean Soc. New South Wales
35, pp. 438, 440.
Csiki 1931, Coleop. Cat., Caraliidae, Harpalinae
5, p. 961.
nigripes Macleay (not Dejean, not Faldermann )
1886, Proc. Linnean Soc. New South Wales
(2) l,p. 140.
biroi Csiki 1931, Coleop. Cat., Carabidae, Har-
palinae 5, p. 948.
Description (for recognition only). Me-
dium sized, depressed; dark, dark-legged,
typically 2-maculate but spots sometimes
lost; unique in genus in New Guinea in
mandibles very short, semicircular; length c.
12-14 mm.
Types. Of maculiger, from Rockhamp-
ton, Australia; probably in Genoa Mus. ( I
did not find it at Melbourne in 1957). Of
nigripes, from Fly R., Papua (implied);
may now be in Macleay Mus., Sydney (not
seen). Of hiroi, as for nigripes (the name
hiroi was proposed to replace nigripes
Macleay, which is preoccupied).
Occurrence in New Guinea. Widely dis-
tributed and common: 127 specimens from
localities including Dobodura and Wau
in all 3 political divisions of New Guinea;
most at low altitudes but reaching at least
1300 and 1500 m at Wau and on the
Bismarck Rge.
Notes. Sloane (1910) has established the
identity of nigripes Macleay (hiroi Csiki)
with maculiger Castelnau.
Outside New Guinea, this species is
known from Australia and New Britain
(Cape Gloucester, Jan.-Feb. 1944, Darling-
ton). It is apparently related to and
probably derived from the same Oriental
stock as Chlaenius tetragonoderus Chaudoir,
which is widely distributed farther west in
the Malay Archipelago, to the mainland
of Asia. C. tetragonoderus batjanicus
Louwerens (1956, Treubia 23, p. 234) of
the northern Moluccas, which varies in
color of legs, may be a transitional form.
An apparently undescribed species of the
group occurs in the Solomons.
The yellow subapical elytral spots are
individually variable in specimens from
New Guinea and are absent or nearly
absent in some individuals. The variation
in spotting apparently occurs throughout
New Guinea.
This is, I think, a rain forest species that
may occur in ordinary leaf litter rather
than in specially wet places, but I have
taken too few specimens to be sure.
Chlaenius gutfula Chaudoir
Chaudoir 1856, Bull. Soc. Nat. Moscow 29, Part
2, p. 216.
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae
5, p. 957 ( see for additional references ) .
Andrewes 1941, Ann. Mag. Nat. Hist. (11) 7, p.
310 (in key).
Louwerens 1953, Verhandlungen Naturforschenden
Gesellschaft Basel 64, p. 313.
csikii Jedlicka 1951, Ann. Mus. National Hungary
1, p. 136 (new synonymy).
astrolabe nsis Jedlicka 1951, Ann. Mus. National
Hungary 1, p. 136 (new synonymy).
immaculata Louwerens 1962, Tijdschrift voor Ent.
105, p. 145 (new synonymy).
Description ( for recognition only ) . Very
small; dull dark bluish, with or without a
small pale spot on suture near apex (see
26
BuUctin Museum of Comparaiive Zoolop^ij, Vol. 137, No. 1
following Notes); see also preceding Key
to Species; length c. 8 mm.
Types. Of gtittiila, from Hongkong; in
Oberthiir Coll., Paris Mus. Of r.s//\// and
astrolahensis, both from Stephansort, Astro-
labe Bay, N-E. N. G.; in Hungarian
National Mus. Of immacuJata, from Ani-
hoina; in Louwerens Coll. (Types not
seen. )
Occurrence in New Guinea. Papua: 7,
Dobodura, Mar.-July 1944 (Darlington);
1, Bisianumu, near Sogeri, 500 m. Mar. 15-
20, 1955 (E. O. Wilson, M.C.Z.), in rain
forest; 1, Brown R., May 23, 1956 (E. J.
Ford, Jr., Bishop Mus.). N-E. N. G.: 1,
Bulolo, 731 m, Aug. 26, 1956 (E. J. Ford,
Jr., Bishop Mus.); 1, Finschhafen (L.
Wagner, M.C.Z.). West N. G.: 3, Hol-
landia, Jan., Apr., Mav 1945 (B. Malkin,
U.S.N.M.); 1, Kota Nika, Res. Hollandia,
Nov. 29, 1957 (R. T. Simon Thomas,
Louwerens Coll.); 1, Maffin Bay, Jan. 1,
1945 (E. S. Ross, California Acad.).
Notes. This species is known from south-
ern Asia, Sumatra, Java, Bali, Celebes,
Timor (Louwerens 1953), the Philippines,
New Guinea, and New Britain (Cape
Gloucester, Darlington, M.C.Z. ). It often
flies to light.
Although most specimens from New
Guinea have a variable ( sometimes minute )
vestige of a subapical sutural pale spot,
several ( not all ) of those from Dobodura
are unspotted.
The characters given by Jedlicka to dis-
tinguish csikii from iiuttuJa seem to me to
be individual rather than spcx'ific, and
"aberration" astrolalmnsis Jedlicka and "var."
inimaculata Louwerens are (I think) un-
necessary names lor imspotted individuals.
Chlaenius amplipennis Chaudoir
Cliaiuloii 1876, Ann. Mus. Civ. Ocnoa 8, p. 252.
Csiki 1931, Coleop. Cat., Carabidac, Ilarpalinac
5, p. 946.
Andrewi's 1911, Ann. Mas. Nat. Hist. (II) 7,
p. 310.
Description (for recognition only). Me-
dium small; dark, dull; uiu(iue among
Chlaenius of New Guinea in labrum deeply
emarginate; length c. 12 mm.
Type. A $ from Java; in Brussels Mus.
(not seen).
Occurrence in New Guinea. N-E. N. G.:
1 6, Bulolo, 2000 ft. (610 m), Mar.-July
1937 (George Rio, Chicago Mus.); 1 9,
Main R., Sepik, Feb. 1965 (R. Hornabrook).
Notes. Chlaenius amplipennis apparently
ranges from Sumatra and Java to the
Philippines, New Guinea, and the Solo-
mons ( Guadalcanal Is., 1944, L. N. Jarcho,
M.C.Z. ). It varies geographically and some
of the geographic fonns may be recogniz-
able subspecies, but I do not have enough
material to decide about this.
Chlaenius flaviguftafus Macleay
Macleay 1825, Annulosa Javanica, p. 14.
Chaiuloir 1876, Ann. Mus. Ci\\ Cenoa 8, p. 52.
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae
5, p. 955 ( see for synonymy and additional
references ) .
Andrewes 1941, Ann. Mati. Nat. Hist. (11) 7,
p. 307 (in key).
Louwerens 1953, Verhandlungen Naturforschenden
Cesellschaft Basel 64, p. 311.
fliittatiis Eschscholtz 1833, Zoologisehen Atlas 5,
p. 26, pi. 25, fig. 8.
iiuniactilipcnuis Jedlieka 1951, Ann. Mus. National
Hungary 1, p. 134 (new synonynn' ).
Description (for recognition onU). Me-
dium sized, rather slender; dull, dark, elytra
2-maculate or immaculate, legs pale usually
with dark knees; most ol upper surface
closely conspicuously punctate; palpi with
apical segments truncate, apices -i or ^-i
wide as lengtli of segment; see also Key to
Species; length c. 11-14.5 mm.
Types. Of flaviL!,uttafus, from Java; in
Briti.sh Mus. (seen). Of iiuttatus. Ma-
nil (l)a; Moscow U. Zool. Mus. (not seen).
Of it)nn(iculi))cnnis. New (hiinea; in Jed-
licka ('oil. (not seen).
Occurrence in Neu Ciuinca. Widely
distributed on Ne\\- Guinea ( including
Doboduia and Wan), and reaching I^iak
and (in tUv .Vdmiralties ) Manns Is.: 283
specimens, most at low altitudes but reach-
ing at least 1300 and 1500 m in places in
the mountains.
The Carabid Beetles of New Guinea • DaiUngton 27
Notes. This common Chlaenius is known
from Sumatra, Java, etc. to New Guinea
and Australia, east to the Philippines,
New Britain, New Ireland, Solomons,
New Hebrides, Fiji, Samoa, and New
Caledonia.
Markings vary indwiduaUij in the series
from New Guinea. Each elytron may have
a conspicuous irregular subapical pale mark,
or fragments of such a mark, or no mark at
all, and the variation occurs in all parts of
New Guinea from which series have been
seen. Unmarked individuals are "aberra-
tion" immacu]ipcnms]ed\\ckA, which I think
is not worth distinguishing.
This species occurs in a variety of wet
places, often in more or less open country.
Chlaenius bimaculafus pongraczi Jedlicka
Jedlicka 1951, Ann. Mus. National Hungary 1,
p. 136.
Description. Generally similar to typical
himaciilatus Dejean in technical characters
( see preceding Key and also Andrewes' key
to Javan Chlaenius, 1941, Ann. Mag. Nat.
Hist. (11) 7, p. 307), but differing some-
what in color and especially in punctation.
Color bluish black, legs testaceous (not
bicolored), antennae and mouthparts red-
dish brown. Punctation of head and pro-
notum reduced but variable: head with or
almost without punctulation (most distinct
posteriorly); pronotum coarsely punctate
only basally, extensively smooth or in part
finely punctulate elsewhere; length c. 12-
14 mm.
Type. From New Guinea; in Hungarian
National Mus. (not seen).
Occurrence in New Guinea. Papua: 9,
Dobodura, Mar.-July 1944 (Darlington);
7, Mt. Lamington, 1300-1500 ft. (c. 400-
450 m), (C. T. McNamara, S. Australian
Mus.); 2, Kokoda, 1200 ft. (366 m). May
& Aug. 1933 (Cheesman); 1, Daradae, near
Javarere, Musgrove R., Oct. 4, 1958
(Gressitt). N-E. N. G.: 3, Sattelberg
(British Mus.); 1, Wareo, Finschhafen
(Rev. L. Wagner, S. Australian Mus.); 1,
Gewak, Salawaket Rge., 1530 m, Sept. 7,
1956 (E. J. Ford, Jr., Bishop Mus.); 1,
Sepik, Maprik area, 160 m, Aug. 26, 1957
(D. Elmo Hardy, Bishop Mus.). West
N. G.: 1, Hollandia, Jan. 1933 (A.M.N.H.);
1, VVaris S. of Hollandia, 450-500 m, July
24-31, 1959 (T. C. Maa, Bishop Mus.); 2,
Ifar, Cyclops Mts., 300-500 m, June 23-25,
Sept. 9, 1962 (Sedlacek); 1, Guega, W. of
Swart Vy., 1200 m, Nov. 14, 1958 (Gressitt);
1, Bodem, 11 km SE. of Oerbefareh, 100 m,
July 7-17, 1959 (T. C. Maa, Bishop Mus.).
Notes. Chlaenius himaculatus Dejean (or
the group of closely related species that
goes under this name) ranges from SE.
Asia to the Philippines and New Guinea
(not Australia). I have ample comparative
material from a number of localities from
SE. Asia to Amboina. Geographic variation
is obvious. The New Guinean form varies
also individually in marking: most individ-
uals have a conspicuous pale spot before
apex of each elytron, but the spot varies
in size and is almost absent in 2 of the
Sattelberg specimens.
My Dobodura specimens were taken in a
grassy bank beside a small river.
Chlaenius olfhofi n. sp.
Description. Form (Fig. 10) of Chlaenius
himaculatus Dejean, slender; head and
pronotum shining green or greenish black;
elytra duller, purplish black, with very fine
c. isodiametric microsculpture; appendages
rufous. Head 0.72 and 0.76 \\'idth prothorax;
eyes large, genae short; antennae with 3rd
segments c. equal to 4th and sparsely
setulose; mandibles average; clypeus and
labrum subtruncate or weakly emarginate;
front with c. punctiform anterior impres-
sions, otherwise almost impunctate (a few
punctules posteriorly); mentum with c.
entire tooth; palpi narrowly truncate at
apex. Prothorax subquadrate, widest at or
slightly behind middle; width/length 1.16
and 1.13; base/apex 1.17 and 1.10; apex not
margined except vaguely at sides; sides
broadly rounded, not or at most faintly
sinuate before obtuse but well defined,
slightly blunted posterior angles; posterior-
28 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
lateral setae c. Vin of prothoracic length be-
fore base, median-lateral setae absent; disc
with impressed middle line, sublinear
baso-lateral impressions; surface extensively
smooth but with a few punctmes mosth'
near base, sides, and along middle; posterior
pronotal hair fringe present. Elytra: width
elytra prothorax 1.44 and 1.45; margins
entire at base, rounded at humeri, with
subapical interruptions; striae moderately
impressed, vaguely punctulate; intervals
slightly convex, moderately closely punctate.
Lower sui'facc shining; proepisterna almost
impunctate; some punctures on metepisterna
including epimera and on base of abdomen
at sides, but much of abdomen smooth or
nearly so; metepisterna long, margined ex-
ternally, margin obsolete anteriorly. Inner
irin^.s full. Lci^s: tarsi c. glabrous above;
5th segments hind tarsi with c. 7 setae each
side below. Secondary .sexual characters:
i front tarsi with 2nd segments c. wide as
long; <^ front tibiae with small tooth below
near base; c^ with usually 1, 9 2 or 3
( unsymmetric in the single 9 ) setae each
sidc> last ventral segment. Aedeagus slender,
open above for much of length. Mea.siire-
ments: length c. 13-14; width c. 4.5-5.0
mm.
Types. Holotype $ (Leiden Mus.) and
i i paratype (M.C.Z., Type No. 31,353)
from Bcrrihard Camp, West N. (i., 50 m,
July-Sept. 1938 (J. Olthof); 1 9 paratype,
same locality, Apr. 12, 1939 (Toxopeus);
1 i paratype, Oro Bay, Pa|»ua, JuK' 12,
1944 (A. II. Mallery, Bishop Mus.). '
Measured specimens. The -' holotype and
9 paratype.
Notes. This seems to be a distinct species
of the himaculatus group, occurring within
the geographic range of ])iinaculalus sub-
species pon^raczi. C. olthof i may be a prod-
uct of an early invasion of a hinuuulatus-
like stock, pon<^raczi of a later one.
Chlaenius occulfus Sloane
Sloane 1907, Dciitsclie Kiit. Zcitscln ill (ni 1!)()7.
p. 467.
Descrij)lion (for recognition onlv). A
medium-sized Chlaenius with subcordate
prothorax; blue-black, sometimes in part
greenish, legs reddish testaceous, antennae
and mouthparts reddish brown; rather
shining, reticulate microsculpture absent or
nearly so on head and pronotum, visible on
elytra especially of female, fine, irregularly
isodiametric; see also Key to Species of
Chlaenius of New Guinea; length c. 14-
17 mm.
Type. From Herbertshohe, New Britain,
"returned to Dr. Horn for Bennigsen's col-
lection" (not seen).
Occurrence in New Guinea. Papua: 7,
Dobodura, Mar.-July 1944 (Darlington);
4, Kokoda, 1200 ft. (366 m), Mav, Aug.
1933 (Cheesman); 1, Laloki, 1909 (F. Muir,
Hawaiian Sugar Planters Association ) ; 5,
Mt. Lamington, 1300-1500 ft. {c. 400-150
m) (C. T. McNamara, S. Australian Mus.);
1, Peria Ck., Kwagira R., 50 m, "No. 7,"
Aug. 14-Sept. 6, 1953 (Geoffrev M. Tate,
A.M.N.H.). N-E. N. G.: 30, vie. Nadzab,
Julv 1944 (Darlington); 1, Busu R., E. of
Lae, 100 m, Sept.'" 14, 1955 (Gressitt); 2,
Wan, 1100, 1200 m, Oct. 30, 1961, Jub-
28-29, 1963 (Sedlacek). West N. G.: 5.
Hollandia, Jan., Apr., Mav 1945 ( B. Malkin,
U.S.N.M.); 1, Humboldt Bay Dist., 1937
(W. Stiiber, British Mus.); 1, Tanahmerah.
Res. Boven Digoel, Apr. 24, 1957 (R. T.
Simon Thomas, Leiden Mus.).
Notes. I have identified this species from
Sloanes description: the size, cordate pro-
thorax, and round(xl humeral margins are
(together) diagnostic in th(^ region in (pies-
tion; other dc^tails agree well enough; and
I have seen a specimen from New Britain
(near Habaul, l''eb. 1929, Pemberton col-
lector, in Goll. Ilawaiiaii Sugar Planters
Association).
This species occurs in New Britain ( [hv
t\iK\ and tlu> specimen Iroin near iiabaul
reierred to abox'c ) and the Soloiiioim
(Guadalcanal; Bougainville) as wtll as
eastern and central New (Guinea. I ha\e
been unable to determine its relationship to
other si)ecies ol (Chlaenius. It is h)un(l
under stones on the banks ol lixcrs.
The Carabid Beetles of New Guinea • Darlington
29
Chlaenius hamifer malcheri Van Emden
\'an Emden 1937, Stettiner Ent. Zeitung 98, pp.
35, 37.
Description (for recognition only). Me-
dium small, moderately broad; usually
very dark with or without slight metallic
tinge, usually without spots but latter some-
times partly developed (see Notes below);
see also Key to Species; length c. 11-12 mm.
Type. From Pauru, New Georgia, Solo-
mon Islands (Fr. Malcher); in Van Emden
Coll., British Mus. (seen).
Occurrence in New Guinea. Papua: 13,
Dobodura, Mar.-July 1944 (Darlington);
1, Oro Bay, July 12, 1944 (A. H. Mallery,
Bishop Mus.); 1, Kokoda, 1200 ft. (366 m),
Aug. 1933 (Cheesman); 1, Port Moresby,
Konedobu, Apr. 20, 1958 (J. J. H. Szent-
Ivany, Dept. Agr. Port Moresby), at light;
1, Popondetta, Aug. 11, 1962 (A. Catley,
Dept. Agr. Port Moresby), at light; 1, Mt.
Lamington, 1300-1500 ft. (c. 400-450 m)
(C. T. McNamara, S. Australian Mus.); 2,
Rouku, Morehead R., Apr. 1962 (W. W.
Brandt, C.S.I.R.O.); 1, Rossel Is. (S.E.
Papua), Oct. 1963 (W. W. Brandt, C.S.I.
R.O.). N-E. N. G.: 2, "No. 2, Oomsis," 22
mi. W. of Lae on Lae-Bulolo Road, 100 m,
Apr. 26, 1959 (L. J. Brass, A.M.N.H.); 2,
Wau, Morobe Dist., 1200 m, Dec. 6, 1961,
Sept. 15-30, 1962 (Sedlacek); 1, Bulolo "G.
T.," (Sedlaceks); 1, 16 km W. of Mumeng,
3000-5000 m, May 1962 (Sedlacek); 1,
Okapa, July 12, 1964 (R. Hornabrook); 1,
lower Busu R., Huon Pen., May 12, 1955
(E. O. Wilson, M.C.Z.). West N. G.: 1,
Hollandia, May 1945 (B. Malkin, U.S.
N.M.); 1, Maffin Bay, June 1944 (E. S.
Ross, California Acad.); 1, Wissel Lakes,
Tage Lake, 1760 m, Aug. 4, 1955 (Gressitt);
1, Wissel Lakes, Enarotadi, 1900-2000 m,
July 2-11, 1962 (N. Wilson, Bishop Mus.).
Notes. This species belongs to a difficult
group of Chlaenius that extends from S.
Asia to NE. AustraHa. The group includes
hamafus Dejean as well as hamifer Chaudoir.
I am not sure whether these two species
really are different, or with which of them
malcheri should go. My treatment of it as
a geographic form of hamifer is tentative.
The range of hamifer is from S. Asia to
Australia.
Apparently only one form of the hamifer-
hamatus group occurs in New Guinea. It
is very dark and usually unmarked, but 2
examples from Dobodura show the posterior
part of a pale "comma" on the apex of each
elytron, and the 1 specimen from Port
Moresby, the 2 from Rouku, and the 1 from
Hollandia have the "commas" complete but
narrow. Chlaenius insulanus Louwerens
(1956, Treubia, 23, p. 234) of the northern
Moluccas is another dark, unmarked form
of the hamifer-hamatus group, but is smaller
and narrower than malcheri.
C. h. malcheri occurs under cover often
in somewhat drier places than most other
Chlaenius except the following (siccus).
Chlaenius siccus n. sp.
Description. Form c. average in genus;
rather shining black, sometimes with slight
greenish or bluish reflections, appendages
rufous; reticulate microsculpture absent on
head and pronotum, fine and c. isodiametric
on elytra. Head 0.68 and 0.67 width pro-
thorax; eyes large, genae short; antennae
with 3rd segments c. equal 4th and scarcely
setulose; mandibles average, rather short,
moderately arcuate; labrum and clypeus
subtruncate; front almost smooth at middle,
punctate at sides and posteriorly, with
slight frontal impressions; mentum with
blunt usually vaguely emarginate tooth;
palpi slender, narrowly subtruncate at apex.
Prothorax subquadrate but rather strongly
narrowed anteriorly; width length 1.28 and
1.28; base/apex 1.33 and 1.31; sides weakly
arcuate for most of length, c. straight and
somewhat converging posteriorly; posterior
angles obtuse, narrowly rounded; margins
narrow anteriorly, wider posteriorly, each
with posterior-lateral setae just before base
and ( at least in some individuals ) median-
lateral setae just before middle; disc ir-
regularly longitudinally impressed each side
30
Bulletin Museum of Comparative Zoology, Vol. 137. No. 1
c. midway between middle and side, with
whole surface rather closely but somewhat
irregularly, coarsely punctate; posterior pro-
notal hair fringe present. Elytra not nar-
rowed anteriorly; width elytra prothorax
1.29 and 1.36; margins entire at base,
arcuate at humeri, interrupted subapically;
striae rather coarsely punctate, intervals
slightly convex, irregularly punctulate.
Lower swface: proepisterna coarsely punc-
tate at least in part, mesepisterna partly
punctate or almost impunctate, sides of
metasterna punctate, abdomen punctulate
at sides and across base but extensively
smooth or nearly so at middle; metepisterna
long, strongly margined ( grooved ) exter-
nally. Inner uin^s full. Le<!,s without obvi-
ous unusual characters; tarsi c. glabrous
above; 5th segments hind tarsi with a. 4
setae each side below. Secondary sexual
characters normal; i front tarsi dilated, 2nd
segment at least as wide as long; 6 front
tibiae not toothed; c^ with 1, 9 2 setae each
side last ventral segment. Measurements:
length e. 11. .5-13.5; width 4.1-4.8 mm.
Types. Holotype S (M.C.Z., Type No.
31,354) and 14 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
additional paratypes as follows. Papua: 2,
Mt. Lamington,' 1300-1500 ft. (c. 400-450
m) (C. T. McNamara, S. Australian Mus.).
N-E. N. G.: 1, Aitape, Aug.-Sept. 1936
(Cheesman); 4, Swart Vv., Karubaka,
1450, 1500, 1550 m, Sept. s', 16, 22, 1958
(Gressitt), some taken in light trap; 8,
Wan. Morobe Dist., 1200 m, dates in Jan.,
Feb., Mar., Aug., 1962-1963 (Sedlacek);
1, Sum-Sum, near Bulolo, Morobe Dist.,
Feb. 7-11, 1966 (Rhonda M. Stevens, Dept.
Agr. Port Moresby); 7, Minj, W. High-
lands, 5200 ft. (c. 1600 m). Mar. 25, May
20, 1960 (J. H. Barrett, Dept. Agr. Port
Moresby). West N. G.: 1, Ilollandia, May
1945 (B. Malkin, U.S.N.M.); 1, Kota Nika,
Res. Ilollandia, Feb. 23, 1956 (R. T. Simon
Thomas, Louwerens Coll.); 4, Ifar, Cyclops
Mts., 300-500 m, June 23-25, 1962 (Sed-
lacek); 1, same locality, 400-800 m, Sept. 9,
1962 (Sedlacek); 2, 'X;. den Iloed. Tfar,"
Dec. 1957 (Louwerens Coll.); 1, Kebar Vy.,
W. of Manokwari, Vogelkop, 550 m, Jan. 4-
31, 1962 (S. & L. Quate, Bishop Mus.).
Measured s))ecimcns. The i holotype and
1 9 paratype from Dobodura.
Notes. In Andrewes' ( 1941, see reference
under genus) key to the Chlaenius of Java,
this runs to leucops Wiedemann, of which
I have specimens from India, Java, and
Luzon, but the present new species is nar-
rower, paler-legged, and more shining than
leucops, with somewhat different sculpture:
e.g., the punctation of the pronotum is
coarser and more irregular than in leucops.
C. siccus is closer to, and may prove to be a
geographic representative of, ophonoides
Fairmaire of Australia ( recorded also from
New Caledonia and New Hebrides ) . How-
ever, siccus is slightly smaller and much
darker than oplionoides, being black with-
out or with only faint metallic tinge while
ophonoides is always plainly greenish black.
In habits, this species resembles the pre-
ceding one (Jiamifcr mah'lieri) and often
occurs with it, under co\er on compara-
tively dry ground in more or less open
places.
Tribe OODINI
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae 5,
p. 1000 (see for synonyiiiN- and additional reter-
enees).
Ooclitac Auet. including Jeannel 1949, Coleop.
Carabicjues de la Region Malgaelu*. Part 3, p.
828.
The members ol this tribe are o\al,
Amara- or even dytiscid-like, and are black
or metallic, usually unmarked. Tlu'ir generic
classification is unsatisfactory (sec^ below.
and see Notes under Anatrichis and Oodes
hu'vissimus). However, the 13 species of
the tribe known from New Cuin(>a obvi-
oush' inc-huU' no striking ciKk'niic genera
and, although diNcrse, (lu'\ are less so than
the oodinc\s ol tlu' Oiiental lu'gion or Aus-
tralia. The Oriental SysfoU)cranius, Ilolco-
coleus, Sinu)us (see under Oodes hievis-
sinuis), etc., and th(> Australian Co])tocarpus
do not reach New (hiinea.
The Carabid Beetles of New Guinea • Darlington 31
The presence or absence of a basal elytral
margin is a useful key character in this
tribe but must be determined with care.
The basal margin is a fine, sharph' marked
line, impressed or formed by a slight eleva-
tion of the elytral surface in front of the
margin, and usually ending inwardly op-
posite the bases of the 3rd striae or 3rd
intervals. It is distinct from the basal de-
pression of the elytra that fits under the
base of the pronotum. It is best examined
under diffused light, for a sharply focused
spotlight makes reflections that simulate a
margin where none is present.
Presence or absence of a small seta-bear-
ing puncture on the posterior edge of the
pronotum on each side near the basal angle
is another useful key character that must
be determined \\'ith care. The setae are
sometimes very small and weak and easily
broken off. The punctures may then be
hard to detect even in clean specimens and
undetectable in dirty ones. Sometimes these
setae and punctures vary within single
species (see Oodes siamensis).
Clean specimens of Anotrichis pusilla,
Oodes exiguus, and O. piceus can be seen
to have a small anterior puncture over each
eye, with or without a small, weak seta.
These punctures are lacking in all other
New Guinean Oodini. This suggests an
actual relationship among the 3 species
named, which is supported by the arrange-
ment of labral setae and by a similarity
of body form, and this in turn suggests
that the conventional distinction between
Anatrichis and Oodes is unnatural. How-
ever, I cannot recharacterize these 2 widely
distributed genera on the basis of the few
I species that occur in New Guinea.
Most oodines are ac[uatic or subaquatic,
living in vegetation or among dead leaves
in or close to standing water, but O.
laevlssimus Chaudoir and probably also the
I 2 related species described below (i.e., the
laevissimus group) live in leaf litter on the
floor of rain forest. This is the habitat of
some Copiocarpus in Australia and of cer-
tain other oodines elsewhere. State of wings
is correlated with habitat. The wings are
fully developed in all known New Guinean
Oodini except the Oodes laevissimus group,
in which the wings are apparently atrophy-
ing as the group leaves subaquatic habitats
and invades terrestrial leaf litter. At the
same time the group is apparently begin-
ning to evolve local flightless species in
different places in New Guinea.
Key to Genera of Oodini of New Guinea
1. Size .small (c. 5 mm); clypeus without seta-
lieaiing punctures; $ front tarsi only slightly
dilated (p. 31) _.__ Anatriclii.s
- Size usually larger; // size small, clypeus
with seta-bearing puncture on each side and
c^ front tarsi wider ( p. .32 ) Oodes
Genus ANATRICHIS Leconte
Leconte 1853, Trans. American Philosophical Soc.
10, p. 391.
Chaudoir 1882, Ann. Soc. Ent. France (6) 2, p.
318.
Sloane 1910, Proc. Linnean Soc. New South Wales
35, pp. 442, 443 (the Australian species).
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae 5,
p. 1003 ( see for synonymy and additional refer-
ences ) .
Diagnosis. Very small Oodini; clypeus
and posterior margin of pronotum without
seta-bearing punctures; labrum with clump
of 4 setae at middle and 1 separate seta each
side; 6 front tarsi typically only slightly
dilated, but variable (see Notes below).
Description. None required here.
Type species. Oodes minutus Dejean, of
North America.
Generic distribution. India and Burma
to Australia; tropical and warm temperate
America.
Notes. The characters and limits of this
genus are doubtful, as suggested in discus-
sion under the tribe (above).
Authorities disagree about the c^ front
tarsi of Anatrichis. Leconte says 4 segments
are slightly dilated and spongiose (with
dense squamae) below. Chaudoir says 3
segments are thus modified. And Sloane
says only 2 segments have squamae. In fact,
different species differ in this respect, and
minor variations of S tarsi may even occur
32 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
within single species (see Notes under A.
pimUa, below ) .
Anafrichis pus'illa Sloane
Sloane 1910, Pioc. Linnean Soc. New South Wales
35, p. 443.
Description ( diagnostic characters only ) .
Small, narrow; pronotum usually with an
almost punctiform impression each side
near base, but these impressions variable
and sometimes almost absent; elytra 7-
striate, striae punctulate; i tarsi slightly di-
lated, with 2 or 3 segments squamulose be-
low ( see following Notes ) ; other characters
given in preceding Key to Genera; length c.
5 mm.
Types. Described from 2 specimens
taken by Sloane near Kuranda, North
Queensland, Australia, June 1906. I here
designate as lectotype the single specimen
now in the Sloane Collection at Canberra.
It is labeled "Kuranda, Q., T.G.S., 6.06" and
"Anatrichis pusilla SI., Id. by T. G. Sloane '
( seen ) .
Occurrence in New Guinea. Papua: 5,
Dobodura, Mar.-July 1944 ( Darlington ) ; 9,
Oro Bay, Dec. 1943-Jan. 1944 (Darling-
ton); 3, Lake Daviumbu, Flv R-, Sept.
11-20 and 21-30, 1936 (Archbold Exp..
A.M.N.H.); 1, Modewa, Modewa Bay, 0-50
m, "No. 17," Dec. 14, 1956 (L. J. Brass,
Fifth Archbold Exp., A.M.N.H.). N-E.
N. G.: 1, Aitape, Aug. 1944 (Darlington).
West N. G.: 24, Ilollandia, July-Sept. 1944
(Darlington); 2, Sarmi, W. of Ilollandia,
July 20-23, 1959 (T. C. Maa, Bishop Mus.);
3, Maffin Bay, Aug. 1944 (Darlington).
Notes. I have a scries of ))UsilJa from
North Queensland, Austraha: from Cairns
(near the type locality), south to Cardwell,
and north to Silver Plains halfway up the
C^ape York peninsula. Specimens from New
(iuinea match Australian ()n(\s well.
A. ptisilla is similar to and may represent
Anatrichis indica (^haudoir of India, and I
ha\'e a related species Iroin Le\ te in the
Philil)pines.
I'hr narrowly dilated ,^ front tarsi seem
to have either 2 or 3 segments squamulose.
I cannot determine whether this is primarily
individual variation, or whether the squamae
are worn off the 3rd segments in some
individuals, or whether the squamae are
sometimes pressed against the soles of the
3rd segments and therefore almost un-
detectable.
Genus OODESBonelli
Bonelli 1810, Observations Ent. 1, table synoptique,
Mem. Acad. Sci. Turin 18, pp. 21-78.
Sloane 1910, Proc. Linnean Soc. New Soutli Wales
35, p. 442 (Australian species).
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae 5,
p. 1006 (see for synonymy and additional refer-
ences ) .
Andrewes 1940, Proc. R. Ent. Soc. London (B)
9, pp. 203 ff. (key to species of India, Burma,
etc. ) .
Jeannel 1942, Faune de France, Coleop. Carabitiucs,
Part 2, p. 980.
Diaii,nosis. No satisfactory one available.
For practical purposes all Ne\^' Guinean
members of the tribe except Anatrichis
pusilla are assigned to Oodes.
Description. None required here.
Tyjx' species. Carahus liclopioidcs Fab-
ricius, of Europe.
Generic distribution. Most of the warmer
parts of the world, but few or none in
South America.
Notes. For comments on classification,
habitats, and state of \\ings see tribe Oodini.
above.
Kiiv TO Species of Oonr.s of New Cuixea
\. Labruni with compact clump of 4 (or
fewer) .setae at nn'ddle and 1 separate seta
each side 2
- Lal)rnni with 6 or 4 separate setae (if 4,
2 adcbtional uu'nutc setae usually present
close to.uethi'r near middle) 3
2. EKtra with 7th striae almost oliliterateil;
length c. 8 mm (p. 33) ))iccu\
- EKtra with 7th striae well impressed;
length r. .") mm {p. 33) cxii^tdis
3. CKpeus without seta-bcariiisi; punctures 4
- Chi^eus with seta-bearing pmnlure each
side - 7
4. Prothorax without basal setae or punctmcs
(p. 33) - nil
Prolliorax witJi seta or small puncture on or
near posterior edue eat'h side ( hicvi.ssitiius
group) 5
The Carabid Beetles of New Guinea
Darlington
33
5. Elytra with hasal margin sharply defined
( see under tribe Oodini ) ; inner wings
full or nearly so (p. 34) laevissimti.s
- Elytra with basal margin obsolete; inner
wings vestigial 6
6. EUtral striae distinctly impressed; pos-
terior pronotal punctures on dorsal surface
at base (West N. G. ) (p. 34) rossi
- Elytral striae reduced to fine superficial
lines; posterior pronotal punctures on basal
edge of pronotum ( N-E. N. G. ) ( p. 35 ) _„_
wiboni
7. Clypeus margined anteriorly, with setae
almost in the angles (p. 36)
siamcnsis vulsus
- Clypeus not margined, with setae behind
the angles 8
8. Prothorax with seta or small puncture each
side on posterior edge 9
- Prothorax without such setae or punctures;
(form stout, convex; c$ luiddle tibiae bent
near base; length c. 13.5-15.0 mm) (p.
36 ) denisonensis
9. Elytra with basal margin sharply defined
(see under tribe Oodini) (p. 36) siccus
- Elytra with basal margin obsolete 10
10. Metepisterna sparsely, vaguely, or not
punctate (p. 37) par
- Metepisterna closely punctate 11
11. Form average, prothoracic width/length c.
1.65 (p. 37) crihristernis
- Form slender, prothoracic width/length 1.44-
1.49 (p. 38) longior
Oodes piceus Nietner
Nietner 1856, J. Asiatic Soc. Bengal 25, p. 526.
Andrewes 19.30, Cat. Indian Insects, Part 18,
Carabidae, p. 238.
1940, Proc. R. Ent. Soc. London (B) 9,
p. 205 (in key).
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae 5,
p. 1010.
Description (for recognition only). A
narrowly oval, convex Oodes with elytra 6-
striate (7th striae absent or faint) and
other technical characters given by An-
drewes (1940); length c. 8 mm. See also
preceding Key and following Notes.
Type. From Colombo, Ceylon; in Stettin
Mus. (not seen).
Occurrence in New Guinea. Papua: 1,
Oro Bav, Dec. 1943-Jan. 1944 (Darlington).
West N. G.: 1, Hollandia, 250 ft., May
1945 (H. Hoogstraal, Chicago Mus.), in
rain forest.
Notes. At the British Museum in 1947-
1948, I compared Oodes piceus with wester-
manni Laferte as identified by Andrewes,
and could find no significant external dif-
ferences except in form of S front tarsi,
which are wider in westermanni. The New
Cuinean specimens are 9 9 , so their as-
signment to piceus is tentative. Oodes
piceus has been recorded from SE. Asia,
Sumatra, Java, the Philippines, and Cele-
bes. O. westermanni occurs in the same
general area.
Oodes exiguus Andrewes
Andrewes 1933, Ent. Monthly Mag. 69, p. 56.
pijginaeus Andrewes 1936, Treubia 15, p. 218
(name used in error for exiguus).
Description ( for recognition only ) . Very
small, size of Anatrichis pusilla but differing
as noted below; see also preceding Key to
Species; length c. 5 mm.
Types. A S from Sumatra, in Deutsches
Entomologisches Mus. (not seen); a 9
"cotype" in Andrewes Coll., British Mus.
( seen ) .
Occurrence in New Guinea. Papua: 71,
Dobodura, Mar.-July 1944 (Darhngton);
12, Oro Bay, Dec. 1943-Jan. 1944 (Dar-
lington). West N. G.: 23, Hollandia, July-
Sept. 1944 (Darlington); 6, Maffin Bay,
Aug. 1944 (Darlington).
Notes. The known range of this species
is now Sumatra (the types), Leyte Is. in
the Philippines (Darlington, M.C.Z.),
Morotai Is. in the Moluccas (Darlington,
M.C.Z.), and New Guinea. It is not known
in Australia.
This small oodine differs from Anatrichis
pusiUa as follows: form wider; only 1 seta
over each eve (2 in pusilla); mandibles
longer, straighter; clypeus with seta-bearing
punctures; elytra with striae not punctulate;
c? front tarsi wider (2nd segments c. long
as wide), with 3 segments squamulose
below. Both A. pusiUa and O. cxii^uus have
elytra with humeri dentate and 3rd inter\'als
2-punctate.
Oodes nil n. sp.
Description. Form (Fig. 11) average, mod-
erately convex; black, appendages slightly
34 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
riifescent; moderately shining, whole upper
surface with microsculpture of fine c. iso-
diametric meshes and also fine punctulation.
Head 0.50 and 0.51 width prothorax; labrum
6-setose, the 2 middle setae small and close
together; clypeus not margined, without
setae; only 1 (posterior) seta over each
eye; front irregular but scarcely impressed;
mentum tooth triangular, not distinctly
emarginate. Prothorax: width length 1.67
and 1.62; base aj^ex 1.75 and 1.80; disc with
fine middle line but transverse and baso-
lateral impressions slight and poorly de-
fined; posterior edge without setae. Elytra:
width elytra /prothorax 1.05 and 1.06; basal
margin present; humeri not dentate; striae
lightly impressed, finely punctate; intervals
nearly flat, 8th wide at base, 3rd 2-punctate.
Inner wings full. Lower surjaee: prosternal
process weakly or not margined between
coxae; sides of body including metepisterna
extensively and closely punctate. Secondary
sexual charaeters: 9 with 1 seta each side
last ventral segment; 6 unknown. Measure-
ments (types); length 10.5-11; width 3.3-
3.4 mm.
Types. Holotype 9 (M.C.Z., Type No.
31,555) from Dobodura, Papua, Mar.-July
1944 (Darlington); 1 9 paratype from Oro
Bay (near Dobodura), Dec. 1943-Jan. 1944
(Darlington).
Additional material. One 9 , Maffin Bav,
West N. G., June 1944 (E. S. Ross, CaH-
fornia Acad.).
Measured specimens. The holotype and
paratyp(\
Notes. For distinguishing characters of
this species see preceding Key to S))ecies,
and Notes under Codes siccus (p. 37).
The specimen from Maffin Bay is larger
than the types (length r. 12.5 mm) but has
the same technical characters.
Oodes laevissimus Chaudoir
Cliauiloir 1882, Aim. Soc. Eiit. France (6) 2, p.
361.
Andrpwes 1924, Ann. Mag. Nat. Hist. (9) II,
p. 588 ( Simons) .
Description f for recognition onl\). I'orm
parallel, depressed; strongly shining; elytra
lightly striate; for technical characters see
preceding Key to Species; length c. 11.5-
12.5 mm.
Types. From Fly R., presumably Papiia,
collected by D'Albertis; the actual type {t.
Andrewes) in Oberthiir Coll., Paris Mus.
(not seen).
Occurrence in New Guinea. Papua: Fly
R. (the types); 22, Dobodura, Mar.-Julv
1944 (Darlington); 1, Kokoda, 1200 ft. (366
m), Aug. 1933 (Cheesman). N-E. N. G.:
19, Aitape, Aug. 1944 (Darlington); 7,
lower Busu R., Huon Pen., Apr. 4, Mav 13,
1955 (E. O. Wilson, M.C.Z.); 2, Erima,
Astrolabe Bay, 1897 (Biro); 1, Sattelberg
(British Mus.); 2, Wareo, Finschhafen (Rev.
L. Wagner, S. Australian Mus.).
Notes. This distinct species is probabl\-
confined to New Guinea, perhaps to the
eastern and central part of the island. An-
drewes referred it to the genus or subgenus
Simous, but I think this was a mistake.
Simous seems to be a natural group of about
9 known species confined to the Oriental
Region including Sumatra, Java, and Borneo,
and characterized by a very short, broad
labrum and a broad, emarginate mentum
tooth. Oodes laevissimus has the labrum
narrower, the mentum tooth narrowcM- and
scarcely emarginate.
The wings in some individuals of this
species look fully developed and fit for
flight but in others they appear slightly
reduced (but still nearly full) and unfit
for flight. It is doubtful if an\- indixiduals
realK' lly. I ha\-e seen none Irom light
traps.
Although all other New Ouinean Oodini
that I have collectt^d are ac^uatie or semi-
acjuatic-, this one is not associated w ith open
water but lives among dead leaves on the
lloor of rain forest. This is probabl\- also
the habitat ol the two related ionns de-
scribed below .
Oodes rossi n. sp.
D('scri))lioii. I'^oiin as in Figure 12. c. as
Idci'issimus. subparallcl. rathc-i" depress(>d;
The Carabid Beetles of New Guinea
Darlington
35
black; tarsi, antennae, and mouthparts more
brownish; moderately shining, entire upper
surface with very fine c. isodiametric micro-
sculpture but without or with only in-
distinct punctulation. Head 0.52 width
prothorax; labrum 6-setose; clypeus not
margined, without setae; 1 (posterior) seta
over each eye; frontal impressions distinct
but poorly defined; mentum with mod-
erately broad c. truncate tooth. Prothorax:
width ^length 1.66; base /apex 1.69; disc
flattened especially posteriorly, with middle
line (and superficial irregularities) but no
other distinct impressions; 1 well impressed
seta-bearing puncture on each side on dorsal
surface just before basal edge. Elytra:
width elytra prothorax 1.09; basal margin
obsolete; humeri not dentate; striae slightly
impressed, faintly punctulatc; intervals
nearly flat, 8th wide at base, 3rd with
2 inconspicuous dorsal punctures. Inner
wings atrophied, reduced to narrow strips c.
Vs long as elytra. Lower surface: prosternal
process not margined between coxae;
metepisterna (and rest of lower surface)
virtually impunctate. Secondary sexual char-
acters: S front tarsi moderately dilated (2nd
segments slightly wider than long), 3 seg-
ments densely squamulose below; 6 with 1
seta each side last ventral segment; 9 un-
known. Measurements: length 14; width
5.9 mm.
Type. Holotype i ( California Acad. )
from Maffin Bay, West N. G., June 14,
1944 (E. S. Ross); the type is unique.
Notes. This species has probably dif-
ferentiated locally, from /f/ei;/.s.s/r7H/.S'-like
stock, by atrophy of the wings, obliteration
of the basal elytral margin, and other small
changes.
Oodes wilsoni n. sp.
Description. Form (Fig. 13) and char-
acters of the preceding species ( rossi ) ex-
cept as follows. Head 0.51 width prothorax.
Prothorax: width/length 1.72; base/apex
1.78; basal seta-bearing punctures on (not
before) basal edge of pronotum. Elytra:
width elytra prothorax 1.06; striae very
fine, superficial. Inner wings reduced to
vestiges c. V-i long as elytra. Secondary
sexual characters: S unknown; 9 with 1
seta each side last ventral segment. Mea-
surements: length 14; width 5.9 mm.
Type. Holotype 9 (M.C.Z., Type No.
31,556) from Ebabaang, Mongi watershed,
Huon Pen., N-E. N. G., 1300-1400 m, Apr.
16-18, 1955 (E. O. Wilson); the type is
unique.
Notes. This is apparently another local-
ized flightless species derived from laevis-
simiis-like stock.
(Oodes siamensis Chaudoir)
Chaudoir 1882, Ann. Soc. Ent. France (6) 2, p.
358.
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae 5,
p. 1011.
/.s.s!/.s' Andievves 1931, J. Federated Malay Mus. 16,
pp. 434, 444, fig. 4 (new synonymy).
alesi Jedlicka 1936, Acta Soc. Ent. Czechoslovakia
33, p. 66 (new synonymy).
Description ( for recognition only ) . Form
average, depressed; black; clypeus mar-
gined, with setae in angles; see siamensis
vulsus in preceding Key to Species of Oodes,
but note basal seta-bearing punctures of pro-
notum usually present in typical siamensis
( see Notes below ) ; length c. 8-9 mm.
Types. Of siamensis, from Bangkok,
Thailand; in Oberthiar Coll., Paris Mus.
( not seen ) . Of issus, from Brunei, Borneo;
in Andrewes Coll., British Mus. (seen). Of
alesi, from Mt. Makiling, Luzon; in British
Mus. ( seen ) .
Occurrence in New Guinea. Represented
only by the following subspecies.
Notes. The synonymy suggested above
is based on examination of the types of
issus and alesi at the British Museum, and
comparison with many specimens from
other localities. They seem to represent
one variable species which is widely dis-
tributed in SE. Asia, Sumatra, Borneo,
the Philippines, New Guinea, and New
Britain, and presumably intervening is-
lands too.
My single specimen of siamensis ( issus )
from Borneo has distinct basal pronotal
36 Bulletin Museum of Comparative Zoology, Vol. 137, Xo. 1
setae on both sides, but they rise from
scarcely detectable punctures that could
hardly be seen if the setae were missing.
Some, but perhaps not all, of my specimens
of this species (alesi) from Leyte have
these setae present too. However, I have
carefully examined both sides of all 36
specimens of the species from New Guinea
and 16 from New Britain, and can find no
trace of basal pronotal setae or punctures
in any of them. This gives a basis for
separating the New Guinea-New Britain
population as a geographical subspecies
(below). First, however, I have had to
discuss siamenms as a whole, in order to
establish the synonymy and distribution of
the species.
Oodes s'tamensis vulsus n. subsp.
Description. Similar to sianieiisis scnsu
stricto (above) but without seta-bearing
punctures on basal edge of pronotum.
Head 0.51 and 0.51 width prothorax. Pro-
thorax: width/length 1.57 and 1.63; base/
apex 1.81 and 1.84. Elytra: width elytra
prothorax 1.08 and 1.07. Measurements:
length c. 8-9; width 3.3-3.7 mm.
Types. Holotype S (M.C.Z., Type No.
31,357) and 13 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington). Ad-
ditional paratypes from West N. G.: 20,
ilollandia, July-Sept. 1944 (Darlington); 1,
Maffin Bay, Aug. 1944 (Darlington); 1,
"Neth. New Guinea," Oct. 20, 1944 (T.
Aarons, California Acad.).
Measured sj)eeiniens. The i holotype and
1 9 paratype from Dobodura.
Notes. This subspecies occurs also in
New Britain ( 16, Cape Gloucester, Dar-
lington, M.C.Z.).
Oodes denisonensis Castelnau
Castelnau 1867, Notes on .Vuslialiaii C^olcoptcra, p.
64.
Sloaiic 1910, Proc. Liimcan Soc. N(nv .South Wales
35, pp. U5, 447.
Csiki 1931, Cok'op. Cat., Carabidac, Haipalinac 5,
p. 1007.
Description (for recognition only). Form
broad, convex; for technical characlcM-s see
Sloane's (1910) key, and preceding Key to
Species of Oodes of New Guinea; length
c. 13.5-15 mm.
Type. From Port Denison (probably
near Bowen, Queensland), Australia; pres-
ent location of type unknown (not seen).
Occurrence in New Guinea. Papua: 1
9 , Rouku, Morehead R., Apr. 1962 {W. \\.
Brandt, C.S.I.R.O.). West N. G.: 1 9,
Merauke (south coast), sea level. Mar. 28,
1955 (L. D. Brongersma, Leiden Mus.).
Notes. The distinctixe characters of
denisonensis are based on the c5 . The two
9 9 from New Guinea agree well in non-
sexual details with specimens from Queens-
land, Australia (from Gayndah, Rockhamp-
ton, Townsville, and Kuranda),
Oodes siccus n. sp.
Description. Form and convexity average;
black, basal angles of prothorax and ap-
pendages slightly more reddish; moderately
shining, whole upper surface finely c. iso-
diametrically microreticulate and iiunctu-
late. Head 0.52 and 0.51 width pro-
thorax; labrum 6-setose; chpeus not mar-
gined, with 1 seta-bearing puncture each
side well behind angle; 1 (posterior) seta-
bearing puncture o\er each eye; front
weakly convex, scarcely impressed an-
teriorly; mentum with rounded-triangular
tooth. Prothorax: width length 1.59 and
1.68; base apex 1.82 and 1.83; disc with
fine middle line, vague wide baso-lateral
impressions, and seta-b(>aring jnmcture each
side on basal edge inside angle. Elytra:
width elytra j^rothorax 1.08 and 1.09; base
margined; humeri not dentat(>; striae mod-
erateK- impressed, laintK punetulate; in-
terxals slightK' convex, 8th wide at base,
3rd 2-punetale. Inner uiii'j^s lull. Lower
surface: prosternal process not margined
between coxae; sides of body iiieluding
metepistcMiia closely punctate. Seco)id(iry
sexual eharaclers: S w itli Ironl tarsi slightly
narrower than usual (2nd segments slightK
longer than wide), with usual 3 s(\gments
densely squamulose; i with 1, 9 2 setae
each side last \("ntr;il setiment. Measure-
The Carabid Beetles of New Guinea • Darlington 37
mcnts: length c. 10-11; width 4.0-4.5 mm.
Types. Holotype $ (M.C.Z., Type No.
31,358) and 12 paratypes from Dobodura,
Papua, Mar.-July 1944 ( DarHngton ) ; and
additional paratypes as follows. Papua: 6,
Lake Daviumbu, Flv R., Aug. 19-30, Sept.
1-10, 11-20, 1936 (Archbold Exp., A.M.
N.H.). West N. G.: 8, Hollandia, July-
Sept. 1944 (Darlington).
Measured speeitnens. The S holotype and
1 9 paratype from Dobodura.
Notes. This is similar to Oodes cribris-
ternis and its allies, but differs in having a
distinet basal elytral margin. It is similar
also to O. nil (described above) but clypeal
and basal pronotal setae are present ( absent
in nil), the elytral striae are less obviously
punctate, and the punctures of the 3rd
intervals are less impressed.
O. siccus occurs also on Morotai Is. in
the Moluccas (Darlington, M.C.Z.).
Oodes par n. sp.
Description. Form ( Fig. 14 ) more quad-
rate than usual, depressed; black, append-
ages in part more rufous; dorsal microsculp-
ture of fine c. isodiametric meshes with very
little additional punctulation. Head 0.57
and 0.59 width prothorax; labrum 6-setose;
clypeus not margined, with seta-bearing
punctiu-e each side behind angle; 1 (pos-
terior) seta over each eye; mentum tooth
entire, bluntly triangular. Prothorax: width/
length 1.57 and 1.61; base/apex 1.62 and
1.51; disc depressed, with moderate middle
line, vague transverse impressions, distinct
but poorly defined rounded baso-lateral
impressions ( sublinear in some lights ) , and
strong seta on basal edge each side inside
angle. Elytra subquadrate; width elytra/
prothorax 1.11 and 1.13; basal margin obso-
lete; humeri not dentate; striae impressed,
punctulate; intervals slightly convex, 8th
wide to base, 3rd 2-punctate. Inner icings
full. Lower sui^ace: prosternal process not
distinctly margined between coxae; sides of
body including metepisterna vaguely or not
punctate. Secondary sexual characters: S
front tarsi moderately dilated (2nd seg-
ments barely wider than long ) , 3 segments
densely squamulose; S with 1, 9 2 seta-
bearing punctures each side last ventral seg-
ment. MeaMirements: length c. 11-12;
width c. 4.6 mm.
Types. Holotype c5 (M.C.Z., Type No.
31,359 ) from Aitape, N-E. N. G., Aug. 1944
(Darlington); and 1 9 paratype, Hollandia,
West N. G., July-Sept. 1944 (Darlington).
Notes. The technical characters, espe-
cially the positions of setae and loss of the
basal elytral margin, suggest that this new
species may be allied to O. crihristernis
and longior, but j)ar differs from both in
being more quadrate and in having the
lower surface including the metepisterna
relatively smooth.
Oodes cribrisfernis Bates
Bates 1892, Ann. Mus. Civ. Genoa 32, p. 323.
Csiki 1931, Coleop. Cat., Carabidae, Harpalinae 5,
p. 1007.
Andrewes 1940, Proc. R. Ent. Soc. London ( B )
9, p. 204 (in key).
Description ( for recognition only ) . Form
moderately slender, depressed; distinguish-
ing characters indicated in preceding Key
to Species of Oodes of New Guinea and in
Andrewes' (1940) key. Prothorax: width/
length 1.67 and 1.63; base/apex 1.63 and
1.58. Elytra: width elytra /prothorax 1.10
and 1.13. Measurements (of New Guinean
specimens ) : length 10.5-13.5; width 4.2-
5.5 mm.
Type. From Burma, in Genoa Mus. (not
seen ) .
Occurrence in New Guinea. Papua: 27,
Mihie Bay, Dec. 1943 (Darlington); 14,
Dobodura, Mar.-July 1944 (Darlington);
1, Mt. Lamington, 1300-1500 ft. (c. 400-
450 m) (C. T. McNamara, S. Australian
Mus.). N-E. N. G.: 1, Lae, Oct. 1944
(Darlington); 2, Aitape, Aug. 1944 (Dar-
lington); 4, Finschhafen, Huon Pen., 10 m,
Apr. 9-16, 1963 (Sedlacek); 1, Wau, Mt.
Missim, Morobe Dist., 880-1050 m, Feb.
8-9, 1963 (Sedlacek). West N. G.: 1,
Maffin Bay, Aug. 1944 (Darlington); 7,
Sansapor, Aug. 1944 (Darlington); 1, "Neth.
38 Bulletin Museum of Comparative Zoologij, Vol. 137, No. 1
New Guinea" without further locality, Oct.
20, 1944 (T. Aarons, California Acad.)-
Measured specimens. A pair (69) from
Dobodura.
Notes. The specimens from New Guinea
here recorded as crihristcrnis possess all
significant characters given in Bates' brief
description and Andrewes' key ( 1940 ) , but
direct comparison will be necessary to con-
firm the identification. The species (if it
is one species ) is now known from Burma,
Sumatra, and New Guinea. O. ohloniius
Castelnau of Australia seems to be allied
but is larger, duller, with finer elytral striae.
Ooc/es longior n. sp.
Description. Form as in Figure 15,
slender, depressed; black, posterior angles
of prothorax and parts of appendages (es-
pecially tarsi, palpi, antennae) ± reddish;
moderately shining, entire upper surface
with fine c. isodiametric microsculpture and
very fine inconspicuous punctulation. Head
0.56, 0.5(S, and 0.59 width prothorax; labrum
6-setose; clypeus not margined, with a
seta each side behind angle; 1 (posterior)
seta-bearing puncture over each eye; men-
tum tooth moderate, impressed or slightly
emarginate. Prothorax: width/length 1.46,
1.44, and 1.49; base/apex 1.70, 1.66, and
1.69; disc with light middle line, no distinct
sulibasal impressions but faintly impressed
each side at extreme base; 1 seta-bearing
puncture on l)asal edge each side near
narrowly rounded basal angles. Elytra:
width elytra prothorax 1.07, 1.09, and 1.09;
basal margin obsolete; humeri not dentate;
apices subangulate near suture (opposite
1st intervals); striae lightly impressed,
faintly punctulat(>; intervals slightly convex,
(Sth slightly narrower than 7th al l)ase, 3rd
2-punctate. huier ivings full. Loner sur-
face: proslernal process not distinctly mar-
gined between coxae; sides ol body below
including metepisterna rather finely, closely
punctate. Secondary .sexual characters: $
fronl larsi moderately dilated (2nd segment
c. wide as long), 3 segments densely
squamnlose b(4ow; ', with 1, 9 2 seta-bear-
ing punctures each side last ventral seg-
ment. Measurements: length c. 14-15;
width 5.2-5.4 mm.
Types. Holotype c^ (M.C.Z., Type No.
31,360) and 1 9 paratvpe from Hollandia,
West N. G., July-Sept! 1944 ( Darlington ) ;
and 1 i para type from Ambunti, Sepik R..
N-E. N. G., May 16, 1929 (Crane-Field
Mus. Pacific Exp., Chicago Mus.).
Measured specimens. The i holotype, 9
paratype from Hollandia, and S paratype
from Ambunti, in this order.
Notes. This new species has the technical
characters (setae, etc.) of crihristcrnis
( above ) but is larger and more slender ( cf.
proportions of criljri.^ternis), with elytra
subangulate at apex.
Although cri1)risfcrnis, like most Oodini,
lives in very wet places, longior may be even
more aquatic. My 2 specimens were taken
in comparatively deep water in floating
debris and vegetation.
Tribe HARPAUNI
Sloane 1898, Proc. Linnean Soc. New South Wales
23, pp. 455, 456 (key to Australian genera).
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 6,
pp. 1023-1268.
Van Emden 1953, Ann. Mag. Nat. Hist. (12) 6,
pp. 513 ff. (discussion in text).
Iliirpalkhic Jeannel 1942, Faune de France, Coleop.
Caraliic}ues, l^ut 2, pp. 57.5-584.
Harpalinae Basilevvsky 1951-1952, Ann. Mus.
Congo Beige (8), Zool., Vols. 6 and 9 (revision
of .Xfrican and Madagascan forms).
The tribe liarpalini contains a large
proportion of the common, medium-sized
(Carabidae that live on the ground in all
climates in all parts of the world. The\' are
very nmnerous in open country, fewer in
rain forest. Those that do li\c' in rain lorest
include many Lccanomcrus in eastern Aus-
tralia and most Trichotichnus in Ne\\-
Ciuinea. The tribe also contains many
smaller species that \i\v in wet places or
b(\side quiet (usually not rapidh' running)
Wider.
(Classification of genera within the Ilar-
jxilini is exceptionalh' difficult, perhaps (I
suspect) because the tribe is relatixcly re-
The Carabid Beetles of New Guinea • Darlington
39
cent in evolution and dispersal. Generic
classifications proposed for members of the
tribe in any one region fail in other regions,
and no usable classification exists for the
genera of the Oriental Region and Malay
Archipelago. The arrangement of genera
in the Junk Catalogue (Csiki, 1932) is said
to follow Schauberger, but he died without
explaining it. The classification that I am
using for the New Guinean forms ( see Key,
below) is based partly on well known
characters that are probably phylogenetic,
but nevertheless the key is partly superficial
and is intended primarily as an aid in
identification, not as a contribution to
harpaline classification.
Characters drawn from the soles of the
male tarsi are fundamental in harpaline
taxonomy but are, of course, useless in the
case of unassociated females. (The single
possible Trichotichmis that I have from
Australia is a female and therefore not
identifiable! ) And characters of the mouth-
parts, including the setae of the penultimate
segments of the labial palpi, are funda-
mental too, but are difficult to see and
understand. Even experienced carabid
specialists make mistakes in placing har-
paline genera. Bates' original placing of
Lamprophonus- and Andrewes' of Carbanus
are examples. Both genera were originally
wrongly characterized and put in the wrong
subtribes. Many of the harpalines that I
have for study from New Guinea were taken
in light traps, and this increases the dif-
ficulty. Light-trap specimens often have
moth scales adhering to and concealing
their mouthparts and tarsal soles, and scales
stuck to the tarsal soles may even counter-
feit sexual squamae.
The distribution of Harpalini over the
world has been misunderstood until re-
cently because of lack of adequate subtribal
and generic classifications, and because of
incorrect assignments of many Australian
and South American species to northern
genera, especially to Harpalus and Aniso-
dactylus. Van Emden (1953), however,
has suggested what I think are natural and
Table 1. Distribution of Principal Subtribes
OF Harpalini (after Van Emden 1953)
1. Anisodactylina: worldwide, but irregularly
distributed; genera in Australia and South America
are prolaably not directly related.
2. Harpalina, Harpali ( Harpalu.s and its im-
mediate allies ) : tliroughout Eurasia, Africa ( and
Madagascar), and North America; absent in Aus-
tralia and South America.
3. Harpalina, Selenophori: most of the world
including South America, but absent in most of
Australia (one or two Oriental genera reach just
the northern edge of Australia ) .
4. Pelmatellina: chiefly Australia and South ( and
Central) America. The genus Nemaglossa may
occur in both Australia and South America but
lias not been adequately studied.
5. Acupalpina: nearly worldwide, with some
genera very widely distributed. The members of
this subtribe are mostly small, water-loving forms
which do not compete with most members of
the other subtribes, except perhaps with small
Pelmatellina in Australia.
useful subtribes and has indicated their
distributions. His arrangement, slightly
modified, is summarized in Table 1.
This outline of harpaline distribution
(Table 1) is, of course, an oversimphfica-
tion. A more detailed study of the distribu-
tion of subtribes of Harpahni would be an
important contribution to insect zoogeogra-
phy.
Within the limits of New Guinea and
Australia, harpaline faunae overlap com-
plexly. Among larger, terrestrial Harpalini
at low altitudes, several primarily Aus-
tralian genera of subtribe Anisodactylina
(Gnathaphanus, Diciphoromerus, Hyphar-
pax) extend to New Guinea and westward
into or across the Malay Archipelago. These
genera live chiefly in relatively open
country, including open Eucalyptus wood-
land, although some of them enter rain
forest too. On the other hand, several pri-
marily Oriental genera of subtribe Har-
palina, especially Trichotichmis and other
Selenophori (but not Harpalus), reach New
Guinea and are dominant there, outnum-
bering the Australian Anisodactylina espe-
cially in rain forest. These genera either
40 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
do not extend to Anstralia or are repre-
sented there by single species on the ex-
treme northern edge of the continent (e.g.,
a CoIeoUssus on Cape York). The Austrahan
genera, chiefly in more open country, and
the Oriental ones, chiefly in rain forest, are
in part ecologically as well as geographi-
cally complementary. This pattern suggests
recent multiple invasions of the rain-
forested areas of New Guinea by Oriental
stocks and of the more open areas by Aus-
tralian stocks, but over a long period some
replacement of Australian by incoming,
competing Oriental groups may have oc-
curred.
At much higher altitudes on New Guinea
is one additional genus of Anisodactylina,
CJiydaeus, which is primarily Asiatic and
has apparently "mountain hopped" across
the Malay Archipelago. This genus does
not reach Australia.
Among smaller, water-loving Harpalini,
primarily Oriental Acupalpina are domi-
nant in New Guinea and several genera
reach the northern half of Australia, but
they decrease or disappear in southern Aus-
tralia. Their place there is taken by small
Pelmatellina {Lecanomenis), which are
ninnerous throughout Australia and a few
of which occur in New Guinea (described
in the following pages) but which are un-
known farther west in the Malay Archipel-
ago. The distributions of Oriental Acu-
palpijia and of small Australian Pelmatellina
are therefore broadly complementary too
in the Australian Region, but with wide and
complex overlapping.
Key to Geneha ok IIaiu'ai.im oi' New Guinea
1. Male front and micldle tarsi with sponge-
like soles of many densely packed, narrow
adhesive hairs 2
- Male front and nsually (not always) middle
tarsi 2-seriately scinanuilose below, or
rarely ( Lyter only) with more than 2
rows of long, narrow seales loosely ar-
ranged - 6
2. Size larger (6-lfi nun); scntellar striae
present (short in llii]>h(ir])(ix); pennltimate
segments labial palpi plnrisetose (Aniso-
dactylina ) 3
- Size smaller ( less than 5 mm in New
Guinea); scntellar striae absent; penulti-
mate segments labial palpi 2-setose
( Pelmatellina ) ( p. 45 ) Lecanameriis
3. Elytra without dorsal pimetures; wings
atrophied; (found only on high mts.) (p.
47 ) Chijdacus
- Elytra each with 1 or more dorsal pime-
tures; wings usually full 4
4. Elytra with se\eral or many conspicuous
dorsal punctures (p. 41) ,-. Gnathaphanus
- Elytra each with 1 dorsal pimcture _ . 5
5. Posterior tarsi long, basal segment much
more than 2X long as wide; hind femora
not strongly curved (p. 42) __ Diaphorovxents
- Posterior tixrsi shorter, basal segment 2X
or less long as wide; hind femora of c5
strongly curved, of 2 less so ( p. 44 )
Hyphen pax
6. Penultimate segment labial palpi with more
than 2 setae anteriorly; often larger ( 5-
11 mm) (Harpalina) 7
- Penultimate segment labial palpi 2-setose
anteriorly; often smaller (2.7-8.0 mm)
(Acupalpina) 12
7. Front tibiae broader, apex % or % wide
as tibial length (by measurement) (p.
59) Harpaloxenns
- Front tibiae narrower 8
8. Entire upper surface pubescent ( p. 48 ) __
PIdti/ntcfopus
- Upper surface not pubescent 9
9. Elytra with 3rd intervals 1-punctate or
impunctate 10
- Elytra with 3rd intervals with several
(very small) punctiues 11
10. Male front and middle tarsi with soles of
long, slender, loose (not 2-seriate) scales;
base of prosternuin and base of abdomen
not pubescent (p. 63) Lytcr
- Male front and (usually) middle tarsi 2-
seriateh' s(iuanndose; b;ise ot pronotnm
and b;ise of abdomen usually short-pubes-
cent (but see Notes under T. nwiliu.s)
(p. 48) -, - riirln>lirlmu.s
11. Last \entral segment with 2 sel;ie e;ich
side in both sexes; eh tra with sutural
angles denticulate ( in New Guinean
species) (p. 64) Coleoli.ssus
- Last ventral seguient with 1 si-ta each side
in both sexes; sutural angles not denticulate
(p. 66) Ily))luicri()u
12. Scutt'llar striae absent; anicrior iii;uginal
line of pronotnm ^\^'^'\^ and entire; length r.
7-8 nun (p. 68) .A/i<)/)/(),i,'r;i;(/.v
- Scntellar striae present; iuilerior m;irgin;il
line of pronotnm line or interruptiil ;it
middle; usualK sm;iller 13
13. Abdomen not pubescent (except for fi.xed
setae ) ( p. 69 ) Egadroina
The Carabid Beetles of New Guinea
Darlington
41
- Abdomen pubescent at least near apex 14
14. Prosternuni without long setae anteriorly
(p. 71) Stcnoloplius
- Prosternuni with several long setae an-
teriorly ( p. 72 ) Acupalpus
Genus CNATHAPHANUS Macleay
Macleay 1825, Annulosa Javanica 1, p. 20.
Chaudoir 1878, Ann. Mus. Civ. Genoa 12, pp.
476, 503.
Sloane 1900, Proc. Linnean Soc. New South Wales
24, p. 553 ( key to some Australian species ) .
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 6,
p. 1041 (see for synonymy and additional refer-
ences ) .
Diaii.no.sis. See preceding Key to Genera
of IlarpaJini of New Guinea.
Description. None required here.
Type species. G. vidneripennis Macleay,
of Java, etc.
Generic distribution. Many species in
Australia, fewer in the Malay Archipelago
and adjacent comer of Asia, with one or
two widely distributed species reaching
India, the Philippines, and islands east to
Samoa and New Caledonia.
Notes. Some species of this genus have
very wide ranges, within the limits given
above. Of the 5 species known in New
Guinea, all are shared with Australia and
several are widespread also on the Malay
Archipelago or islands of the western
Pacific. These insects are often common in
open coimtry including grassland and open
woodland, but are not often found in rain
forest. All species of the genus that I know
are fully winged and probably fly.
Key to Species of Gnathaphanus of New Guinea
1. Elytra with intervals 3, 5, and usually 7 (at
least posteriorly) with dorsal punctures con-
spicuously impressed; (black, legs black;
elytra deeply sinuate and acuminate at apex)
(p. 41) licinoidcs
- Elytra with fewer, less impressed dorsal
punctures 2
2. Elytra with series of dorsal punctures on
outer edges of intervals 3 and ( at least pos-
teriorly) 5; (legs yellow) (p. 41) upolcnsis
- Elytra with series of dorsal punctures only
on 3rd intervals ( single punctures sometimes
present on other intervals) 3
3. Smaller ( c. 9-10 mm ) ; more shining ( es-
pecially the S ) ; piceous, legs brownish
yellow ( p. 42 ) picipes
- Larger (c. 12-13 mm); dull black or me-
tallic, legs black 4
4. Head and prothorax green, elytra cupreous
(except in discolored individuals); pos-
terior angles of prothorax distinct, bluntly
obtuse or very narrowly rounded (p. 42) ..
pulcher
- Dull black; posterior angles of prothorax
broadly rounded (p. 42) philippcnsis
Gnathaphanus licinoides Hope
Hope 1842, Ann. Mag. Nat. Hist. 9, p. 427.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 6,
p. 1042 ( see for synonymy and additional refer-
ences ) .
Description. None required here; see
Key, above; length c. 10 mm.
Type(s). From Port Essington, northern
Australia; presumably in Hope Mus., Ox-
ford (not seen).
Occurrence in New Guinea. Papua: 24,
Dobodura, Mar.-July 1944 (Darlington); 1,
Kokoda, 1200 ft. (366 m), Sept. 1933
(Cheesman); 1, Wakaiuna, Sewa Bay, Nor-
manby Is., Jan. 1-8, 1957 (W. W. Brandt,
Bishop Mus.). N-E. N. G.: 18, Wau, 1200
m, dates in Jan., Mar., Apr., June, July,
Sept., Nov., Dec. 1961-1963 (Sedlaceks);
1, Stephansort, Astrolabe Bay, 1899 ( Biro ) ;
1, Aitape, Aug. 1944 (Darlington). West
N. G.: 1, Hollandia, May 1945 (B. Malkin,
U.S.N.M.); 1, same area, Cyclops Mts., 50-
100 m, June 22-24, 1959 (Gressitt, T. C.
Maa, Bishop Mus.), in light trap.
Notes. Tlie known range of licinoides in-
cludes northern Australia, New Britain,
the Solomons, New Hebrides, and New
Caledonia, as well as New Guinea.
Gnafhaphanus upolensis (Csiki)
Csiki 1915, Denkschriften Akad. Wiss. Wien,
Math-Nat. 91, p. 163 (Diortjche).
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 6,
p. 1044 (see for synonymy and additional refer-
ences ) .
impressipennis Castelnau 1867, Notes on Aus-
tralian Coleop., p. 100 ( in Harpalus, but not
Harpahis iiu))ressipennis Motschulsky 1844).
Chaudoir 1878, Ann. Mus. Civ. Genoa 12, p. 510.
Description. None required here; length
c. 8-9 mm.
42 BuUetin Museum of Coynparative Zoology, Vol. 137, No. 1
Types. Of impressipennis, from Rock-
hampton, Australia; in Genoa Mns. Of
upolensis, from Upolu, Samoa; in Vienna
Mils, (not seen).
Occurrence in New Guinea. Common
( 175 specimens seen ) probably throughout
New Guinea at low altitudes including
Dobodura, up to 1200 m at Wau and to
2300 m on Mt. Kaindi ( near Wau ) . Speci-
mens taken in every month.
Notes. This very^ common carabid occurs
usually in relatively open country, includ-
ing grassland and open Eucalyptus wood-
land, from the Malay Peninsula across the
Malay Archipelago to New Guinea and
Australia, east at least to the Philippines
and Samoa, and New Caledonia.
Gnafhaphanus picipes (Macleay)
Macleay 1864, Trans. Ent. Soc. New South Wales
1, p. 117 (Harpahis).
Chaudoir 1878, Ann. Mus. Civ. Gencxi 12, p. 509.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 6,
p. 104.3 (see for synonymy and additional refer-
ences ) .
De.<icription. None required here; length
c. 9-10 mm.
Types. From Port Denison (Bowen),
Queensland, Australia; probably in Mac-
leay Mus., Sydney (not seen).
Occurrence in New Guinea. Papua: 12,
Port Moresby, Jan., Feb., Mar., May, Aug.,
Oct., Dec. (various collectors; M.C.Z.,
British Mus., Bishop Mus., U.S.N.M., Dept.
Agr. Port Moresby), some under logs in
FAicali/))lus country, some at light; 2, Browu
K., May 22, 25, 1956 (F. J. Ford. Jr., Bishop
Mus.).'
Notes. This uortheastcrn Australian
species apparently extends only to the
southern edge of New Guinea.
Gnafhaphanus pulcher (Dejean)
Dejean 1829, Species Cleneral Coleop. 4, p. 282
( llai))altis).
Cliaiidoir 1878, .Ann. Mus. Civ. Cenoa 12, p. .505.
Csiki 1932, Coleop. Cat., Caral)idae, Harpalinae 6,
p. 1043 (.see for synonymy and additional refer-
ences ) .
Description. None required here; length
r. l.'3-16 mm.
Types. From "Nouvelle-Hollande" ( =
Australia); presumably in Oberthiir Coll.,
Paris Mus. (not seen).
Occurrence in New Guinea. Papua: 13,
Port Moresby area, various dates in Jan..
Feb., Mar., May (various collectors; Dept.
Agr. Port Moresby); 3, Bisianumu, 1600 ft.
(485 m), Feb. 12,'l966 (J. H. Barrett, Dept.
Agr. Port Moresby ) .
Notes. This Australian species apparently
reaches only the southern part of New
Guinea, perhaps only the open-wooded
Eucalyptus areas where many other Austra-
lian insects occur. It is represented on the
Lesser Sunda Islands, west to Bali, by
subspecies extrarius Schauberger.
Gnafhaphanus philippensis (Chevrolat)
Chevrolat 1841, Revue Zool., p. 221 {Amhhjgmi-
thus).
Chaudoir 1878, Ann. Mus. Civ. Genoa 12, p. 511
(as laeviceps Macleay).
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 6,
p. 1043 ( see for synon>my and additional refer-
ences ) .
Description. None required here; length
c. 12-16 mm.
Type(s). From "Manille" (Manila); in
Hope Mus., O.xford (not seen).
Occurrence in New Guinea. Papua: 1,
Kokoda, 1200 ft. (366 m). May 1933
(Cheesman); 3, Rouku, Morehead R.,
\Vest Papua, Apr. 1962 (W. W. Brandt.
C.S.I.R.O.).
Notes. Tliis species ranges from SE. Asia
to Australia, east to Philippines, but is
surprisingly scarce and perhaps localized
ill New (hiinea.
Genus DIAPHOROMERUS Chaudoir
Chaudoir 18 1.), Hull. Soc. \at. Moscow Ifi, Part
2, p. 402.
1878, Ann. Mus. Civ. Ccnoa 12. p. 17(S.
Csiki 1932, Coleop. Cat., Caral)i(lae, llarpaliuar (i,
p. 1044 (see lor additional rclerences ).
DiaL!,nosi,s. See Key to Genera of llar-
paJini of New Guinea.
Description. None rtciuircd hen-.
Type species. D. iridipennis ('haudoir, ol
.Xustralia including ("ape York (CMiaudoir
ISTS).
The Carabid Beetles of New Guinea • Darlington 43
Generic distribution. Primarily Austra-
lia, with species also on New Zealand, New
Caledonia, New Guinea, the Molueeas
(Amboina), and Timor, and with 2 New
Guinean species extending to New Britain.
Notes. Many species of this genus in
Australia inhabit open Eucah/pius wood-
land or grassland. The two species in New
Guinea occur in rain-forested parts of the
island, but I do not know their exact
habitats. The New Guinean species of
Diaphoromerus, like most Australian ones,
are winged.
Key to Species of Diaphoromerus of
New Guinea
1. Larger (8.5-10.5 mm); posterior angles of
prothorax (narrowly) rounded (p. 43) ._.
papuensis
- Smaller (6.0-7.5 mm); posterior angles of
prothorax obtusely angulate, scarcely blunted
(p. 43) papuclhis
Diaphoromerus papuensis (Macleay)
Macleay 1876, Proc. Linnean Soc. New South
Wales 1, p. 168 {Harpalu.s).
Csiki 1932, Coleop. Cat., Caral:)idae, Harpalinae 6,
p. 1043 (Gnathaphanus).
hasilewsktji Louwerens 1962, Tijdschrift voor Ent.
105, p. 139 (Gnatluiplianus) (new synonymy).
Description. None required here; length
c. 9-10 mm.
Types. Of papuensis, from Hall Sound,
Papua; presumably in Macleay Mus.,
Sydney (not seen). Of basilewskyi, from
Amboina Is., Moluccas, 70 m altitude
(A. M. R. Wegner), at light; holotype in
Louwerens Coll. (not seen), 2 paratypes
now in M.C.Z.
Occurrence in Neiv Guinea. Common
probably throughout New Guinea: 119,
from widely scattered lowland localities,
from Port Moresby and Dobodura to
Manokwari, up to 1300 m at Wau, and (SOO
m at Araucaria Camp, West N. G. Speci-
mens collected in every month except
August.
Notes. Macleay 's statement that the
third elytral interval is punctate on inner
side before apex places this species in
Diaphoromerus rather than Gnathaphanus,
and the length (4y2 lines = 9 mm) is diag-
nostic of this species in New Guinea.
Closely related species in Australia prob-
ably include mehnarius Dejean and iridi-
pennis Chaudoir. D. papuensis occurs also
in New Britain (Cape Gloucester, Dar-
lington) and the Moluccas (types of
basilewskyi) .
Diaphoromerus papuellus n. sp.
Description. Form as in Figure 16, rather
small, convex; brownish piceous, append-
ages testaceous or brownish testaceous;
moderately shining, ? scarcely duller, both
sexes with reticulate microsculpture iso-
diametric or slightly transverse on head,
more transverse on pronotum and elytra.
Head 0.69 and 0.69 width prothorax; eyes
prominent; front weakly impressed; mentum
toothed; ligula slightly shorter than para-
glossae, latter separate at apex. Prothorax
transverse-subquadrate; width/length 1.43
and 1.44; base/apex 1.41 and 1.39; sides
rounded anteriorly, nearly straight, con-
verging, sometimes slightly sinuate before
slightly obtuse but distinct and scarcely
blunted basal angles; disc formed as usual,
basal impressions sublinear, weak, mar-
gined at base but not or indistinctly punc-
tate. Elytra: width elytra prothorax 1.20
and 1.19; sides slightly sinuate before apex;
striae impressed; intervals slightly convex,
subequal, 3rd 1 -punctate on inner side near
apex. Inner wings full. Legs: 1st segment
hind tarsi elongate. Secondary sexual char-
acters: S front and middle tarsi dilated
( 2nd and 3rd segments of front tarsi slightly
wider than long, of middle tarsi narrower ) ,
with densely pubescent soles. Measure-
ments: length 6.0-7.5; width 2.0-2.8 mm.
Types. Holotype $ (British Mus.) and
6 paratypes (some in M.C.Z. , Type No.
31,361) from Kokoda, Papua, 1200 ft. (366
m), Aug. (except one specimen May) 1933
(Cheesman); and additional paratypes as
follows. Papua: 1, Kerema, May 3-9, 1959
(C. D. Michener, Bishop Mus.); 2, Kiunga,
Fly R., July 4-8, Aug. 8-10, 1957 (W. W.
Brandt, Bishop Mus.); 1, Rouku, Morehead
44 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
R., West Papua, Apr. 1962 (W. W. Brandt,
C.S.I.R.O.); 3, Yule Is. (Hungarian Na-
tional Mus.); 39, "Papua" without further
locality ( Hungarian National Mus. ) . West
N. G.': 2, Merauke, Apr. 6, 1952 (L. D.
Brongersma, Leiden Mus.); 1, same locality,
Jan. 26-Feb. 10, 1960 (T. C. Maa, Bishop
Mus.); 1, Kepi, Res. Mappi, Oct. 15, 1957
(R. T. Simon Thomas, Louwerens Coll.);
4, Wasian, Sept. 27, 1939 (R. G. Wind,
California Acad.). Also 1 paratype, Koitaki,
1500 ft. (455 m), New Guinea (diyision
unknown), Oct.-Noy. 1928 (Pemberton,
H.S.P.A.).
Measured specimen.^. The i holotype and
1 9 paratype from Kokoda.
Notes. I haye seen a specimen of this
species from Kerayat, New Britain (E. J.
Ford, Jr., Bishop Mus.).
This is eyidently a member of the
Diuphoromerus austral is group. As com-
pared with austral is itself, the present new
species has better defined posterior pro-
thoracic angles. In this character it agrees
with D. aereus Dejean, of SW. Australia,
but papueUus lacks the obyious punctation
of the base of the pronotum of aereus. As
compared with queen.slandicus Csiki (man-
dihularis Castelnau), papueUus is larger,
with more obtuse posterior prothoracic
angles.
Genus HYPHARPAX Macleay
Macleay 1825, Annulosa Javanica 1, p. 22.
Chaudoir 1878, Ann. Mus! Civ. Genoa 12, p. 496.
Sloanc 1898, Proc. Liniican Snc. New South Wales
23, pp. 458-159.
Csiki 1932, Coleop. Cat., Carahidae, Ilarpalinae 6,
p. 1051 ( se(" lor s> ii()n\ni\' and additional refer-
ences ) .
D/V/ij/fo.s/.v. See preceding Key to Genera
of llarpali)ii of New Guiiiea.
Description. None required here.
Type .species. //. lateralis Macleay ( =
dentipes Wiedemann), of Jaya.
Generic distribution. Chiefly Australia.
extending to New Zealand, and west in the
Malay Ar« •hijH'hi'io to Java and Sumatra.
Notes. See Notes under following species.
Hypharpax dentipes (Wiedemann)
Wiedemann 1823, Zool. Magazin 2, p. 54 (Harpalus).
Chaudoir 1878, Ann. Mus. Civ. Genoa 12, p. 500.
Andrewes 1919, Trans. Ent. Soe. London for
1919, p. 158.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae
6, p. 1052 (see for synonymy and additional
references ) .
Description. None required here. This
is the only species of the genus known
from New Guinea. Length c. 7-10 mm.
Type. From Java; in Copenhagen Zool.
Mus. (not seen).
Occurrence in New Guinea. Papua:
39, Port Moresby and vie, May, Sept.,
Oct., Dec. (yarious collectors; M.C.Z., Brit-
ish Mus., Bishop Mus., Dept. Agr. Pt.
Moresby), some under logs in Eucalyptus
country, some at light; 6, Yule Is., Nov. 7
and 16, 1933 (R. V. Oldham, British Mus.);
2, same locality (Fry Coll., British Mus.);
1, Lake Daviumbu,' Fly R., Sept. 11-20,
1936 (Archbold E.xp., A.M.N.H.); 1, Rouku.
Morehead R., West Papua, Mar. 1962 (W.
W. Brandt, C.S.I.R.O.). N-E. N. G.: 3, Lae
and yic. Mar. 1963, Aug. 1964 (Sedlacek);
18, Sum-Sum, 64 km N. of Wau, 580 m,
Feb. 15, 1963 (H. W. Chssold, Bishop Mus.);
4, Bulolo, 720 m, Aug. 13, 19, 24, 27, 1956
(E. J. Ford, Jr., Bi.shop Mus.), 2 of these
taken in light trap; 1, Wau, 1200 m. May
1-15, 1962 (Sedlacek) in light trap. Also
7 specimens from Papua, "British N.
Guinea," and New Guinea without exact
localities.
Notes. The sexes of dentipes differ con-
siderably: males not only have the front
and middle tarsi dilated, with spongy soles,
but also ha\e the hind femora more or less
dentate and the hind tibiae more or l(\ss
cur\'ed. The deyelopment of the femoral
tooth and the degree of cur\ature of the
tibiae xar)' indi\idually in males from sin-
gle localities and also vary geographicalK",
and the si/e of the insect varies geograph-
icalK'. The species therefore has received
several names. The synonymy has not been
fully worked out, but my impression is
that a single \ariabl(^ spcx'ies of Hypharpax,
The Carabid Beetles of New Guinea • Darlington
45
for which dentipes is the oldest name, oc-
curs in Sumatra, Java, Celebes, and New
Guinea, and that it occurs also in NE.
Australia under the name kreffi Castelnau.
This tentative conclusion should be tested
by more rigorous study, for which I now
have neither the material nor the time.
In New Guinea this species has been
found onlv in the eastern half of the island,
especially but not exclusively in the more
open Eucalyptus country of southern Papua.
Genus LECANOMERUS Chaudoir
Chaudoir 1850, Bull. Soc. Nat. Moscow 23, Part 1,
p. 446.
Sloane 1920, Proc. Linnean Soc. New South Wales
45, pp. 132, 137 (as synonym of Nemaf!,lossa).
Csiki 1932, Coleop. Cat., Carabidae, Haipalinae 6,
p. 1058 ( as synonym of Nemaglossa ) ( see for
additional references ) .
Thenarotes Bates 1878, Cistula Ent. 2, p. 319.
Diagnosis. Small Harpalini ( length under
5 mm in New Guinea); elytra without
scutellar striae (in New Guinean species);
penultimate segments labial palpi con-
spicuously 2-setose; S front and middle
tarsi with densely pubescent soles.
Description (characters common to New
Guinean species ) . Form c. as in Figures 17,
IS; more compact and convex than in most
Acupalpina, with margins of prothorax and
elytra relatively narrow. Head: mandibles
moderately long, straight posteriorly, curved
apically; eyes not very large but almost
contiguous with sides of mouth below;
frontal impressions deep, curved, sharply
defined; mentum with triangular tooth;
ligula 2-setose, with paraglossae attached,
longer than ligula; palpi short, apical seg-
ments subconical, penultimate segments of
labial palpi 2-setose. Prothorax subquadrate
or subcordate; disc convex, median longi-
tudinal line impressed, baso-lateral impres-
sions shallow and poorly defined, surface
of disc punctate across base, almost impunc-
tate elsewhere. Elytra: humeri prominent;
basal margin entire, rounded or obtusely
subangulate at humeri; striae impressed,
entire, not distinctly punctate; scutellar
striae lacking; 3rd intervals 1-punctate on
inner edge behind middle. Inner icings
full. Lotcer surface including abdomen
virtually glabrous except for "fixed" setae.
Secondary sexual characters: i front and
middle tarsi moderately dilated, with
densely pubescent soles; 2 setae each side
apex last ventral segment in both sexes.
Type species. Of Lecanomerus, L. in-
sidiosus Chaudoir, of SW. Australia; of
Thenarotes, T. tasmanicus Bates, of Tas-
mania.
Generic distribution. Species of Lecano-
merus (sensu lato) are diverse in Australia,
less so in New Zealand, New Caledonia,
and New Guinea. For further details see
Notes, below.
Notes. The supposed identity of Lecano-
menis (including Thenarotes) of Australia
and NemagJossa of Chile is doubtful.
Sloane ( 1920 ) , who suggested it, did so
without what would now be considered
critical study, and I have not been able
to make the comparisons necessary to con-
firm it. I shall therefore tentatively treat
Lecanomerus as distinct from NemagJossa
and confined to the Australian Region. The
genus does not have an "Antarctic" distribu-
tion pattern. Species are numerous and
diverse along the whole eastern edge of
Australia north to Cape York. Five species
are reported from Tasmania (Sloane), but
4 of them occur on the Australian mainland
too, and the 1 species endemic to Tasmania
is not much differentiated.
The 3 small, compact Lecanomerus found
in New Guinea resemble, but are specifi-
cally distinct from, certain unidentified
species that I found common on the Cape
York Peninsula of Australia in 1958. The
New Guinean forms occur in rain-forest
areas, not in Eucahjptus country. They
probably live among dead leaves and under
vegetation on the ground near standing
water or perhaps sometimes in leaf litter on
the floor of rain forest. However, I did not
distinguish them in the field and cannot be
sure of their habitats.
46 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Key to Species of Lecanomervs of
New Guinea
1. Prothorax narrowly subcortlate ( width /length
1.22 and 1.18); '(Hollandia, West N. G.)
( p. 46 ) angustior
- Prothorax wider; { Papna ) 2
2. Brown: slightly narrower {cf. proportions in
Description.^); piinetation of base of pronotnm
discontinnous, with middle of base virtnalK'
impnnctate (p. 46) incdius
- Black; relatively sHghtly wider; punctation
somewhat irregular but c. continuous across
base of pronotnm (p. 46) latior
Lecanomerus angustior n. sp.
Description. With characters of genus;
fomi (Fig. 17) narrowly compact; color
brownish piceous, prothoracic and elytral
margins and suture usually slightly rules-
cent, appendages testaceous; moderately
shining, reticulate microsculpture faint,
slightly transverse on front and on pronotal
disc, more transverse on elytra. Head 0.69
and 0.69 width prothorax; eyes smaller than
average, genae slightly rounded-oblique.
Prothorax: width length 1.22 and 1.18;
base apex 1.13 and 1.15; sides weakly
rounded anteriorly, converging and usually
sinuate posteriorly before c. right posterior
angles; base and apex unmargined at least
at middle; base of pronotum punctate at
sides, scarcely so at middle. Elytra: width
elytra/prothorax 1.47 and 1.45. Measure-
ments: length 3.6-4.0; width 1.6-1.7 mm.
Types. Holotype i (M.C.Z., Type No.
31,362) and 6 paratypes all from Hollandia,
West N. G., July-Sept. 1944 (Darlington).
Meamired specimens. The .', holotype and
1 9 paratype.
Notes. See preceding Key to S))ecies for
diilerential characters. This is the western-
most known species ol the genus. It prob-
ably represents medius (below) ol eastern
New Guinea. Perhaps additional related
forms are still to be found in western West
N. G.
Lecanomerus medius n. sp.
Description. With charactc-rs of genus;
form average; color brownish i)ice()us, mar-
gins of jirothorax and elytra slighth' or not
rufescent, appendages testaceous, antennae
slightly browner except at base; shining,
reticulate microsculpture faintly indicated,
meshes scarcely distinct at 50 X. Head
0.64 and 0.65 width prothorax; eyes mod-
erate, genae short, rounded. Prothorax:
width length 1.34 and 1.34; base apex 1.26
and 1.27; sides broadly rounded except
nearly straight and converging posteriorly
to obtuse but finely denticulate posterior
angles; base and apex unmargined at least
at middle; base punctate at sides, not at
middle. Elytra: width elytra prothorax
1.36 and 1.34; humeri broadh' rounded.
Secondary .sexual characters as for genus.
Measurements: length 3.5-4.3; width 1.6-
1.8 mm.
Types. Holotype i (M.C.Z., Type No.
31.363) and 20 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington), and
9 paratypes, Oro Bay, near Dobodura, Dec.
1943-Jan. 1944 (Darlington).
Measured specimens. The i holotype and
1 9 paratype from Dobodura.
Notes. See Notes under preceding and
following species.
Lecanomerus latior n. sp.
Description. With characters of genus;
form ( Fig. 18 ) compact, relati\ely broad;
black, appendages brownish testaceous;
shining, c. without reticulate microsculpture.
Head 0.68 and 0.67 width prothorax; eyes
slightly larger than in nwdius (above),
genae short, forming c. right angles with
neck. Prothorax: width length 1.36 and
1.39; base apex 1.24 and 1.16; sides broadl\-
rounded except c. straight and conxerging
posteriorly to obtuse but minutel\ dentic-
ulate posterior angles; apex margined but
marginal line sometimes laint at middle;
base not margined; entire base ol pionotum
punctate, but punctures sparser at middle
ol base. Elytra: width el\tra prothorax
1.44 and 1.38; humeri obtuseK' sometimes
vagueK- subangulate. Measurements: length
3.6-3.7; width 1.6-1.7 mm.
Tyj)es. Holotype 6 (M.C.Z., Type No.
31.364) and 1 9 paratope from l^obodura.
The Carabid Beetles of Ne\\' Guinea • Darlington 47
Papua, Mar -July 1944 (Darlington); and
additional paratypes from Papua as fol-
lows: 1, Bisianumu, near Sogeri, 500 ni.
Mar. 15-20, 1955 (E. O. Wilson, M.C.Z.),
taken in rain forest; 1, Kokoda, 1200 ft.
( 366 m ) , Aug. 1933 ( Cheesman ) .
Measured speeimens. The 6 holotype and
$ paratype from Dobodura.
Notes. Distinguished from the 2 pre-
ceding species by broader form, black
color, and pronotum with entire apical
marginal line and more extensive basal
punctation.
Genus CHYDAEUS Chaudoir
Chaudoir 1854, Bull. Soc. Nat. Moscow 27, Part 1,
p. 343.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 6,
p. 1080.
Diagnosis. See Key to Genera of Harpalini
of New Guinea.
Description. None required here.
Tijpe species. C. obscurus Chaudoir, of
India.
Generic distribution. Tlie Himalayas
(Sikkim, etc.), Formosa, Sumatra, Java,
the Philippines, and New Guinea; usually
at high altitudes.
Notes. This is the clearest case I know of
an Asiatic stock of Carabidae that has
"mountain hopped" to New Guinea. All
species of the genus are generally similar
and probably closely allied. The wings of
some species have atrophied, but those of
others are still fully developed, and C.
bakeri Andrewes is dimorphically winged
at Baguio on Luzon. Flying individuals
may therefore have dispersed from moun-
taintop to mountaintop and from island
to island across the Malay Archipelago
rather recently, in terms of evolutionary
time.
Chydaeus papua n. sp.
Description. Form as in Figure 19, stout,
convex; black, legs brownish, antennae and
mouthparts irregularly brownish testaceous;
both sexes moderately shining, upper sur-
face irregularly punctulate but reticulate
microsculpture faint or absent. Head 0.76
and 0.74 width prothorax, c. as in Chydaeus
obscurus Chaudoir; mentum toothed; ligula
free at apex, truncate; paraglossae arcuate,
narrow, c. long as ligula but widely sepa-
rated from it. Prothorax broadly subcordate;
width/length 1.48 and 1.49; base apex 1.12
and 1.09; sides broadly rounded through
much of length, sinuate before well defined
c. right posterior angles; pronotum strongly
convex (more so than in obscurus), base
margined, basal impressions poorly de-
fined, surface of disc more closely and
coarsely punctate at sides and especially
base than at middle. Ehjtra: width elytra/
prothorax 1.21 and 1.25; humeri subdentate;
apices weakly sinuate; striae entire, rather
lightly impressed; 3rd intervals without
dorsal punctures. Inner wings vestigial.
Lower surface and legs without obvious
special characters. Secondary sexual char-
acters: S front tarsi moderatelv and middle
tarsi narrowly dilated, densely squamulose
below; 6 with 1, 9 2 setae each side last
ventral segment. Measurements: length
9.2-10.6; width 3.6-4.4 mm.
Types. Holotype i (M.C.Z., Type No.
31,365) and 6 paratypes from Mt. Wilhelm,
Bismarck Rge., N-E. N. G., above 10,000
ft. (above 3000 m), Oct. 1944 (Darlington),
in open country above timberline; and
additional paratypes as follows, all from
the Bismarck Rge.: 2, Mt. Wilhelm, 2800-
2900 m, July 6, 1963 (Sedlacek); 1, "No. 5,"
Piunde-Aude Camp, east slopes Mt. Wil-
helm, June 13, 1959 ( L. J. Brass, Sixth
Archbold Exp. to Papua, A.M.N.H.); 1,
Lake Aunde, 3400-3500 m, July 4, 1963
(Sedlacek); 1, Lake Sirunki, 2800-2900 m,
June 15, 1963 (Sedlacek); 6, Mt. Otto Sum-
mit, Nov. 1965 ( Dept. Agr. Port Moresby ) .
Additional material. One, Murray Pass,
Papua, 2400-2800 m, Nov. 6, 1965 (Sed-
laceks); 1 i , Camp E. of Mt. Wilhelmina,
Snow Mts., West N. G., 3600 m, Sept. 1938
( Toxopeus ) .
Measured specimens. The 6 holotype and
1 ? paratype from Mt. Wilhelm.
48 BuUctin Museum of Comparative Zoology, Vol. 137, No. 1
Notes. This geographically isolated Chy-
daeus is similar to obscurtis Chaudoir (of
Sikkini, etc.) but has a slightly wider head
and differs in other details.
The Snow Mts. specimen may represent
an independent population, distinguished
by wider prothorax and perhaps by other
characters, but more material is necessary
to decide this.
Genus PLATYMETOPUS Dejean
Dejean 1929, Species General Coleop. 4, p. 68.
Csiki 1932, Coleop. Cat., Carabidae, Ilaipalinae 6,
p. 1205 ( see for synonymy and additional refer-
ences ) .
Schanberger 1938, Arl^eiten niorphologische nnd
taxonomische Ent. 5, p. 41 (see for comments
on some species of the Malay Archipelago ) .
Basilewskv 1950, Ann. Mus. Congo Bcfge (8),
Zool.. 6. p. 141.
Dkiii,nosis. Medium-sized, dull black Ilar-
palini distinguished from all other members
of the tribe in New Guinea by dorsal surface
entirely coarsely punctate and pubescent.
Description. None required here.
Tij))c -species. P. vestitus Dejean, of
Africa.
Generic distribution. Africa, the Cape
Verde Islands, and Madagjascar; SE. Asia,
Japan, and the Malay Archipelago to the
Philippines and New (iuinea (not Aus-
tralia ) .
Notes. A single widely distributed species
of this genus reaches New Guinea.
Plafymetopus iaficeps Dejean
Dejean 1829, Species Ceneral Coleop. 4, p. 70.
Csiki 1932, Coleop. Cat., (^aral)idae, Ilarpalinae 0,
p. 1200 (see tor additional rcterenccs and for
"varieties" ) .
Description. None re([uired here. See
DiciiS.nosis of genus. Length r. 8 mm.
Type(s). From the Philippines; in
Obcrthiir Coll., Paris Mus. (not seen).
Occurrence in Netv Guinea. Gollectcnl
only in the western part of West N. (i.:
4, Biak Is., dates in Jan., Feb., Mar., Apr.
1952 (L. 1). Bronger.sma, Leiden Mus.), at
light; 2, Wong R., Feb. 9, 1957 ( 1{. T. Simon
Thomas, Louwerens Coll.), at liirht; 1,
Sorong-Doom, Feb. 9, 1957 (R. T. Simon
Thomas, Louwerens Coll.), at light.
Notes. Closely related fomis of this
genus, some treated as varieties of Platy-
metopus jlaiiJahris (Fabricius) by Csiki,
are widely distributed in SE. Asia and the
Malay Archipelago. Their taxonomy is a
problem. The problem, however, lies mainly
in the Oriental Region rather than New
Guinea, and I cannot undertake to solve it
now.
Whatever the final taxonomic arrange-
ment, it seems clear that one, dark-legged
form of Phitymetopus (all surely New
Guinean individuals are dark-legged ) has
reached New Guinea recently from the
west and may perhaps still be confined to
the western end of the island. Its absence
elsewhere in New Guinea is suggested by
the facts that members of this genus are
usually common where they occur at all
and that they fly to light, but that none
has been found in light trap material from
central and eastern New Guinea. P. hiti-
ceps has been prexiously known from Bnrn
( specimen in Andrewes Coll. ) and the
Philippines.
Besides the dark-legged indixiduals re-
corded above, I have seen two yellow-
legged ones labeled "Dor)" and "Dorey."
They were probabK' collected by Wallace
and are presumably really from ('elebes
(see Part I of the present work, pp. 330-
331). They arv probably referable to P.
sidjrugosus Schanberger (see reference cited
under genus, above ) of Celebes. This
species should not be listed from New
Guinea.
Genus TRICHOTICHNUS Morawitz
Nh)ra\vil/. 1803, Mem. Acad. Sci. St. Petersburg
(7) 0, No. 3, p. 03.
Csiki 1932, Coleop. (>at., C^araliidac, Ilarpalinae
0, pp. 1210, 1217 (see for additional refer-
ences, subgenera, and synonx'nn).
l^asilcwskx 1950, .\nii. Mus. C'ongo Beige, Zool.
0, p. 87.
Di(iii,nosis. See Key to Gei\era of IlarpaJini
of New Gui)u'(i.
The Carabid Beetles of New Guinea • Darlington
49
Description ( important characters shared
liy New Guinean species). Form of or-
dinary, medium-sized Harpalini; upper sur-
face not pubescent. Head smooth except
for deep, obhque, usually linear frontal
impressions; 1 seta over each eye; mentum
toothed; labial palpi with penultimate seg-
ments with more than 2 setae. Prothoiox
subcordate or transverse; side margins each
with 1 seta-bearing puncture, before mid-
dle; disc usually extensively punctate espe-
cially across base, with punctation finer and
usually sparser across middle. Elytra with
striae entire, impressed, impunctate; 3rd in-
tervals 1-punctate near inner edge at or
slightly behind middle ( punctures some-
times obscured or absent on one or both
elytra). Inner wings usually full, rarely
dimorphic (some populations of nigricans
and altiis). Lower sw^acc: prosternum
anteriorly with short pubescence (reduced
in mcdiiis). Legs: front tibiae with apex
less than Vi wide as tibial length; hind
tarsi moderate or long. Secondary sexual
characters: 6 front and usually middle tarsi
2-seriately squamulose; 2 setae each side
last ventral segment in both sexes. See also
Notes, below.
Type species. T. longitarsis Morawitz, of
Japan.
Generic distribution. Temperate and
tropical Eurasia and the Malay Archipel-
ago to New Guinea, etc. (probably not
Australia); eastern North America. The
genus in a broad sense, including Hyparpa-
his, occurs also in Africa (Csiki, probably
following Schauberger), but Basilewsky con-
siders llyparpalus a separate genus and
does not recognize Trichotichnus in Africa
south of the Sahara.
Notes. I have had difficulty with both
the generic and the specific classifications
of the 15 New Guinean species that I now
assign to this genus. Several of the species
might go in Lampetes {Lamprophonus) or
Carbanus, but I have not found satisfactory
characters to distinguish these genera from
Trichotichnus. However, I do not intend to
reduce them to synonymy now. They need
further study based on Oriental as well as
New Guinean forms. This study will re-
quire more time and material than I now
have.
The descriptions of species in the follow-
ing pages are brief, and allowance must be
made for individual variation, which is
surprisingly great in some characters. For
example, the punctation of the outer elytral
intervals is variable in some cases (e.g., in
mixtus). The form of the elytral apices is
sometimes variable (e.g., in denarius and
altus). And the form of the apex of the
aedeagus is surprisingly variable in some
species. I have figured it in some cases
but have usually not used it as a diagnostic
character. But see under alius and dux
(Figs. 172, 173).
All New Guinean species of Trichotichnus
are fully winged and probably capable of
flight (some of them have been taken in
light traps ), with 2 exceptions. T. nigricans,
although apparently always fully winged
at low altitudes, is dimorphically winged
on the Bismarck Range. And some pop-
ulations of T. alius include individuals with
slightly shortened and weakened wings, al-
though other individuals of this species fly.
Some Trichotichnus in other parts of the
world have atrophied or dimorphic wings.
Most or all of the common, unspotted
Trichotichnus in New Guinea probably live
on the ground in rain forest, but I did not
distinguish the different species in the field
and cannot be sure of their exact habitats.
The following Key to Species of Tri-
chotichnus of New Guinea works reason-
ably well for series of clean specimens, but
it is not perfect. I myself have had trouble
placing some single specimens. In order to
simplify identifications and reduce need for
using the key, I give the following notes for
recognizing several of the commoner, dark
( unmarked ) species.
If large (7.3-S.3 mm), rufo-piceous,
shining, and without pubescence at front of
prosternum (but some setae at apex of
50 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
prosternal process ) : see Lyter, second
genus after TrichoticJinus.
If large (8-9 mm), broad, and with
paiiiol raised 10th intervals in elytral mar-
'j^ins: denarius.
If large ( c. 8-10 mm ) , less broad, with-
out raised 10th intervals, and found at con-
siderable elevations in mountains (usually
over 1200 m ) : probably altus.
If small ( 6.5-7.5 mm ) , dark, and without
reticulate microsculpture on elytral inter-
vals: probably nigricans.
If small (6.3-7.0 mm), dark, and with
reticulate microsculpture on elytral inter-
vals: probably semimas (which lacks
squamules on 6 middle tarsi ) .
Key to Species of Trichotichnus of
New Guinea
1. Head c. % width prothorax (H/P 0.66 and
0.68); prothorax transverse with broadly
rounded sides; length 5.0-6.3 mm (p.
50 ) siraneoi
- Head c. % or more width prothorax (by
measurement); prothorax more subcordate;
usually larger 2
2. Elytra with partial raised 10th intervals in
marginal channels; (no dorsal markings;
length 8-9 mm) (p. 51) denarhi.s
- Elytra without partial raised 10th inter-
vals 3
3. Male with only anterior (not middle)
tarsi squamulose; elytral intervals micro-
reticulate; ( no dorsal markings; length 6.3-
7.0 mm) (p. .52) .semimas
- Male with middle as well as anterior tarsi
with sf|uamides; elytra often (not always)
without microrcticuhition 4
4. Elytra without subapical sutural spot or
sutural intervals pale 5
- Elytra with eonnnon subapical sutural spot
pale, or sutural intervals pale near apex 11
5. Prothorax and elytra without pale margins;
abdomen usually without (•(>)is))icu()us pale
spots or margins; femora not coii.s))icuotisly
paler than abdomen 6
- Protliorax and elytra usually with narrow
pale margins; abdomen usually with con-
spicuous pale lateral spots or margins;
femora conspicnotisUi pale 10
6. Length 6. .5-7. 5 mm; pronotum not iiiucli
depressed at sides toward base; (basal
margin of pronotum usually incomplete)
(p. 52) )u^iic(nis
- Larger, or sides of pronotum more de-
pressed toward base 7
7. Eyes slightly larger, separated from mouth
below by c. '/s diameter of an eye; apex
of aedeagus short; (length c. 8-10 mm)
(p. 53) modiLs
- Eyes slightly smaller, more distant from
mouth below; apex of aedeagus longer,
slender 8
8. Length lL.5-13.0 nun (p. ,53) dii.x
- Length 7.2-10.0 mm 9
9. Prosternal pubescence more abundant; size
usually larger (c. 8-10 mm); pronotum
more punctate, less shining (p. 54) _ (dtus
- Prosternal pui^escence usualK' scanty, but
variable; size usually smaller (7.2-8.5
mm); pronotum less punctate, more shining;
(direct comparison necessary to determine
some specimens) (p. 55) mediii.s
10. Elytral intervals not obviously microreticu-
late; abdomen with pale spots usually
largest and most conspicuous at sides of
subapical segment; (length 8.3-9.3 mm)
(p. .56) hrcnidti
- Elytral intervals microreticulate; abdomen
usually more extensively pale margined;
(length 8..5-9.4 mm) (p. .56) ohscurus
11. Length usually 7.6-8.5 mm ( rarely slightly
smaller); subapical sutural pale spot dis-
tinct, reaching 3rd intervals (p. 57) _
guttuhi
- Smaller; sutural pale spot \ariable, some-
times smaller or less distinct 12
12. Prothorax wide at base ( base, ape.\ 1.34);
(length 7.5 nun) (p. 57) mongi
- Prothorax narrower at base 13
13. Sides of elytra (intervals 8, 9) rugose-
punctate; (length 5.3-6.8 mm) (p. 58) __
semirugosiis
- Sides of elytra not rugose, although some-
times punctulate 14
14. Length 5.8-7.5 mm; pronotum depressed
at sides toward base; ( and see Notes under
this species) (p. 58) mixtu.s
- Length .5. .3-5. 8 mm; sides of pronotum
scarcely deprcssi'd (p. 59) dclicatu.s
Trichotichnus sfraneoi (Louwerens)
Louwercus 1962, Tijdschrift \c)or Ent. 105, p. 1 12,
iig. 7 {C(irhanu.s).
Dc.scri))lion. With charactt^rs of genus;
small, form (Fig. 20) broad; brownish
black, sides ol piouotum and c']\ tr;i \aguely
translucent, lower snrlacc and appendages
more rufous; elytr;i fainth- iridescent but
not distinctK' microreticulate (at 50x).
Head small, 0.66 and 0.68 width prothorax;
eyes ku-gc. s('p;uated from month below b\
The Carabid Beetles of New Guinea • Darlington
51
c. ^i-j or less width of an eye. PiotJiomx
transverse with broadly rounded sides;
width length 1.59 and 1.59; base/apex 1.41
and 1.39. Elytra: width elytra prothorax —
and 1.30. Secondary sexual characters nor-
mal for Trichotichnus. Measurements (in
New Guinea ) : length c. 5.0-6.3; width c.
2.3-2.7 mm.
Types. Holotype $ (Louwerens Coll.),
allotype, 6 paratypes all from Amboina Is-
land^ Moluccas, 70 m (A. M. R. Wegner),
at light; a paratype now in M.C.Z. (Type
No. 31,149) (holotype not seen).
Occurrence in New Guinea. Papua: 3,
Brown R., May 21, 23, 24, 1956 ( E. J.
Ford, Jr., Bishop Mus.); 1, Mts. between
Agamoia and Ailuluai, Ferguson Is., 900 m,
"No. 4," June 5-17, 1956 (L. J. Brass,
U.S.N.M.). N-E. N. G.: 13, Wau, Morobe
Dist., 1200, 1300 m, various dates (Sedla-
cek); 1, Wantoat, Finisterre Mts., 4000 ft.
(1220 m), Sept. 9, 1957 (Monroe and
Holland, Canadian Nat. Coll.); 1, Elipta-
min Vy., 1350-1665 m, June 23-30, 1959
(W. W. Brandt, Bishop Mus.). West
N. G.: 1, Hollandia, July-Sept. 1944 (Dar-
lington); 1, Cyclops Mts., Sabron, 2000 ft.
(610 m), June 1936 (Cheesman); 1, Hol-
landia area, W. Sentani, Cyclops Mts., 50-
100 m, June 22-24, 1959 (Gressitt & T. C.
Maa, Bishop Mus.), in light trap; 2, Star
Rge., Sibil, 1260 m, May 16, June 16, 1959
(Leiden Mus.), at Hght.
Measured .specimens. A pair ( i 9 ) from
Brown R.
Notes. Andrewes placed Carhanus in the
wrong subtribe of Harpalini, erroneously
considering it a member of the Acupalpina.
The arrangement of setae on the labial palpi
seems to me to place it with the Haipalina,
and I can find no positive character to
distinguish it from Trichotichnus. How-
ever, I do not intend to synonymize Car-
hanus now. It requires further study. The
name can be used for a group of small,
mutually similar species (lautus Andrewes
of Burma, flavipes Andrewes of Java,
pliilippinus Jedlicka of the Philippines, and
straneoi Louwerens of the Moluccas, New
Guinea, etc. ) that may eventually be sepa-
rated from Trichotichnus.
T. straneoi extends to New Britain and
New Ireland (specimens in Bishop Mus.).
Nothing is recorded of its habitat or habits
except that it flies to light.
Trichotichnus denarius n. sp.
Description. With characters of genus;
form slightly broader than usual; black or
piceous, appendages browner, sides of ab-
domen with small pale marks (variable);
rather shining, most of upper surface with-
out visible microreticulation (at 50x ) but
elytra silky in some lights. Head 0.77 and
0.77 width prothorax; eyes large, separated
from mouth below by e. ^o diameter of
an eye. Frothorax transverse-subcordate;
width/length 1.59 and 1.63; base/apex 1.14
and 1.10; sides converging and usually
broadly and slightly sinuate before distinct
but obtuse posterior angles; basal marginal
line fine or interrupted at middle; disc
weakly convex, moderately depressed at
sides especially posteriorly, extensively
punctate but with punctation finer and
slightly sparser at middle. Ehjtra broad;
width elytra/prothorax 1.36 and 1.36; mar-
ginal gutters wider than usual, with ir-
regular raised 10th intervals in c. middle
V3 of length; apices usually slightly dehis-
cent and c. pointed, but variable; outer
intervals (8, 9, and 10) usually slightly,
finely punctulate. Secondary sexual char-
acters as for genus. Measurements: length
c. 8.0-9.0; width 3.3-3.8 mm.
Types. Holotype S (M.C.Z., Type No.
31,366) and 111 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
additional paratypes as follows. Papua:
20, Kokoda, 1200 ft. (366 m), June, July,
Aug., Sept., Oct. 1933 (Cheesman); 1, same
locality, Mar. 28-29, 1956 (Gressitt); 2,
Biniguni, Gwariu R., 150 m, "No. 3," July
27-Aug. 14, 1953 (Geoffrev M. Tate,
A.M.N.H.); 5, Peria Ck., Kwagira R., 50 m,
"No. 7," Aug. 14-Sept. 6, 1953 (Geoffrey
M. Tate, A.M.N.H.); 1, Kokoda-Pitoki, 400
m, Mar. 23, 1956 (Gressitt). N-E. N. G.:
52 BiiUefin Museum of Comparative Zoology, Vol. 137, No. 1
3, lower Busu R., Huon Pen., May 4, 1955
(E. O. ^^'ilson, M.C.Z.), in lowland rain
forest; 1, Simbang, Huon Gulf, 1899 (Biro).
West N. G.: 1, Wamoro (ex Coll. G.
Hauser, British Mus.).
Measured .speeimens. The c5 holotype and
1 9 paratype from Dobodura.
Notes. The partial 10th intervals in the
elytral margins immediately distinguish
denarius. The species is very common in
eastern New Guinea ( apparently much
less so in the west), presumably in ram
forest. One of Miss Cheesmans specimens
was taken at light and so apparently were
the Peria Creek individuals, which have
scales and wing fragments of other insects
on them.
Trichofichnus semimas n. sp.
Description. With characters of genus;
form rather slender; black or piceous, ap-
pendages brown, abdomen with some ( vari-
able) small pale marks at sides; moderately
shining but elytra with transverse micro-
reticulation distinct at 50x. Head 0.79 and
0.79 width prothorax; eyes moderate, sepa-
rated from mouth below by nearly % diam-
eter of an eye. Prothorax subcordate;
width length 1.48 and 1.48; base/apex 1.18
and 1.16; sides rounded anteriorly, con-
verging and sinuate before distinct c. right
(slightly obtuse) posterior angles; basal
marginal line faint or interrupted at middle;
disc very little depressed at sides toward
base, extensively punctate except almost
impunctate at middle. Ehjlra normal;
width elytra prothorax 1.23 and 1.26; outer
intervals not distinct])' punctulate. Second-
ary sexual characters normal except only
front (not middle) tarsi of 6 scjuamulose.
Measurements: length 6.3-7.0; width 2.5-
2.8 mm.
Tijpes. Holotype i (M.C.Z., Type No.
31,367) and 19 paratypes from Dobodura,
Pajma, Mar.-July 1944 (Darlington); 4
paratypes, Kokoda, Papua, 1200 ft. (366
m), Oct., Sept. 1933 (Cheesman); 3 para-
types, same locality. Mar. 20, 28-29, 1956
(Gressitt), in light trap; 2, Xormanb) Is.,
W'akaiuna, Sewa Bav, Nov. 1-10, 1956 and
Jan. 1-8, 1957 (W. \\'. Brandt, Bishop
Mus.). N-E. N. G.: 1, Erima, Astrolabe
Bav, 1896 (Biro); 2, Madang ("Friedrich-
\^m.-hafen"), 1901 (Biro); 1, Bulolo, 730
m, Aug. 27, 1956 (E. J. Ford, Jr., Bishop
Mus.), in light trap; 1, Sum-Sum, 64 km N.
of Wan, 580 m, Feb. 15, 1963 (Sedlacek).
West N. G. : 4, Hollandia area, W. Sentani,
Cyclops Mts., 50-100, 100, 150-250 m, June
(various dates) 1959 (Gressitt and T. C.
Maa, Bishop Mus.); 2, Ifar, Cyclops Mts.,
450-500, 400-800 m, Sept. 7,' 7-9, 1962
( Sedlacek ) ; 3, Dojo, Res. Hollandia, Apr.
1957, 1958 (R. T. Simon Thomas, in Lou-
werens Coll.); 1, Maffin Bay, Aug. 1944
(E. S. Ross, California Acad.); 1, Wasian,
Vogelkop, Sept. 1939 (Wind. M.C.Z.).
Measured specimens. The i holotype and
1 9 paratype from Dobodura.
Notes. Males of semimas are imique
among New Guinean Trichofichnus in
lacking the usual sexual squamules of the
middle tarsi. Fc^nales resemble nii^ricans
( below ) but differ by presence of reticuhite
microsculpture on the elytra. T. semimas
probably lives in rain forest, and has been
taken at light near Hollandia as wi-ll as at
Kokoda.
Trichofichnus nigricans Schauberger
SLlianhermT 1935, Eiit. An/.cimT 15, p. 34.
De.ivription. \\'ith characters of genus;
form rather small and slender; black,
appendages brownish, sides of abdomen
with or without small, often vague pale
marks; upper surlace including elytra with-
out distinct reticulate microsculpture. Head
0.76 and 0.75 width jirothorax; eyes usually
moderati- and separated Irom mouth below
by c. ':. diam(4er ol an c\c. but e\"es smaller
and more distant Irom mouth in short-
wing(>d indi\iduals hom Bismarck Range.
Prothorax subcordate. with sides slightK
or not sinuate belore usnalK well defined
but obtuse basal angles; width IcMigth 1.43
and 1.49; base apex 1. 2 1 and 148; basal
inaruinal line usualK ineomplete at middle;
The Carabid Beetles of New Guinea • Darlington
53
disc only slightly depressed at sides basally,
extensively punctate, punctation finer and
sparser at middle. Elytra nomial; width
elytra prothorax 1.27 and 1.25; outer inter-
vals not distinctly punctulate. \V/ng.s full
in lowland populations, dimorphic on Bis-
marck Range (see Notes, below). Second-
ary sexual characters normal. Measure-
ments: length c. 6.5-7.5; width c. 2.5-2.8
mm.
Types. (Holo)type (Andrewes Coll.,
British Mus.) and 2 "cotypes" all from
Sattelberg, N-E. N. G. (G. Hauser). I saw
the type in London in 1948 and made a
satisfactory comparison with it.
Occurrence in Neiv Guinea. Common
and widely distributed: 180 specimens
from localities well distributed over New
Guinea and on Woodlark and Nonnanby
Is.; most from low altitudes, but series from
Chimbu Vy., Bismarck Rge., between 5000
and 7000 ft. (c. 1500-2100 m), and from
Wau, 1200 m. Specimens taken in every
month.
Measured specimens. A pair { $ 9 ) from
Dobodura, Papua.
Notes. T. nigricans occurs also on New
Britain, and related fonns, compared by
Schauberger (1935, p. 36), are known from
Java, Sumatra, and Celebes.
This species probably occurs in rain
forest, although the Chimbu specimens
were taken in open country.
I have not examined the \\'ings of every
specimen, but all or most of those from low
altitudes are winged, and they often fly.
They have been taken in light traps at
several localities. However, of my Chimbu
series, only 3 specimens (both sexes) have
full wings, and 11 (both sexes) have the
wings reduced to thin strips c. % as long
as the elytra. I have not examined the
wings of the 41 specimens from Wau be-
cause some or all were collected in light
traps, which would select only winged in-
dividuals.
Besides the 180 specimens that I assign
to this species without much doubt, 7
specimens from various localities in New
Guinea are assigned doubtfully, because of
slight differences in various characters.
Trichotichnus modus n. sp.
Description. With characters of genus;
form ( Fig. 21 ) average, somewhat variable
( see proportions ) ; black, legs brownish, ab-
domen ± brownish (apical segment darker)
with lateral pale areas absent or not sharply
defined; shining, elytra subiridescent but
without visible reticulate microsculpture.
Head 0.72 and 0.76 width prothorax; eyes
slightly larger than usual, separated from
mouth below by c. Vs diameter of an eye. Pro-
thorax transverse-subcordate; width/length
1.43 and 1.53; base/apex 1.30 and 1.20;
sides rounded anteriorly, converging and
usually slightly sinuate before well defined
but obtuse posterior angles; basal marginal
line faint or incomplete at middle; disc
moderately depressed at sides, extensively
punctate except c. impunctate at middle.
Elytra: width elytra /prothorax 1.32 and
1.36; marginal channels narrow, without
10th intervals; outer intervals not distinctly
punctulate. Secondary sexual characters
normal; apex aedeagus short, ± hooked
dorsally. Measurements: length 8.0-10.0;
width 3.4-3.9 mm.
Types. Holotype S (M.C.Z., Type No.
31,368) and 6 paratypes all from Dobodura,
Papua, Mar.-July 1944 (Darlington).
Additional material. Papua: 1, Nor-
manby Is., Wakaiuna, Sewa Bay, Jan. 1-8,
1957 (W. W. Brandt, Bishop Mus.). West
N. G.: 1, Wissel Lakes, Tage L., 1760
m, Aug. 4, 1955 (Gressitt). These speci-
mens are assigned to modus doubtfully.
Measured specimens. The c5 holotype and
1 9 paratype.
Notes. This species resembles denarius
in size and color, but differs by lack of
partial 10th elytral intervals.
Trichotichnus dux n. sp.
Description. With characters of genus;
fonn c. average, very large; black, append-
ages dark brown, abdomen without distinct
pale areas; upper surface finely micro-
reticulate, meshes c. isodiametric on head,
54 Bulletin Museum of Comparative Zoology, Vol 137, No. 1
increasingly transverse on pronotum and
elytra. Head 0.76 and 0.77 width prothorax;
eyes separated from mouth below by c.
% width of an eye (but eyes more deeply
covered than usual ^^'ith transparent win-
dow-like material so edges not precisely
defined). Fiothorax subcordate; width/
length 1.41 and 1.46; base/apex 1.19 and
1.23; sides rounded anteriorly, converging
and straight or slightly sinuate before well
defined but obtuse posterior angles; basal
and apical marginal lines usually faint or
interrupted at middle; disc moderately de-
pressed at sides posteriorly, finely but ex-
tensively punctate or punctulate, the punc-
tation strongest basally. Elytra: width
elytra/prothorax 1.39 and 1.30; margins
narrow; outer intervals not distinctly punc-
tulate. Inner icings full (see following
Notes). Secondary .sexual characters nor-
mal; apex aedeagus as in Figure 173. Mea-
.mrements: length 11.5-13.0; width 4.4-
5.1 mm.
Types. Holotype $ (Bishop Mus.) and
4 paratvpes (2 in M.C.Z., Type No. 31,369),
from Edie Creek, 14 km S^^^ of Wau, N-E.
N. G., 1900 and 2000 m, Oct. 4-10, 1961,
and Feb. 13, 1962 (Sedlacek). Additional
paratypes from N-E. N. G.: 3, Wau, 1700,
2400 m, Jan. 9-12, Oct. 6, 1962 ( Sedlacek ) ;
6, Kepilam, 2400 and 2500 m, June 21, 20-
22, 21-23, 1963 (Sedlacek); 1, Tambul, 2200
m, May 27-June 7, 1963 (Sedlacek); 1,
Laiagam, W. Highlands, Mar. 23, 1960 (J.
H. Barrett, Dept. Agr. Port Moresby), at
light; 1, Moke, Okapa Subd(istrict), E.
Ibghlands, 6400 ft. ( 1950 m), Apr. 17, 1962
(J. II. Barrett, Dept. Agr. Port Moresby).
2, Okapa, June 12, 1964, Jan. 10, 1965
(Hornabrook).
Mea.surcd specimens. The i holotype and
1 9 paratype from Edie Creek.
Notes. The large size distinguishes this
species from all odiers of the genus in New
Guinea. All specimens of the type series
are fully winged, but most were taken in
light traps which, of course, select winged
individuals.
Trichotichnus alius n. sp.
Description. With characters of genus;
form of c. average large New Guinean
Trichotichnu.s; black or piceous, appendages
dark brownish, sides of abdomen with in-
distinct or poorly defined (variable) pale
areas; moderately shining, elytra with trans-
verse microreticulation faintly or not visible
at 50x. Head 0.76 and 0.75 width prothorax;
eyes separated from sides of mouth below
by c. % width of an eye. Prothorax sub-
cordate; width length 1.40 and 1.50; base
apex 1.15 and 1.18; sides converging and
straight or slightly sinuate before well de-
fined but obtuse posterior angles; disc mod-
erately depressed at sides posteriorly, vari-
ably but often extensively punctate, most
conspicuously so across base and least so
across middle; basal marginal line entire,
faint, or interrupted at middle (variable).
Ehjtra: width elytra prothorax 1.25 and
1.31; margins narrow, without 10th inter-
vals; apices pointed or blunted (variable);
outer intervals not distinctly punctulate.
Inner wings dimorphic on the Bismarck
Range (type series), full and strong in
some individuals, slightly shortened (but
still folded at apex) and with slightly
weakened venation in other indixiduals.
Secondary sexual characters normal; i cop-
ulatory organs as in Figure 172. Measure-
mcnts: length 8.0-10.3; width 3.3-3.9 mm.
Types. Holotype 6 (M.C.Z., Type No.
31370) and 42 paratvpes from Chimbu
Vy., Bismarck Rge., N-E. N. G., 5000-7500
ft. (c. 1500-2300 m), Oct. 1944 (Darling-
ton); and 16 paratypes, Tomba, S. .slope of
Mt. Ilagen (Bismarck Rge.), 2450 m. May
22-24, 1963 ( Sedlacek ) .
Additional material. See Notes, below.
Measured specimens. The 6 holotype and
1 9 paratype from Chimbu Vy.
Notes. T. altus and its allies {dux, above,
and medius, below) are the common moun-
tain-living Tricliotichnus of New Guinea.
These 3 species seem clearU distinct and
have differcMit, but in part o\(>rlapping,
ranges: dux, on the mountains of the
The Carabid Beetles of New Guinea • Darlington 55
Morobe area; typical alius, on the Bismarck traps, and such material usually includes
Range; and typical medius, on the Torri- only fully winged individuals and is not
celli Mts. However, I have seen many satisfactory for study of wing dimorphism,
additional specimens of alius or closely This is another reason for not attempting
related forms from other localities, as fol- a more detailed study of alius and related
lows. Papua: 9, Mt. Giluwe, 2500 m. May species with the material available now.
1 and 27, June 6, 1963 (Sedlacek); 1,
Dimifa, SE. of Mt. Giluwe, 2200 m, Oct. Trichotichnus medius n. sp.
11, 1958 (Gressitt); 1, Owen Stanley Rge.,
Goilala, Bome, 1950 m. Mar. 8-15, 1958 Description. With characters of genus;
(W. \y. Brandt, Bishop Mus.); 1, Mafulu, form average; black, appendages brownish
4000 ft. (1220 m), Jan. 1934 (Cheesman). testaceous, abdomen with or without poorly
N-E. N. G.: 270 specimens (in addition to defined pale lateral areas (variable), hind
the type series) from localities including femora not sirikino^ly paler than abdomen,
Morobe Dist.; Kratke Mts.; W. Highlands; tibiae paler than femora; shining, elytra
and (S. of the Markham-Ramu Vy.) Sala- not visibly microreticulate at 50x. Head
waket Rge; Mongi Watershed; and Huon 0.78 and 0.79 width prothorax; eyes sepa-
Pen. West N. G.: 35 specimens, from rated from mouth below by % or Vs diam-
localities including the Star Rge.; Wissel eter of an eye. Pwihorax subcordate; width/
Lakes; and Snow Mts. (Top Camp; lebele length 1.49 and 1.52; base/apex 1.18 and
Camp; Mist Camp; Baliem Camp). Most 1.14; sides converging and straight or
specimens are from altitudes of 1200 to slightly sinuate before well defined but
2700 m, but a few, from within 200 m of obtuse basal angles; fine basal marginal
sea level. Individuals have been taken in line usually complete; disc moderately de-
every month. Most specimens are in the pressed at sides basally, extensively punc-
British Mus., Bishop Mus., A.M.N.H., tate across base, punctation much finer
Leiden Mus., C.S.I.R.O. Coll., and M.C.Z. and somewhat sparser across middle and
I have restricted the type series of alius anteriorly. Elyira: width elytra prothorax
and its close relatives to specimens from 1,31 and 1.26; margins without 10th inter-
single localities or restricted areas because vals; outer intervals not distinctly punc-
the species of this group obviously vary tulate. Inner icings full. Lower sutiace:
geographically as well as individually. In flnfen'or part of prosternum with pubescence
general, specimens from north of the Mark- usually sparse (but variable). Secondary
ham-Ramu Valley average larger, those sexual characters normal. Measurements:
from south of the valley smaller, except length c. 7.5-8.5; width 3.1-3.3 mm.
that some specimens from Wau are as small Types. Holotype S ( Bishop Mus. ) and
as some of the types of medius. Two dis- 22 paratypes (some in M.C.Z., Type No.
tinct forms, a very large one (dux) and a 31,371) from Mokai Village, Torricelli Mts.,
smaller one (tentatively referred to alius), N-E. N. G., 750 m, various dates in Dec.
occur at Edie Creek, showing that the 1958 and Jan. 1959 (holotype, Jan. 1-23,
species of this group are not entirely al- 1959) (^^^ W. Brandt); and additional
lopatric. The characters, variation, and dis- paratypes, all from Torricelli Mts., as fol-
tribution of these species need more study lows: 19, Mobitei, 750 m, dates in Feb.,
than I can give them now. Mar., Apr. 1959 (W. W. Brandt, Bishop
T. alius is known to have dimorphic wings Mus. ) ; 3, Wantipi, Nov. 30-Dec. 8, 1958
(see Description) only on the Bismarck (W. W. Brandt, Bishop Mus.).
Range. The specimens in question were Additional material. Some specimens
not collected at light. Much of the other among those summarized under Trichoti-
material listed above was taken in light chnus alius may prove referable to medius,
56
Bulletin Museum of Coiupuratwe Zoologij, Vol. 137, No. 1
especially the smaller ones from south of
the Markham-Ramii Valley and from Wau.
Measured specimens. The 6 holotype and
1 9 paratype from Mokai.
Notes. This species is difficult to define
exactly. It is larger than niii,ric(U]s, with
basal marginal line of pronotum usually
entire (usually interrupted in ni'^ricans).
It is smaller than modus and (dtus, with
eyes intermediate in size. The sparseness
of pubescence on the anterior part of the
prosternum is an aid in identification, but
it is not infallible. This species is more
shining than most Trichofichnus, and ap-
proaches Lijter (p. 63) in appearance, but
the clothing of the male front and middle
tarsi of mcdius consists of 2 rows of broad
scales as usual in Trichotichnus. Neverthe-
less, a Tricliolicluius like the present one
may have been ancestral to Lytcr.
Trichotichnus brandti n. sp.
Description. With characters of genus;
form average; black or piceous, not marked
above except lateral margins of elytra and
sometimes of prothorax narrowly incon-
spicuously rufesccnt or translucent; reddish
piceous below with epipleurae and narrow
maigin of abdomen paler, the pale marks
usually widest and most conspicuous at
sides of subapical ventral segment; ap-
pendages brownish testaceous, femora con-
spicuously paler than abdomen; shining,
elytra without distinct reticukite micro-
sculpture. Head 0.75 and 0.76 width pro-
thorax; eyes separated from mouth below
by about 'c diameter of an vyv. Prothorax
transverse-subcordate; width/length 1 .43
and 1.45; base/apex 1.20 and 1.21; sides
converging and straight or slightly sinuate
before obtuse but well formed ( sometimes
slightly bhuited) basal angles; disc de-
pressed at sides near base, extensively punc-
tate, least so near middle; basal marginal
line faint or interrupted at middle. Klijlrtr.
width elytra prothorax 1.36 and 1.38; Stli
and 9th int(>rvals usually with a little sparse,
fine punctulation. Secotidanj sexual char-
acters normal. Measurements: length 8.3-
9.3; width 3.4-3.8 mm.
Ti/pcs. Ilolotvpe S (Bishop Mus.) and
3 paratypes (2 in M.C.Z., Type No. 31,372)
from Feramin, N-E. N. G., 1200-1500 m.
May 11-22 (holotype), 23-31, 1959 (W.
W. Brandt); and additional paratypes as
follows, all from N-E. N. G.: 3, Torricelli
Mts., Mobitei, 750 m. Mar. 16-31. Apr. 1-
15, 16-22, 1959 (W. W. Brandt, Bishop
Mus. ); 2, Eliptamin Vy., 1665-2530 m, June
23-30, and 2, same localitx, 1200-1350 m,
Aug. 16-30 and Sept. 1-15, 1959 (W. W.
Brandt, Bishop Mus.); 15, Wau, Morobe
Dist, 1100, 1200 (most), 1300, and 1700-
1800 m. Mar., Apr., and all months from
June to Dec., 1961-1963 (Sedlacek).
Additional material. The following addi-
tional specimens are tentatixely assigned
to this species. Papua: 1, Dogon, Amazon
Bay Dist., 2400 ft. (c. 730 m), Sept. 1962
(W. W. Brandt, C.S.I.R.O.); 1, Kokoda,
1200 ft. (366 m), July 1933 (Cheesman),
at light. N-E. N. G.: 1, Tuwep. Salawaket
Rge.\ 1350 m, Sept. 9, 1956 (E. J. Ford, Jr..
Bishop Mus.), in light trap. West N. G.:
1, Cyclops Mts., Sabron Camp 2. 2000 ft..
June 1936 (Cheesman); 1, "Neth. New
CTuinea' without further locality, Oct. 20.
1944 (T. Aarons, California Acad.).
Measured s})ecimens. The ^ holotype and
1 9 paratN'pe from Feramin.
Notes. See under lollowing species
(obscurus).
Trichotichnus obscurus n. sp.
Description. With characters of genus;
ionn broad-average; brownish piceous, lat-
eral margins of prothorax and cKtra \aguel\-
paler or translueeut; abdomen broadK' mar-
gin(>d with yellow, the \eIlow margins
widest anteriorh'; appendages testaceous,
hind leinora strikingK jiale; inodeiateK'
shining, elytia witli transNcrse reticulate
microsculpture \isible in both sexes (at
50 ■ ). Head 0.77 and 0.76 width prothorax;
eyes separated lioin mouth below by Vi
or less width o( an e\e. Prothorax trans-
The Carabid Beetles of New Guinea • Darlington 57
verse-subcordate; width length 1.40 and
1.44; base/apex 1.21 and 1.20; sides con-
verging, slightly, broadly sinuate before dis-
tinct but obtuse posterior angles; disc mod-
erately depressed at sides basally, extensively
but rather finely punctate except almost
impunctate at middle. Elytra: width
elytra/prothorax 1.34 and 1.27; 8th and 9th
intervals usually not distinctly punctulate.
Secondary sexual characters normal. Mea-
siiremeuts: length 8.5-9.4; width 3.3-3.7
mm.
Types. Holotype S (Bishop Mus.) and
2 paratypes (1 in M.C.Z., Type No. 31,373)
from Saidor, Matoko, Finisterre Rge., N-E.
N. G., Aug. 29-Sept. 5 and Sept. 6-24,
1958 (W. W. Brandt); and additional
paratypes as follows: Papua: 1, S. High-
lands^ Dimifa, SE. of Mt. Giluwe, 2200 m,
Oct. 11, 1958 (Gressitt). N-E. N. G.: 2,
Wau, Morobe Dist., 1200 m, Nov. 1-20 and
Dec. 1961 (Sedlacek); 7, Edie Creek, 14
km SW. of Wau, 2000 m, Feb. 13, 1962
(Sedlacek); 1, Eliptamin Vv., 1200-1350
m, Aug. 1-15, 1959 (W. W. Brandt, Bishop
Mus.).
Measured speciynens. The <^ holotype and
1 9 paratype from Saidor.
Notes. Although this species is superfi-
cially similar to the preceding one {hrandti),
I think it is distinct, differing most obvi-
ously by presence of elytral microsculpture.
The 8th and 9th elytral intervals are usually
less punctulate in ohscurus than in hrandti.
Jrichotichnus gutfula n. sp.
Description. With characters of genus;
form average; brownish black above, sides
of elytra ( and to some extent of prothorax )
testaceous, elytra with a conspicuous com-
mon testaceous subapical sutural spot
(reaching 3rd intervals), abdomen either
with irregular broad testaceous margins or
wholly rufescent, appendages brownish
testaceous; moderately shining, elytra usu-
ally with distinct transverse microreticula-
tion. Head 0.78 and 0.79 width prothorax;
eyes large, separated from mouth below
by c. Vs width of an eye. Prothorax sub-
transverse; width/length 1.48 and 1.43;
base/apex 1.19 and 1.21; sides rounded
anteriorly, converging and nearly straight
or slightly sinuate before obtuse but well
defined basal angles; disc depressed at
sides basally; basal marginal line usually
indistinct at middle; surface of disc exten-
sively punctate, the punctation finer and
less dense at middle. Elytra: width elytra/
prothorax 1.32 and 1.33; 8th and 9th inter-
vals not or not much punctulate. Secondary
sexual characters normal. Measurements:
length 7.6-8.7; width 3.1-3.6 mm.
Types. Holotype 6 (Bishop Mus.) and
31 paratvpes (some in M.C.Z., Type No.
31,374) 'from Wau, Morobe Dist., N-E.
N. G., 1200 m, Feb., Mar., June, Aug.,
Sept., Oct., Nov., Dec. 1961-1963 (Sedla-
ceks); 1 paratype, same locality, 1700-
1800 m, Nov. 17, 1961 (Sedlacek).
Additional material. Twenty-four speci-
mens from 13 widely scattered localities
including Dobodura, in all 3 political divi-
sions of New Guinea; altitudes, near sea
level to c. 2000 m. Specimens taken in
every month except May and June.
Measured specimens. The i holotype and
1 9 paratype from Wau.
Notes. This species is characterized by
its size plus presence of a conspicuous pale
subapical sutural spot. It is apparently
widely distributed especially in the foot-
hills and lower mountains of New Guinea.
Specimens of this or a closely related
species have been seen also from New
Britain and New Ireland (Bishop Mus.).
Trichotichnus mongi n. sp.
Description. With characters of genus;
form as in Figure 22, differing from other
Trichotichnus by subquadrate prothorax,
strongly narrowed anteriorly; piceous, lat-
eral margins of prothorax and elytra nar-
rowly translucent or pale, sutural intervals
reddish toward apex; shining, elytra sub-
iridescent, without distinct reticulate micro-
sculpture. Head 0.72 width prothorax;
eyes rather small, separated from mouth
below by more than V^ width of an eye.
58
Bulletin Museum of Comparative Zoology. Vol. 137, No. 1
Prothorox subquadrate except strongly nar-
rowed at extreme front; width length 1.40;
base/apex 1.35; sides rounded anteriorly,
broadly sinuate before c. right ( slightly
obtuse) basal angles; basal marginal line
fine but entire; disc scarcely depressed at
sides, extensively punctate, the punctures
finer and less dense across middle. Elytra
convex ( more so than usual ) ; width elytra/
prothorax 1.46; outer intervals with a little
sparse punctulation. Secondary .sexual char-
acters of 9 normal; i unknown. Measure-
ments: length c. 7.5; width 3.3 mm.
Type. Holotype 9 (M.C.Z., Type No.
31,375) from Tumnang, Mongi Watershed,
Huon Pen., N-E. N. G., 1400-1600 m, Apr.
14-15, 1955 (E. O. ^^^ilson); the type is
unique.
Notes. The unique form of this species
makes it worth describing, even though
I have only one specimen of it and do not
know the male.
Trichotichnus semirugosus n. sp.
Description. With characters of genus;
form c. average, small; brownish piceous,
margins of prothorax and elytra narrowly
testaceous, elytra with common subapical
sutural pale spot usually reaching 3rd in-
tervals; lower surface in part dark but with
extensive, irregular testaceous areas; anten-
nae brownish, legs brownish testaceous;
rather shining, elytra usually with faint
tran.sverse microreticulation. Head O.H\ and
0.84 width prothorax; eyes rather large,
separated from mouth below by <". 's width
of an eye. Prothorax subcordate; width
length 1.51 and 1.4(S; base apex 1.12 and
1.16; sides broadly round(>d anteriorly, con-
verging and straight or nearly so posteriorly;
posterior anglers distinct but obtuse, some-
times minutely denticulate; base more
oblique at sides than usual, not or indis-
tinctly margined; disc weakly depressed at
sides, extensively punctate, the punctation
finer and less dense across middle. Elytra:
width elytra /prothorax 1.39 and 1.44; 8tli
and 9th intervals rugosely punctate, punc-
tation present but less dense at bases of 6th
and 7th intervals. Secondary sexual char-
acters normal. Measurements: length 5.3-
6.8; width 2.3-2.8 mm.
Types. Holotype 9 (M.C.Z., Type No.
31,376) and 2 (99) paratypes from
Dobodura, Papua, Mar.-July 1944 (Dar-
lington); and additional paratvpes as fol-
lows: N-E. N. G.: 1, Wau, 1200 m, No\'.
21, 1961 (Sedlaceks); 7, Finschhafen, Huon
Pen., 10 m, Apr. 9-16, 1963 (Sedlacek), in
mercurv vapor light trap; 3. Torricelli Mts.,
Mobitei, 750 m, Feb. 28-Mar. 4, Mar. 16-
31, 1959 (W. W. Brandt, Bishop Mus.).
West N. G.: 1, Cyclops Mts. (no further
details) (Cheesman); 2, Hollandia area.
W. Sentani, Cyclops Mts., 50-100 m, June
22-24, 1959 (Gressitt), in light trap; 2,
Ifar, Cvclops Mts., 450-500 m, Sept. 7 and
9, 1962 (Sedlacek).
Measured specimens. One i paratype
from Mobitei and the 9 holotype, in this
order.
Notes. The coloration and the dense
punctation of the ehtral margins would
place this species in Lampetes, if Lampetes
were distinguished from Trichotichnus. The
new species is in fact close to Lampetes
isahellinus Louwerens of Amboina (Tijd-
schrift voor Ent. 105, 1962, p. 140). Ho\\'-
ever, comparison with paratopes of isa-
hellinus shows that scmiruii.osus has the
outer elytral intervals more completeK' and
more densely rugose, although the differ-
ence is not great.
Trichotichnus mixtus n. sp.
Descriplion. With characters of genus;
lorm average, rather small; brownish i^ice-
ous, lateral margins ol prothorax and elytia
transluc(Mit or pale, and sutural and some-
times 2nd interxals ol clxtia paler befori'
apex (variable), abdomen broadK" but ir-
regularly pale-margined, legs and antennae
irregularly brownish ti'staceous; shining,
elytra with or without light transverse
mieroretieulation. llccul 0.79 and 0.79 width
piothorax; eyes rather large, separated Irom
mouth below by c. 'is diameter ol an eye. Pro-
ihonix transNcrse-subeordate; width length
The Carabid Beetles of New Guinea • Darlington 59
1.54 and 1.54; base apex 1.15 and 1.13;
sides broadly rounded anteriorly, nearly
straight and converging posteriorly to ob-
tuse, sometimes slightly blunted posterior
angles; basal marginal line usually inter-
rupted at middle; disc depressed at sides
basally, extensively punctate, the puncta-
tion finer and sparser across middle. Elytra:
width elytra prothorax 1.36 and 1.36; 7th
and 8th intervals variabK' punctate (8th
varying from almost impunctate to almost
rugose); other intervals sparsely or not
punctulate. Secondary sexual characters
normal. Measurements: length 5.8-7.5;
width 2.8-3.2 mm.
Types. Holotype S (Bishop Mns.) and
4 paratypes (2 in M.C.Z., Type No. 31,377)
from Torricelli Mts., Mobitei, N-E. N. G.,
750 m. Mar. 5-15, Apr. 16-22 (holotype
with latter date), 1959 (W. W. Brandt).
Additional material. Twenty-one speci-
mens from 11 localities (including Wau)
in all 3 political divisions of New Guinea
are assigned to mixtus but not as types.
They vary considerably in several char-
acters.
Measured specimens. The c5 holotype and
1 9 paratype.
Notes. Because of the variation of this
species ( if it is all one species ) I have con-
fined the type series to specimens from one
locality. In general, the species should be
recognizable by size; sutural intervals pale
before apex; prothorax usually relatively
wide (wider and a little more depressed
at sides toward base than in scmirug,o.sus);
and outer elytral intervals usually punc-
tulate but not rugose, although this last
character is surprisingly variable even in
the type series.
Trichotichnus delicafus n .sp.
Description. With characters of genus;
form ( Fig. 23 ) slender-average, very small;
brownish piceous, prothorax with margins
narrowly pale or translucent, elytra with
margins and common subapical sutural spot
(including small parts of 2nd intervals)
testaceous or rufescent; abdomen with or
without well defined pale margins; af)pend-
ages testaceous; shining, elytra without
reticulate microsculpture. Head 0.74 and
0.77 width prothorax; eyes moderate, sepa-
rated from mouth below by c. '^i\ or Vs width
of an eye. Prothorax subcordate; width/
length 1.50 and 1.44; base/apex 1.11 and
1.21; sides broadly slightly sinuate before
obtuse, usually slightly blunted posterior
angles; basal marginal line vague or incom-
plete at middle; disc scarcely depressed at
sides even basally, less densely punctate
than usual, with middle of disc least punc-
tate. Elytra: width elytra/ prothorax 1.30
and 1.31; outer intervals without or with
only sparse punctulation. Secondary sexual
characters normal. Measurements: length
5.3-5.8; width 2.2-2.4 mm.
Types. Holotype 9 (Hungarian Nat.
Mus.) and 1 9 paratype (M.C.Z., Type
No. 31,378) from I. Deslacs (Garove Is.),
N-E. N. G., 1901 (Biro); and additional
paratypes as follows: Papua: 1, Woodlark
Is. (Murua), Kulumadau Hill, Mar. 9-12,
1957 (W. W. Brandt, Bishop Mus.). West
N. G.: 1, Hollandia, Dec. 1944 (W. T.
Nailon, Fenton Coll.); 1, Res. Hollandia,
Dojo, 2nd Strip, July 12, 1957 ( R. T. Simon
Thomas, Louwerens Coll.).
Measured specimens. A i paratype from
Woodlark Is. (the only 6 of the species
seen) and the 9 holotype, in this order.
Notes. T. delicatus is characterized b\'
small size, markings, and scarcely depressed
sides of pronotum.
Genus HARPALOXENUS Schauberger
Schauberger 1933, Ent. Anzeiger 13, p. 154.
Diagnosis. Characters as for Trichotichnus
(preceding genus) except anterior tibiae
wider; form characteristic (Fig. 24); upper
surface without distinct reticulate micro-
sculpture, but 8th and 9th elytral intervals
closely punctulate (except in fortis), other
intervals sparselv or not punctulate; wings
full.
Description. None required here.
60
Bulletin Museum of Cotnporaiive Zoology, Vol. 137, No. 1
Type species. 11. javanus Schauberger, of
Java.
Generic (listriJ)iifion. Java and Ando-
nare Is., Celel>es, Philippines, Moluccas
(Halmahera), New Guinea, and (unpub-
lished) Solomon Islands and New
Hebrides.
Notes. The species assigned to this genus
seem to form a natural group. However,
the group is apparently closely allied to
TricJioticlinus and further study may show
that it is not worth generic separation.
Key to Species of Harpaloxenus of
New Guinea
1. Anterior tibiae with apex c. Mi wide as
til)ial leii.irtli: head relatively wider (more
than 0.80 width prothorax) - ^ - 2
- Anterior tibiae with apex c. M wide as tibial
len.qth; head relatively smaller (usually less
than 0.80 width prothorax) -- 3
2. Male with front but not middle tarsi
squamulose (p. 60) fortis
- Male with front and middle tarsi squamulose
(p. 61) ma.s
3. Larj^er (length c. 11 mm) (p. 62) wan
- Smaller (length c. 10 mm or less) - 4
4. Frouotum with sides usually slightly de-
pressed toward 1)ase, base usually more
eoarsely and extensiveh' punetate, posterior
angles usually better defined, and median-
lateral setae usually c. Vi of prothoracic
length before apex (p. 61) celehen.sis
- Pronotum with sides not depressed, base
usualK' more finely punetate especially at
middle of base, posterior angles usually more
obtuse, and median-lateral setae usually c.
'':! of prothoracic length before apex (p.
62 ) sedlaccki
Harpaloxenus fortis n. sp.
Dcscri])lio)i. With characters of genus;
form as in Figure 24, heavily built; ])rownish
piceous, lateral margins ol pronotum and
elytra including 9th intervals ±: yellowish,
lower surhice with extensive yellowish areas
especially laterally, appendages brownisli
testaceous. Head wide, ().<S(S and ().(S5 w idth
prothorax; antennae stout, middle segments
scarcely longer than wide. Proihorax
broadly cordate; width/length 1.57 and
1.58; base/apex 0.93 and 0.99; sides con-
verging and usually l)roadl\' but not strongly
sinuate before obtuse but distincl basal
angles; disc slightly depressed at sides pos-
teriorly, basal impressions weak and ir-
regular, base punctate chiefly toward sides.
Elytra elongate-quadrate; width elytra pro-
thorax 1.14 and 1.14; outer intervals (8.
9) less punctulate than usual in genus; 3rd
intervals either with or (usually) without
minute puncture on inner edge behind
middle. Le^.s: front tibiae very wide in
both sexes, apex c. Va wide as tibial length,
and apex usually sinuate-emarginate with
outer angle slightly produced; middle tibiae
slightly wider and more arcuate than in
most other species of genus. Secondary
sexual characters: 6 front tarsi slightly di-
lated, with segments 1 (apex only) to 4 2-
seriately squamulose below; middle tarsi
not perceptibly dilated and not squamulose.
Measurements: length 8.5-10.3; width 3.1-
3.7 mm.
Types. Holotype S (M.C.Z., Type No.
31,379 ) and 17 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
additional paratypes from Papua as fol-
lows: 8, Kokoda, 1200 ft. (366 m). May.
July, Aug. 1933 (Cheesman); 1, same lo-
cality. Mar. 20, 1956 (Gressitt), in light
trap; 2, Saputa, near Buna, 1943-44 ( R. B.
Speiry, Chicago Mus.); 1, Deria, Amazon
Bay Dist., Dec. 1962 (W. W. Brandt.
C.S.I.R.O.); 1, Mt. Lamington, 1300-1500
ft. (c. 400-450 m) (C. T. McNamara, S.
Australian Mus.).
Additional material. West N. G.: 2.
(99), Ilollandia, Apr. 1945 (B. Malkin.
U.S.N.M.); 3 (99), Hollandia area, W.
Sentani, Cyclops Mts., 50-100, 150-250 m,
June 17, 22-24, 1959 (Gressitt and T. C.
Maa, Bishop Mus.), 2 of thes(> specimens
teneral and taken in light trap; 1 s" , Kota
Nika, Res. Ilollandia, Jan. 25, 1956 ( R. T.
Simon Thomas, Louwerens (^oU.); 1 9,
Wasian, Vogelkop, Sept. 1939 (Wind.
M.C'.Z. ). Thes(> specimens unfortunat<:4\
are all 9 9 . The\' arc referred to the
present spc>cies rather than the following
one (mas) because the 8th and 9th ehtral
inter\als are almost impunctate.
The Carabid Beetles of New Guinea • Darlington 61
Measured specimens. The i holotype and
1 9 paratype from Dobodura.
Notes. The present species differs from
all others known in the genus by absence
of squamules on the male middle tarsi.
However, I do not think that this justifies
making a separate genus or subgenus. This
species seems otherwise to be a well char-
acterized Harpaloxenus, and it apparently
is closely related to the following (mas),
which has the male middle tarsi normally
squamulose.
Harpaloxenus mas n. sp.
Description. With characters of genus;
form c. as in preceding species (foiiis)
but slightly less heavily built; characters as
in preceding species except as follows.
Head relatively slightly smaller, 0.81 and
0.S2 width prothorax. Prothorax with sides
converging but not or scarcely sinuate pos-
teriorly, and with posterior angles slightly
more obtuse; width/length 1.52 and 1.45;
base/apex 1.00 and 0.9cS. Elytra: width
elytra/prothorax 1.24 and 1.29; outer inter-
vals (8, 9) closely punctulate at least an-
teriorly, sometimes in part rugose. Sec-
ondary sexual characters: i front and middle
tarsi slightly dilated, 2-seriately squamu-
lose. Measurements: length c. 9.5-10.5;
width 3.5-3.7 mm.
Types. Holotype $ (Bishop Mus.) and
3 ( (5 9 9 ) paratypes (pair in M.C.Z., Type
No. 31,380) from Finschhafen, Huon Pen.,
N-E. N. C, 10 m, Apr. 9-16, 1963 ( Sedla-
cek), in mercury vapor light trap; 1 i
paratype, Wau, 1050 m, Nov. 4, 1961
(Sedlacek); 1 i paratype. Hoi Maffin, near
Sarmi, West N. G., July 18, 1959 (T. C.
Maa, Bishop Mus.).
Measured specimens. The 6 holotype and
1 9 paratype from Finschhafen.
Notes. This species resembles the pre-
ceding one in width of front tibiae but
differs in details of form (slightly nar-
rower head, slightly differently .shaped pro-
thorax) and in extensive punctulation of
outer elytral intervals. The paratype from
Hoi Maffin has these intervals less punc-
tulate than in the Finschhafen specimens
but still more punctulate than in any fortis
that I have seen.
Harpaloxenus celebensis Schauberger
Schauberger 1933, Ent. Anzeiger 13, pp. 155, 157.
Louwerens 1953, Verhandlungen Naturforschenden
Gesellschaft Basel 64, p. 306.
1956, Treubia 23, p. 222.
Description ( for recognition only ) . Form
average, rather variable; prothorax and
elytra usually narrowly yellow-margined.
Head 0.75 and 0.81 width prothorax. Pro-
thorax trans verse-subcordate; width length
1.50 and 1.52; base/apex 1.20 and 1.06 (ex-
ceptionally variable); sides rather weakly
converging, not or only slightly sinuate be-
fore distinct but slightly obtuse or blunted
posterior angles, with anterior-lateral setae
usually c. V4 of prothoracic length from
apex. Elytra: width elytra/prothorax 1.24
and 1.21; dorsal punctures usually present,
on inner edge 3rd intervals against 2nd
striae (sometimes absent on one or both
elytra). Legs: front tibiae with apex c. %
wide as tibial length. Secondary sexual
characters: S front and middle tarsi squamu-
lose. Measurements: length 8.5-10; width
3.2-3.8 mm.
Type. From South Celebes; probably
in Schauberger Coll. (not seen).
Occurrence in New Guinea. Papua: 7,
Dobodura, Mar.-July 1944 (Darlington);
2, Dogon, Amazon Bay Dist., 2400 ft. (c.
740 m), Oct.-Nov. 1962 (W. W. Brandt,
C.S.I.R.O.) N-E. N. G.: 3, Wau, Morobe
Dist, 1200 m, Dec. 18, 1961 (Sedlacek);
1, Finschhafen, 10 m, Apr. 9-16, 1963
(Sedlacek), in hght trap; 2, Torricelli Mts.,
Mobitei, 750 m. Mar. 5-15, 16-31, 1959
{\y. W. Brandt, Bishop Mus.). West N. G.:
6, Hollandia and vicinity including Cyclops
Mts. (various dates and collectors); and 5,
doubtfully identified, from localities farther
west in West N. C, including Biak Is.
Measured specimens. A pair ( <^ 9 ) from
Dobodura.
Notes. Louwerens records celebensis
from Java, Sumba, and Halmahera, as
62 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
well as Celebes, and the present records
extend its range to New Guinea. However,
variation is considerable and I am not sure
of my identifications in some cases. Small
specimens of this species can be confused
with certain Trichotichnus, especially with
discolored individuals of ii.uttula, but the
particular Trichotichnus in question do not
have the Sth and 9th elvtral intervals closelv
punctulate and have, of course, slightly
narrower front tibiae. See under the follow-
ing species (sedlaceki) for further com-
parisons.
Harpaloxenus sedlaceki n. sp.
Description. With characters of genus;
form average; black or piceous, sides of pro-
notum not or vaguely pale, elytra narrowly
pale-margined, abdomen pale-spotted at
sides, appendages brownish testaceous.
Head 0.75 and 0.75 width prothorax.
Prothorax transverse-subquadrate; width
length 1.50 and 1.49; base apex 1.19 and
1.21; sides moderately converging and nearly
straight posteriorly but usually not sinuate,
with median-lateral setae usually c. M', of pro-
thoracic length before apex; posterior angles
obtuse, ± blunted; disc rather strongly almost
evenly convex, not depressed at sides pos-
teriorly, baso-lateral impressions slight and
poorly defined; base (rather finely) punctate
especially at sides. Elytra: width elytra, pro-
thorax 1.22 and 1.20; outer intervals (8, 9)
extensively closely punctulate; dorsal punc-
tures of 3rd intervals usually c. midway be-
tween 2nd and .3rd striae (see Notes, be-
low). Le^s: front tibiae with apex c. Vl
wide as tibial length. Secondary sexual
characters: 6 front and middle tarsi sliglith
dilated, 2-seriately scpiamulose. Mea.surc-
menls: l(>ngth 8.0-9.0; width 3.1-3.5 mm.
'Types. Holotype $ (Bishop Mus.) ami
20 paratypes (some in M.C.Z., Type No.
31,381) from Wau, Morobe Dist., 'l20() m,
N-E. N. G., dates in Apr., Aug., Oct., Nov.,
Dec. 1961-1963 (Sedlacek).
Additional material. Twenty-two speci-
mens from numerous localities, from Mo-
dewa and Dobodura in Papua to Hollandia
in West N. G. Some of these specimens
are identified only doubtfully.
Measured s))ecimens. The i holotype and
1 9 paratype.
Notes. This species is similar to celehensis
(above) but differs in having the pronotum
slightly more convex with sides usually not
at all depressed toward base, and in other
ways indicated in the preceding Key to
Species of Harpaloxenus of Neic Guinea.
In the types of sedlaceki the dorsal punc-
ture of the 3rd interval is usually midway
between the 2nd and 3rd striae, not close
to the 2nd stria. Of the 21 specimens from
Wau, only 1 has the puncture close to the
2nd stria (on the inner edge of the 3rd
interval) on both elytra. Two have the
puncture close to the 2nd stria on one
elytron but near the middle of the 3rd
interval on the other. One has the puncture
close to the 3rd stria on one elytron. And
17 have the puncture near the middle of the
3rd interval (but somewhat variable in
position ) on both elytra. However, this un-
usual position of dorsal punctures may be
characteristic of the local population at
Wau rather than of the species as a \\'hole.
Harpaloxenus wau n. sp.
Description. With characters of genus;
form nearly as in celehensis and sedlaceki,
but larger; pronotum and ehtra not or nar-
rowly and faintly pale-margined, abdomen
with irregular testaceous marks at sides,
appendages brownish testaceous. Head 0.73
and 0.75 width prothorax. Prothorax trans-
verse-subquadrate; width length 1.48 and
1.51; base apex 1.17 and 1.24; sides slightly
converging posteriori), nearly straight but
not sinuate before slightK obtuse-blunted
posterior angl{\s; disc usualK' slightly de-
pressed at sides posteriorh', baso-lateral im-
pressions \agu(\ base finch punctate espe-
cialh- toward sides. Elytra: width elytra
prothorax 1.30 and 1.28; outer intervals (8,
9) extensively punctulat(\ 3rd intervals with
dorsal punctur(> usualK' by 2nd stria behind
middle. Leii,s: Iront tibiae with ajiex c. 'i
wide as tibial length. Secondary sexual
The Carabid Beetles of New Guinea • Darlington
63
characters: S front and middle tarsi slightly
dilated, 2-seriately squamulose. Measure-
ments: length c. 11; width c. 4.2 mm.
Types. Holotype c^ (Bishop Mus.) and
29 paratypes (some in M.C.Z., Type No.
31,382) from Wau and vicinity, Morobe
Dist., N-E. N. G., 1100, 1200 (most), 1700-
1800 m, dates in every month, 1961-1964
( Sedlacek ) .
Measured specimens. The 6 holotype and
1 9 paratype.
Notes. The large size distinguishes this
species from other similar ones. As com-
pared with sedlaceki, with which it occurs,
wau is not only larger but has the sides of
the pronotum usually slightly depressed
toward base and the dorsal puncture of
the 3rd intervals usually adjacent to the 2nd
stria. Of the 8 types, only 1 has this punc-
ture distant from the 2nd stria on both
elytra. Two have the puncture against the
2nd stria on one elytron but distant from it
on the other elytron. And 5 have the punc-
ture against or very near the 2nd stria on
both elytra.
LYTER n. gen.
Diapiosis. Form and characters of me-
dium-sized Trichofichnus, but c$ front and
middle tarsi below with more than 2 rows
of long, slender scales forming a loose
vestiture (not a dense sole); prosternum
glabrous anteriorly; 3rd elytral intervals
without or with only faint traces of dorsal
punctures.
Description. Head: eyes separated from
mouth below by c. Vs diameter of an eye;
antennae rather short, middle segments c.
IVaX long as wide; front smoothly convex
except frontal suture shaiply impressed,
with impressed lines extending diagonally
back to above eyes; mentum with triangular
tooth; labial palpi with penultimate seg-
ments plurisetose; ligula long, emarginate,
2-setose outside middle of length; para-
glossae shorter than ligula. Frothorax with
1 lateral seta each side about Vi prothoracic
length from apex. Elytra: margins entire
at base, obtusely subangulate at humeri,
sinuate before apex; marginal channels nar-
row, without partial 10th intei-vals; striae
entire; scutellar striae long, at base 2nd
intervals; 3rd intervals impunctate or with
vestige of puncture by 2nd striae behind
middle (position as in Trichotichnus) .
Lower surface: prosternum glabrous an-
teriorly but with several setae at apex pro-
sternal process. Inner wings full. Legs:
front tibiae irregularly truncate, apex c. V:,
wide as tibial length, with principal (inner
apical) spur not much expanded; hind tarsi
with 1st segment 2x or more as long as wide
at apex, 5th segment with 2 accessory setae
each side. Secondary sexual characters: S
front tarsi slightly dilated, 4 segments
loosely clothed below with slender long
scales; middle tarsi scarcely dilated, with
some (fewer) similar scales; 2 setae each
side last ventral segment in both sexes;
S copulatory organs as in Figure 174, with
apex of middle lobe not produced beyond
orifice.
Type species. Lyter glaher n. sp. (below).
Generic distribution. The single species
is confined to New Guinea, so far as
known.
Notes. The relationships of this new
genus are doubtful. It may be derived
from an ancestor like Trichotichnus medius
(p. 55), from which it differs most obvi-
ously in the clothing of the 6 tarsi (2-
seriately squamulose in Trichotichnus). The
new genus is notable also for its relatively
long ligula, for absence of pubescence on
anterior part of prosternum, and for virtual
suppression of dorsal elytral punctures.
The name Lyter, from the Greek, signifies
one who loosens ( the squamae of the S front
tarsi ) .
Lyter glaber n. sp.
Description. With characters of genus;
form as in Figure 25; reddish piceous, ap-
pendages redder; shining, reticulate micro-
sculpture absent on front of head and disc
of pronotum, faint and somewhat transverse
on elytral intervals. Head 0.75 and 0.77
width prothorax. Prothorax: width length
64 Bulletin Museum of Comparative Zoolofiy. Vol. 137, No. 1
1.45 and 1.49; base/apex 1.12 and 1.16;
sides rounded anteriorly, c. straight and
converging in posterior half, narrowly mar-
gined; posterior angles slightly obtuse,
blunted; base and apex with or without
faint marginal lines; disc slightly depressed
at sides basally, the depressed areas finely
but not closely punctate. Elytra: width
elytra prothorax 1.21 and 1.22. Measure-
ments: length 7.3-8.3; width 2.9-3.3 mm.
Types. Holotype $ (California Acad.)
and 32 paratypes (some in M.C.Z., Type
No. 31,383) from Finschhafen, N-E. N. G.,
various dates in April and May (holotype.
May 1) (E. S. Ross); and additional para-
types as follows. Papua: 21, Kokoda,
1200 ft. (366 m), Apr., Aug., Sept. 1933
(Cheesman); 1, Owen Stanley Rge., Goilala,
Tapini, 975 m, Nov. 16-25, 1957 (W. \V.
Brandt, Bishop Mus.); 1, Dogon, Amazon
Bay Dist., 2400 ft. (730 m), Oct.-Nov.
1962 (W. \V. Brandt, C.S.I.R.O.); 1, Mt.
Lamington, 1300-1500 ft. {c. 400-450 m)
(C. T. McNamara, S. Australian Mus.).
N-E. N. G.: 5, "No. 14," Umi R., Markham
Vy., 480 m, dates in Nov. 1959 (Sixth
Archbold Exp., A.M.N.H.); 1, Lae, July
1924 (F. E. Skinner, Bishop Mus.); 3,
Bulolo, 730, 1170 m, Aug. 15, 19, 21, 1956
(E. J. Ford, Jr., Bishop Mus.); 58, Wau,
Morobe Dist., 1200 m, dates in every
month except June, 1961-1963 (Sedlacek);
2, same locality, 1700-1800 m, Nov. 17,
1961 (Sedlacek); 4, Sum-Sum, 64 km N. of
Wau, 580 m, Feb. 15, 1963 ( Sedlacek ) ; 1,
Karimui, S. of Ooroka, 1000 m, June 3,
1961 (Cre.ssitt), in light trap. West N. <;.:
3, Ilollandia area, \V. Sentani, Cvclops
Mts., 50-100, 150-250 m, June 17, 22-24,
1959 (Gressitt); 1, Kota Nika, Res. Ilol-
landia, Feb. 14, 1956 (R. T. Simon Thomas,
Louwerens Coll.), in light trap.
Measured speeimens. The i holotype and
1 9 paratype Irom I'^inschhalen.
Notes. Although this carabid is appar-
ently common in some places, I hiiled to
find it and do not know its habitat. The
k)calities suggest that it lives in rain forest.
Specimens Irom Bulolo and Ilollandia were
taken in light traps, which implies that the
insect flies.
Genus COLEOUSSUS Bates
Bates 1892, Ann. Mus. Civ. Genoa 32, p. 338.
Csiki 1932, Coleop. Cat., Caral)idae, Harpalinae 6,
p. 1217 (as suligenus of TriclioticJiniis) .
Andrewes 1939, Ann. Maj?. Nat. Hist. (11) 3, p.
132.
Diaiinosis. See Key to Genera of Har))alini
of New Guinea.
Deseription ( for recognition only ) . Form
of broad medium-sized Harpalini; upper
surface ( in New Guinean species ) \'er)-
shining, without reticulate microsculpture.
Head: eyes relatively large (compared
with most Triehotichnus), almost contigu-
ous with sides of mouth below; frontal
impressions deep, subpunctiform or curving
toward eyes posteriorly; mentum toothed;
ligula rounded, 2-setose; paraglossae at-
tached to ligula but longer, with narrowly
rounded apices; penultimate segments labial
palpi with more than 2 setae anteriorh'.
Prothorax as in Trieliotiehnus. Elytra:
sutural angles denticulate ( in New Ciuinean
species); striae entire; sutural striae long;
3rd intervals seriate-punctate. Inner winiis
full. LeiS,s: tarsi slender. Secondary .sexual
characters: see Descriptions of species.
Ty))e species. llypolitJius perlueens Bates,
of Kashmir, etc. (fixed In- Andrewes, 1939).
Generic distribution. India and Ceylon,
Sikkini, Burma, etc., to Java, Borneo,
Philippines (Negros), Cele]>es, Burn,
Moliiecas (Amboina), New (Guinea, Solo-
mons, New Hebrides, and ihc Cape York
Pen. of Australia (occurrence in Philii")-
pines. New Hebrides, and Australia bas(>d
on unpublished records ) .
Notes. Members of this genus seem to
be rare insects, usualK' taken onl\ one or
two indi\iduals at a time, although the\
are winged and IK to light. I do not know
their habitat.
Key io Species of Coleolisscs of
Ni:\\ (".riNKA
I. Onter an,ul(\s of cKtia not (Iclincd. hroaill)
ronndecl (p. 65) jHijitui
The Carabid Beetles of New Guinea
Darlington
65
- Outer angles of elytra ( before subapical
sinuations) well defined, right or obtuse (p.
65 ) angulatns
Coleolissus papua n, sp.
Description. With characters of genus;
form as in Figure 26, large, broad; lilack,
appendages irregularly reddish piceous;
very shining, elytra iridescent. Head 0.73
and 0.72 width prothorax; front faintly
punctulate. Prothorax transverse; width/
length 1.54 and 1.56; base/apex c. 1.11 and
1.11; sides broadly rounded to rounded
posterior angles; lateral margins broader
and more depressed posteriorly; apex mar-
gined, base indistinctly so; disc depressed;
baso-lateral impressions broad but poorly
defined, closely punctate; other parts of
disc sparsely or not punctulate. Elytra
wide; width elytra prothorax 1.30 and 1.39;
base margined; humeri rounded; outer sub-
apical angles rounded; apices slightly sinu-
ate before denticulate sutural angles; striae
impunctate; intervals slightly convex, finely
sparsely (scarcely detectably) punctulate,
3rd with c. 7 small punctures irregularly
spaced along most of length of inner edge.
Lower siwface: prosternum and abdomen
with a little fine, short, sparse pubescence
(scarcely detectable); prosternal process
setose. Secondary sexual characters: S un-
known; 9 with 2 setae each side last
ventral segment. Measurements: length
12.5-13.5; width 4.8-5.3 mm.
Types. Holotype 9 (Bishop Mus.) from
Kiunga, Fly R., Papua (W. W. Brandt);
and 1 9 paratype (M.C.Z., Type No.
31,384) from Hollandia area, VV. Sentani,
Cyclops Mts., West N. G., 150-250 m, June
23, 1959 (T. C. Maa).
Notes. This species is close to Coleolissus
leveri Van Emden of the Solomons ( I have
a specimen compared with the type) and
even closer to C. kalisi Louwerens of Celebes
( I have 2 paratypes received by courtesy of
Mr. Louwerens). The New Guinean insect
is slightly larger and broader than kalisi,
with broader baso-lateral prothoracic im-
pressions. I collected a single specimen of a
Coleolissus very similar to pai)ua on the
Cape York Pen., Australia, in 1958.
Coleolissus angulatus n. sp.
Description. With characters of genus;
form as in Figure 27; black, elytra subirides-
cent, appendages reddish brown. Head 0.72
and 0.72 width prothorax; front virtually
impunctate. Prothorax transverse-subcor-
date; width/length 1.51 and 1.56; base/apex
1.12 and 1.09; sides converging but usually
not sinuate (sometimes slightly so) before
± rounded posterior angles; lateral margins
moderately broad and reflexed; apex mar-
gined, base not; disc weakly convex, baso-
lateral impressions not sharply defined,
rather closely but irregularly punctate; disc
otherwise less closely but extensively punc-
tate especially across base and at sides,
scarcely punctate at middle. Elytra: width
elytra prothorax 1.30 and 1.32; basal mar-
gin entire, vaguely obtusely angulate at
humeri; outer apical angles well defined,
usually right, sometimes obtuse; sutural
angles denticulate; striae deep, impunctate;
intervals slightly irregular but not distinctly
punctate, 3rd with c. 7 very small punctures
irregularly spaced on inner edge along
most of length of intei^vals. Lower suiiace:
prosternum not pubescent except for setae
at apex of prosternal process; abdomen not
pubescent except for usual "fixed" setae.
Secondary sexual characters: $ front tarsi
moderately dilated, middle tarsi scarcely
so, both pairs 2-seriately squamulose; 2
setae each side last ventral segment in
both sexes. Measurements: length c. 7.5-
8.5; width 3.1-3.3 mm.
Types. Holotype S (M.C.Z., Type No.
31,385) and 2 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
additional paratypes as follows. N-E. N. G.:
3, Finschhafen, 10 and 180 m, Apr. 9-16,
1963 (Sedlacek); 1, Aitape, Aug. 1944
(Darlington). West N. G.: 2, Hollandia
area, W. Sentani, Cyclops Mts., 150-250 m,
June 25, 1959 (Gressitt and T. C. Maa,
Bishop Mus.); 1, Ifar, Cyclops Mts., 150-
66
Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
500 m, Sept. 6-9, 1962 (Sedlacek); 1, "Neth.
New Guinea" without further loeality, Dec.
10, 1944 (T. Aarons, California Acad.).
Measured specimens. The <:5 holotype and
1 9 paratype from Dobodura.
Notes. The deep elytral striation and
sharply defined outer elytral angles well
characterize this Coleolissus. The Dobodura
specimens were, I think, taken at light, and
some specimens from other localities are
evidently from light trap material.
Genus HYPHAEREON Macleay
Macleay 1825, Annulosa Javanica, p. 22.
Andrewes 1919, Trans. Ent. Soc. London for
1919, p. 156.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 6,
p. 1221 (see for additional references).
Diapiosis. Among New Guinean Har-
palina this genus is recognizable by: form
rather NebrioAike; elytra with series of
(small) punctures on inner edge 3rd inter-
vals; only 1 seta each side last ventral seg-
ment in both sexes.
Description ( for recognition only ) . Form
c. Nehria-like, convex; reticulate micro-
sculpture faint or absent on head and
pronotum, fine and transverse ( ± visible at
50x ) on elytra; elytra subiridescent. Head:
eyes moderate, narrowly separated from
mouth below; mentum toothed; ligula sub-
truncate, 2-setose; paraglossae rounded, at-
tached to but longer than ligula. Frothorax
subcordate; anterior marginal line fine but
usually entire. Elytra with margins sinuate
before apex; striae entire; 3rd intervals
seriate-punctate. Inner uini!,s: see under
species. Secondary sexual characters: i
front and middle tarsi slightly dilated, 2-
seriately squamulose; 1 seta each side apex
last ventral segment in both sex(\s.
Type species. II. reflexus Macleay, of
Java.
Generic distrihtition. Known from Su-
matra, Java, Flores, Celebes, and New
(riiinea (not Australia).
Notes. Of the 3 New Guinean species of
Ilyphaercon, timid us is most like the type
species, with whicli I shall compare it (in
Notes under timidus). Calathomijnus, which
resembles Ilyphaercon in form and in having
only 1 pair of setae on the last ventral
segment in both sexes, and which also oc-
curs in the Malay Archipelago (but not
New Guinea ) , is probably closely related
to Hyphacreon but differs in having strongly
angulate humeri.
Key to Species of Hyphaereon of
New Guinea
1. Humeral margins broadly evenly rounded;
elytral striae shallow, inter\'als flat; setae of
apical ventral segment distant from margin
by more than ^/io length of segment (p.
66 ) levis
- Humeral margins obtusely subangulate; elytral
striae deeper; setae of last ventral segment
less than %o length of segment from margin - 2
2. Prothorax less cordate, with slightly broader
base (width of base/width of head 1.20 and
1.17); wings dimorphic, often much reduced;
lowland-living (p. 67) timidus
- Prothorax more cordate, with narrower base
(width of base of prothorax/ width of head
1.14 and 1.12); wings large, folded; moun-
tain-living (p. 68) cordcHfi
Hyphaereon levis n. sp.
Description. With characters of genus;
fonn ( Fig. 28 ) average; black, legs testa-
ceous, antennae and mouthparts brown.
Head 0.67 and 0.66 width prothorax.
Prothorax subcord;ite-subquadrate; wddth
length 1.38 and 1.38; base/apex 1.19 and
1.16; base/head 1.10 and 1.07; sides con-
verging and very slightly sinuate before
obtuse, blunted posterior angles; baso-kit-
eral impressions poorh' defined; disc ex-
tensively pimctate especially across base
and at apex, almost impunctate across mid-
dle. Elytra: width elytra prothorax 1.26
and 1.26; details ;is usu;il in genus except
humeral margins broadly evenly rounded;
striae entire^ but less deep than usual, hmer
u'inis,s lull in both specimens. Measure-
ments: l(Migth c. 9.0; ^^■idth 3.3-3.4 mm.
Types. Ilolotvpe $ (Leiden Mus. ) and
1 5 "paratype (M.C.Z., Type No. 31,386)
both from Sibil, Star Rge., West N. G..
1260 in, |une 1959 (Neth. New Gnincvi
Kxp.).
The Carabid Beetles of New Guinea • Darlington
67
Notes. For comparisons, see preceding
Key.
Hyphaereon timidus n. sp.
Description. With characters of genus;
form (Fig. 29) of small, rather broad
Nebria; black or piceous, appendages testa-
ceous, antennae in part brown. Head 0.64
and 0.65 width prothorax. Prothorax sub-
cordate-subquadrate; width /length 1.42 and
1.39; base/apex 1.20 and 1.17; base/head
1.20 and 1.17; sides converging and usually
slightly, broadly sinuate before obtuse,
slightly blunted posterior angles; anterior
marginal line entire or not (variation in-
dividual); baso-lateral impressions poorly
defined; disc finely irregularly punctate
basally, c. impunctate elsewhere. Elytra:
width elytra prothorax 1.19 and 1.23;
humeri obtusely but usually distinctly angu-
late; striae deep, intervals convex. Wings
dimorphic or polymorphic (see Notes, be-
low). Secondary sexual characters as for
genus. Measurements: length c. 6-7;
width 2.6-2.9 mm.
Types. Holotype i (M.C.Z., Type No.
31,387) and 55 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); 20
paratvpes from Oro Bay (near Dobodura),
Dec. 'l943-Jan. 1944 (Darlington).
Additional material. N-E. N. G.: 17,
Nadzab, July 1944 (Darhngton); 1, same
locahty, June 1944 (Krombein, U.S.N.M.);
1, Erima, Astrolabe Bay, 1896 (Biro); 1,
Busu R., "12 km," Sept. 21, 1956 (E. J.
Ford, Jr., Bishop Mus.). West N. G.: 34,
Hollandia, July-Sept. 1944 ( Darlington ) ; 6,
same locality. May 1945 (B. Malkin,
U.S.N.M.); 5, Sabron, Cyclops Mts., 930
ft. (c. 280 m), Apr. 1936 \ Cheesman ) .
Measured specimens. The £ holotype and
1 9 paratype from Dobodura.
Notes. This new species is similar to
Hyphaereon reflexus Macleay (the type
species of the genus) of Java, but timidus
differs slightb' in proportions (e.g., the
base of the prothorax is relatively narrower
than in reflexus) and the pronotum of
timidus is less extensively punctate.
The wings of this species vary (Figs. 29,
A, B), and the variation is complex, being
partly individual, partly geographic, and
partly correlated with Iwdy size. Of the
specimens from Dobodura, 5 have wings
large and folded at apex; 51, strongly re-
duced. However, both the long- and the
short-winged forms are variable in the
Dobodura series. Among the long-winged
individuals, some have wings about 10%
shorter than others and with slightly weak-
ened venation, and among the short-winged
ones, the wing vestiges vary from about %
to about V2 the length of an elytron. In
the series from Oro Bay (only a few miles
from Dobodura) the proportion of long-
and short-winged individuals is different:
10 are long-winged, 9 short-winged. Seven-
teen specimens from Nadzab and 3 from
other localities in N-E. N. G. are all fully
winged or at least have wings long and
folded at apex. But my series from Hol-
landia is again dimorphic: 7 specimens
are long-winged, 27 short-winged. All the
long-winged specimens from Dobodura, Oro
Bay, and Hollandia are large. Some short-
winged individuals are equally large, but
there is much more variation in size among
the short-winged ones. I do not remember
noting this correlation in any other Ca-
rabidae. I have not studied state of wings
in specimens not collected by myself be-
cause I do not know how they were taken,
and method of collecting may have favored
getting one wing class more than another.
It is doubtful if even the long-winged
form of this species flies. Individuals are
common at some localities where much
collecting has been done, but few have
been obtained except by myself (on the
ground), and no specimen is labeled as
taken at light. The variation and use of
wings in this species would be an interesting
subject for study in the field.
Although my field notes are scanty, I
think my series of this species were taken
among dead leaves and vegetation on the
ground near water.
68
Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Hyphaereon cordons n. sp.
Description. With characters of genus;
form N ebria -\ike; black or piceous, append-
ages irregularly brown, darker than in other
species. Head 0.67 and 0.67 width pro-
thorax. ProfJiora.x cordate; width/length
1.35 and 1.43; base apex 1.23 and 1.18;
base/head 1.14 and 1.12; sides converging
and broadly sinuate before slightly obtuse
(almost right), slightly blunted posterior
angles. Elytra c. %o or more wider than pro-
thorax (elytra prothorax 1.32 and 1.32);
humeri ± subangulate; striae deep, intervals
convex. Win^is fully developed, or at least
long and folded at apex, in all specimens.
Seconckiry se.xual cJiaracters as for genus.
Measurements: length c. 7-8.5; width 2.7-
3.4 mm.
Types. Holotype £ (M.C.Z., Type No.
31,388) and 77 paratypes all from Chimbu
Vy., Bismarck Rge., N-E. N. G., 5000-7500
ft. (c. 1500-2300 m), Oct. 1944 (Dar-
lington ) .
Additional material. N-E. N. G.: 1,
Kainantu, 1650 m, Oct. 20-26, 1959 (T. C.
Maa, Bishop Mus.); 1, Wau, Morobe Dist.,
1200 m, Dec. 4-5, 1961 (Sedlacek), in
mercury vapor light trap.
Measured specimens. The <^ holotype and
1 9 paratype.
Notes. Although this species is evidently
sometimes common, and although all speci-
mens are fully winged, they are rarely
taken in light traps, which suggests that
even this winged species rarely flies.
Whether it is a real species or a local form
of timidus is not possible to say from
museum specimens. In any case it is
clearly distinguishable as indicated in the
preceding Key.
Genus ANOPLOGENIUS Chaudoir
Chaiuloir 1852, Bull. Soc. Nat. Moscow 25, 1, p.
88.
Csiki 1932, Colcop. Cat., Carabidae, liarpaliiiae 6,
p. 1236.
Schauberger 1937, Ent. Rundscbau 54, p. 272.
Basilewsky 1951, Ann. Mus. Congo Belj^c (8),
Zoo!., 9, p. 122 (see for synonymy and additional
refcronces).
Diagnosis. Relatively large Acupalpina;
anterior marginal line of pronotum entire
and deeply impressed; scutellar striae ab-
sent.
Description. None required here.
Type species. Stenoloplius alaccr Dejean,
of Africa.
Generic distribution. The warmer parts
of the Old World; in the Oriental-Aus-
tralian area, from China and Japan to
northern Australia.
Notes. The species of this genus are
among the most aquatic of Carabidae, oc-
curring as a rule in vegetation that is float-
ing in water. They are active and winged
and are common in some places, including
the Philippines, although unaccountably
rare or local in New Guinea.
Anoplogenius marginafus (Macleay)
Macleay 1888, Proc. Linnean Soc. New South
Wales (2) 3, p. 472 (HarpJancr).
?iticisus Andrewes 1926, Ann. Mat;. Nat. Hist. (9)
18, p. 279.
?poJitus Schauberger 1937, Ent. Rundschau 54, p.
273.
Description. None needed here. See
Notes below; length c. 6.5-8.0 mm.
Types. Of murginattis, from King's
Sound, Australia; probably in Macleay
Mus., Sydney (not seen). Of incisus, from
Fort de Kock, Sumatra; in Briti.sh Mus.
(seen). Of politus, from "Tigerinsel (New
Guinea)" (? = Matjan, Pulau-Pulau, which
is really not off New Guinea but south of
Celebes); in Stockholm Mus. (not seen).
Occurrence in New Guinea. The only
(supposedly) New Guinean specimens of
Anoplogenius known to me are: 1 labeled
simply "Papua," presumably collected by
Biro, now in Hungarian National Mus.;
scNcral Irom "Dorey, New (Guinea," pre-
sumabK- collected by Wallace and perhaps
actual]) from Celebes (see Part I of the
present work, pp. 330-331); and the types
ol ])olHus from "Tigerinsel," probably off
(Celebes rather (liaii New (Guinea.
Notes. The Oriental- Australian species of
Anoplogenius are taxonomicalK difficult at
b(\st, and in the case of this New (Uiinean
The Carabid Beetles of New Guinea • Darlington
69
species the difficulty is increased by lack
of adequate material and by doubt about
localities, as indicated above. The three
authors concerned published their descrip-
tions without reference to each other, but
Andrewes and Schauberger both compared
their species with A. cyanescens Hope, and
Macleay's Harplaner marginatus is appar-
ently an Anoplogenius near cyanescens (B.
P. Moore, personal communication, 1965).
I therefore tentatively conclude that incisus,
politiis, and marginatus are probably all
one species which ranges at least from
Sumatra to New Guinea and northern
Australia. This species is narrowly or
indistinctly pale-margined, with relatively
distinct (narrowly rounded) posterior pro-
thoracic angles, and with baso-lateral im-
pressions of pronotum extensively punc-
tate. I have specimens with these charac-
ters from Morotai Island in the Moluccas,
and from Townsville and Rockhampton,
Australia.
Genus EGADROMA Motschulsky
Motschulsky 1855, fitude Ent. 4, p. 43.
Csiki 1932, Coleop. Cat., Carabidac, Harpalinae 6,
p. 1239 (as subgenus of Actipalpus) (see for
additional references ) .
Jeannel 1942, Faune de France, Coleop. Carabiques,
Part 2, p. 699.
Basilewsky 1951, Ann. Miis. Congo Beige (8),
Zoo!., 9, p. 144.
Diagnosis. See Key to Genera of Harpalini
of New Guinea.
Description. None required here.
Type species. Carabus smaragduhis Fab-
ricius, below.
Generic distribution. The warmer parts
of the Old World: Africa and Madagas-
car ( 1 species reaching the Mediterranean
part of Europe); southern Asia north to
Japan, and across the Malay Archipelago
to the Philippines and Australia.
Notes. The Oriental-Australian species of
Egadroma are exceptionally difficult. They
are closely related or at least very similar
among themselves; they vary geographically
and individually; and some species are
widely distributed and very common, so
that many specimens fell into the hands of
early taxonomists who described them in-
adequately and failed to understand their
interrelationships. I do not pretend fully
to understand them now, but can make
the following comments on the species that
occur in New Guinea. Three common
species occur there, distinguishable by both
external and genitallic characters. (A fourth,
endemic species is known from a single ? . )
All three are widely distributed outside
New Guinea and at least two of them have
received different names in different places.
To fix the synonymy of these species out-
side New Guinea would be a major, time-
consuming undertaking, and is beyond my
power now. I shall therefore simply use
for each species the name that I think ap-
plies in New Guinea, with tentative notes
on synonymies.
Although I did not always distinguish
the species in the field, my notes suggest
that quinquepustulata and smaragdula oc-
cur in wet places usually by standing water,
but that robusta occurs principally in drier
habitats, especially under cover in open
grassland. All these species are winged,
and all fly to light.
Key to Species of Egadroma of New Guinea
1. Size larger (c. 6-7 mm); elytra conspicu-
ously 3- or 5-maculate, and very shining;
apex of aedeagus long (p. 70 )
quinquepustulata
- Usually smaller (less than 6 mm, except in
Cyclops); usually with reduced or no mark-
ings, and often ( not always ) less shining;
apex of aedeagus shorter (except unknown
in Cyclops) 2
2. Large (7.4 mm) (p. 70) cyclops
- Smaller ( less than 6 mm ) _., 3
3. Form relatively narrower, with relatively
wider head ( head usually more than 0.74
width prothorax, prothoracic width/length
usually less than 1.45); elytra rather strongly
iridescent; aedeagus finely notched at sides
(p. 70) smaragdula
- Broader, with relatively narrower head (head
usually less than 0.74 width prothorax, pro-
thoracic width/length usually more than
1.45); elytra less iridescent, usually with
distinct microreticulation; aedeagus not
notched at sides (p. 71) robusta
70 Bulletin Museum of Comparative Zoolofiy, Vol. 137, No. 1
Egadroma quinquepusfulafa (Wiedemann)
Spiistiilatiis- Wiedemann 1823, Zoologisches Maga-
zin (2) 1, p. 58 (Badi.stcr) .
Csiki 1932, Coleop. Cat., Caral)idae, Harpalinae
6, p. 1240 (see for additional references).
Habu 1961, in Kira and ITniesao, Nature and Life
in Southeast Asia (Kyoto) 1, p. 276, fig. 4
( <5 genitalia ) .
Description. None needed here; see pre-
ceding Key to Species and following Notes;
length c. 6-7 mm.
Type(sj. From Bengal, India; in Copen-
hagen U. Mus. (not seen).
Occurrence in New Guinea. Widely dis-
tributed at low altitudes, fairly common:
41 specimens from localities over most of
the length of New Guinea (Papua to the
Vogelkop); most at low altitudes, but 1,
Chimbu Vy., Bismarck Bge., 5000-7500 ft.
(c. 1500-2300 m), and 1, Wau, 1200 m.
Notes. This relatively large, clearly
marked, and therefore comparatively easily
recognized Efradroma ranges from SE.
Asia including Japan and Formosa across
the Malay Archipelago to North Queens-
land, Australia. In the past, the species
has often been treated as a variety of
smaraiiduki but it is unquestionably distinct
by genitallic as well as external characters.
Specimens from New Guinea vary in
elytral markings, the variation being in-
dividual, not geographic. Conspicuous post-
humeral pale spots are always present, and
a variable (sometimes Faint) subapical
sutural mark is always present too, but
antcapical spots on the 7th and 8th intervals
are variably developed and often absent.
Egadroma cyclops n. sp.
Descri])lio)i. Form as in T'igure 30, larg(\
broad; side margins of j^rothorax testaceous-
translucent, ol elytra scarcely so; append-
ages reddish testaceous, antennae darker
from 3rd segment; shining, front with iso-
diametric reticulate microsculpture, discs
of pronotum and elytra not visibly micro-
reticulale (at 5()X ) but moderately irides-
cent. Head 0.6.S width prothorax; formed
as usual in genus. Prothorax transverse;
width length 1.45; base/apex 1.22; sides
rounded, with moderate reflexed margins;
basal angles rounded; base not margined,
apical marginal line interrupted at middle;
baso-lateral impressions broad, shallow, c.
rugose-punctate, with punctation finer and
sparser at middle of base. Elytra: width
elytra/prothorax 1.33; humeri prominent
but rounded; striae deep, impunctate;
scutellar striae long; intervals finely sparsely
punctulate, 3rd with 1 puncture on inner
edge less than % from apex. Winii,s full.
Lower surface: prostemum with some short
pubescence; abdomen not pubescent at
apex. Secondary sexual cJiaractcrs: i un-
known, 9 normal. Measurements: length
7.4; width 3.3 mm.
Type. Holotype 9 (Bishop Mus.) from
Hollandia area, W. Sentani, Cyclops Mts..
West N. G., 50-100 m (Gressitt and T. C.
Maa), in light trap.
Notes. This new species resembles
smaraiidula but is larger (a large smaragdula
is less than 6 mm long), with relatively
narrower head. I feel sure it is a distinct
species although reprc-sented b\- onl>- a
single 9 specimen.
Egadroma smaragdula (Fabricius)
Fabricius 1798, SuppUineutuni Eut. Systematicac.
p. 60 (Camhits).
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 6.
p. 1241 (see for additional references).
Jedlicka 1935, Acta Soe. Ent. Czechoslovakia 32,
p. 1 13 (in key).
Habu 1961, in Kira and Umesao, Nature and Lite
in Southeast Asia (Kyoto) 1, p. 275, fig. 3 ( S
genitalia).
Descri))liou. None required here; see
preceding Key to Species: k-ngth ± 5 mm.
Typc(s). From "India ori«Milali"; in
CoiXMihagen V\n\. Mus. (not sec-n).
Occurrciu-e in New Guinea. Common at
low altitud(\s probably throughout New
(i^uinea: 121 specimens from Miln(> Ba\
and Port Moresb>- to the X'ogelkop; most at
low altitudes but 2 from Wau and 1 from
Rattan Ciuup at 1200 in.
Notes. This is the common, unmarked
(at most with a faint rufesc(Mit area along
The Carabid Beetles of New Guinea • Darlington 71
suture posteriorly) Egadroma of the Ori-
ental-Australian area. It apparently ex-
tends from Asia to northern Australia but,
liecause of difficulty in distinguishing it
from similar forms, I have not tried to fix
the exact limits of its range.
Egadroma robusta Sloane
Sloane 1907, Deutsche Ent. Zeitschrift for 1907, p.
469.
Andrewes 1927, Ann. Mag. Nat. Hist. (9) 19, p.
110 (as synonym of dingo Castelnau).
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 6,
p. 1242 (as synonym of vestigialis) .
Description. None required here. See
preceding Key to Species and Notes, be-
low; length ± 5 mm.
Types. From Gazelle Pen., New Britain;
should now be in Deutsches Ent. Mus.,
Berlin-Dahlem (not seen).
Occurrence in New Guinea. Very com-
mon throughout New Guinea at low alti-
tudes: 368 specimens from many localities
from Milne Bay and Normanby, Ferguson,
Rossel, and Sudest Islands, Papua, to west-
ern part of West N. G. Although this species
does not commonly occur at altitudes of
more than a iew hundred meters, I ha\e
seen 4 from Wau, 1200 m, and 1, Chimbu
Vy., 5000-7500 ft. (c. 1500-2300 m). Speci-
mens have been collected in every month.
Notes. Most individuals of this species
from New Guinea have the elytra un-
marked or with only small posthumeral
pale marks on the 6th intei^vals, rarely
extending to the 5th and 7th intervals.
Similar unmarked individuals occur in New
Britain (types of robusta) and Cape York,
Australia (collected by me in 1958).
However, a few specimens from New
Guinea, mostly from the far west including
Biak Island, have also pale subapical
sutural dashes and variable subapical pale
spots, best developed on the 7th intervals.
These specimens may be referable to
Egadroma quadrimaculata (Macleay), which
was described from Australia, but which
may extend across the Malay Archipelago
at least to Java. In other words, robusta
may be an unspotted form ( occurring prin-
cipally but not exclusively on New Guinea )
of a more widely distributed maculate
species, tentatively identified as quadrimac-
ulata Macleay. This case requires further
study.
Most individuals of robusta are easily
separable from smaragdula by proportions
(indicated in the preceding Key) and
duller surface. Apparent intermediates do,
rarely, occur. I do not know whether they
are hybrids or individual variants. These
two species probably occupy different habi-
tats: smaragdula, wet places; robusta, rel-
atively dry ones.
Genus STENOLOPHUS Stephens
Stephens 1828, iHustrations British Ent., Man-
dibulata 1, pp. 67, 165.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae
6, p. 1259 (see for synonymy and additional
references )
Jeanne] 1942, Faune de France, Coleop. Carabiques,
Part2, p. 693.
Basilewsky 1951, Ann. Mus. Congo Beige (8),
Zool., 9, pp. 118, 213 (in text).
Diagnosis. See Key to Genera of Harpalini
of New Gui7iea.
Description. None required here.
Type species. Carabus teutonus Schrank,
of Europe, etc. (see Jeannel, 1942).
Generic distribution. Eurasia and North
America, and probably the Oriental Re-
gion and Malay Archipelago to Australia;
(probably not Africa below the Sahara, al-
though closely related genera occur there).
See Notes, below.
Notes. Jeannel and Basilewsky have
divided Stenolophus, and Basilewsky sug-
gests that the genus in a strict sense may be
confined to the Holarctic Regions. How-
ever, the following t\\'o New Guinean
species seem to fit reasonably well in
Stenolophus according to characters given
by Basilewsky (1951, p. 213). Moreover,
one of these two species (gonidius) has
the first segment of the posterior tarsi
plainly carinate while the other (volucer)
has not, which suggests that this character,
which has been used in dividing St e nolo-
72 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
phus, is less important than has been
thought. I shall therefore leave both New
Guinean species in Stenolophus, where in
fact earlier authors put them.
Both the following two species are ap-
parently widely distributed in the Malay
Archipelago, and I have specimens prob-
ably representing both from North Queens-
land, Australia, but because of doubt about
identifications I prefer not to state their
distributions in detail.
Key to Species of Stenolophus of
New Guinea
1. Sides of prothorax sinuate before base, with
liasal angles nearly right and scarcely blunted
(p. 72) volucer
- Sides of prothorax c. straight, converging
but not or scarcely sinuate posteriorly, with
basal angles obtuse, narrowly rounded (p.
72 ) gonidiiis
Stenolophus volucer Andrewes
Andrewes 1930, Arkiv for Zoologi 21A, No. 29, p.
5.
Description. None required here; length
c. 5.5-6.0 mm.
Types. Five, from Sumatra; actual
"type" in Stockholm Mus. (not seen).
Occurrence in Neio Guinea. Papua: 52
specimens from various localities, including
a series from Dobodura. N-E. N. G.: 1,
Nadzab, Julv 1944 (Darlington). West
N. G.: 1, River Tor (mouth) 4 km E.
Hoi Maffen, July 19, 1959 (T. C. Maa,
Bishop Mus.).
Notes. See Notes under the genus. I
have identified this species from descrip-
tion, and am not ((uite sure of it. It is, as
Andrewes says, about the si/c and color of
ii^onidius (below) but with lateral borders
of elytra darker and with differently formed
prothorax.
Stenolophus gonidius Bates
Bates 1890, Ann. Mus. C:iv. Ceiioa 27, p. 101.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 6,
p. 1262 (see for additional references).
Andrewes 1933, Cat. Carabidae Sumatra, p. 317.
Description. None required here- l(>ngth
c. 5.5-6.5 mm.
Types. From Burma; in Genoa Mus.
(not seen).
Occurrence in New Guinea. Papua: 31
specimens from Dobodura, Oro Bay, Port
Moresby, Fly R., and Ferguson Is. N-E.
N. G.: 1, Finschhafen, Huon Pen., 80 m,
Apr. 16, 1963 (Sedlacek), in Malaise trap.
( No specimens from ^^^est N.G. )
Notes. See Notes under the genus. Al-
though I have seen a cotype of gonidius
in the British Museum and have made com-
parisons with it, I am not quite sure of the
identity of the New Guinean specimens.
Genus ACUPALPUS Latreille
Latreille 1829, in Cuvier, Regne Animal, ed. 2,
4, p. 391.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 6,
pp. 1238, 1242.
Jeannel 1942, Faune de France, Coleop. Carabiqucs,
Part 2, p. 712.
Basilewsky 1951, Ann. Mus. Congo Beige (8),
Zool., 9, pp. 232, 233.
Diap,nosis. See preceding Key to Genera
of UarpaJini of Neic Guinea and following
Notes. I ha\'e tentatively assigned to this
genus all the small Harpalini of New
Guinea ( 6 species ) that possess lonii,
sparse prosternal setae. All are winged.
The front tarsi of the S are not or at most
(in furvinus) slightly dilated, with squamae
thin, transparent, difficult to detect, and
perhaps absent in some cases.
Description. None required here.
Type .■species. Carahus nicridianus Lin-
naeus, of Europe.
Generic distribution. Now considered to
include all principal regions of the world.
but the generic classification of these small
Harpalini is not lully worked out. At least
3 stocks of this genus reach nortliern tropi-
cal Australia, but the\ aiiparenth' do not
extend far into southern tempcMate Aus-
tralia, where tluMr place is taken In small
species of Lccanonwrtis (see discussion un-
der Tribe Harpalini).
Notes. Several species ol this genus (>ither
range widely in the Oriental-Australian area
or b(4()ng to wide-ranging groujis of closely
interrelaled species. Their nomenclature is
The Carabid Beetles of New Guinea • Darlington 73
difficult, and synonymies outside New
Guinea remain to be determined.
Key to Species of Acu palpus of
New Guinea
1. Anterior margin of clypeus not notched or
impressed at sides ( at most slightly sinuate ) ;
base of prothorax c. squarely truncate, with
posterior angles c. rectangular 2
- Anterior margin of clypeus notched or im-
pressed at sides; posterior angles of prothorax
usually more obtuse or rounded . 3
2. Larger (c. 3.7 mm); elytral margins behind
humeri faintly serrate at 50 X ( P- 73) __ cxactus
- Smaller (c. 2.7 mm); elytral margins not
visibly serrate at 50x (p- 73) -- exactellus
3. Prothorax with baso-lateral impressions not
punctate; posterior angles (narrowly) roimded
(p. 74) ftirvinus
- Prothorax with baso-lateral impressions punc-
tate; posterior angles usually more distinct ^ 4
4 Color brown with darker head; prothorax
with base more oblique at sides and posterior
angles more obtuse (p. 74) hrunnicolor
- Color darker, more uniform; prothorax with
base less oblique at sides and basal angles
more nearly right 5
5. Smaller (3.0-3.3 mm) (p. 75) ustus
- Larger (3.5-4.2 mm) (p. 75) papiia
Acupalpus exactus n. sp.
Description. With characters of genus;
form ( Fig. 31 ) relatively slender, elytra
subparallel; piceous, clypeus etc. reddish,
margins of prothorax rather broadly and
indefinitely reddish, suture and margins of
elytra narrowly but conspicuously reddish,
appendages testaceous; shining, dorsal
microsculpture faint or absent. Head 0.84
and 0.85 width prothorax; eyes large, promi-
nent; frontal impressions deep, converging
anteriorly, ending at deep clypeal suture;
anterior margin of clypeus slightly sinuate
but not distinctly notched at sides; man-
dibles long; mentum not toothed; ligula
slender, free at apex, 2-setose; paraglossae
slightly longer than ligula, narrowly
rounded; palpi with apical segments sub-
conical. Prothorax broadly subcordate;
width/length 1.34 and 1.30; base/apex 1.10
and 1.07; sides broadly rounded anteriorly,
slightly converging and broadly sinuate to
c. right, sharply defined posterior angles;
base and apex not margmed; lateral mar-
gins moderate, not crenulate; baso-lateral
impressions large, deep, irregular but not
distinctly punctate; disc normal, impressed
median line groove-like at base. Elytra
long; width elytra/prothorax 1.47 and 1.52;
humeri prominent but rounded; margins
behind humeri visibly serrate (at 50x);
striae deep, entire; intervals convex, 3rd
with puncture on inner edge well behind
middle. Measurements: length c. 3.7;
width 1.4-1.5 mm.
Types. Holotype 9 (M.C.Z., Type No.
31,389) from Hollandia, West N. G., July-
Sept. 1944 (Darlington); and 1 9 paratype
(Bishop Mus.) from Kiunga, Fly R., Papua,
July 15-21, 1957 (W. W. Brandt); 1 para-
type, Popondetta, Papua, 25 m. May 1966
( Shanahan-Lippert, Bishop Mus.), light
trap.
Notes. This New Guinean species resem-
bles and is probably related to Acupalpus
horni Andre wes of SE. Asia but is darker,
with lateral margins of prothorax not
crenulate (faintly crenulate in horni), and
with baso-lateral impressions of pronotum
less linear. Other apparently related forms
occur in the Philippines, Moluccas ( Morotai
Is.), and North Queensland in Austraha.
Acupalpus exactellus n. sp.
Description. With characters of genus;
smaller and relatively shorter than exactus
( above ) ; piceous, margins of prothorax and
elytra not or not conspicuously paler, ap-
pendages testaceous; moderately shining,
reticulate microsculpture (slightly trans-
verse ) visible on front anteriorly but absent
on discs of prothorax and elytra. Head 0.75
and 0.76 width prothorax; eyes relatively
much smaller than in exactus, with genae
more oblique; front similarly impressed;
clypeus without distinct notches at sides
anteriorly; mouthparts as in exactus. Pro-
thorax broadly subcordate with wide base;
width length 1.40 and 1.42; base/apex 1.24
and 1,16; sides broadly arcuate, slightly
converging and broadly sinuate to c. rec-
tangular posterior angles; base and apex
not margined; lateral margins narrow an-
74 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
teriorly, l:)ioader posteriorly, not crenulate;
baso-lateral impressions broad, deep, ir-
regular, but not distinctly punctate; disc
normal, with middle line deeply impressed
basally. Elytra: \\'idth elytra prothorax
1.41 and 1.45; humeri broad but rounded;
margin behind humeri not visibly serrate
(at 50x ); striae and dorsal punctures as in
exacttis. Secondary sexual characters: S
front tarsi not dilated and apparently with-
out squamae ( see Notes, below ) . Measure-
uicuts: length c. 2.7; width 1.3 mm.
Types. Holotype $ (M.C.Z., Type No.
31,390) and 1 9 paratype from Hollandia,
West N. G., July-Sept. 1944 (Darlington).
Notes. So far as I know, the only species
with which the present one need be com-
pared is exacttis (above), with which com-
parison is made in the Key and preceding
Description.
Although the type is a $ ( dissected ) , the
anterior tarsi are slender and without the
expected squamae, so far as I can deter-
mine under highest power of my stereo-
scopic microscope.
Acupalpus furvinus n. sp.
Descri])tion. With characters of genus;
form rather elongate; reddish piceous,
sides of prothorax and suture and sidt\s of
elytra morc^ reddish, appendages irregularly
testaceous, antennae brownish except at
base; shining, but front with light iso-
diametric microsculpture, pronotum with
more transverse microreticulations at most
hiintly indicated, elytra with distinct trans-
verse meshes (at 50x ) but sHghtK' irides-
cent. Head 0.71 and 0.73 width prothorax;
eyes moderate; front impressed as usual;
clypeus with anterior margin notched or
impressed at sides; mouthparts as in
e.xactus. Prothorax subquadrate; width
length 1.31 and 1.31; base/apex 1.10 and
1.11; sides broadly rounded anteriorly,
nearly straight (or slightly rounded) and
converging posteriorly to narrowly round(>d
posterior angles; base and apex not mar-
gined; margins moderate, broader jioste-
riorl\ and rniming into irregular, rounded.
not distinctly punctate baso-lateral impres-
sions; disc as usual. Elytra: width elytra/
prothorax 1.38 and 1.39; striae entire, well
impressed; 3rd intervals 1-punctate on inner
edge behind middle. Secondary .sexual char-
acters: i front tarsi slightly dilated, 4th
segments deeply emarginate, 4 segments 2-
seriately squamulose. Measurements: length
3.5-4.7; width 1.5-1.8 mm.
Types. Holotype $ (M.C.Z., Type No.
31,391 ) and 6 parat>'pes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
additional paratypes as follows. Papua: 1,
Lake Daviumbu, Flv R., Sept. 21-30, 1936
(Archbold Exp., A.M.N.H.). N-E. N. G.:
1, Aitape, Aug. 1944 (Darlington).
Measured s))eci7nens. The 6 holotype and
1 9 paratype from Dobodura.
Notes. According to my notes made at the
British Museum in 1948, this species rep-
resents a widely distributed Oriental-Aus-
tralian group of Acupalpus which includes
annarncnsis Bates and jurvus Andrewes of
SE. Asia, as well as various named forms
from the Malay Archipelago. Most of them
are testaceous with dark elytral clouds, but
jurvus is more uniformly colored, as is
furvimis. However, the latter differs from
jurvus in haxing more prominent eyes and
less rounded posterior prothoracic angles.
A representati\ e of this species group oc-
curs in North Queensland, Australia.
Acupalpus brunnicolor (Sloane)
Sloane 1S98, Proc. Linnean Soc. New South Wales
23, p. 466 (Thcnarotcs).
Andrewes 1930, Cat. Indian Inseets, Part 18,
Caral)i(Iai', p. 10.
Description (for recognition only). A
brown Acupalpus characterized in preced-
ing Key to Species: head 0.82 and 0.82
widtli prothorax; prothoracic width length
1.44 and 1.42, base apex 1.13 and 1.16;
width elytra prothorax c. 1.43 and 1.38;
length c. 3.8, width 1.5-1.6 nun.
Types. From Behn Ri\er, Wcslcru Aus-
tralia, collected 1)\' Helms; t\pe retinned
to Lea, should be in Sonlh Australian Mns.
(not secMi).
The Carabid Beetles of New Guinea • Darlington 75
Occurrence in Neiv Guinea. Papua: 3,
Port Moresby, Oct. 1944 (Darlington); 1,
Orionio River, Feb. 17, 1967 ("H. C,"
Bishop Mus.), light trap.
Measured specimens. A pair ( c5 9 ) from
Port Moresby.
Notes. According to notes that I made
at the British Museum in 1947-1948, brun-
nicolor of Avistralia may be a form of a
widely distributed species that has received
the name sinuellus Bates in SE. Asia and
punctafus Jedlicka in the Philippines. How-
ever, the classification of these small har-
palines is still so doubtful that I do not
wish to declare synonymies, but shall say
only that hrunnicolor probably represents a
widely distributed species group that may
have reached Australia recently (perhaps
by way of the Lesser Sunda Islands) and
that may then have spread from Australia
to the Eucalyptus country of southern New
Guinea.
Acupalpus ustus Andrewes
Andrewes 1930, Zool. Mededelingen 13, p. 195.
Description ( for recognition only ) . With
characters of genus; form rather stout;
piceous, moderately shining. Head 0.76
and 0.77 width prothorax; eyes average.
Prothorax subcordate; width/length 1.38
and 1.37; base apex 1.12 and 1.10; sides
variably sinuate before obtuse (almost
right) posterior angles; disc normal, baso-
lateral impressions punctate, punctation ab-
sent or sparse across middle of base. Elytra:
width elytra/prothorax 1.43 and 1.44. Mea-
surements (in New Guinea): length 3.0-
3.3; width l..'3-1.4 mm.
Types. From Sumatra and Borneo; the
(holo)type, from Borneo, in Andrewes
Coll., British Mus. (seen).
Occurrence in Neiv Guinea. Papua: 29,
Dobodura, Mar.-July 1944 (Darlington).
West N. G.: 1, Hollandia, July-Sept. 1944
( Darlington ) .
Measured specimens. A pair { i 9 ) from
Dobodura.
Notes. My identification is tentative, al-
though based on comparison of specimens
with the type. As in other species of this
genus, I am not absolutely sure of synon-
ymies and make none, but only state that
this species appears to be widely distributed
in the Malay Archipelago.
Acupalpus papua n. sp.
Description. With characters of genus;
form (Fig. 32) rather broad, with sides of
elytra shghtly arcuate; piceous, apices of
elytra sometimes rufescent, appendages
testaceous, antennae browner from 3rd seg-
ments; moderately shining, reticulate micro-
sculpture distinct and c. isodiametric on
front, faint ( and more transverse ) or absent
on disc of pronotum, not visible (at 50x )
on slightly iridescent elytra. Head 0.82 and
0.80 width prothorax; eyes moderate; frontal
impressions extending onto clypeus; anterior
edge of clypeus finely notched at sides;
mouthparts c. as in exactus. Prothorax
broadly subcordate; width/length 1.43 and
1.46; base/apex 1.15 and 1.10; sides broadly
rounded anteriorly, converging and slightly,
broadly sinuate before slightly obtuse but
well defined posterior angles; margins
moderate anteriorly, broader posteriorly,
with baso-lateral areas depressed and punc-
tate, punctation not extending across middle
of base. Elytra: \\'idth elytra/prothorax
1.46 and 1.48; humeri broadly rounded;
margins behind humeri not distinctly serrate
at 50X; striae moderately impressed; 3rd
intervals 1-punctate on inner edge as usual.
Measurements: length 3.5-4.2; width 1.5-
1.8 mm.
Types. Holotype S (M.C.Z., Type No.
31,392) and 4 paratypes from Dobodura,
Papua, Mar.-July 1944 (DarHngton); and
additional paratypes as follows. Papua: 1,
Oro Bay (near Dobodura), Dec. 1943-Jan.
1944 (Darlington); 1, Fly R., Lake Davi-
umbu, Sept. 1-10, 1936 (Archbold Exp.,
A.M.N.H.). West N. G.: 2, Hollandia,
July-Sept. 1944 (Darlington).
Measured specimens. The c^ holotype and
1 ? paratype from Dobodura.
76 BuUetin Museum of Comparative Zoology. Vol. 137, No. 1
Notes. Except that it is close to tistiis
( above ) , I cannot state the relationships of
this new species. As compared with iistu-s,
papim is larger, with relatively slightly
w ider head and wider prothorax.
Tribe ANAULACINI
Csiki 1932, Coleop. Cat. Carabidae, Harpalinae 7,
p. 1287 ( see for synonymy and additional refer-
ences ) .
Jedlicka 1963, Ent. Abhandlungen 28, p. 283.
MiLsoreini Auct.
Andrewes 1930, Cat. Indian Carabidae, p. XIII.
Jeannel 1949, Coleop. Carabiques de la Region
Malgache, Part 3, p. 860.
Masorcitdc jeannel 1942, Fainie de France, Coleop.
Carabiqnes, Part 2, p. 1012.
Ma.sorcinac Basilewsky 1953, Exploration Pare Na-
tional rUpeniba, Fasc. 10, Carabidae, pp. 17,
118.
This is a small, pan-tropical tribe of
obscure ('arabidae some of which, includ-
ing all those found in New Guinea, super-
ficially resemble Nitidulidae. They may be
recognized by appearance ( Figs. 33, 34 ) ;
short, strongly arcuate, flattened mandibles;
rather long tibial spurs; and other technical
characters given by authors cited above.
y\ll the species that I know are winged,
and the widely distributed Aephnidius
adelioides flies to light, but Odontomasoreus
has not been taken at light and perhaps
do(\s not fly. The few species that I have
collected in New Guinea and Australia live
in leaf litter on the ground in rain forest.
Four genera of the tribe are known from
New Guinea.
Kkv to Genera of Anaulacini of
New Guinea
1. iiimicii dentate; (antennae short, not reacli-
ing base of prothorax; labrnni rounded;
mentnni lobed or snlidentate; chtra pale-
spotted) (p. 76) Odoiilomd.soreus
- Ilnnieri not dentate 2
2. Antennae sliort, not reaching base of pro-
tliorax (p. 77) Anaiilcicu.s
- Antennae longer, reaching or passing base
of prothorax 3
3. Mcntuiii not loolhed; size larger, c. 5 mm
(p. 77) Acp]iniditis
- Mentuni toothed: smaller, less than 3 mm
(p. 78) Caphoia
ODONTOMASOREUS n. gen.
Diagnosis. Rather small, convex Anau-
lacini; immediately distinguished from other
genera of tribe by humeri dentate.
Description. Form nitiduloid (Fig. 33),
convex; color piceous with pale elytral
marks; reticulate microsculpture isodiamet-
ric on head, slightly and irregularly trans-
verse on pronotum and elytra. Head nor-
mal, c. as in Aephnidius adelioides except
labrum rounded; antennae short, reaching
r. middle of prothorax, with median seg-
ments wider than long; mentum blimtly
toothed or obtusely prominent at middle.
Prothora.x normal; anterior angles mod-
erately advanced; base sinuate but scarcely
lobed; disc with fine, abbreviated middle
line. Elytra: humeri finely dentate; mar-
gins weakh' sinuate near apex; striae in-
dicated but scarcely impressed, scutellar
striae not visible. Secondary .sexual char-
acters: i tarsi slightly dilated, 3 segments
2-seriately squamulose; i copulatory organs
as in Figure 175.
Ty))e .species. Odontomasoreus humcralis
( below).
Generic distribution. Known (^ily from
New Guinea, thus far only from the east-
ern and central parts of the island.
Notes. I recognize only 1 species of this
new genus, with 2 subspecies.
Key to Subspecies of
Odontomasoreus fii mfrai.is
1 . Larger ( 3.4-4.0 mm ) ; humeri broadh pale
(Papua) (p. 76) humcralis ,9.,9.
- Smaller (3.1-3.5 mm); humeri dark or at
most with \ague or small pale areas (N-E.
N. G. and eastern \\'e.st N. G.) (p. 77) ^
subsp. rcdurliis
Odonfomasoreus humeralis n. sp.
Descri])lion. With cliaraclers o\ genus;
form as in Figure 33; brownish piceous,
humeri, base of elytra, and an elongate mark
on 2n(l inter\al of each elytron near apex
testaceous; mouth parts and ai)pendages
brownish or testaceous. Jlcad 0.60 and
0.63 width prothorax. ProlJiorax widest
near base, narrowed anteriorh-; width/
The Carabid Beetles of New Guinea • Darlington 77
length 1.81 and 1.81; base/apex 1.42 and
1.39; sides weakly arcuate, very narrowly
margined, each with usual 2 setae, at base
and c. -i from apex; base finely margined,
apical marginal line weak or interrupted
at middle; disc vaguely impressed each
side before base. Elytra: width elytra/
prothorax 1.10 and 1.10. Measurements:
length 3.4-4.0; width 1.7-1.8 mm.
Types. Holotype $ (M.C.Z., Type No.
31.393) and 21 paratypes all from Dobo-
dura, Papua, Mar.-July 1944 (Darlington).
Measured specimens. The i holotype and
1 9 paratype.
Notes. Most specimens of the type series
were taken in flood water in rain forest
after heavy rainfall. The insect evidently
inhabits leaf litter and perhaps loose soil
in rain forest.
Odonfomasoreus humeralis reductus
n. subsp.
Description. Similar to typical humeralis
but smaller, with the basal pale areas of
elytra reduced or absent but subapical
marks distinct. Head 0.62 and 0.62 width
prothorax. Prothorax: width/length 1.80
and 1.86; base/apex 1.45 and 1.41. Elytra:
width elytra prothorax 1.08 and 1.08. Mea-
surements: length 3.1-3.5; width 1.5-1.6
mm.
Types. Holotype $ (M.C.Z., Type No.
31.394) and 6 paratypes from Hollandia,
West N. G., July-Sept. 1944 (Darlington).
Additional paratypes from N-E. N. G.: 1,
Astrolabe Bay, 1898 ( Biro ) ; 5, Aitape, Aug.
1944 (Darlington).
Measured specimens. The $ holotype and
1 9 paratype from Hollandia.
Genus ANAULACUS Macieay
Macleay 1825, Annulosa Javanica, p. 22.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1292 (see for additional references).
Jedlicka 1963, Ent. Abhandlungen 28, p. 286.
Diagnosis. See preceding Key to genera.
Description. None required here.
Type species. Anaulacus sericeipennis
Macleay, of Java.
Generic distribution. Tropical Asia to
the Philippines and New Guinea; South
Africa.
Notes. One species (in fact, only 1 in-
dividual) of this genus has been taken in
New Guinea.
Anaulacus siamensis Chaudoir
Chaudoir 1876, Bull. Soc. Nat. Moscow 51, Part 2,
p. 25.
'hterhai Jedlicka 1934, Sbornik Ent. Odd. Nar.
Mus. Prague 1934, p. 119.
?kendengensis Louwerens 1952, Treubia 21, p. 215.
Description ( for recognition only ) . With
characters of genus; fonn (Fig. 34) of
Aephnidius but antennae relatively short;
reddish piceous without well defined mark-
ings. Head 0.66 width prothorax. Prothorax:
width/length 1.74; base/apex 1.34. Elytra:
width elytra/ prothorax 1.08. Measurements:
length c. 4.5; width c. 1.9 mm.
Type. From Siani; in Oberthiir Coll.,
Paris Mus. (not seen).
Occurrence in Neiv Guinea. West N. G.:
1, Geelvink Bay, 1878 (Raff ray & Maindron,
Paris Mus.).
Notes. This individual is identified from
description and from notes, made at the
British Museum in 1947-1948, on a speci-
men from the Andaman Islands identified
as siamensis by Andrewes. The unique
type of sterbai, from Malinao, Tayabas,
Philippine Is., is in the British Museum
too; it does not differ significantly from
the Andaman siamensis. I have a paratype
of kendengensis from Java, and it too is
very close to siamensis. All these names
probably apply to one species that ranges
from the southeastern corner of Asia to the
Philippines and New Guinea, but the
material seen is too limited to justify a
final decision.
Genus AEPHNIDIUS Macleay
Macleay 1825, Annulosa Javanica, p. 23.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1288 (see for additional references and list
of species).
Jeannel 1949, Coleop. Carabiques de la Region
Malgache, Part 3, p. 861.
Jedlicka 1963, Ent. Abhandlungen 28, p. 284.
78 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Diag,mms. See preceding Key to genera.
Description. None required here.
Type species. AepJmidiiis adelioides Mac-
leay (below).
Generic distribution. All principal trop-
ical and some adjacent warm-temperate
areas of world.
Notes. A single widely distributed species
of the genus occurs in New Guinea.
Aephnidius adelioides Macleay
Macleay 1<S25, Animlosa javaiiita, p. 23, pi. 1,
fig. 7.
Andrewes 1930, Cat. Indian Insects, Part IS,
Caral)idae, p. 11.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1288 (see for synonymy and additional refer-
ences ) .
Description. None required here; length
± 6 mm.
Type. From Java; in British Mus. (seen).
Occurrence in New Guinea. Widely dis-
tributed: 29 specimens from Papua (in-
cluding Dobodura), N-E. IS. G., and West
N. G., at low and moderate altitudes, up
to 1200 m (at Wau).
Notes. This species ranges from SE.
Asia, Japan, and Formosa to northern
Australia, east to the Philippines, New
Britain, and New Ireland. Seven speci-
mens from Wau and 1 from near Hollandia
are labeled as taken in light traps.
Genus CAPHORA Schmidt-Goebel
Schmidt-CJoebel 1846, l''aiinula (Coleop. Hiiiiuiniac,
p. 91.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1293 (see lor additional relerences).
Jedlicka 1963, Knt, Abliandlnni^'en 28, p. 288.
Diapiosis. Very small Anaulacini ( under
3 mm); characterized in the preceding Key
to (kniera of the tribe.
Description. None refjuired lu^e.
Type species. Capltora liuniilis Scliuiidl-
Goebel ( below ) .
Generic distrihntion. SE. Asia, Sumatra,
Java, the Philippin<>s, New (>iiinea, and
Cajie York. Australia (see following
species ).
Caphora humilis Schmidt-Goebel
Schmidt-Goebel 1846, Fannula Coleop. Birnianiae,
p. 91, pi. 3, fig. 8a-b.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1293 (see for additional references).
Jedlicka 1963, Ent. Abhandlnngen 28, p. 289.
Description. None required here. The
very small size distinguishes this species
from all other members of the tribe found
in the region in question. Length c. 2.5 mm.
Type. From Burma; should be in Prague
Mus. (not seen).
Occurrence in New Guinea. Papua: 1,
Brown River, May 24, 1956 (E. J. Ford, Jr.,
Bishop Mus.), in light trap.
Notes. This species is recorded from
India and Burma to Sumatra and Java
and occurs also on the tip of Cape York,
Australia (collected by me in 1958). My
Cape York specimens were found in the
Lockerbie rain forest, in leaf litter mixed
with bird droppings under a large tree in
which colonial birds had nested. Tlie
beetles were in company with Peri<i.ona
niiiriceps, which is often carried by com-
merce, and this suggests that CapJiora too
may sometimes be carried by man.
Tribe CYCLOSOMINI
This is another small tribe, repr(\sented
in all the warmer rt>gions of the world.
The name to use for it is doubtful but not
worth detailed discussion here. The mem-
bers of the tribe, although apparently re-
lated to Anaulacini, are superficialh' Lel)ia-
like but diffcT from Lebia iu having very
long tibial spurs. The only genus of the
tribe that occurs iu New Guinea is
Sarotlirocrepis.
Genus SAROTHROCREPIS Chaudoir
Chandoir 1876, linll. Soc. Nat. Mo.scow 51, Tart 2.
p. 76.
(>siki 1932, Coleop. ('at., ("arabidae, Harpalinae 7,
p. 1302 (see lor s\n()n\Mn\', additional refer-
ences, and list of species ) .
Jedlicka 1963, Knt. Abliandlunuen 2S. p. 290.
Diaii,)}osis. Sec undc^r tribe.
Dcscri))ti()n. None reciuired lieic.
The Carabid Beetles of New Guinea • Darlington
79
Type species. Carahus corticalis Fab-
riciiis, of Australia.
Generic distribution. Represented in
Australia by numerous, diverse species; 1
species group extends to New Guinea,
some Lesser Sunda Islands, Celebes, and
the Philippines.
Notes. Although most other genera of
this tribe are (I think) ground-living,
Sorotlirocrepis is arboreal. Many Australian
species live on the trunks of Eucalyptus
trees, but a few live in foliage, as does the
single New Guinean species. The tarsal
claws tend to vary with habitat. In the
foliage-living New Guinean species and also
its immediate Australian relatives, each
claw has several long teeth. In some Aus-
tralian tree-trunk-living forms, the claw
teeth are shorter or irregular.
Sarothrocrepis papua n. sp.
Description. Form as in Figure 35; ir-
regular bro\\'nish yello^^', elytra with vari-
able post-median dark brown mark usually
irregularly triangular or M -shaped; reticu-
late microsculpture isodiametric on front,
scarcely or slightly transverse on pronotum,
more transverse on elytra. Head 0.67 and
0.67 width prothorax. Frothorax: width
length 1.45 and 1.50; h-Ase head 1.41 and
1.41 (apex of prothorax too rounded for
exact measurement ) ; margins moderate an-
teriorly, much wider posteriorly ( as usual in
genus), each with seta at basal angle and
another c. ''.•! from apex; base finely mar-
gined, apical marginal line interrupted at
middle; median line lightly impressed, sub-
apical transverse impression weak, subbasal
transverse impression and posterior-lateral
impressions slight. Elytra: width elytra
prothorax 1.52 and 1.48; striae deeply im-
pressed, not punctate; a seta-bearing punc-
ture at base each 2nd interval, usually an
inconspicuous puncture on inner edge each
3rd interval near apex, and sometimes an
apparent minute puncture on outer edge
3rd interval c. V-i from base. Inner u'inii,s
full. Lower swface and legs: no note-
worthy characters except tarsal claws each
with 4 long teeth, the inner one smaller and
sometimes difficult to see. Secondary sexual
characters: i front tarsi slightly dilated, 3
segments 2-seriately squamulose (4th seg-
ments with soles of non-sexual adhesive
hairs in both sexes); S (not 9 ) apical
ventral segment deeply acutely notched at
middle; 1 seta each side apex last ventral
segment in both sexes. Measurements:
length 4.6-6.4 (usually 5.0-5.5); width 2.3-
3.0 (usually c. 2.7) mm.
Types. Holotype c^ (M.C.Z., Type No.
31,395) and 120 paratypes all from Dobo-
dura, Papua, Mar.-July 1944 (Darlington).
Additional material. Thirteen specimens
from other localities in Papua, N-E. N. G.,
and West N. G.; and 2 specimens from
Cape Gloucester, New Britain, Jan.-Feb.
1944 (Darlington).
Measured .specimens. The holotype and
1 9 paratype.
Notes. Sarothrocrepis papua resembles S.
fasciota Macleay of North Queensland, Aus-
tralia, but is larger, with prothorax narrower
and elytral markings slightly different.
Three similar species occur in the Malay
Archipelago. S. m-miorum Jordan, from
Tenimber (and in Andrewes Coll. also
from Sumbawa, Sumba, and Andonare Is. ) ,
has prothoracic margins narrower than
papua and the dark M-mark usually better
defined (but the mark is variable in papua).
S. javanica Van Emden has prothoracic
margins narrower and basal impressions of
pronotum more linear. And S. andrewesi
Jedlicka, of the Philippines, has the elytral
marks different (3 dark stripes on yellow
background) and basal impressions of
pronotum better defined, sublinear. These
names may all be based on forms of one
widely distributed variable species, but I
prefer to treat them as separate species
for the time being.
S. papua was very common in under-story
foliage of rain forest at Dobodura, espe-
cially in clumps of dead leaves still attached
to low branches. It is probably mainly
diurnal, although I have seen 2 specimens
taken in light traps.
80
Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Tribe LEBIINI
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1305 ff. (see for synonymy and additional
references ) .
Jedlicka 1963, Ent. Abhandlungen (Dresden) 28,
p. 295.
Habu 1967, Fauna Japonica, Carabidae, Trun-
catipennes Croup, p. 57.
Lehiidae Jeannel 1942, Faune de France, Coleop.
Carabiques, Part 2, p. 1017.
Teannel 1949, Coleop. Caraliiques de la Region
Malgache, Part 3, p. 876.
Lehiinae Basilewsky 1953, Exploration Pare Na-
tional rUpemba, Fasc. 10, Carabidae, p. 184.
Lebiini are medium sized and small
Carabidae usually recognizable by general
appearance. Tlie elytra are usually (but
not always) obliquely truncate or sinuate-
truncate at apex (and often spined too),
and the insects have additional technical
characters, including genitalic ones, given
by authors cited above.
The tribe is a very large one. It is best
represented in the tropics and includes a
large proportion of arboreal forms especially
in the tropics.
Arboreal Lebiini divide into two ecologic
groups, one living on tree trunks and the
other in foliage. The tree-trunk dwellers
are numerous in rain forest and some occur
on fallen trees and logs as well as the trunks
and larger branches of living trees. Some
are nocturnal, some diurnal. Some of them
can be trapped under sacking tied around
tree trunks or laid over logs. Lebiini living
on tree trunks and logs in New Guinea
include Stcnotchis, Mi.scclus, Mimtthodcs,
C(it(isco))iis\ Pcricahis, Coptodcra (some),
Moclifhcrus, and Strickhindia. (Other lebi-
ine genera, especially PJidopldocus, A'^ono-
cliila, and Australian Demetrida, live on the
trunks of Eucalijpius trees in more open
woodland in Australia. ) The foliage dwel-
lers in New Guinean rain forest include
Aristolehia, Lehia, Dolichoctis, Cclacncphcs,
and especially New Guinean Demetrida.
(The ecology of Demetrida is discussed in
more detail in Note.s' imder the genus. )
Different foliage-inhabiting Lebiini in New
(Uiinea probabK inhabit dilfcrent sj^jccial
niches. Some species are commonly taken
by sweeping under-story plants in rain
forest but other species are not, and these
may live at higher levels in the trees. They
may be difficult to collect except when trees
are felled, unless they fly to light. Be-
sides the arboreal forms, a few Lebiini in
New Guinea ( and relatively more in colder
climates) live on the ground, especially in
leaf litter in rain forest. In New Guinea,
these include some Coptodcra, probably
Sijntomus- and Microlestes and Apri.stus, and
certainly Anomotarus and Nototonis. The\
can be collected in small numbers, labori-
ously, by sifting or in Berlese traps, or in
larger numbers and more easily by washing
out debris and loose earth from the forest
floor or by sorting and washing flood debris
from rain forest.
Most Lebiini are winged and many of
them fly actively to escape danger or to
disperse (see Notes under Catascojm.s),
and some, presumably mostly nocturnal
species, fly to light at night. The only
known New Guinean lebiine with atrophied
wings is Nototariis papua.
Although Lebiini are most numerous and
diverse in the tropics, the tribe as a whole
is virtualK' cosmopolitan. A few genera,
including Lehia and Coptodera, are very
widely distributed too, but most other
genera have restricted distributions.
Thirty genera of Lebiini are known in
New Guinea and at least 3 additional
genera probably occur. Many of the genera
are shared with, better represented in. and
probably derived from the Oriental Region.
These include Lehia, Catascopus, Coptodera.
Dolichoetis, and more than a do/en smaller
genera. Genera shared maiuK with and per-
haps derived from Australia are fewer but
include Aiionochda. TriiionotJiops, Phloeo-
earahii.s, Aiioniofarus, Xototarus. and espe-
cially Denielridu. The genera Miniithodes.
Striekkindia, and Miseelus now center on
New Guinea and may have originated there.
The onl\- l(>biine genus actualK confined
to New (Guinea is monotxpic Minitj)hJoeus.
The uenus Demetrida seems to hv in the
The Carabid Beetles of New Guinea • Darluii'ton
81
very midst of an explosive evolutionary
radiation, which is discussed under the
genus. I know no other case quite like it
among Carabidae.
The following key to the genera of
Lebiini of New Guinea is practical, not
phylogenetic. Genera that occur together
in the key are not necessarily closely re-
lated, and the key is designed for onJy the
New Guinean species of some genera. I
haN'e used form of the whole insect as a key
character of some genera. The form is
characteristic in some cases, and the form
of the whole is surely no less important
than the form of a part.
After the key, the genera are treated in
the order of the Coleopterorum Ccitalogus
(Csiki 1932), not in the order in which
they are keyed out.
An enormous supposed lebiine, Holopon-
erus godeffroiji (Fairmaire), has been de-
scribed from New Britain. I do not know
this insect, but I think it is probably not
a member of the Lebiini but of the tribe
Helluodini, under which I shall discuss it in
more detail.
Key to Genera of Lebiini of New Guinea
1. Fourth segments of hind tarsi deeply eniar-
ginate, with lobes \-2 or more total length
of segment 2
- Fourth segments of hind tarsi more shal-
lowly emarginate or subtnincate 10
2. Form usually broad Lebia-\ike ( Figs. .37-
41 ); base of prothorax ± lobed; upper sur-
face not pubescent; tarsi not puljescent
above; S middle tibiae excised on inner
edge near apex 3
- Form not LebiaAike, usually more slender;
l)ase of prothorax often ( not always ) with-
out lobe; tarsi often (not always) pubes-
cent above; S middle tibiae usually not
excised ( but tuberculate-serrate in many
Demetrida ) „ 4
3. Outer-apical angles of elytra sharply
formed and prothorax ± hemispheric; S
middle tibiae with 2 excisions on inner
edge near apex ( p. 83 ) Ari.stolehia
- Outer-apical angles of elytra either rounded
or sharply formed, but if latter, prothorax
not hemispheric (in New Guinean species);
$ middle tibiae with 1 excision on inner
edge near apex (p. 85) Lebia
4. Small (4-4.5 nun); protliorax trapezoidal,
widest at base (Fig. 87) (p. 134)
Peliocypas
- Larger; prothorax usually not as de-
scribed 5
5. Upper surface coarsely rugose and pubes-
cent; prothorax strongly lobed at base;
(form as in Fig. 42; length c. 8 mm)
( p. 89 ) Lachnodenna
- Upper surface not coarsely rugose; pubes-
cent or not, ])ut if pubescent, prothorax
not lobed at base 6
6. Fifth elytral intervals with coarse seta-
bearing puncture near base; prothorax with
( very short ) basal lobe ( form as in Fig.
95; length c. 8-9 mm) (p. 139) ____ Anclmta
- Fifth elytral intervals without coarse punc-
ture near base; prothorax usually (not
always) without basal lobe 7
7. Form (Figs. 97-109) usually slender; apex
of elytra sinuate-truncate or angulate or
spined but not broadly and strongly
rounded; tarsi pilose above (p. 140)
Demetrida
- Form usually less slender; apex of elytra
broadly and strongly rounded or weakly
sinuate-truncate with outer angles broadly
rounded; tarsi above pilose or not 8
8. Prothorax with (slight) basal lobe; tarsi
not pilose above (p. 184) Trigonothops
- Prothorax without basal lobe (but base
slightly oblique near angles); tarsi usually
pilose above (pilosity slight in some
Parena) 9
9. Upper surface not pubescent (p. 138) .. Paremi
- Upper surface pubescent ( p. 140 )
Endtjnomena
10. Form (Fig. 37) characteristic, broadly
oval with outer elytral angles sharply
formed; £ middle tibiae with 2 excisions
on inner edge near apex; ( length c. 9.5—
11.0 mm — smaller species of same genus
key out in couplet 3) (p. 83) ..^^ Aristolebia
- Form not as above; S middle tibiae usu-
ally not 2-excised _-_ 11
11. Form (Fig. 36) characteristic; small (c.
4 mm or less); upper surface pubescent;
prothorax with extra anterior-lateral setae;
brown with single broad transverse dark
band across elytra (p. 82) ._- (SomotiicJtus)
- Not as above in one or more ways 12
12. Very small (less than 4 mm); color black
or (rarely) transversely fasciate 32
- Larger or, if length less than 4 mm, with
color pattern of longitudinal lines 13
13. Form (Fig. 88) characteristic, slender,
with rounded elytral apices; mentum with-
out tooth and claws not toothed; (black;
length c. 7 mm) (p. 135) Celaenephes
- Not as above 14
82
Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
14. Claws simple, not toothed; size usually
large; color often metallic _ - 15
- Claws each with several teeth; size often
(not always) smaller; color rarely metal-
lic 17
15. Form (Fig. 45) characteristic, slender,
subcylindric, with long genae and small
eyes, and with rounded-truncate elytral
apices; often (not always) only 1 seta
over eye; (length 9.5-14.5 mm) (p. 91) _.
Miscelus
- Form not as above; 2 setae over each
eye 16
16. Labrum long, notched at apex; size larger
( 8-22 mm ) ; color metallic or rarely brown,
without geometric marks (p. 101)
Catascopus
- Labrum shorter, not notched; smaller (7-
8 mm in New Guinean species); elytra
with geometric marks (p. 110) „.. Pericohis
17. Mentum not toothed 18
- Mentum toothed 19
18. Third elytral intervals with 2-4 dorsal
punctures, lint if 2, not as described be-
low; i middle tibiae usually excised on
inner edge before apex (p. 110) .. Coptodera
- Third elytral intervals with 2 minute non-
seta-bearing punctures behind middle, or
without recognizajjle dorsal punctures; $
middle tibiae not excised (p. 124)
_ Dolichoctis
19. Elytra spined 20
- Elytra not spined 21
20. Prothorax without extra lateral setae; form
(Fig. 44) not .strikingly broad (p. 90) _
Stciiotcltis
- Prothorax with many extra lateral setae;
form (Fig. 86) strikingly broad (p. 132) ..
Stricklatulia
21. i""orm ( F'ig. 43) characteristic, broad, with
broadly rounded elytral apices; (dull
black; lengtli c. lO-ll mm) (p. 90) _..
Siniiriis
- Not as above 22
22. Elytral apices very strongly sinuate-
truncate; (slender; color green, blue, or
coppery; length in New Guinea c. 8-9
mm ) ( p. 94 ) Holcodcrus
- Not as above __ __ 23
23. Form (Figs. 47-58) characteristic, very
broad, with wide head but relatively small
eyes, prothorax usually c. 2x wide as long,
elytra short-cjuadrate; (pubescence and
color diver.se; length c. 4-6.5 mm ) ( p.
95) Miuutlwdes
- Form not as above (if c. similar bul labrum
notched, .see Minupldoeiis, below) 21
24. Labrum notched at apex; pronotiun with
numerous lateral setae; ( shining black;
length 7.5-8.0 mm) (p. 117) ._.. Minuphloeti.s
- Labrum not notched; pronotum with 2 lat-
eral setae each side 25
25. Third elytral intervals with 3 or 4 dorsal
punctures or (in some Agonochila) these
pimctures lost amid other coarse puncta-
tion and short pubescence ^._ 26
- Third elytral intervals with 2 or rarely 1
dorsal punctures 27
26. Surface conspicuously short-pubescent and
(at least on elytra) roughened (p. 118) __
Agonochila
- Surface not distinctly pubescent, not
roughened; (3rd elytral intervals with 3
pimctures) (p. 122) Oxijodontus
27. Labial palpi slender 28
- Labial palpi with apical segments ±
widened, usually subtriangular 29
28. Third elytral inter\als 2-punctate; pro-
notum setulose (p. 122) Mochthcrtis
- Third elytral intervals 1-punctate; pro-
notmn not setidose ( p. 123 ) ^- (Mochtheroides)
29. Antennae and tarsi relatively short and
thick; i middle tiliiae arcuate and with
shallow excision at middle of length; (color
brown; length c. 8 mm) {p. 138)
( Ploch ion us )
- Antennae and tarsi more slender; S middle
tibiae not as described 30
30. Eyes abruptly prominent, genae short and
forming c. right angles with neck ( p.
183 ) Pldococarahus
- Eyes less prominent, genae longer and
forming obtuse angles with neck 31
31. Side pieces of metasternum long; inner
wings full; color pattern usualh' present
(p. 186) -- Anomotarits
- Side pieces of metasternum scarcely longer
tliau wide; inner wings vestigial; color r.
uniform brownish black (p. LSo) Xototcini.s
32. Claws simple, not tootlicd; ( incntuin with
entire tooth) (p. 137) A;)n.s7(/.s
- Claws each with se\ eral [ sometimes weak )
teeth .-.- 33
33. Mentum with (emarginate) tooth {p.
135) ___ _ _ Syntoimi.s
- Nhntuin not toothed (p. 136) .... Microlc'stcs
(Genus SOMOTRICHU5 Seidlitz)
Seidlitz 1887, Fauna lialtica, 2nd cil,, Cattungcn.
p. 7.
Nhiteu 1963, Ann. Mus. C.W. Genoa 74, pp. 131 ff.
(See also references under following species)
Didi^iiosis. Form as in I'imire 36. .small.
The Carabid Beetles of New Guinea • Darlington
83
subparallel, with eyes rather small and
\\idely separated (in this tribe); upper
surface pubescent; pronotum with several
strong setae each side; wings full; 4th hind-
tarsal segment weakly emarginate.
Description. None required here.
Type species. Carahus elevatiis Fabricius
(below).
Generic distribution. One species has
been dispersed over the wanner parts of
the world by man. A second species is
known only from Madagascar.
(Somotrichus elevatus (Fabricius))
Fabricius 1787, Mantissa Insectorum 1, p. 198
( Carahus).
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1308 (see for synonymy and additional refer-
ences ) .
Jeannel 1942, Faune de France 40, Coleop.
Carabiques, Part 2, p. 1032.
Description ( for recognition only ) . With
characters of genus; brown with broad,
regular, darker brown fascia across middle
of elytra; length c. 3.5-4.0 mm.
Types. From tropical America; now in
Hunter Coll. (Glasgow) and Fabricius
Coll. (Kiel) (not seen).
Occurrence in New Guinea. Not recorded
but may occur.
Notes. Somotrichus elevatus is sup-
posedly native in tropical Africa but has
been carried over much of the world by
commerce. It is often found in seaport
cities. In the Malay Archipelago it has
been collected on Java, Celebes, and
Batjan ("Batchian") in the Moluccas, and
I have a specimen before me from Peleliu
in the Palau Is. Its occurrence in New
Guinea is therefore likely. It has not yet
been found in Australia.
Genus ARISTOLEBIA Bates
Bates 1892, Ann. Mus. Civ. Genoa 32, p. 428.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1308 (see for additional references, synonymy,
and list of species ) .
Jedlicka 1963, Ent. Abhandlnngen 28, p. 311.
Diagnosis. Similar to large Lehio with
prothorax c. hemispheric and outer elytral
angles sharply defined; wings full; 4th seg-
ments middle and hind tarsi emarginate or
lobed (see following Notes); claws with
9-11 long teeth in larger species but only
5-7 teeth in smaller species; c^ middle tibiae
each with 2 (not 1 as in Lebia) excisions
close together on inner edge near apex; most
other characters including those of mouth-
parts as in Lebia.
Description. None required here.
Type species. Aristolebia quadridentata
Bates, of Bunua.
Generic distribution. Southern India
(specimens in M.C.Z.), southern China,
Burma, etc., to the Philippines, New
Guinea, and the tip of Cape York, Aus-
tralia (see under A. wau, below).
Notes. The smaller species described be-
low are in some ways transitional between
Aristolebia and Lebia, but Aristolebia seems
to me to be a natural group worth distin-
guishing from Lebia, which is an enormous,
unwieldy genus. The tarsal lobes in
Aristolebia vary, but the variation shows
continuity. In large Asiatic species of the
genus the 4th segments of the hind and
middle tarsi are relatively weakly emar-
ginate. In the large New Guinean species
{papua) the lobes of the 4th segments are
rather short on the hind but longer on the
middle tarsi. And in the smaller New
Guinean species the lobes of the 4th seg-
ments are long, more than Vl' the segments'
length even on the hind tarsi, and are rela-
tively longer in capitis than in icau.
In New Guinea, Aristolebia occurs chiefly
at mid-altitudes. It is probably arboreal
(in rain forest) and probably diurnal, al-
though a few individuals have been taken
in light traps at Wau.
Key to Species of Aristolebia of New Guinea
1. Larger, 9.5-11.0 mm (p. 84) papua
- Smaller, 5.5-6.5 mm 2
2. Entirely yellow or brownish yellow; sutural
angles distinct and usually subdenticulate (p.
84 ) tvau
- Elytra dark with broad stripes or spots pale;
sutural angles (narrowly) rounded (p. 85) ..
capitis
84 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Arisfolebia papua n. sp.
Description. With characters of genus;
form as in Figure 37; black, sides of pro-
notum broadly and of elytra narrowly pale,
elytra with variable pale marks or some-
times wholly dark, lower surface, mouth-
parts, and appendages reddish brown;
rather shining, microsculpture as described
below. Head 0.76 and 0.73 width prothorax;
front irregularly slightly impressed and
rugulose anteriorly, rather sparsely punc-
tulate, with c. isodiametric microreticulation
especially posteriorly. Prothorox: width/
length 1.67 and 1.60; base/apex c. 1.95
and 1.98 (figures approximate because an-
terior angles not defined ) ; margins narrow
anteriorly, broad posteriorly, each with seta-
bearing puncture at basal angle and before
middle; base and apex with entire impressed
marginal lines; disc slightly transversely
rugulose, sparsely punctulate, in part lightly
microreticulate. Elytra ample; width elytra/
prothorax 1.70 and 1.67; outer-apical and
sutural angles acute and denticulate; striae
entire, impressed, faintly or not punctulate;
intervals with slightly transverse microretic-
ulation and sparse fine punctulation, 3rd
with 2 dorsal punctures on outer edge c. %
from base and less than Vi from apex
(slightly variable in position). Leii,.s: 4th
segments middle and hind tarsi as in Figure
166; claws broadly triangular, each with c.
10 long teeth. Secondary sexual characters:
6 front tarsi scarcely dilated but with nar-
row, irregularly 2-seriate squamules; i mid-
dle tibiae 2-excised; i with 2, 9 c. 4 setae
each side before apex last ventral segment.
Measurements: length 9.5-11.0; width 4.5-
5.3 mm.
Types. Holotype i (Bishop Mus.) and
17 paratypes (some in M.C.Z., Type No.
31,396) from Wau, Morobe Dist., N-E.
N. G., 1100 to 1300 m, dates in Jan.. Feb.,
Apr., May, Aug., and Sept., 1961-1963
(Sedlaceks) (holotype, 1200-1300 m. May
7, 1963); and additional paratypes as fol-
lows. N-E. N. G.: 1, Swart Vy., Karubaka,
1500 m, Nov. 11, 195S (Gressitt). West
N. G.: 1, -'Humbolt Bay" (N. A. Doherty,
British Mus.).
Additional material. Papua: 1, W . Dis-
trict, Oriomo Govt. Station, Oct. 26-28,
1960 (Gressitt).
Measured specimens. The i holotype and
1 9 paratype from Wau.
Notes. This may prove to be a geographic
form oi Arisfolebia davaonis (Heller) of the
Philippines, but the color is different
( davaonis has the prothorax rusty red, not
black) and other details are probably dif-
ferent, although I cannot be sure about
them from Heller's description of his single
specimen. A form of davaonis, or a related
species, has been found also on Salajar Is.
off Celebes (specimens received from
Louwerens ) .
The single individual from Papua is the
only one in the New Guinean series with
whollv dark elvtra, and it differs slightly
from the type series in other ways. It may
prove to be a distinguishable geographic
form. Other variation in elytral pattern is
individual in the series from Wau.
Arisfolebia wau n. sp.
Description. With characters of genus;
form c. as in preceding species {))apua):
usually entirely reddish yellow, rarely with
faint dusky areas especially at base of
elytra; upper surface with light, irregular,
c. isodiametric or slightK- transverse micro-
reticulation. Head 0.80 and 0.77 width
prothorax. Prothorax: width length 1.56
and 1.67 (difference due parth to slight
abnormal extension of basal lobe in the first
individual); lateral margins wide, flattened
or weakly reflexed especially posteriorly,
each with usual 2 setae; base and apex
with (>ntire marginal lines, but apical line
weak at middle; disc irregularh' ± trans-
versely rugulose. Elytra ample, convex;
width elytra i)rothorax 1.72 and 1.76; outer-
apical angles obtuse but well defined and
sometimes subdenticulate, sutural angles
slightK dehiscent, ± subdenticulate; striae
entire, impressed, not distinctly punctulate;
The Carabid Beetles of New Guinea • Darlington
85
intervals convex, 3rd with 2 inconspicuous
punctures on outer edge before middle and
c. Vi from apex. Legs: 4th segments hind
tarsi deeply emarginate but lobes shorter
than usual in Lebia, 4th segments of middle
tarsi with longer lobes; claws each with c.
7 long teeth. Secondary sexual characters:
i front tarsi with slender squamae probably
in 2 series but often disarranged; S middle
tibiae with 2 excisions close together on
inner edge near apex; c^ with apparently 2,
9 3 setae each side near apex last ventral
segment. Measurements: length 5.5-6.5;
width 2.7-3.2 mm.
Types. Holotype i (Bishop Mus.) and
22 paratvpes (some in M.C.Z., Type No.
31,397) all from Wau, Morobe Dist., N-E.
N. G., 1100-1500 m, dates in Jan., Feb.,
Mar., Apr., May, June, Sept., Nov., Dec,
1961-1963 ( Sedlaceks ) .
Measured specimens. The S holotype
and 1 9 paratype.
Notes. I took a single 9 of this species
at Lockerbie, near the tip of Cape York,
in January 1958, thus extending the known
range of Aristolchia to Australia.
Aristolebia capitis n. sp.
Description. With characters of genus;
fonn and characters as in wau (above)
except slightly wider; elytra dark with
either large humeral and smaller subapical
marks reddish yellow or very broad reddish
yellow stripes running from humeri to apex.
Head 0.72 and 0.72 width prothorax. Pro-
thorax: width/length 1.64 and 1.59. Elytra:
width elytra/pro thorax 1.60 and 1.69; outer-
apical angles sharply defined but sutural
angles narrowly rounded. Legs with 4th
segments middle and hind tarsi strongly
lobed (Fig. 167); claws each with 5 long
teeth and sometimes a 6th (inner) tooth
that is difficult to see. Measurements:
length c. 6.0-6.5; width 3.1-3.3 mm.
Types. Holotype S (A.M.N.H.) and 1
9 paratype (M.C.Z., Type No. 31,398)
both from Mar Village, west Vogelkop,
West N. G., Nov.-Dec. 1944 (V. S.
Mallory ) .
Notes. The rounding of the sutural angles
and the small number of claw-teeth of
capitis are Le1)ia-\ike, but the form is that
of an Aristolchia ( some Lehia approach
this form too ) and the $ middle tibiae are
decisively 2-excised.
Genus LEBIA Latreille
Latreille 1802, Hist. Nat. Crustaceorum et In-
sectorum 3, p. 85.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1310 (see for additional references, synonymy,
subgenera, and list of species).
Jeannel 1942, Faune de France, Coleop. Carabiques,
Part 2, p. 1028.
Jedlicka 1963, Ent. Abhandlungen 28, p. 314.
Diagnosis. See Key to Genera of Lebiini
of New Guinea.
Description (characters of New Guinean
species only). Form broad but variable
(Figs. 38-41); coloration variable, upper
surface not pubescent. Head: eyes promi-
nent, genae short; 2 setae over each eye;
clypeus transverse, truncate or broadly
slightly emarginate, with 1 seta-bearing
puncture each side; labrum ± transverse,
sometimes slightly rounded anteriorly, 6-
setose; mentum strongly toothed; ligula
rather broad, 2-setose; paraglossae attached
to ligula, broad, setose. Prothorax ± lobed
at base, anterior angles broadly rounded
(so base/apex ratio not determinable);
lateral margins broad posteriorly, ± reflexed,
each with 2 setae, at or near basal angle and
before middle; disc with usual impressed
middle line, weak (or obsolete) anterior
and deeper posterior transverse impressions,
and weak transverse strigulation. Elytra
wide but variable in form; striae entire; 3rd
interval 2-punctate on outer edge. Inner
wings full. Lower surface with some short,
often inconspicuous pubescence. Legs: 4th
segments middle and hind tarsi very deeply
emarginate, with long lobes; 5th segments
with accessory setae; claws with 4-6 long
teeth. Secondary sexual characters: S an-
terior tarsi not or scarcely dilated, with
slender squamae in 2 series, often disar-
ranged; S middle tibiae with 1 small deep
86
BiiUetin Museum of Comparative Zoology, Vol. 137, No. 1
excision on inner edge just before apex; $
with apparently 1 or 2 (rarely 3), 9 2 or
more seta-bearing punctures each side near
apex last ventral segment (but these punc-
tures often difficult to identify amid other
punctures and pubescence ) .
Type species. Carahus haemorrhoidaJis
Fabricius {— Lehia maiiiinato (Fourcroy)),
of Europe.
Generic distribtition. Nearly world-wide
except absent in some cold regions and on
some remote islands. Species are numerous
in most tropical regions but are relatively
few (7) in New Guinea and still fewer in
Australia. Tliis suggests that the genus
has entered the Australian Region recently,
from the direction of tropical Asia.
Notes. The New Guinean representatives
of this huge, widely distributed genus are
probably all arboreal. Some or all of them
live in the lower foliage of rain forest. They
are probably diurnal, being rarely taken
in light traps.
Key to the Species of Lebia of New Guinea
1. Outer-apical angles of elytra well defined ._^_ 2
- Outer-apical angles of elytra rounded 3
2. Color piceous; form of Endynoiucna (Fig.
3cS) (p. 86) ciulynoiuena
- Color yellow; form more of typical Lcbia
(Fig. 39) (p. 86) externa
3. Elytra with conspicuous black "anchor"
mark on testaceous background, or dark with
anterior lunules and apex testaceous (p.
87 ) ____ karenia
- Elytra differently marked or not marked 4
4. Elytra dark with large conunon cordate
area testaceous (p. 87) cordifcr
- Not tlius marked . 5
5. Brown, elytra sometimes vaguely darker or
with vague discal cloud hut not sharply
bicolored, and head and pronotum not or
only lightly microrelieulate (p. 88) i>(i])iicUii
- Eillier hicolored or with head and pronotum
heavily microreticulate 6
6. Not sharply hicolored, brown, elytra often
with disc darker; head and pronotum heavily
microreticulate (p. 88) harda
- Bicolored, head and prothorax red-testace-
ous, elytra entirely black or piceous; liead
and pronotimi not or lightly microreticulate
(p. 89) _ iii.siihinni)
Lebio endynomena n. sp.
Description. With characters of genus;
form (Fig. 38) more of Endynomena than
of typical Lebia; piceous, reflexed margins
of prothorax and ( narrowly ) of elytra trans-
lucent testaceous, appendages reddish tes-
taceous; shining, reticulate microsculpture
absent or faint on front and pronotal disc,
distinct and strongly transverse on elytra.
Head 0.82 width prothorax; front weakly
impressed at middle and on each side an-
teriorly, irregularly rather sparsely punc-
tate. Prothorax subcordate; width/length
1.69; base apex not determinable; base and
apex margined. Elytra c. % wider than pro-
thorax, narrowed anteriorly; width elytra
prothorax 1.72; apices slightly obliquely
sinuate-truncate with outer angles well de-
fined and almost subdenticulate and sutural
angles irregularly narrowly rounded; striae
impressed, not distinctly punctulate. Sec-
ondary sexual characters as for genus, in-
cluding (^ middle tibiae with 1 deep
excision on inner edge just before apex; $
with 2 or 3 seta-bearing punctures before
apex each side last ventral segment (punc-
tures unsymmetric in the single specimen);
9 unkno\\n. Measurements: length c. 7.7;
width c. 3.9 mm.
Type. Holotype S (Bishop Mus.) from
Bubia, Markham Vy., N-E. N. G., 50 m,
Sept. 19, 1955 (Gressitt); the type is
unique.
Notes. This species differs in form and
appearance from any other Lchia kno\\n
to me, but the generic characters, including
the excision of the i middle tibiae, are
clearly those of Lebia.
Lebia externa n. sp.
Description. With characters of genus;
form as in Figure 39; reddish yelkn\', ap-
pendages slightl)' paler; rather shining,
reticulate microsculpture absent or faint
on Iront and pronotal disc, distinct and
transverse on ebtra. Head 0.92 and 0.92
width prothorax; Iront wt'akK impressed at
middle and on each side antcriorlx , slighth'
The Carabid Beetles of New Guinea • Darlington
87
irregularly punctate. ProtJwrax rather small,
not hemispheric but transversely subquad-
rate with anterior angles broadly rounded;
width/length 1.51 and 1.52; base margined,
apex not margined at middle; disc rather
strongly transversely strigulose and vaguely
punctulate. Elytra almost 2x wide as pro-
thorax; width elytra/pro thorax 1.98 and — ;
rather strongly narrowed anteriorly; apices
obliquely truncate-emarginate with outer
angles obtuse but distinct and sutural angles
narrowly rounded; striae impressed, not
punctulate. Secondary sexual characters:
S front tarsi with squamae ( if present ) not
easily distinguishable (worn off?); S mid-
dle tibiae with 1 deep excision on inner
edge just before apex; c^ with 2, 9 4 setae
each side near apex last ventral segment.
Measurements: length 7.0-7.3; width c.
3.2-3.4 mm.
Types. Holotype $ (Bishop Mus.) from
Pinciiu, Huon Pen., N-E. N. G., Apr. 20,
1963 (Sedlacek); 1 $ paratype (M.C.Z.,
Type No. 31,399) from Wau, Morobe Dist.,
Mt. Missim, 880-1050 m, Feb. 8-9, 1963
(Sedlacek); 1 9 paratype, Popondetta,
Papua, 60 m, Sept. 3-4, 1963 (Sedlacek).
Notes. Except for the distinct outer-
apical elytral angles, this species resembles
large individuals of Lebia papiicUa, de-
scribed below.
Lebia karenia Bates
Bates 1892, Ann. Mns. Civ. Genoa 32, p. 426.
Andrewes 1933, Ent. Series Indian Forest Records
18, Part 5, pi. 3, fig. 9.
Description (of New Guinean individ-
uals). With characters of genus; form c.
of typical Lebia; head, prothorax, and lower
surface usually reddish testaceous (head
and prothorax sometimes infuscate), elytra
varying from dark with posthumeral lunules
and apices testaceous (as figin-ed by An-
drewes) to testaceous with broad sutural
anchor mark; appendages reddish or tes-
taceous; microreticulation light and irregu-
lar on front, isodiametric or slightly trans-
verse on pronotum, more transverse on
elytra. Head 0.83 and 0.79 width prothorax.
Prothorax not hemispheric but transverse-
subquadrate with anterior angles broadly
rounded; width/length 1.54 and 1.61; base
margined, apex with marginal line weak or
interrupted at middle. Elytra less than 2x
width prothorax, narrowed anteriorly; width
elytra/prothorax 1.83 and 1.84; apices
obliquely sinuate-truncate with outer and
sutural angles narrowly rounded; striae
deep, impunctate. Measurements: length
6.0-7.5; width 2.8-3.8 mm.
Types. From Burma, in Genoa Mus.
(not seen).
Occurrence in Neio Guinea. Probably
throughout New Guinea at low altitudes
and in the lower mountains up to 1200 m
(at Guega W. of Swart Valley); 20 speci-
mens seen, from all 3 political divisions of
the island.
Measured specimens. A i from Dobodura,
Papua, and 9 from TorriceHi Mts., N-E.
N. G.
Notes. My identification of this species
is based on comparison with Andrewes'
material at the British Museum.
Lebia cordifer n. sp.
Description. With characters of genus
(but 5th segments missing from all tarsi);
form (Fig. 40) of typical rather narrow
Lebia, piceous above with clypeus and
labrum, side margins of prothorax, narrow
reflexed margins of elytra, and large com-
mon heart-shaped area on elytra (extend-
ing from inside humeri to apical Vi at 2nd
intervals and reaching 6th intervals lat-
erally) testaceous; lower surface and ap-
pendages brownish to testaceous; shining,
reticulate microsculpture absent or faint
on front and on disc of pronotum (but
these areas sparsely punctulate), trans-
verse on elytra. Head 0.89 width prothorax;
eyes large and very prominent; front with
V-shaped impression at middle and im-
pressed each side anteriorly. Prothorax
relatively small, not hemispheric; width/
length 1.58; sides strongly rounded, then
strongly sinuate just before c. acute but
blunted posterior angles; base with broad,
88
Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
strong, truncate lobe, weakly margined;
apex subtruncate, not margined at middle.
Elytra rather narrow but almost 2x width
of ( small ) prothorax, slightly narrowed
anteriorly; width elytra prothorax 1.95;
apices obliquely sinuate-truncate, with
outer angles broadly and sutural angles
narrowly rounded; striae impressed, not
distinctly punctulate. Secondary sexual char-
acters of i as described for genus; i with
2 setae each side before apex last ventral
segment; 9 unknown. Measurements: length
c. 5.7; width c. 2.7 mm.
Type. Holotype $ (Leiden Mus.) from
Bivak 39 A, Star Rge., West N. G., 1500 m,
July 12, 1959 (Neth. New Guinea Exp.);
the type is unique.
Notes. This is distinguished from other
New Guinean species in the preceding Key
to Species, but I do not know its real rela-
tionships.
Lebia papuella n. sp.
Description. Form of typical Lehia with
relatively small prothorax; entirely brownish
yellow, elytra sometimes with faint darker
cloud; shining, reticulate microsculpture c.
absent on front and on disc of pronotum,
present on elytral intervals as transverse
impressions not forming regular reticula-
tions. Head 0.92 and 0.88 width prothorax;
front scarcely impressed. Prothorax small,
transversely subcjuadrate; width length 1.54
and 1.56; basal and apical marginal lines
faint or interrupted at middle; disc with
anterior transverse impression subobsolete.
Elytra much wider than prothorax, nar-
rowed anteriorly; width elytra jMothorax
1.92 and 1.91; apices oblicpiely slightK-
sinuately truncate, with outer-apical angles
broadly and sutural angles more narrowly
rounded; striae impressed, not punctate.
Secondary sexual characters as for genus;
6 with apparently 2, $ 2 or more apical
ventral setae each side. Measurements:
length 4.1-5.5; width 2.0-2.7 mm.
Types. Holotype i (M.C.Z., Type No.
31,400) and 25 paratypes all from Dobo-
dura, Papua, Mar.-July 1944 (Darlington).
Additional material. Thirtv, from nu-
merous localities in all 3 political divisions
of New Guinea, from lowlands to 1700 m
(above Wau). Some of these specimens
are assigned to this species doubtfully.
Measured specimens. The i holotype and
1 9 paratype.
Notes. Lebia papuella seems closely re-
lated to a species from Queensland, Aus-
tralia, that I identify as picipennis Macleay,
but ])apuella has less pronotal microsculp-
ture and less sinuate elytra! apices than
picipennis. Similar (but not identical) un-
determined species occur in the Philippines.
Besides the type series, I have one ex-
ceptionally large 9 from Dobodura that
seems to be papuella. (Exceptional outsize
individuals occur in some other, American,
species of Lehia.) Its proportions and mea-
surements are: head 0.87 width prothorax;
prothoracic width/length 1.54; width ehtra
prothorax 1.94; length 6.5; width 3.3 mm.
My specimens (the types) were taken
by sweeping and beating undergrowth and
low foliage in rain forest.
Lehia barda n. sp.
Description. With characters of genus;
form of typical Lebia except prothorax
tending toward hemispheric; yellow, elytra
with ± distinct common dorsal plagia dark,
the dark area sometimes extending almost
to sides of elytra; lower surface and ap-
pendages yellow; whole upper surface rel-
atively dull, with deeply impressed iso-
diametric microsculpture becoming slightK
transverse on elytra. Head 0.81 and 0.81
width prothorax; front with 2 small im-
pressions anterior!). Prothorax: width
Icngtli 1.60 and 1.70; base margined; apex
not distinctly margined at middle. Elytra
narrowed anteriorb'; width elytra prothorax
1.80 and — ; apices weakly sinuate-truncate,
witli outer and sutural angles rounded;
striae deep, not distinctK' punctulate. Sec-
ondary sext(al characters as for genus; S
apparenth w itli 2, 9 3 setae each side last
ventral segment. Mcd.surcnwnls: length
4.4-5.8; width 2.2-2.8 mm.
The Carabid Beetles of New Guinea • Darlington
89
Types. Holotype i (M.C.Z., Type No.
31,401) and 2 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
additional paratypes as follows. Papua: 1,
between Laloki R. and Brown R., 25 m.
Mar. 16, 1956 (Gressitt); 1, Normanby Is.,
Wakainna, Sewa Bay, Nov. 21-30, 1956
(W. W. Brandt, Bishop Mns.). N-E. N. G.:
1, Busu R., E. of Lae, 100 m, Sept. 13, 1955
(Gressitt); 1, Wewak, 2-20 m, Oct. 11,
1957 (Gressitt). West N. G.: 1, Hollandia,
Apr. 1945 (Malkin, U.S.N.M.); 1, same
locality, 100 m, Ang. 24, 1955 (Gressitt).
Measured specimens. The S holotype and
1 9 paratype from Dobodura.
Notes. This species may be related to the
preceding ( papiiello ) but has the prothorax
more hemispheric, a more distinct elytral
cloud, and much heavier dorsal microsculp-
ture. It is somewhat similar also to Lehia
mehnota Chaudoir of Australia and Java
(but not New Guinea! ) but is much smaller,
with more hemispheric prothorax, and with
the dark dorsal elytral mark less defined.
Lebia insulorum n. sp.
Description. With characters of genus;
form (Fig. 41) of typical Lehia with rather
wide prothorax; bicolored, head and pro-
thorax red, elytra piceous; lower surface
red with sides of abdomen piceous; ap-
pendages reddish testaceous; shining, re-
ticulate microsculpture absent or faint on
front and pronotum, distinct and moderately
transverse on elytra. Head 0.80 and 0.79
width prothorax; front with trace of large
but indistinct (perhaps variable) V-shaped
impression. Prothorax transverse, not hemi-
spheric; width length 1.82 and 1.87; sides
broadly rounded, slightly sinuate before
slightly obtuse, blunted posterior angles;
base margined, apex weakly or not mar-
gined at middle. Ehjtra slightly narrowed
anteriorly; width elytra/ prothorax 1.72 and
1.70; apices obliquely weakly sinuate-
truncate with outer angles broadly and
sutural angles narrowly rounded; striae
deep, not distinctly punctulate. Second-
ary .sexual characters of S as described for
genus; i with 2 setae before apex each
side last ventral segment; 9 unknown.
Measurements: length c. 7.5; width c. 3.4
mm.
Types. Holotype S (Bishop Mus.) from
Nonnanby Is., Wakaiuna, Sewa Bay,
Papua, Jan. 1-8, 1957 (Gressitt); and 1 i
paratype (C.S.I.R.O., Ganberra, Australia)
from Rossel Is., SE. Papua, Oct. 1963 (W.
W. Brandt).
Notes. Although this distinct species is
placed in relation to others in the preceding
Key to Species, I do not know its real rela-
tionships.
Genus LACHNODERMA Macleay
Macleay 1873, Trans. Ent. Soc. New South Wales
2, p. 321.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1347 ( see for additional references and list
of species ) .
Jedlicka 1963, Ent. Abhandlungen 28, p. 302.
Diagnosis. Form (Fig. 42) diagnostic;
and see preceding Key to Genera of Lebiini
of New Guinea.
Description. None required here.
Type species. Lachnoderma cinctum
Macleay, of Australia.
Generic di.sfribution. SE. Asia including
India and Japan, and across the islands
to the Philippines, New Guinea, and
Australia.
Notes. I do not know how the different
species of this genus are related to each
other, and I do not know their habitats and
habits.
Lachnoderma foveolafum Sloane
Sloane 1915, Proc. Linnean Soc. New South Wales
40, p. 472.
Description. None required here: the
only species of the genus in New Guinea;
readily recognized by form (Fig. 42), color
(see Notes below), very coarse sculpture,
and pubescence; wings full; length (to apex
of elytra) c. 8 mm.
Type. From Gairns District, North
Queensland, Australia; in Sloane Goll.,
Canberra (seen).
90
Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Occurrence in New Guineo. Only in
Papua: 1, Yule Is. (Van Emden Coll.,
British Miis.); 1, Port Moresby, Sept. 24,
1955 (Gressitt), in light trap; 1, Kiunga,
Flv R., Aug. 1-3, 1957 (W. W. Brandt,
Bishop Mus.); 2, Laloki, 1909, 1910 (F.
Muir, H.S.P.A.); 2, Dogura, Oct. 20-Nov.
19, 1955 (E. L. Cassidy, Bishop Mus.); 1,
Goilala, Loloipa, Owen Stanley Rge., Jan.
1-15, 1958 (W. W. Brandt, Bishop Mus.).
Notes. Sloane's (Australian) type had the
prothorax red and elytra wholly blue-black.
Some Papuan specimens are similar but
others have the sides of the prothorax
blackish and the suture more or less red.
The variation is apparently individual.
Genus SINURUS Chaudoir
Chaudoir 1869, Ann. Soc. Ent. Belgium 12, p. 129.
Jedlicka 1963, Ent. Abhandlungen 28, pp. 298,
368 ( with key to the 3 known species ) .
Dia^nosi.s. Form (Fig. 43) diagnostic;
and see preceding Key to Genera of Lehiini
of New Guinea.
Description. None required here, but
note labrum long, emarginate, 6-setose;
mentum with short tooth; ligula very wide
(or fused with paraglossae), 4-setose; 4th
hind-tarsal segments small, simply emar-
ginate; claws with c. 4 teeth; <:5 front tarsi
with 3 segments each with 2 slender
s(iuamae at apex; S 9 both with 1 seta
each side last ventral segment.
Generic dislrihnt ion. SE. Asia (Burma,
etc.) across the islands to the Philippines
and New (guinea.
Type species. Si]uinis ojxiciis C^haudoir
( below ) .
Notes. "Sintirns?" ohsctiriis Sloane, from
Sattelberg, N-E. N.G., is transferred to
MochtJicrus (cj. v. ).
Sinurus somewhat resembles but is ap-
parently not related to Co])to^lossus of Aus-
tralia.
Sinurus opacus Chaudoir
Chaudoir 1869, Ann. .Soc. Ent. Iklgiuni 12, p. 130.
Jedlicka 1963, Ent. Al)handlungen 28, p. 368.
Louwerens 1964, Ent. Tidskiift 85, p. 188.
Description (selected characters only).
With characters of genus; form as in Figure
43; dull black; not setulose (except elytral
margins very finely setulose) but entire
upper surface heavily, finely, c. isodiamet-
rically microreticulate. Head 0.75 and 0.72
width prothorax. Protliorax variable in
shape and proportions; width length 1.27
and 1.43; base/apex 1.21 and 1.11; sides
slightly (variably) angulate near middle.
Elytra: width elytra prothorax 1.65 and
1.59; striae entire, well impressed, with
long, impressed scutellar striae. Measure-
ments ( New Guinean specimens ) : length
c. 10-11; width c. 4.4-5.2 mm.
Type. From Borneo; in Oberthiir Coll.,
Paris Mus. (not seen).
Occurrence in Neio Guinea. Papua: 1,
Popondetta, 25 m. May 1966 (Shanahan-
Lippert, Bishop Mus.), in light trap. N-E.
N. G.: 1, lower Busu R., Huon Pen., Mar.
28, 1955 (E. O. Wilson, M.C.Z.), in low-
land rain forest. West N. G.: 1, Araucaria
Camp, 800 m. Mar. 1939 (Toxopeus); 1,
Mt. Gyifrie, sea level-1000 ft. (-c. 300 m).
Apr. 1939 (Cheesman, S. Australian Mus.
(sic)); 3, Waigeu Is.. Camp 1, Mt. Nok.
2500 ft. (r. 760 m). May 1938 (Cheesman).
Measured specimens. Two { i 9 ) from
^^'aigeu.
Notes. The known range of opacus is
from Perak (Malay Pen.) and perhaps
Burma to the Philippines and New
(iuinea. The 7 New Guinean specimens
\'ar\ in shape and proportions of prothorax.
I cannot separate them satisfaetoriK- from
1 from Perak and 4 from the Philippines
that I have for comparison.
The few .specimens of this species that
1 ha\(> collect(xl (in the Philippines) were,
1 think, among fermenting leaves on the
ground in rain lorest.
Genus STENOTELUS Bouchard
Bouchard 1903, Ann. .Soc. Ent. France 72, p. 171.
Jedlicka 1963, Ent. .Ahliandlnnuen 28, p. 371.
Diuii^no.sis. See k'igure 44, and Key to
Genera of Lehiini of \cu- Guinea.
The Carabid Beetles of New Guinea • Darlington
91
Description. None required here, but
note labrum rather narrow, subtruncate,
not or at most faintly emarginate, 6-setose;
ligula, 4th hind-tarsal segment, claws, and
secondary sexual characters c. as described
for Sinurus (above).
Ty))e species. Stenotelus opactis Bouc-
hard.
Generic distribution. Malay Pen.,
Greater Siinda Islands, and Philippines
(opaciis); Celebes (piceiis Louwerens 1952,
Treubia 21, p. 217); and now New Guinea
(new species described below).
Notes. The species of this genus live on
tree trunks in rain forest and are probably
nocturnal.
Stenotelus spinosus n. sp.
Description. With characters of genus;
form as in Figure 44; black, appendages
dark; upper surface not pubescent, but
elytral margins very finely short-setulose;
rather shining, reticulate microsculpture c.
isodiametric on front, somewhat transverse
on disc of pronotum, more transverse on
elytra; lower surface with sparse, irregular,
short pubescence. Head 0.88 and 0.88
width prothorax; front \\'eakly impressed
each side anteriorly. Prothorax cordate
with sides angulate before middle and
strongly sinuate posteriorly (but sinuation
less than in opacus); width length 1.44 and
1.51; base/apex 1.07 and 1.07; apex mar-
gined, base not distinctly so; side margins
strongly reflexed, each with a seta at angu-
lation and at (blunted) basal angle; disc
with usual middle line and transverse im-
pressions, and faintly transversely strigu-
lose. Elytra: width elytra/prothorax 1.67
and 1.72; humeri rounded but prominent;
outer-apical angles spined, sutural angles
acutely toothed; striae entire, moderately
impressed ( but scutellar striae faint ) ; 3rd
intervals each with 2 conspicuous seta-
bearing punctures on inner edge slightly
behind middle and near apex. Inner winfi.s
full. Lei!,s slender; 4th hind-tarsal segment
long, slender, scarcely emarginate; 5th seg-
ment with short, weak accessory setae;
claws 4-toothcd, the innermost tooth small.
Secondary sexual characters: S front tarsi
scarcely dilated but with 3 segments 2-
seriately squamulose (squamae often dis-
arranged); 6 with 1, 9 2 setae each side
near apex last ventral segment. Measure-
ments: length 7.4-8.5; width 3.1-3.7 mm.
Types. Holotype S (M.C.Z., Type No.
31,402) and 13 paratypes from lower Busu
R., Huon Pen., N-E. N. G., May 4, 1955
(E. O. Wilson), in lowland rain forest; and
additional paratypes as follows. Papua: 5,
Dobodura, Mar.-July 1944 (Darlington);
1, Kiunga, Fly R., Aug. 24-27, 1957 (W. W.
Brandt, Bishop Mus. ); 4, Normanby Is.,
Wakaiuna Bay, Dec. 1-10, 1956, and Jan.
1-8, 1957 (W. W. Brandt, Bishop Mus.).
West N. G.: 1, Mt. Nomo, S. of Mt.
Bougainville, 700 ft. (c. 210 m), Feb. 1936
(Cheesman); 2, Waigeu Is., Camp 1, Mt.
Nok, 2500 ft. {c. 760 m), May 1938 (Chees-
man ) .
Measured specimens. The S holotype and
1 9 paratype from Dobodura.
Notes. S. spinosus is probably related to
S. piceus Louwerens of Celebes (see under
Generic distriI)ution, above) but piceus is
described as pubescent, with outer-apical
angles of elytra only strongly toothed, while
spinosus is not pubescent and has these
angles spined, although the length of the
spines varies.
The few specimens of this species that I
collected were taken on trunks of standing
and fallen trees in rain forest, mostly under
burlap bands put out to trap nocturnal
Carabidae.
Genus MISCELUS Klug
King 1834, Jalirbuchern Insectenkunde 1, p. 82.
Sloane 1907, Deutsche Ent. Zeitschrift for 1907,
p. 473.
1923, Trans. Ent. Soc. London for 1923,
p. 250.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae
7, p. 1359 (see for synonymy and additional
references ) .
Andrewes 1935, Fauna British India, Coleop.,
Carabidae 2, p. 3.
Jeanne! 1942, Faune de France, Coleop. Ca-
92 Bulletin Museum of Coivpomtive Zoology, Vol. 137. No. 1
rabiques, Part 2, p. 1017 (footnote: included
in "Pericalidae").
Jedlicka 1963, Ent. Alihandhingen 28, p. 398.
Diagnosis. Form (Fig. 45) characteristic
(note form of eyes and genae, and of
elytral apices); 1 or 2 setae over each eye
(see Notes below); clypeus emarginate;
labrum long, strong!)' rounded at apex, 6-
setose, emarginate in some species but not
in others; mentum toothed; ligula truncate,
with usually 4 setae at apex, and additional
setae in 2 irregular rows posteriorly; para-
glossae longer than ligula, rounded, with-
out setae; mesosternum wide between coxae;
metasternum with longitudinal row of small
tubercles each side of middle; wings full;
4th hind-tarsal segments small, oval, weakly
emarginate; 5th segments with weak ac-
cessory setae; claws not toothed; 6 front
tarsi scarcely dilated but each with 3 seg-
ments 2-seriately squamulose below; c^ with
small patch of dense pubescence on lower
edge of front femur near base; i with 1, $
2 setae each side last ventral segment, the
inner setae in 9 distant from margin.
Descri])tion. None required here.
Type species. Miscehis javanus King.
Generic distribution. SE. Asia (includ-
ing (leylon and India) to the Philippines,
New (iuinea, and part of Cape York,
Australia.
Notes. The taxonomic position of this
remarkable genus is doubtful, but will not
be debated here. Sloane (1907) suggested
a separate tribe for it, but one of the char-
acters he stressed (the presence of only
1 seta over each eye) is inconstant within
the genus (see below), and Sloane later
(1923) doubted it tribal separation was
valid. Andrewes (1935) did give it tribal
rank.
The variation in number of setae over
each eye in this genus is remarkable, it
has been noticed before, but has not been
ade(}uately described. Some ot the species,
including the type of the genus (javanus
Klug), have only 1 seta oxer each eye
(Fig. 169), while'others have 2 (Fig. 16cS).
Many species ol Carabidae belonging to
genera that normally have 2 pairs of setae
over the eyes are known to have lost the
anterior pair, but the posterior setae then
usually remain in their original position,
between or slighth' behind the posterior
corners of the eyes. But in the Miseelus
with a single seta over each eye, the seta is
between the positions of the 2 original ones,
and appears to correspond to the single
seta over each eye of the tribe Harpalini.
The New Guinean Miseelus with 1 and with
2 setae over each eye are apparenth' dif-
ferent species, but they are so similar that
some authors ( not noticing the setae ) have
failed to separate them or have treated them
as "varieties." Intermediates do not usually
occur: each individual has either 2 setae
over each eye or 1 seta in intermediate
position. The only exception 1 have found
is a 9 unicolor from Geelvink Bay (Paris
Mus. ) with 1 seta each side in intermediate
position and also, but only on the left side,
an additional seta posteriorly. Most com-
mon species of Miseelus have 1 seta over
each eye, but forms with 2 occur in Ceylon
and southeastern Asia as well as in New-
Guinea. I plan to consider this case in
more detail in Part I\' of the present work,
in discussion of variation of taxonomic
characters.
The variation of the labrum, entire or
emarginate in different members of this
genus, is noteworthy too.
The species of Miseelus that I ha\(' col-
lected in New Guinea and the Philippines
were on or under the bark of [vcv trunks or
logs in rain barest.
Key to Species ok Miscelus of Nkm- Guinea
1 . Ehtral iiitorvals 3, 5, 7 carinati' at liaso;
( 2 setae ()\ cr each eye; prothora.v more
(juadrate; len.uth 14.5 mm ) (p. 93) lit(tuit\us
- Ehtral interxals not carinale at l)a.se 2
2. Two setae o\er eaeli eye; lalirnm with ape.\
emariiiiiate; onler-apieal el>tral angles more
narrowly rounded (p. 93) .sihliit^
- One seta o\er eacli eye; lahruni iidt em;u-
jiinate; outer-apical angles oi eKtri more
l)roadl\' rounded - -^
3. Not spotted (p. 93) tDtirolor
- Elytra with sul)apical sutural red spot (p.
94 ) ( jdvatui.s)
The Carabid Beetles of New Guinea • Darlington 93
Miscelus luctuosus Putzeys
Putzeys 1875, Ann. Mus. Civ. Genoa 7, p. 725.
Andrewes 19.35, Fauna British India, Coleop.,
Carabidae 2, p. 3, footnote.
Description. A large Miscelus with rela-
tively .square prothorax and with elytral
intervals 3, 5, and 7 carinate at base; length
14.5 mm; other distinguishing eharaeters in-
cluding number of supraocular setae and
emargination of labrum not noted by
Putzeys, but Andrewes specifies 2 setae
over each eye in this species.
Type. From Andai, Papua, New Guinea
(Beccari and D'Albertis, Genoa Mus.) (not
seen ) .
Occurrence in Netc Guinea. Apparently
known only from the type.
Notes. I have seen no Miscelus with
carinate elytral intervals from New Guinea,
although carinate forms do occur elsewhere.
I think the species is probably distinct.
It should be easily recognizable.
Miscelus sibling n. sp.
Description. With characters of genus;
form (Fig. 45) as usual; black, not spotted.
Head 0.83 and 0.78 width prothorax; 2
setae over each eye; labrum emarginate at
apex. Prothorax subcordate; width/length
1.24 and 1.22; base/apex 0.94 and 0.95;
basal transverse impression very deep.
Elytra: width elytra/prothorax 1.33 and
1.28; outer-apical angles narrowly rounded;
intervals not carinate at base. Win'^s full.
Secondary sexual characters as for genus.
Measurements: length 12.0-14.5; width c.
4.1^.5 mm.
Types. Holotype S ( Bishop Mus. ) and
4 paratypes (2 in M.C.Z., Type No. 31,403)
from Wan, Morobe Dist., N-E. N. G., 1100-
1200 m, dates in Sept., Oct., 1961, 1962
(holotype, 1100 m, Oct. 13, 1961) (Sedla-
ceks); and additional paratypes as follows.
Papua: 2 ( $ $ ), Dobodura, Mar.-July
1944 (Darlington); 3, Goilala, Loloipa,
Owen Stanlev Rge., (1 specimen 975 m),
Nov. 16-25, ' 1957 and Jan. 16-30, 1958
(W. W. Brandt, Bishop Mus.). N-E. N. G.:
2, Sattelberg, Huon Gulf, 1899 (Biro); 1,
same locality (British Mus.); 1, Karimui,
1080 m, July 14-15, 1963, (Sedlacek); 1,
Okapa, Aug. 6, 1965 ( Hornabrook ) . West
N. G.: 1, Tami, May 11, 1903 (Paris Mus.).
Measured specimens. The 6 holotype and
1 9 paratype from Dobodura.
Notes. This and the following species
(unicolor) are sympatric, occurring at sev-
eral of the same localities, and both occur
also in New Britain.
Miscelus unicolor Putzeys
Putzeys 1875, Ann. Mus. Civ. Genoa 7, p. 725.
?stijgicus Putzeys 1875, Ann. Mus. Civ. Genoa 7,
p. 726.
Sloane 1907, Deutsche Ent. Zeitschrift for 1907,
p. 474.
?morioformis Macleay 1876, Proc. Linnean Soc.
New South Wales 1, p. 168.
Sloane 1907, Deutsche Ent. Zeitschrift for 1907,
p. 474.
Description. None required here. This
insect, whatever its proper name (see dis-
cussion below), is the common, smaller,
unspotted Miscelus of New Guinea, with
1 seta over each eye; labrum not emar-
ginate; outer-apical angles of elytra broadly
rounded; length (in New Guinea) 9.5-13.0
mm.
Types. Of unicolor, from Java, should
be in Brussels Mus.; of stygicus, from
Andai, Papua, now in Genoa Mus.; of
morioformis, from Hall Sound, Papua, pre-
sumably in Macleay Mus., Sydney (none
seen ) .
Occurrence in Neiv Guinea. Common
and widely distributed: 69 specimens,
from numerous localities in all three polit-
ical divisions of New Guinea; most at low
altitudes, but reaching 1200 m at Wan.
Notes. The application of the name
unicolor to this species in New Guinea is
conventional. Without revising the whole
genus, which I cannot do, I cannot decide
the relationships of the New Guinean pop-
ulation to populations farther west, nor
can I decide the relationship of the un-
spotted populations to spotted javanus.
This species {"morioformis" ) is recorded
94 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
from Coen, halh\'a\- up tlie Cape York-
peninsula, Australia, by Sloane (1907).
(M/sce/us javanus Klug)
King 1834, Jahrbiichern Insectenkunde 1, p. S2, pi.
1, fig. 9.
Csiki 19.32, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1359 (see for many additional references and
conventional synonymy ) .
Description. None required here. If the
typical form of this species occurs in New
Guinea, it is the only spotted Mi.scehis
there. Length c. (S.. 5-1 1.0 mm.
Types. From Java; now should be in
Berlin U. Zool. Mus. (not seen).
Occurrence in Neic Guinea. Doubtful:
New Guinea has sometimes been included
in the range of this species, but the synon-
ymy is confused and old published records
are doubtful, and I have seen no specimens
from the island.
Notes. The supposed unspotted form of
javanus, unicolor Putzeys, which may or
may not really be conspecific, does occur
in New Ciuinea and is treated above.
Genus HOLCODERUS Chaudoir
(Jliaudoir 1869, Ann. Soc. Ent. Belgium 12, p. 153.
Csiki 1932, Coleop. Cat., Caral)i(lae, Harpalinae 7,
p. 1360 (see for additional references and list
of species).
Jcdlicka 1963, Knt. Ahliandlnngen 28, p. 396.
Dia!J,iU)sis. Form (of New Guinean
species) as in Figur(> 46 (but form diverse
in species outside New Guinea); color
metallic; pronotum with 1 or more strong
setae at or near each anterior angle; elytral
apices unarmed but very strongly sinuate-
emargiiiat(>; length c. 8-9 mm.
Description (selected additional char-
acters only). Not pubescent above (sparsely
so below). Uead: labruiu moderately long,
subtiiiiicatc or slightly cmarginate, 6-setose;
inculuiii toodicd; labium 4-set()se, para-
glossac distinct, longer than labium, with-
out setae. Prolhorax: pronotum with mid-
dle line coarse. Elytra: 3rd intervals with
3 or inoic punctures, anterior punctme on
outer and middle and posterior |)uncturcs
on inner edge of intervals. Inner ir//)g.s full.
Legs: 4th hind-tarsal segments scarcely
longer than wide, shallowly emarginate;
5th segments with accessory setae; claws
with c. 4 weak teeth grouped near middle.
Secondary .sexual characters: i front tarsi
with 3 segments 2-seriately squamulose
(apical squamules of 3rd segment over-
lapping but not attached to 4th segment);
2 setae each side last ventral segment in
both sexes.
Type species. Holcodertis praemorsiis
C>haudoir, of Ceylon.
Generic distriJ)ution. SE. Asia (includ-
ing (a'vIoii and India) and across the
islands to the Philippines, New Guinea,
and northern Australia.
Notes. This genus is relatively diverse in
the western part of the Malay Archipelago.
A single species group extends eastward to
New Guhiea and Australia ( see Notes under
following species ) .
Holcoderus elongatus (Saunders)
Saunders 1863, Trans. Ent. Soc. London (3) 1,
p. 466, pi. 18, fig. 5a-h (Catascoput;) .
Wallace 1863, in Saunders paper cited above, p.
460 ( Cata.sc(>))iis).
Csiki 1932, Coleop. C^at., Carabidae, Harpalinae 7,
p. 1360 (see for additional references).
Andrewes 1946, I'roc. R. Ent. Soc. London (B) 15,
p. 87.
Description. None required here. See
Figure 46, characters stated under genus,
the following Notes, and Andrewes' (1946)
detailed rcdescription. Length c. 8-9 mm.
Type. From Dorey, West N. (i., col-
lected by Wallace; typ(^ in berlin U. Zool.
Mus. (not seen).
Occurrence in New Guinea. ProbabK
throughout New Guinea: 24 spcximcMis,
from all 3 political divisions ol the island;
most from low altitudes ( including Dobo-
dura), but 2 from Wan, 1150. 1200 m, and
1 from W'aigeu Is., 2500 ft. ( r. 760 m).
Notes. The \ariabilit\- of this species (if
it is all one species) is remarkable. The
h)rm is relatively constant, hut toloi \aries
from wholK- blue or green or copper\ to
bicolored with blue or green elytra and
The Carabid Beetles of New Guinea • Darlington 95
bright copper or violet prothorax. The
punctation of the pronotal disc varies: the
disc is always closely punctate in part, but
a variable area centered near or behind
the middle is usually less punctate. And
the lateral prothoracic setae vary in number
and position : at least 1 strong seta is always
present (unless broken off) at each pos-
terior angle, at the angulation of the pro-
thoracic margin near or just before the
middle on each side, and at each anterior
angle, but some individuals have additional
lateral setae of different sizes between the
anterior and median setae, and the occur-
rence of these extra setae is sometimes
strikingly unsymmetric.
This variation makes exact definition of
the species and comparison with other
species difficult. I think, however, that all
New Guinean specimens of the genus can
be referred to elongatus, that the latter is
probably confined to New Guinea and adja-
cent small islands, and that closely related
forms occur both in the western Malay
Archipelago (e.g., gracilis Oberthi.ir) and
in tropical northeastern Australia ( coerulei-
pennis Sloane).
I do not know the habits of Holcoderus
but I suspect that elongatus may inhabit
tree tops. Tliis would account for my
failure to find the species' natural habitat.
My single specimen from Dobodura was
taken at light, but this seems to be ex-
ceptional. No other specimens are labeled
as from light traps, and the bright color
suggests partly diurnal habits. However,
Wallace (1863) says that elongatus flies at
dusk.
Genus MINUTHODES Andrewes
Andrewes 1941, Ann. Mag. Nat. Hist. (11) 7,
p. 317.
Phitia Chaudoir 1869, Ann. Soc. Ent. Belgium 12,
p. 155 (not Platia Hiibner 1820, et al) .
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1361.
Andrewes 1939, Ann. Mag. Nat. Hist. (11) 3, p.
137.
Diagnosis. Usually immediately recog-
nizable by form (head very wide but eyes
smaller than usual in tribe, prothorax usu-
ally c. 2x wider than long, and elytra short
and subquadrate), small size (4-6.5 mm),
and other characters given in the Key to
Genera of Lehiini.
Description. Form as indicated above
and in Figures 47-58; upper surface espe-
cially of elytra often (not always) with
short pubescence, and elytra often (not
always) with color patterns of many pale
lines or pale blotches. Head wide but with
relatively small eyes; antennae rather short;
2 setae over each eye; front slightly im-
pressed each side anteriorly; clypeus sub-
truncate, with 1 seta each side; labrum
rather long, irregularly rounded or subtrun-
cate anteriorly, 6-setose; mentum toothed;
ligula with 2 principal setae and 1 or more
much smaller setae; paraglossae attached to
ligula, longer, broadly rounded, without
setae. ProtJiorax very wide, scarcely lobed
at base, very broadly emarginate anteriorly,
wdth wide, depressed or slightly reflexed
lateral margins; each margin with a seta at
basal angle and at or before middle of
length; disc with usual middle line, weak
anterior transverse impression, deeper sub-
basal transverse impression. Elytra very
wide and short; humeri prominent but
rounded; apices obliquely sinuate-truncate;
striae entire; 3rd interval with 3 dorsal
pvmctures at least in some species, but these
punctures often difficult to distinguish
amid other punctation and pubescence.
Inner wings full. Lower suiiace not or not
extensively pubescent. Legs rather slender;
tarsi sparsely setose above; 4th hind-tarsal
segment weakly emarginate; 5th segment
with accessory setae; claws each with c.
3 short, weak (vestigial?) teeth. Secondary
sexual characters: 6 front tarsi slightly
dilated, with numerous narrow squamae
not arranged in 2 series; 2 setae each side
near apex last ventral segment in both
sexes; and see under ^L sexualis for special
secondary sexual characters of this species.
Type species. Platia lineella Chaudoir,
96
Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
fixed by Andrewes 1939, p. 137. Andrewes
designated this species as the type of
Platia Chaudoir, and it is therefore also
the type of MinutJiodes, proposed as a new
name for preoccupied Platia.
Generic distrihuiion. Nine species on
New Guinea and neighboring small is-
lands, fewer on the Moluccas, Celebes,
New Britain, and northern Australia;
none known elsewhere.
Notes. This is a very distinct genus, con-
fined to a limited geographic area ( above ) .
The insects live wholly or chiefly on tree
trunks and fallen logs in rain forest. Al-
though they are winged, they do not often
fly to light, which suggests that they are
mainly diurnal.
The striking secondary sexual characters
of the 9 of se.xuali.s ( and of the related
hraclnjdera Chaudoir of the Moluccas) are
unique, so far as I know.
The Greek ending -odes does not indi-
cate gender, and Andrewes did not specify
gender when he proposed Minuthodes to
replace Platia. I therefore tentatively treat
the name as feminine, to make the gender
consistent with Platia.
Key k) Si'KciEs of Mi.wrnonKS ok New Guinea
1. Ehtra luarki'd witli iiuiiicrous loiiyitiuliiial
pak' liut's, soiiK'tiiiK s iiiucli intcrniptcd 2
- Elytra differently marked or not marked 5
2. Median-lateral protlioraeie setae before mid-
dle, c. '/•; of iirotlioraeic length from apex
(p. 96) . p(i))uaii(i
- Median-lateral setae near nn'ddle of pro-
thoraeie length 3
3. Elytra dnll; (length c. 6.5 mm) (p. 97)
rossi
- Elytra shining (under pubeseenee) 4
4. Smaller (c. 4.8 mm) (p. 97) _._- scdlucikoiuni
- Larger (c. 6.2 mm) (p. 97) .suhnitcn.s
5. Metallie bine blaek (p. 9,S ) tiwlallica
- Not metallie, black with or witlioni reddish
yellow spots 6
6. Elytra not plainly pubescent ( pubesci'iiee
actually present but very short, scarcely
\isil)le); 9 last \entral segment usually
with s(iuare <'.\cision at apex, and 9 hind
femur with flange or tooth near apex an-
teriorl> ; (shiin'ng black, unspottixl or 2-
or 4-spotted, Iml il spotted at least 1 pair
of s]iols elongate) scxualis
6a. Elytra not spotted, or each with a
single pale dash behind middle (Fig.
•55) (Papua) (p. 98) scxiudls s. s.
6b. Ehtra either each with a single based
dash, or 4-spotted with posterior spots
elongate ( Figs. .56, A ) ( central and
western New Guinea) (p. 99)
_- subsp. signata
- Elytra plainl\- pubescent; 9 not as de-
scribed 7
7. lilaek, not spotted; (Goodenough Is.) (p.
99 ) si\n])h'x
- Each eh'tron with 2 rather large red spots;
(mainland of New Guinea) 8
8. Smaller (4.0-5.3 nnn); elytral spots c.
regular in outline (Fig. 57) (p. 100) __ rcgularis
- Larger ( 5.5-5.8 mm ) ; elytral .spots irregu-
lar in outline (Fig. .58) (p. 100) __ irregularis
Minuthodes papuana (Sloane)
Sloane 1917, Proc. Linnean Soe. New South W'ales
42, p. 4,33 (Platia).
Agonochda lineella Sloane 1907, Deutsche Ent.
Zeitsehrift for 1907, p. 182 (not Platia lineella
Ghaudoir 1869).
Description. With characters of genus;
form as in Figure 48; black or brownish
black, appendages irregularly brown, elytra
with complex, \ariable pattern of pale lines
(Figs. 48, A); head and pronotum mod-
erately shining although closely punctate,
elytra roughened and duller, and upper sur-
face especialh' ehtra with short but distinct
pubescence. Head 0.78, 0.79, 0.81, and 0.80
width prothorax. Prothorax: width length
2.00, 2.04, 2.00, and 1.96; base apex 1.04.
1.06, 1.05, and 1.04; median-lateral setae c.
':i prothoracic length before apex. Eh/tra:
width elytra jirothorax 1.46, 1.47, 1.45, and
1.44; outer-apical angles moderateh and
sutinal angles more narrowh' rounded. Sec-
ondary sexual characters as lor genus; 9 last
ventral segment and hind lemora not modi-
fied, ^leasnrenients: li'iigth 4.4-5.2; width
2.2-2.6 mm.
Ty))e. From lierbcrlshohe, "New Pom-
crania" ( = New Britain); should be in
Deutsches Ent. Institut, Berlin (not seen).
Occurrence in Neic Guinea. Common
and widely distributed al low altitudes
throughout New (Guinea, and octnrring
also on Nornianb\ , (ioodenousj;!!. ;in(l Hos-
The Carabid Beetles of New Guinea • Darlington 97
sel Is.: 142 specimens seen in all; reaches
at least 1200 m at Wau.
Measured specimens. A S 9 from Dobo-
dura and c^ $ from Normanby Is., figures
listed in this order.
Notes. This species occurs on New
Britain as well as New Guinea, and it
apparently represents a group of species
(or subspecies?) that includes JinceUa
( Chaudoir ) of the Moluccas ( I have a
series from Morotai Is.) and qucenslandica
( Sloane ) of North Queensland, Australia ( I
have specimens from near Cairns and from
the Rocky Scrub, Cape York Pen.). The
different forms of this group are distin-
guished mainly by elytral color pattern:
linceUa has a relatively simple pattern of
3 pale lines on each elytron ( Fig. 49 ) ;
queenslandica, a complex pattern of short
lines, with 1 or 2 longer lines formed by
fusion of short ones (Fig. 50); and papuana,
a c. intennediate but very variable pattern
(Figs. 48, A). Some specimens from New
Guinea have elytral markings like those of
the type (from New Britain) as described
by Sloane.
The elytral pattern of papuana may be
genetically dimorphic at some localities
(cf. the dimorphism of markings described
for se.xualis), but the variation as a whole
is so complex that I have been unable to
analyze it satisfactorily.
Minuthodes rossi n. sp.
Description. With characters of genus;
form as in Figure 51; brownish piceous,
elytra with pattern of many short narrow
longitudinal pale lines in 3 transverse series;
head and prothorax moderately shining al-
though closely punctate, elytra roughened
and duller, and upper surface especially
elytra with short pubescence. Head 0.71
width prothorax, narrower than usual in
genus. Prothorax: width/length 1.79; base/
apex 1.21; sides irregularly broadly rounded,
almost subangulate at middle, slightly
sinuate before well defined but slightly
obtuse basal angles; median-lateral setae
near middle of prothoracic length. Elytra:
width elytra/prothorax 1.37; outer-apical
angles broadly rounded, apices subangulate
c. opposite ends 2nd intervals, sutural
angles narrowly rounded. Secondary sexual
characters of i as for genus; 9 unknown.
Measurements: length 6.5; width 3.2 mm.
Type. Holotype $ (California Acad.)
from Maffin Bay, West N. G., Sept. 1944
(E. S. Ross); the type is unique.
Notes. This seems to be a distinct species
although known from a single specimen
from a well collected lowland locality.
Minuthodes sedlacekorum n. sp.
Description. With characters of genus;
form as in Figure 52; irregular reddish
piceous with complex elytral pattern of
many short longitudinal pale lines in 3
irregular transverse series, appendages ir-
regular testaceous and brown; upper sur-
face including elytra shining although
pubescent and moderately closely punctate.
Head 0.74 width prothorax. Prothorax:
width/length 1.78; base apex 1.23; sides
broadly arcuate, slightly sinuate before well
defined posterior angles; median-lateral
setae near middle of prothoracic length.
Elytra: width elytra/prothorax 1.49; outer-
apical angles broadly rounded, sutural
angles narrowly rounded; striae coarsely
but irregularly punctate, intervals more
finely punctate. Secondary sexual char-
acters of S as described for genus; $ un-
known. Measurements: length 4.6-4.8;
width 2.3 mm.
Type. Holotype c^ (Bishop Mus.) from
Wau, Morobe Dist., N-E. N. G., 1050 m,
Sept. 16, 1961 (Sedlaceks); 1 £ paratvpe
(M.C.Z., Type No. 31,588), Pindiu, Huon
Pen., N-E. N. G., 870-1300 m, Apr. 21-22,
1963 ( Straatman ) .
Notes. More material may show that
this is a (distinct) geographic representa-
tive of the preceding species, rossi.
Minuthodes subnitens n. sp.
Description. With characters of genus;
black, elytra with pattern (Fig. 53) of
many short longitudinal pale lines in 3
98
Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
irregular transverse series, appendages red-
dish testaceous; rather shining although
whole upper surface rather closely punctate
and short-pubescent. Head 0.74 width pro-
thorax; as usual in genus except labrum
broadly emarginate at apex (an individual
rather than specific character?). Protliorax:
width length 1.84; base apex 1.25; base
more lobed than usual; sides broadly arcu-
ate, sinuate before c. right posterior angles,
with median-lateral setae near middle of
length. Elytra: width elytra prothorax
1.44; outer-apical angles broadly and sutural
angles more narrowly rounded; striae im-
pressed but not more coarsely punctate
than intervals. Secondary sexual characters
of 6 as for genus; 9 imknown. Measure-
ments: length 6.2; width 2.(S mm.
Type. Holotype $ (British Mus.) from
Mt.'Baduri, Japen Is., West N. G., 1000 ft.
(305 m), Aug. 1938 (Cheesman); the type
is unique.
Notes. This may (or may not) be a
(distinct) geographic representative of the
2 preceding species, ro.ssi and sedlacekorum.
Minufhodes metallica n. sp.
Description. With characters of genus;
lorm as in Figure 47; black, elytra with
strong blue-purple reflections, appendages
dark brown; shining but short-pubescent,
head and disc of pronotum sparsely punc-
tulate, elytra rather closely punctate as
well as punctulate. Head 0.79 width pro-
thorax. Frothorax: width length 1.98; base
ajiex 1.15; sides rather strongly rounded
anteriorly, nearly straight and conxerging
posteriorly until abruptly sinuate just be-
fore c. right posterior angles; median-lat-
eral setae c. V-s of prothoracic length from
apex. Elytra: width elytra prothorax 1.41;
outer-apical angles broadly and sutural
angles more narrowly rounded; striae
obsolete. Secondary sexual characters of
6 as lor genus; 9 unknown. Measure-
ments: IcngUi 5.0; width 2.6 mm.
Type. Holotype,' ( Briti.sh Mus.) from
Kokoda, Papua. 1300 ft. (c. 400 m). Sept.
1933 ( (>hecsinan j; 1 ,^ paratope ( S. Aus-
trahan Mus.), Mt. Lamington, Papua,
1300-1500 ft. (c. 400-460 m) (McNamara).
Notes. This is the only metallic Minu-
thodes know n from New Guinea.
Minufhodes sexualis n. sp.
Description. With characters of genus;
fonn as in Figure 55; black or brownish
black, appendages dark, elytra either with-
out markings or each with 1 pale dash on
5th interval behind middle; shining, pubes-
cence of most of upper surface absent or
so short as scarcely to be visible. Head
0.88 and 0.86 width prothorax; front
sparsely punctulate. Prothorax wide but
with relatively narrow base; width length
2.03 and 2.03; base apex 0.98 and 1.00;
median-lateral setae c. % of prothoracic
length from apex; disc sparsely punctulate.
Elytra: width elytra/prothorax 1.35 and
1.34; outer-apical angles broadly rounded,
sutural angles blunted or subdenticulate
(slightly variable); striae impressed and
punctate; intervals con\ex, without distinct
reticulate microsculpture, sparsely irregu-
larly punctate or punctulate. Secondary
sexual characters: S as for genus; 9 usu-
ally with last \entral segment with con-
spicuous c. square excision at apex, and
9 always with a short ridge or blunt tooth
on anterior edge hind femur near apex.
Measurements: length 4.5-5.6; width 2.0-
2.5 mm.
Types. Holotype 9 (M.C.Z., Type Xo.
31,404) and 1 9 paratype from Dobodma,
Papua, Mar.-July 1944 (Darlington); and
additional paratopes as lollows, all from
Papua: 7, Oro Ba\ near Dobodura, Dec.
1943-Jan. 1944 (Darlnigton); 2. Kokoda-
Pitoki, 450 m. Mar. 24, U)56 (C;ressitt); 1.
Mafuhi, 4000 ft. (1220 m), Dc>c. 1933
(C:heesman); 1, "Daradac iTn," 80 km N.
Port Moresby, 500 m, St«pt. 6, 1959 ( T. C.
Maa, Bishoi:) Mus.); 1, Koitakinunui, Apr.
1, 1918 (|. T. Ziminir, Chicago Mus.);
10, Mt. Lamington, 1300-1500 It. ( r. 400-
460 m) (McNamara. S. Australian Mus.).
Measured s))ecinicns. A 6 paratNj^e from
Kokoda-Pitoki and the ? hol()t\ii(\
The Carabid Beetles of New Guinea
Darlington
99
Notes. This species is evidently closely
related to M. bradujdcra Chaudoir of the
Moluccas (described from Batjan Is. and
represented by a series from Morotai Is.
in the M.C.Z.), but sexualis lacks the
metallic tone of the elytra of J)i(icliydcm,
and the ridge or tooth of the 9 femur, not
quite apical in scxiialis, is fully apical in
hrachijdera. These 2 forms, with the "sub-
species" described below, may eventually
be considered conspecific, but I prefer to
treat the New Guinean populations as a
separate species until their interrelationships
are better understood.
The material before me suggests that
sexualis may be dimorphic in two ways.
The pale dash on the elytron is either
present or absent but never partially de-
veloped in all specimens seen, and is some-
times present or absent in different in-
dividuals from single localities, for example
in those from Oro Bay. And, although most
females have a square excision on the last
ventral segment as described, 1 of 2 fe-
males from Dobodura has the last ventral
segment only acutely emarginate.
Minufhodes sexualis signata n. subsp.
Description. As typical sexualis (above)
except for markings ( Figs. 56, A ) : elytra
each with a broad posthumeral spot and
usually also a narrow stripe behind middle
(chiefly on 5th interval but bent inward
posteriorly) reddish or yellow^ (some in-
dividuals from Wau have only the post-
humeral stripe, as noted below). Head
0.89 and 0.90 width prothorax. Prothorax:
width/length 1.96 and 2.02; base/apex 1.03
and 1.02. Elytra: width elytra prothorax
1.41 and 1.42. Secondary sexual characters
as in typical sexualis. Measurements:
length 4.;3-5.8; width 2.0-2.9 mm.
Types. Holotype 9 (M.C.Z., Type No.
31,405) and 3 paratypes from Sambeang,
Mongi Watershed, Huon Pen., N-E. N. G.,
400 m, Apr. 21, 1955 (E. O. Wilson); and
additional paratypes as follows, all from
N-E. N. G.: 1, Butala, Mongi R., Huon
Pen., Apr. 22, 1955 (Wilson, M.C.Z.); 2,
lower Busu R., Huon Pen., Apr. 22 and May
12, 1955 (Wilson, M.C.Z.), in lowland rain
forest; 2, Finschhafen, Apr. 17 and May — ,
1944 (E. S. Ross, California Acad.); 1,
Wantoat, Finisterre Rge., 4000 ft. (1220
m), Sept. 9, 1957 (Munroe & Holland,
Canadian National Coll.); 1, Lae, 10 m,
July 5, 1962 (Sedlacek); 9, Wareo, Finsch-
hafen (L. Wagner, S. Australian Mus.);
16, Simbang, Huon Gulf, 1898 (Biro).
Additional material. N-E. N. G.: 11,
Wau, Morobe Dist., altitudes from 1050 to
1200 m, dates in Jan., Mar., Aug., Sept.,
Oct., 1961-1963 (Sedlaceks). West N. G.:
42, from localities scattered from Hollandia
to the Vogelkop.
Measured specimens. A c5 paratype from
Finschhafen and the 9 holotype.
Notes. Because this species varies geo-
graphically, I have restricted the type series
to specimens from a few localities in a
comparatively small area.
The elytral markings are essentially con-
stant, with only minor variation, in all
specimens except those from Wau, of which
only 4 have typical markings, while 7 have
markings reduced to a single posthumeral
dash on each elytron (Fig. 56 A). I have
seen no intermediates between these two
patterns. Inheritance of marking in this
case, as in typical sexualis, may be simply
Mendelian.
Minufhodes simplex n. sp.
Description. With characters of genus;
form as in Figure 54; black, not marked,
appendages brown; surface shining but
short-pubescent, head and prothorax punc-
tulate, elytra more closely punctate. Head
0.79 width prothorax. Prothorax: width/
length 2.0; base apex 1.15; sides irregularly
rounded anteriorly, nearly straight and con-
verging posteriorly, abruptly sinuate just
before c. right posterior angles; median-
lateral setae c. Vs of prothoracic length from
apex. Elytra: width elytra prothorax 1.45;
outer-apical angles broadly rounded, apices
bluntly subangulate opposite ends 2nd in-
tervals, sutural angles narrowly rounded;
100
BuUetin Museum of Comparative Zoology, Vol. 137, No. 1
striae impressed, not well defined, not
specially punctate. Secandanj .sexual char-
acters of 6 unknown; of 9 normal, with-
out special characters of sexualis. Measure-
ments: length 4.7; width 2.4 mm.
Type. Holotype $ (Manson Valentine
Coll.) from Goodenough Is., Papua, Oct.
14, 1943 (W. B. Jones); the type is unique.
Notes. I do not know whether this in-
sular species is represented on New Guinea
proper.
Minufhodes regularis n. sp.
Description. With characters of genus;
black or brownish black, appendages
brownish testaceous, elytra each with c.
regular posthumeral and subapical spots
reddish yellow (Fig. 57); rather shining al-
though surface pubescent and head and
pronotum irregularly punctulate or punc-
tate and elytra more closely punctate. Head
0.81 and 0.79 width prothorax. Prothorax:
width/length 1.96 and 1.96; base/apex 1.13
and 1.18; sides rounded anteriorly, c.
straight and converging posteriorly, briefly
but often abruptly sinuate before c. right
or slightly blunted posterior angles; median-
lateral setae c. Vy. of prothoracic length
from apex. Elytra: width elytra/prothorax
1.46 and 1.44; outer-apical angles broadly
and sutural angles narrowly rounded; striae
impressed but not sharj^ly limited and not
more coarsely punctate than intervals. Sec-
ondary sexual characters as for genus. Mea-
surements: length 4.0-5.3; width 2.1-2.8
mm.
Types. Holotype $ (M.C.Z., Type No.
31,406) and 3 jiaratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
additional paratypes as follows. Papua: 1,
Fly R. 5 miles l)elow Palmer R., May 23-
31, 1936 (Archbold Exp., A.M.N.H.). N-E.
N. {,.: 1, Saidor, Gabumi Village, Finis-
terre Rge., July 1-21, 1958 {\\ . \\ . Brandt,
Bishop Mus.);'2, Wan, Morobe Dist., 1150,
1200 in, Sept. 7, 1961, Sept. 26-27, 1964
(Sedlaceks); 1, Swart Vy., Karubaka, 1500
m, Sept. 20, 1958 (Gressitt), in light trap;
1, W'cwak, 2-20 m. Oct. 11, 1957 (Gressitt).
West N. G.: 1, vie. Hollandia, July-Sept.
1944 (Darlington); 1, same locality, 60 m,
Nov. 26, 1954 (L. D. Brongersma, Leiden
Mus.); 1, Maffin Bay, Aug. 1944 (E. S.
Ross, California Acad.); 1, Sibil, Star Rge.,
1260 m, Aug. 24, 1959 (Leiden Mus.); 1.
mountain slope above Bernhard Camp, 100
m, Apr. 1939 (Toxopeus).
Measured .specimens. The i holotype and
1 $ paratype from Dobodura.
Notes. M. rcfiularis is apparently widely
distributed in New Guinea at moderate
altitudes.
Minufhodes irregularis n. sp.
Description. With characters of genus;
black, elytra each with 2 (posthumeral
and subapical ) irregular reddish yellow
spots (Fig. 58), antennae and palpi red-
dish testaceous, legs much darker; rather
shining although surface short-pubescent,
head punctulate at middle and strigose at
sides, pronotal disc ± punctulate, elytra
more closely punctate and in part faintly
microreticulate. Head 0.75 and 0.77 width
prothorax, as described for gt^nus except
strigose at sides and with labrum distinctly
emarginate (both specimens). Prothorax:
width length 1.97 and 1.86; base/apex 1.12
and 1.11; sides broadly rounded, converging
posteriorly, briefly sinuate before c. right
posterior angles; median-lateral setae c. 'a
(or slightly more) of prothoracic length
from apex. Elytra: width elytra prothorax
1.34 and 1.42; outer-apical angles broadly
and sutural angles narrow I\ rounded; striae
impressed but not sharj-)!) limited and not
more coarseK' punctate than intervals. Sec-
ondary sexual characters oi S normal; 9
unknown. Mca.suroncnts: length 5.5-5.8;
w idth 2.8-2.9 mm.
Types. Holotype 6 (U.S.N. M.) and 1 S
ixuatype (M.C.Z., Type No. 31,407) both
from Hollandia, West N. (;.. May 1945
(B. Malkin).
Notes. This and th(> preceding .species
{reij,tdaris) are supcrlicialK sinu'lar, but
fh(> two are s\'miiatric and diHcr in sig-
The Carabid Beetles of New Guinea • Darlington 101
nificant details, and they may not be
closely related.
Genus CAJASCOPUS Kirby
Kirby 1825, Trans. Linncan Soc. London 14, p. 94.
Wallace 1863, in Saunders, Trans. Ent. Soc. Lon-
don (3) 1, pp. 460-461 (habits).
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1362 (see for additional references).
Andrewes 1937, Proc. R. Ent. Soc. London ( B )
6, pp. 187 ff. (key to species of India, etc.).
Jeannel 1949, Coleop. Carabiques de la Region
Malgache, Part 2, p. 1007 (in text).
Jedlicka 1963, Ent. Abhandlungen 28, p. 397.
Diapwsis. See Key to Genera of Lehiini
of New Guinea. In practice most Catascoptis
can be recognized by their medium to large
size (in the tribe), form (with prominent
eyes, etc.), and usually metallic coloration,
without geometric elytral markings.
Deseription (characters common to New
Guinean species of the genus, with ex-
ceptions noted). Form variable (Figs. 59-
64), slender and convex to broad and
depressed; color metallic (except in bnm-
neus), usually green, sometimes partly or
wholly blue or purple; size c. 8-22 mm;
upper surface not pubescent, more or less
shining (elytra sometimes dull), with micro-
sculpture present or absent, if present c.
isodiametric on head, somewhat transverse
on pronotum and elytra. Head with promi-
nent eyes; 2 setae over each eye; front
longitudinally impressed each side; clypeus
± emarginate, 1-setose each side; labrum
long, rounded at apex, emarginate at apex,
6-setose; antennae with 4 segments glabrous
except for tactile setae and a little pubes-
cence at apex 4th segment; mentum toothed;
ligula 4-setose, paraglossae much longer,
not setose; palpi slender. Prothorax quadrate
or subcordate; base not lobed; lateral
margins variable, each with 1 seta at base
and 1 or more near or before middle; base
with entire margin (except in dobodiira),
apex at middle not margined or weakly so;
disc with impressed middle line, deep pos-
terior transverse impression, and usually
weak (but variable) anterior transverse
impression. Elytra with humeri prominent
but rounded (humeral margins slightly
thickened in laevigatiis); apices variable, as
described for separate species, often toothed
or spined; striae entire, punctation vari-
able; 7th intervals usually and 5th some-
times raised or carinate at base; .3rd inter-
vals usually .3-punctate (2-punctate in
Jatiis), with punctures often near middle of
intervals (not on edges) but position vari-
able. Inner icings full. Lower surface with
some inconspicuous, short, sparse pubes-
cence (much more pubescence along mid-
line in tcaUacei group); last ventral seg-
ment usually slightly, broadly (variably)
emarginate in both sexes. Legs slender;
4th hind-tarsal segments small, weakly
emarginate; 5th segments with accessory
setae; claws not toothed. Secondary sexual
characters: i front tarsi slightly ( scarcely )
dilated, with 3 segments 2-seriately squamu-
lose below; i with 1, $ 2 or 3 setae each
side last ventral segment ( except i as well
as 9 with 2 or 3 setae each side in strigicol-
lis).
Type species. C. hardwickei Kirby, of
India.
Generic distribution. Represented in 3
separate tropical areas: numerous in trop-
ical Asia and the Malay Archipelago ( and
a few in tropical Australia); fewer in
tropical Africa (absent in Madagascar);
and probably represented also in tropical
South and Central America (but Jeannel
doubts whether the American species
should be included in the genus).
Notes. Although Catascopus occurs also
in Africa and probably in tropical America,
its headquarters are in tropical Asia and
the Malay Archipelago. The greatest num-
bers of species are on the Malay Pen. and
the western part of the Archipelago, but
the genus is well represented east to New
Guinea, where 14 species are now known.
Of these 14 species, elegans and smaragdulus
range from the mainland of Asia across the
islands to northern Australia; facialis, from
Asia to western New Guinea but not Aus-
tralia; and laevigatus is common to the
102 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Moluccas and New Guinea. Most other
New Guinean species of the genus are
endemic, and one group of striking species
(the icaUacei group) has probably evolved
on New Guinea and is now represented
there by at least 5 species. Only 5
Catascoptis (2 of them endemic) occur
in Australia, and they seem to represent 5
separate invasions from New Guinea. So,
tlie distribution of the genus suggests
multiple dispersal eastward across the
Malay Archipelago, with considerable spe-
ciation and some secondary radiation on
New Guinea, and minor invasions of north-
ern Australia.
All the Catascopus that I know live on
tree trunks and fallen logs in rain forest.
They are all winged, and very active. Con-
cerning their habits, Wallace (1863) says,
"The species of the yeniis Catascopus are senii-
iiocturnal in their hal)its, never flying except
at ni^ht. The species taken at Dorev (viz.,
WaUacei, W. W. S.; elongatus, W. ' W. S.
[— Ilolcodcrus]; Anwnsis, W. W. S.; amncmis,
Clniud. [= clcfi.ans]) flew against me at dusk.
The greater part of the species and individuals
I have taken have, however, been captured
under the decaying bark of fallen trees.
"As soon as the bark of a tree splits and
cracks so as to separate it from the wood,
the Cata-icnpi frecjuent it, I)ut I could scarcely
ever capture them in that position, owing to their
great activity and the force required to tear
off the bark. After a tree has lain about a year
the bark Ix'coines rotten and can be easily
broken off, and tlicii, 1)\ thi' assistance of a
net, the insects which lurk beneath it can be
more easily caplincd. 'I'lu> larger species found
in Malacca, Borneo and Singapore used fre-
(juently to be seen coursing along the surface
of some immense fallen trees, from one crack
to another, their brilliant lioihes glittering
splendidly in the sunlight.
"To capture them was by no means easy, as
they would get under the trunk where it touched
the ground, if closely pursued and no friendly
crevice was at hantl. Many an hour ha\e 1
pleasantly spent in hunting them in tiie dcMise
swampy forests of Borneo, hi Malacca and
Singapore the spice of fear and danger would be
added to the interest of the .sport, owing to the
jirobable vicinity of tigers, who might at any
moment be watching us as eagerly and with as
di-atlly a puipose as we were watching the poor
Catascopi.
"However closely pursued 1 ha\e never seen
one of these insects fly in the day time, neither
do they come out at all into the light, except
to \'isit some part of the trunk they reside in,
to which the subcortical passages do not ex-
tend. . . . The species and indi^■iduals of this
genus are much more abundant in Malacca and
Borneo than in the equally luxuriant forests of
the Molucas and New Guinea."
Key to Species of Catascopus of New Guinea
1. Elytral apices without acute teeth or spines
at or near sutural angles 2
- Elytral apices acutely toothed or spined at
or near sutural angles 4
2. Outer elytral angles rounded or \ery
obtusely angulate 3
- Outer elytral angles right or (if obtuse)
very well defined, sometimes denticulate;
length c. 10.5-13.5 mm (p. 103) ._ facialis
3. Color metallic green or blue; length c. 8.5-
10 mm (p. 103) elegans
- Color brown or bronze; length c. 12-13
mm ( see also Notes under this species )
(p. 104) Inuniieus
4. Prothorax with 2 or more lateral setae
near or before middle on each side ( if
setae broken off, positions shown by punc-
tures ) ; Unm relatixely broad and de-
pressed 5
- Prothorax with only 1 median-lateral seta
each side; form variable but often more
slender and convex 7
5. Two or 3 setae near or before middle each
side; length 17.5 mm ( see also Descrip-
tion) (p. 104) latus I
More (often 6) such setae each side ,. (•>
6. Elytral striae lightb' impressed; ehtral
margins wider than usu;il near middle;
length r. 10-11 mm (p. 104) _ laeiifJiattis
- Elytral striae deepiT; ehtnil margins less
wide; length r. 12-13 mm (.see also
l)('scri))tii»i ) (p. 105) siiliis
7. Outer ehtral angles blunt or angulate
hut not si^ined; relati\ely small species,
usu;d!> untler 1 1 mm 8
- Outer elytral angles spined; larger species,
13-21 mm {icaUacci group) 10
8. Outer el\ tr;il ;uigles rounded or obtuse;
smaller, 'r. 7.5-8.0 nun (p. 105) _. .
_ stnaragduhis
- Outer elytral angles u.sually c. right or
acute, or if obtuse, size larger 9
9. I'rothoraeic margins narrow ( almost as in
dedans); basal marginal line of pronotum
c. obsolet(>; reticulate microsculpture oii-
.solcle on disc of el\tra; length r. 9-10
nnii (p. 106) . - clohoduro
- I'rothoracic margins slightly wider; b;isal
nuirginal line ol prothorax impres.sed;
The Carabid Beetles of New Guinea • Darlington 103
reticulate microsculptuie distinct on disc
of elytra; length 8.7-9.3 mm (p. 106) . hiroi
10. Prothoracic margins moderate 11
- Prothoracic margins wider (see Descrip-
tions ) 13
1 1 . Fifth elytral inter\ als not or not much
raised near base; length c. 13-15.5 mm
(p. 107) aruensis
- Fifth as well as 7th elytral inter\'als
raised near base; usually larger 12
12. Prothorax more rjuadrate with blunter
posterior angles; head and prothorax ±
green, elytra blue-purple (note head colored
as prothorax); length c. 15-18 mm (p.
108 ) strigicollis
- Prothorax more cordate, with more acute
posterior angles; prothorax green or
cupreous, head and elytra blue-purple
(note head colored as elytra); length c.
15-20 mm ( p. 108 ) wallacei
13. Prothorax narrower (width/length c. 1.40);
head less depressed posteriorly; head dark,
pronotum green cupreous, elytra blue-
purple or blue-green; length c. 17-22
mm (p. 109) taylori
- Prothorax wider (width/length c. 1.70);
head more depressed posteriorly; head as
well as prothorax green, elytra blue-green;
length c. 20 mm (p. 109) rex
Cafascopus facialis (Wiedemann)
Wiedemann 1819, Zoologisches Magazin 1, 3, p.
165 (Corahus).
Csiki 1932, Coleop. Cat., Carabidae, Haipalinae 7,
p. 1364 (see for synonymy, "varieties," and
many additional references not concerned with
New Guinea ) .
Jedlicka 1963, Ent. Abhandlungen 28, pp. 382, 395
( "fascialis" ) .
Description ( for recognition only ) . With
characters of genus; form rather compact;
green or blue and green; elytra with outer-
apical angles well defined, apices sometimes
subangulate (variable) near suture, striae
deeply impressed and strongly punctate,
and 5th and 7th intervals raised; length (in
New Guinea) c. 10.5-13.5 mm.
Tijpe(s). From "Bengalia," in Copen-
hagen Univ. Mus. (not seen).
Occurrence in New Guinea. West N. G. :
1, Maffin Bay, Aug. 1944 (Darlington); 1,
"Dorey" (Paris Mus.). Also 1 specimen
labeled only "N. guin" ( British Mus. ) .
Notes. If my identifications are correct,
this species ranges from SE. Asia to the
Philippines, Moluccas, and (western)
New Guinea but does not reach Australia.
It is variable, and its full synonymy and
subspecies (if any) remain to be worked
out. It is rare in New Guinea and may be
confined to the western part of the island
(perhaps it has recently arrived from the
west). I found it common on Morotai Is.
in the Moluccas.
Cafascopus elegans (Weber)
Weber 1801, Observations Entomologicae, p. 45
(Elaphnis).
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1363 (see for additional references and ex-
tensive synonymy ) .
Andrewes 1937, Proc. Ent. Soc. London for 1937
(B) 6, p. 189.
Van Emden 1937, Stettiner Ent. Zeitschrift 98,
p. 35 (as subsp. australasiac Hope).
Jedlicka 1963, Ent. Abhandlungen 28, pp. 380,
385.
amoenus Chaudoir 1861, Berliner Ent. Zeitschrift
5, p. 120.
obliquatus Fairmaire 1881, Le Naturaliste 3, p.
381 ( new synonymy ) .
Description ( for recognition only ) . With
characters of genus; form convex; green or
partly coppery; prothoracic margins nar-
row; elytral apices unarmed; in general
without striking characters; length c. 8.5-
10 mm.
Types. Of elegans, from Sumatra (col-
lected by Doldorf ) , present location of type
unknown; of amoenus, from Dorey, West
N. G., now in Oberthiir Coll., Paris Mus.;
of obliquatus, from New Britain, presum-
ably now in Paris Mus. (none seen).
Occurrence in New Guinea. Very com-
mon (about 200 specimens) throughout
New Guinea, chiefly at low altitudes (in-
cluding Dobodura), but reaching 1700 m
near Wan.
Notes. The range of elegans, including
its supposed subspecies and varieties
(which need further study), is from SE.
Asia to Australia, east at least to the
Philippines and Solomons.
The name obliqiuitus Fairmaire has been
104 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
overlooked by most authors, and the cita-
tion in Csiki is incorrect. The description
clearly is based on a small specimen of
the present species, which is common in
New Britain.
Cafascopus brunneus n. sp.
Description. With characters of genus;
form as in Figure 59, compact and convex
(in genus); brown, subaeneous, append-
ages brown; rather shining, reticulate micro-
sculpture faint on head, light on pronotum
and elytra. Head large, 0.97 and 0.97 width
prothorax; front irregularly sculptured and
in part sparsely punctulatc. ProtJiorax
square-cordate; width/length 1.38 and 1.37;
base/apex 1.01 and 0.96; margins moderate;
disc lightly transversely strigulose and
punctulatc. Elytra: width elytra/prothorax
c. 1.53 and 1.59 (but elytra warped so mea-
surements inexact); humeri very prominent,
almost subangulate ( narrowly rounded ) an-
teriorly; apices oblique, scarcely sinuate,
with outer angles scarcely indicated (very
l)roadly rounded) and sutural angles nar-
rowly rounded and sometimes minutely
denticulate; striae well impressed, faintly
punctulatc; no intei'vals specially elevated
at liase. Measurements: length c. 12-13;
width c. 4.4-5.0 mm.
Types. Holotype $ ( Bishop Mus. ) and
1 9 paratype from Goilala, Tapini, Owen
Stanley Rge., Papua, 975 m, Nov. 16-25,
1957; and 2 additional paratypes (M.C.Z.,
Type No. 31,408) from Goilala, Loloipa,
Owen Stanley Rge., Jan. 16-30, Feb. 1-15.
1958 ( all these sp(>cimens, W. W. Brandt ) ;
1 paratype (S. Australian Mus.), W'areo,
Finsclihafcn, N-E. N. G. (L. Wagner).
Measured specimens. The 6 holotype and
1 9 paratype from Loloipa.
Notes. This species is unusual in its
rather compact form, j)]ain brown-aeneous
color, and simple elytral apices (excejit lor
minute, variable denticles near sutural
angles). So far as I know, il is not closely
related to any previously described species.
Characters distinguishing it from other
species are given in the preceding Key.
Cafascopus lafus n. sp.
Description. With characters of genus;
form as in Figure 60, very broad, depressed;
head and pronotum dark green, elytra
purple, lower surface and appendages red-
dish black; head and pronotum shining with
reticulate microsculpture absent or faint,
elytra dull and closely microreticulate.
Head 0.87 width prothorax; front flat,
broadly irregularly impressed. Prothorax
wide-subcordate; width length 1.77; base/
apex 0.97; side margins rather narrow (in
relation to width of prothorax ) , moderately
reflexed, left with 3, right with 2 formerly-
seta-bearing punctures at and before mid-
dle. Elytra: width elytra prothorax 1.36;
humeri broad but margin not thickened
and not subangulate; margins rather nar-
row; outer-apical angles prominent, slightly
acute; apices with moderate spines c. op-
posite ends sutural striae; striae moderateh'
impressed, scarcely punctulatc; intervals
not elevated at base, punctulatc especially
along middle, 3rd with onl\' 2 dorsal punc-
tures, less than ^4 from base and near or
behind middle (position unsymmetric).
Measurements: length 17.5; width 6.3 mm.
Type. Holotype 9 (British Mus.) from
W. Tami R., Pukusan-Humboldt Bav Dist.,
West N. G., June 1937 {\\ . Stiiber); the
type is unique.
Notes. This striking and thoroughh' dis-
tinct species is snfficientK' compared with
others in the Key to Species of Catascopus
of New Guinea.
Catascopus laevigatus Saunders
SauiKlns lcS63, Trans. Kiit. Sov. Loiulon (3) 1,
p. 458, pi. 18, fi,t,^ 2a-l).
Csiki 1932, Colcop. C"at., Caiabidae, Harpalinac 7,
p. 1365 (sor for additional rcfrrcnces ).
Description ( for recognition only). With
charact(M-s of genus; wide and d(^pr(\ssed;
green, shining; el\lra with sutural angles
spined, outer-apical ehtral angles c. right;
length c. 10-1 1 mm.
Ty))es. From ''Batch ian. Ternate and
Am. Wallace," type now in ObcM'thiir
C^oll. I'aiis Mus. (not seen).
The Carabid Beetles of New Guinea • Darlington
105
Occurrence in Neiv Guinea. Twenty-four,
from numerous localities in all 3 political
divisions of New Guinea; occurs at Do-
bodura and up to 1200 m at Wau.
Notes. I have seen specimens also from
the Am Is. and from Biiru, Cerain, and
Halniahera (Jilolo) in the Moluccas. The
closely related C. laticoUis Macleay of North
Queensland (Kuranda and Atherton Table-
land, and Coen-Rocky Scrub areas) repre-
sents the species in Australia.
Caiascopus sidus n. sp.
Description. With characters of genus;
form as in Figure 61; rather wide but less
depressed than laevig,atus; green, elytra
blue purple with green humeri ( at Wau ) or
c. wholly green (Star Rge.) or c. wholly
purple (Japen Is.), lower surface and ap-
pendages dark brown; shining, reticulate
microsculpture faint on front and on disc
of pronotum, distinct on elytra. Head 0.93,
0.90, 0.92 width prothorax; front irregularly
impressed at middle, sparsely minutely
(scarcely detectably) punctulate. Prothorax
transverse-cordate with wide base; width/
length 1.57, 1.61, 1.47; base/apex 1.14, 1.13,
1.13; side margins broader and more re-
flexed than in lacvi^atus, each with c. 6
strong setae (or punctures) in anterior %;
disc almost without transverse strigulation,
faintly and sparsely (hardly detectably)
punctulate. Elytra: width elytra/prothorax
c. 1.49, 1.54, 1.57; humeri prominent but
with margins rounded (not widened and
subangulate as in laevigatiis); outer-apical
angles well defined, c. right or nearly so;
apices with short spines not quite at sutural
angles; striae well impressed, scarcely punc-
tate ( more impressed but less punctate than
in laevigatas ) ; intervals scarcely elevated at
base. Secondarij sexual characters as de-
scribed for genus. Measurements: length c.
12-13; width 4.6-5.1 mm.
Types. Holotype $ (Bishop Mus.) from
Wau, Morobe Dist., N-E. N. G., 1200 m,
Sept. 15-30, 1962 (Sedlacek); 1 i paratype,
same locality, 1250 m, Sept. 16, 1962 ( Sed-
laceks); 1 paratype, Mt. Missim (near Wau),
1600 m, Mar. 17, 1966; 7 paratypes, Wau
Ck., 1200-1500 m, Sept. 16-18, 1964 (M.
Sedlacek) (some paratypes in M.C.Z., Type
No. 31,409).
Additional material. West N. G. : 1 ^ ,
Sibil, Star Rge., 1260 m. May 16, 1959
(Leiden Mus.), at light; 1 S, Mt. Baduri,
Japen Is., 1000 ft. (305 m), Aug. 1938
( Cheesman ) .
Measured specimens. The i holotype and
the c^ S from Star Rge. and Japen Is., in
this order.
Notes. Although similar to laevigatas,
sidus is more convex, with wider and more
reflexed prothoracic margins, and other dif-
ferential characters noted in the preceding
description. The single specimens from
Star Rge. and Japen Is. differ from the
types in color of elytra (see Description,
above) but I do not wish to call them
subspecies without seeing more material.
Caiascopus smaragdulus Dejean
Dejean 1825, Species General Coleop. 1, p. 331.
Csiki 1932, Coleop. Cat., Carabidae, Haq:)alinae 7,
p. 1366 (see for additional references and
synonymy, which do not concern New Guinea).
Description (for recognition only). With
characters of genus; small, rather broad;
green or partly coppery; prothorax with
margins wider than in clegans and set off
by submarginal longitudinal swellings;
elytra with outer-apical angles rounded or
bluntly obtuse, apices each with an acute
tooth or short spine; intervals not elevated
at base; length 8 mm or less.
Type(s). From Java; now in Oberthiir
Coll., Paris Mus. (not seen).
Occurrence in New Guinea. Seventeen
specimens from localities in all 3 political
divisions of New Guinea and from Nor-
manby Is.; at low altitudes only.
Notes. C. smaragdulus ranges from the
southern corner of Asia (Burma, etc.) to
New Guinea, New Britain, and the north-
eastern corner of Australia (specimens
from the Rocky Scrub, mid-peninsular
Cape York, taken by me in June 1932).
In this species the width of the prothorax
106 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
and the development of elytral spines vary
both individually and, I think, geographi-
cally, but I do not have enough material
from outside New Guinea to define satis-
factory subspecies.
Catascopus dobodura n. sp.
Description. With characters of genus;
form ( Fig. 62 ) c. average with elytra rather
wide but convex; green, elytra sometimes
greenish castaneous or purplish laterally,
lower surface almost black, appendages
dark brown; shining, reticulate microsculp-
ture absent or nearly so on front, pronotum,
and disc of elytra, indicated toward sides
and apex of eUtra. Head 1.01 and 1.01
\\'idth prothorax; front slightly depressed
anteriorly and longitudinally impressed each
side. Prothorax quadrate-subcordate; width
length 1.44 and 1.42; base/apex 1.08 and
1.04; sides broadly rounded anteriorly with
anterior angles only a little advanced,
strongly sinuate c. % of length before right
or slightly acute basal angles; side margins
very narrow, not set off by longitudinal
swc^llings, each with usual seta at basal
angle and 1 median-lateral seta just before
middle; basal transverse impression deep
(as usual) but basal marginal line obsolete
at middle; disc with faint transverse strigae
and faint sparse punctulation. Elytra :
width elytra/prothorax — and 1.59 (elytra
of S too spread to measure); lateral mar-
gins moderate; outer-apical angles c. right
or slightly obtuse but distinct, apices each
will) spine c. opposite end 2nd interval;
striae moderately impressed, faintly punc-
tulale; intervals no! elevated at base. Mea-
surements: length e. 9-10; width c. 3.4-
3.8 mm.
Ti/i)es. llolotype i (M.C.Z., Type No.
31,410) from Oro Bay, Papua, Dec. 1943-
jaii. 1944 ( Darlington ); and 1 9 paratyjK'
Iroin Dobodura (near Oro Bay), Mar.-Julv
1944 (Darlington).
Other material. One 9 , Kiunga, Fly R.,
July 2.3-25, 1957 (\V. W. Brandt. Bishop
Mus.).
Measured specimens. The types.
Notes. It is surprising to find a new,
medium-small, green Catascopus at low
altitudes in Papua, but the species seems
clearly distinct. In form ( except that it is
a little broader) and narrow prothoracic
margins it resembles ele<ians but is im-
mediately distinguished by spined elytral
apices, reduction of microreticulation of
elytra, and in other ways. In form of elytral
apices and reduction of microreticulation
it somewhat resembles laevii!,atus but is
more slender and convex, with only 1
median-lateral pronotal seta on each side.
The virtual obliteration of the middle part
of the posterior marginal line of the pro-
notum is diagnostic of this new species.
Catascopus hiroi n. sp.
Description. With characters of genus;
form as in preceding species (dobodura):
gr(.>en or blue-green, \\'ith some coppery
color at sides of elytra especially behind
humeri, lower surface and appendages
brown or brownish black; moderately
shining, front and disc of pronotinn with
reticulate microsculpture absent or vc^rx'
light, but disc of eh'tra entirely (trans-
versely) microreticulat(\ Head 0.99 and
1.00 width prothorax; front flat, slightK'
irregularly depressed, and with (usual)
longitudinal impression each side. Pro-
tliorax quadiati'-subcordatc; \\'idth IcMigth
1.41 and 1.34; base apex 1.13 and 1.08;
sides broadly arcuate anteriorlw sometimes
laintly angulate at median-lateral seta,
sinuat(> slightly less than ^i before right or
slightly acute basal angles; lateral margins
narrow but paralleled b\' slightly swollen
ridges acccMituating the marginal channels,
each with usual seta at basal angle and 1
median-lateral seta slightK' before middle;
basal marginal line entire in all specimens.
Elytra: width eKtra inothorax 1.63 and
1.58; ontei-apical angles sharplx delined,
right or slightK acute; apices each with
short spine near but not (juite at sutural
angle; striae well im]ii(\sse(l, slightK" iiune-
The Carabid Beetles of New Guinea • Darlington
107
tulate; intervals convex, not elevated at
base, punctulate. Measurements (of types):
length 8.7-9.3; width 3.5-3.7 mm.
Types. Holotype S ( Hungarian National
Mus.) and 2 paratypes from Stephansort,
Astrolabe Bay, N-E. N. G., 1897 (Biro);
and 1 paratype, Erima, Astrolabe Bay,
1897 (Biro). (Two paratvpes now in
M.C.Z., Type No. 31,411.)
Additional material. West N. G.: 1,
Waigeu Is., Camp 1, Mt. Nok, 2500 ft. (c.
760 m ), May 1938 ( Cheesman ) . This speci-
men is a (5 larger than the types (c. 11
mm ) and \\\\h disc of pronotum more
distinctly microreticulate, but it seems
clearly referable to hiroi.
Measured speeimens. The c5 holotype and
1 9 paratype from Stephansort.
Notes. This is another medium-small,
green species presumably related to the
preceding one (dobodura) but differing
in a number of details, including entire
basal marginal line of pronotum, presence
of distinct reticulate microsculpture on disc
of elytra, and position of elytral spines,
which are closer to the suture in bird than
in dobodura.
Cafascopus wallacei group
Catascopus wallacei Saunders and its im-
mediate relatives, including all the remain-
ing New Guinean species of the genus,
treated below, fonn an apparently natural
group of large, often conspicuously colored
species characterized by having both sutural
and outer-apical elytral angles acutely
toothed or spined and by having a longi-
tudinal zone of dense, conspicuous pubes-
cence along the midline of the body, from
prosternum almost to the tip of the ab-
domen in the i but mainly on the sterna
in the $ . Otherwise the species of this
group share the characters stated under the
genus, with minor exceptions.
The uallacei group of Catascopus centers
on New Guinea, where 5 species are now
known. Most of them are sympatric: 4 of
the 5 species have been found at Wau. Of
the 5 New Guinean species, aruensis and
wallacei each reach one or more small
islands to the west (Aru Is., Waigeu, Mysol),
and aruensis reaches also New Britain,
New Ireland, and Cape York in Australia.
A sixth species of the group (chaudoiri
Castelnau) is endemic in northern Aus-
tralia.
Some species of this group vary in-
dividually in form especially of the pro-
thorax, in degree of elevation of the 5th
elvtral intervals, and in some other details.
Although I can clearly recognize only the
5 species treated below, Straneo (see refer-
ences under the species) has distinguished
others, and he may be right. A thorough
study of long series will be required to
decide this, including study of genitalic
characters, which are indicated by Straneo.
I am, incidentally, very much indebted to
Prof. Straneo for loan of paratypes of his
3 species of this group.
Catascopus aruensis Saunders
Saunders 1863, Trans. Ent. Soc. London (3) 1,
p. 458, pi. 17, fig. 5a-b.
Csiki 1932, Coleop Cat., Carabidae, Harpalinae 7,
p. 1362 (see for additional references and
partial synonymy ) .
Straneo 1943, Ann. Mus. Civ. Genoa 61, p. 302.
Jedlicka 1963, Ent. Abhandlungen 28, pp. 382,
393.
cupricoUis Chaudoir 1883, in R. Oberthiir, Coleop-
terorum Novitates 1, p. 24 (not cupricoUis
W'aterhouse 1877).
brevispinosus Sloane 1910, Proc. Linnean Soc.
New South Wales 35, pp. 398, 400 (new
synonymy ) .
aeneicollis Andrewes 1919, Ann. Mag. Nat. Hist.
(9) 3, p. 481 (new name for cupricoUis
Chaudoir ) .
Andrewes 1924, Ann. Mag. Nat. Hist. (9) 14,
p. 593.
?dalbertisi Straneo 1943, Ann. Mus. Civ. Genoa
61, p. 304.
Description. With characters of genus
and of wallacei group (above); green or
blue-green (elytra rarely purplish); pro-
thorax more square (less cordate) than in
wallacei, but somewhat variable; elytra
with outer-apical angles spined, sutural
angles with shorter spines or simply acute
108 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
(variable); 5th intervals not or not much
elevated near base (slightly variable), 7th
intervals subcarinate at base; length c. 13-
15.5 mm.
Types. Of aruensis Samiders, from "Aru
[Is.]. Wallace," now in Oberthih- Coll.,
Paris Mus. Of cupricollis Chaudoir, from
Fly R., New Guinea, now also in Oberthiir
Coll., Paris Mus. Of brevispinostis Sloane,
from Coen, Cape York, Australia, now in
Sloane Coll., Canberra. Of dalhcHisi
Straneo, holotype from Hatam, Papua, in
Genoa Mus., and allotype from Andai,
Papua, in Straneo Coll. (See Notes, below.)
Occurrence in New Guinea. Widely
distributed but much less common than
wollaeei (below): 27 specimens, from all
3 political divisions of New Guinea and
from Normanby Is.; most from low alti-
tudes but reaching 1200 m at \Vau.
Notes. Outside New Guinea this species
occurs on the Aru Is. (type locality), New-
Britain, New Ireland, and Cape York,
Australia (types of hrevispinosus) . Pos-
sibly some of the outlying populations may
hv distinguishable as subspecies.
Of the types listed above, I have seen
only those of hrevispinosus (briefly in
1957, but Dr. B. P. Moore has sent me
additional notes on them) and the allotype
of (hilhertisi ( through the kindness of Prof.
Straneo ) .
Catascopus walJocei Saunders
Saunders 1863, Trans. Ent. Soc. London (3) 1, p.
462, pi. 17, fiK. 4a-l).
Csiki 1932, Coloop. Cat., Carabidae, Harpalinac 7,
p. 1367 (see for additional references).
Straneo 1943, Ann. Mus. Civ. Cenoa 61, p. 302,
fi.u;. a.
ledlicka 1963, Ent. Ahliandlunuen 28, pp. 382,
393.
?heccarn Straneo 1943, Ann. Mus. Civ. (ienoa 61,
p. 303, fig. h.
Description. With characters of gcinis
and of ualldcei group (above); usually
brightly bicolored, with head and elytra
purple and prothorax brassy or copjKMy,
but coloration sometimes duller; prothorax
a little more subcordat(> (less square) than
in aruensis, with posterior angles more
prominent and more acute; elytra with
sutural as well as outer-apical angles spined;
5th and 7th intervals elevated near base
(slightly variable, especially the 5th);
length c. 15-20 mm.
Tijpes. Of icallacei Saunders, from "Wa-
giou [Waigeu], Dorey and Mysol"; actual
(holo)t\'pe now in Oberthiir Coll., Paris
Mus. (not seen). Of beccarii Straneo, holo-
type from Hatam, Papua, in Genoa Mus.,
and allot\'pe from Andai, Papua, in Straneo
Coll. (allotype seen).
Occurrence in New Guinea. Common
probably throughout New Guinea: 176
specimens before me (including 82 from
Dobodura ) ; most from low altitudes, but
reaching 1300 m near Wau.
Notes. This beautiful carabid is appar-
ently confined to New Guinea and zoogeo-
graphically closely associated islands in-
cluding Aru Is., Waigeu, and Mysol. It
apparently does not reach the Moluccas
proper, nor New Britain, nor Australia.
Most of the indi\iduals from Dobodura
were trapped imder strips of burlap laid
across the trunks of fallen trees in rain
forest.
Catascopus strigicoHis Straneo
Straneo 1943, .\nn. Mus. Ci\ . Cenoa 61, p. 305,
fig. c.
Description. With characters of genus
and of udllacei group, except S as well
as 9 with 2 or 3 setae each side last ventral
segment; bicolored, head and prothorax
green or slighth' copper)", ebtra purple or
(especialK- basally) bluish or greenish; pro-
thorax almost square except sides sinuate
posteriorly (as usual in group); c^Ktra with
rather short spines at outer-apical angles
and still shorter (slightl\- variable) ones
at sutural angles; 5th as well as 7th interxals
raised near bas(>; length r. 1.5-18 mm.
Types. IIolot)pe (GcMioa Mus.) and
allot\iK> (Straneo Coll.) both from .\ndai,
Papua, Aug. 1872 (D'Albertis). I have
examined the allotype, loaned b\' courtesy
of Prof. Straneo.
The Carabid Beetles of New Guinea • Darlington
109
Occurrence in New Guinea. Papua: the
types. N-E. N. G.: 21, Wau, Morobc Dist.,
altitudes from 900 to 1500 m, dates in Mar.,
Apr., May, July, Aug., Sept., Oct., Dee.,
1961-1964 (Sedlaceks); 1, Sattelberg, Huon
Gulf, 1899 (Biro); 1, Wareo, Finschhafen
(L. Wagner, S. Australian Mus.).
Notes. This seems to be a distinct species,
immediately distinguished from wallacci by
head colored like pronotum ( not like
elytra), elytral spines shorter, and other
details, and from aruensis by 5th elytral
intervals raised, size usually larger, and S
with additional apical ventral setae. It
may occur mainly in mountains rather than
in lowlands, and it may be confined to part
of eastern New Guinea, but further collect-
ing is needed to confirm these possibilities.
Cafascopus taylori n. sp.
Description. With characters of genus
and of wallacei group; form (Fig. 63) of
large wallacei; head black or very dark blue,
prothorax brassy or slightly coppery, elytra
blue purple, lower surface and appendages
dark; head and pronotum rather shining
with reticulate microsculpture absent or
weak, elytra duller with close, slightly
transverse reticulate microsculpture. Head
0.77 and 0.81 width prothorax, impressed
across base; front with usual 2 longitudinal
impressions and slightly sculptured and
irregularly punctulate posteriorly. Prothorax
quadrate-subcordate; width length 1.55 and
1.51; base/apex 1.08 and 1.10; sides broadly
arcuate anteriorly with anterior angles flat-
tened and roundly produced, very broadly
sinuate posteriorly to right or acute slightly
denticulate posterior angles; side margins
rather wide (at widest point of prothorax,
width of the flattened margin is c. Vi
width from outer edge of margin to mid-
line of pronotum ) , flattened, reflexed, each
with usual seta at basal angle and 1 before
middle; disc finely transversely strigulose,
sparsely and faintly punctulate. Elytra long,
c. as in wallacei; width elytra/prothorax
1.42 and 1.34; outer-apical angles spined,
sutural angles acutely produced or spined
(individual variation); striae well im-
pressed, faintly punctulate; 5th and 7th
intervals elevated near base. Secondary
sexual characters normal for genus; c^ with
1, 2 2 setae each side last ventral segment.
Measurements: length c. 17-22; width c.
5.8-7.1 mm.
Types. Holotype S (M.C.Z., Type No.
31,412) from Aiura, N-E. N. G., 1900 m,
July 1962 (R. W. Taylor, #2147), in rain
forest; additional paratypes as follows.
N-E. N. G.: 4, Mt. Missim, Wau, Morobe
Dist., 950-1000, 1500, 1600-2000 m, Dec.
28, 1961, Aug. 10, Sept. 21-24, 1964 ( Sedla-
ceks); 1, Eliptamin Vy., 1200-1350 m, Aug.
16-30, 1959 (W. W. Brandt, Bishop Mus.);
4, Wareo, Finschhafen (L. Wagner, S.
Australian Mus.); 1, Moife, 15 km NW. of
Okapa, 2100 m, Oct. 7-14, 1959 (T. C. Maa,
Bishop Mus.); 2, Okapa, E. Highlands,
Apr. 20, 1964 ( Hornabrook ) ; 2, 13 km
SE. Okapa, 1650-1870 m, Aug. 26, 1964
(Sedlaceks); 2, Morae, Kukukuku [Rge.],
E. Highlands, 6000 ft. (c. 1850 m). Mar. 1,
1964 (Hornabrook). West N. G.: 3, Wissel
Lakes, Arabu Camp, 1800 m, Oct. 7, 1939
(H. Boschma, Leiden Mus.); 1, Wissel
Lakes, Enarotadi, 1800-1900 m, Aug. 10,
1963 (Sedlacek).
Measured specimens. The S holotype and
9 paratype from Enarotadi.
Notes. This new species seems close to
wallacei, from which it differs mainly in its
wider prothoracic margins. The difference
is striking on comparison of specimens.
Mainly because the prothoracic margins are
wider, the present new species has a rela-
tively narrower head and wider prothorax
as shown by measurements: in a measured
6 of wallacei the head is 0.90 width pro-
thorax and the prothoracic width/length is
1.40. Also, taylori averages larger than
wallacei and usually occurs at higher alti-
tudes.
Cafascopus rex n. sp.
Description. With characters of genus
and of wallacei group; form as in Figure
64; broad with very broad prothorax; green,
110 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
front and elytra bluish green, lower surface
and appendages brownish black; head
roughened and closely microreticulate pos-
teriorly, more shining but slightly strigulose
anteriorly, pronotum and elytra duller,
closely slightly transversely microreticulate.
Head large, but only 0.78 and 0.83 width
prothorax; depressed across base, with usual
longitudinal impression each side anteriorly.
Prothorax transverse-cordate, very wide but
with relativelv narrow base; width length
1.76 and 1.67;' base apex 0.90 and 0.89; side
margins rather broad especially anteriorly
and with anterior angles flattened and ad-
vanced, each with posterior and 1 median-
lateral seta, latter slightly farther forward
than usual. Elytra: width elytra/prothorax
1.23 and 1.29; outer-apical and sutural
angles both with short spines; striae well
impressed, scarcely punctulate; intervals
convex, slightly punctulate, 7th carinate at
base, others slightly humped but not cari-
nate. Secondary sexual characters of S nor-
mal for genus; 9 unknown. Measurements:
length c. 20; width 6.7-6.8 mm.
Types, llolotype i (Bishop Mus.) from
Mokai Village, Torricelli Mts., N-E. N. G.,
750 m, Dec. 8-15, 1958 (W. W. Brandt);
and 1 6 paratype (M.C.Z., Type No.
31,413) from Kiunga, Fly R., Papua, Sept.
24-25, 1957 (W. W . Brandt).
Notes. Within the uallacei group, the
large size and very broad, cordate pro-
thorax immediately distinguish this striking
species.
Genus PERICALUS Macleay
Maclcay 1825, Annulosa Javanica, p. 15.
Csiki 1932, Coleop. C^at., Caial)i(la(', I lurpalinac 7,
p. 1368 (sfc tor synonymy and adtlitional icln-
ences ) .
Jcannci 1919, Clolrop. C^arahiciucs dc la Rruion
Malj^aclic, Part 2, p. 1007 (in text).
Jedlicka 1963, Ent. Al)liandlungen 28, p. 373.
Dia<j,nosis. Similar to Catascopus (labrum
emarginate, ligula-paraglossae similar, 4(h
hind-larsal scgineuts scarcely emarginate,
claws simple, etc. ) but eyes more abruptl\-
jiroTuinetit; cl\]icus tnmcate; elytra usually
with geometric color pattern; size usual!)
smaller.
Description. None required here.
Type species. Pericalus cicindeloides
Macleay, of Ja\a, etc.
Generic distribution. Confined to, but
widely distributed and diverse in, tropical
Asia and the Malay Archipelago, reaching
New Guinea and New Britain but not
Australia.
Notes. The genus is represented in New
Guinea by only the following species
(fii^uratus), and on New Britain by a dif-
ferent, endemic species (klapperichi Jed-
Hcka 1953, Ent. Blatter 49, p. 145).
Pericalus figurafus Chaudoir
Chandoir 1861, Berliner Ent. Zeitschrift 5, p. 124.
Description. None required here; the
form (Fig. 65) and elytral markings make
this insect unmistakable, in New Guinea;
length c. 7-8 mm.
Type. Supposedly from Gelehes, col-
lected by Wallace; now in Oberthiir Coll.,
Paris Mus. (not seen).
Occurrence in New Guinea. Common
( more than 150 specimens ) probabK
throughout the island at low altitudes, and
occurring up to 1320 m nc>ar Wau.
Notes. So far as I know, this insect has
not been found in Celebes since Wallace's
time, and it has not been recorded from the
Moluccas. I think it is possible that the
type really cam(> from New Guinea and
that the species is endemic there. It li\es
on tree trunks and reccMith' fallen logs in
rain forest.
Genus COPTODERA Dejean
Dejean 1825, Species GeTieral (Coleop. 1, p. 273.
Csiki 1932, Coleop. Cat., Carahidae, llarpalinae 7.
p. 1370 (see for adilitional references, s>non-
ymy, and list of .species).
JcaiiiKd 1949, Coleop. Carabiqnes de la Retjion
\laluache. Part 3, pp. 921. 926.
jcdiida 19(i3. i:nt. \l)liand]nn,uen 28, p. 341.
Ecliiuxhild (;liand()ir 1883, Coleopterornni Xovi-
(atcs 1, p. 21 ( n(>\v s\i)on\in>' ).
?rri(h()C(>i)t(>(lcia Lonwerens 1958, Trenhia 24, p.
255.
The Carabid Beetles of New Guinea • Darlingion
111
Diag,nosis. See Keij to Genera of Lebiini
of Neic Guinea.
Description. Form (Figs. 66-70) broad,
± depressed; upper surface not pubescent
(in New Guinean species). Head: eyes
prominent; 2 setae over each eye; front im-
pressed each side anteriorly; clypeus with 1
seta each side; hibrum variable, usually
rather long, subtruncate or slightly emar-
ginate at apex, 6-setose; antennae with 3V-2
segments glabrous; mentum without tooth;
ligula 2-setose, paraglossae attached to but
much longer than ligula, broadly rounded,
without setae at apex but with small setae
at sides. Prof/jora.v broadly subcordate (ex-
cept in <!,rossa), with base sometimes lobed,
usually not; 2 setae each side, at basal angle
and before middle; disc with usual im-
pressions. Elytra broad; humeri broadly
rounded but rather prominent; apices with
outer-apical angles rounded, sutural angles
variable; striae entire, not distinctly punc-
tate; intervals convex but none specially
elevated; 3rd intervals with 2-4 seta-bearing
punctures (if 4, near base on outer edge,
c. Vi from base on outer edge, behind
middle on inner edge, and near apex usu-
ally on inner edge), but one or both inter-
mediate punctures missing in some species.
Inner icings full. Lower surface: proster-
num usually with a little sparse pubescence,
abdomen not pubescent. Legs slender; 4th
hind-tarsal segments simply emarginate; 5th
tarsal segments with accessory setae; claws
with 3 or 4 teeth. Secondary sexual char-
acters: i front tarsi slightly dilated, with
3 segments 2-seriately squamulose, and c^
middle tarsi with 2 segments squamulose
in some (not all) species; i middle tibiae
with 1 or 2 excisions on inner edge near
apex in most species (see Notes, below);
2 setae each side apex last ventral segment
in both sexes.
Type species. Of Coptodera, C. f estiva
Dejean, of Cuba; of Ectinocliila, E. tes-
selata Chaudoir [= aurata (Macleay)], of
Australia; of Trichocoptodera, T. maculata
Louwerens, of Celebes.
Generic distribution. In a broad sense,
the genus is pan-tropical. (In Jeannel's
restricted sense, Coptodera proper is con-
fined to the Americas, and related Old
World forms are divided into several
genera. ) In the Asiatic- Australian area,
species of the genus {sensu lato) are
numerous from southeastern Asia including
Japan across the whole Malay Archipelago,
and a few occur in Australia and New
Caledonia. For further details see Notes,
below.
Notes. Jeannel (1949) divides Cop^of/era
(sensu lato) and its immediate allies into
a number of small genera based primarily
on genitallic characters. It seems to me that
in this case, as in many others, Jeannel has
carried generic splitting beyond the limit
of usefulness. I have not attempted to
check the genitallic characters, which would
require dissection of many species from
many parts of the world. But I can say that,
if Jeannel's concept of genera were applied
to the New Guinean species, I would have
to divide Coptodera into about 5 genera,
2 or 3 of which would be new. The new
names would be meaningless except to
extreme specialists, and the fine splitting
would hide the broader relationships and
geographic patterns of the group. By treat-
ing the diverse New Guinean species as
members of one genus, I emphasize what
I think is a fact, that the group is a natural
one even though the species are diverse,
and that it has a pan-tropical distribution.
The most useful taxonomic treatment in
the end may be to retain Coptodera in a
broad sense but to divide it into a reason-
able number of natural subgenera. This
should, of course, be done on a worldwide
basis, not in a local faunal work.
Certain characters do vary remarkably in
this genus. The larger New Guinean species,
which are more typical of Coptodera, have
the base of the prothorax subtruncate, some-
times slightly oblique toward the sides but
not lobed. However, in 2 smaller New
Guinean species {papueUa and wau), the
112 Bulletin Museum of Comparative Zoology. Vol. 137, No. 1
base of the prothorax does have a distinct
short basal lobe. And in the Australian
"Ectinochih" tesseluta, the base of the
prothorax is more strongly lobed. All these
species have similar, diagnostic mouthparts
( mentum without tooth, and ligiila and
paraglossae as described), and the small
New Giiinean species are transitional in
other ways: they have wider prothoracic
margins and look more like Coptodem than
Ectinochila tesseJata does, and papueUa has
elytral markings like some more-typical
Coptodera, liut both papucUa and iiau ap-
proach Ectinochila in dense dorsal micro-
sculpture, and uau has Ectinocliila-Wke
elytral markings.
The dorsal elytral punctures confirm this
relationship. The number of punctures
varies in Coptodera. The full number is 4
on each 3rd interval, placed c. as noted in
the preceding Description. This is the ar-
rangement in the type species of the genus
(C. f estiva Dejean, of Cuba) and in some
of the more or less typical New Guinean
species, e.g., cyanclla and cluta. However,
^rossa and lincolata have the 3rd intervals
3-pimctate (puncture at basal % missing),
and oxijj)tera has the 3rd intei-vals only 2-
punctate (both median punctures missing,
leaving only the subbasal and subapical
ones). But tlie EctiiiocJiila-likv New Guinean
species (papuclla and wan) and also the
Australian E. tcssclata have the 3rd intervals
4-punctate as in typical Coptodera.
The excisi(ms of the i middle tibiae also
confirm the relationship of Ectinochila to
Coptodera. The 6 middle tibiae have a
single small excision (like that in lochia)
in inner edge near apex in the Cuban type
species of Coptodera {f estiva), in most New
(iuinc^an species of the genus including the
Ectinoclula-hkc ones, and in the Australian
E. tesselata. However, 2 mm-EctinocJiila-
like New Guinean species are different:
Coptodera ox\iptera has 2 small excisions
on each i middle tibia (like Aristolehia),
and C. ornati))ennis has none.
Most Coptodera ha\(> the dorsal surface
glabrous, but "Trichocoptodera^ maculata
Louwerens of Celebes has the pronotum
sparsely pilose. Coptodera ornatipennis
Louwerens of the Moluccas seems closely
related, and a paratype of it (which I owe
to the generosity of Mr. Louwerens) has a
few inconspicuous fine hairs still on the
pronotal disc. Specimens that I assign to
this species from New Guinea seem to
lack pronotal pubescence, but the hairs
may be rulibed off ( in light-trap speci-
mens ) or be adhering invisibly to the
pronotal surface (in specimens mounted
from alcohol). However, although I have
listed Trichocoptodera as a possible syn-
onym of Coptodera, it may eventually prove
worth recognition as a separate genus or
subgenus, distinguished by i middle tibiae
without excisions and perhaps by other
characters.
The (S species of Coptodera in New-
Guinea represent 7 stocks with different,
independent geographic distributions. ( 1 )
C. grossa is endemic and without close
relatives anywhere, so far as I know. (2) C.
ornatipennis occurs in the Moluccas as well
as New Guinea, with an apparent relati\ e
on Celebes. (3) C. cyanelki represents the
flexuosa group, which ranges from SE.
Asia to Australia (the Australian species
being r/?/.sYr^///.s- Chaudoir). (4) C lincolata
ranges from Celebes to New Guinea and
New Britain, and an apparently related
species {mastersi Macleay) is in Australia.
(5) C. cluta apparently occurs from SE.
Asia to New Guinea and New Britain,
and (6) C. oxyptcra, from Celebes to New
Guinea, New Britain, and New Ireland;
these species are not representc^d in Aus-
tralia. And (7) C. papucUa and nan are
endemic to New Guinea, probably related
to each other, and less closeK' related to
"Ectinocliihr aurata of Australia.
Most ol the common CU)pt()dcr(i in Xcw
Guinea inhabit tree trunks and recentK'
lallcn logs in rain lorcst. I low ever, a lew
species ol the genus elsewhc-re li\e among
dead leaxcs on the Uronnd. and this max* be
The Carabid Beetles of New Guinea
Darlington
113
the habitat of some of the less eommon
New Guinean ones.
Key to Species of Coptodera of New Guinea
1. Prothora.x not lobed at l)ase; dorsal micro-
reticulation moderate or partly absent; larger
( usually 5 mm or more, excepting small
individuals of lincolata ) ___ 2
- Prothorax lobed at base; dorsal micro-
reticulation close, heavily impressed; smaller
(3.5-4.8 mm) 7
2. Very large (8.5-9.5 mm); form as in Figure
66, with very long mandibles and transverse
prodiorax (p. 113) grossa
- Smaller; mandibles relatively shorter; pro-
thorax it subcordate 3
3. Head and disc of pronotum not microreticu-
late; each elytron with 2 irregidar, ± trans-
verse pale blotches ( Fig. 67 ) ; c^ middle
tibiae without excisions (p. 113) oniatipennis
- Head and disc of pronotum microreticulate
( lightly so in oxijptera ) ; c5 middle tibiae
with excision(s) on inner edge near apex _ . 4
4. Elytra with sutural angles blunt, narrowly
rounded; elytra usually (not always) con-
spicuously spotted or striped with pale 5
- Elytra with sutural angles acute, often dentic-
ulate; elytra unmarked or with only a few
inconspicuous minute pale flecks 6
5. Each elytron with 3 irregular pale blotches
sometimes joined to form an irregular longi-
tudinal stripe (p. 114) cyanella
- Elytra usually with numerous, more or less
separate, longitudinal pale lines (Fig. 68)
but pale pattern somewhat variable, some-
times almost obliterated (p. 114) ___. lineolata
6. Very broad; elytral striae less impressed;
color dark without pale markings; 3rd
elytral intervals with only 2 ( subbasal and
subapical) punctures (p. 115) oxijptcra
- Less Inroad; elytral striae deeply impressed;
elytra usually with minute pale flecks; 3rd
intervals 4-punctate (p. 115) cluia
7. Front of head heavily microreticulate but
not longitudinally rugulose; elytra irregularly
2-fasciate with pale (p. 115) papucUa
- Front of head in part longitudinally rugulose
as well as microreticulate; elytra with a
large, common, irregular X-shaped pale area
(Fig. 70) (p. 116) ivau
Copfodera grossa n. sp.
Description. With characters of genus;
form as in Figure 66; very large; reddish
black, appendages brown; .shining, elytra
faintly silky or subiridescent, reticulate
microsculpture absent or faint on front and
on disc of pronotum, distinct (but lightly
impressed) and transverse on elytra. Head
0.73 and 0.72 width prothorax; mandibles
exceptionally long, nearly straight; clypeus
rounded at sides, sinuately emarginate at
middle; labrum very long, narrowed an-
teriorly, obtusely emarginate; front almost
smooth posteriorly, slightly punctate an-
teriorly, as is clypeus. Vrothorax very wide,
formed as in Figure 66; width/length 1.93
and 1.82; base/apex 1.49 and 1.44; base
not lobed; side margins narrow, each with
seta almost at basal angle and less than
Vi from apex (farther forward than usual);
basal and apical marginal lines entire; disc
almost without punctation or strigae. Elytra
wide; width elytra prothorax c. 1.67 and
1.65; apices slightly obliquely sinuate, outer-
apical angles rounded, sutural angles blunt
or subdenticulate; striae well impressed,
punctulate; intervals convex, finely and
sparsely punctulate, 3rd with subbasal and
subapical seta-bearing punctures and 1 in-
termediate puncture on inner edge behind
middle. Secondary sexual characters: as of
genus except i with squamae on front tarsi
only ( not on middle tarsi ) ; i middle tibiae
with 1 excision; i copulatory organs as in
Figure 176. Measurements: length c. 8.5-
9.5; width 4.0-4.7 mm.
Types. Holotype $ (M.C.Z., Type No.
31,414) from Dobodura, Papua, Mar.-July
1944 (Darlington). Paratypes from N-E.
N. G.: is (Bishop Mus.), Wau (Mt.
Missim), Morobe Dist., 1100 m, July 22,
1961 (Sedlaceks); 2, Karimui, 1080 m, July
13, 1963 (Sedlacek); 1, Wareo, Finschhafen
(L. Wagner, S. Australian Mus.).
Measured specimens. The S paratype from
Wau and the 9 holotype.
Notes. Although this species scarcely
looks like a Coptodera, it has the essential
characters of the genus. It is distinguished
from other New Guinean species in the
preceding Key.
Coptodera ornatipennis Louwerens
Louwerens 1962, Tijdschrift voor Ent. 105, p. 146,
fig. 9.
Description. With characters of genus;
114 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
form and markings as in Figure 67; head
and pronotum reddish testaceous or reddish
piceous, elytra dark with pale marks as
shown, but marks somewhat variable;
reticulate microsculpture absent or faint
on front and on disc of pronotum, present
(but light) and transverse on elytra. Head
0.85 and 0.86 \\idth prothorax. ProtJiurax
wide-subcordate; width length 1.71 and
1.71; base apex 1.19 and 1.12; base not
lobed; disc with a little faint sparse punc-
tulation, not pubescent but margins with a
few short hairs near anterior angles (see
following Notes). Elytra: width elytra,
prothorax 1.61 and 1.62; sutural angles
blunt; 3rd intervals with subbasal and sub-
apical seta-bearing punctures but no inter-
mediate punctures. Secondary sexual char-
acters: 6 front tarsi \\'ith .3 segments with
squamae (as usual); 6 middle tarsi with
paired squamae at apex 1st segment and
on 2nd segment; 6 middle tibiae not ex-
cised. Measurements: length c. 5.0-6.5;
width 2..3-2.9 mm.
Types. From Amboina, Moluccas; in
Louwerens Coll. (1 paratype seen).
Occurrence in Neiv Guinea. Papua: 1,
Dobodura, Mar.-July 1944 (Darlington); 1,
Kokoda, 1200 ft. (366 m), Aug. 1933
(Cheesman). N-E. N. G.: 3, Finschhafen,
Huon Pen., 10 m, Apr. 9-16, 1963 (Sedla-
cek), in mercurv vapor light trap; 1, Pindiu,
Huon Pen., 890 m, Apr. 17, 1963 (Sedlacek).
in mercury vapor light trap. West N. (i.:
1, "Neth. New Guinea" [probably vie. Hol-
landia], Nov. 10, 1944 (T. Aarons, Cali-
fornia Acad. ) .
Measured sj)ecimens. Ai from Dobodura
and 9 from Finschhafen.
Notes. This species occurs in the MoIik*-
cas (the types) as well as in New (Guinea.
and it seems closely related to "Tricho-
coptodera" nuiculata Louwerens of C'elebcs.
My Moluccan paratype of ornalipennis
actually shows vestiges ol pronotal pubes-
cence. I can see no sign of it on the Ne\\
Guinean specimens, but the latter are prob-
ably all either from alcohol or Irom light
traps. New Guinean specimens do ha\c a
few inconspicuous short setae on the pro-
thoracic margins anteriorly, but such setae
are present in C. oxyptera too, and very
short ( vestigial? ) stubs of setae are visible
at 80x in some other species of Coptodera.
I suspect that this is a ground-living
rather than arboreal species.
Copfodera cyanella Bates
Bates 1869, Ent. Monthly Mag. 6, p. 74.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1370 (see for synon>niy and additional refer-
ences ) .
Lonwerens 1956, Treuliia 23, p. 225 (Moluccas).
Description. None required here; see
preceding Key: length ± 6-7 mm.
Type(s). From New (iuiiiea, collected
by Wallace (if really from New Guinea,
presumably collected at Dorey); now in
Oberthiir Coll., Paris Mus. (not seen).
Occurrence in New Guinea. Common
throughout New Guinea and on Normanby
Is.: 181 specimens (including 81 from
Dobodura and Oro Bay); most from low-
altitudes but up to 1200 m at Waw.
Notes. This species ranges west to the
Moluccas, ("elebes, and Borneo, and east
to New Britain. It is apparently related to
C. flcxuosa Schmidt-CToebel, which occurs
from SE. Asia to the Philippines, Celebes,
etc., overlapping the range of cyanella. C.
australis Chaudoir, of eastern Australia, is
apparently a distinct but related species.
Coptodera lineolafa Bates
Rates 1869, Knt. Mdiithly ^lat,^ 6, p. 74.
Csiki 19.>2, Coleop. (^at., Carabidae, Harpalinae 7,
p. 1371 (see lors\non\ni\ and additional refer-
ences ) .
Louwerens 1956, Trenhia 23, p. 225 (Moluccas).
Description. None required lure; see
preceding Key and Figure 68; length ± .5-6
iiiin.
'iypes. I'rom New (Fuinea, "collect(>d in
numbers | jiresumabK' at Dorey] b\' Mr.
\\ allace"; [)rcsumed t\'pe now in Oberthiir
Coll., Paris Mus. (not seen).
Occurroicc i)i Xcu- Guiwi'd. Common
l^robabK' throughout N<'w <^uinea and on
I^iak and Xonnanby Is.: 231 specimens
The Carabid Beetles of New Guinea • Darlington
115
(including 66 from Dobodura and Oro
Bay); most from low altitudes, but reaching
1200-1500 m at Wau and 1400 m at Ka-
rubaka. Swart Valley.
Notes. C. lineolata ranges from Celebes
to New Guinea and New Britain, and C.
mastcrsi Macleay of eastern Australia is
closely related.
New Guinean specimens vary in size
individually (not geographically) from c.
4.5 to 6.3 mm in length. The pale elytral
marks vary individually and perhaps also
geographically, although I cannot now de-
fine useful subspecies. Individuals from
Biak Is. have the markings notably reduced,
but variably so.
Copfodera eluta Andrewes
Andrewes 1923, Trans. Ent. Soc. London for 1923,
p. 30.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1370 ( see for synonymy and additional refer-
ences ) .
Lonwerens 1956, Treubia 23, p. 225 (Molnccas).
1964, Ent. Tidskrift 85, p. 181 (Borneo).
Jedlicka 1963, Ent. Aljhandlmmen 28, pp. 342, 349,
fiH. 103.
interrupta Chaiidoir 1869, Ann. Soc. Ent. Belsinni
12, p. 194 ( not interrupta Schniidt-Coebel
1846).
Description. None needed here; see Key
to Species of Coptodera of Neic Guinea,
and following Notes; length ± 6.5 mm.
Types. Both Chaudoir and Andrewes had
this insect from several different localities,
and neither designated a type. Its selection
should await careful study of specimens
from all pertinent localities, for the species
is variable, perhaps polytypic (Jedlicka,
1963), and often misidentified.
Occurrence in New Guinea. Twenty-six
specimens from numerous localities in all
3 political di\'isions of New Guinea; most
at low altitudes, but one at 1200-1300 m
at Wau.
Notes. This species apparently ranges
from SE. Asia to the Philippines, New
Guinea, and New Britain. Most New
Guinean individuals have the elytra slightly
flecked with pale, but some are almost
unmarked. These resemble C. oxyptera in
dark color and acute sutural angles but
differ strikingly in narrower form, deep
elytral striae, 3rd intervals 4-punctate, and
6 middle tibiae with only 1 subapical
excision.
Coptodera oxyptera Chaudoir
Chaudoir 1869, Ann. Soc. Ent. Belgium 12, p. 175.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1371 (see for additional references).
Louwerens 1956, Treubia 23, p. 225 (Moluccas).
Description ( for recognition only ) . Form
broad; color black, not marked; prothorax
usually with a few short fine setae on mar-
gins near apical angles; elytra with sutural
angles acute or acutely denticulate but not
spined; 3rd intervals with only subbasal and
subapical seta-bearing punctures; S front
but not middle tarsi squamulose, and S
middle tibiae each with 2 small excisions on
inner edge near apex; length c. 5.5-7.0 mm.
Type. From Celebes (Wallace), now in
Oberthiir Coll., Paris Mus. (not seen).
Occurrence in New Guinea. Common
probably throughout New Guinea: 145
specimens (including 77 from Dobodura);
most at low altitudes but reaching at least
1300 m at Wau and 1200 at Rattan Camp,
Snow Mts.
Notes. I have not seen specimens from
Celebes, but Chaudoii's description fits the
present species, specifying (partly by ref-
erence to his description of testrastigma)
pointed-denticulate but not spined elytral
apices and presence of only subbasal and
subapical punctures of the 3rd elytral inter-
vals. For comparison with eluta, see Notes
under that species, above. C. oxyptera, as
I identify it, occurs on Celebes, the Moluc-
eas, New Guinea, New Britain, and New
Ireland. It is not represented in Australia.
Coptodera papuella n. sp.
Description. With characters of genus;
form (Fig. 69) c. as in wau (below),
Coptodera -like but with prothorax lobed
or subpedunculate at base; dull green,
margins and much of base and apex of
prothorax and margins and markings of
116 Bulletin Museum of Comparative Zoology, Vol. 137, \o. 1
elytra testaceous, the elytral markings Ijeiiig surface. The basal lobe of the prothorax
2 transverse series of longitudinal lines of is intermediate between the usual (lobe-
varying length on the intervals, appendages less ) condition in Coptodcra and the very
irregularly testaceous; entire upper surface strong lobe in E. aiirata. In this and in
with close, heavily impressed, reticulate some other characters (see Notes under
microsculpture c. isodiametric on head and Coptodcra ) the present new species and the
disc of pronotum, scarcely transverse even following one ( icau ) connect Coptodcra
on elytra. Head 0.82 and 0.81 width pro- and Ectinochila and (I think) justify re-
thorax; mandibles rather short (in genus); ducing Ectinochila to synonymy,
labrum usually weakly emarginate at apex. Coptodcra papucUa is common on trunks
Prothorax wide-subcordate; width length and large branches of standing and recently
1.71 and 1.68; base/apex 1.14 and 1.19; fallen trees in rain forest.
base lobed, subpedunculate; base and apex
not margined at middle; disc with usual Copfodero wau n. sp.
impressions and also impressed each side. Description. With characters of genus
Elytra wide; width elytra prothorax 1.70 and (except as follows) of preceding species
and 1.68; apices obliquely sinuate-truncate (papucUa); form as in Figure 70; color c.
with outer and sutural angles rounded; as in papuclla except pale marks of elytra
striae impressed, not distinctly punctulate; fused to form a broad X, with anterior
intervals moderately convex, 3rd 4-punctate arms of X extending (narrowly) to humeri
as described for genus. Secondary .sexual and posterior arms more or less connected
characters: S front tarsi very narrowly across suture. Head 0.82 and 0.83 width
squamulose, middle tarsi not squamulose; £ prothorax; front longitudinally rugose an-
middle tibiae with 1 small excision or im- teriorly especially at sides, more irregularly
pression on inner edge near apex; 2 setae rugose posteriorly. Prothorax: width length
each side last ventral segment in both 1.69 and 1.70; base/'apex 1.26 and 1.19;
sexes. Measurements: length 3.5-3.9; width disc impressed each side (as in papuclla),
1.8-2.0 mm. with an area before middle relatively
Types. Holotype i (M.C.Z., Type No. shining and transverscK' microreticulate.
31,415) and 4 paratypes from Dobodura, Elytra: width cKtra prothorax 1.65 and
Papua, Mar.-July 1944 (Darlington), and 1.67; elytra slighth' humped near base, the
56 additional paratypes from Oro Bay raised area relatively shining and with
(near Dobodura), Dec. 1943-Jan. 1944 confused microsculpture, and other dark
(Darlington). areas of elytra slightK more shining than
Additional material. Forty-five, from nu- pale areas. Measurements: length 4.4—4.8;
merous localities in all 3 political divisions width 2.0-2.3 mm.
of New Guinea; most at low altitudes but Types. Holotype £ (Bishop Mus. ) and
up to 1150 m at Wau. Because the type 12 paratypes (some in M.C.Z., T> pe Xo.
series is adequate, because I expect to 31,416) from Wau, Morobe Dist., N-E.
distribute paratypes to all museums con- N. G., altitudes from 1200 to 1500 m, dates
cemed, and because some geographic vari- in jvme, Sept., Dee.. 1961-1962 (Sedlaceks),
ation (of markings) seems to occur, I have and additional paratxpes as follows. N-E.
restricted the type series to specinKMis Irom N. (i.: 1, Kaiiiantu, 1250 ni, Jan. 8, 1965
Dobodura and Oro liay. (Sedlacek); 3, ()kai)a, dattvs in Jan.. lune.
Measured specimens. The c5 holotype antl Sept., 1961, 1965 ( llornabrook ). West
1 9 paratype from Dobodura. N. G.: 6, I^uarotadi, Wissel Lakes, 1850-
Notes. This species resembles the Aus- 1900 in. liil\ 28, 1962 (Sedlacek).
tralian Ectinochila aurata (Macleay) in Measured specimens. The i holotype and
small size and dull, liea\ily microreticulate 1 ',■ i)aral\pi' bom Wau,
The Carabid Beetles of New Guinea • Darlington
117
Notes. For distinguishing characters and
place of this species among other New
Guinean Coptodcro, see Description above.
Notes under the preceding species (pop-
tiella ) , and tlie Key to Sj)ecies of Coptodcra
of New Guinea.
MINUPHLOEUS n. gen.
Diagnosis. See Key to Genera of Lehiini
of New Guinea., Figure 71; and Notes, be-
lo^^^
Descri})tion. Form broad, depressed;
some very short pubescence present above
and below, the hairs longer at sides of pro-
thorax and elytra. Head broad; eyes rather
small but prominent; 2 setae over each eye;
cKpeus subtruncate, 1-setose each side;
labrimi subparallel, rounded at sides an-
teriorly, notched at middle, 6-setose; an-
tennae short, reaching not or not much
beyond base of prothorax, pubescent from
apex 4th segment; mandibles ordinary;
mentum strongly toothed; ligula rather nar-
row, 4-setose; paraglossae attached to ligula
but longer, rounded, not setose; palpi
slender. Frothorax wide-subcordate; sides
of disc widely, irregularly depressed but
actual margins moderate or narrow, with
numerous lateral setae; basal marginal line
entire, apical marginal line weak or inter-
nipted at middle; disc with middle line
well impressed, basal transverse impression
very deep, anterior transverse impression
almost obsolete. Elytra: humeri prominent
but rounded, strongly margined; apices
slightly obliquely sinuate-truncate, with
outer angles broadly and sutural angles
narrowly rounded; striae entire; intervals
not elevated at base, 3rd with c. 4, 5th
with 1 or 2 (near base), 7th with c. 4 or
5 apparent special seta-bearing punctures
variable in position and difficult to identify
among other punctures. Inner wings full.
Legs: tarsi sparsely pilose above; 4th hind-
tarsal segment rather small, weakly emar-
ginate; 5th segment with accessory setae;
claws with c. 5 or 6 small teeth. Secondary
sexual characters: S front tarsi very little
dilated but with 4 segments squamulose,
the squamules slender, rather numerous,
not paired; c^ middle tarsi without squamae;
S middle tibiae with small excision on inner
edge near apex; 2 setae each side near apex
last ventral segment in both sexes; i copu-
latory organs as in Figure 177.
Type species. Minuphloeus mixttis, be-
low.
Generic distribution. That of the single
known species, below.
Notes. This insect differs from Minti-
thodes in form; the labrum is notched (not
notched in Minuthodes); and the lateral
pronotal setae are more numerous. It looks
a little like some Fhilophloeus (an Aus-
tralian genus unknown in New Guinea),
but the antennal pubescence is different
(antennae pubescent from middle of 3rd
segments in Fhilophloeus, from apex of 4th
in Minuphloeus); the labrum is different
(not notched in Fhilophloeus); etc. It
slightly resembles some Coptodera, but the
mentum is toothed (not in Coptodcra), and
the ligula is 4-setose (2-setose in Coptodcra).
Minuphloeus even resembles some wide,
depressed Catascopus, but the toothed
claws, excised i middle tibiae, and other
characters differentiate it from that genus.
I am therefore forced to treat the insect as
a new monotypic genus, exact relationships
undetermined, occurring ( so far as known )
only in a small area in New Guinea.
The name of the new genus is formed
by combining the first two syllables of
Minuthodcs with the last two of Fhilo-
phloeus.
Minuphloeus mixtus n. sp.
Description. With characters of genus;
form as in Figure 71; black, shining, most
of upper surface without reticulate micro-
sculpture but extensively punctate. Head
0.84 and 0.83 width prothorax; front ir-
regularly impressed, irregularly punctate,
with short longitudinal ridge each side in-
side position of anterior supraocular setae.
Frothorax: width/length 1.82 and 1.80;
118 BuUetin Museum of Comparatwc Zoology, Vol. 137, No. 1
base/apex 1.04 and 1.04; margins with
numerous strong setae irregularly spaced in
whole length; dise finely, sparsely, irregu-
larly punctate. EJijtro: width elytra/pro-
thorax 1.51 and 1.55; striae moderately im-
pressed, closely punctate; intervals sHghtly
con\ex, sparsely punctate. Measurements:
length c. 7.0-8.5; width c. 3.3-4.0 mm.
Types. Holotype S (Bishop Mus.) and
32 paratypes (some in M.C.Z., Type No.
31,417) from Wan (including Mt. Missim,
Kunai Ck., Mt. Kaindi), Morobe Dist., N-E.
N. G., 900, 1200, 1300, 1400, 1500, 1500-
1800, 1600-2000 m, dates in Jan., Feb.,
May, June, July, Aug., Sept., Nov., Dec,
1961-1964 (Sedlaceks), and additional para-
types as follows. N-E. N. G.: 1, Moife,
15 km NW. of Okapa, 2100 m Oct. 11-13,
1959 (T. C. Maa, Bishop Mus.); 8, Okapa,
dates in Apr., Aug., Oct., 1964, Mar. 1965
(Hornabrook). West N. G.: 19, Enarotadi,
Wlssel Lakes, 1800, 1800-1900, 1850-1950
m, dates from July 19 to Aug. 4, 1962
( Sedlacek ) .
Measured specimens. The i holotype and
1 9 paratype from VVau.
Notes. See Notes under genus. The
insect looks as if it lived on tree trunks or
under bark, but its actual habitat is not
recorded.
Genus AGONOCH/LA Chaudoir
Chaudoir 1848, Bull. Soc. Nat. Moscow 21, Part 1,
p. 119.
1869, Asm. Soc. Eiit. B(.'l,<riuin 12, p. 223.
Csiki 1932, Colcop. Cat., Caiabidae, Harpalinae 7,
p. 1379 (sec for additional references and list
of species ) .
Sloane 1898, Proc. I^inncan Soc. New South Wales
23, p. 494 (in key to Australian .uencra of
Lebiini ).
Diaii,nosis. See Key to Genera of Lebiini
of New Guinea.
Description: characters common lo the
New Guinean species of the genus (Aus-
tralian species are more diverse). Form as
in Figures 72-76; small, broad, ± convex;
short-pubescent above and below, and part
or all of upper surface also closely punctate
or (at least elytra) roughened. Head: eyes
prominent but not large; 2 setae over each
eye; clypeus subtruncate, 1-setose each side;
labrum broadly rounded or subtruncate, 6-
setose; mentum toothed; ligula with 2 long
and usually 2 short setae, paraglossae c.
as long as or slightly longer than and at-
tached to ligula; palpi, especially penulti-
mate segments, short. Protliorax variable
in form (see Figs, cited); base ± arcuate
at middle but not strongly lobed; lateral
margins narrow to \\'ide, each with seta at
base and at (usually) or slightly before
middle of length; base and apex with lightly
impressed marginal lines sometimes faint or
interrupted at middle; disc with moderate
middle line and transverse impressions.
Elytra: humeri moderately prominent,
rounded; apices obliquely sinuate-truncate,
with outer angles broadh' and sutural angles
narrowly roimded or blunted; striae entire
or nearly so but usually lightly impressed,
not sharply defined; 3rd intervals apparently
usually 3- or 4-punctate, but dorsal punc-
tures difficult to identify amid other punc-
tation and pubescence. Inner icings full.
Legs slender; 4th tarsal segments weakly
emarginate; 5th segments with accessory
setae; claws with c. 4 short teeth. Secondary
sexual characters: i front tarsi slightK'
dilated, soles formed of many squamae not
arranged in 2 series ( in all species of which
S 6 in satisfactory condition are axailable);
i middle tibiae with small notch or im-
pression on innc>r edge just before apex
(except in expa)isa); 2 setae each side last
ventral segment in both sexes.
Tyj)c sj)ecies. A. guttata Chaudoir, of
southern Australia (only spc^cies mentioned
by Chaudoir in 1848).
(U'ncric distribution. Many species in
Australia; 1 Australian species also (intro-
duced?) in Now Zralaiul; 7 small species
in New Guinea, chieih in lower nioimtains.
Notes. The 7 New Guinean species that
I assign to this genus difler among them-
seKcs, but they all seem to belong to one
small group ol (he genus that may be
The Carabid Beetles of New Guinea • Darlington 119
restricted to New Guinea and the adjacent
tropical part of Australia. Described Aus-
tralian species of the group probably in-
clude Ag,o nod lil a ovalis Sloane and intricata
Sloane ( both described 1923, Proc. Linnean
Soc. New South Wales 48, p. 39), and I
have specimens representing one or more
forms of this group from North Queensland,
Australia, from rain forest on and near the
Atherton Tableland. This group of small,
pubescent species, with notched 6 middle
tibiae, jnay prove to be worth generic sepa-
ration from A'j^onochila, but the Australian
Agonochila need much more study before
division of the genus is undertaken.
Most Australian A<ionochila live on tree
trunks, especially on shaggy-trunked Eii-
ccihjptus trees. The habitat of the New
Guinean ones is not recorded but is prob-
ably in rain forest.
Key to Species of Agonochila of New Guinea
1. Elytra with pattern of many pale longitudinal
dashes in 3 irregular transverse series which
cover nearly the whole elytra (Fig. 72) 2
- Elytra differently marked or not marked 3
2. Prothorax not depressed at sides (p. 119) ____
viinuthoides
- Prothorax depressed at sides (p. 119) . ,.
duplicata
3. Prothorax not subcordate; anterior prothoracic
angles broadly rounded-in 4
- Prothorax broadly subcordate 6
4. Elytra dark with single c. regular, common,
red or testaceous area behind middle (p.
120 ) gressitti
- Elytra not marked as described 5
5. Elytra with markings varying from isolated
pale flecks to irregular X-pattern ( Figs.
74, A, B) (p. 120) variabilis
- Color entirely red, without elytral markings
( p. 120 ) nifa
6. Elytra with 2 irregular transverse pale fasciae
behind middle (Fig. 75) or with markings
expanded (Fig. 75A); length 5.3-5.7 mm
(p. 121) cxpansa
- Elytra with a large common pale area (Fig.
76) or single broad post-median fascia;
length 6.0-6.7 mm (p. 121) dorsata
Agonochila minuthoides n. sp.
Description. With characters of genus;
form as in Figure 72; irregular dark reddish
brown, elytra with complex pattern of short
pale lines in 3 irregular transverse series,
appendages testaceous; most of upper sur-
face irregularly punctate or roughened but
surface of head and pronotum shining be-
tween punctures. Head 0.81 and 0.80 width
prothorax. Prothorax transverse-quadrate;
width length 1.57 and 1.53; base/apex 1.28
and 1.28; side margins very narrow, with no
flattened areas inside margins. Elytra:
width elytra/prothorax 1.64 and 1.66. Mea-
surements: length 4..3-4.8; width 2.2-2.4
mm.
Tiipes. Holotype $ (M.C.Z., Type No.
31,418) from Didiman Ck., Lae, N-E. N. G.,
Mar. 27, 1955 (E. O. Wilson), in lowland
rain forest; 1 $ paratype, Busu R., E. of
Lae, 100 m, Sept. 14, 1955 (Gressitt); and
1 9 paratype, Sattelberg, Huon Gulf, N-E.
N. G., 1899 (Biro).
Measured specimens. The $ holotype and
9 paratype.
Notes. The color pattern and very nar-
row prothoracic margins distinguish this
species. The 3 known specimens are all
from a rather small area in northern N-E.
N. G., but it would be unsafe to assume
that the species is really so localized.
The complex color pattern of this small
lebiine is so like that of some Minuthodes
and of Coptodcro lineolata as to suggest
mimetic convergence.
Agonochila duplicata n, sp.
Description. With characters of genus;
form as in Figure 73; irregular dark reddish
brown, elytra with complex pattern of short
pale lines in 3 irregular transverse series
(much as in preceding species, minuthoides);
appendages testaceous; much of upper sur-
face irregularly punctate, but surface shin-
ing between punctures. Head 0.71 width
prothorax. Prothorax wide; width/length
1.77; base/apex 1.25; margins broadly de-
pressed. Elytra: width elytra/prothorax
1.50. Measurements: length c. 4.5; width
2.2 mm.
Type. Holotype i (Hungarian National
120 BuUetin Mitscuiu of Comparative Zoology, Vol. 137, No. 1
Mus.) from Sattelberg, Huon Gulf, N-E.
N. G., 1899 (Biro); the type is unique.
Notes. Although the individual deseribed
above is colored much like the preceding
species (miniiflioidcs) and occurs within
the range of the latter, I think it is distinct.
The wider prothoracic margins are striking,
and the greater width they give the pro-
thorax is reflected in the proportions, the
head being relatively smaller, the prothorax
wider, and elytra relatively narrower in
dupJicata than in minuthoidc.s.
Agonochila gressitti n. sp.
Description. With characters of genus;
form slender; head and prothorax red or
reddish brown, elytra slightly darker ( often
nearly black) \\'ith large, common, red or
testaceous area behind middle varying in
size and shape but always with relatively
regular margin (compared with some fol-
lowing species); appendages testaceous;
most of upper surface punctate but mod-
erately shining between punctures. Head
0.65 and 0.68 width prothorax. Frothorax:
width length 1.72 and 1.76; base/apex not
calculated (anterior angles too rounded-in
for exact measurement of apex); sides
arcuate through most of length, sometimes
faintly subangulate at inedian-lateral setae;
posterior angles obtuse, slightly blunted;
margins moderate. Elytra: width elytra/
prothorax 1.53 and 1.62. Measurements:
length 4.2-5.5; width 2.1-2.7 mm.
Types, flolotype S (Bishop Mus.) and
5 paratypes (2 in M.C.Z., Type No. 31,419)
all from Swart Vy., Karubaka, N-E. N. G.,
1500-1550 m, dates in Nov. 1958 (Gressitt).
Addidoiial material. N-E. N. G.: 2,
Adalbert Mts., Wannma, 800-1000 ni, Oct.
26, 27, ]958 (Gressitt, 1 specimen bearing
his number 3222); 1, W'um, l^pper jiniuii
Vy., 840 m, July 16, 1955 (Gressitt)."
Measured specimens. The A holotype and
1 9 paratype.
Notes. The lorm plus coloration of this
species are diagnostic, in New Ciuiiiea.
The specimens listed under Addilional ma-
terial are slightly smaller and less sharply
bicolored than the types but seem to be
conspecific.
Agonochila rufa n. sp.
Description. With characters of genus;
form c. as in va rial) ills (following species);
entirely rufous, not marked; most of upper
surface moderately punctate but head and
pronotum shining between punctures, elytra
duller; appendages testaceous. Head 0.71
and 0.69 width prothorax. Prothorax:
width length 1.64 and 1.65; base apex not
calculated ( anterior angles rounded-in ) ;
sides faintly angulate at median-lateral
setae, slightly sinuate before c. right
(slightly obtuse) basal angles; margins
rather narrow. Elytra: width elytra pro-
thorax 1.66 and 1.63. Measurements: length
c. 4.0-4.5; width 2.1-2.3 mm.
Types. Holotype 9 (Bishop Mus.) from
Bisianumu, E. of Port Moresby, Papua, 500
m, Sept. 22, 1955 (Gressitt); and paratypes
as follows. Papua: 2 9 9 (1 in M.C.Z.,
Type No. 31,420), Kokoda-Pitoki, 450 m.
Mar. 23, 24, 1956 (Gressitt); 1 i , Keparra-
Sangi, nr. Kokoda, 500 m, ^hlr. 26, 1956
(Gressitt), "Sago palm." N-E. N. G.: 1.
Wareo, Finschhafen (L. W'agner, S. Aus-
tralian Mus.).
Measured specimens. The 9 holotype and
1 9 paratype from Kokoda-Pitoki.
Notes. The plain rufous color is diagnostic
for this species in this genus in New
Guinea.
Agonochila variabilis n. sp.
Description. With characters of genus;
form as in Figure 74; irregularh- brownish
black with \arial)le elytral markings pale
( Figs. 74, A, B ) ; appendages brownish
testaceous; most oi upper surhice punctate
but shining between punctures, elytra
slightK' l(\ss shining. Head 0.70 and 0.71
width prothorax. Prothorax: width length
1.62 and 1.67; base aj^ex not ealc-ulated
(anterior angles rounded-in ); margins mod-
erate. Elytra: width elytra prothorax 1.58
and 1.63. MeasuremcnI.s: length c. 4.0-
4.5; width 2.1-2.3 mm.
The Carabid Beetles of New Guinea • Darlington 121
Types. Holotype S ( Bishop Miis. ) and
13 paratypes (some in M.C.Z., Type No.
31,421 ) all from Wissel Lakes, West N. G.,
with following additional details: holotype
and 1 9 paratype, Uiapnra, Kamo Vy.,
1530 m, Aug. 11, 15, 1955 (Gressitt); 1
paratype, Wagete, Tigi L., 1700 m, Aug. 17,
1955 (Gressitt); 10 paratypes, Enarotadi,
altitudes from 1750 to 1900 m, dates in
Aug. 1955 (Gressitt) and July, Aug. 1962
(Sedlaeek); 1, Moanemani, Kamo V., 1500
m, Aug. 13, 1962 (Sedlaeek).
Additional material. Papua: 2, Mafulu,
4000 ft. ( c. 1230 m ) , Jan. 1934 ( Cheesman ) .
N-E. N. G.: 24, Wau, Morobe Dist., alti-
tudes from 1100 to 1450 m, dates in all
months except Apr., June, Nov., 1961-1963
(Sedlaeek); 1, Mt. Mis(s)im, Morobe Dist.,
5850 ft. ( c. 1780 m ), Apr. ( Stevens, M.C.Z.).
Measured specimens. The <5 holotype and
9 paratype from Urapura.
Notes. The specimens from Wissel Lakes
vary surprisingly in elytral pattern (Figs.
cited). Of the 2 from Mafulu, 1 has mark-
ings comparable to those of the most heavily
marked Wissel Lakes individual, and the
other is even more heavily marked. The
Wau and Mt. Mis(s)im individuals are
heavily marked ( Fig. 74B ) but somewhat
variable. The variation is obviously partly
individual, but heavy markings are ap-
parently commoner in eastern than in west-
ern New Guinea.
Agonochila expansa n. sp.
Description. With characters of genus
except as noted below; form (Fig. 75)
broad, with wide-subcordate prothorax;
black, elytra with 2 irregular, interrupted
pale fasciae behind middle, the posterior
one narrower and more interrupted, the
fasciae sometimes partly joined and ex-
tended anteriorly on each elytron (Fig. 75 A);
appendages irregularly blackish brown; en-
tire upper surface punctate but moderately
shining between punctures, and head also
obliquely-longitudinally rugulose at sides
between eyes. Head 0.76 and 0.76 width
prothorax. Prothorax: width/length 1.89
and 1.80; base/apex 1.18 and 1.18; side
margins broadly depressed, with median-
lateral setae before middle of prothoracic
length. Elytra: width elytra prothorax 1.52
and 1.59. Secondary sexual characters as
for genus except i middle tibiae not ex-
cised or impressed near apex. Measure-
ments: length 5..'3-5.7; width 2.5-2.8 mm.
Types. Holotype 9 ( Bishop Mus. ) from
Finisterre Rge., Saidor, Kiambavi Village,
N-E. N. G., Aug. 1-28, 1959 (W. W.
Brandt), and paratypes as follows. N-E.
N. G.: 1 S in poor condition (M.C.Z.,
Type No. 31,422), Wau, Morobe Dist.,
1400-1500 m, Dec. 20, 1961 (Sedlaeek); 6,
Okapa, xMar. 23, Apr. 4, 1964 (Homabrook);
1 6 (with expanded markings), 11 km S.
of Mt. Hagen (town), N-E. N. G., 2000-
2300 m. May 20, 1963 (Sedlaeek).
Measured specimens. The S paratype
from Wau and the 9 holotype.
Notes. In form and markings this species
looks more like a Coptodera than an Agono-
chila, but it has the characters of the latter
genus, as here defined. The middle tibiae
lack excisions in both 6 6 listed, but this is
probably a specific ( not generic ) character,
for the follo\\'ing species (dorsata), which
seems close in most ways to the present
one, has the notch present, but weak,
Agonochila dorsata n. sp.
Description. With characters of genus;
fonn as in Figure 76; black or irregularly
reddish black, elytra either with large testa-
ceous area as figured or the pale area re-
duced to a single transverse post-median
fascia; entire upper surface closely punc-
tate, but ± shining between punctures.
Head 0.82 and 0.82 width prothorax; front
especially at sides slightly rugulose as well
as punctate. Prothorax broadly cordate;
width length 1.84 and 1.80; base apex 1.13
and 1.15; sides broadly depressed, with
median-lateral seta slightly before middle.
Ehjtra: width elytra/ prothorax 1.58 and
1.65; sutural angles better defined than
122 Bulletin Museum of Comparatwe Zoology, Vol. 137, No. 1
usual, scarcely blunted. Secondonj sexual
characters as for genus, including 6 middle
tibiae impressed on inner edge near apex.
Measurements: length 6.0-6.7; width 2.9-
3.3 mm.
Types. Holotype S (Bishop Mus.) and
11 paratvpes (some in M.C.Z., Type No.
31,423) from Kepilam, N-E. N. G., 2420-
2540 m, June 21 and 23, 1963 (Sedlacek).
Additional material N-E. N. G.: 3, 11
km S. of Mt. Hagen (town), 2000-2300 m.
May 20, 1963 (Sedlacek); 1, Edie Ck.,
Morobe Dist., 2000-2100 m, Oct. 5-10, 1963
(Sedlaceks); 1, Kainantu, 2150 m, Jan. 8,
1965 (Sedlacek).
Measured specimens. The i holotype and
1 $ paratype.
Notes. This is the largest Agonochila in
New Guinea, and it occurs at relatively
high altitudes. The form plus markings are
diagnostic. The testaceous area of the elytra
varies geographically: it is large (c. as in
Fig. 76) in the whole type series, but
reduced to a (broad) transverse fascia
(Fig. 76A) in all specimens listed under
Additional material. However, I do not
wish to make subspecies without seeing
more material from more localities.
Genus OXYODONTUS Chaudoir
C:liaucl()ir 1869, Ann. Soc. Ent. Beljj;iuni 12, p. 239.
Diagnosis. See Keij to Genera of Lebiini
of Neic Guinea; note especially form, small
size, long acute mentum tooth, rounded-
oblique elytral apices, and plainly 3-punc-
tate 3rd elytral intervals.
Descri})tion. Form as in Figure 77; part
of surface including pronotum and sides of
elytra very inconspicuously setulose. Head:
eyes prominent, 2 setae over each eye;
labrum ± rounded, 6-setose; mentum with
long, acute tooth; ligula narrow, with 2
long and 2 shorter setae; paraglossae r.
long as ligula, attached, wide, without
setae. Vrothorax with usual 2 setae each
side. Elytra formed as figured; apices
rounded-oblique; striae entire, moderately
impressed; 3rd intervals strongly 3-punctate,
with punctures c. Vi from base on outer
edge, and near middle and apex on inner
edge. Inner icings full. Legs slender; 4th
tarsal segments small, weakly emarginate;
5th segments with accessory setae; claws
with c. 3 teeth. Secondary sexual characters:
i front tarsi slightly dilated, 3 segments
with narrow squamae not in 2 regular series;
c5 middle tibiae with minute but deep
excision on inner edge just before apex; 6
with 1, 9 2 setae each side near apex last
ventral segment.
Ty})e species. O. tripunctatus Chaudoir
( below ) .
Generic distribution. That of the single
species.
Notes. The relationships of this incon-
spicuous genus are not clear.
Oxyodontus tripunctatus Chaudoir
Chaudoir 1869, Ann. Soc. Ent. Belgium 12, p. 239.
Louwerens 1956, Treubia 23, p. 226 (Moluccas).
Description. None required here. See
under genus, of which this is the only known
species, and see Figure 77; length c. 4-4.5
mm.
Types. Two specimens from Celebes,
collected by \\'allace; type now in Ober-
thiir Coll., Paris Mus. (not seen).
Occurrence in Netc Guinea. Common
probably throughout New Guinea: 78
specimens (including 53 from Dobodura),
from localities in all 3 political dixisions of
New Guinea; chiefK' at low altitudes, but
to 1300 m at Wixu.'
Notes. This species has been prc^xiousK
recorded from Celebes and the Molueeas,
and I have a series of it (or of a closcK'
related species) also from Leytc and Lu/on
in the Philippines. It is not known in New
Britain or Australia. I think it lives in
und(>rst()ry foliage in rain forest, but my
scanl\- field notes are not ch^ar about this.
Genus MOCHTHERUS Schmidt-Goebel
Scliniitlt-(;oc])C'l 1846, Fannnia Colcop, i^irniaiiiae,
p. 76.
Csiki 1932, Coleop. Cat., Carahitlac Harpalinac 7,
The Carabid Beetles of New Guinea • Darlingion 123
p. 1382 ( see for additional references, synonymy,
and list of species).
Jedlicka 1963, Ent. Abhandlun.ucn 28, p. 352.
Dkipiosis. See Key to Genera of Lehiini
of New Guinea., and note form (Fig. 78),
unarmed elytral apices, and minutely setu-
lose pronotal disc.
Description. Form c. as in Figure 78.
Head: eyes large; 2 setae over each eye;
labiiim subtruncate, not emarginate, 6-
setose; mentum weakly, usually obtusely
(variably?) toothed; ligula and paraglossae
subequal, attached, together wide, 4-setose;
palpi slender. Prothorax cordate, with usual
2 setae each side. Elytra wide, unarmed;
apices slightly obliquely sinuate-truncate;
3rd intei'vals 2-punctate, the punctures on
inner edge behind middle and near apex.
Inner wings full. Legs slender; 4th tarsal
segments small, scarcely emarginate; 5th
segments with (weak) accessory setae;
claws each with 2 long and 1 shorter tooth.
Secondary sexual characters: $ front tarsi
scarcely dilated, 3 segments with paired
squamae; i middle tibiae not excised; i
with 1, 9 2 setae near apex each side last
ventral segment.
Type species. M. angiilatiis Schmidt-
Goebel {= tetraspilotus Macleay) of SE.
Asia, etc.
Generic distribution. SE. Asia including
Japan, and across the Malay Archipelago
to the Philippines and New Guinea (not
Australia), with one species recorded (in-
troduced?) also on Christmas Is. and
Samoa.
Notes. A single common species of the
genus occurs on New Guinea.
Mochtherus obscurus (Sloane)
Sloane 1907, Deutsche Ent. Zeitschrift for 1907,
p. 183 {?Simtms).
Andrewes 1927, Ann. Mag. Nat. Hist. (9) 19, p.
110.
immacuJatus Maindron (not Redtenbacher) 1908,
Nova Guinea 5, p. 299.
Description ( for recognition only ) . With
characters of genus; form as in Figure 78;
dull black, surface minutely short-setulose;
length c. 6-7 mm.
Type. From Sattelberg, N-E. N. G.;
should be in Deutsches Entomologisches
Institut, Berlin-Dahlem (seen by An-
drewes ) .
Occurrence in New Guinea. Common
probably throughout New Guinea ( 130
specimens, including 79 from Dobodura),
most at low altitudes, but reaching 1200
m at Wau ( only 1 specimen at this altitude )
and 1530 m on the Salawaket Rge. (2 speci-
mens). Found also on Normanby Is. (2
specimens ) and Waigeu Is. ( 1 ) .
Notes. This species occurs also on New
Britain (6 specimens including 3 from
Gazelle Pen.) and New Ireland (3).
The relationship of the New Guinean
obscurus to asemus Andrewes (recorded
from the Moluccas by Louwerens 1956,
Treubia 23, p. 226) and to other species
farther west in the Malay Archipelago re-
mains to be determined. In general the
"species" seem very closely inter-allied in
this genus, and some may prove to be
geographic subspecies.
This insect lives on and under the bark
of tree trunks and recently fallen logs in
rain forest.
(Genus MOCHTHEROIDES Andrewes)
Andrewes 1923, Trans. Ent. Soc. London for 1923,
p. 50.
Jedlicka 1963, Ent. Abhandlungen 28, p. 352.
Diagnosis. See Key to Genera of Lebiini
of New Guinea.
Description. Form c. as in Figure 79.
Head: eyes moderate; 2 setae over each
eye; mandibles moderate; labrum rounded-
truncate, not (or scarcely) emarginate, 6-
setose; mentum obtusely prominent at
middle but scarcely toothed; ligula wide,
4-setose, with paraglossae not attached (ex-
cept at base), longer and narrower than
ligula. Prothorax subcordate, with very
narrow margins, each with usual 2 setae.
Ehjtra with apices obliquely sinuate-trun-
cate, unarmed; striae entire; 3rd intervals
with 1 seta-bearing puncture on inner edge
at extreme apex but othei-wise impunctate.
124 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Legs slender; 4th tarsal segments emar-
ginate for less than half of length; 5th
segments with accessory setae; claws each
with c. 4 very small teeth. Inner icings
full. Sceonckmj sexual charoeters: 6 front
tarsi wider than in Moehfhcnts, with 3 seg-
ments 2-seriately squamulose; c5 middle
tibiae not excised; i with 1, 9 with 2
setae near apex each side last ventral seg-
ment.
Type species. Masoreiis sericans Schmidt-
Goebel, of Burma, etc.
Generic distribution. Known from
Burma, Singapore, Sumatra, Philip-
pines, and New Britain; not recorded from
New Guinea, but may occur there.
Notes. Mochthcroides superficially re-
sembles MochtJierus but the two genera are
probably not related. They differ in mouth-
parts, punctures of 3rd elytral intervals,
claw teeth, etc.
(Mochtheroides niger Jedlicka)
[t'dlic'ka 1934, Acta Soc. Ent. Prai^ue 31, p. 122.
'■ 1963, Ent. Abhandlungfii 28, p. 352.
Description ( for recognition only ) . With
characters of genus; form as in Figure 79;
black, most of surface (except prosternum)
not setulose. ProtJwrax with margins very
narrow. Elytra with 3rd intervals with only
1 (apical) seta-bearing puncture; length c.
4.5-4.8 mm.
Type. From Sibu>'an Is., Philippines;
in Andrewes Coll., British Mus. (se(Mi).
Occurrence in Netc Guinea. Not yet
found, but may occur.
Notes. Three specimens that T collected
at Cape Clouccster, New Britain (under
th(^ bark of a small dead tree) seem in-
distinguishable from the Philippine type,
with which I compared them in 194S. This
distribution suggests that the species will
be louud in New Guinea too.
Genus DOUCHOCTIS Schmidt-Goebel
Schmidt-Goebel 1846, Faunula Cok-op. Bimiainac,
p. 62.
Andrewes 1931, Zoo). Mcdedelingcn 14, pp. 62-64
(key to SuiiKitraii species).
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1383 (see for additional references and list
of species ) .
Louwerens 1958, Treubia 24. pp. 258, 259 (com-
ments on some species ) .
Jedlicka 1963, Ent. Abhandlunsen 28, p. 356.
Diagnosis. Small Lebiini, rather diverse
in form; not pubescent; mentum without
tooth; ligula and paraglossae fused into a
broadly rounded whole, with usually 2
principal and several slightly smaller setae;
each 3rd elytral interval usually \N'ith 2
minute punctures behind middle, these
punctures without setae.
Description. Form variable (Figs. 80-
85); not pubescent above, with or without
reticulate microsculptme, latter ( if present)
c. isodiametric on front, transverse on
pronotum, more transverse on elytra. Head:
eyes moderately prominent (abruptly so
in distorta), with 2 seta-bearing pimctures
over each eye except anterior puncture
absent in distorta and reduced in aculeata
group to a small impressed puncture \\'ith-
out seta; clypeus c. truncate, 1-setose each
side; labrum rather long, subtruncate or
slightly arcuate at apex except slightK
emarginate in microdera, 6-setose; mentum
without tooth, at most slighth' arcuatcK
prominent at middle; ligula and maxillae
fused, together broadly rounded, \\\ih usu-
ally 2 principal setae slighth' bc>fore apex
and additional smaller setae at apex ( setae
often difficult to distinguish); palpi rath(>r
short, but apical segments not widened; an-
tennae moderate, pubescent from 5th seg-
ments, sometimes a little pubescence on
apex of 4th. Prolhorax: .setac> at basal
angles present, median-lateral setae present
or absent; base not margined at middle,
apex usualK' with fine marginal line entire;
discal impressions usually present, some-
times almost obsolete. Elytra varying in
form and in pi-esence or absence of apical
spines; striation entire, varying in depth
and in pnnctalion; .'^rd inlcrxals nsnall\ w itli
2 niinutt> punctures without setae placed
irregularly in posterior half of elytral length
(see Notes. b(4ow). Inner wings bill. Legs
The Carabid Beetles of New Guinea • Darlington 125
slender; 4th tarsal segments weakly emar- elytra too. The species of this group differ
ginate; 5th segments with few, weak ac- among themselves in color, elytral micro-
cessory setae; claws with c. 3 to 5 teeth, sculpture, and form of elytral striae. This
Secondary sexual characters: 6 front tarsi group is thus far known only from the
very little dilated, 3 segments with paired eastern half of New Guinea, and the species
squamae at least near apex; S middle tarsi are partly allopatric: divisa and hiion seem
without squamae; c^ middle tibiae not ex- not to occur together, and neither do
cised; S with 1, 9 1 or 2 setae each side castanea and polita.
last ventral segment. Although the New Guinean Dolichoctis
Type species. Dolichoctis .striata Schmidt- are rather diverse, they are less so than
Goebel (below). the Oriental members of the genus. Some
Generic distribution. Numerous from Oriental groups, including Menartis (a
SE. Asia across the islands to New Guinea, group of small convex species ) , are not
a few species reaching New Britain, New represented in New Guinea at all. The
Ireland, the Solomons, and northern general pattern of distribution of the genus
Australia. suggests that 3 or 4 stocks have reached
Notes. The 13 New Guinean species of New Guinea at different times, probably
Dolichoctis can be arranged in 5 groups. D. all from the direction of tropical Asia, and
striata and microdera represent separate that 1 or 2 of the older stocks have radiated
species groups which arc widely distributed moderately on the island,
outside New Guinea and which have prob- The 2 minute impressed punctures, with-
ably reached New Guinea comparatively out setae, on each 3rd elytral interval pos-
recently from the west. D. distorta is unique teriorly are present in most Dolichoctis but
and forms a group of its own; it may be may be absent in distorta (in which these
derived from either of the following species punctures, if present, are lost in the general
groups or from a common ancestor. Six punctation of the intervals) and are dif-
of the remaining New Guinean species form ficult to see and perhaps sometimes absent
what I call the aculeafa group, character- in the polita group. These minute punctures
ized by anterior supraocular seta-bearing are best seen in carefully cleaned speci-
punctures reduced to small impressed mens under diffused light. Wlien I have
points without setae, median-lateral pro- been able to see them clearly in the first 1
notal setae lost, elytra dentate or spined, or 2 specimens of a series, I have credited
and reticulate microsculpture present on that species wath possessing them, without
entire upper surface. Excepting dentata, attempting to clean and examine whole
which is satisfactorily distinct, the species series.
of this group are very similar to each other Although my field notes do not distin-
and difficult to define because of occur- guish most species of this genus, I know
rence of intermediates. They are sympatric that most of them (except rriicrodera) are
— all 6 species occur at Dobodura — and arboreal, living in understory foHage in
do not seem to be differentiating geographi- rain forest. They are usually collected by
cally. This group is represented also out- sweeping or beating. They do not often
side New Guinea. Finally, 4 species form fly to light, which suggests that they may
what I am naming the polita group, which be largely diurnal. However, one species,
is like the aculeata group in form and in distorta, is apparently known only from
spined elytra, but anterior supraocular seta- light-collected specimens. It may be noc-
bearing punctures are present, and reticu- turnal and may occupy a habitat that col-
late microsculpture is absent on head and lectors do not often reach, perhaps tree-
pronotum and in some cases absent on tops in rain forest.
126 BitUetin Museum of Comparative Zoology, Vol 137, No. 1
Key to Species of Dolichoctis of New Guinea
1. EKtia obliquely truncate at apex, not
spined or denticulate; elytra usually
spotted 2
- Elytra spined or acutely denticulate at
apex; not spotted, except sutural area some-
times red 3
2. Prothorax wide and widely margined (p.
126 ) striata
- Prothorax narrow, narrowly margined (p.
127 ) inicrodera
3. Head distorted, eyes small but abniptly
prominent, front swollen on each side;
prothorax semicircular, more than 2X wide
as long ( p. 127 ) distorta
- Head normal; prothorax less than 2x wide
as long _^ 4
4. Head and pronotum (and elytra) with
reticulate microsculpture; anterior seta-
bearing punctures over eyes reduced to
minute punctures without setae {acidcata
group ) 5
- Head and pronotum without reticulate
microsculpture ( elytra with or without it ) ;
anterior ( as well as posterior ) seta-bearing
punctures present over eyes ( polita
group ) 10
5. Form broader, more Agomim-MVe (Fig.
82 ) ; prothorax relatively smaller and nar-
rower; elytra dentate at apex (p. 128) -._.
dentata
- Form more oval or fusiform; prothorax
usually relatively larger and wider; elytra
spined at apex 6
6. Elytral striae very lightly impressed, 7th
striae reduced to very fine superficial
lines; elytral spines usually very long (but
varial)le) (p. 128) .v/j/'/io.sr/
- Elytral striae including 7th well impressed;
elytral spines usually shorter 7
7. Suture or sutural area red; form usually
relatively narrow; length 4.6-5.6 mm. (p.
129) sutundis
- Suture not red; form variable; size often
larger - __ _ 8
8. Sides of prothorax ± strongly sinuate;
elytral striae moderately impressed (p.
129) .._^ andcala
- Sides of prothorax not or only .slightly
simiale; elytral striae often ilceper 9
9. i'rothorax wider (width/length 1.78 and
1.80), with sides more rnuiidcd and wiUi
wider margins (p. 130) siihrottiiKia
- Prothorax narrower (width length 1.61 and
1.69), with sides less rounded and with
narrower niargins (p. 130) sidxpiddrdtd
10. Strikingl)- bicolorcd, head and prothorax
red, elytra black or piceous; often larger
(5.8-7.4 mm) II
- Not or at most \aguely bicolored; often
smaller ( 5.4-6.5 mm ) ._„ 12
11. Elytra with grooved striae and distinct
reticulate microsculpture (p. 131) .._. divi.sci
- Elytra with striae formed by rows of small
punctures and elytral disc without reticu-
late microsculpture (p. 131) huon
12. Elytra with reticulate microsculpture (p.
131 ) castanea
- Elytral disc without reticulate microsculp-
ture (p. 132) polita
Dolichoctis striata Schmidt-Goebel
Schmidt-CIoebel 1846, Faunula Coleop. Birmaniae,
p. 62.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1384 (see for synon\'my and additional refer-
ences ) .
Louwerens 1958, Treubia 24, p. 258.
Jedlicka 1963, Ent. Abhandlungen 28, p. 357.
Description ( for recognition only ) . With
characters of genus; form broad, with elytra
unarmed; black or piceous, elytra usually
with red spots; 2 setae over each eye; pro-
thorax with 2 setae each side (see Notes,
below); length c. 4—4.5 mm.
Ttjpe(s). From Burma, in Prague Mus.
(not seen).
Occurrence in Netv Guinea. Common
probably throughout New Guinea: 200
specimens (by count), including examples
from Normanbv, Woodlark, Biak, and \Vai-
geu Is.; most at low altitudes (including
Dobodura), but a few at 1050, 1100, and
1200 m at Wau.
Notes. The recorded range of striata is
from SE. Asia (including Ceylon and
Japan) to the Philippines, Ne^ (ruinea.
and Xorth Queensland, Australia, ;uid I
have specimens also from New Britain and
New Ireland. \Miether populations from
all these places are in hut conspecific is
a (]uestion for future study.
Most individuals from New CJuinea are
either 4-spottcd (each clvtron with a pale
spot near bas(> and another near apex), 2-
spottcd (with oiiK the subapical spots), or
intermediate (with consiiicuons subapical
and fainter subb;isal spots — note that the
subbasal elytral spots \ ;ny in distinctness
more than in si/(0. The siiiule iiidiv idujil
The Carabid Beetles of New Guinea
Darlington
127
seen from Woodlark Is. is the only un-
spotted one in the New Gninean series.
However, 4-spotted, 2-spotted, and un-
spotted individuals are said to occur else-
where in the species' range (Louwerens
1958).
Of the 200 New Guinean individuals, all
that are in condition to examine have 2
seta-bearing (or formerly seta-bearing)
punctures over each eye, and all have both
a basal and a median-lateral seta (or
puncture) in each prothoracic margin ex-
cept that 2 specimens from Nabire, West
N. G. (Bishop Mus. ), lack the median-
lateral seta and puncture on one side.
These specimens have the pronotum slightly
unsymmetric: angulate on the side with
median seta, evenly arcuate on the side
without seta. Numerous other specimens
from the same locality have the median
seta and puncture present on both sides.
This species lives in understory foliage
in rain forest.
Dolichoctis microdera Andrewes
Andrewes 1930, Ann. Mag. Nat. Hist. (10) 6, p.
665.
1931, Zoologische Mededelingeii 14, p. 63.
Louwerens 1956, Treubia 23, p. 226 ( Moluccas ) .
1964, Ent. Tidskrift 85, p. 184 (Borneo).
Description ( for recognition only ) . With
characters of genus; form (Fig. 80) rela-
tively slender, with narrow, narrowly mar-
gined prothorax; black or piceous, each
elytron with 2 pale spots; 2 setae over each
eye; prothorax with basal but not median-
lateral setae; length c. 4.5-5 mm.
Type. From Sumatra; in Andrewes
Coll., British Mus. (seen).
Occurrence in New Guinea. Papua: 6,
Dobodura, Mar.-July 1944 (Darlington).
N-E. N. G.: 1, Nadzab, July 1944 (Dar-
lington); 1, Torricelli Mts., Siaute, sea
level, Nov. 9-17, 1958 (W. W. Brandt,
Bishop Mus.).
Notes. Comparison (made in 1948)
shows that New Guinean specimens differ
slightly from the Sumatran type, but the
latter is unique. More material from more
localities is needed to show whether the
differences are individual or geographic.
The known range of the species now in-
cludes Sumatra, Borneo, the Moluccas,
and New Guinea.
My New Guinean specimens were (I
think) taken among dead leaves on wet
ground, a unique habitat for members of
this eenus in New Guinea.
Dolichocfis distorta n. sp.
Description. With characters of genus;
form as in Figure 81; irregularly reddish
piceous, appendages irregularly brown;
shining, dorsal reticulate microsculpture
lacking but most of surface irregularly,
rather finely punctate. Head 0.64 and 0.65
width prothorax; eyes abnormally small but
abruptly prominent, with a channel over
each eye running diagonally forward; pos-
terior seta-bearing puncture high above
each eye, anterior puncture absent; front
strongly swollen each side of median longi-
tudinal channel, each swollen area im-
pressed near middle; sides of head behind
eyes longitudinally multisulcate. Frothorax
very wide; width/length 2.13 and 2.18;
base/apex 1.43 and 1.40 (base measured
across seta-bearing punctures); sides very
broadly rounded into base, with posterior
angles not defined; margins very widely
depressed, slightly reflexed; posterior-lateral
setae present, median-lateral setae absent;
base and apex not margined; disc with
median line and posterior and anterior trans-
verse impressions. Elytra: width elytra/
prothorax 1.16 and 1.17; outer-apical angles
distinct, c. right or minutely acute; apices
each with short spine c. opposite ends 2nd
intervals; striae impressed, not punctate;
intervals punctate, 3rd with usual 2 small
punctures doubtfully distinguishable behind
middle. Secondary sexual characters as for
genus; $ with 1, 9 2 setae each side last
ventral segment. Measurements: length
5.7-6.5; width 2.5-2.7 mm.
128 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Types. Holotype 9 (M.C.Z., Type No.
31,424) from Dobodura, Papua, Mar.-July
1944 (Darlington); 3 paratypes, Kokoda,
Papua, Mar. 28-29, 1956 (Gressitt), taken
in light trap; and 1 paratype, same locality,
1200 ft., June 1933 (Cheesman).
Measured specimens. A i paratype from
Kokoda ( British Mus. ) and the 9 holotype.
Notes. In spite of its unique modifica-
tions, this species is clearly a Dulichoctis.
All known specimens were probably taken
at light: Gressitt's are so labeled; Miss
Cheesman's specimen has the scales of
Lepidoptera stuck to it; and mine was
taken on a lighted window.
Dolichocfis acuieafa group
Dolichoctis aculeata Chaudoir and its
immediate relatives form a well defined
group with the following characters in addi-
tion to characters of the genus: form usu-
ally suboval or fusiform ( but broad
AiS,onum-\\Ve in dentata); entire upper sur-
face microreticulate; posterior seta-bearing
punctures over eyes present, anterior pimc-
tures reduced to minute points without
setae; prothoracic margins with seta-bearing
punctures at basal angles, without median-
lateral punctures; elytra with outer-apical
angles well defined (except in dentata)
and apices acutely dentate or spined c.
opposite ends of 1st striae or 2nd intervals;
last ventral segment with 1 seta each side
in both sexes.
Besides aculeata itself (as I identify it),
the lollowing 4 closely related new species
occur in New Ciuinea: sj)in()sa, suturalis,
suhrotiinda^ and subijuadrata. These 5
species (including (/rj//rY//r/ ) apparenlK iii-
tergrade to some extent, and their status
is tliereh)re doubtlul. D. dentata is more
distinct. The species of this group arc all
sympatric in New Guinea.
Although most species ot the acideata
group are New Guinean, the group is repre-
sented west at least to ('elebes (by typical
aculeata), on New Britain, New Ireland,
and the Solomons, and in North Queens-
land, Australia. The group is apparenth'
not represented in the Philippines.
Dolichocfis dentafo n. sp.
Description. With characters of genus
and of aculeata group; form as in Figure
82; broad-Ai;o;Hn?!-like with relati\ely small
prothorax; brownish black, margins and
legs paler brown, antennae and mouthparts
testaceous. Head 0.82 and 0.83 width pro-
thorax. Protliora.x rather small, quadrate-
subcordate; width length 1.58 and 1.60;
base apex 1.23 and 1.21; side margins
moderateK' wide and reflexed. Elytra:
width elytra prothorax 1.71 and 1.73; outer-
apical angles rounded, apices acutely den-
tate; striae moderately impressed, impunc-
tate. Measurements: length 6.5-7.0; width
2.8-3.1 mm.
Types. Holotype 6 (M.C.Z., Type No.
31,425) and 21 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
6 additional paratypes from Oro Bav
(near Dobodura), "Dec. 1943-Jan. 1944
( Darlington).
Additional material. N-E. N. G.: 1, Sur-
prise Ck., Nh)robe Dist., Oct. 7 (Stevens,
M.C.Z.); 1, Simbang, Huon Gulf, 1898
(Biro); 2, Torricelli Alts., Wantipi Village,
Nov. 30-Dec. 8, 1958 (W. W. Brandt,
Bishop Mus.). West N. G.: 1. Ilollandia,
Jan. 1945 (Malkin, U.S.N.M.); 1, llijob, 25
m, Sept. 10, 1956 ( Neth. New Guinea Exp.,
Leiden Mus.); 1, Wasian, \\)gelkop, Sept.
1939 (Wind, M.C.Z.).
Measured specimens. The i holotvpe and
1 9 paratype from Dobodura.
Notes. I know no \c'r\' close relati\es of
this species. It is, ol course, placed in rela-
tion to others in tlu' iireceding Key lo
Species.
Dolichocfis spinosa n. sp.
Description. With characters of genus
and ol (icutcala gioup; black. ai')pendages
dark brown; niicrosculpturc more transverse
than usual on pronotum and elytra, latter
sliuhtK iiidcsceut. Head 0.74 and 0.76
The Carabid Beetles of New Guinea • Darlington
129
width prothorax. Prothurax transverse-cor-
date; width/length 1.67 and 1.64; base/apex
1.26 and 1.25; sides depressed but margins
not well defined. Elytra: width elytra
prothorax 1.61 and 1.72; outer-apical angles
well defined but obtuse, apices with long
(but variable) slightly dehiscent spines;
striae, especially outer ones, very lightly
impressed. Measurements (types only);
length c. 6.0-7.5 (including spines); width
2.6-3.2 mm.
Types. Holotype $ (M.C.Z., Type No.
31,426) and 9 paratypes from Dobodura,
Papua, Mar.-July 1944 (DarHngton). Ad-
ditional paratypes as follows, all from
Papua: 3, Kokoda, 1200 ft., June, Aug.,
Sept. 1933 (Cheesman); 1, Palmer R. at
Black R., June 7-14, 1936 (Archbold Exp.,
A.M.N.H.).
Additional material. Eighteen from vari-
ous localities in all 3 political divisions of
New Guinea; some at low altitudes, some
at 1200 (at \\'au), 1300, and 2000 m.
Measured speeimens. The i holotype and
1 9 paratype from Dobodura.
Notes. Typical specimens of this new
species are easily recognized by very light
elytral striation and very long elytral spines,
but some individuals listed under Additional
material have shorter spines and vary
toward one or another of the following
species.
Dolichoctis sufuralis n. sp.
Deseription. With characters of genus
and of aeuleata group; form c. as in aculeata
(following species) but more slender,
smaller; castaneous with suture or sutural
area reddish, appendages brownish testa-
ceous. Head 0.69 and 0.74 width prothorax.
Prothorax: width length 1.79 and 1.74;
base/apex 1.40 and 1.39; sides broadly
rounded anteriorly, slightly sinuate before
somewhat obtuse, blunted posterior angles;
margins rather widely depressed especially
posteriorly. Elytra rather narrow; width
elytra prothorax 1.44 and 1.52; outer-apical
angles well defined but obtuse, apices
with short spines; striae moderately im-
pressed, not distinctly punctate. Measure-
ments: length c. 4.6-5.6; width e. 2.0-2.4
mm.
Types. Holotype i (M.C.Z., Type No.
31,427) and 23 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
6 paratypes from Oro Bay (near Dobodura),
Dec. 1943-Jan. 1944 (Darlington).
Additional material. Thirty-one (some
doubtfully identified), from numerous lo-
calities including all 3 political divisions of
New Guinea and Normanby Is.; most from
low altitudes ( usually below 500 m ) but 1
from Finisterre Rge. at 1200, and 1, Upper
Jimmi Vy. at 1300 m.
Measured specimens. The i holotype and
1 9 paratype from Dobodura.
Notes. In the aculeata group of New
Guinean Dolichoctis, only relatively small,
slender individuals have reddish sutures.
This correlation of size, form, and color
suggests that suturalis is a real species, al-
though the distinguishing characters are
slight.
Dolichoctis aculeata Chaudoir
Chaucloir 1869, Ann. Soc. Ent. Belgium 12, p.
251.
Andrewes 1930, Treubia 7, Supplement, p. 336.
Louwerens 1956, Treubia 23, p. 226 (Moluccas).
Description. With characters of genus
and of aculeata group. Head 0.72 and 0.67
width prothorax. Prothorax: width/length
1.80 and 1.75; base/apex 1.30 and 1.42
( proportions notably variable ) ; sides rather
broadly depressed. Elytra: width elytra/
prothorax 1.50 and 1.49; outer-apical angles
well defined, almost right ( slightly obtuse ) ,
apices spined; striae moderately impressed.
Measurements: length c. 5.0-6.5; width c.
2.2-3.0 mm.
Types. From Celel>es, collected by
W^allace; type now in Oberthiir Coll., Paris
Mus. (not seen).
Occurrence in New Guinea. Common
probably throughout the island: 120 speci-
mens (including 65 from Dobodura and
Oro Bay), from all 3 political divisions of
130 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
New Guinea and Rossel and Woodlark Is.;
most at low altitudes but reaching 1200 to
1400 m at some localities.
Measured specimens. A pair { i 9 ) from
Dobodura.
Notes. My identification of this species
is based on comparison with specimens
identified by Andre wes in his collection.
I collected speciinens that I refer to this
species at Iron Range and Rocky R. in
the mid-peninsular rain forest of Cape
York, Australia, in 1958.
Dolichoctis subrotunda n. sp.
Description. With characters of genus
and of aculeata group; similar to oculeata,
differing principally in form of prothorax
(Fig. 83), with broadly rounded sides not
or scarcely sinuate posteriorly. Head 0.65
and 0.66 width prothorax. Prothorax:
width/length 1.78 and 1.80; base/apex 1.42
and 1.33; sides flattened but not strongly re-
flexed. Elytra: width elytra/prothorax 1.49
and 1.49; apical spines moderate; striae mod-
erately impressed. Measurements (types):
length c. 5.0-6.5; width c. 2.3-2.9 mm.
Types. Holotype 6 (M.C.Z., Type No.
31,428) and 26 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
2 additional paratypes from Oro Bay ( near
Dobodura), Dec. 1943-Jan. 1944 (Dar-
lington ) .
Additional material. Sixty (some doubt-
fully identified) from numerous localities
in all 3 political divisions of New Guinea
and Normanby and Woodlark Is.; most at
low altitudes, but recorded above 1000 m
at several localities including Wau and at
2500 m in the Chimbu area.
Measured specimens. The i holotype and
1 9 paratype from Dobodura.
Notes. Individuals assigned to this spe-
cies vary considerably in si/e, depth of
elvtral striae, etc. A single specimen from
Waigeu Is. (Camp Nok, 2500 ft. (c. 770
m), Apr. 1938, Cheesman) differs from all
specimens from the mainland of New
Cuinea in having a poorly defined sub-
apical red spot on (>ach elytron near suture.
Doiichocfis subquadrato n. sp.
Description. With characters of genus
and of aculeata group; form similar to
aculeata except prothorax smaller and sub-
quadrate (Fig. 84). Head 0.75 and 0.70
width prothorax. Prothorax: width length
1.61 and 1.69; base apex 1.36 and 1.44;
sides usually slightly sinuate near base;
margins scarcely depressed anteriorly, more
broadly so posteriorly. Elytra: width elytra/
prothorax 1.60 and 1.54; apices with mod-
erate spines; striae usually deeply impressed
(deeper than in aculeata); 7th intervals
slightly elevated at base. Measurements
(types only): length 5.7-6.7; width 2.5-2.8
mm.
Types. Holotype S (M.C.Z., Type No.
31,429) and 3 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); 2
additional paratypes from Oro Bay (near
Dobodura), Dec. 1943-Jan. 1944 (Darling-
ton); and 4 paratypes from Milne Bay,
Papua, Dec. 1943 (Darlington).
Additional material. Four from widely
scattered localities in New Guinea; and 3,
Aru Is. ( British Mus. ) . Also several doubt-
fully identified from Wau.
Measured specimens. The i holotype and
1 9 paratype from Dobodura.
Notes. This species seems clearly distinct
from the preceding one (subrotunda): the
proportions of head prothorax and pro-
thoracic width length reflect the relativeK'
large, wide prothorax of subrotunda and the
smaller and narrower one of subquadrata.
However, aculeata is intermediate. These
3 species together form a bewildering, vari-
able complex that includes many individ-
uals wliieh I cannot ]ilace satislactoriK'.
Dolichoctis polifa group
The following four species lorni a group,
apparently confined to Ni'w Cuinea, char-
acterized as follows: form c. as in aculeata
group l)ul more skMider; microsculpture
absent on head and pronotum, prc\scMit or
absent on elytra; 2 setae over each eye;
pronotum with setae* (or inmetures) at basal
The Carabid Beetles of New Guinea • Darlington 131
angles but median-lateral setae absent;
elytral apices dentate or spined; 1 seta
each side last ventral segment in both sexes.
Dolichoctis divisa n. sp.
Description. With characters of genus
and of polita group; form as in Figure 85;
head and prothorax red, elytra black and
slightly silky, legs dark, antennae pale;
elytra with transverse microsculpture. Head
0.63 and 0.67 width prothorax. Prothorax:
width/length 1.52 and 1.57; base/apex 1.41
and 1.38; sides slightly sinuate before c.
right (narrowly rounded) basal angles;
sides of disc slightly depressed. Elytra:
width elytra prothorax 1.35 and 1.37; outer-
apical angles c. right, apices with moderate
spines; striae well impressed, not punctate.
Measurements: length 6.6-7.4; width 2.6-
2.9 mm.
Types. Holotype $ (M.C.Z., Type No.
31,430) and 1 9 paratype from Dobodura,
Papua, Mar.-July 1944 (Darlington), and
additional paratypes as follows. Papua: 2,
Bisianumu, E. of Port Moresby, 500 m,
Sept. 23, 1955 (Gressitt); 1, Kokoda, 1200
ft. (366 m), Aug. 1933 (Cheesman); 1,
Milne Bay, Dec. 1943 (Darlington); 1,
Brown River, 20 km N. of Port Moresby,
Apr. 29, 1960 (C. W. O'Brien, Bishop
Mus.); 1, Popondetta, 60 m, Oct. 18, 1963
(Shanahan, Bishop Mus.); 4, Mt. Laming-
ton, 1300-1500 ft. (c. 400-460 m) (C. T.
McNamara, S. Austrahan Mus.).
Measured specimens. The pair ( i 9 )
from Dobodura.
Notes. This strikingly bicolored species
is apparently confined to a small part of
eastern New Guinea.
Dolichoctis liuon n. sp.
Description. With characters of genus
and of polita group; head and prothorax
red, elytra usually darker ( castaneous ) ,
sometimes scarcely darker; legs and anten-
nae dark; whole upper surface without
reticulate microsculpture. Head 0.65 and
0.66 width prothorax. Frothorax: width
length 1.66 and 1.67; base/apex 1.40 and
1.41; sides slightly sinuate before slightly
obtuse (nearly right) basal angles; sides of
disc slightly depressed. Elytra: width
elytra /prothorax 1.39 and 1.40; outer-apical
angles c. right, apices short-spined or
acutely toothed; striae scarcely impressed,
formed by rows of small punctures. Mea-
surements: length 5.8-6.9; width 2.3-2.8
mm.
Types. Holotype i (Bishop Mus.) and
6 paratypes from Pindiu, Huon Pen., N-E.
N. G., 500-600, 750-850, 870-1300 m, Apr.
19, 20, 21, 21-22, 1963 (Sedlacek). Addi-
tional paratypes as follows, all from north-
ern part of N-E. N. G.: 4, Finschhafen, 10,
80 m, Apr. 12, 16, 1963 (Sedlacek); 1, Lae,
10 m. May 12, 1966 (Gressitt); 1, Busu R.,
E. of Lae, 100 m, Sept. 15, 1955 (Gressitt);
1, Torricelli Mts., Mobitei, 750 m. Mar. 5-
15, 1959 (W. W. Brandt, Bishop Mus.).
Some paratypes now in M.C.Z. (Type No.
31,431).
Measured specimens. The S holotype and
1 9 paratype from Finschhafen.
Notes. The dark antennae, punctate
elytral striae, and absence of elytral micro-
sculpture clearly distinguish this species
from divisa (above). These 2 species are
apparently allopatric, confined to different
small areas of eastern New Guinea.
Dolichoctis costonea n. sp.
Description. With characters of genus
and of polita group; reddish castaneous,
prothorax sometimes slightly paler, append-
ages reddish brown; elytra with transverse
reticulate microsculpture. Head 0.66 and
0.68 width prothorax. Prothorax: width/
length 1.62 and 1.63; base/apex 1.32 and
1.31; sides broadly rounded, not or slightly
sinuate before usually obtuse basal angles;
disc slightly depressed at sides. Elytra:
width elytra prothorax 1.33 and 1.45; outer-
apical angles distinct but obtuse and slightly
blunted, apices short-spined or acutely
dentate; striae well impressed, not distinctly
punctate. Measurements: length 5.4-6.5;
width 2.1-2.5 mm.
Types. Holotype 6 (M.C.Z., Type No.
132
Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
31.432) and 2 paiatypes from Dobodiira,
Papua, Mar.-July 1944 (Darlington); and
additional paratvpes as follows. Papua:
3, Kokoda, 1200 ft., July, Aug. 1933
(Cheesman); 1, Kokoda-Pitoki, 450 m. Mar.
24, 1956 (Gressitt); 2, Mt. Lamington, 1300-
1500 ft. (c. 400-460 m) (C. T. McNamara,
S. Australian Mus.); 1, Brown R., May 25,
1956 (E. J. Ford, Jr., Bishop Mus.). N-E.
N. G.: 15, Pindiu, Huon Pen., 500-600,
860, 870-1300 m, Apr. 19-22, 1963 (Sed-
lacek); 1, Busu R. E. of Lae, 100 m, Sept.
14, 1955 (Gressitt); 1, Bubia, Markham Vy.,
50 m, Sept. 20, 1955 (Gressitt); 1, Sattel-
berg, Huon Gulf, 1899 (Biro); 1, Madang
(Friedrich-Wilh.-hafen), 1901 (Biro).
MecLsurcd specimens. The S holotype and
1 9 paratype from Dobodura.
Notes. This speeies is sympatrie with the
two preceding ones, but perhaps allopatric
with the following (polita).
Dolichoctis polifa n. sp.
Description. With characters of genus
and of polita group; reddish castaneous,
prothorax sometimes slightly paler, append-
ages not or slightly paler; entire upper
surface without reticulate microsculpture.
Head 0.66 and 0.66 width prothorax. Vro-
thorax: width/length 1.64 and 1.78; base/
apex 1.38 and 1.36; sides usually not sinuate
before obtuse, sometimes blunted posterior
angles; sides of disc moderately depressed.
Elytra: width elytra/ prothorax 1.33 and
1.36; outer-apical angles distinct but obtuse
and slightly blunted; apices short-spined or
acutely toothed; striae well impressed, im-
punctate. Measurements: length 5.5-6.5;
width 2.1-2.6 mm.
Tijpes. Holotype i (Bishop Mus.) and
33 paratypes (some in M.G.Z., Tvp(> No.
31.433) from Wan, Morobe Dist, N-E.
N. G., altitudes from 1050 to 1500 m, dates
in Jan., Feb., Mar., June, July, Sept., Oct.,
Dec, 1961-1964 (Scdlacek);' and 2 para-
types, Upp(>r Watut R., 24 km W. oi
Bulolo, N-E. N. G., 760 m. Mar. 5-6, 1963
(Scdlacek).
Measured specimens. The i holotype and
1 9 paratype from Wau.
Notes. This may be a geographic form
(confined to the Morobe area) of Dolicho-
ctis castanea (aboye), distinguished pri-
marily by absence of clytral microsculp-
ture, but I do not care to recognize sub-
species in this genus until relationships and
geographic patterns are better understood.
Genus STRICKLANDIA Macleay
Macleay 1886, Proc. Linnean Soc. New South
Wales (2) 1, p. 138.
Diagnosis. Form ( Fig. 86 ) characteristic,
large, yery broad, depressed; prothorax
strongly cordate, with numerous extra lat-
eral setae anteriorly; elytra yery wide, each
2-spined. See also Key to Genera of Lebiini
of New Guinea.
Description. Form broad, depressed;
black, moderately shining; not obyiously
pubescent but pronotum and sometimes
other parts of upper surface yery incon-
spicuously sparsely setulose; reticulate mi-
crosculpture absent or indistinct on head
and disc of pronotum ( but pronotal disc
transyersely rugulose), yisible but meshes
imperfect and irregular on elytra. Head:
eyes rather small but prominent; 2 setae
oyer each eye; front flattened, weakl\- de-
pressed; clypeus subtruncate \\ ith rounded
angles, 1-setose each side; labrum long,
apex subtruncate or slighth broadly emar-
ginate, 6-setose; mandibles moderately long,
not strongly arcuate, longitudinalb' striate
aboye at middle oF length; antennae slender,
pubescent from middle 4th segments;
mentum subtruncate in sinus, sli^htK' lobed
or with short blunt tooth; ligula wide at
apex, with 2 or 3 large and 2 or more smaller
setae, and paraglossae attached to and
slightly longer than ligula, without setae;
palpi slendei-, apical segments labial palpi
with longitudinal row of numcMous setae
ab()\-e. Prothorax cordate; base not lobed
but irregularly obliciucK rounded to basal
angles; sides angulate or scalloped at mid-
dle. r(41exed, with priiuipal setae at basal
The Carabid Beetles of New Guinea • Darlington 133
angles and near middle and several addi-
tional often smaller setae anteriorly; median
longitudinal and basal and apical transverse
areas impressed; base and apex not dis-
tinctly margined. Elytra very wide, widest
near base; humeri rounded but very promi-
nent; outer-apical and sutural angles both
spined; margins finely serrate and setulose;
striae entire, punctate; 3rd intervals with 1
or 2 seta-bearing punctures behind middle.
Inner winfis full. Lefis slender; 4th seg-
ments middle and hind tarsi narrow, scarcely
emarginate; 5th segments with accessory
setae minute (vestigial?); claws with c. 4
small teeth, in basal half of claw length.
Secondary sexual characters: 6 front tarsi
scarcely dilated but 3 segments with small
2-seriate squamae; S middle tarsi without
squamae; i middle tibiae not excised; S
with 1, ? 2 setae each side last ventral seg-
ment.
Type species. Stricklandia pericalloides
Macleay.
Generic distribution. New Guinea (2 or
more species ) ; Moluccas ( 1 species, from
Batjan Is., Louwerens 1956, Treubia 23, p.
241 ) ; New Britain ( 1 probably unde-
scribed species); and North Queensland,
Australia (1 species). The members of this
genus that I have collected live on tree
trunks and fallen logs in rain forest.
Notes. I do not know the relationships or
geographic origin of this primarily New
Guinean genus.
Key to Species of Stricklandia of New Guinea
1. Prothorax narrower (usually c. 1.5 X wide
as lonfj; at middle, but sometimes wider),
with relatively narrow margins (reflexed
margins often less than V4 as wide as distance
from midline to lateral trough, but some-
times wider) (p. 133) pericalloides
- Prothorax very wide (c. 1.9x wide as long
at middle), with very wide margins (re-
flexed margins more than V2 as wide as
distance from midline to lateral trough)
(p. 133) hta
Stricklandia pericalloides Macleay
Macleay 1886, Proc. Linnean Soc. New South
Wales (2) 1, p. 139.
Description. See generic Diagnosis and
Description. Head 0.79 and 0.78 width
prothorax. Prothorax: width/length 1.48
and 1.56; l^ase/apex 1.18 and 1.15; reflexed
margins relatively narrow. Elytra: width
elytra/ prothorax 1.55 and 1.54. Measure-
ments (Dobodura series): length c. 11.5-
13.5 (including elytral spines); width 4.5-
5.1 mm.
Type. From Fly R., Papua; presumably
in Macleay Mus., Sydney (not seen).
Occurrence in New Guinea. Common
probably throughout New Guinea: 96
specimens (some doubtfully identified, see
following Notes), from all 3 political divi-
sions of the island; most at low altitudes
but reaching c. 1500 to 2000 m at several
localities including Wau.
Measured specimens. A pair { i 9 ) from
Dobodura, Papua.
Notes. Some individuals tentatively as-
signed to pericalloides have prothoracic
margins relatively wide (but not so wide as
the following species) and may be specifi-
cally distinct, but I do not wish to describe
them at present. Mr. Louwerens may refer
these individuals to a species he will prob-
ablv describe from New Britain.
Stricklandia lata n. sp.
Description. With characters of genus;
form as in Figure 86, extraordinarily wide;
color and surface as described for genus,
but elytral microsculpture more transverse
than in pericalloides. Head 0.64 and 0.68
width prothorax. Frothorax wide-cordate;
width length 1.89 and 1.89; base/apex 1.06
and 0.99; margins very wide {c. V2 wide as
distance from inner edge of margin to
middle line), with outer edge irregular.
Elytra: width elytra prothorax 1.29 and
1.36. Measurements: length c. 15-16; width
c. 6.5 mm.
Types. Holotype S (Leiden Mus.) from
Arabu Camp, Wissel Lakes, West N. G.,
1800 m, 1939 (H. Boschma), and additional
paratypes from Wissel Lakes as follows:
Is (M.C.Z., Type No. 31,434), Digitara,
Oct. 1938 (P. J. Eyma); 1 9 , Wagete, Tigi
134 Bulletin Museuin of Comparative Zoology, Vol. 137, No. 1
L., 1700 m, Aug. 17, 1955 (Gressitt); 3,
Enarotadi, 1850, 1850-1900, 1850-2050 m,
dates in July, Aug. 1962 (Sedlacek).
Measured specimens. The c5 holotype and
9 paratype from Wagete.
Notes. Distinguished from pericaUoides
by much wider prothorax and other dif-
ferences of proportion shown l:)y ratios in
the Descriptions.
Genus PEUOCYPAS Schmidt-Goebel
Schmidt-Goebel 1846, Faunula Coleop. Birnianiae,
p. 33.
Jeannel 1949, Coleop. Carabiques de la Rejiion
Malgache, Part 3, p. 991.
Demetrias Csiki 1932 (in part), Coleop. Cat.,
Carabidae, Haipalinae 7, p. 1386 (see for addi-
tional references ) .
Risoplulits Jedlicka 1963 (not Leach), Ent. Abhand-
lungen 28, p. 401.
Diagnosis. In New Guinea, the form
(Fig. 87), small size (under 5 mm), and
long-lobed 4th tarsal segments are diag-
nostic.
Description. None required here. See
detailed description of following new
species.
Type species. P. sutiiraJis Schmidt-Goebel,
of Burma, etc.
Generic distribution. Southern and east-
ern Asia to the Philippines and New
Guinea ( not Australia ) ; Africa, Mada-
gascar.
Notes. Generic distinctions and applica-
tions of generic names have been confused
in the group of genera to which this genus
belongs. In my present use of Teliocijpas
I am following Jeannel, although I do not
like his multiplication of higher categories.
Peliocypas papua n. sp.
Description. Form as in Figure 87;
brown, appendages slightly paler; not
pubescent; rather shining, reticulate micro-
sculpture lightly impressed and irregular, c.
isodiametric on head and elytra, slightly
transverse on disc of pronotum. Head 0.97
and 0.97 width prothorax (measured at
middle); eyes moderately prominent; 2
setae over each eye; front slightly impressed
at sides between eyes and at sides an-
teriorly; frontal suture indicated but not
impressed; clypeus subtruncate, 1 -setose
each side; labrum transverse, subtruncate
with rounded angles, 6-setose; mentum \\'ith
strong triangular tooth; ligula rounded-
subtruncate, apparently 2-setose, with para-
glossae of c. same length, apparently at-
tached, narrowly rounded, without setae.
Prothorax subquadrate, widest at base, with
anterior angles rounded; width (at middle)/
length 1.15 and 1.24; base apex 1.38 and
1.47; base and apex subtruncate (base
slightly sinuate), not margined; side mar-
gins narrow, broader basally and reflexed
and running into deep baso-lateral impres-
sions, each margin with setae at basal angle
and
V.
from apex; disc with usual me
dian line and transverse impressions and
lightlytransverselystrigulo.se. Elytra: width
elytra /prothorax 2.04 and 2.14; humeri
broadly rounded but not much narrowed;
apices obliquely sinuate-truncate, outer
angles rounded, sutural angles blunted;
striae entire but light, not punctate; 3rd
intervals with 2 conspicuous dorsal punc-
tures c. Vi from base and ^4 from apex.
Inner nings full. Legs slender; 4th seg-
ments of middle and hind tarsi with long
lolies; 5th segments with accessory setae;
claws each with 1 long tooth outside and
2 smaller teeth inside middle of length.
Secondary sexual characters: i front tarsi
with squamae (if present) not clearly dif-
ferentiated; last ventral segment with apex
deeply notched in i , entire in 9 ; c^ w ith
1, 9 2 setae each side apex last ventral
seguKMit. Measurements: length c. 4.0-
4.5; width c. 1.8-1.9 mm.
Types. Holotype S (Hungarian National
Mus.) and 5 paratypes (2 in M.C.Z., Tvpe
No. 31,435) all from Madang ("Friedrich-
\\'ilh.-hafen"), N-E. N. G., 1901 (Biro).
Measured specimens. The i holotype and
1 9 paratype.
Notes. This is the (easternmost species of
a genus or group of genera \'er\' w(41
The Carabid Beetles of New Guinea • Darlington 135
represented in the Orient. In Jedlicka's
(1963, pp. 401-402) key to the species of
"R/.wp/n'/us," papiia rnns to conplet 5 but
fits neither species there named, being
narrower-headed than tinicolor JedHcka
and smaller than vimmcri Jedlicka.
New Britain, New Ireland, the Solomons,
Fiji, Samoa, and New Caledonia (speci-
mens seen from all these islands). It lives
in foliage and may (I think) have been
carried eastward into the Pacific by man,
perhaps in thatching material.
Genus CELAENEPHES Schmidt-Goebel Genus SYNTOMUS Hope
Schmidt-Goebel 1846, Faunula Coleop. Binnaniae,
p. 77.
Csiki 1932, Coleop. Cat., Carabidae, Flarpalinae
7, p. 1412 (see for synonymy and additional
references ) .
Jedlicka 1963, Ent. Abhandlungen 28, p. 399.
Diap}osis-. See Key to Genera of Lebiini
of New Guinea and Description of following
species.
Description. None required here.
Type species. Celaenephes parallelus
Schmidt-Goebel ( below ) .
Generic distribution. That of the single
species.
Notes. I do not know the relationships
of this monotypic genus.
Celaenephes parallelus Schmidt-Goebel
Schmidt-Goebel 1846, Faunula Coleop. Birmaniae,
p. 77.
Van Emden 1937, Stettiner Ent. Zeitung 98, p. 35.
Andrewes 1947, Arkiv for Zool. 38A, No. 20, p. 12.
Louwerens 1956, Treubia 23, p. 225 (Moluccas).
See additional references under genus.
Description ( for recognition only ) . Form
as in Figure 88; slender, with elytral apices
simply rounded-truncate; plain black or
piceous; mentum not toothed; claws not
toothed; length c. 6.5-7.5 mm.
Type(s). From Burma; in Prague Mus.
(not seen).
Occurrence in New Guinea. Common
throughout New Guinea and on Normanby
Is.: 206 specimens, most at low altitudes
but a few up to 1550 and 1700 m; found at
Dobodura and Wau.
Notes. This easily recognized carabid
ranges at least from Ceylon, extreme NE.
India (not peninsular India, according to
Andrewes), Burma, etc. to the Philip-
pines and northern Australia, and east to
Hope 1838, Coleop. Manual 2, p. 64.
Jeanne! 1942, Faune de France, Coleop. Carabiques,
Part 2, p. 1075.
Metabletus Schmidt-Goebel 1846, Faunula Coleop.
Birmaniae, p. 38.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1413 (see for additional references, synonymy,
and list of species ) .
Jedlicka 1963, Ent. Abhandlungen 28, p. 420.
Diafinosis. Known among New Guinean
Lebiini by small size, form (Fig. 89),
mentum with emarginate tooth, and tarsal
claws with 2 or 3 minute inconspicuous
oblique teeth.
Description. None required here.
Type species. Of Syntomtis, Carabus
truncatellus Linnaeus, of Europe; of Metab-
letus, M. obscuroguttatus Schmidt-Goebel,
of Burma, etc.
Generic distribution. Temperate and
tropical Eurasia and across the islands
to North Queensland, Australia; North
Ameriea; parts of Africa.
Notes. Only one, widely distributed
species of this genus reaches New Guinea.
Syntomus quadripunctatus (Schmidt-Goebel)
Schmidt-Goebel 1846, Faunula Coleop. Birmaniae,
p. 39 {Metabletus).
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1418 (see for synonymy and additional refer-
ences ) .
Description ( for recognition only ) . With
characters of genus; form as in Figure 89;
black; upper surface dull but not pubescent;
elytra with 3rd intervals 2-punctate; length
c. 3.5 mm.
Type(s). From Burma; in Prague Mus.
(not seen).
Occurrence in New Guinea. N-E. N. G.:
4, Wau, Morobe Dist., 1250 m, dates in
Jan., Feb., Sept. 1961, 1962 (Sedlacek); 1,
136 BiiUetin Museum of Comparative ZooIos,ij, Vol. 137, No. 1
Mt. Missim (near Wau), 1050 m, Dec. 27,
1962 (Sedlaceks); 1, Mt. Missim (Stevens,
M.C.Z.); 1, Finschhafen, Apr. 1944 (E. S.
Ross, Cal. Acad.). West N. G.: 1, Eramboe,
80 km ex Merauke, Jan. 29, 1960 (T. C.
Maa, Bishop Mus.).
Notes. The known range of S. quodri-
punctotus is from SE. Asia including
Ceylon, Burma, and Japan across the
Malay Archipelago to the Philippines,
New Guinea, and the NE. corner of Aus-
tralia. Occurrence in AustraUa is based on
a single teneral 9 that I collected N. of
Mareeba, North Queensland, Feb. 1958.
Genus MICROLESTES Schmidt-Goebel
Schmidt-Goebel 1846, Faunula Coleop. Birmaniae,
p. 41.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1420 (see for additional references, synonymy,
and list of species).
Jeanncl 1942, Faune de France, Coleop. Carabiques,
Part 2, p. 1084.
Mateu 1959, Rev. fran^-aise d'Ent. 26, pp. 135 ff.
( species of tropical Asia ) .
1963, Ann. Mus. R. I'Afrique Central, Ser.
in-8°, No. 121, pp. 1-149 ( niono,ii;raph of
African species).
Jedlicka 1963, Ent. Abhandlunsen 28, p. 425.
Diagnosis. Distinguished among New
CHiinean Lebiini by small size, form (note
genae not swollen, prothorax lobed at base ),
mentum without tooth, and tarsal claws
toothed (teeth few and minute in curtatus).
DcscripiUm. None required here.
Tijpc species. Microlestes inconspicuus
Schmidt-Goebel, of Burma, etc.
Generic distribution. Warm-temperate
and tropical Africa and Evirasia and is-
lands to Australia; North America; scat-
tered records elsewhere.
Notes. Two unr(4ated speci(\s occur in
New Ciuinea, one with Oriental and the
other with apparent Australian relation-
.ships.
Kky io Spi:c:n:s ok Microlestes ok Nkw Guinea
1. Hclativcly broad (protlioracic widtli Iciiutli
1.50); elytra with 2 incomplete pale iastiac;
length 3.8 mm (p. 136) ..._ cinctu.s
- Narrower (protlioracic width /]ensj;th 1.18
and 1.28); dull black, not marked; length
not over 3.2 mm (p. 136) curtatus
Microlestes cinctus n. sp.
Description. Form as in Figure 91; black,
in part slightly brownish, elytra with 2 in-
complete transverse fasciae testaceous, ap-
pendages irregularly brownish, bases of
femora and of antennae slightly darker;
rather shining, upper surface with reticulate
microsculpture of meshes isodiametric on
head and pronotum, less regular and slightly
transverse on elytra. Head 0.79 and 0.77
width prothorax. Prothorax \\'ide-subcor-
date; width length 1.50 and 1.55; base
apex 1.12 and 1.14; sides rather narrowly
margined, each margin with setae at base
and c. % from apex; usual discal impres-
sions present but weak. Elytra: width
elytra/prothorax 1.67 and 1.56; striae lightly
impressed, minutely irregular or faintly
punctulate; intervals sparsely minutely punc-
tulate, 3rd with 1 seta-bearing puncture, on
inner edge c. Vi from apex. Legs: claws
each with c. 4 distinct, oblique teeth. Mea-
surements: length 3.6-.3.8; width 1.7 mm.
Ti/pe. Holotype 6 (Bishop Mus.) from
Feramin, N-E. N. G., 1200-1500 m, Mav
11-22, 1959 (W. W. Brandt); 1 S para-
type, Okapa (Okasa), N-E. N. G., July 8,
1965 (Hornabrook), "pine forest, leaf mold."
Notes. Of other species known to me,
this is most like M. atrifasciatus Sloane of
NE. Australia (base of Cape York Pen. to
northern New South Wales), but the color
pattern is different, the ehtra in atrifasciatus
being testaceous with a dark irregular post-
median fascia and subapical and sublateral
dark spots.
Microlestes curtatus n. sp.
Description. i'"onn as in Figure 90;
slender, \\ itli el\ tra niuch shoiter than abdo-
men; dull brow nish black, apjicMidages dark;
entire upinr surface \\ith reticulate micro-
sciilplure irregular (partK' longitudinal) on
head, slightly transverse on pronotiun and
(4ylra. Head 0.92 and 0.89 w idlh i-)rothorax.
Prothorax narrow-subcordatc; width length
The Carabid Beetles of New Guinea • Darlington 137
1.18 and 1.28; base/apex 1.08 and 1.04;
side margins very narrow, each with setae
at basal angle and c. V4 from apex; disc
with median line impressed, transverse im-
pressions scarcely indicated. Elytra very
short, narrowed anteriorly; width elytra/
prothorax 1.68 and 1.64; striae lightly in-
dicated, sometimes scarcely visible, irregu-
lar but not distinctly punctate; 3rd intervals
with 2 punctures, before middle and c. V-t
from apex. Legs: claws each with c. 2
small oblique teeth, easily overlooked. Mea-
surements: length to apex elytra 2.4-2.6,
to apex abdomen 2.8-3.2; width 1.0-1.1 mm.
Types. Holotype S (M.C.Z., Type No.
31,436) and 28 paratypes all from central
plains of Luzon, Philippine Is., Feb.-Sept.
1945 (Darlington).
Occurrence in New Guinea. West N. G. :
2, Dor(e)y (probably collected by Wallace,
British Mus.; this locality is, of course,
somewhat doubtful ) .
Measured specimens. The S holotype and
1 9 paratype from central plains of Luzon.
Notes. I have based this new species on
Philippine individuals because of doubt
about Wallace's locality "Dorey" (see Part
1 of mv work on New Guinean Carabidae,
pp. 330-331).
M. curtatus is similar to exilis Schmidt-
Goebel but has shorter elytra. This species
(curtatus) with very short elytra is not
represented in the Andrewes Collection and
was evidently not known to Mateu (1959)
or Jedlicka ( 1963 ) . It is unknown in Aus-
tralia.
Genus APRISTUS Chaudoir
Chaudoir 1846, Enumeration des Carabiques . . .
Caucase . . ., p. 42.
Csiki 1932, Coleop. Cat., Carabidae, Haipalinae 7,
p. 1432 ( see for additional references, synonymy,
and list of species ) .
Jeannel 1942, Faune de France, Coleop. Carabiques,
Part 2, p. 1083.
Jedlicka 1963, Ent. Abhandlungen 28, p. 427.
Diagnosis. Very small Lebiini, recogniz-
able (in New Guinea) by form (Fig. 92);
surface not pubescent but all or part (at
least elytra) dull and heavily microreticu-
late; genae not swollen; mentum with
entire tooth; claws not toothed.
Description. None required here.
Type species. Apristus suhaeneus Chau-
doir, of the Caucasus and Mediterranean
region.
Generic distribution. \\^arm-temperate
and tropical Eurasia and the Malay Archi-
pelago to the Philippines and New
Guinea (not Australia); part of Africa
(not Madagascar); North and Central
America, Cuba.
Notes. American species of this genus,
which are the only ones I have collected,
live on the ground, usvially on sand or
gravel near water.
Key to Species of Apristus of New Guinea
1. Color brownish bronze; entire upper surface
dull; length 3.0-3.5 mm (p. 137) ^ hiroi
- Color bluish black; front of head and middle
of pronotimi relatively shining, elytra dull;
lengtli 3.5-3.9 mm (p. 137) sedlaccki
Aprisfus biroi n. sp.
Description. With characters of genus;
form as in Figure 92; brownish bronze,
including appendages; entire upper surface
dull, heavily microreticulate. Head 0.90 and
0.90 width prothorax. Prothorax: width/
length 1.28 and 1.30; base/apex 0.91 and
0.90. Elytra: width elytra/prothorax 1.64
and 1.70. Measurements: length 3.0-3.5;
width 1.2-1.5 mm.
Types. Holotype S (Hungarian National
Mus.) and 5 paratypes (2 in M.C.Z., Type
No. 31,437) all from Madang ("Friedrich-
Wilh.-hafen"), N-E. N. G., 1901 (Biro).
Measured specimens. The $ holotype and
1 9 paratype.
Notes. Similar to A. louwerensi Andrewes
of Java, but with elytra more narrowed
anteriorly and with fainter striae.
Apristus sedlaceki n. sp.
Description. With characters of genus;
form c. as in preceding (biroi) except
sides of prothorax more rounded anteriorly
and much more strongly sinuate c. % from
138 Bulletin Museum of Comparative Zoology. Vol. 137, No. 1
base; l)liiish black, appendages dark; front
of head shining with reticulate microsculp-
tiire faint and fragmentary, middle of
pronotal disc ± shining, rest of upper sur-
face including elytra (except edges of
suture) microreticulate and dull. Head 0.92
and 0.91 width prothorax. ProtJiorax:
width length 1.22 and 1.25; base/apex 0.94
and 0.91. Elytra: width elytra prothorax
1.82 and 2.00. Measurements: length 3.5-
3.9; width 1.5-1.8 mm.
Tifpcs. Holotvq^e 6 (Bishop Mus.) and
2 paratypes ( 1 in M.C.Z., Type No. 31,438)
from Tobo-Salembeng, Huon Pen., N-E.
N. G., Apr. 26, 1963 (Sedlacek); 1 para-
type, Golden Pines, Bulolo, N-E. N. G.,
600 m, Feb. 19, 1962 (Sedlacek); and 1
paratvpe, Zengaren, N-E. N. G., 1500 m,
Apr. 28, 1963 (Sedlacek).
Measured speeimens. The S holotype and
1 9 paratype from Tobo-Salembeng.
Notes. This may be related to A.
cuprascens Bates (described from Japan
and identified from the Philippines by An-
drewes), but the color of sedlaeeki is
bluish rather than cupreous, the front is
more shining than in euprascens, and com-
parison of specimens shows slight differ-
ences of form not worth describing in de-
tail here.
(Genus PLOCHIONUS Latreille & Dejean)
Latrcille & Dcjeaii 1824, llistoire KaturcIU- et
Icono^iaphie Coleop. d'Europe 1, p. 150.
Csiki 1932, Coleop. Cat., Carabidac, Ilarpalinae 7,
p. 1451 (see for additional references, synonymy,
sul)fienera, and list of species).
Jeannel 1942, Faune de France, Coleop. Caralii(iu(S,
Fart 2, p. 10:33.
Diaij,ru)sis. See Key to Genera of Lehii)u.
and Description of following species.
Descriplion. None rccjuired h('rc\
Type species. Caral)u.s pcdlens l^'abricius
( below ) .
Generic distribution. Native in tropical
and subtropical Aiiirrica, with th(> follow-
ing species now c. cosmoiMjIilaii.
Notes. A supposed ciKleiiiic PlocJiiinius
in New Caledonia needs confirmation.
{Piochionus pollens (Fabricius))
Faliricius 1775, Systenia Ent., p. 244 (Carahns).
Britton 1948, Proc. Hawaiian Ent. Soc. 13, p. 237
( Hawaii).
Jedlicka 1963, Ent. Alihandlungen 28, p. 450.
See also references under trenus.
Description ( for recognition only ). Fonu
as in Figm-e 93; brown; not pubescent; $
front and middle tarsi slighth' dilated, 2-
seriately squamulose; S middle tibiae
arcuate, lower edges broadly shallowly
emarginate below near middle of length;
length c. 7-9.5 mm.
Type. From Europe ("Habitat Dres-
dae"), now presumed lost (not seen).
Occurrence in Neic Guinea. Not yet
found, but likely to occiu".
Notes. This species, probably originally
from America, has been carried b\' luan
to most of the warmer parts of the world.
In the Asiatic-Pacific region it is known
from SE. Asia, Sumatra, Java, New Brit-
ain, New Ireland, New Hebrides, Fiji, and
Polynesia including Hawaii.
Genus PARENA Motschulsky
Motschulsky 1859, etude Eut. 8, p. 31.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1453 ( see for additional references, synonymy,
and list of species ) .
Jeannel 1949, Coleop. Carahiciues de la Rejj;ion
Malyache, Part 3, pp. 948, 971.
Jedlicka 1963, Ent. AMiandlunsien 28, p. 439.
Diaii,nosis. See Key to Genera of Lehiini
of New Guinea and Figm-e 94; note form
stout, sm-face not pubescent, 4th tarsal seg-
mcMits long-lobed; length c. 8-10 mm.
Dcscri))lio)i. None required here.
Ty))c six'cies. Paroui hicolor Motschulsky,
ol Java.
Generic dislrihution. Most species in
area from SK. Asia (including Japan) to
northern Australia, fewer in Africa and
Madagascar.
Notes. Tile ') species that ha\-e been
found in New (Guinea represent 3 indepen-
dent, w idel\ distributed stocks, f'urther
stn(l\- is needed to clarify their geographic
Nariation and noinciulatui-e; in\' ]')r(\s(>nt
The Carabid Beetles of New Guinea • Darlington 139
material is inadequate. I shall therefore
treat the species only briefly.
All members of this genus that I know
are winged and arboreal.
Key to Species of Parena of New Guinea
1. Color entirely testaceous (with sometimes
vague posterior elytral cloud pale brown);
tarsi and antennae contrastingly black (p.
139 ) testacea
- Color partly or wholly darker; tarsi and
antennae reddish testaceous 2
2. Color testaceous or reddish testaceous with
very liroad, well defined black elytral fascia
(p.' 139) fasciata
- Color irregular rufo-piceous, without well
defined elytral marking (p. 139) picea
Parena fesfacea (Chaudoir)
Chaudoir 1872, Ann. Soc. Ent. Belgium 15, p. 178.
(Crossoglossa).
Description. None required here; see
preceding Key, length (in New Guinea) c.
10 mm.
Types. From the Deccan, India; now in
Oberthiir Coll., Paris Mus. (not seen).
Occurrence in New Guinea. N-E. N. G.:
4, Wau, Morobe Dist., 1200 m, June 25,
Oct. 11-18, Nov. 19, Dec. 5-6, 1961
( Sedlaceks ) .
Notes. This species is now known from
India, (China?), Sumatra, Java ("variety"
cruralis Andrewes), and New Guinea (not
Australia ) .
Parena fasciafa (Chaudoir)
Chaudoir 1872, Ann. Soc. Ent. Belgium 15, p. 179
( Crossoglossa ) .
Jedlicka 1963, Ent. Abhandlungen 28, pp. 440,
443, fig. 154.
sloanei Csiki 1932, Coleop. Cat., Carabidae, Har-
palinae 7, p. 1455 (new synonymy).
plagiata Macleay 1876, Proc. Linnean Soc. New
South Wales 1, p. 167 (Phlocodromius) (new
synonymy ) .
Description ( for recognition only ) . Form
as in Figure 94; yellow or reddish yellow
with conspicuous, broad, transverse, black
elytral fascia; length (in New Guinea) c.
8-9 mm.
Types. Of fasciata, from the Moluecas,
now in Oberthiir Coll., Paris Mus.; of
plagiata, from Yule Is., Hall Sound, Papua,
in Macleay Mus., Sydney; of sloanei (new
name), as for plagiata Macleay (none seen).
Occurrence in New Guinea. Papua: Yule
Is. (type of plagiata). N-E. N. G.: 1, Lae,
July 1944 (F. E. Skinner, Purdue U. Coll.,
Bishop Mus.); 1, Busu R. E. of Lae, 100 m,
Sept. 13, 1955 (Gressitt); 1, Bulolo, 732 m,
Aug. 18, 1956 (E. J. Ford, Jr., Bishop Mus.),
in light trap; 1, Finschhafen, Huon Pen.,
180 m, Apr. 16, 1963 (Sedlacek); 1.
Mumeng, 600 m, Mar. 10, 1962 (Sedlacek).
West N. G.: 2, Hollandia, 250 ft.. May 4,
Nov. 3, 1944 (Hoogstraal, M.C.Z.).
Notes. I have seen specimens that I refer
to this species from Java, Borneo, the
Philippines (including Luzon), Celebes,
the Moluccas, New Britain, and northern
Australia, as well as New Guinea.
Parena picea (Macleay)
Macleay 1871, Trans. Ent. Soc. New South Wales
2, p. 86 (Phloeodromius).
Description. None required here; see
preceding Key; length (in New Guinea) c.
9-10 mm^.
Types. One specimen from Gayndah,
South Queensland, Australia (probably
now in Macleay Mus., Sydney) is presum-
ably the actual type (not seen), although
Macleay mentions also "a few specimens
from other portions of Queensland."
Occurrence in New Guinea. N-E. N. G.:
1, Wau, Morobe Dist., 1200 m, Feb. 25,
1963 (Sedlaceks). West N. G.: 1, Nabire,
S. Geelvink Bay, 10-40 m, Sept. 1-4, 1962
( Sedlacek ) .
Notes. I have no specimens from Aus-
tralia and have identified the New Guinean
ones from the original description. I also
tentatively assign to this species single indi-
viduals from New Britain and Manus Is.
( Bishop Mus. ) .
Genus ANCHISTA Nietner
Nietner 1856, J. Asiatic Soc. Bengal 6, p. 523.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1455 (see for additional references, synonymy,
and list of species ) .
Jedlicka 1963, Ent. Abhandlungen 28, p. 449.
140 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Diag,nosis. See Key to Genera of Lebiini
of New Guinea, and under following
species.
Description. None required here.
Type species. Lehia hrunnea Wiede-
mann, of India and Ceylon.
Generic distribution. Tlie few known
species are confined to SE. Asia including
Ceylon and Japan, except that one (below)
is widely distributed on the Malay Archi-
pelago and islands of the western Pacific.
Notes. I know nothing about the habitat
or habits of members of this genus.
Anchisfa binofata (Dejean)
Dejean 1(S25, Species Ceneiai Coleop. 1, p. 252
(Plucluoiiiis).
See also references under genus.
Description { for recognition only ) . Form
as in Figure 95; brownish piceous, each
elytron with longitudinal testaceous area
centered before middle; surface not pubes-
cent; 5th intervals with conspicuous seta-
bearing piuicture at base; length c. 8-9 mm.
Type(s). From the Marianas; now in
Oberthiir Coll., Paris Mus. (not seen).
Occurrence in New Guinea. Papua: 1,
Hagita, near Milne Bay, Aug. 10, 1919
(J. T. Zimmer, Chicago Mus.).
Notes. This species has now been foimd
in SE. Asia (India to Japan), the Anda-
man Is., Sumatra, Java, Borneo, the
Philippines, Buru, New Guinea, and the
Marianas. It has probably been dispersed
partly by man.
Genus ENDYNOMENA Chaudoir
Cliaudoir J 872, Ann. Soc. Knt. Bcliiium 15, p. 1S6.
Csiki 1932, Coleop. Cat., Carabidae, llarpalinae 7,
p. 1457 (see for additional references, synonymy,
and list of species ) .
jedlicka 196.3, Ent. Ahliandlungcn 28, p. 308.
Diuii^nosis. See Kcij to Genera of Jj'hiini
of New Guinea.
Descri))lion. None required here.
Type species. Plochionus })radieri Fair-
maire (below).
Generic distribution. SE. Asia including
Japan, with the following species very
widely spread over the islands of the
Pacific presumably carried by man.
Notes. The habitat and habits of this
genus too are unknown to me.
Endynomena pradieri (Fairmaire)
Fairmaire 1849, Revue and Magazine Zool. 1, pp.
34, 281.
See also references under genus.
Description ( for recognition only ) . Form
as in Figure 96; brown or piceous; surface
with short pubescence; length c. 8 mm.
Type. From Tahiti; in Oberthiir Coll.,
Paris Mus. (not seen).
Occurrence in Neiv Guinea. N-E. N. G. :
1, Sepik, Maprik area, 160 m, Aug. 29, 1957
(Hardy, Bishop Mus.), at light.
Notes. This insect has been recorded
from parts of SE. Asia, Sumatra, the Phil-
ippines, Fiji, Samoa, Tonga, New Cale-
donia, Tahiti, Hawaii, and other remote
Pacific islands, and I have seen a speci-
men from New Britain (Bishop Mus.).
Genus DEMETRIDA White
White 1846, Voyage Erebus & Tenor, p. 2.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1459 (as subgenus of Xanthophoca) (see
for additional references).
Britton 1941, Proc. R. Ent. Soc. London (B) 10.
p. 188.
Xanthophoca Chaudoir 1848, Bull. Soc. Nat.
Moscow 21, Part 1, p. 73.
Diaiinosis. Among New Guinean Lebiini
of the same general form (Figs. 97-109)
and si/e (5..5-12.0 mm), the species of
Demetrida are distinguished by tarsi pubes-
cent (si^arsely pilose) above, with 4th seg-
ment loug-lobed and tarsal claws with sev-
eral or many long t{>eth; ligula and
paraglossae joiiucl, rounded-truncate, usu-
ally 4-setose ( sometimes with 2 additional
smaller setae); palpi not widely e.xpand(xl;
and ' middle tibiac> usualK' (not al\\a\s)
with inner edge in apical '-t or '2 ol length
with a row of several low tubercles.
Descriplion ( aiiplieablc to all New
Guinean species). l'\)rni usualK slender
(broadest in itnilalrix) , convex; color di-
verse, brown ()!■ black or metallic', nniloim
The Carabid Beetles of New Guinea • Darlinaton 141
or bicolored or tesselated (but pattern not sharply defined, usually subpunctate, but
geometric and not simply 2-maculate ) ; disc otherwise c. smooth or at most sparsely
upper surface with short or long pubescence punctulate. Elytra with humeri rounded,
or not pubescent; reticulate microsculpture margined; outer apical angles rounded or
variable, rarely present on whole upper angulate or denticulate; actual apices
surface, often present only on elytra, some- obliquely truncate or sinuate-emarginate or
times absent; elytral reticulations c. iso- angulate, denticulate, or spined c. opposite
diametric or slightly transverse when not ends of 2nd intervals or 2nd striae; striae
otherwise described. Head narrower than entire, usually well impressed, sometimes
or wider than prothorax; eyes prominent punctulate; intervals flat or moderately
but varying from species to species; genae convex, 3rd with 1 (in tenuis only) or 2 or
usually shorter than eyes and oblique, more dorsal, usually seta-bearing punctures,
rarely angulately prominent; 2 setae over and 5th intervals rarely with similar punc-
each eye, the posterior often distant from tures; when 3rd intervals 2-punctate, the
posterior comers of eyes; front impressed punctures often Vi or less from base on outer
each side anteriorly, often also flattened edge and V-i or less from apex on inner edge
and/or weakly impressed or subpunctate of intervals, but positions vary; additional
at middle; mandibles short, strongly curved; punctures (if present) on 3rd intervals
clypeus 1-setose each side; labrum trans- sometimes smaller and more irregular than
verse, 6-setose anteriorly, with additional the 2 primary punctures. Inner wings full
smaller depressed setae at rounded angles; in all New Guinean species (reduced in
antennae slender, pubescent from ( part of ) some New Zealand and Australian ones ) .
4th segments, first 3 segments more sparsely Lower surface variable, often sparsely or
or not pubescent; mentum with entire partly pubescent (least so in imitatrix);
tooth; ligula and paraglossae equal in prosternal process variable in profile. Legs
length, united, forming a rounded-truncate slender; tarsi pubescent (sparsely pilose)
structure with 4 principal setae and some- above; 4th tarsal segments very deeply
times 2 additional smaller setae; palpi emarginate, with long lobes; 5th segments
pubescent, not widely expanded. Protliorax with accessory setae; claws each with 3
cordate or quadrate or trapezoidal, often to 8 long teeth ( not counting apex of claw )
(not always) as long or longer than wide, and sometimes additional smaller teeth, the
sometimes (not usually) wider at base number varying from species to species
than at middle (but width of prothorax and also varying a little individually, some-
always measured at widest midpoint in com- times different on the 2 claws of one tarsus,
puting proportions); side margins varying Secondary sexual characters: S front tarsi
in width in different species, each margin not or not much dilated but with 3 segments
with a seta at or before middle and some- narrowly 2-seriately squamulose below ( the
times also a seta at or near basal angle, squamae sometimes disarranged and not
and in seticoUis with additional setae an- obviously 2-seriate ) ; 6 middle tarsi without
teriorly (setae often broken off, but their sexual squamules; either i middle tibiae
positions marked by characteristic punc- each with a row of 3 to 9 low tubercles on
tures ) ; disc convex, more so in some species inner edge in outer % or V2 of length, the
than in others; anterior and posterior mar- tibial edge being thus subsinuate or sub-
ginal lines absent or incomplete, middle serrate in profile ( Fig. 160 ) ( this condition
line very coarse and deep (except finer in called tuberculate-serrate), or S middle
kokoda), but subbasal and subapical trans- tibiae modified in some other way, or S
verse impressions weak or obsolete; baso- middle tibiae either straight or slightly
lateral impressions usually present but not bent-in at apex but without tubercles (Fig.
142 BuUetin Muscuui of Comparative Zoology, Vol. 137, No. 1
161) (see Notes below); last ventral seg-
ment with 2 to 4 apical setae each side in 6
(possibly only 1 each side in 6 tenuis), 3
to 8 or more in 9 ( except only 2 each side
in 9 tenuis), the number in each sex usually
fairly constant in a species but varying
somewhat individually and sometimes un-
symmetric with ( for example ) 2 setae on
one side and 3 on the other in a c5 , or 5 on
one side and 6 on the other in a 9 .
Type species. So far as I know, type
species have not been strictly designated
for either Demctrida or Xanthophoea.
These genera were based on New Zealand
and Australian forms respectively, and
type species should be selected during
work on the New Zealand and Australian
members of the group. I therefore make
no designations now.
Generic distribution. Numerous in Aus-
tralia and New Guinea, fewer in New
Zealand, New Caledonia, New Britain,
and the Moluccas (Amboina, Batjan) (oc-
currence in New Britain and Moluccas
based on undescribed material before me ) .
Notes. As Britton points out, Demetrida
has priority over Xantho))]i(>ea. The genus
as a whole is diverse. Perhaps it can be
usefully divided, but this will require re-
vision of the many Australian species, which
seem to bridge the gap between the flight-
less (ground-living?) Demetrida of New
Zealand and the winged (arboreal) species
of New Guinea.
The New Guinean species of Demetrida
may all be interrelated but different ones
differ remarkably in many details. Varia-
tion of some characters within the genus
is indicated in the preceding Description,
and some species groupings are suggested
in the Key to Sjjecies. llowevt^r, diff entices
in the 6 middle tibiae, which may distin-
guish natural species groups, are worth
describing in more detail. Of the 56 species
of the genus now recogni/ed Ironi New
Guinea, l)oth sexes are known ol 47, onK
the 6 of 4, and onlx' the 9 of 5. In most
species of whieli the ■ is known, and also
in at least some Australian and New Zea-
land species, the i middle tibiae have the
inner edge tuberculate-serrate ( see Descrip-
tion, above). The nimiber of tubercles
varies from c. 3 to 9 in different species
(with some individual variation too), and
the tubercles vary in prominence, being
sometimes poorly developed and difficult
to see. The tuberculate-serrate 6 middle
tibiae probably characterize most Demetrida
throughout the genus' range. However,
variations from this pattern occur among
New Guinean species. In D. ni<j,ripennis
( and perhaps also in prima, of which the i
is unknown) the c^ middle tibiae have the
inner edge weakly 2-emarginate. In i
imitatrix each middle tibia has a long
tubercle on inner edge separated from the
apex by an emargination. And the following
16 species have the i middle tibiae straight
or slightly bent-in ( slightly bent-out in
reversa) at apex but not or not distinctly
tuberculate-serrate: tripuncta, genicula, an-
gulata, reversa, kiunga, recta, rex, brunnea,
fumipes, ni<^ricej)s, saidor, divisa, liumcralis.
viridibasis, mafulu, and sibil.
Because of the large number of species
and because many characters are shared by
related species or convergent in unrelated
ones, most New Guinean Demctrida can be
defined only by combinations of characters.
However, D. imitatrix is unique in fonn
( relatively broad ) and in form of 6 middle
tibiae. D. viti^il is uni(iue in abrupt promi-
nence of eyes. D. kokoda is unifiue in form
and in fineness ol impressed niiddli' line of
pronotum. D. seticoUis is unique (among
the nonpubescent species) in possessing
extra lateral pronotal setae* anteriorly. And
D. tenuis is uni{|ue in sculpture of front, in
having only 1 seta-bearing pvmelure on 3rd
elytral inter\al, and in ha\ing oiiK 1 apical
seta on each side in the •" , and only 2 in
the 9 . Besides these* single species with
iiiiiciue eharaeters, the lollowing pairs or
small groups of species share special char-
acters. Among New Ciuincnm Demetrida,
oiilx rcltild. riridilxisis. and niafiiln have
The Carabid Beetles of New Guinea • Darlingion 143
almost the whole upper surface micro-
reticulate; only tripuncta and genicuJa have
the genae stronghj angulately prominent,
although tenuis and some other species
ha\e the genae subprominent; only scriato
and nubicola (of nonpubescent species)
have special seta-bearing punctures on 5th
elytral intervals; and only nig,ripennis and
perhaps prima have the i middle tibiae
2-sinuate on inner edge.
Some New Guinean species of Dcmetrida
are remarkably variable. Great individual
variation is indicated by differences in
proportions of the Measured specimens of
some species. And Mendelian dimorphism
is suggested in some cases. For example,
dimorphism or polymorphism of color ap-
parently occurs in Demetrida diversa (mark-
ings black or green, legs dark or pale) and
in mafnhi (markings present or absent),
and color differences among some other
species may be Mendelian, and presence or
absence of certain prothoracic and elytral
setae may be Mendelian too, as is the case
among some other Carabidae. (Genetic
dimorphism of these and other Carabidae
will be considered in more detail in Part
IV of my work on the carabid beetles of
New Guinea.) This situation suggests that
the explosive evolution of Demetrida in
New Guinea, discussed below, is correlated
with great genetic variability of some
species, as would be expected. Different
species may still share homologous genes,
and characters that have become stabilized
in some species may still be dimorphic or
polymorphic in other species.
I think that Demetrida is in fact in the
very midst of an evolutionary explosion in
New Guinea. This is suggested by the
diversity of superficial differences among
many apparently closely interrelated species
and by the great variability of some species.
Apparently one or more ancestors have
recently invaded an open or incompletely
occupied habitat in New Guinea — primarily
the low foliage of rain forest — where other
predaceous beetles of this size are few.
This habitat is occupied in other tropical
regions by Carabidae of the genus Calleida,
which many of the New Guinean Demetrida
resemble in size, form, and even color, al-
though the two genera are well differen-
tiated taxonomically by differences in
mouthparts, tarsal pubescence, etc. That
Demetrida and Calleida are geographically
complementary is true but an oversimplifi-
cation. The situation is complicated in
many ways, for example by the presence
of many species of Lebia in some other
tropical regions but few in New Guinea.
The ecology of Demetrida seems con-
sistent with a recent independent radiation
of the New Guinean species. While most
New Guinean species apparently live in
fohage in rain forest, most Australian
species live on shaggy-barked tree trunks
(especially of Eucalyptus trees) in rela-
tively open woodland, and the Australian
tree-trunk forms and the New Guinean
rain-forest-foliage forms have evidently
radiated independently. Although this is
true, it is another oversimplification. A few
northern Australian species of Demetrida
do inhabit rain-forest foliage, but they are
very few, uncommon, and probably ecolog-
ically unimportant. Perhaps they represent
the ancestral stock(s) from which the New
Guinean rain-forest forms have evolved.^
^ The following key characterizes 3 species of
Demetrida from North Queensland, Australia, that
are or may be members of the New Guinean
radiation of the genus. D. angulata, described in
the present paper, is the only species known to be
common to Australia and New Guinea. The other
2 Australian species named in the key are distin-
guished by 1-punctate 3rd intervals from all known
New Guinean si^ecies except tenuis.
Key to Certain Australian Demetrida
1. Elytral apices sinuate-truncate; 3rd intervals
1-punctate 2
- Elytral apices obtusely angulate; 3rd intervals
2-punctate (1 9 , Rocky R. on Cape York;
New Guinea) angulata (n. sp. )
2. Prothorax with posterior-lateral setae (11,
middle Cape York, Cairns, Kuranda, etc.,
vie. Brisbane, Clarence R. )
longicollis Macleay
144 BiiUetin Museum of Comparative Zoology, Vol. 137, No. 1
Also, a few Australian Demetrida live in
long grass, and a few unrelated New
Guinean species (perhaps pallen.s?) may
have invaded grassland independently, at
high altitudes — but this is a guess, based
on the insects' appearance; the actual habi-
tats of the New Guinean species in question
are not recorded. However, these excep-
tions and doubts do not change the general
fact: Demetrida has radiated ± indepen-
dently in the New Guinean rain forest, and
the radiation may be continuing explosively
now. The radiation of these beetles parallels
in some ways the radiation of birds of
paradise in the same forests.
The geographic distribution of different
species of Demetrida in New Guinea is
not yet very well known, and the ecologic
distribution of the species within the rain-
forest-foliage habitat is hardly known at
all. Some species of the genus are ap-
parently localized in parts of New Guinea,
and geographic replacement may occur in
some cases. But many other species are
evidently wide-ranging on the island, and
many species sometimes occur together at
one locality or in a very limited area. For
example, I found 8 species at Dobodura,
at relatively low altitudes. And 19 species
have been found at or near Wau on the
Morobe Plateau, at mid-altitudes. The
genus as a whole ranges in New Guinea
from sea level to or above timber line but
is evidently best represented at mid-alti-
tudes, where it is a]:)parently dominant,
and where most of the strikingly colored
species occur. Some species are evidently
confined to or specialK characteristic of
either low, middle, or high altitudes, and
related species may replace each other at
different altitudes in some cases.
- Prothorax without posterior-lateral setae 3
3. Protliorax narrower ( width length 0.84 and
0.93); front snhearinate (New (liniiea
only) (/('/II//.V n. sp. )
- Prothorax wider (width/length 1.07); front
not snhearinate (1 9, Cairns)
_ ferruginca Cha ndoi r
As to their ecolog\ , the bright color of
some New Guinean Demetrida and the
fewness of individuals taken at light sug-
gest that most species are diurnal. How the
various species that occur together, for ex-
ample at Wau, di\ide the niches within
the rain-forest-toliage habitat can only be
guessed at now. A few may have become
nocturnal. Some species certainly live in
understory vegetation in the rain forest,
but some may live at mid-levels and some
77uiy live in the actual tree tops. Different
species tmiy specialize in narrower habitats,
or they 7nay specialize in different kinds
or sizes of prey. But I should repeat that
this is mainly guesswork. There is an op-
portunity here for exciting work in the
field, on the ecologic radiation of a donii-
nant group of insects that is radiating
structurally.
Some New Guinean Demetrida may be
mimics. Evolution of mimetic relationships
would, I think, be consistent with the genus
being now in the midst of an evolutionary
explosion, with many species geneticalh
variable, read)' to respond to special selec-
tion pressures. Demetrida imitatri.x re-
sembles and may mimic the common New
Guinean carabid Viola <i,onum violaeeum
(Ghaudoir), and some other brighth- colored
Demetrida ma\' mimic other (^arabidae
(perhaps certain C^olliurini) and other
Ix'etles.
An extraordinar\ eircumstanee is that,
although many species of Demetrida occur
in New Guinea and although some of them
are common ( 1 have examined a total of
about 1250 individuals) all 56 New Ciuinean
si)ecies seem to be undeseribed! Howe\er,
this should not be interpretc^d as e\id(Mice
ol e\()lution within historic tinu-s. Most
ol the common species occur at mid-alti-
tudes in the mountains, where not much
carabid collecting was done until [']\'elyn
Gheesmans time, in the If).3()'s, and where
really extensiNC eolleetions of ('arabidae
have been made only recentK, by Dr.
Co-(\ssitt. the Se(llae(>ks. and other l^ishop
The Carabid Beetles of New Guinea • Darlington 145
Museum entomologists. Andrewes, during Beetles of New Guinea," Bull. Mus. Comp.
his work on Oriental Carabidae, did see a Zool. 126, No. 3, pp. 328-330).
few older specimens of Demctricki from In drawing descriptions in this genus I
New Guinea, including one or two of the have used "c." (circa, meaning approxi-
strikingly colored forms, but he refrained mately) even more often than usual, as a
from describing them; he did not know stratagem for saving space where I do not
what genus to put them in! So, I think think exact or detailed statements are use-
failure of earlier authors to describe New ful. I have also sometimes used it as a
Guinean species of Demctrida was a result warning that variation probably occurs al-
partly of the inaccessibility of the habitats though my material is too limited to show it.
of most common species, partly of com- A statement of my procedure in attacking
mendable caution on the part of taxonomists the particularly difficult problem presented
including H. E. Andrewes, and probably by the New Guinean Demetrida may be
partly just of chance. useful to future taxonomists. My method
Methods. My specific descriptions in has been to alternate between the general
Demetrida follow a special, slightly modi- and the particular, with first a general
fied form designed to characterize the sorting of individuals into apparent species
species adequately without wasting space, and preparation of a very preHminary key.
Characters covered in the generic Descrip- then drawing of detailed descriptions
tion are not repeated, but each specific species by species to determine characters
description begins with a statement that and variation, then preparation of an im-
the species shares the generic characters. In proved general classification and an im-
addition to the usual proportions, the ratio proved key, then further checking of details
of width of base of prothorax width of head and variation, and eventually (by a much
is given; it is especially useful in distin- longer process than this! ) preparation of a
guishing some species of Demetrida. The final key and descriptions emphasizing
headings Inner icings. Lower swface, and characters that have proved significant and
Leii,s are omitted; these subjects are suf- emphasizing variation, and last of all com-
ficiently covered in the Description of the pletion of introductory and explanatory ma-
genus. A special heading Claws is added terial, including the present statement. This
because number of claw teeth may prove is the general method that taxonomists use
to be diagnostic of some species. in classifying any unknown animals, but
Secondary sexual characters, especially the process has been much more complex
modifications of the S middle tibiae, have in Demetrida than usual. Specific problems
been examined carefully and used in char- have been numerous and difficult. In some
acterizing species. The tibiae are best seen cases I cannot be sure from available ma-
against an illuminated white background, terial whether differences in color, presence
To see a tibia clearly at the proper angle or absence of setae, or length of elytral
it is often necessary to straighten a middle spines are specific characters, cases of
leg, and this can usually be done without Mendelian dimorphism, or other individual
relaxing the specimen, by pulling the tibia variations. I have had to decide these
straight with a pin point and putting a cases arbitrarily, and my groupings of
minute drop of glue on the articulation to species are partly arbitrary. Tlie resulting
hold the straightened tibia in place. How- classification is at best an approximation,
ever, I have not examined the 6 copulatory Of course this is true of most classifications,
organs. This is a task for third-stage taxono- but I am more than usually conscious of
mists, far beyond what I have time to do the fact in this case,
now ( see Part I of my "The Carabid The question may be raised, why publish
146 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
a classification that is only a doubtful ap-
proximation? The answer is that it presents
an exciting situation in the only way that it
can be presented now. Further work on
Demetrido, including field work (which is
essential), requires some sort of classifica-
tion. In this and many other cases where
a classification is needed, w^e can only
follow what I think is a basic rule of the
trade: a taxonomist must do the best he
can with the available material in the avail-
able time. Actually, this imperfect treat-
ment of Dcmetrida may prove to be the
most important part of my taxonomic work
on New Guinean Carabidae.
The following Key to Species of Demetrida
of New Guinea is complicated and at some
points difficult to use. This is inevitable in
the case of an "exploding" group in which
some species are exceptionally variable and
others connected by intergrades. The key
is designed primarily for identification. It
is partly but not wholly phylogenetic:
species that are closely grouped in the key
are likely to be related but are not neces-
sarily so. A few dimorphic or exceptionally
variable species are run out at two different
points in the key, but I have had to limit
such multiple treatments. A few individuals
arc therefore unidentifiable by key char-
acters and have to be placed by comparison
of specimens. I might be able to construct
a multiple-tn-atmcnt kc-y that would iden-
tify every iiidixidual variant of eacli
Demetrida now known from New Guinea,
but the key would be impossibly complex
even lor present use and it would not take
care of new material, which will surely
include new variants of many species. The
key must be used with care and discretion.
Proportions must be calculated from inea-
sinements. Much variation must be allowed
for, more than I have been able to indicate
in detail. Alternatives must be tried when
specimens do not key out clearly. The test
of a key like this is whether it works rea-
sonably well in practice. First-time users
will probabK' Hud it \{'r\' difficult. Persons
who become more familiar with it will, I
hope, find shortcuts, in part suggested by
section headings inserted in brackets.
For comparisons of New Guinean De-
metrida with Australian species see espe-
cially Footnote .3 ( p. 143 ) and under D.
j)rima (p. 150).
Key to the Species of Demetrida of
Neav Guinea
[Pubescent]
1. Most of upper surface including sides of
head behind eyes plainly pubescent 2
- Surface not pubescent or (.scriata and
nuhicola only) pubescence very sparse,
fine, scarcely detectable 8
2. Pubescence short; elytra truncate or sinuate-
truncate at apex 3
- Pubescence long, sparse-pilose: elytra usu-
ally lobed or spined at apex ( scarcely so
in pallen.s) 6
3. Posterior-lateral prothoracic setae present,
at basal angles; smaller, length under 7.5
nmi (p. 149) aitape
- Posterior-lateral prothoracic setae absent;
size larger 4
4. Eyes more prominent, head nearly as wide
as prothorax ( width head/prothorax 0.98
and 0.96); 9 with 3 ( .^probably 2) apical
ventral setae each side; (length c. 9 mm)
( p. 149 ) goroka
- Eyes less prominent, head relatixely nar-
rower; apical ventral setae more numer-
ous 5
5. Color entirely brown; length usualK' more
than 9 nun (p. 150) priiiui
- Bicolored, head and prothorax brown,
elytra nearly black: length usually less
tliiui 9 mm (p. 151) __ fi/grZ/K'/ui/.v
(1 b^Ktral apices siiniate-truucate or we;ikl\'
lobed; color irreguLir p;ile brown (Y>.
151 ) pallcus
- Elytral apices spined 7
7. brown, el\ tra uitli pale speckles (p.
151 ) tcsscldtd
- Almost black, el\ tra faintly or not speckled
(p. b52 ) crcpciii
[I'UijtKi tiuncdic]
(S. l']|ytral apices oblicjuc^b' truncate or sinu;ite-
truucate (Figs. 97, 99) 9
- Elytral apices ;mgulate, toollied, spined,
or at le;ist subaugnhitely lobed (Fig. 102)
c. opposite ends of 2n(l striae or 2nd
iuter\als 15
9. TImHI and iisn;ill\ 5tli eblial intcrxals e;ieli
with se\('r;il st't;i-be;n ing punetuies; upper
suilaee with ;i little sp;irse, line, scarcely
(leteetiible jMibescence 10
The Carabid Beetles of New Guinea
Darlington
147
- Third intervals with 1 or 2 and 5th inter- 20.
vals without seta-bearing punctures; upper
surface without such pubescence; (color -
brown) 11 21.
10. Color entirely brown (p. 153) seriata
- Bicolored, reddish brown with base of
elytra black; (see also Description) (p.
153 ) mibicola
11. Large (over 10 mm) and prothorax
strongly narrowed in front and outer 22.
angles of elytra distinct, only slightly
blunted (p. 154) magna
- Either smaller or with prothorax differently
shaped or with outer elytral angles
rovmded 12
12. Prothorax wider than long at middle (by
measurement); posterior-lateral prothoracic 23.
setae usually present 13
- Prothorax longer than wide; posterior-
lateral prothoracic setae alisent 14 _
13. Larger (7-9.8 mm); outer angles of elytra
distinct although sometimes slightly blunted 24.
(p. 155) truncata
- Smaller (5.6-6.3 mm); outer angles of
elytra bhuited or rounded (p. 155) _^.- minor
14. Third elytral intervals 2-punctate; apical
ventral setae 3 each side in i , probably 25.
more in 9 (p. 156) std)temti.s
- Third elytral intervals 1-punctate; apical
ventral setae 1 each side in S , 2 in $ ( p.
156 ) tenuis
[Elytra ungulate, toothed, or spined]
15. Pronotum without posterior-lateral seta-
bearing punctures 16
- Pronotum with posterior-lateral seta-bear-
ing punctures at or near posterior angles --. 57
16. Color above brown (testaceous to piceous),
reddish, black, or bicolored black-and-
paler, but not in any part metallic 17
- Color above partly or wholly metallic
blue, blue-black, green, or purple, often
but not always bicolored 50
17. Not distinctly bicolored above, usually ±
uniform lirown, sometimes grading into og
darker brown or piceous on some parts _
of body ( some doubtful species are run
both ways in the key) 18 £7
- Sharply bicolored above, partly brown or _
reddish, partly (sometimes only broad
humeral areas) black 39 28.
18. Genae angulately or roundly prominent; _
(3rd intervals of elytra 3-punctate) 19 29.
- Genae oblique or weakly rounded, not
prominent 20
19. Elytral apices obtusely angulate (p.
157) tripuncta -
- Elytral apices acutely toothed ( p. 158 ) -^_.
_._ genicula
EKtral apices weakly lobed or angulate
with the angles blunted, obtuse, or right 21
EKtral apices acutely dentate or spined __ 23
Prothorax subcordate, wider than head and
much wider than long ( width/length c.
1.35); length 5.5-7.1 mm (p. 158) __ latangula
Prothorax subcjuadrate, usually (not always)
narrower than head and not or not much
wider than long; size larger 22
Prothorax slightly narrower (cf. descrip-
tions); i middle tibiae slightly bent-in
at apex; length 7.5-8.9 mm; (Papua) (p.
159 ) angulata
Prothorax slightly wider; $ middle tibiae
slightly bent-out at apex; length 8.5-9.2
mm; (West N. G. ) (p. 159) reversa
Median impressed line of pronotum fine;
(form as Fig. 103, very elongate; length c.
10-11 mm) (p. 160) kokoda
Median impressed line of pronotum
coarse 24
Prothorax at middle usually wider than or
equal to width of head, or only slightly
narrower 25
Prothorax much narrower ( usually by %o
or more) than width of head 33
Prothorax more cordate, usually wider (c.
% to 'h-i wider than long, with base often
wider than head) with sides more rounded
and more evenly rounded anteriorly, and
with wider margins, and 3rd elytral interval
with only two dorsal punctures, and color
entirely brown, and length usually less
than 7 mm 26
Either prothorax more quadrate and nar-
rower (less than Mi wider than long, with
base usually narrower than head) with
sides often but not always less evenly
rounded anteriorly and with narrower
margins, or 3rd elytral interval with more
than t\\Q dorsal punctures ( additional ones
sometimes smaller than the 2 primary
ones), or color darker (at least partly
blackish), or size larger 27
Elytral apices spined (p. 160) moda
Elytral apices acutely angulate or short-
toothed (p. 161) suhmoda
Prothorax with wider margins, ± cordate 28
Prothorax with narrower margins, quadrate
or more narrowly cordate 31
Smaller, length less than 8 mm 29
Larger, length 8 mm or more 30
Eyes more prominent; base of pronotum
less punctate; elytra usually with (some-
times faint) reticulate microsculpture (p.
162 ) hollandia
Eyes less prominent; base of pronotum
more punctate; elytra without reticulate
microsculpture (p. 162) toau
148 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
30. Third elytral interval 2-punctate (p. 163) _ 41.
similis
- Third elytral interxal with more than two -
punctures (p. 164) dupliccita
31. Smaller, length 6.7-7.6 mm; ± piceous; 42.
base of prothorax more punctate; 3rd
elytral interxals with more than two punc- -
tures (p. 164) suhpunctaia
- Larger, length 7.7-9.2 mm; browii; base 43.
of prothorax less punctate; 3rd elytral in-
tervals 2-punctate 32 _
32. Male middle tibiae tuberculate-serrate;
length 7.7-9.2 mm (p. 165) dohodura
- Male middle tibiae not tuberculate-serrate;
length c. 10.8 mm (p. 166) kiiin^a
[Protliorax much uanoucr than head]
33. Elytral 3rd intervals 2-punctate, and color
reddish brown \\ ith legs pale except knees
.sometimes dark, and length not over 10
mm 34
- Elytral 3rd intervals 3-punctate, or ( // 3rd
interval 2-punctate) color in part darker
with legs partly or wholly dark, or size
over 10 mm 36
34. Most of upper surface distinctly micro-
reticulate (p. 166) mafidii
- Microreticulation distinct (if at all) ouK
on elytra 35
35. Male middle tibiae tuberculate-serrate;
sides of prothorax usually more rounded-in
at apex (p. 167) forma
- Male middle tibiae bent-in at apex but not
tuberculate-serrate; sides of prothorax usu-
ally almost straight anteriorly (p. 167) recta
36. Very large, length 10.2-11.4 nun, and
basal angles of prothorax very prominent
(Fig. 105) (p. 168) L rex
- Smaller; l:)asa! angles of prothorax usualK
less promiui'iit -. 37
37. Legs red or brown, not darker llian disc
of elytra (p. 169) hitntnca
- Legs partly or wliolly dark _ 38
38. Third inter\als of elytra usually 3-pune-
tate; pronotum witlumt reticnkitc micro- _
sculpture (p. 169) .. . fumipcs ^
- Third intervals usually 2-punctate; pro-
notum ( h'glitly ) mierori'ticulate ( [). 170)
vclata
[Bieolorcd hut not mclalJic] 52.
39. Head and pronotum retldisli, cl\ tr;i en-
tirely dark or with only apices slightly -
reddish 10 r>:].
- Pattern not as described; elytra usually
(not always) bicolored 42 -
40. Lh Iral striae very lightly impressed; le.gs
l)lack (p. 171) ni^ri))e.s
- Elvtral striae well impressed; legs usually 54.
paler _ 41
44.
45.
46.
47.
48.
49.
[A/<
50.
El\tra usualK with reticulate microsculp-
tnre (p. 162) hoUandia
Elytra without reticulate microsculptiu'e
(see also couplet 29) (p. 162) wait
Prothorax wider (c. V-, or more wider than
long ) 43
Prothorax narrower ( c. long or longer than
wide ) 44
Elytra black with large connnon discal
area red (p. 171) dorsali.s
Elytra reddish brown with dark base
(western N. C;.; for individuals from
central and eastern N. C. see 'Notes imder
duplicata, p. 164) (p. 172) hasalis
Elytral apices angulatc: (color diverse,
see Description ) (p. 172 ) diversa
Ehtral apices spined (spines sometimes
short ) 45
Eyes abrupt (Fig. 107); (anterior angles
of protliorax more rounded than usual ) ( p.
173 ) vigil
Eyes normal, prominent but less abrupt 46
Head and pronotum black or piceous, or
at least darker than elytral disc 47
Head and pronotum red or brown 48
Elytra wholly brown, long-spined (p.
174) nigriceps
Elytra brown with darker humeri, shorter-
spined (p. 174) saidor
Elytra red or brown with base wholly
black, except suture sometimes red to base;
le.gs pale, sometimes with dark knees ( p.
175 ) ilivisa
EK tra with onl\- liiuntai dark; legs dark or
l)icolorcd 49
Dark humeral ari'as wider, usualK in-
chiding parts of 5th inter\als; lengtli 9..3-
10.8 mm (p. 175) humeralis
Dark humeral areas narrower; length 8.3-
9.4 mm (p. 169) _.. ftimi))cs
iaUic at least in jiart]
Broad ( bro;idcst i)i inctrida) : color en-
tircK bhic-black (p. 176) imitatrix
More slender; color not ;is described 51
ilc;id ;in(l prothor;ix red, eKtr;i entirely
green cxcipt sometimes jiurplisli posteri-
orK (p. 177) -- I iridipi-nni.\
(]()l(ii' I II it as dcsciibcd 52
Clolor entircK' green, .urecn ;uid black, or
gi'cen ;uKi purple 53
i^icolorcd, cKtrii in part icd or brown 54
Linger (9.2-10.8 mm); cncs more iibrupt;
eKtra lon.g-spined (p. 177) U'pida
Smaller (7.0-9.0 mm); eyes less ;ibrupt;
(Kli.i nsiialK sliorliT-spiiH d (p. 178) ...-
std)Iepida
llc;id :in(l j^roiiotum led or blown, not
I'l.iiiiK nu'lallic- 55
The Carabid Beetles of New Guinea • Darlingtun
149
- Pronotuni and sometimes also head metallic
green 56
55. Elytral apices angnlate (p. 172) __._ diversa
- Elytral apices short-spined (p. 179)
viridibasis
56. Elytral apices acutely annulate; most of
upper surface microreticulate (p. 166) iiiafulii
- Elytral apices long-spined; only elytra
(faintly) microreticulate (p. 179) sibil
{Posterior-lateral setae presciit]
57. Color hrown-piceous (head and pronotum
usually darker than elytra); prothoracic
margins with extra setae anteriorly (p.
180 ) seticollis
- Color wholly or mainly black, blue-black,
or green-black; prothoracic margins with-
out extra setae anteriorly 58
58. Legs red testaceous (p. 181) pallipes
- Legs dark 59
59. Elytra black with common discal area red
(p. 181) discoidalis
- Elytra metallic blue-black or green-black
without discal red area 60
60. Elytra short-spined ( p. 182 ) .— sedlacekorum
- Elytra long-spined (p. 182) hrandti
Demetr'ida aitape n. sp.
Description. With characters of genu.s;
form nearly as in following species {fS,oroka,
Fig. 97), slender, eyes prominent, elytral
apices obliquely subtruncate; entirely brown;
surface short-pubescent, without r(>ticulate
microsculpture but sparsely punctulate.
Flead 0.91 and 0.90 width prothorax; eyes
moderately prominent, genae shorter and
oblique. PwtJiora.x narrowly subcordate;
width/length 1.10 and 1.13; base/apex 1.36
and 1.34; base/head 0.94 and 0.91; sides
broadly rounded in anterior %, broadly
sinuate before right or slightly acute some-
times minutely blunted posterior angles;
margins rather wide in proportion to width
of insect, each with seta before middle
and at or just before basal angle; baso-
lateral areas slightly depressed and more
closely punctate than disc. Elytra long;
width elytra/prothorax 1.69 and 1.61; apices
slightly obliquely sinuate-truncate, with
outer angles moderately and sutural angles
narrowly rounded; striae deep, subpunctate
(or with sides of intervals slightly irregu-
lar); 3rd interval with c. 4 special seta-
bearing punctures (difficult to distinguish
amid other punctation). Claws with c. 6
teeth. Secondary sexiial characters: S tarsi
as genus; i middle tibiae tuberculate-ser-
rate ( c. 5 small tubercles ) ; 6 with 3 or 4,
$ 4 or 5 apical ventral setae each side.
Measurements (types only): length 5.8-
7.2; width 1.8-2.3 mm.
Types. Holotype $ (M.C.Z., Type No.
31,439) and 3 paratypes from Aitape, N-E.
N. G., Aug. 1944 (Darlington); 6 para-
types, Mt. Lamington, Papua (C. T.
McNamara, S. Australian Mus.).
Additional jnaterial. Papua: 1, Do-
bodura, Mar.-July 1944 (Darlington). N-E.
N. G.: 1, Erima, Astrolabe Bay, 1896 (Biro).
West N. G.: 1, Dojo [near Hollandia],
Apr. 1958 (G. den Hoed, Louwerens Coll.,
to Leiden Mus. eventually).
Measured specimens. The c^ holotype and
1 9 paratype.
Notes. The relatively small size and
presence of posterior-lateral prothoracic
setae distinguish this species from other
short-pubescent ones in New Guinea. The
individuals listed under Additional material
are slightly larger than the types and vary
slightly in form, but seem to be conspecific.
x\ll have setae at the posterior angles of
the prothorax as well as before middle,
and all are 9 9 with 5 setae each side last
ventral segment.
Demetrida goroka n. sp.
Description. With characters of genus;
form as in Figure 97, slender, eyes promi-
nent, elytral apices obliquely subtruncate;
reddish brown, elytra darker but not black,
appendages brown; surface short-pubes-
cent, without reticulate microsculpture but
sparsely irregularly punctate. Head 0.98
and 0.96 width prothorax; eyes rather
abruptly prominent, genae slightly shorter,
sinuate-oblique; front flattened, irregularly
impressed. Frothorax narrowly subcordate;
width length 1.04 and 1.02; base apex 1.26
and 1.38; base/head 0.86 and 0.91; sides
broadlv rounded in anterior %, broadly
150 BuUetin Museum of Comparative Zoology, Vol. 137, No. 1
sinuate before sharply acute posterior
angles; margins rather wide in proportion
to width of insect, each with seta slightly
before middle but none at base; baso-
lateral areas irregularly impressed, punc-
tate. Elytra long; width elytra /prothorax
1.69 and 1.68; apices slightly obliquely
sinuate-truncate, with outer angles blunted
or rounded, sutural angles narrowly
rounded; striae deep, irregularly subpunc-
tate; 3rd intervals each with 4 special
punctures in type, the punctures in part
obscured or absent in paratype. Chas with
6 or 7 teeth. Secondary sexual characters:
$ unknown; 9 with 3 apical ventral setae
each side in both specimens. Measure-
ments: length ± 9.0; width 2.8 mm.
Types. Holotype 9 (Bishop Mus.) and
1 9 "paratype (M.C.Z., Type No. 31,440)
both from Goroka, N-E. N. G., 1500 m.
May 22, 1961 (J. L. & M. Gressitt), taken
in light trap.
Notes. D. <:oro1<a superficially resembles
prima and nin,ripennis (below) but has
wider head \\\\\\ more prominent eyes,
slightly narrower prothorax ( with more dis-
tinctly lobed base), and only 3 apical ventral
setae each side in 9 (probably only 2 in $),
while 9 9 of prima and nigripennis have
5 or 6 such setae each side.
Demetrida prima n. sp.
Descri])tion. With characters of genus;
form nearly as in preceding species (goroka.
Fig. 97), slender, but with eyes less promi-
nent than in goroka, elytral apices obliquely
subtruncate; reddish brown; surface short-
pubescent, without reticulate microsculp-
ture but sparsely punctulate. Head 0.91
and 0.87 width prothorax; eyes less promi-
nent than usual in genus, genae long-
{)bli((uc. Prothorax narrowly subcordate;
width/length 1.03 and 1.08; base/apex 1.24
and 1.11; base/head 0.96 and 0.93; sides
weakly arcuate anteriorly, broadly sinuate
before right or slightK' acute well defined
basal angles; margins moderate, (\ich ap-
parently with special seta-bearing punc-
ture near middle of length but not at base
(these setae and punctures difficult to
distinguish amid general pubescence); baso-
lateral impressions punctate. Elytra long;
width elytra prothorax 1.64 and 1.53; apices
truncate or weakly sinuate-truncate, with
outer angles moderately and sutural angles
more narrowly rounded; striae deep, ir-
regularly punctate; 3rd interval with ap-
parently 2-5 special seta-bearing punctures
( sometimes difficult to distinguish amid
other punctation ). Clans with 5 or 6 teeth.
Secondary sexual characters: 6 unknown;
9 with c. 5 apical ventral setae each side.
Measurements: length 9.0-9.8; width 2.8-
3.0 mm.
Types. Holotvpe 9 (Bishop Mus.) from
Wau, Morobe Dist., N-E. N. G., 1200 m.
Mar. 23, 1963 (Sedlacek), in mercury vapor
light trap. Additional (99) paratypes
from N-E. N. G. as follows: 1, Maprick,
160 m, Dec. 29, 1959-Jan. 17, 1960 (T. C.
Maa, now in M.C.Z., Type No. 31,441); 1,
Torricelli Mts., Siaute, sea level, Nov. 9-17,
1958 (W. W. Brandt, Bishop Mus.); 1,
Mumeng, 600 m. Mar. 9, 1962 (Sedlacek).
Measured specimens. The 9 holotype and
9 paratype from Mumeng.
Notes. Among New Guinean Demetrida,
prima should be easily known by form,
including form of elytral apices, rather large
size, nearly uniform reddish brown color,
and short-pubescent surface.
Superficially prima resembles some Aus-
tralian species of Demetrida. For example
it is somewhat similar in form to grandis
(Chaudoir) of southern Australia but has
shorter antcnmae, smaller eyes, less promi-
nent genae, prothorax narrower anteriorly
with narrower margins, and elytra uni-
formly brown (not strip(>d as in grandis).
D. prima also somewhat resembles con-
stricticeps (Sloane) of southwestern Aus-
tralia in lonn and is similar in color, but
]nima has shorter antennae, much less
prominent genae, less strongU' simiate sides
ol prothorax, and dilleis in other details.
And prima differs Ironi l)()th the Aus-
The Carabid Beetles of New Guinea • Darlinaton
151
tralian species named and from all other
Australian species known to me in amount
and character of pubescence.
Demetrida nigripenn'is n. sp.
Description. See Plate 1, figure I; with
characters of genus; form, elytral apices,
pubescence, punctation, and other asexual
characters c. same as in preceding species
(prima), but color brownish red with
elytra black or nearly so, and size smaller.
Head 0.88 and 0.89 width prothorax. Pro-
thorax: width/length 1.06 and 1.07; base/
apex 1.20 and 1.18; base/head 0.90 and
0.98. Elytra: width elytra/prothorax 1.56
and 1.64; 3rd intervals with apparently 1-4
principal seta-bearing punctures (difficult
to distinguish). Secondary sexual char-
acters: S tarsi as genus; 6 middle tibiae
weakly 2-emarginate on inner edge near
apex; S with c. 4, 9 c. 6 apical ventral
setae each side. Measurements: length
8.3-9.0; width 2.5-2.8 mm.
Types. Holotype S (Louwerens Coll.,
eventually to Leiden Mus.) and 1 S para-
type (M.C.Z., Type No. 31,442) from Dojo
[near Hollandia], West N. G., Apr. 1958
(G. den Hoed); and 1 9 paratype, Hol-
landia, May 1945 (B. Malkin, U.S.N.M.).
Measured specimens. The i holotype and
9 paratype.
Notes. This may prove to be a geographic
form of the preceding species ( prima ) , but
more material of both sexes from more
localities is needed to clarify the relation-
ship. The form of the c^ middle tibiae is
unique among known members of the
genus.
Demetrida pailens n. sp.
Description. See Plate 1, figure II; with
characters of genus; eyes prominent, pro-
thorax small, cordate-quadrate, and elytral
apices sinuate-subtruncate and usually sub-
angulate c. opposite ends 2nd striae; color
irregular testaceous brown, elytra irregu-
larly tesselated with small paler spots; sur-
face long-pubescent, without reticulate mi-
crosculpture, punctate as described below.
Head 1.10 and 1.08 width prothorax; eyes
prominent, genae short-oblique. Prothorax:
width/length 1.09 and 1.03; base/apex 1.36
and 1.38; base/head 0.85 and 0.91; side
margins moderate, entirely fringed with
long setae; disc irregularly sparsely punc-
tate. Elytra: width elytra/prothorax 2.07
and 2.17; apices usually as figured, sub-
angulate or weakly lobed (simply sinuate-
truncate in 1 specimen), outer angles
broadly and sutural angles less broadly
rounded; striae moderately impressed, in
part slightly interrupted, irregularly sub-
punctate; intervals all with series of coarse
seta-bearing punctures among which special
dorsal punctures are not distinguishable.
Claws with 7 or 8 teeth. Secondary sexual
characters: 6 tarsi as genus; S middle
tibiae tuberculate-serrate (c. 6 small tuber-
cles); c^ with c. 4 apical ventral setae each
side; 9 unknown. Measurements: length
8-9; width 2.9-3.3 mm.
Types. Holotype c5 (Leiden Mus.) and
4 $ S paratypes (2 in M.C.Z., Type No.
31,443) all from Moss Forest Camp (Snow
Mts.), West N. G., 2800 m, Oct. 9-Nov. 5,
1938 (Toxopeus).
Measured specimens. The $ holotype and
1 9 paratype.
Notes. This very distinct species occurs
at a higher altitude than any other
Demetrida known to me. The coloration,
which superficially resembles that of some
high-altitude Agonini (some Mactdagonum),
suggests that the insect lives in grass, al-
though the specimens were taken at "Moss
Forest Camp."
The (slight) variation in form of elytral
apices is one of many examples of individ-
ual variation in this remarkably variable
genus.
Demetrida fesselata n. sp.
Description. With characters of genus;
form ( Fig. 98 ) c. average, with eyes promi-
nent, prothorax small, elytra spined; color
irregular dark reddish brown, elytra with
152 BiiUetin Museum of Comparative Zoology, Vol. 137, No. 1
numerous small pale flecks forming rows
most conspicuous on ( but not confined to )
odd intervals, legs pale; surface sparsely
long-pubescent, without reticulate micro-
sculptme, irregular but scarcely punctate
except for punctures (variable in size) from
which hairs rise. Head 1.16 and 1.08 width
prothorax; eyes prominent, genae nearly as
long as eyes, oblique. Prothorax small, nar-
ro\N']y cordate-subquadrate; width length
1.03 and 1.07; base apex 1.31 and 1.35;
base head 0.81 and 0.88; sides weakly
rounded anteriorly, often subangulate near
middle of length; side margins moderate,
irregularly fringed for entire length with
long setae; baso-lateral depressions poorly
defined, irregular but scarcely punctate.
Elytra: width elytra prothorax 2.28 and
2.23; apices spined, outer angles rounded
or obtusely blunted (variable), sutural
angles obtusely blunted; striae lightly im-
pressed, in part interrupted or reduced to
rows of punctures; intervals flat but ir-
regular, odd intervals with series of seta-
bearing punctures of moderate size, each
puncture usually on posterior side of a
broad low tubercle. Claws with c. 6-8 teeth.
Secondary sexual characters: 6 tarsi as
genus; i middle tibiae tuberculate-serrate
(c. 5 widely spaced small tubercles); i
with 3 (or more?), 9 5 or 6 apical ventral
setae each side. Measurements: length 8.7-
9.8; width 3.0-3.4 mm.
Types. Holotype $ (Bishop Mus.) and
1 9 paratype (M.C.Z., Type No. 31,444)
from Mt. Kaindi, N-E. N. G., 2350 in, Jan.
10 and June 9, 1962 (Sedlaceks), "the
paratype taken in mercury xapor light
trap; and 1 additional paratype i without
head, same locality, 2400 m, Jan. 28, 1963
(Sedlacek); 3 paratypes, Wau, 2400 m, Jan.
9-12, 1962 (Sedlaceks); 2 paratypes, 32 km
S. of Wau, Bulldog Rd., 2850 m. May 29-
30, 1962 (Sedlacek), light trap.
Additional material. Papua: 1, Mt. Tafa,
8500 ft. (c. 2600 m), Mar. 1934 (Cheesman).
N-E. N. (;.: 1 ,5, Edie Creek, 14 km S\A'
of \Yau, 2000 m, Fvh. 13, 1962 (S(>dlacek);
1 9 , Enarotadi, 2000 m, Aug. 1962 ( Sed-
lacek). West N. G.: I $, Swart Valley,
\V. ridge 1800-2000 m, Nov. 19, 1958
( Gressitt ) .
Measured specimens. The i from Edie
Creek and the 9 holotype, figures given
in this order.
Notes. The specimens before me vary not
only in size and color but also in fonn of
outer-apical elytral angles, depth of striae,
presence or absence of low rounded tuber-
cles on odd elytral intervals, and in other
ways. Some of this variation is surely in-
dividual, but some may be geographic.
Only additional series from several localities
can decide this.
Demefrida crepera n. sp.
Description. Form and characters c. as in
preceding species ( fesselata ) except color
piceous or slightly reddish piceous without
distinct pale flecks on elytra. Head 1.04
and 1.05 width prothorax. Prothorax:
width length 1.08 and 1.06; base apex 1.30
and 1.25; base head 0.85 and 0.85. Elytra:
width elytra prothorax 2.05 and 2.12; sculp-
ture somewhat variable but in general like
that in preceding species (tesselata), in
\\'hich the sculpture varies too. Secondary
sexual characters: 6 tarsi as genus; 6 mid-
dle tibiae tuberculate-serrate ( c. 6 tuber-
cles); 6 with c. 4, 9 c. 6 apical \entral
setae each side. Measurements: length 9.5-
10.4; width 3.1-3.5 mm.
Types. Holotype $ (A.M.N.H.) and 6
paratypes (2 in M.C.Z., Type No. 31,445)
from N. slope, Mt. l^ayman, Maneau Rge.,
Papua (the h()lot\pe and 4 parat\pes at
"No. 4,"' 2230 m, Ma>- 19-June 19, 1953; 1
paratype, same data except June 1-7; 1
paratype same except "No. 5," 1550 m. lune
30-Ju'lv 13) (all specimens collected 1953
by Ceoffrey M. Tate).
Additional material. N-E. N. G.: 1, "No.
10," Purosa Camp, Okapa area, 1950 m,
Sept. 29, 1959; 1, "No. 6," Pengagl Camp,
east slopes Mt. W'ilhelm, 2770 m, July 3,
1959 (both specimens Sixth Archbold Exp.,
L. 1. Hrass. A. M.N. 11.).
The Carabid Beetles of New Guinea • Darlington 153
Measured specimens. The S holotype and cles widely spaced ) ; S with 3, 9 4-6
1 5 paratype. apical ventral setae each side. Measure-
Notes. This form is apparently a geo- mcnts: length 5.6-7.6; width 2.0-2.8 mm.
graphic representative (perhaps eventually Types. Holotype i (Bishop Mus.) and
to be considered a subspecies) of the pre- 4 paratypes (2 in M.C.Z., Type No. 31,446)
ceding species (tesseJata) but is almost from Eramboe, 80 km ex Merauke, West
black rather than brown, is not distinctly N. G., holotype Feb. 1, paratypes Jan. 29,
pale-speckled, is slightly larger, and differs Feb. 5, 1960 (T. C. Maa).
slightly in proportions, especially in having Additional material Papua: 1 £ , Aroa
relatively narrower elytra. Estate, W. of Redscar Bay, 1 m, Sept. 29,
All individuals of the type series have 1958 (Gressitt); 1 $ , Bisianumu, E. of Port
moth scales stuck to them, indicating that Moresby, 500 m, Sept. 23, 1955 (Gressitt);
they were taken in light traps. 1, Daradae nr. Javarere, Musgrove R., 100
m?, Oct. 2, 1958 (Gressitt); 4, Mt. Laming-
Demetrida seriata n. sp. ton, 1300-1500 ft. {c. 400-460 m) (C. T.
Description. With characters of genus; McNamara, S. Australian Mus.).
form c. average, with prominent eyes, sub- Measured specimens. The 6 holotype and
cordate prothorax, elytra with sinuate- 1 ? paratype.
truncate apices and usually slightly nar- ^otes. This species is characterized by
rowed to\N'ard base; color brown or relatively numerous dorsal elytral seta-
testaceous; surface not obviously pubescent bearing punctures. Notable also is pres-
( actually very sparsely and inconspicuously ence of a little sparse, inconspicuous pubes-
so); reticulate microsculpture present (faint) cence (not visible on sides of head behind
only on elytra. Head 1.06 and 0.98 (some- eyes) and of several weak outward-directed
times less) width prothorax; eyes promi- hairs on margins of prothorax near anterior
nent, genae short-oblique, not prominent, angles. The species is very distinct and
Prothorax narrowly subcordate; \\'idth/ probably ranges over the whole length of
length 1.06 and 1.04 (wider in some speci- New Guinea although it has been found
mens); base/apex 1.32 and 1.30; base/head thus far only in two widely separated areas
0.87 and 0.87; sides broadly rather weakly near opposite ends of the island,
arcuate in anterior %, broadly sinuate before
c. right posterior angles; each side with Demetnda nub/co/a n. sp.
seta before middle and at posterior angle Description. See Plate 1, figure III; with
(all specimens) and additional weaker setae characters of genus; head, prothorax, and
directed more to side than upward near posterior part of elytra dark red, basal %
anterior angle; most of disc virtually im- of elytra black with black color extending
punctate. Elytra: width elytra /prothorax farther back at sides than at suture ( suture
1.92 and 1.97; apices obliquely sinuate- narrowly red), lower surface red (yellow-
truncate, with both outer and sutural angles ish on abdomen ) with metastema and con-
rounded or blunted; striae impressed, tiguous parts of epipleurae dark, femora and
scarcely punctulate; intervals convex, ir- outer edges of tibiae dark, tarsi and an-
regularly sparsely punctulate, 3rd with usu- tennae reddish yellow; not obviously pubes-
ally 6 ( sometimes fewer ) larger seta-bear- cent but with some sparse very inconspicu-
ing punctures, and 3 or 4 similar punctures ous hairs; reticulate microsculpture absent,
usually present on 5th intervals. Claws Head 1.06 width prothorax; eyes prominent,
with c. 5 teeth. Secondary sexual char- genae shorter, obliquely rounded into neck;
acters: S tarsi as genus; 6 middle tibiae front slightly convex, \\'ith 2 impressions
weakly tuberculate-serrate (c. 6 small tuber- anteriorly ( as usual in genus ) and ir-
154 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
regularly slightly impressed at middle. Pro-
thorax siibquadrate with base slightly
broader and apex narrower than usual;
width/length 1.03; base/apex 1.48; base/
head 0.92; sides weakly arcuate for much of
length, subangulate at median-lateral setae,
strongly sinuate before slightly acute promi-
nent posterior angles; margins moderate,
each with seta-bearing puncture c. % from
apex and at basal angle, and with several
much finer hairs directed laterally near an-
terior angles; disc moderately (not strongly)
convex, baso-lateral impressions present
but irregular, subpunctate. Elytra: width
elytra/prothorax 2.11; apices sinuate-trun-
cate, outer angles broadly rounded, sutural
angles blunted; striae well impressed, finely
punctulate; intervals convex, 3rd with c.
7 and 5th with 4 or 5 seta-bearing punc-
tures. Claws with c. 5 teeth. Secondary
sexual characters: i unknown; 9 with 4
apical ventral setae each side. Measure-
ments: length 7.8; width 2.8 mm.
Type. Holotype $ (Leiden Mus.) from
Lower Mist Camp [Snow Mts.], West
N. G., 1550 m, Jan. 31, 1939 (Toxopeus);
the type is unique.
Notes. This distinct species is the only
known Demetrida that combines unarmed
elytral apices with dual (black and red)
coloration. The sparse, very inconspicuous
pubescence and the extra seta-bearing
punctures of 3rd and 5th elytral intervals
are noteworthy too. The form of elytral
apices and the character of pubescence and
setae suggest a relationship with seriata,
but nuhicola is specifically distinct not only
in color but also in form of prothorax.
Demetrida magna n. sp.
Description. With characters of genus;
form large, slender, with large eyes, long-
quadrate or trapezoidal [)r()tliorax, elytra
sinuate-truncate at apex; color entirely red-
dish brown; surface not jMibescent, reticu-
late microsculpturc distinct (and sliglitly
transverse) only on elytra, punctation as
described below. Head 0.94 and 0.98
width prothorax; eyes prominent, genae
short and oblique. Prothorax long, sub-
parallel or trapezoidal; width length 0.98
and 1.00; base/apex 1.46 and 1.58; base/
head 1.01 and 1.01; sides broadly arcuate
anteriorly, usually broadly sinuate before
right or slightly acute usually blunted pos-
terior angles; margins moderate, each with
seta at basal angle and before middle; disc
fainth' or not punctulate except irregularly
subpunctate in baso-lateral depressions.
Elytra long; width elytra prothorax 1.92
and — ( elytra of 2nd specimen too spread
for measurement ) ; apices obliquely sinuate-
truncate, outer angles well defined (some-
times slightly blunted), sutural angles nar-
rowly rounded or blunted; striae impressed,
faintly punctulate; intervals convex, sparsely
punctulate, 3rd with 2 seta-bearing punc-
tures (present in all specimens but varying
in position). Cdows with c. 7 or 8 teeth.
Secondary sexual characters: S tarsi as
genus; $ middle tibiae weakly tuberculate-
serrate; £ with 3, 4, or 5 (number some-
times unsymmetric), 9 c. 6 apical ventral
setae each side. Measurements: length
10.3-12.0; width 3.3-4.2 mm.
Types. Holotype S (Bishop Mus.) from
Finschhafen, Huon Pen., N-E. N. G., 20-
150 m, Apr. 15, 1963 (Sedlacek). Para-
types as follows: N-E. N. G.: 2 ( ,$ 9 ),
Pindiu, Huon Pen., 870-1300 m, Apr. 20,
21-22, 1963 (Sedlacek, M.C.Z., Type No.
31,447); 1 9 , Adalbert Mts., Wanuma, 800-
1000 m, Oct. 25, 1958 (Gressitt); 1, Mark-
ham R., 10 m, Jan. 18, 1961 (Sedlaceks).
Papua: 1, Kokoda, 1200 ft. (366 m). May
1933 (Cheesman); 1, Owen Stanlev Rge.,
Goilala, Loloipa, Feb. 1-15, 1958 (w. W.
Brandt. Bishop Mus.); 1, Mt. Lamington,
1300-1500 ft. {c. 400-460 m) (C. T.
McNamara, S. Australian Mus.). West
N. G. : 1 ^ , Guega, \\\ of Swart Valley,
1200 m, Nov. 15, i958 (Gressitt).
Measured s])ecimens. The i holotype and
9 paratype from Adalbert Mts.
Notes. Comparative characters of mapia
are gi\(>n in tlie preceding Key. The species
The Carabid Beetles of New Guinea • Darlington 155
appears to be widely distributed at low
altitudes in New Guinea, but not common.
Demetrida truncata n. sp.
Description. With characters of genus;
form c. average, but variable; entirely
reddish brown; not pubescent, reticulate
microsculpture present ( but light and vari-
able) only on elytra, and surface not or
not much punctulate. Head 0.96 and 0.91
width prothorax; eyes moderately promi-
nent, genae much shorter, oblique. Pro-
thorax rather long, variable in shape (nar-
rowly subcordate to trapezoidal); width/
length 1.07 and 1.05; base apex 1.37 and
1.38; base/head 0.95 and 1.02; sides vari-
ably arcuate anteriorly, broadly sinuate be-
fore right or slightly acute blunted or well
defined posterior angles; margins moderate,
each usually with seta at or near basal
angle and before middle (but see Notes
below); disc smooth at middle, slightly
wrinkled or subpunctate at base and sides.
Elytra long; width elytra prothorax 1.80
and 1.78; apices obliquely sinuate-truncate,
outer angles well defined (c. right but
shghtly variable), sutural angles narrowly
rounded; striae impressed, punctulate; in-
tervals ± punctulate, often with an irregu-
lar row of small punctures near middle each
interval, 3rd with 2 dorsal punctures in all
specimens. Claws with 7 or 8 teeth. Sec-
ondary sexual characters: $ tarsi as genus;
S middle tibiae finely tuberculate-serrate
(about 9 slight tubercles); S with 3 or 4,
9 5 or 6 apical ventral setae each side.
Measurements: length 7.0-9.8; width 2.5-
3.5 mm.
Types. Holotype $ (Bishop Mus.) from
Wau, Morobe Dist., N-E. N. G., 1150 m,
Oct. 16, 1961 (Sedlacek); and paratypes as
follows. N-E. N. G.: I 6, Wau, Mt. Mis-
sim, 880-1050 m, Feb. 8-9, 1963 (Sedlacek);
1 9 , Busu R., E. of Lae, 100 m, Sept. 14,
1955 (Gressitt); 1 9, Finschhafen, Huon
Pen., 180 m, Apr. 16, 1963 (Sedlacek); 1 i ,
Torricelli Mts., Mobitei, 750 m, Feb. 28-
Mar. 4, 1959 (W. W. Brandt, Bishop Mus.).
West N. G.: 1 9 , Hollandia, Nov. 21, 1944
(H. Hoogstraal, M.C.Z.); 1 9 , Wans, S. of
Hollandia, 450-500 m, Aug. 16-23, 1959
(T. C. Maa, Bishop Mus.); 1 9, Jutefa
Bay, Pim, sea level-100 ft. (30 m), Feb.
1936 ( Cheesman ) ; 1 $ , mountain slope
above Bernhard Camp, 750 m, Mar. 1939
(Toxopeus). (Some paratypes in M.C.Z.,
Type No. 31,448.)
Additional material N-E. N. G.: 1 9,
Finisterre Rge., Saidor: Aiyawa Village,
June 16-23, 1958 (W. W. Brandt, Bishop
Mus.). West N. G.: 1 teneral 9, Hol-
landia, 250 ft. (c. 75 m), Nov. 3, 1944
(H. Hoogstraal, M.C.Z.); 1 9, Camp 1,
Mt. Xok, Waigeu Is., 2500 ft. (c. 760 m).
May 1938 (Cheesman).
Measured specimens. The i holotype and
the 9 paratype from Busu R.
Notes. This species (if it is all one
species) is widely distributed at low alti-
tudes in central and western New Guinea.
It is not recorded in Papua and may be
replaced there by the following species
(minor). Much more material from many
localities is needed to establish the specific
limits and geographic variation of these
forms.
Although truncata usually has a seta-
bearing puncture at or near each posterior
prothoracic angle, the individual from
Jutefa Bay has a well developed seta on
the right but no trace of seta or puncture
on the left, and seta and puncture are
lacking on both sides in the individual from
the Finisterre Range. The types and other
specimens listed above vary in other ways
the significance of which cannot be deter-
mined without more material. For example,
the prothorax is narrowly subcordate or
subquadrate in most of the types while
the individual from Waigeu Is. has the
prothorax strikingly trapezoidal, but the
extremes are connected by intermediates.
Demetrida minor n. sp.
Description. With characters of genus;
form as in truncata except outer apical
elytral angles c. rounded; reddish brown;
not pubescent, reticulate microsculpture in-
156 BuUetin Musciiiu of ConijMirafivc Zoology. Vol. 137. No. 1
distinct or lightly indicated on elytra, sur-
face very little punctulate. Head 0.89 and
1.02 width prothorax; eyes prominent,
genae short and oblique. Frothorax quad-
rate-subcordate; width/length 1.17 and 1.08;
base/apex 1.52 and 1.44; base/ head 1.03
and 0.97; sides broadly rounded in anterior
■")4, broadly sinuate before right-obtuse,
slightly blunted basal angles; margins mod-
erate, each with seta at base and before
middle (posterior seta-bearing puncture
present on both sides in all specimens);
disc scarcely punctate. Elytra: width
elytra/prothorax 1.71 and 2.02; apices
obliquely sinuate-truncate, outer angles
blunted or rounded, sutural angles narrowly
rounded; striae impressed, faintly or not
punctulate; intervals sparsely inconspicu-
ously punctulate, 3rd with 2 dorsal punc-
tures. Claics with c. 5 teeth. Secondary
sexual characters: i tarsi as genus; c^
middle tibiae weakly tuberculate-serrate;
S with 2, 9 4 apical ventral setae each
side. Measurements: length 5.6-6.3; width
2.0-2.2 mm.
Types. Holotype 6 (M.C.Z., Type No.
31,449) and 1 S paratype from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
1 9 paratype from Brown R., Papua, 5 m,
Oct. 23, 1960 (Gressitt).
Measured specimens. The A holotype and
9 paratype.
Notes. This species is probably related to
truncata but is smaller, with more rounded
outer apical elytral angles and fewer apical
vc^ntral setae.
Demetrida subfenuis n. sp.
Descriplion. With characters of genus;
form (Fig. 99) c. as in truncata and minor
but more slender; reddish brown; not
pubescent, reticulate microsculpture at most
faintly indicated on elytra, surface not
Diueli puiictiilate. Head 1.09 and 1.04
width prothorax; eyes proiniiiciit. genae
short and oblicjue, not jiioniinent. Pro-
tliorax el()ngate-su1)(ina(hal('; width length
0.89 and 0.94; base apex 1 .,36 and 1.40;
base/head 0.88 and 0.96; sides weakly
arcuate in much of length, broadly sinuate
well before c. right, scarcely blunted basal
angles; margins narrow, each with seta-
bearing puncture before middle but none
at base. Elytra: width elytra prothorax
2.08 and — ( elytra of 2nd specimen too
spread to measure); apices obliquely sinu-
ate-truncate, with outer angles obtuse and
slightly blunted or narrowly rounded and
sutural angles blunted; striae impressed,
faintly punctulate; intervals slightly punc-
tulate, 3rd with 2 dorsal punctures. Claics
with 5 or 6 teeth. Secondary sexual char-
acters: 6 tarsi as genus; i middle tibiae ±
bent-in and weakly tuberculate-serrate; S
with 3 apical ventral setae each side (both
sides both specimens); 9 unknown. Mea-
surements: length c. 7.0; w idth c. 2.3 mm.
Types. Holotype c5 (Bishop Mus.) from
Wum, Upper Jimmi Valley, N-E. N. G.,
840 m, July 17, 1955 (Gressitt); and 1
broken i paratype (M.G.Z., Tvpe No.
31,450), vie. Hollandia, West N. G., July-
Sept. 1944 (Darlington).
Notes. D. suhlcnuis resembles minor
(above) but is much narrower and lacks
posterior-latt>raI prothoracic setae, which
are present in minor.
Demetrida tenuis n. sp.
Dcscri})tion. With characters of genus;
form (Fig. 100) c. as in preceding species
(suhtcntii.s) but even more slender; brownish
led, legs slightly paler; not pubescent,
reticulate microsculpture visible (but very
light) only on elytra, surface not much
punctulate. IIc(ul 1.09 and 1.07 width pro-
thorax; eyes large, moderatcK' prominent,
genae much short(M-, oblicpie but conxexly
prominent; Iront carinate at middle anteri-
orl\- (all sj)i'eimens). ProtJiorax elongate-
(juadrate; width length 0.84 and 0.93; base/
apex 1.21 and 1.19; base head 0.87 and 0.83;
sides very wcakK' irregularK' anuulate, verv
broadly sinuate belore right or slightly
obtuse but well deliiicd basal angles; mar-
gins rati It r narrow, each with seta-bear-
The Carabid Beetles of New Guinea
Darlington
157
ing puncture before middle but without
posterior seta or puncture; disc faintly
pinictulate, wrinkled or subpunctate in
baso-lateral areas. Elytra: width elytra,
prothorax 2.36 and 2.32; apices sinu-
ate-truncate, outer and sutural angles ±
rounded; striae impressed, punctulate; in-
tervals slightly convex, faintly sparsely
punctulate, 3rd with 1 dorsal puncture, c.
Vi or less from apex (both sides all ex-
amples). Claws with c. 4 teeth. Secondary
sexual characters: 6 tarsi as genus; 6
middle tibiae tuberculate-serrate (4 tuber-
cles); S with 1, 9 2 apical ventral setae
each side. Measurements: length c. 6.0-
6.5; width 2.0-2.3 mm.
Types. Holotype $ ( Bishop Mus. ) from
Aroa Estate, W. of Redscar Bay, Papua,
1 m, Sept. 29, 1958 (Gressitt);' and 1 9
paratype (M.C.Z., Type No. 31,451) from
Owen Stanley Rge., Papua, Goilala: Lo-
loipa, Feb. 1-15, 1958 (W. W. Brandt); 1
c5 paratvpe, Brown R., 20 km N. of Port
Moresby, Apr. 29, 1960 (C. W. O'Brien,
Bishop Mus.).
Measured specimens. The 9 holotype and
9 paratype.
Notes. D. tenuis is characterized by small
size, very narrow form especially of pro-
thorax, carination of front, 1-punctate 3rd
elytral intervals, and small number of setae
of apical ventral segment. The last three
characters are unique among New Guinean
Demetrida, but some Australian species
have 1-punctate 3rd intervals, as indicated
in Notes under the genus and in Footnote 3
(p. 143).
Demetrida tripuncfa n. sp.
Description. With characters of genus;
form c. average, except genae angulate,
elytral apices obtusely angulate; reddish
brown; not pubescent, reticulate micro-
sculpture distinct ( but light ) only on elytra,
surface not much punctulate. Head 1.06
and 0.94 width prothorax; eyes large, promi-
nent (slightly variable), genae shghtly
shorter than eyes, subangulately prominent;
front flattened and irregularly impressed
anteriorly. Prothorax subquadrate; width/
length 1.04 and 1.06; base/apex 1.28 and
1.14; base head 0.80 and 0.87; sides
broadly irregularly arcuate in c. anterior %,
broadly sinuate before c. right but blunted
posterior angles; margins moderate, each
with seta before middle but none at
posterior angle; surface punctate-wrinkled
in baso-lateral impressions and margins.
Elytra: width elytra/prothorax — (elytra
spread) and 1.91; apices obtusely angulate,
with outer angles right or slightly obtuse,
sutural angles obtuse; striae impressed,
sometimes finely punctulate; intervals con-
vex, sparsely punctulate, 3rd with 3 dorsal
punctures (all specimens). Claws with c.
6 long teeth and sometimes an additional
minute one. Secondary sexual characters:
5 tarsi as genus; 6 middle tibiae slightly
bent-in near apex but not tuberculate-ser-
rate; i with apparently 3, 9 6-8 apical
ventral setae each side. Measurements:
length c. 8-9; width 3.0-3.3 mm.
Type. Holotype 9 (M.C.Z., Type No.
31,452) from Hollandia, West N. G., Nov.
21, 1944 (Hoogstraal).
Additional material. Papua: 1 S , Ori-
omo Govt. Sta., W. District, Oct. 26-28,
1960 (Gressitt), taken in Malaise trap; 1
9 , Brown R., 5 m, Oct. 23, 1960 (Gressitt),
taken on palm. N-E. N. G.: 1 9, Bulolo,
730 m, Aug. 15, 1956 (E. J. Ford, Jr.,
Bishop Mus.), taken in light trap.
Measured specimens. The 6 from Papua
and the 9 holotype, in this order.
Notes. The subangulate genae distin-
guish this species from all the preceding
ones except tenuis, which is very different
in many ways (see preceding Key to Species
of Demetrida).
The four specimens listed above agree in
a general way and in such important char-
acters as prominence of genae, 3-punctate
3rd intervals, and obtusely angulate elytral
apices, but they are from scattered localities
and they differ in many details. The single
6 is teneral and warped so that width of
158 Bulletin Miiscinii of Comparative Zoology, Vol. 137, No. 1
elytra cannot be measured, and some other
characters are difficult to see. More ma-
terial is needed to show whether all these
specimens really are conspecific.
Demefrida genicula n. sp.
Description. With characters of genus;
form (Fig. 101) as in preceding species
( tripuncta ) but elytral apices acutely
toothed; reddish brown; not pubescent,
reticulate microsculpture distinct (but light)
only on elytra, surface not much punctulate.
Head 1.08 and 1.03 width prothorax; eyes
prominent, genae subangulately prominent;
front flattened and irregularly slightly im-
pressed anteriorly. Prothorax quadrate-
subcordate; width length 0.97 and 1.04;
base/apex 1.24 and 1.19; base/head 0.(S3
and 0.S3; sides irregularly weakly arcuate in
anterior 'n, strongly sinuate before right or
slightly acute slightly blunted posterior
angles; margins moderate, each with seta-
bearing puncture just before middle but
none at base; disc weakly strigulose or sub-
punctate especially laterally. Elytra: width
elytra/prothorax 1.89 and 1.98; apices with
short spines or acute teeth, outer angles
sharply defined, right or slightly acute,
sutural angles obtuse; striae impressed,
scarcely punctulate; intervals convex,
slightly sparsely punctulate, 3rd usually 3-
punctate (4-punctate on left side only in in-
dividual from above Bernhard Camp).
Claw.') with 5 or 6 teeth. Secondary sexual
characters: i tarsi as genus; i middle
tibiae bent-in at apex but not tuberculate-
serrate; 6 with .3 or 4, 9 4 or 5 apical ventral
setae each side. Measurements: length 8.2-
9.2; width 2.6-3.0 mm.
Types, [lolotvpe 6 (U.S.N.M.) from
Iloliandia, West N. G. (J. W. Bongberg);
and paratypes as follows. West N. G.: 1
9 , Mountain slope above Bernhard Camp,
100 m, Apr. 19.39 (Toxopeus). N-E. N. (;.:
1 9 , Wan, Morobe Dist., 1200 m, Oct. 29,
1961 (Sedlacek); 1$, Erima, Astrolabe
Bay, 1896 (Biro). Pa|Mia: 1, Daradae,
near Javarere, Musgrove R., 100 m, Oct. 4,
1958 (Gressitt).
Additional material. N-E. N. G.: 1 9,
Tsenga, Upper Jimmi Vallev, 1200 m, Julv
14, 1955 (Gressitt).
Measured specimens. The i holotype and
the 9 paratype from Wau.
Notes. The acutely dentate or short-
spined rather than obtusely angulate elytral
apices distinguish this from the preceding
species (tripuncta). More material, espe-
cially a good series taken at one time and
place, is needed to show whether the dif-
ference is in fact specific.
Demefrida lafongula n. sp.
Description. With characters of genus;
form (Fig. 102) small and moderately broad
(in genus); reddish brown; not pubescent,
reticulate microsculpture present (some-
times faint, and slightly transverse ) only
on elytra, surface not much punctulate.
Head 0.89 and 0.91 width prothorax; eyes
prominent, genae short, not prominent;
front slightly irregularly impressed or with
punctiform impression before middle. Pro-
thorax subcordate, wide; width length 1.35
and 1.35; base apex 1..38 and 1.39; base'
head 0.97 and 0.94; sides broadly rounded
anteriorly, sinuate before well defined right
(sometimes slightly obtuse or acute) basal
angles; margins moderately wide, each with
seta before middle but none at base; disc
sometimes slightly wrinkled or subpunctate
basally and laterally. Elytra rather short
and broad (in genus); width elytra pro-
thorax 1.85 and 1.78; apices obtusely an-
gulate, with outer angles obtuse and usually
slightly blunted, sutural angles blunted;
striae imjiresscHl, \ aguely or not punctulate;
inter\'als convex, si)arsel\- pimctulate, 3rd
with 2 dorsal punctures. Clans with 3 or
4 teeth (and sometimes a small 5th one).
Secondary sexucd characters: i tarsi as
genus; ,5 middle tibiae strongK' tubercu-
late-serratc>; .s w ith 2 or 3, 9 3 or 4 apical
\ciitral setae each side. Measurements:
length 5.5-7.1; width 2.3-2.9 mm.
Types. Ilolotype $ (Bishop Mus.) from
Bisianumu, Iv of Port Mor(\sbv, Pajma,
500 m, Sejit. 3, 1959 (T. C. Maa); and
The Carabid Beetles of New Guinea • Darlington
159
paratypes as follows. Papua: 1 ? , Brown
R., E. of Port Moresby, 100 m, June 8,
1955 (Gressitt, now in M.C.Z., Type No.
.31,453); 1 9, Brown R., May 21, 1956
(E. J. Ford, Jr., Bishop Mus.); 1, Mt.
Lamington, 1300-1500 ft. (c. 400-460 m)
(C. T. McNamara, S. Australian Mus.); 1,
Buna Bay (C. T. McNamara, S. Australian
Mus.). N-E. N. G.: 1 9, Huon Pen.,
Pindiu, Apr. 20, 1963 (Sedlacek). West
N. G.: 1 c^ , Maffin Bay, Sept. 1944 (E. S.
Ross, Cal. Acad.).
Measured specimens. The i holotype and
1 9 paratype from Brown R.
Notes. This apparently widely distributed
lowland species is characterized by small
size, relatively broad form, and obtuse an-
gulation of elytral apices. See preceding
Key to Species of Demetrida for further
differential characters.
Demetrida angulata n. sp.
Description. With characters of genus;
form slender-average, with obtusely angu-
late elytral apices; reddish browai; not pubes-
cent, reticulate microsculpture distinct (light
and usually slightly transverse) only on
elytra, surface not much punctulate. Head
1.07 and 1.08 width prothorax; eyes promi-
nent, genae short, oblique, not prominent.
Prothorax subquadrate, narrow; width/
length 1.01 and 1.04; base/apex 1.40 and
1.31; base/head 0.88 and 0.87; sides weakly
irregularly arcuate (sometimes almost par-
allel) in c. anterior %, broadly sinuate be-
fore ± right but blunted posterior angles;
margins moderate, each with seta-bearing
puncture before middle but none at base;
disc variably wrinkled or subpunctate pos-
teriorly and laterally. Elytra: width elytra
prothorax 1.88 and 2.08; apices obtusely
angulate, outer angles sharply defined but
varying from slightly obtuse to acute,
sutural angles blunted; striae impressed,
usually faintly punctulate; intervals convex,
sparsely inconspicuously punctulate, 3rd
with 2 dorsal punctures. Claws with 5 or
6 teeth. Secondary sexual characters: S
tarsi as genus; i middle tibiae scarcely
modified, slightly bent-in near apex, not
distinctly tuberculate-serrate; S with 3 or 4,
9 5 or 6 setae each side last ventral
segment. Measurements: length 7.5-8.9;
width 2.5-3.1 mm.
Types. Holotype 6 (Bishop Mus.) from
Brown R., Papua, Sept. 30, 1959 (T. C.
Maa), taken sweeping; and paratypes as
follows (some in M.C.Z., Type No. 31,454).
Papua: 1 9 , same data as type except
dated Aug. 30, 1959; 1 9 , Brown R., E. of
Port Moresby, 100 m, June 8, 1955 (Gres-
sitt); 1, same locality, Apr. 27, 1960 (C.
W. Obrien, Bishop Mus.); 1 9, Laloki, nr.
Port Moresby, Aug. 30-Sept. 2, 1959 (T. C.
Maa, Bishop Mus.); 1 9, Kiunga, Fly R.,
July 11-14, 1957 (W. W. Brandt, Bishop
Mus.); 1 9, Daradae, nr. Javarere, Mus-
grove R., 100 m, Oct. 2, 1958 (Gressitt).
Measured specimens. The i holotype and
1st 9 paratype from Brown River.
Notes. See Key to Species of Demetrida
of New Guinea for distinguishing charac-
ters of an<i,ulata. It is the only New Guin-
ean Demetrida known to occur also in
Australia ( 1 9 , Rocky R., mid-peninsular
Cape York). In New Guinea, it has been
found only in Papua.
Demetrida reversa n. sp.
Description. With characters of genus;
form of preceding species (angulata) but
slightly less narrow; reddish brown; not
pubescent, reticulate microsculpture distinct
(and somewhat transverse) only on elytra,
but much of surface sparsely punctulate.
Head 1.03 and 1.07 width prothorax; eyes
prominent, genae short and oblique, not
prominent. Prothorax subquadrate; width/
length 1.10 and 1.12; base apex 1.32 and
1.31; base head 0.94 and 0.98; sides nearly
straight and subparallel or slightly con-
verging anteriorly, subangulate at setae,
broadly sinuate before right or slightly
acute basal angles; margins moderate, each
with seta-bearing puncture at or before
middle but none at base; disc slightly ir-
regularly subpunctate at base and laterally.
Elytra: width elytra/prothorax 1.85 and
160
Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
1.83; apices ungulate (the angles c. right,
but variable), outer angles right or acute,
sharply formed; sutural angles obtuse-
blunted; striae impressed, faintly punc-
tulate; intervals sparsely punctulate, 3rd
with 2 dorsal punctures. Claws with 6 or 7
teeth. Secondary .sexual chaiacter.s: S tarsi
as genus; i middle tibiae slightly bent out
near apex, inconspicuously or irregularly
tuberculate-serrate (Fig. 162); i with c.
3, 9 c. 6 setae each side last ventral seg-
ment. Measurements: length 8.5-9.2; width
3.0-3.4 mm.
Types. Holotype 6 ( Bishop Mus. ) and
7 paratypes (3 in M.C.Z., Type No. 31,455)
from Guega, W. of Swart Valley, West
N. G., 1200 m, Nov. 14, 15, 1958 (Gressitt),
and 1 paratype, Swart Valley, W. Fork,
1300-1350 m, Nov. 17, 1958 (Gressitt).
Measured specimens. The 6 holotype and
1 9 paratype from Guega.
Notes. Among similar species with angu-
late but not spined elytral apices, this is
distinguished by quadrate prothorax, pro-
portions as given, and especially by form
of 6 middle tibiae, slightly bent outaard
at apex. Nevertheless the present species
may be closely related to the preceding one
[an^ulata), which is known only from
Papua (and Australia), while the present
one is known only from a restricted area
of West New Guinea.
Demefrida kokoda n. sp.
Descri])(ion. With characters of genus;
form as in Figure 103, large, slender;
reddish brown; not pubescent, reticulate
microsculpture present (light or iaint) onK-
on elytra, surface not much |)unctulate.
Head 0.99 and 0.99 width prothorax; exes
slightK' smaller than usual but prominent,
genae scarcely distinct from neck. Prothorax
cordate-subquadrate; widtli knigth 1.11 and
1.12; base/apex 1.22 and 1.23; base head
0.84 and 0.87; sides strongh' rounded in
anterior %, strongly sinuate before c. right
or slightly acute but blunted [)()st('rior
angles; margins narrow, each with seta
slightly before middle but none at base;
disc with middle line finer than usual in
genus, baso-lateral areas slightly punctate.
Elytra very long; width elytra ^ prothorax
1.72 and c. 1.86 (elytra spread); apices
with moderate spines, outer angles c. right
or slightly acute, sharply formed, sutural
angles slightly obtuse, sometimes denticu-
late; striae impressed, punctulate; intervals
only slightly convex, scarcely punctulate,
3rd with 2 seta-bearing punctures. Claws
with 7 or 8 teeth. Secondary .sexual char-
acters: 6 tarsi as genus; i middle tibiae
weakly tuberculate-serrate; i with c. 4, 9
numerous (up to 9) apical ventral setae
each side. Measurements: length c. 10.0-
11.0; width 2.9-3.5 mm.
Types. Holotype i ( British Mus. ) and
5 paratypes (2 in M.C.Z., Type No. 31,456)
from Kokoda, Papua, 1200 ft. (366 m).
May, Aug. (holotype), Sept., Oct., 1933
( Cheesman ) ; 1 paratype, Popondetta,
Papua, 25 m, June 1966 ( Shanahan-Lip-
pert. Bishop Mus.).
Additional material. N-E. N. G.: 1 9,
Wau, Morobe Dist., 1050 m, Apr. 30, 1962
(Sedlacek). West N. G.: 1 9, Waris. S.
of Ilollandia, 450-500 m, Aug. 24-31, 1959
(T. G. Maa, Bishop Mus.); 1 9 , Hollandia.
J:m. 1945 (B. Malkin, U.S.N.M.).
Measured sj)ecimens. The c5 holotype and
1 9 paratype.
Notes. The form ol this species is uni({ue
in the genus, so far as I know, and other
characters including the relatixely fine
middle line ol the pronotum are distinctixe.
Demetrida moda n. sp.
Description. See Phite 1, figure I\'; with
characters of genus; reddish brown; not
pubesci'ut, reticulate microsculpture distinct
(but light) onl\' on elytra, parts ol upper
siu"lace sparsely j)unctulate. Head 0.85
and 0.84 width prothorax; e\'es prominent,
genae short, obliciue. not jirominent. Vro-
thorax subcordalc; width length 1.46 and
1.33; base /apex 1.40 and 1.37; base/head
1.02 and 1.04; sides arcuate^ anteriorly,
strongly sinuate belorc sharply defined right
or acute posteiioi- angles; margins moderate.
The Carabid Beetles of New Guinea • Darlington
161
each with seta before middle but none at
base; disc slightly punctate basally. Elytra:
width elytra prothorax 1.69 and 1.77; apices
with moderate spines, outer angles obtuse,
sutinal angles obtuse; striae impressed,
faintly punctulate; 3rd intervals 2-punctate.
Claws with c. 4 teeth. Secondary sexual
characters: 6 tarsi as genus; 6 middle
tibiae strongly tuberculate-serrate ( c. 4
rounded tubercles, Fig. 160); i with 2, 9
3 apical ventral setae each side (number
may vary). Measurements: length c. 5.5-
6.5; width 2.1-2.5 mm.
Types. Holotype S (M.C.Z., Type No.
31,457) and 4 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington), and
additional paratvpes as follows. Papua: 8,
Kokoda, 1200 ft.' (366 m), Aug., Sept., Oct.,
1933 (Cheesman). N-E. N. G.: 2, Sattel-
berg, Huon Gulf, 1899 (Biro).
Additional material N-E. N. G.: 2,
Pindiu, Huon Pen. (1 labeled 500-600 m),
Apr. 19, 20, 1963 (Sedlacek); 1, Finschhafen,
May 7, 1944 (E. S. Ross, Gal. Acad.); 1,
Bubia, Sept. 1949 ( N. L. H. Krauss, Bishop
Mus.).
Measured specimens. The 6 holotype and
1 9 paratype from Dobodura.
Notes. This species, as the name moda
is intended to suggest, is the first of several
generally similar forms which differ among
themselves slightly in proportions and color
and more significantly in length of elytral
spines and punctation of 3rd elytral inter-
vals. Some of these forms may be geo-
graphically limited and allopatric and may
eventually be considered subspecies. The
present species seems to be confined to the
eastern half of New Guinea. Its differential
characters are given in the preceding Key
to species.
Demefrida submoda n. sp.
Description. With characters of genus;
form of preceding species (rnoda) except
elytra acutely toothed, not spined, and
proportions slightly different, with head rel-
atively slightly wider and base of prothorax
narrower; color, microsculpture, etc. as in
moda. Head 0.89 and 0.91 width prothorax;
eyes prominent, genae short and oblicjue.
Prothorax subcordate; width/length 1.40
and 1.37; base/apex 1.43 and 1.45; base/
head 0.97 and 0.95; sides rounded anteriorly,
strongly sinuate before c. right posterior
angles; margins rather wide, each with seta
near or before middle but none at base;
disc slightly irregular or subpunctate baso-
laterally. Elytra: width elytra prothorax
1.70 and 1.81; apices acutely angulate or
denticulate, outer angles obtuse or blunted,
sutural angles obtuse; striae impressed, not
distinctly punctulate; intervals convex,
slightly inconspicuously punctulate, 3rd 2-
punctate. Claws with c. 5 teeth. Secondary
sexual characters: 6 tarsi as genus; 6 mid-
dle tibiae tuberculate-serrate (4 tubercles);
6 with 2, 9 3 apical ventral setae each
side. Measurements: length 6.0-7.0; width
2.3-2.8 mm.
Types. Holotype S (Bishop Mus.) and
17 paratypes (7 in M.G.Z., Type No.
31,458) from Wau and vicinity (including
Mt. Missim), Morobe Dist., N-E. N. G.,
altitudes from 950 to 1400 m, dates in Jan.,
Feb., Mav, Julv, Aug., Sept., Nov., Dec,
1961-1964 (Sedlaceks, T. G. Maa) (holo-
type, 1250 m. May 3, 1963); and additional
paratvpes from N-E. N. G. as follows: 2,
Bulolo, 1065 m, Aug. 15, 16, 1956 (E. J.
Ford, Jr., Bishop Mus.); 1, Upper Watut
R., 24 km W. Bulolo, 760 m. Mar. 5-6,
1963 (Sedlacek).
Additional material. N-E. N. G.: 1, vie.
Nadzab, July 1944 (Darlington); 2, Kas-
sem, 48 km E. of Kainantu, 1350 m, Nov.
7, 1959 (T. G. Maa, Bishop Mus.); 1,
Kumun, Upper Jimmi Valley, 1000 m, July
13, _ (Gressitt); 1, Maprik, 150 m, Dec.
29-Jan. 17, 1960 (T. G. Maa, Bishop Mus.);
1, Eliptamin ^^^, 1200-1350 m, July 16-31,
1959 (W. W.' Brandt, Bishop Mus.); 1,
Goroka, 1550 m, June 19, 1955 (Gressitt),
"pigeon peas cane." West N. G.: 1, Hol-
landia, Dec. 15, 1944 ( Hoogstraal, M.G.Z.).
Measured s})ecinwns. The 6 holotype and
1 9 paratype from Wau.
Notes. This species differs from moda
162 Bulletin Museum of Coinparative Zoology, Vol. 137, No. 1
as indicated in tht' preceding Description.
It may prove to be only a subspecies of
moda, and has thus far been found only in
the central-eastern part of the north side
of New Guinea, chieflv in the lower moun-
tains.
Demetrida hollandia n. sp.
Description. With characters of genus;
form c. of moda and suhmoda (above) but
color darker, reddish brown with elytra
darker brown or brownish black with apical
% or less often paler, the pale apical area
varying in distinctness and extent; micro-
sculpture, etc. c. as in moda, with reticula-
tions faint but usually visible on elytra.
Head 0.89 and 0.(S8 width prothorax, eyes
prominent, genae short. Prothorax sub-
cordate, slightly narrower than in moda;
width length 1.32 and 1.31; base/apex 1.36
and 1.34; base head 0.97 and 0.95; sides
slightly irregularly rounded, often subangu-
late at lateral setae; margins moderately
wide, each with seta near or before middle
but none at base; siuface not or ver\ Httle
punctate. Elytra: width elytra prothorax
1.86 and 1.85; apices short-spined, outer
angles obtuse but more distinct than in
moda, sutural angles blunted; striae im-
pressed, not distinctly punctulate; intervals
convex, scarcely punctulate, 3rd with 2
principal punctures and often (not alwa\s)
with 1 or more intermediate i^unctures
which vary in size and sometimes do and
sometimes do not bear setae. CUaus with
c. 4 te(>th. Secondary sexital characters: i
tarsi as genus; i middle tibiae strongly
tuberculate-serrate (c. 4 tubercles); i with
2, 9 3 apical ventral setae each side. Mea-
surements: length 5.8-7.6; width 2.1-2.9
mm.
Types. Tlolotype S (M.C.Z., Type No.
31,459) and 33 paratypes from vie. ITol-
laudia. West N. (i., July-Sept. 1944 (Dar-
lington ).
Addilional material. West N. (',.: 11,
Hollandia and vicinity including C]yclops
Mts., at low altitudes (not over 500 m),
\ari()us dates and collectors; 10. Maffin
Bav, dates in June, July, Aug., Sept., Oct.
1944 (E. S. Ross, Cal. Acad.); 5, Nabire,
S. Geelvink Bay, 5-50 m, Aug. 25-Sept. 5.
1962 (Sedlacek); 1, \\'a.sian (Vogelkop),
Sept. 1939 (Wind, M.C.Z.); 1, Fac Fac,
June 1939 (Wind, M.C.Z.). N-E. N. G.:
33, various localities including Fluon Pen.;
Torricelli Mts.; Sepik Dist.; Wewak; Lae;
Bulolo; Wau. Papua: 1 teneral 6 , doubt-
fully identified, from Bisianumu, E. of Port
Moresby, 500 m, Sept. 24, 1955 (Gressitt).
Measured specimens. The c5 holotype and
1 9 paratype.
Notes. D. hollandia seems closely related
to moda and suhmoda but is distinguished
by characters given above. Some of the
specimens listed under Additional material
\ar\- toward icaii (see below).
Demetrida wau n. sp.
Description. With characters of genus;
form c. as in moda, sidmwda, and hollandia
but more slender with eyes less prominent
and genae slightly longer and less abrupt
than in the species named; reddish brown,
eU'tra brownish black, usualK' not paler
at apex; not pubescent, upper surface in-
cluding elytral disc without reticulate micro-
sculpture but in part (especialK- cKtra)
sparseK' punctulate. Head 0.89 and 0.89
width prothorax; e>'es and genae as indi-
cated abo\e. Prothorax subcordate; width/
length 1.21 and 1.22; base apex 1.44 and
1.44; base head 0.99 and 0.99; sides rounded
anteriorly, often subangulate at setae (as in
hollandia), strongly sinuate before c. right
basal angles; margins rath(M- wid(\ each
with seta near or beloic middle but none
at base; disc more punctate basalh' and
laterally than in the '^^ preceding species.
Elytra: width elytra prothorax 1.7.3 and
1.73; a|Mees short-spined (or with long
acute teeth), with outer angles obtuse
but distinct, sutural angles blunted; striae
impressed, punctulate; interxals eou\('x,
punctulate. 3i(l olten 4-i')unetate but inter-
mediate |iuiutui('s \aiiabl(' in si/e and
sometimes absent and with oi' without
setae. Clans with c. ,5 teeth. Secondary
The Carabid Beetles of New Guinea • Darlingtofi
163
sextial cJiamcters: 6 tarsi as genus; 6
middle tibiae weakly tuberculate-serrate (c.
3 tubercles distinct); 6 with 2, 9 3 apical
ventral setae each side. Measurements:
length 6.4-7.9; width 2.4-2.9 mrn.
Types. Holotype c^ (Bishop Mus.) and
118 paratypes (some in M.C.Z., Type No.
31,460) all from Wau, Morobe Dist., N-E.
N. G.; altitudes from 1000 to 1450 m;
dates in everv month, 1961-1963 (holotvpe,
1200 m, July '22, 1961 ) ( Sedlaceks ) .
Additional material. N-E. N. G. : 9 addi-
tional teneral, broken, or atypical speci-
mens from Wau; 1, Jim(m)i R., E. High-
lands, Julv-Sept. 1961 (W. W. Brandt,
C.S.I.R.O.); 1, Upper Watut R., 24 km W.
Bulolo, 760 m, Mar. 5-6, 1963 (Sedlacek);
1, Erima, Astrolabe Bay, 1897 (Biro). West
N. G.: 2, Hollandia, May, June 1945 (B.
Malkin, U.S.N.M.); 1, Waris, S. of Hol-
landia, 450-500 m, Aug. 16-23, 1959 (T. C.
Maa, Bishop Mus.); 1, Ifar, 400-550 m,
June 23, 1959 (T. C. Maa, Bishop Mus.).
Measured specimens. The S holotype and
1 9 paratype.
Notes. D. wau may be primarily a geo-
graphic representative of liollandia, but the
long type series seems distinct; the speci-
mens listed above from Hollandia and
Waris are plainly wau, not hollandia; and
the one from Ifar seems to be wau except
that the elytra are distinctly microreticulate.
Apparent intermediates do occur at some
other localities, however. They are tenta-
tively placed with Additional material un-
der hollandia. See also Notes under D.
subpunctata (3rd species below).
One 9 of wau, from Wau, is a note-
worthy abnormalit)', \\4th the posterior pro-
thoracic angles irregularly widened and
each \\'ith 2 setae, although normal in-
dividuals of wau lack posterior-lateral
setae.
Demefrida similis n. sp.
Description. With characters of genus;
form of moda, etc. but larger, rather slender,
with prothorax narrowly subcorclate and
elytra spined or acutely dentate and with
outer angles sharply formed; reddish
brown, elytra not or only slightly darker;
not pubescent; reticulate microsculpture
visible (often faint) only on elytra. Head
0.87 and 0.90 width prothorax; eyes promi-
nent, genae shorter, oblique. Prothorax
subcordate; width/length 1.26 and 1.22;
base/apex 1.35 and 1.34; base/head 0.98
and 0.98; sides broadly sometimes slightly
irregularly arcuate in more than % of length,
strongly sinuate before right or slightly
acute usually slightly blunted posterior
angles; margins rather wide, each with
seta-bearing puncture before middle but
none at base; disc slightly punctate at
base and sides. Elytra long; width elytra/
prothorax 1.61 and 1.69; apices short-
spined (rarely only acutely toothed), outer
angles well formed, varying from slightly
obtuse to acute, sutural angles blunted-
obtuse; striae impressed, finely punctulate;
intervals slightly convex, sparsely finely
punctulate, 3rd 2-punctate ( all specimens ) .
Claws with 6 or 7 teeth. Secondary sexual
characters: 6 tarsi as genus; S middle
tibiae tuberculate-serrate (c. 4 tubercles);
S with 2, 9 3 or 4 apical ventral setae each
side. Measurements: length 8.8-10.8; width
3.0-3.5 mm.
Types. Holotype £ (M.C.Z., Type No.
31,461 ) and 3 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
additional paratvpes as follows. Papua: 4,
Kokoda, 1200, 1300 ft. {c. 366, 400 m),
June, Aug., Sept. 1933 (Cheesman), 1
labeled also 'Tn fungus. A, & under bark
behind it," and 1 "At light"; 1, same locality,
380 m. Mar. 20, 1956 (Gressitt), in light
trap; 2, Kokoda-Pitoki, 400 m. Nhu". 23,
1956 (Gressitt); 2, Mt. Lamington, 1300-
1500 ft. (c. 400-460 m), (C. T. McNamara,
S. Australian Mus.).
Additional material. Papua: 1, Kiunga,
Fly R., Oct. 1-7, 1957 (^^^ ^^^ Brandt,
Bishop Mus.). N-E. N. G.: 1, Ebabaang,
Mongi W\itershed, Huon Pen., 1300-1400
m, Apr. 16-18, 1955 ( E. O. Wilson, M.C.Z.).
Measured specimens. The i holotype and
1 9 paratype from Dobodura.
164 Bulletin Museum of Couiparativc Zoology, Vol. 137, No. 1
Notes. Characters distinguishing this
species from moda, etc. are given in the
Description, above; and see also Notes
under the following species.
Demefrida duplicafa n. sp.
Description. With characters of genus;
form c. as in similis (above); reddish
brown, elytra not or not much darker; not
pubescent, reticulate microsculpture visible
only on elytra, more transverse than in
similis, surface in part sparsely punctulate.
Head 0.92 and 0.91 width prothorax; eyes
prominent, genae shorter, oblique. Pro-
tJiorax quadrate-subcordate; width length
1.27 and 1.29; base/apex 1.31 and 1.31;
base/head 0.94 and 0.96; sides (usually a
little irregularly) rounded anteriorly, sinu-
ate before c. right but usually blunted pos-
terior angles; margins rather wide, each
with seta at or slightly before middle but
none at base; disc not much punctate
even basally. Elytra: width elytra/ pro-
thorax 1.71 and 1.75; apices with moder-
ate spines, outer angles sharply defined
and sometimes acutely denticulate, sutural
angles blunted-obtuse; striae moderately
impressed, scarcely punctulate; intervals
slightly convex, 3rd with 2 principal and
usually one or more smaller intermediate
dorsal punctures. Claws with c. 5 teeth.
Secondary sextial characters: i tarsi as
genus; 6 middle tibiae tubereulate-serrate
{c. 6 small tubercles); 6 with 2, 9 3
apical ventral setae each side. Measiire-
incnts: length c. 8.0-9.0; width 2.9-3.3
mm.
ry))es. llolotype $ (M.C.Z., Type No.
31,462) and 10 paratypes from Dobodiua,
l^apiia, Mar.-July 1944 ( Darhiiglon ); 3
[)aratyp('s bom Kokoda, Papua, 1200 It.
(366 m), Apr., June, Aug. 1933 (Cheesman);
94 paratypes, Mt. Lamington, Papua, 1300-
1500 ft. {c. 400-460 m) (C. T. McNamara,
S. Australian \bis. ).
AddHio)ud nialcriid. Si.\t)-one (includ-
ing 44 from W'au), from 12 localities, in all
3 political divisions of New (guinea ( from
Milne lia\ to mountain slojX' al)o\c l^crn-
hard Camp), altitudes from near sea level
to at least 1500 m (at Wau), various dates
and collectors.
Measured specimens. The i holotype and
1 9 paratype from Dobodura.
Notes. D. duplicata is much like similis
and occurs at some of the same localities
l)ut differs constantly (at least at Dobodura)
by having the elytra more distinctly and
more transversely microreticulate, with 3rd
intervals with more than 2 dorsal punctures.
The additional punctures vary in size and
sometimes do and sometimes do not bear
setae.
At Dobodura, duplicata (like similis) is
uniformly brown, but individuals with base
of elytra ± darker occur with brown in-
di\ iduals at many localities including Wau.
The color is not obviously dimorphic but
apparently continuously variable. A re-
lated population in \\hich the elytra are
always dark at base occurs in West N. G.
(see hasalis, p. 172). The specimens sum-
marized above under Additional material
vary in other ways which cannot profitably
be discussed in detail here.
Demefrida subpuncfafa n. sp.
Description. ^Vith characters of genus;
fomi c. as in moda, udti. etc., but slightly
more slender; dark reddish brown, elytra
darker (dark castaneous), legs browii; not
pubescent; microsculpture \isible ( faint,
distinctK' trans\cM"se ) ouK on eUtra, but
much of upper surface finely sparsely
punctulate. Head 1.00 and 0.96 width i^ro-
thorax; eyes moderate, genae slightb'
shorter, obliciue. Prothorax narrowly sub-
cordate; width length 1.16 and 1.16; base
apex 1.37 and 1.3(S; base head 0.93 and
().9(S; sides weakly irregularly arcuate in
anterior "i or more, strongK' sinuate before
right or slightly acute i)()sterior angles;
margins narrower than in moda and irau.
each with seta before middle but none at
base; sinface rather closely punctate across
base and in margins. Elytra: width elytra
prothorax 1.8(S and I.SS; ai:)ices with short
spines, outer angles distinct but obtuse ami
The Carabid Beetles of New Guinea • Darlington 165
sometimes slightly blunted, sutural angles
blunted or narrowly rounded; striae im-
pressed, faintly punctulate; intervals con-
vex, 3rd usually 3- ( rarely 4- ) punctate but
intermediate puncture(s) variable in size
and sometimes indistinguishable. Claws
with c. 4 teeth. Sccondanj sexual characters:
i tarsi as genus; i middle tibiae tubercu-
late-serrate ( c. 4 rounded tubercles ) ; S
with 2, 9 3 apical ventral setae each side.
Measurements: length 6.7-7.6; width 2.3-
2.7 mm.
Types. Holotype S (M.C.Z., Type No.
31,463) and 5 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
additional paratypes from Papua as fol-
lows: 7, Kokoda, 1200 ft. (366 m), May,
June, Julv, Aug., Oct. 1933 (Cheesman);
5, Kokoda-Pitoki, 450 m. Mar. 24, 1956
(Gressitt); 1, Bisianumu, E. of Pt. Moresby,
500 m, Sept. 23, 1955 (Gressitt); 1, "Papua,"
(Hungarian Nat. Mus. ); 3, Mt. Lamington,
1300-1500 ft. (c. 400-460 m) (G. T. Mc-
Namara, S. Australian Mus.).
Measured specimens. The S holotype and
1 9 paratype from Dobodura.
Notes. This may ( or may not ) be the
Papuan representative of the hollandia-icau
group of central and western New Guinea.
D. suhpunctata most resembles wau but is
slightly more slender, with narrower pro-
thoracic margins ( which distinguish it also
from hollandia), and with distinct elytral
microsculpture.
Demetrida dobodura n. sp.
Description. With characters of genus;
form c. as in moda and similis but slightly
more slender; brown (not dark), elytra not
or not much darker; not pubescent, reticu-
late microsculpture visible ( light or faint )
only on elytra, surface (except of elytra)
not much punctulate. Head 0.98 and 0.96
width prothorax; eyes prominent, genae
short, oblique. Prothorax subquadrate;
width/length 1.11 and 1.12; base/apex 1.34
and 1.30; base/head 0.95 and 0.96; sides
weakly irregularly arcuate in % or more of
length, weakly sinuate before c. right but
blunted posterior angles; margins narrower
than in moda and .similis, each with seta at
or slightly before middle but none at base;
surface weakly punctate across base and in
margins. Elytra: width elytra/prothorax
l.ScS and 1.87; apices with moderate spines,
outer angles acutely denticulate (or right
but sharply formed in some individuals
listed under Additional material), sutural
angles blunted-obtuse; striae moderately im-
pressed, finely punctulate; intervals slightly
convex, punctulate, 3rd with 2 dorsal punc-
tures. Cdans with c. 5 teeth. Secondary
sexual characters: i tarsi as genus; i
middle tibiae tuberculate-serrate (3 or 4
\\'ell spaced small tubercles ) ; S with 2 or
3, 9 5 to 7 apical ventral setae each side.
Measurements: length 7.7-9.2; width 2.6-
3.3 mm.
Types. Holotype S (M.C.Z., Type No.
31,464) and 21 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
9 paratypes, Kokoda, Papua, 1200 ft. (366
m), Aug., Sept. 1933 (Gheesman).
Additional material. Papua: 1, Kiunga,
Fly R., Aug. 8-10, 1957 (^^^ \\'. Brandt,
Bishop Mus.); 1, Koitaki, 1500 ft. (c. 450
m), Oct.-Nov. 1928 (Pemberton, H.S.P.A.).
N-E. N. G.: 1, Pindiu, Huon Pen., 500-600
m, Apr. 19, 1963 (Sedlaeek). Also 1 old
specimen, i , labeled "New Guinea. Sayer,"
"probably N. gen. near Euproctus," and
"Gen. probably near Gtenodactylus" (the
last 2 labels probably by Andrewes ) .
Measured specimens. The i holotype and
1 9 paratype from Dobodura.
Notes. Among other brown, spined
Demetrida. this should be recognizable by
prothorax subquadrate c. wide as head and
slightly wider than long by measurement,
by the rather large size, and the 2-punctate
3rd elytral intervals. See also Notes under
following species (kiunga).
The specimens listed under Additional
material are doubtfully identified. D.
dobodura is therefore known with certainty
only from Dobodura and Kokoda, in
Papua.
166 BiilJctin Mu-scmn of Companitive Zoology, Vol. 137, No. 1
Demefrida kiunga n. sp.
Description. With characters of genus;
form c. of preceding (dohocJum) but
larger; reddish brown, elytra slightly but
not much darker; not pubescent, reticulate
microsculpture distinct only on elytra, sur-
face not much punctulate. Head 1.04
and 1.00 \\idth prothorax; eyes moderately
prominent, genae shorter and oblique.
Pmtliorax subquadrate; width length 1.00
and 1.02; base/apex 1.31 and 1.24; base/
head 0.90 and 0.89; sides very weakly
arcuate anteriorly, slightly subangulate at
setae, strongly sinuate well before slightly
acute sometimes slightly blunted basal
angles; margins rather narrow, each with
seta at or before middle but none at base;
disc slightly irregular or subpunctate at base
and in margins. Elytra long; width elytra/
prothorax 1.87 and 1.80; apices long-spined,
outer angles c. right and sharply formed
but not denticulate, sutural angles obtuse;
striae impressed, faintly punctulate; inter-
\als slightly convex, 3rd 2-punctate. Claws
with c. 7 teeth. Secondanj sexual ehar-
aeters: S tarsi as genus; c^ middle tibiae
unmodified, virtually straight, not tubercu-
late-serrate; i with 3 or 4,9 4 or 5 ( un-
symmetric in both individuals ) setae each
side last ventral segment. Measurements:
lengdi 10.8; width 3.4 mm.
Tijpes. Holotype <^ (Bishop Mus.) and
1 9'paratype (M.C.Z., Type No. 31,465)
from Kiunga, Fly R., Pajma, Aug. 14-17,
18-23, 1957 (W. W. Brandt).
Notes. Among other jilain reddish brown
Demetrida with (piadrate prothorax and
spined elytra, this is distinguished by rel-
atively large si/e, proportions, simple $
middle tibiae, and oilier characters given
in the Key to Species.
As compared with dohodtira, the present
species is larg(>r, with longer elytral spines
but less produc{>d outer elytral angles, as
well as with differcMit ^ tibiae. The single
individual of dobodura seen from Kiunga
is a i with all the characters of dobodura:
smaller si/.e, denticulate outer elytral angles.
and plainly tubcrculate-serrate middle
tibiae.
Demefrida mafuiu n. sp.
Description. See Plate 2, figure V; with
characters of genus; color dimorphic, either
dark red with prothorax and basal -.-. of
elytra green with green color extending
farther back at sides than at suture or
entirely irregular dark reddish brown, legs
either dark with paler tarsi or entirely
brown, antennae brown in both cases; not
pubescent, reticulate microsculpture pres-
ent, c. isodiametric on head and elytra and
transverse on pronotum, surface not much
punctulate. Head 1.11 and 1.07 width pro-
thorax; eyes prominent, genae shorter,
oblique, not prominent; front flattened and
irregularly slightly impressed before mid-
dle. Prothorax subquadrate; width length
0.96 and 0.99; base apex 1.35 and 1.35;
base/head 0.84 and 0.87; sides weakly
arcuate for much of length, scarcely angu-
late at setae, sinuate before prominent c.
right basal angles; margins narrow, each
with seta-bearing puncture slighd>' before
middle but none at base; disc more convex
than usual, baso-lateral impressions irregu-
larly punctate. Elytra: width elytra/pro-
thorax 1.96 and 2.03; apices acutely angu-
late or dentate, outer angles sharply formed,
c. right or obtuse, sutural angles obtuse-
striae well impressed, fineU' punctulate;
intervals slighth' con\ex, faintl)' sparsely
punctulate, 3rd 2-punctate. Clatvs \\'ith c.
6 t(>eth. Secondary sexual characters: $
tarsi as genus; S middle tibiae hcut in at
apex but not tubcrculate-serrate; S with
4, 9 c. 5 apical ventral setae each side.
Measuremenis: length 9.3; width 2.9 mm.
Type. Ilolotvpc $ (British Mus.) from
Mafuiu, Papua, 4000 ft. (1220 m), Dec.
1933 (Cheesman); and 1 9 paratype (also
British Mus.) widi .same data v\cv\A datcnl
Ian. 1934.
Notes, (.'omparisoii with direr.sa (p.
172) suggests that the color dimorphism ot
nuifulu is simply MeiuU'lian. not sexual.
These 2 species max l)e related, but mafuUi
The Carabid Beetles of New Guinea • Darlington 167
seems surely distinct ])y form, greater
convexity of pronotum, and more distinct
reticulate microsculpture of much of the
upper surface. The 2 individuals of majuJu
share these characters and, except in color,
differ only slightly in other ways: e.g..,
the 9 has the elytral apices more acutely
toothed but the outer angles more obtuse.
The 2 color forms of mafulu are keyed out
separately in the Keij to Species.
Demetrido forma n. sp.
Description. With characters of genus;
form as in Figure 104; reddish brown,
legs testaceous; not pubescent, microsculp-
ture present ( weak or faint ) only on elytra,
surface not much punctulate. Head 1.12
and 1.10 width prothorax; eyes prominent,
genae short, oblique. Prothorox subquad-
rate, long; width length 0.99 and 1.01; base
apex 1.37 and 1.23; base/head 0.82 and
0.81; sides weakly irregularly rounded
through much of length, moderately sinu-
ate posteriorly before c. acute but blunted
basal angles; margins narrow, each with
seta-bearing puncture slightly before mid-
dle but none at base; surface in part ir-
regular or weakly punctate posteriorly and
laterally. Elytra rather long; width elytra/
prothorax 2.11 and 2.18; apices spined, outer
angles c. right, sharply formed, sutural
angles right or slightly obtuse, sometimes
slightly blunted; striae impressed, weakly
punctulate; intervals convex, 3rd with 2
dorsal punctures. Claws with 7 or 8 teeth.
Secondary sexual characters: S tarsi as
genus; 6 middle tibiae tuberculate-serrate
(c. 3 or more tubercles); 6 with 2 or 3, $ c.
4 apical ventral setae each side. Measure-
ments (type series): length 9.3-9.6; width
3.1-3.3 mm (specimens listed under Addi-
tional material 6.8-9.6 mm long).
Types. Holotype $ (Bishop Mus.) and
7 paratypes (3 in M.C.Z., Type No. 31,466)
all from Pindiu, Huon Pen., N-E. N. G.,
500-600, 870-1300 m; dates in Apr. 1963
(holotype, 500-600 m, Apr. 19) (Sedlacek).
Additional material. Twenty-three speci-
mens from 9 localities in Papua, N-E.
rV. G., and eastern West N. G. Because
of variations (see following Notes) and
doubtful identifications these specimens
are not recorded in detail.
Measured specimens. The i holotype and
1 9 paratype.
Notes. Under this species I have tenta-
tively placed all nonpubescent, brown,
pale-legged New Guinean Demetrida with
spined elytra, prothorax elongate-subquad-
rate and considerably narrower than head
( width head prothorax usually but not
always c. 1.10 or more), and 3rd intervals
2-punctate ( but see below ) . The specimens
thus assembled vary considerably in size,
prominence of eyes, exact form of prothorax,
and length of elytral spines. Several species
may be represented but, if so, I cannot
separate them now.
Although the 3rd intervals are 2-punctate
on both elytra in most individuals, in 3
cases a 3rd (intermediate) puncture is
present on one side only, and an individual
from Wau which I tentatively assign to
forma is 3-punctate on both sides. This
individual is the only forma (if it is this
species) seen from Wau. It is a small S
with tuberculate-serrate middle tibiae.
Demetrido recto n. sp.
Description. With characters of genus;
form c. as preceding species (forma) except
elytra short-spined; reddish brown, legs
pale with dark knees; not pubescent, reticu-
late microsculpture distinct only on elytra,
but surface in part finely sparsely punc-
tulate. Head 1.14 and 1.20 width prothorax;
eyes prominent, genae shorter, oblique, not
prominent (but see Notes below). Pro-
thorax long-quadrate; width length 0.92
and 0.88; base apex 1.28 and 1.28; base/
head 0.85 and 0.84; sides virtually straight
anteriorly or weakly angulate at setae,
sinuate well before slightly acute basal
angles; margins narrow, each with seta
slightly before middle but none at base;
baso-lateral impressions weak, subpunctate.
168 Bulletin Miiscuin of Comparative Zoology, Vol. 137, No. 1
Elytra: width elytra prothorax 2.14 and
2.30; apices shoit-.spined or acutely toothed,
outer angles sharply formed, acute, inner
angles obtuse; striae impressed, punctulate;
intervals convex, 3rd 2-punctate. Clows
with c. 6 or 7 teeth. Secondary .sexual
characters: 6 tarsi as genus; i middle
tibiae bent in at apex but not tuberculate-
serrate; 6 with 3, 9 4 or 5 setae each
side last ventral segment. Measurements:
length 8.4-9.7: width 2.7-3.3 mm.
Types. Holotype c^ (Bishop Mus.) and
2 9? paratypes (1 in M.C.Z., Type No.
31,467) from Wan, Morobe Dist, N-E.
N. G., 1200 (liolotypc), 1050, and 1090 m,
dates in Jan. 1963 (holotype, Jan. 8-10)
( Sedlacek ) .
Additional material. West N. G. : 1 c^ ,
Hollandia, May 1945 (Hoogstraal, M.C.Z.).
Measured sj^ccimens. The J, holotype and
1 9 paratype.
Notes. D. recta resembles forma but is
distinguished by straighter sides of pro-
thorax and especially by bent-in but not
tuberculate-serrate i middle tibiae. D.
recta may be more closely related to
l<iun£ia but is smaller, more slender, with
shorter elytral spines. The real interrela-
tionships of these and other more or less
similar species are doubtful.
Tliis species emphasizes that the ratio
base/apex of prothorax must be interpreted
with caution. The ratio of 1.28 in recta
suggests that the apex is considerably nar-
rower than the base, and this is true when
the apex is measured in the standard way,
between the most advanced points of the
angles. Nevertheless, the prothorax appears
\ iitually rectangular.
Th(> genae of the lK)l<)t\'|)e are unsym-
metrie: the right one is normal, as de-
scribed above and as in the other speci-
mens of the species, wlule the left one is
subangulate just behind the eye, although
not so jirominent as in tri})uncfa and
iS,enicula. This slight angulation of the left
gena in one specimen only ol recta is
presumably an abnormality.
Demetrida rex n. sp.
Description. With characters of genus;
form as in Figure 105; reddish brown; not
pubescent, reticulate microsculpture faint
or absent even on elytra, but much of
upper surface sparsely inconspicuously
punctulate. Head 1.06 and 1.08 width pro-
thorax (at middle); eyes prominent, genae
shorter, oblique. Prothorax trapezoidal,
wider at base than at middle; width ( at
middle) length 1.06 and 1.02; base, apex
1.41 and 1.34; base/width at middle 1.04
and 1.06; base head 0.98 and 0.98; sides
shaped as figured, narrowly margined, each
with seta-bearing puncture at or slightly
before middle but none at base; surface
irregularly slightly punctate at base and
sides. Elytra ample; width elytra prothorax
(at middle) 2.00 and 2.12; apices spined,
outer angles acutely denticulate, sutural
angles obtuse-blunted; striae impressed,
punctulate; intervals slightly convex, 3rd
with 3 dorsal punctures (all specimens).
Claws with 7 or 8 teeth. Secondary sexual
characters: 6 tarsi as genus; c5 middle
tibiae slightly bent-in toward apex but not
tuberculate-serrate; 6 with 2 or 3, 9 c. 5
apical ventral setae each side. Measure-
ments: length 10.2-11.4; width 3.5-4.0 mm.
Types. Holotype i ( Bishop Mus. ) from
Mokai Village, Torricelli Mts., N-E. N. G.,
750 m, Dec. 8-15, 1958 (^^^ W . Brandt);
and additional paratypes as follows. N-E.
N. G.: 1 9, Eliptaniin Vy., 1200-1350 m.
June 19-30, 1959 (W. \V. Brandt. Bishop
Mus.); 1 9, Adalbert Mts.. Wanunia, 800-
1000 m, Oct. 24, 1958 (Gressitt); 1 i , Pin-
diu, Huon Pen., Apr. 20, 1963 (Sedlacek).
Papua: 1 i , Dogon, Amazon Ba\ Dist..
2400 ft. ( c. 730 m), Oct.-NoN . 1962 (W. W.
Brandt, C.S.I.H.O.); 1 9, Owen Stanley
Rge., Ooilala, B()m(\ 1950 m. Apr. 16-30.
1958 (\\. W. Brandt. Bishop Mus.). (Thc>
]")aratvpes from Adalbert Mts. and Pindiu
now in M.C.Z.. Type No. 31,468.)
Measured specimens. The ■: holotype and
9 paratype from Eliptamin Valley.
Notes. Although the 6 speeiinens listi-d
The Carabid Beetles of New Guinea
Darlington
169
above come from 6 different localities, they
seem to be conspecific and to represent a
very distinct species, characterized by large
size, form of prothorax, presence of an
acute tooth ( almost a short spine ) at outer-
apical elytral angle, and 3-punctate 3rd
intervals. See Key to Species for place of
rex among other New Guinean Demetrida.
Demefrida brunnea n. sp.
Description. With characters of genus;
form average, with spined elytra; brownish
testaceous, head and prothorax usually
slightly darker than elytral disc, legs pale;
not pubescent, reticulate microsculpture
visible (light) only on elytra, surface not
much punctulate. Head 1.12 and 1.13 width
prothorax; eyes prominent, genae shorter,
oblique. Prothorax quadrate; width length
1.02 and 0.99; base/apex 1.23 and 1.26;
base/head 0.83 and 0.87; sides subparallel
or weakly irregularly arcuate in anterior %
or more, subangulate at lateral setae,
broadly sinuate before right or slightly
acute posterior angles; margins narrow,
each with seta at or slightly before middle
but none at base; surface irregular or
slightly punctate at base and sides. Elytra:
width elytra prothorax 2.06 and 2.14; apices
spined, outer angles denticulate, sutural
angles right or slightly obtuse, slightly
blunted; striae impressed, faintly punctu-
late; intervals slightly convex, 3rd usually
3-punctate. Claws with c. 6 teeth. Sec-
ondary sexual characters: 6 tarsi as genus;
6 middle tibiae slightly bent in at apex but
not tuberculate-serrate (Fig. 161); 6 with
usually 3, 9 5 or 6 apical ventral setae each
side. Measurements: length 8.5-10.0; width
2.9-3.4 mm.
Types. Holotype $ (British Mus.) and
8 paratypes (3 in M.C.Z., Type No. 31,469)
from Mt. Baduri, Japen Is., West N. G.,
1000 ft. (305 m), Aug. 1938 (Cheesman);
and the following additional paratypes
from West N. G.: 1, R. Manai-Undei,
Japen Is., 500 ft. (c. 150 m), Oct. 1938
(Cheesman); 3, Mt. Lina, Cvclops Mts.,
3500-4500 ft. (c. 1070-1370 mj, Mar. 1936
(Cheesman); 6, Sibil, Star Rge., 1260 m,
dates in May, June 1959 ( Neth. N. G. Exp.,
Leiden Mus.), at hght; 2, Sibil Vy., Star
Mts., 1245 m, Oct. 18-Nov. 8, 1961 (S.
Quate, Bishop Mus.); 1, Bivak 36, Star
Rge., 1220 m, July 29, 1959 (Neth. N. G.
Exp., Leiden Mus.).
Additional material N-E. N. G.: 2,
Eliptamin Vy., 1200-1350 m, June 19-30,
Aug. 1-15, 1959 (W. W. Brandt, Bishop
Mus.); 1, Feramin, 1200-1500 m, June 1-6,
1959 (W. W. Brandt, Bishop Mus.); 1,
Pindiu, Huon Pen., 1200-1450 m, Apr. 18,
1963 (Sedlacek). Papua: 3, Mafulu, 4000
ft. (c. 1220 m), Jan. 1934 (Cheesman); 1,
\\'akaiuna, Sewa Bay, Normanbv Is., Dec.
11-20, 1956 (W. W. Brandt, Bishop Mus.).
Measured specimens. The i holotype and
1 9 paratype from Japen Is.
Notes. D. brunnea resembles forma in
most key characters, but brunnea has oviter
apical elytral angles acutely denticulate (c.
right in forma), 3rd intervals usually 3-
punctate (2-punctate in forma), and S
middle tibiae slightly bent-in but not
tuberculate-serrate as in forma.
Actually, the punctures of the 3rd inter-
vals vary slightly. Two individuals of
brunnea (the holotype and the paratype
horn Bi\ak 36) have 3 punctures on one
and 2 on the other side, although all other
brunnea listed above are 3-punctate on
both sides.
Demetrida fumipes n. sp.
Description. See Plate 2, figure VI; with
characters of genus; form slender, with
prominent eyes and short-spined elytra;
reddish brown, elytra ± paler on disc but
with sides behind humeri blackish, legs
pale with outer edges of tibiae and apices
of femora dark or legs more extensively
dark; not pubescent, reticulate microsculp-
ture faint or light even on elytra, surface
not much punctulate. Head 1.12 and 1.18
width prothorax; eyes prominent, genae
shorter and not prominent. Prothorax
quadrate; width length 1.05 and 0.99; base/
apex 1.21 and 1.17; base/head 0.83 and
170 Biilletin Museum of Companitive Zoology, Vol. 137, No. 1
0.82; sides almost straight or weakly arcuate
in anterior %, sinuate before c. right but
irregular basal angles; margins narrow, each
with seta-bearing puncture before middle,
none at base; surface scarcely punctate
even baso-laterally. Ehjtiri: width elytra
prothorax 2.07 and 2.26; apices short-spined,
outer angles acute, sutural angles slightly
blunted; striae impressed, slightly punc-
tulate; intervals slightly convex, faintly
sparsely punctulate, 3rd usually 3-punctate.
Claws with c. 5 or 6 teeth. Secondary sexual
characters: i tarsi as genus; 6 middle
tibiae slightly bent-in at apex but not
tuberculate-serrate; 6 with 2 or 3, 9 4-6
apical ventral setae each side. Measure-
ments: length 8.3-9.4; width 2.7-3.2 mm.
Tifpes. Holotype S (Bishop Mus.) and
17 paratypes (6 in M.C.Z., Type No. 31,470)
all from Wan and vicinity (including Mt.
Missim), Morobe Dist., N-E. N. G., alti-
tudes from 1100 to 1500-1900 m, dates in
Feb., Mar., Apr., July, Sept., Nov., 1961-
1963 (holotype, Wan, 1200-1300 m, Apr. 6,
1963) (Sedlaceks).
Additional material. Papua: 3, Doveta,
Amazon Bay Dist., 2400 ft. (730 m), Aug.
1962 (W. W. Brandt, C.S.I.R.O.). N-E.
N. G.: 1 teneral, Wau, 1200 m, Sept. 2,
1961 (Sedlacek).
Measured specimens. The S holotype and
1 9 paratype from Wau.
Notes. Among species that are not ob-
viously bicolored, fumipes is closest to
forma but has legs in jiart darker, sides of
elytra behind humeri darker, and 3rd in-
tervals 3-punctate (2-punctate in forma).
Among bicolored species, fumipes is nearest
humcraJis but is smaller, witli humeri less
extensively black. See also comparative
notes under following species.
Demefrida veiafa n. sp.
Description. With characters of genus;
lonn as in preceding species (fumi))es);
reddish brown, disc of elytra ± paler but
sides ol elytra narrowly blackish behind
humcMi, legs in part dark (at least darker
than elytral disc); not pulK\scent, reticulate
microsculpture faintly indicated on pro-
notum and sometimes on part of head and
distinct (but light) on elytra, surface not
much punctulate. Head 1.14 and 1.08
width prothorax; eyes prominent, genae
shorter, not prominent. ProtJiorax quadrate,
long; width length 0.94 and 1.00; base apex
1.21 and 1.34; base head 0.84 and 0.88;
sides weakly irregularly arcuate anteriorly,
strongly sinuate before right or slightly
acute basal angles; margins narrow, each
with seta-bearing puncture before middle
but none at base; baso-lateral areas irreg-
ularly subpunctate. Elytra: width elytra/
prothorax 2.05 and 2.16; apices short-spined
or acutely toothed, with outer angles ± right
and sharply defined, sutural angles obtuse
or slightly rounded; striae impressed, faintly
punctulate; intervals slightly convex, faintly
sparsely punctulate, 3rd usually 2-punctate.
Claws with 5 or 6 teeth. Secondary sexual
characters: i tarsi as genus; i middle
tibiae tuberculate-serrate (c. 6 low tuber-
cles ) ; S with 3 ( rarely 4 ) , 9 c. 5 setae
each side last ventral segment. Measure-
ments: length 8.4-8.8; width 2.7-3.0 mm.
Types. Holotype i ( Bishop Mus. ) from
Saidor, Kiambavi Village, Finisterre Rge.,
N-E. N. G., July 22-29, 1958 (W. \\\
Brandt ) , and 9 paratypes from Finisterre
Rge. (3 in M.C.Z., Type No. 31,471) as
follows: 1, same data as holotype except
Aug. 1-28; 7, Saidor, Ahitoko (N'illage),
Aug. 29-Sept. 5, Sept. 6-24, 1958 (all col-
lected by W. W. Brandt).
Measured specimens. The i holot>pi' and
1 9 paratype from Matoko Village.
Notes. This apparent relati\c> of fornui
is distinguished from the latter b\- dark
elytral edges and daik legs and from
fumipes b\' usualK 2-punctate rather than
3-punctate 3rd intervals, and celata differs
from th(\se and from other similar species
also in Inning ic'ticiilate microsculpture
N'isible, although hiint, on pronotum (and
sometimes part ol head ) as well as elytra.
D. veldid may be more closely related to
diversa but is less distinctK bicolored and
The Carabid Beetles of New Guinea
Darlington
171
more distinctly microreticulate, w ith usually
better developed (but still short) elytral
spines.
The 3rd intervals are 2-punctatc on both
sides of all individuals except that an extra
(3rd) puncture is present on one side only
in two individuals.
One specimen of the type series has
moth scales stuck to it and is presumably
from light-trap material.
Demefrida nigripes n. sp.
Description. With characters of genus;
form as in Figure 106; head and prothorax
red, elytra black, legs and antennae exten-
sively dark with pale bases, tarsi paler; not
pubescent, reticulate microsculpture virtu-
ally absent in i , present (moderateh'
transverse ) on elytra in 9 , surface not
much punctulate. Head 0.98 and 0.94 width
prothorax; eyes prominent, genae shorter
and oblique. Prothorax cordate-subquad-
rate; width length 1.20 and 1.28; base/apex
1.39 and 1.34; base head 0.94 and 0.98;
sides broadly slightly irregularly arcuate in
more than anterior n, moderateh' sinuate
before e. right posterior angles; margins
rather wide, each with seta-bearing punc-
ture slightly before middle but none at
base; disc slightly punctate at sides and
base. Elytra parallel; \\'idth elytra pro-
thorax 1.77 and 1.71; apices with short
spines, outer angles well defined but
slightly obtuse, sutural angles obtuse-
blunted; striae less impressed but more
punctulate than usual; intervals slightly or
not convex, 3rd with 3 dorsal punctures
(middle puncture sometimes doubtful).
Claics with 5 teeth. Secondary sexual char-
acters: i tarsi as genus; 6 middle tibiae
tuberculate-serrate (3 or 4 rounded tuber-
cles); 6 with 2, 9 3 apical ventral setae
each side. Measurements: length 7.4-7.7;
width 2.3-2.7 mm.
Types. Holotype £ (Bishop Mus.) from
Swart Vy., Karubaka, N-E. N. G., 1350 m,
Nov. 18, 1958 (Gressitt); and 1 9 para-
type (Bishop Mus.), Daradae, nr. Javarere,
Musgrove R., Papua, 100 m ?, Oct. 2,
1958 (Gressitt).
Notes. Although the 2 specimens listed
above are from different localities, they
agree in so many ways ( in spite of disagree-
ment in a few details ) that I feel sure they
are conspecific and that they represent an
unusually distinct species, characterized by
form, color, and relatively light but strongly
punctulate elytral striae, as well as by other
characters given in the Key to Species of
Demetrida of New Guinea.
Demefrida dorsalis n. sp.
Description. See Plate 2, figure VII; with
characters of genus; black, elytra with large
common red area centered behind middle,
appendages dark; not pubescent, reticulate
microsculpture absent or indistinct, but
parts of upper surface ( especially elytra )
sparsely punctulate. Head 0.93 and 0.89
width prothorax; eyes prominent, genae
shorter, oblique; front slightly convex, im-
pressed each side anteriorly, slightly punc-
tate at middle. ProtJu^rax subcordate;
width length 1.33 and 1.39; base apex 1.34
and 1.33; base/head 0.95 and 0.96; sides
broadly arcuate in more than anterior %,
strongly sinuate before right or slightly
acute posterior angles; margins rather wide,
each with seta-bearing puncture at or
slightly before middle but none at base;
surface slightly punctate at base and sides.
Elytra: width elytra prothorax 1.77 and
1.78; apices angulate, the angulations right
or slightly acute, outer angles obtuse or
narrowly rounded, sutural angles obtuse;
striae well impressed, scarcely punctulate;
intervals convex, sparsely but distinctly
punctulate, 3rd with c. 4 dorsal punctures.
Chiics with c. 5 teeth. Secondary sexual
characters: S tarsi as genus (squamae of
front tarsi disarranged, but probably in 2
series ) ; S middle tibiae tuberculate-serrate
(c. 4 low rounded tubercles); S with 2 or
3, 9 3 or 4 apical ventral setae each side.
Measurements: length 6.8-8.2; width 2.7-
3.1 mm.
172 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Ti/pes. Holotype S (Bishop Miis.) and
7 paratypes (3 in M.C.Z., Type No. 31,472)
all from Wau and vicinity (inchiding Mt.
Missim and Nami Creek), Morobe Dist.,
N-E. N. G., altitudes from 1100 to 1650
m, dates in Jan., Feb., Mar., 1962, 1963
(holotype. Wan, 1100 m, Jan. 31, 1963)
( Sedlacek ) .
Measured specimens. The i holotype and
1 5 paratype.
Notes. Although known only from a
single locality, this species seems a distinct
on(\ characterized by form, color, angulate
but not spined elytral apices, and virtual
absence of dorsal elytral reticulate micro-
sculpture, as well as by other key char-
acters.
Demetrida basalis n. sp.
Description. With characters of genus;
form and most characters of duplicata ( p.
000); red or brown with base of elytra
(sometimes only humeri) black, legs and
antennae reddish with part of femora and
tibiae usually darker; not pubescent, micro-
sculpture visible (sometimes faint or in-
distinguishable ) only on elytra, surface in
part sparsely punctulate. Head 0.93 and
1.02 width prothorax; eves prominent, genae
shorter, oblique. Prothorax ({uadrate-sub-
cordate; width/length 1.21 and 1.17; base/
apex 1.36 and 1.30; base ^head 0.92 and
0.87; sides irregularly arcuate in more than
anterior 'V\, sometimes subangulate at setae,
strongly sinuate before c. right or slightly
acute sometimes slightly blunted posterior
angles; margins moderately wide, each with
seta-bearing puncture at or slightly before
middle but none at base; disc slightK
punctate across base and in margins.
Ell/Ira: width elytra/prothoiax 1.S6 and
2.10; apices spined, outer angles ± right and
sharply defined, sutmal angles slightly
obtuse, blunted; striae impressed, scarcely
punctulat(\ intervals slightly convex, slightly
punctulate, 3rd with 2 principal and 1 or 2
intermediate smaller dorsal punctures (all
specimens). Clans with r. 5 teeth. Sec-
ondary sexiad chdraclcrs: i tarsi as genus;
4 middle tibiae tuberculate-serrate (c. 6
small tubercles); i with 2 or 3, 9 3 (or
more?) apical ventral setae each side. Mea-
surements: length 8.4-10.0; width 2.9-3.5
mm.
Types. Holotype $ (Bishop Mus.) from
Swart Vv., West N. G., W. ridge, 1800-
2000 m,^ Nov. 19, 1958 (Gressitt); and
paratypes as follows. West N. G.: 4(2 in
M.C.Z., Type No. 31,473), Swart Vy.,
Karubaka, 1500 m, Nov. 11, 20, 1958 (Gres-
sitt); 2, Wissel Lakes, Kamo Vy., Itouda and
Moanemani, 1500-1700 m, Aug. 18, 16, 1962
(Sedlacek); 1, Star Rge., Sibil Vy., 1245
m, Oct. 18-Nov. 8, 1961 (S. Quate, Bishop
Mus.); 1, same locality, 1260 m, June 16.
1959 (Neth. N. G. E.xp., Leiden Mus.), at
hght; 1, Star Rge., Bivak 39, 1300 m, June
28, 1959 (Neth. N. G. Exp., Leiden Mus.);
1 teneral, Araucaria Gamp, 800 m, Apr. 2,
1939 (Toxopeus).
Additional material. One teneral, Karu-
baka, 1450 m, Nov. 16, 1958 (Gressitt),
light trap.
Measured specimens. The 6 holotype and
1 9 paratype from Karubaka.
Notes. This ma>- be a geographic ( west-
ern ) representatix'e of duplicata ( p. 164 )
with elytra black at base rather than en-
tirely reddish brown (but intermediate
color fonns occur as noted under duplicata )
and with elytra Ic^ss distinctb microreticu-
late. Its placc^ among other similarly bi-
colored species is indicated in the Keij to
Species.
Demefrida diverso n. sp.
Description. See Plate 2, figure MIL
with cliaracters ot genus; lorm slender,
with prominent eyi's and acutcK angulate
or short-spined el\ tral apices; color diverse,
brownish red with either whole base of
elytra (except suture ) or onl\ humeri either
black or green (indi\idu;il \ariation), legs
either entireK' red or almost entiicly black
(h()lot\pe, basal '.: ol ehtra black, legs
red); not pubeseint. reticulate microscul]!-
ture indistinc-t or light rwn on elytra, sur-
laee not much piuulnlate. Head 1.07 and
The Carabid Beetles of New Guinea • Darlington 173
1.14 width prothorax; eyes prominent, genae
shorter and obHque. Prothorax subquadrate
or trapezoidal with base varying from nar-
rower than to wider than widtli at middle;
width (at middle) length 1.04 and 0.95;
base apex 1.39 and 1.46; base head 0.88
and 0.94; sides weakly irregularly arcuate
in c. anterior %, sinuate before c. right or
acute but blunted or narrowly rounded
posterior angles; margins rather narrow,
each with seta-bearing puncture before mid-
dle but none at base; surface irregularly
punctate basally and in margins. Elytra:
width elytra/ prothorax (at middle) 1.97
and 2.25; apices with slightly obtuse or
acute angulations or very short spines,
outer angles well defined but varying from
slightly obtuse to acute, sutural angles
obtuse; striae moderately impressed, slightly
punctulate; intervals slightly convex, 3rd
usually 2- sometimes 3-punctate ( see Notes,
below). Claws with c. 5 or 6 teeth. Sec-
ondary sexual characters: 6 tarsi as genus;
S middle tibiae tuberculate-serrate (c. 7
small tubercles); 6 with 2 or 3, 9 4 or
more apical ventral setae each side. Mea-
surements: length 7.5-9.5; width 2.7-3.3
mm.
Types. Holotype c^ (Bishop Mus.) and
27 paratypes (some in M.C.Z., Type No.
31,474) all from Wau and vicinity (in-
cluding Mt. Missim), Morobe Dist., N-E.
N. G., altitudes from 1090 to 1700 m, dates
in Jan., Feb., Mar., May, June, July, Sept.,
Oct., Nov. 1961-1963 (holotype, 1250 m.
May 3, 1963) (Sedlaceks).
Additional material. N-E. N. G.: 3,
Eliptamin Vy., 1200-1350, 1665-2530 m,
June 23-30, July 1-15. 16-31, 1959 (W. W.
Brandt, Bishop Mus.); 1, Korop, Upper
Jimmi Vy., 1300 m, July 12, 1955 (Gres-
sitt), in light trap; 1, Swart Vy., Karubaka,
1550 m, Nov. 8, 1958 (Gressitt, No. 3145);
1, Jim(m)i R., E. Highlands, July-Sept.
1961 (W. W.Brandt, G.S.I.R.O.).
Measured specimens. The S holotype and
1 9 paratype.
Notes. The series from Wau shows the
entire range of variation indicated in the
preceding Description. Occurrence together
of such diverse individuals in what seems
to be one population suggests Mendelian
dimorphism of color (elytral bases black
or green, legs red or black ) , and exceptional
genetic variation of some other characters.
The different characters vary indepen-
dently. For example, leg color is not cor-
related with color or extent of basal elytral
marks. Of the type series, 15 individuals
have 2 punctures on each 3rd interval, 4
(including the holotype) have 2 on one
side and 3 on the other, and 2 individuals
have 3 punctures on each side.
For distinguishing characters of diversa
see the Key to Species of Demetrida of New
Guinea.
Demetrida vigil n. sp.
Description. With characters of genus;
form as in Figure 107; head and prothorax
brownish red, elytra red with basal V.\
black, antennae red, legs mainly black; not
pubescent, reticulate microsculpture faint
and irregular even on elytra in i ( possibU'
more distinct in 9 ), surface not much
punctulate. Head 1.03 width prothorax;
eyes not larger than usual but exceptionally
abruptly prominent, joining neck posteriorly
with virtually no genae. Vrothorax long-
quadrate with anterior angles virtually ob-
literated; width/ length 0.99; base/apex 1.39;
base/head 0.91; sides weakly arcuate,
sinuate before c. right basal angles; margins
narrow, each with seta-bearing puncture
at or slightly before middle (an extra ad-
ventitious puncture on left) but none at
base; disc very convex, slightly rugulose
and punctulate especially in baso-lateral
depressions. Elytra: width elytra prothorax
1.92; apices spined, outer angles acute-
denticulate, sutural angles obtuse-blunted;
striae slightly impressed, faintly punctulate;
intervals slightly convex, 3rd 2-punctate.
Claus with 4 or 5 teeth. Secondary sexual
characters: 6 tarsi as genus; S middle
tibiae weakly tuberculate-serrate (c. 4 low
tubercles); c^ with 3 apical ventral setae
174 Bulletin Muscidu of Comparative Zoology, Vol. 137, No. 1
each side; 9 unknown. Mcasiiiements:
length 7.8; width 2.5 mm.
Type. Holotype S (C.S.I.R.O., Can-
berra) from Doveta, Amazon Bay Dist.,
SE. Papua, 2400 ft. (c. 730 m), Aug. 1962
(W. W. Brandt); the type is unique.
Notes. The eyes, more abruptly promi-
nent than in anv other Denietrida that I
know, distinguish vigil from such similarly
colored species as diversa and divisa.
Demefrido nigriceps n. sp.
Description. With characters of genus;
form ( Fig. 10(S ) e. average, with prominent
but not abrupt eyes, rather narrow pro-
thorax, and spined elytra; head and pro-
thorax black, elytra entirely brown, ap-
pendages brown with antennae darker
outwardly; not pubescent, microsculpture
indicated (faint and irregular) on elytra
only. Head 0.96 and 1.01 width prothorax;
eyes prominent, genae shorter, oblique.
Prothorax subquadrate, long, with base
sometimes wider than middle; width length
0.96 and 1.01; base/apex 1.30 and 1.30; base/
head 0.(S9 and 0.85; sides subparallel in ante-
rior 'Yi, faintly angulate at lateral setae,
broadly sinuate before right or slightly acute
posterior angles; lateral margins narrow,
each with seta-bearing puncture at or
slightly before middle but none at base; sur-
face scarcely or slightly punctate in luargins.
Elytra: width elytra prothorax 2.21 and
2.14; apices spined, outc-r angles acute
and subdenticulate, sutural angles obtuse-
blunted; striae slightly iiupressed, slightly
pimctulate; intervals scarcely convex, 3rd
2-punctate. ('lans with 6 or 7 teeth. Sec-
ondary sexual characters: ', tarsi as genus;
$ middle tibiae not inodificxl ( r. straight,
not tuberculate-serrate, in both s|:>ecimens ) ;
i with 2-4 apical ventral setae each side
(holotype, 4 on each side; paratype, 2 on
one side, 3 on other j; v unknown. Mea-
surements: length e. 10.0; width 3.2 mm.
Types. Holotype S (Bishop Mus.) and
1 ^'paratypc^ (M.C.Z., Type No. 31,475)
both Iroin Sibil Vall(>v, Star Mts.. Wrsi
N. G., 1245 m, Oct. 18-Xov. 8, 1961 (S.
Quate), the holotype at light, the paratype
in Malaise trap.
Notes. The unmodified 6 tibiae distin-
guish this species among other similar ones,
and other differential characters are given
in the Key to Species of Demetrida of New
Giunea.
The sex { i S ) of both specimens has
been determined by dissection as well as
by examination of the front tarsi.
Demetrida saidor n. sp.
Description. With characters of genus;
form slender, with prominent but not
abrupt eyes and spined elytra; head and
prothorax brownish black, elytra brownish
testaceous with humeri and sometimes en-
tire base narrowly black, appendages ir-
regularly dark with paler tarsi; not pubes-
cent, reticulate microsculpture visible ( faint
or light) only on elytra, surface not much
(slightly, finely, sparsely) punctulate. Head
1.25 and 1.11 width prothorax; eyes promi-
nent, genae short, oblique. Prothorax long-
quadrate with relatively wide base; width/
length 0.93 and 1.02; base apex 1.43 and
1.30; base/head 0.85 and 0.88; sides sub-
parallel or slightly arcuate in c. anterior ^^i,
sinuate before usually acute but slightly
blunted posterior angles; margins narrow,
each \\'ith seta-bearing puncture at or
slightly before middle but none at base;
surface slightK' punctate at base and in
margins. Elytra: width ehtra prothorax
2.28 and 2.17; apices spined, outer angles
acute, sharpK defined, sutural angles ob-
tuse; striae lightK impressed, punctulate;
intervals I hit or slightK' comcx, 3rd 2-
punctate (all specimens), the posterior
puncture fai- back. Clatrs with c. 6-7 teeth.
Seconchiry sexii(d characters: S tarsi as
g{>nus; -; middl(> tibiae not or \ery littU^
modilied, not bent in at apex and not
tuberculate-serrate; ,^ with 3, 9 c. 6 apical
ventral setae each side. Measurements:
length 8.6-10,0; width 2.8-3.3 mm.
I'yiics. ?Tolotype S (Bishop Mus.) from
The Carabid Beetles of New Guinea
Darlington
175
Saidor, Matoko Village, Finisterre Rge.,
N-E. N. G., Sept. 6-24, 1958 (W. W.
Brandt); 1 6 paratype (M.C.Z., Type No.
31,476) from Saidor, Kiambavi Village,
Aug. 1-28, 1958 (W. W. Brandt); 2(69)
paratypes from Sepalakembang, Salawaket
Rge., N-E. N. G., 1920 m, Sept. 11-14, 12,
1956 (E. J. Ford, Jr., Bishop Mus.), in
light trap.
Additional material. West N. G.: 1 ten-
eral 6 , Wissel Lakes, Moanemani, Kamo
Vy., 1500 m, Aug. 19, 1962 (Sedlacek).
Measured specimens. The c$ holotype and
the 9 paratype from Sepalakembang.
Notes. Among more or less similar species
(fiimipes, velata, nigriccps) this is distin-
guished by combination of polished ( not
microreticulate) black head and pronotum,
black humeri and legs, and simple 6 middle
tibiae.
Demefrido divisa n. sp.
Description. With characters of genus;
form c. average, with prominent but not
abrupt eyes, rather narrow prothorax,
spined elytra; red or yellowish with basal
% or V-i of elytra black, lower surface red-
dish yellow with metepisterna mainly dark,
legs testaceous with dark knees, antennae
brown; not pubescent, reticulate micro-
sculpture distinct (light) only on elytra,
surface not much punctulate. Head 1.14
and 1.11 width prothorax; eyes prominent,
genae oblique and shorter than eyes, some-
times slightly convex in profile but not
very prominent. Prothorax quadrate, long;
width length 0.95 and 0.95; base apex 1.30
and 1.32; base/head 0.88 and 0.95; sides
nearly straight (except subangulate at
setae) in more than anterior ■'4, sinuate
before c. right or acute, slightly blunted
posterior angles; margins rather narrow,
each with seta-bearing puncture before
middle but none at base; surface vaguely
subpunctate baso-laterally. Elytra: width
elytra prothorax 2.05 and — (elytra spread);
apices variably spined (spines usually short),
outer angles sharph' defined, right or acute,
sutural angles obtuse; striae impressed,
scarcely punctulate; intervals slightly con-
vex, sparsely slightly punctulate, 3rd with
2 or 3 dorsal punctures (see following
Notes). Claws with c. 7 teeth. Secondary
sexual characters: 6 tarsi as genus; S
middle tibiae scarcely modified, at most
slightly bent in at apex, not tuberculate-
serrate; $ with c. 4, 9 5 or 6 apical ventral
setae each side. Measurements: length
9.5-11.5; width 3.1-3.9 mm.
Types. Holotype $ (Bishop Mus.) and
1 <i 'paratype (M.C.Z., Type No. 31,477)
from Tsenga, Upper Jimmi Vy., N-E. N. G.,
1200 m, July 14, 1955 (Gressitt); and addi-
tional paratvpes as follows. N-E. N. G.: 2,
W'au, iMorobe Dist., 1200 m, Sept. 27, 1961,
May 1-15, 1962 (Sedlacek), in light trap;
5, Okapa ( Hornabrook ) ; 1, Swart Vy.,
Karubaka, 1300 m, Nov. 7, 1958 (Gressitt);
1, Sattelberg (British Mus.); Papua: 1,
Kokoda, 1200 ft. (366 m). May 1933
(Cheesman); 2, Dogon, Amazon Bay Dist.,
2400 ft. {c. 730 m), Sept., Oct.-Nov. 1962
(W. W. Brandt, C.S.I.R.O.). West N. G.:
1 9 , Bomberi, Vogelkop, 700-900 m, June
5, 1959 (Gressitt).
Measured specimens. The 6 holotype and
9 paratype from Sattelberg.
Notes. D. divisa resembles one of the
color forms of diver.sa, but divisa is larger,
with elytra at least short-spined and S
middle tibiae not tuberculate-serrate as in
diversa.
In the 6 holotype and S paratype from
Tsenga and also the 9 from Bomberi the
3rd intervals are 2-punctate; in all other
specimens, 3-punctate; but I find no other
characters to suggest that this is a specific
difference.
The specimen from Sattelberg is labeled
by Andrewes, "Genus mihi ignotum."
Demefrida humerolis n. sp.
Description. With characters of genus;
form c. average, with prominent eyes,
quadrate prothorax, and short-spined elytra;
reddish brown, humeri black, legs black or
bicolored; not pubescent, microsculpture
176 Bulletin Muacuni of Coiup<irativc Zoology. Vol. 137, No. 1
faint even on elytra, surtace not much
punctulate. Head 1.07 and l.OS width pro-
thorax; eyes prominent, genae shorter and
obHque. ProtJiorax sul)Ciuadrate with rather
broad base; width length 1.06 and 1.00;
base /apex 1.33 and 1.23; base/head 0.89
and 0.90; sides weakly irregularly arcuate,
usually subangulate at setae, broadly sinu-
ate before right or slightly acute posterior
angles; margins rather narrow, each with
seta-bearing puncture at or before middle
but none at base; disc subpunctate across
base and in margins. Elytra: width elytra
prothorax 1.99 and 2.16; apices short-spined.
outer angles sharply defined, usually dentic-
ulate, sutural angles c. right or slightly
ol)tuse; striae moderately impressed, faintly
punctulate; intervals slightly convex, 3rd
usually 3-punctate. Claws with 5 or 6 teeth.
Secondary sexual characters: 6 tarsi as
genus; i middle tibiae scarcely modified,
at most slightly bent in at apex but not
tuberculate-serrate; 6 with 3, 9 c. 6 apical
ventral setae each side. Measurements:
length 9.3-10,8; width 3.0-3.5 mm.
Types. Holotype 6 (Bishop Mus.) and
12 paratypes (some in M.C.Z., Type No.
31,478) all from Swart Vy., Karubaka, N-E.
N. G., 1300 to 1600 m, dates in Nov. 1958
(holotype, 1300 m, Nov. 7) (Gressitt).
Additional material. N-E. N. G.: 2, Kas-
sem, 48 km E. of Kainantu, 1350 m, Nov. 7,
1959 (T. C. Maa, Bishop Mus.); 1, Tsenga,
Upper Jimmi Vy., 1200 m, July 13, 1955
(Gressitt); 1, Jim(m)i R., E. Highlands,
Julv-Sept. 1961 (W. W. Brandt, C.S.I.R.O.).
West N. G.: 1, Sibil, Star Rge., 1260 in.
May 24, 1959 (Neth. N. G. Exp., Leiden
Mus. ) .
Mea.sured s])eciniens. The •; holot\pc and
1 9 paratyjie from Karubaka.
Notes. P. huntcralis differs Irom the
preceding species (divisa) i)riucipall\ in
color, having less black on eUlral bas(>s
but darker legs. It is close also to fut)ti))es
but is larger and more lu>a\ily marked.
The interr(4ationships ol these forms are
still not clear.
The 3rd inter\als are usually .3-punetate
in humeralis but are only 2-punctate in the
indixidual from Sibil.
Demefrida imitafrix n. sp.
Description. See Plate 3, figure IX; with
characters of genus; relatively wide; dark
blue-black with dark appendages; not
pubescent, reticulate microsculpture absent
or faint, but surface finely sparsely punc-
tulate. Head 0.84 and 0.88 width prothorax;
eyes prominent, genae much shorter and
oblicpie. Frothorax subcordate; width
length 1.39 and 1.36; base apex 1.36 and
1.40; base head 1.02 and 1.02; sides
broadly slightly irregularly rounded an-
teriorly, strongly sinuate before c. right
slightly l)lunted posterior angles; margins
moderately wide, each with seta-bearing
puncture at or slightly before middle but
none at base; disc subpunctate baso-later-
ally. Elytra short and wide; width ebtra
prothorax 1.74 and 1.80; apices spined, outer
angles obtuse or blunted, sutural angles
obtusely blunted; striae impressed, slighth'
or scarcely punctulate; intervals nearly flat
or slightly convex, 3rd 2-punctate. Claics
with 4 or 5 teeth. Secondary sexual cliar-
acters: 6 tarsi as genus; o middle tibiae
with inner edge swollen or thickened before
apex, the swollen portion separated from
the apex b\' a broad emargination; i with
2 or 3, 9 c. 4 apical ventral setae each side.
Measurements: length 7.4-8.5; width 3.0-
3.5 mm.
Types. IIolotN'iie i (Bishop Mus.) from
Karimui, S. of Goroka, N-E. N. (i., 1000 m,
Inne 2, 1961 (Gressitt), taken in light trap;
and paratopes as follows. Papua: 1 broken
V, Dobodura, Mar.-Jul\ 1944 (Darlington)
(M.G.Z., Type No." 31,479), taken on a
lighted window; 1, Kokoda, 1200 ft. (366
m), Aug. 1933 (Gheesman); 1. Dogon,
Amazon Bay Dist., 2400 ft. ( r. 730 m).
Sept. 1962 (W. W. Brandt, C;.S.1.R.(). ); 1,
Misima Is., No\ . 1963 (\V. \V. Brandt,
C:.S.I.R.O.). West N. <;.: 1. Gamp 2,
Sabron. Gyclops Mts., 2000 ft. (610 m),
|nl\ 1936 ((>heesman).
The Carabid Beetles of New Guinea
Darlington
177
Measured specimens. The 6 holotype and
the 9 paratype from Dogon.
Notes. Among the New Guinean species
of Demetricla, this one is unique in its
broad form and in form of i tibiae. Never-
theless it has the essential characters of
Demetrida and I do not think it should be
separated from that genus, at least not
unless the genus as a whole is divided.
Superficially, D. imitatrix resembles Vio-
higonum violaceiim (Chaudoir), which
is very common at low altitudes in New
Guinea. This may be an example of
Batesian mimicry.
Demefrida viridipennis n. sp.
Description. See Plate 3, figure X; with
characters of genus; not pubescent; head
and prothorax red, elytra bright green usu-
ally shading to purple toward apex, ap-
pendages reddish yellow; reticulate micro-
sculpture visible ( light ) only on elytra, but
surface in part slightly sparsely punctu-
late. Head 1.08 and 1.11 width prothorax;
eyes moderately prominent, genae shorter
and oblique, not prominent. Prothorax
subquadrate; width length 1.08 and 1.03;
base/ apex 1.32 and 1.31; base head 0.88
and 0.86; basal angles c. right, c. blunted;
margins rather narro\\', each \\\i\\ seta near
or slightly liefore middle but none at base;
disc subpunctate across base and in margins.
Elytra: width elytra/prothorax 1.99 and
2.17 (latter spread by pin?); apices spined,
outer angles sharply defined, c. right ( some-
what variable), sutural angles obtuse; striae
deeply impressed, slightly punctulate; in-
tervals slightly convex, faintly punctulate,
3rd with 2 dorsal punctures. Claws with c.
5 teeth. Secondary sexual characters: S
tarsi as genus; c^ middle tibiae tuberculate-
serrate (c. 6 low tubercles); c5 with 2, $
3 setae each side last ventral segment.
Measurements: length 6.9-8.5; width 2.5-
3.1 mm.
Types. Holotype $ (Bishop Mus.) from
Wau, Morobe Dist., N-E. N. G., 1200-1300
m. Mar. 14, 1963 (Sedlacek); and para-
types (some in M.C.Z., Type No. 31,480)
as follows. N-E. N. G.: 4, Wau, 1200,
1220-1250, 1300 m, Nov. 12, 1961, Feb. 11,
Jan. 23, 1963, Oct. 14, 1965 (Sedlaceks);
1, Bulolo (near Wau), 1005 m, Aug. 25,
1956 (E. J. Ford, Jr., Bishop Mus.); 1,
Mt. Missim, 1600-2000 m, Sept. 21-24, 1964
(M. Sedlacek); 1, Karimui, 1080 m, Julv
14-15, 1963 (Sedlacek); 1, Okapa, Apr. 19,
1965 ( Hornabrook ) ; 1, Kainantu, July 9
(Sedlaceks); 1, \\\un. Upper Jimmi Valley,
840 m, July 17, 1955 (Gressitt); 1, Finis-
terre Rge., Saidor, Funvende, 1200 m, Sept.
24-30, 1958 (W. W. Brandt, Bishop Mus.);
1, Adalbert Mts., Wanuma, 800-1000 m,
Oct. 26, 1958 (Gressitt). Papua: 1, Ko-
koda, .200 ft. (366 m), June 1933
(Cheesman); 2, Owen Stanley Rge., Goilala
(Loloipa, Nov. 25-Dec. 10^ and Tapini,
975 m, Nov. 16-25, 1957) (W. W. Brandt,
Bishop Mus.); 1, Mt. Lamington, 1300-
1500 ft. (c. 400-460 m) (C. T. McNamara,
S. Australian Mus.).
Measured specimens. The i holotype and
1 9 paratype from Wau.
Notes. Form, color, and deep elytral
striae set this distinct species off from all
others of the genus kno\\'n to me. It is
evidently widely distributed in at least the
eastern half of New Guinea at moderate
altitudes.
Demetrido lepida n. sp.
Description. See Plate 3, figure XI; with
characters of genus; head and prothorax
black, elytra green-purple (variable, often
more green to\\'ard base and more purple
toward apex, sometimes slightly reddish on
disc), appendages dark; not pubescent,
reticulate microsculpture faint even on
elytra, surface not much punctulate. Head
1.24 and 1.12 width prothorax; eyes rather
abruptly prominent, genae c. long as eyes,
slightK' convex in outline but not very
prominent; front wide, irregularly flattened
and impressed or subpunctate at middle.
Prothorax subquadrate; width length 0.95
and 1.05; base apex 1.30 and 1.22; base/
head 0.74 and 0.77; sides arcuate anteriorly,
sinuate before c. right but blunted or nar-
178 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
rowly rounded posterior angles; margins
narrow, each with seta-bearing puncture at
or shghtly before middle but none at base;
disc convex, baso-lateral impressions almost
obsolete, surface faintly subpunctate across
base and in margins. Elytra: width elytra
prothorax — (elytra spread) and 1.95; apices
long-spined, outer angles acute or denticu-
late, sutural angles obtuse; striae lightly
impressed, lightly punctulate; intervals flat
or slightly convex, .3rd 3-punctate. Claws
with c. 7 teeth. Secondary sexual char-
acters: S tarsi as genus; S middle tibiae
slightly tuberculate-serrate {c. 4 spaced
tubercles); S with 3, 9 4-6 apical ventral
setae each side. Measurements: length
9.2-10.8; width 3.0-3.3 mm.
Types. Holotype 9 ( Bi.shop Mus. ) and
10 paratypes (some in M.C.Z., Type No.
31,-1(S1) from Swart Vy., Karubaka, N-E.
N. G., altitudes from 1300 to 1600 m, dates
in Nov. 1958 (holotype, 1450 m, Nov. 12)
(Gressitt); and additional paratypes as
follows. West N. G.: 17, Wissel Lakes,
Enarotadi, altitudes from 1750 to 1900 m,
dates in July, Aug. 1962 (Sedlacek); 1,
W'isscl Lakes, Itouda, Kamo Vv., 1500-1700
m, Aug. 18, 1962 (Sedlacek); 1, Wissel
Lakes, Kamo-Debei div., 1700 m, Aug. 13,
1955 (Gressitt); 1, Lower Mist Camp,
1700 m, Jan. 17, 1939 (Toxopeus). Papua:
1, Owen StanlcN' Hge., Goilala, Loloipa,
Feb. 1-15, 1958' (W. W. Brandt, Bishop
Mus.).
Measured specimens. A 6 paratype from
Swart Valley and the 9 holotype.
Notes. The bright color, rather abruptly
prominent eyes, and long elytral spines
characterize this fine species. It appears to
be widely distributed in New Guinea at
moderate altitudes. Ot the 19 specimens
seen, only 3 are i $ .
Two Karubaka individuals and one from
JMiarotadi are labe](>(l as taken in light
traps.
Demefrida sublepida n. sp.
Description. With characters of genus;
form r. as in preceding species {lepida)
but eyes less abruptly prominent and elytral
spines shorter; head and prothorax green,
elytra green-purple (variable); appendages
dark, tarsi paler; not pubescent, reticulate
microsculpture visible (faint) only on elytra,
surface not much punctulate. Head 1.16
and 1.11 width prothorax; eyes prominent
but not abrupt, genae shorter and oblique.
Prothorax subquadrate; width length 1.01
and 1.03; base apex 1.20 and 1.18; base/
head 0.80 and 0.86; sides weakly slightly
irregularly arcuate in more than anterior
■'4, usuallv stronglv sinuate before usuallv
acute posterior angles; margins narrow, each
with seta-bearing puncture slightly before
middle but none at base; baso-lateral im-
pressions subobsolete, disc slightly trans-
versely wrinkled, vaguely subpunctate
across base and in margins. Elytra: width
elytra prothorax 2.11 and 2.19; apices
spined, outer angles c. right or obtuse,
sutural angles obtuse; striae well impressed,
scarcely punctulate; intervals convex, finely
sparsely punctulate, 3rd 2-punctate. Cdaws
with 5 or 6 teeth. Secondary sexual cJiar-
acters: i unknown; 9 with 3 or 4 apical
ventral setae each side last ventral segment.
Measurenients: length 7.0-9.0; width 2.5-
3.1 mm.
Tyj)es. Holotype 9 (Bishop Mus.) from
Wissel Lakes, Enarotadi, West N. G., 1850
m, Aug. 1, 1962 (Sedlacek); and addi-
tional paratypes as follows. West N. (i.: 7
(some in \LC.Z., Type Xo. 31,482), Enaro-
tadi, 1750 to 1900 m, dates in July, Aug.
1962 (Sedlacek); 1, Wissel Lakes, IVapura,
Kamo Vv., 1530 m, Aug. 11, 1955 (Gressitt).
N-E. N.'g.: 1, Swart Vy., Karul)aka, 1300
m, Nov. 7, 1958 (Gressitt); 1, Wan, Morobe
Hist., 1.300 m. June 15, 1961 (Gressitt),
on Pii:)turus. All specimens are 9 9 .
Meastircd six'cinwns. Tlu' 9 holotxpe and
9 paratype from Enarotadi.
Soles. Although perhaps related to the
preceding spc^cies (lepida), sid)lcpid(i differs
in a surjirising number ol characters in-
cluding less abrupt eyes, head and pro-
notum green rather tlian black. el\ tral
spines slioitei-, elytial striae deeper, claws
The Carabid Beetles of New Guinea • Darlington 179
with fewer teeth, and size smaller. These
species evidently occur together at some
localities, and they may be involved in some
sort of mimicry.
Demetrida viridibasis n. sp.
Description. With characters of genus;
form slender, with ± prominent eyes and
short-spined elytra; red, c. basal Va of elytra
bright green with the green color extending
back more at sides than at middle, femora
and parts of tibiae dark; not pubescent,
reticulate microsculpture usually visible
(but light) on front of head and on pro-
notum as well as on elytra, much of upper
surface also sparsely finely punctulate.
Head 1.05 and 1.02 width prothorax; eyes
prominent, genae c. long as eyes and
slightly convex in outline but not very
prominent. Frothorax subquadrate, long;
width/length 1.00 and 1.09; base/apex 1.25
and 1.23; base/head 0.85 and 0.85; sides
nearly straight for much of length except
slightly subangulate at setae, broadly sinu-
ate before c. right slightly blunted posterior
angles; margins narrow, each with seta-
bearing puncture slightly before middle but
none at base; disc less convex and with
more distinct baso-lateral impressions than
in k'pida and suhJepida, subpunctate across
base and in margins. Elytra moderately
long; width elytra prothorax 1.98 and 1.86;
apices short-spined, outer angles c. right,
sharply formed, sutural angles obtuse; striae
impressed, faintly punctulate; intervals con-
vex, 3rd 3-punctate. Cdaics with c. 5 teeth.
Secondary sexual characters: i see Notes,
below; 9 with 5 or more apical ventral
setae each side. Measurements: length
8.5-10.3; width 2.9-3.5 mm.
Types. Holotype 9 (C.S.I.R.O., Can-
berra ) from Dogon, Amazon Bay Dist., SE.
Papua, 2400 ft. (c. 730 m), ^Sept. 1962
(W . \V. Brandt); 1 9 paratype (C.S.I.R.O.)
with same data except collected Oct.-Nov.;
1 9 paratype (M.C.Z., Type No. 31,483),
Doveta, Amazon Bay Dist., 2400 ft. (c. 730
m), Aug. 1962 (W. W. Brandt).
Additional material. N-E. N. G.: 3,
Swart Vy., Karubaka, 1300, 1500 m, Nov. 7,
11, 20, 1958 (Gressitt); 1, Finisterre Rge.,
Saidor, Kiambavi Village, Aug. 1-28, 1958
(W. W. Brandt, Bishop Mus.). West N. G.:
1, Cyclops Mts., 3400-4500 ft. (c. 1040-
1370 m). Mar. 1936 (Cheesman).
Measured specimens. The 9 holotype and
9 paratype from Doveta.
Notes. This species seems close to the
green-marked form of diversa but has
longer elytral spines. Some specimens
listed under Additional material are doubt-
fully identified. Most are 9 9 ; the only S ,
from Swart \'y., has middle tibiae slightly
bent in toward apex but not tuberculate-
serrate, and c. 4 apical ventral setae each
side.
Demetrida sibil n. sp.
Description. With characters of genus;
form slender, with prominent eyes and
spined elytra; head, prothorax, and c. basal
Vi of elytra dark greenish, the dark color
extending farther back at sides of elytra
than at middle, and suture sometimes red
almost to base, rest of elytra red, femora
and outer edges of tibiae greenish black,
antennae brown, lo^^'er surface greenish
black in anterior half, abdomen red; not
pubescent, reticulate microsculpture faint
even on elvtra. Head 1.19 and 1.10 width
prothorax; eyes moderately abruptly promi-
nent, genae nearly as long as eyes, oblique;
front flattened, irregularly slightly im-
pressed and subpunctate at middle. Pro-
thorax subquadrate; width length 0.94 and
0.98; base apex 1.38 and 1.28; base head
0.81 and 0.82; sides weakly irregularly
arcuate for much of length, rather abruptly
sinuate before right or slightly acute some-
times slightly blunted posterior angles;
margins narrow, each with seta-bearing
puncture slightly before middle but none at
base; disc strongly convex, with baso-lat-
eral impressions weakly indicated, surface
slightly punctate across base and in margins.
Ehjtra: width elytra/ prothorax 2.09 and
1.98; apices spined, outer angles acute or
denticulate, sutural angles obtuse; striae
180 Bulletin Museum of Co7y}parative Zoology, Vol. 137, No. 1
moderately impressed, finely punetulate; in-
tervals slightly convex, very sparsely in-
conspicuously punetulate, 3rd 3-punctate
(except intermediate puncture lacking on 1
side in 1 paratype). CAciws with c. 6 teeth.
Secondary sexual character: 6 tarsi as
genus; 6 middle tibiae bent-in at apex but
not tuberculate-serrate; c5 with apparently
4, 9 c. 8 or 9 apical ventral setae each side.
Measurements: length 9.4-10.8; width 3.0-
3.4 mm.
Types. Holotype $ (Leiden Mus.) and 5
paratypes (2 hi M.C.Z., Type No. 31,484)
from Sibil, Star Rge., West N. G., 1260 m,
dates in May and June, 1959 (holotype,
June 17) (Neth. New Guinea Exp.), taken
at light; and additional paratypes as fol-
lows. West N. G. : 2, preceding locality
("Star Mts. Sibil Val."), 1245 m, Oct. 18-
Nov. 8, 1961 (S. Quate, Bishop Mus.), at
light. N-E. N. G.: 1, Feramin, 1200-1500
m, June 15-18, 1959 (W. W. Brandt,
Bishop Mus.).
Measured specimens. The i holotype and
1 9 paratype from Sibil.
Notes. The color (head and pronotum as
well as elytra! bases green ) and long-spined
elytra distinguish this from other species of
the diversa complex.
Demetrida seticollis n. sp.
Description. With characters of genus;
form as in Figure 109; brown, with head
and prothorax and sometimes base of elytra
slightly darker; appendages brown; not
pubescent, reticulate inicrosculpture visible
(but very light) only on elytra. Head 1.20
and 1.19 width prothorax; eyes \('r\' promi-
nent, genae {)bli(|ue and not sharply distinct
from neck; front irregularly flattened or
impressed and subpunctate at middle. Pro-
thorax (piadrate-trapezoidal; width /length
1.02 and 0.97; base apex 1.45 ;uid 1.29;
base/head 0.86 and 0.90; sides wcakK
arcuate or c. straight in "'i or more ol length,
strongly sinuate beh)re prominent, c. light
or acute (somewhat varia])le ) basal angles;
margins narrow, each \\ itli seta before mid-
dle and at base and additi<)n;il iisualK
smaller setae anteriorly; disc moderately
convex, with irregular baso-lateral impres-
sions, surface irregular or subpunctate
across base and in margins. Elytra ample;
width elytra/prothorax 2.20 and 2.22; apices
long-spined, outer angles acute-denticulate,
sutural angles obtuse; striae lightly im-
pressed, finely punetulate; intervals slightly
convex or c. flat, 3rd 3- or 4-punctate ( vari-
able, sometimes unsymmetric). Claws with
6-8 teeth. Secondary sexual characters: $
tarsi as genus; i middle tibiae tuberculate-
serrate ( c. 4 widely sometimes irregularly
spaced tubercles); 6 with 2-A, 9 c. 6
apical ventral setae each side. Measure-
ments: length 8.7-10.5; width 2.9-3.4 mm
( except 1 i from Wissel Lakes, doubtfully
identified, 11.3 X 3.7 mm).
Types. Holotvpe 6 ( Bishop Mus. ) from
Wissel Lakes, Enarotadi, West N. G., 1900
m, Aug. 21, 1955 (Gressitt); 61 paratypes
(some in M.C.Z., Type No. 31,485) from
the Wissel Lakes area (Enarotadi, Moane-
mane, Itouda, Urapura, Okaitadi, "Paniai-
Kamo div."), 1500-2050 m, dates in Jul\-,
Aug., 1955, 1962 (Gressitt, Sedlacek );" and
4 additional paratvpcs from the same area,
Arabu Camp, 1800 m, Oct. 7, 8, 12, 17,
1939 (H. Boschma, Leiden Mus.).
Additional material. West N. G. : 1 very
large 6 , data as holot)pe except 1500 m,
Aug. 14, 1962 (Sedlacek); 2, Juliana Bivak,
1800 m, Aug. 30, Sept. 5, 1959 (Neth. N. G.
Exp., Leiden Mus.); 1, Star Rge., Bivak
39A, 1500 m, July 2, 1959 (Neth. N. G.
Exp., Leiden Mus.); 1, Swart Vv., \^^ ridge,
1800-2000 m, Nov. 19, 1958' (Gressitt).
N-i:. ^, (;.: l, Gewak, Salawaket Hge.,
1530 ni, Sept. 6, 1956 ( E. J. Ford. Jr..
Bishop Mus.), in light trap. Papua: 1,
Owen Stanle\- Hge., Cioilala, Bom(\ 1950 m.
Mar. 8-15, 1958 ( W. \\ . Brandt, Bishop
Mus.).
Measured specimens. The S holotype and
I 9 paratype from liinarotadi.
Notes. The extra seta of the prothoracic
margins antciioily distinguish this species
hoin all other noiniubescent Demetrida
known lo ww. These setae aic much
The Carabid Beetles of New Guinea • Darlington 181
stronger and more erect than the fine extra
marginal hairs of D. seriata and niibicola.
When the setae are ]:)roken off, or perhaps
lacking in aberrant individuals, the species
is still recognizable by form especially of
prothorax, color, and long elytral spines.
D. scticoUis apparently ranges widely in
New Guinea at considerable altitudes (not
known below 1500 m ) on the higher moun-
tain ranges. The fact that it has not been
found on the Morobe Plateau (Wau, etc.)
is noteworthy.
Demefrida pollipes n. sp.
Description. With characters of genus;
form slender, with moderately prominent
eyes and strongly spined elytral apices; head
and pronotum reddish piceous, elytra
blackish with small discal area usually
reddish, legs testaceous, antennae brown,
lower surface dark with metepisterna paler;
not pubescent, reticulate microsculpture
absent or indistinct, surface not much punc-
tulate. Head 1.14 and 1.16 width prothorax;
eyes prominent, genae shorter, oblique.
P rot J} o rax subquadratc, long; width length
1.03 and 0.96; base apex 1.35 and 1.34;
base/head 0.84 and 0.86; sides very weakly
arcuate in c. anterior %, strongly sinuate
before right or acute usually slightly
blunted posterior angles; margins narrow,
each with seta-bearing puncture before
middle and at basal angle; disc with mod-
erate baso-lateral impressions, scarcely
punctate. Elytra: width elytra /prothorax
2.04 and 2.05; apices strongly spined, outer
angles acute or denticulate, sutural angles
obtuse; striae lightly impressed, finely ir-
regularly punctulate; intervals almost flat,
3rd usually 2-punctate, sometimes 3-punc-
tate on 1 side. Claws with c. 5 teeth. Sec-
ondary sexual characters: £ tarsi as genus;
S middle tibiae weakly tuberculate-serrate
(c. 4 low tubercles); S with 3, 9 5 or 6
apical ventral setae each side. Measure-
ments: length 8.4-9.6; width 2.5-2.9 mm.
Types. Holotype S (Bishop Mus.) and
19 paratypes (some in M.C.Z., Type No.
31,486) from Wau, Morobe Dist., N-E.
N. G., altitudes from 1100-1500 m, dates
in Jan., Feb., Mar., Apr., Sept., Dec, 1961-
1966 (holotype, 1450 m, Feb. 6, 1963)
(Sedlacek, 1 paratype T. C. Maa); 1 para-
type, Mt. Missim,' 1600-2000 m, Sept. 21-
24, 1964 (M. Sedlacek).
Measured specimens. The S holotype and
1 9 paratype.
Notes. See Key to Species for place of
D. pallipes among other New Guinean
Dcmetrida.
Demetrido discoidalis n. sp.
Description. With characters of genus;
form c. as in preceding species i))allipes);
black, elytra with large elongate common
red area centered behind middle, lower
surface and appendages dark; not pubes-
cent, reticulate microsculpture absent or
indistinct on elytra, surface not much
(sparsely inconspicuously) punctulate. Head
1.06 and 1.13 width prothorax; eyes promi-
nent, genae shorter, oblique. Frothorax sub-
quadrate, but anterior angles rounded to
neck; width length 1.04 and 1.00; base/
apex 1.51 and 1.47; base/head 0.90 and
0.91; sides arcuate through much of length,
sinuate before c. right slightly blunted pos-
terior angles; margins narrow, each with
seta-bearing puncture before middle and at
base; disc with baso-lateral impressions
weak, surface slightly irregular or sub-
punctate across base and in margins.
Elytra: width elytra/prothorax 1.91 and
2.02; apices ( variably ) spined, outer angles
sharply defined, varying from acute to
slightly obtuse, sutural angles obtuse;
striae lightly impressed, punctulate; inter-
vals flat, 3rd 2-punctate. Claws with 6 or
7 teeth. Secondary sexual characters: S
tarsi as genus; 6 middle tibiae tuberculate-
serrate (c. 4 low rounded tubercles); S
with 2 or 3, 9 4 or 5 apical ventral setae
each side. Measurements: length 9.0-11.5;
width 3.0-3.7 mm.
Types. Holotype S (Bishop Mus.) from
Sibii Vy., Star Rge., West N. G., 1245 m,
182 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Oct. 18-Nov. 8, 1961 (S. & L. Quate), in
Malaise trap; and paratypes as follows.
West N. G.: 1, Sibil, Star Rge., 1260 m,
June 1959; 2, Bivak 36, Star Rge., 1220 m,
July 28, 1959; 1, Bivak 39A, Star Rge.,
1550 m, July 5, 1959 (these 4 paratypes all
Neth. N. G. Exp., Leiden Mus.)'. N-E.
N. G.: 1, Eliptamin Vv., 1665-2530 m,
June 19, 1959 (W. W. Brandt, Bishop Mus.);
1, Feramin, 1200-1500 m, June 15-18, 1959
(W. W. Brandt, Bishop Mus.). (Some
paratypes in M.C.Z., Type No. 31,487. )
Measured specimens. The S holotype and
9 paratype from Bivak 39A.
Notes. This may be a geographic repre-
sentative of the preceding species ( paUipes)
from which it differs only slightly in form
but more in color, with larger red area on
elytra and dark rather than pale legs. It
resembles dorsaJis in color but differs in
form (much narrower than dorsaJis), pres-
ence of posterior-lateral prothoracic setae,
and in other ways: in fact, these 2 species
are not closely related.
Demetrida sedlacekorum n. sp.
Description. With characters of genus;
form slender, c. as in preceding species
{paUipes, discoidaJis) ])ut elytra short-
spined; black with bluish tone especially
on elytra, appendages dark; not pubescent;
reticulate microsculpturc> absent or indis-
tinct, but upper surface in part with \er)'
fine, sparse, inconspicuous punctulation.
Head 1.14 and 1.08 width prothorax; eyes
prominent, genae shorter, oblique. Pro-
thorax subcjuadrate, long, with rathcM- broad
base; width length 0.98 and 0.99; base apex
1.41 and 1.43; base/head 0.86 and 0.87;
sides weakly arcuate in c. Ti of length,
sinuate before c. right but variabl(\ blunted
posterior angles; margins rather narrow,
each with seta-bearing imncturc^ before
middle and at l:)asal angle; disc with baso-
lateral impressions deep but small, sub-
|)unctat('. i'Jijtrd: width el\tra prothorax
2.03 and 1.93; apices short-spined, outer
angles sharply defined, usually acute, su-
tural angles obtuse; striae lightly impressed.
punctulate; intervals flat or slighth- convex,
third usually 2-, sometimes 3-punctate.
Claws with 6 or 7 teeth. Secondary sexual
characters: S tarsi as genus; i middle
tibiae weakly tuberculate-serrate (margin
wavy); i with 2 or 3, 9 4 or 5 apical
ventral setae each side. Measurements:
length 8.5-9.8; width 2.6-3.1 mm.
Types. Holotype c^ (Bishop Mus.) and
28 paratypes (some in M.C.Z., Type No.
31,488) all from Wau, Morobe Dist., N-E.
N. G., altitudes from 1180 to 1500 m, dates
in Jan., Feb., Mar., Apr., June, Sept., Nov.,
1961-1964 (holotype, 1220-1250 m, Jan. 23,
1963) (Sedlaceks).
Additional material. N-E. N. G.: 2,
Jim(m)i R., E. Highlands, July-Sept., 1961
"(W. W. Brandt, C.S.I.R.O.). Papua: 1,
Ow^en Stanlev Rge., Goilala, Tororo, 1560
m, Feb. 21-24, 1958 {W . W. Brandt,
Bishop Mus.).
Measured specimens. The i holotype and
1 9 paratype.
Notes. See final couplets of Key to
Species for place of sedlacekortim among
other New Guinean Demetrida.
Demetrida brandti n. sp.
Description. See Plate 3, figure XII;
with characters of genus; form c. of paUipes
and discoidalis, slender, with long-spined
elytra; color entirely bhu^-black. with dark
appendages; not pubescent, microsculpture
\irtuallv absent even on elvtra, surface not
much (very finely, sparsely, inconspicu-
ously) punctulate. Uead 1.09 and 1.07
width prothorax; c\es prominent, genae
shorter, oblicine. Prothorax ({uadrate, long;
width ItMigth 0.98 and 1.03; base ;ipex 1.44
and 1.43; base head 0.87 and 0.88; sides
weakly arcuate in c. ant(Mior 'H, broadly
sinuate before c. right or slightK" acute
posterior angles; margins rather narrow
each with seta near or before middle and
at base; baso-l;iteral impressions moderate,
subpunctate. Elytra: width elytra pro-
thorax 1.98 and 1.94; apices with moderately
long spines, outer angles wc-ll defined, ±
ritiht, sutural angles blunted; striae sli<j:htl\
The Carabid Beetles of New Guinea • Daiiin<'toii
183
impressed, faintly punctulate; intervals
slightly convex, 3rd 2-punctate. Claws with
c. 6 teeth. Secondary sextial characters: i
tarsi as genus; 6 middle tibiae tuberculate-
serrate ( c. 4 tubercles ) ; i with 3, 9 4 or 5
apical ventral setae each side. Measure-
ments: length 8.8-10.2; width 2.8-3.3 mm.
Types. Holotype i (Bishop Mus.) and
2 paratypes from Finisterre Rge., Saidor,
Kiambavi Village, N-E. N. G., 1400 m,
July 22-29 (holotype), Aug. 1-28 (para-
types); 1 paratype, Saidor, Funyende, 1200
m, Sept. 24; 2 paratypes, Saidor, Matoko,
Aug. 29-Sept. 5, Sept. 6-24 (all collected
1958 by W. W. Brandt for Bishop Mus.;
some paratypes now in M.C.Z., Type No.
31,489).
Additional material. N-E. N. G.: 2, Swart
Vy., Karubaka, 1500 m, Sept. 20, 1958
(Gressitt); 1, Gewak, Salawaket Rge., 1530
m, Sept. 6, 1956 (E. J. Ford, Jr., Bishop
Mus.), in light trap. West N. G.: 2,
Wamena, 1700 m, Feb. 10-25, 1960 (T. C.
Maa, Bishop Mus.). Papua: 1, Purosa
Camp, Okapa area, 1950 m, Sept. 23,
1959 (L. J. Brass, Sixth Archbold Exp.,
A.M.N.H.).
Measured specimens. The i holotype and
1 9 paratype from Kiambavi.
Notes. This will probably prove to be a
geographic subspecies of sedlacekorum dis-
tinguished mainly by longer elytral spines.
Genus PHLOEOCARABUS Macleay
Macleay 1871, Trans. Ent. Soc. New South Wales
2, p. 85.
Sloane 1898, Proc. Linnean Soc. New South Wales
23, p. 499.
Csiki 1932, Coleop. Cat., Caraliidae, Harpalinae 7,
p. 1488 (see for additional references, synonymy,
and list of species ) .
Diagnosis. See Key to Genera of Lebiini
of New Guinea and Figure 110.
Description ( characters common to the
2 New Guinean species). Form c. as in
Figure 110; color diverse; not pubescent.
Head: eyes large, prominent; 2 setae over
each eye; antennae pubescent from middle
of 4th segments; front with long, slightly
curved costa on each side passing inside
position of anterior seta; clypeus transverse,
1-setose each side; labrum wide, arcuate-
truncate, 6-setose; mentum with long, entire
tooth; ligula subtruncate with 2 principal
setae, paraglossae attached to ligula, nar-
rowed and rounded to apex of ligula; palpi
rather short, apical segments of labial palpi
widened, c. triangular. Prothorax transverse,
arcuately narrowed anteriorly, slightly lobed
at base; margins rather wide, flat, scarcely
reflexed, each with usual 2 setae; disc with
impressed middle line and weak transverse
impressions; base with fine marginal line
entire or nearly so, apex not margined at
middle. Elytra with rounded, slightly nar-
rowed humeri; apices obliquely sinuate-
truncate, with outer angles broadly and
inner angles narrowly rounded; striae entire,
moderately impressed, not punctate; in-
tervals not specially elevated at base, 3rd
2-punctate with punctures before middle
on outer edge and behind apical % near
inner edge. Inner icings full. Legs mod-
erate; 4th segments middle and hind tarsi
emarginate; 5th segments with accessory
setae; claws each with c. 4 rather long
teeth. Secondary .sexual characters: i front
tarsi scarcely dilated, 2-seriately squamu-
lose; i middle tarsi also squamulose; i
middle tibiae not excised; 6 with 1 prin-
cipal (sometimes a 2nd smaller), 9 2 setae
each side last ventral segment.
Type species. P. mastersi Macleay, of
Australia.
Generic distribution. Australia, with 1
Australian species extending to New
Guinea and New Britain, and an addi-
tional species endemic in New Guinea.
Notes. The 2 species here assigned to
Phloeocarahus are very different super-
ficially but share the technical characters
of the genus.
Key to Species of Phloeocarabvs of
NE^v Guinea
1. Color black or piceous, elytra with testa-
ceous marks (p. 184) nigricollis
- Strikingly bicolored, head and prothorax red,
elytra bine (p. 184) -— euplenes
184 Bulletin Museuni of Comparative Zoology, Vol. 137, No. 1
Phloeocarobus nigricoHis (Macleay)
Macleay 1864, Trans. Ent. Soc. New South W'ak-s
1, p. Ill {Trigoiwthoiis) .
See also references under yenus.
hasalis Sloane 1907, Deutsche Eut. Zeitschrift for
1907, p. 182 (new synonymy).
Description. None required here; length
c. 6-8 mm.
Types. Of nipicoUis, from Port Denison
( Bowen ) , Queensland, Australia, presum-
ably in Macleay Mus., Sydney; of basalis,
from the Gazelle Pen., New Britain, should
be in Deutsche Ent. Institut, Berlin-
Dahlem ( none seen ) .
Occurrence in New Guinea. Thirty-four
specimens, from localities covering almost
the whole length of New Guinea, most at
low altitudes but records up to 1300 (at
Wau), 1400, and 1500 m.
Notes. Sloane distinguished hasalis from
niii,rieollis by a slight color difference,
which does not hold in the series before me:
5 specimens from New Britain include both
individuals with base of elytra entirely dark
{hasalis) and individuals with the pale
marks reaching the elytral base (nigricollis).
The series from New Ckiinea is even more
variable, with elytra ranging from almost
wholly reddish testaceous to almost wholly
piceous with subbasal pale marks scarcely
indicated. The pronotum also varies, from
reddish piceous with pale margins to red-
dish testaceous, and the variation is partly
independent of the variation of elytral
pattern. All these color forms seem to me
to be one species. The variation of pattern
may prove to be partly geograjihic, but the
material liefore me is not suHicient to
establish this.
Phloeocarobus euplenes n. sp.
Descrij)tion. With characters of genus;
lonn as in T^igure 110; strikingly bicolored,
head and prothorax red, elytra blue, lower
surface and appendages reddish testaceous;
moderateh' shining, reticulate niierosculp-
ture absent or hunt on bout and on disc
oi pronotum, distinct and slightK trans-
verse on cKlra. Head: O.Tf) and 0.76 w idlh
prothorax; front scarcely impressed, faintly
sparsely punctulate. Prothorax: width/
length 1.58 and 1.62; base/apex 1.67 and
1.62; margins broadly flattened, not much
reflexed; disc sparsely irregularly punctu-
late and faintly transversely strigulose.
Elytra: width elytra prothorax 1.42 and
1.47; striae well impressed, faintly punctu-
late; intervals slightly convex, each with an
irregular row of punctules along middle.
Secondary sexual characters: i as for genus;
9 unknown. Measurements: length 5.0-
5.2; width 2.1-2.2 mm.
Types. Holotype 6 (Bishop Mus.) from
Torricelli Mts., Wantipi Village, N-E. N. G.,
Nov. 30-Dec. 8, 1958 (W. W. Brandt);
and 1 i paratvpe (M.C.Z., Type No.
31,490) from Kiunga, Fly R., Papua, T'llv
11-14, 1957 (W. W. Brandt).
Notes. This new Phloeocarahus is strik-
ingly different in color from any other
member of the genus known to me. In
form and color it resembles and may mimic
New Guinean species of the agonine genus
Eui)lenes.
Genus TRIGONOTHOPS Macleay
Macleay 1804, Trans. Eut. Soc. New South Wales
1, p. 110.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1488 (see for additional references and list
of species).
DiaiS.nosis. Sec^ Key to Genera of Lehii)}i
of New Guinea and Notes under the follow-
ing species.
Description. None rt>quired here.
Type s))ecies. Calleida pacifica Erichson
of Australia (original designation).
Generic distribution. TT(>retofore known
onK' horn Australia including Tasmania
and oilier elose-King islands; range now
extended lo New (>uinea.
Notes. The Australian numbers ot this
geinis li\(" on tree trunks. 1 think.
Trigonothops lateralis n. sp.
Description, i-'onii as in f'igure 111; en-
tireU (slightly reddish) xellow except h:)r
a wide biowiiish black stripe along the
The Carabid Beetles of New Guinea
Darlinp,ton
185
outer side of each elytron (not including
the reflexed margin) from humerus nearly
to apex; not pubescent; moderately shining,
with lightly impressed reticulate micro-
sculpture isodiametric on front, slightly
transverse on disc of pronotum and elytra.
Head 0.86 width prothorax; 2 setae over
each eye; front slightly impressed at middle,
longitudinally impressed each side anteri-
orly; clypeus broadly rounded anteriorly
( a small notch at middle may be abnormal ),
1-setose each side; labrum transverse, ir-
regularly broadly rounded in front, 6-setose;
mentum with triangular tooth; ligula thick-
ened, blunt, probably originally setose but
setae broken short; paraglossae c. long as
or slightly shorter than ligula, attached
except at extreme apex, apparently with-
out setae; maxillary palpi not or not much
thickened, labial palpi with apical seg-
ments wider, not quite ^2 wide as long,
narrowly obliquely truncate. Prothorax
subcordate but with broad base; width/
length 1.44; base/apex 1.54; base broadly
briefly lobed, margined; apex broadly emar-
ginate, not distinctly margined at middle;
side margins broad and reflexed especially
toward base, each with 2 seta-bearing punc-
tures, at basal angle and c. % from apex;
disc with middle line deep, transverse im-
pressions less well defined, baso-lateral
foveae deep but not sharply limited; surface
of disc with faint weak transverse strigula-
tion, almost impunctate. Elytra: width
elytra/prothorax 1.94; humeri slightly nar-
rowed and rounded but not obliterated;
reflexed lateral margins moderate; apices
slightly obliquely truncate and very slightly
sinuate, with outer angles broadly and su-
tural angles narrowly rounded; striae entire,
moderately impressed, faintly irregular but
not distinctly punctulate; intervals slightly
convex, not distinctly punctate except 3rd
with 2 small dorsal punctures, on inner edge
just before middle and behind apical Vi (sub-
basal puncture, if present, minute and not
surely detectable). Inner wing,s full. Legs:
hind tarsi missing; middle tarsi with 4th
segments very deeply emarginate, with
lobes much longer than V2 length of seg-
ment; claws (of front tarsi) each with 4
long teeth and apparently an additional
very short tooth toward base; 5th segments
(of front tarsi) with accessory setae. Sec-
ondary sexual characters: i unknown; $
with last ventral segment subtruncate,
slightly subsinuate at middle, with 2 setae
near apex each side. Measurements: length
6.7; width 2.9 mm.
Type. Holotype 9 (Leiden Mus.) from
Wissel Lakes, central West N. G., Arabu
Camp, 1800 m, Oct. 12, 1939 (H. Boschma);
the type is unique.
Notes. Even without the 6 I am rea-
sonably sure that this insect is a Trigono-
thops. It agrees with T. pacificus Erichson
in form and in significant characters in-
cluding the mouthparts (see preceding
Description ) and position of the dorsal
elytral punctures. Moreover, the elytral
color pattern is derivable from that of
Trigonotliops pacificus: lateral stripes like
those of hiteralis would be left if the inner
portion of the dark elytral pattern of
pacificus were erased.
This individual had lost most of its legs
before I received it. Only the left front leg
is still complete, and the left middle leg is
complete except for the 5th segment. But
these are enough to show essential char-
acters, and the specimen is in good condi-
tion otherwise.
Genus NOTOTARUS Chaudoir
Chaudoir 1875, Bull. Soc. Nat. Moscow 49, Part
2, p. 19.
Sloane 1898, Proc. Linnean Soc. New South
Wales 23, p. 494.
Diagnosis. As Aiu)motarus (following
genus) but side pieces of metasternum short,
scarcely longer than wide; and (in the New
Guinean species) genae short-setulose; an-
tennae with segments 2 and 3 more or less
pubescent; tarsi pubescent (sparsely short-
pilose) above; i middle tibiae not modi-
fied.
186 Bulletin Mit.'icum of Coinparatwv Zoolof^y, Vol. 137, No. 1
Description. See Notes (below) and de-
tailed Description of following species.
Ti/j)e species. Nototariis australis Chau-
doir, of Western Australia.
Generic distribution. Previously known
only from Australia; range now extended
to New Guinea.
Notes. Chaudoir described Nototarus as
without a mentum tooth, and the tooth is
certainly difficult to see in some Australian
species, but it may be depigmented rather
than absent. The characters and generic
classification of this group of Carabidae
need further study, which will have to be
based on the Australian rather than New
Guinean forms. Some Australian species
assigned to Nototarus by Sloane do have
the mentum toothed, and the single New
Guinean species (below) is evidently
closely related to some of them.
Nototarus papua n. sp.
Description. Form as in Figure 112;
brownish black, humeri broadly paler
brown, appendages brownish testaceous;
reticulate microsculpture lightly impressed,
c. isodiametric on front, slightly transverse
on pronotum, more irregular on elytra, and
surface irregularly rather sparsely punctu-
late. Head 0.81 and 0.80 width prothorax;
eyes moderately prominent, genae rounded-
oblique, short-setulose; 2 setae over each
eye; front longitudinally rugulose each side;
clypeus slightly emarginate-truncate, 1-
setose each side; labrum wide, slightly
emarginate-truncate, 6-setose; mandibles
rather short, curved; antennae moderate,
pubescent from middle of 3rd segments;
mentum w ith long, narrowly rounded tooth;
ligula 2-setose, paraglossae attached, e(iual
in length, wide, not setose; maxillary palpi
slender, labial palpi with apical segments
wide. Prothorax cordate, short-lobed at
base; width length 1.31 and 1.30; base
apex 0.88 and 0.92; margins narrow, each
with seta at basal angle and c. ' i from apex;
basal and apical marginal lines inti-rrupted
at middle; disc with middle line coarse,
almost entire, transxcrse impressions almost
obsolete. Elytra short, narrowed toward
base, connate; width elytra/prothorax 1.49
and 1.51; striae entire, impressed, faintly
punctulate; intervals convex especially
toward base, 3rd 2-punctate, the punctures
near middle of length and c. V4 from apex.
Inner icings vestigial. Lower swjace not
obviously pubescent ( in part with very
short inconspicuous sparse pubescence).
Le^s moderate; tarsi short-pilose above;
4th segments middle and hind tarsi slighth"
emarginate; 5th segments with long ac-
cessory setae; claws each with 3 or 4 teeth.
Secondary sexual characters: labial palpi
with apical segments wider in c^ (truncate
apex wider than length of inner edge),
narrower in 9 (truncate apex narrower
than inner edge); 6 front tarsi slightly
( scarcely ) dilated, 3 segments 2-seriately
squamulose; i middle tarsi not squamu-
lose; i middle tibiae not modified; $ with
1, 92 seta-bearing punctures each side
last ventral segment. Measurements: length
4.6-5.3; width 1.9-2.3 mm.
Types. Holotype 6 (M.C.Z., Type No.
31,491) and 20 paratypes all from Dobo-
dura, Pa|>ua, Mar.-July 1944 (Darlington).
Measured specimens. The c^ holot\'pe and
1 9 paratype.
Notes. This is the onl\' Nototarus thus
far found in New Guinea. It may be re-
lated to N. morosus Sloane of Port Darwin,
Australia, but has thc^ prothorax exidently
more narrow(>d posteriori), with sinuate
sides. Other, apparently related, unde-
scribed spicies occur in North Queensland.
This species is one of the very few strictly
flightless Garabidae found at low altitudes
in New Guinea. Most or all of in\- speci-
mens were, I think, taken in flood debris
from the floor of rain forest. The rain-forest-
floor habitat and the insect's tlightlessness
pi'rhai)s explain w h\ other collectors have
not tound it.
Genus ANOMOTARUS Chaudoir
Chaiuloii I,S7.^, linll. Soc. Nat. Moscow U), Part 2,
p. \H.
The Carabid Beetles of New Guinea • Darlington 187
Sloane 1898, Pioc. Linnean Soc. New South Wales
23, p. 494.
Csiki 1932, Coleop. Cat., Caraliidae, Harpalinae 7,
p. 1493 ( see for additional references, synonymy,
and list of species ) .
Tedlicka 1963, Ent. Abhandlungen 28, pp. 300,
450.
Diap,nosis. See Key to Genera of Lebiini
of New Guinea.
Deseription (based on New Giiinean
species only). Fomi as in Figures 113-
117; small, slender, depressed; elytra usu-
ally (not always) with characteristic pale
marks; not pubescent. Head: eyes mod-
erately prominent, genae rounded, more or
less prominent ( usually less so than eyes ) ;
2 setae over each eye; antennae moderate,
pubescent from 4th segment (3rd segment
with only usual apical setae); clypeus trans-
verse with rounded angles, 1-setose each
side; labrum transverse, broadly emargi-
nate, 6-setose; mandibles short, strongly
curved; mentum with long tooth; ligula
rather broad, 2-setose, paraglossae attached
and c. equal in length, wide, not setose;
maxillary palpi moderate, not widened;
labial palpi with apical segments widened.
Frothorax cordate, with base briefly lobed;
side margins moderate or narrow, with
setae at basal angle and c. V-i or % from
apex; disc with impressed middle line and
less distinct transverse impressions. Elytra
with apices simple; striae entire, not dis-
tinctly punctate; 3rd intervals 2-punctate,
with punctures near or before middle and
c. Vi from apex; 8th intervals usually finely
carinate on inner edge near base (carinae
sometimes so fine as to be scarcely detect-
able). Inner wings iuW. Lotver swiace not
extensively pubescent; side pieces of meta-
sternum long. Legs slender; tarsi not pubes-
cent above; 4th segments middle and hind
tarsi slightly emarginate; 5th segment with
long accessory setae; claws with c. 4 or 5
teeth. Secondary sexual characters: i front
tarsi slightly (scarcely) dilated, 3 segments
2-seriately squamulose; last segment i labial
palpus wider (truncate outer edge almost
as long as inner side), of 9 less wide; 6
middle tarsi without squamae; i middle
tibiae with inner edge tuberculate-serrate
( cf . Demetrida ) , with c. 3 or 4 low tuber-
cles in row toward apex ( in all New
Guinean species of which the c5 is known ) ;
i with 1, 9 2 setae each side last ventral
segment.
Type species. Anomotarus olivaceus
Chaudoir, from Melbourne, Australia.
Generic distribution. Southern Asia
(Ceylon, India, Japan, etc.) to Australia
and Tasmania, and New Caledonia.
Notes. Although I recognize 8 (closely
interallied) species of Anomotarus in New
Guinea, material is scanty and almost
nothing is known of their habits. I think
most of them probably live among dead
leaves on the ground in rain forest. A few
specimens have been taken at light.
Key to Species of Anomotarus of New CIuinea
1. Each elytron either with longitudinal post-
humeral stripe (outside 4th stria) and sub-
apical-sutural spot pale, or with only the
subapical spot, or unmarked 2
- Each eh tron with a more or less incomplete
oblique or transverse (not longitudinal) spot
or band before middle and usually (not
always ) a subapical-sutural spot pale 5
2. Very slender (prothoracic width/length
1.15); subsericeous black, unmarked (p.
188 ) gressitti
- Less slender (prothoracic width/length c.
1.20 or more); elytra with at least subapical-
sutural pale spot(s) 3
3. Elytra with distinct post-humeral stripes
(as well as subapical-sutural spot(s)) pale;
prothorax less narrowed basally (base/apex
e. 1.10 or more) (p. 188) (stignnila)
- Post-humeral stripes indistinct or absent
(subapical-sutural spot(s) distinct); pro-
thorax more narrowed basally (base/apex
1.05 or less) 4
4. Brown, with subapical-sutural spots small and
separated; prothorax narrowed (width length
1.22 and 1.23); elytra more narrowed toward
base (p. 188) —_ tvallacei
- Black, with subapical-sutural spots united in
conspicuous square plagia; prothorax wider
(width length 1.35); elytra less narrowed
toward base ( Eig. 115) (p. 189) phigifer
5. Each elytron with transverse-oval pale spot
before middle but without subapical spot
(Eig. 114) (p. 189) ocellatus
188 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
- Elytra with aiitt'iior and posterior pale
spots 6
6. Elytra with an almost strictly transverse,
rej^ular, c. entire pale fascia before middle
( as well as an incomplete siibapical fascia ) ;
elytral margins bicolored; femora bicolored;
surface in part sericeous; length 5.5-6.0 mm
(p. 189) tratisversus
- Anterior elytral marks more oblicjue and/or
more irregular and/or more interrupted;
elytral margin usually entirely translucent-
testaceous; femora not bicolored; surface not
or not so strongly sericeous, more shining;
size usually smaller 7
7. Anterior elytral marks usually more oblique;
femora pale ( Moluccas and western antl
central New Guinea) (p. 190) onuitii.s
- Anterior elytral marks more nearly trans-
verse; femora darker (central and eastern
New Guinea) 8
8. Prothorax wider (width/length 1.38 and
1.41); markings wide (p. 190) fuscipcs
- Prothorax narrower (width/length 1.27);
markings n;irrower (Fig. 117) (see also
Dc'scii)>tiou) (p. 191) uaii
Anomofarus gressitti n. sp.
Description. With cliaractcrs of geiui.s;
form as in Figure 113; .slender; lilack, not
marked, but margins of prothorax and espe-
eially of elytra more or less pale translucent;
surface dull, closely punctulate and micro-
reticulate, elytra subalutaceous; legs testa-
ceous, antennae and mouthparts brownish
testaceous. Head 0.80 width prothorax.
Frothorax: width length 1.15; base, apex
1.00. Elytra: width elytra prothorax 1.73;
striae lightly iinpressed, faintly punctulate;
intervals c. flat, 6th, 7th, and (Sth with
inner edges finely carinate tor increasing
distances from base. Measiireitient.s: length
c. 5.0; width 1.9 mm.
Type. Holotype 9 (Bishop Mus.), from
Maprik, N-E. N. G., 160 m, Oct. 14, 1957
(Gressitt); the type is uniciue.
Notes. The slender form and dull bkuk
color, without markings, should nuikc this
species easy to recogni/e.
(Anomofarus stigmula (Chaudoir))
Ghaudoir 18.52, Bull. Soc. Nat. Moscow 25. Part 1,
p. 57 {Ci/min(Jis) .
Andrewes 19.30, C^at. Indian insects 18, (^arabidac,
p. 28.
Csiki 1932, Goleop. Gat., Garabidae, Ilarpalinae 7,
p. 1493 (see for synonymy and additional refer-
ences ) .
Louwerens 1953, Verhandlungen Naturforschcnden
Gesellschaft Basel 64, p. 316.
Jedlicka 1963, Ent. Abhandlungen 28, p. 451.
Description. With characters of genus;
form, including elytra, elongate; brown,
elytra each with longitudinal humeral mark
( outside 4th stria ) and variable apical mark
pale; appendages including femora testa-
ceous; rather shining, but surface finely
microreticulate and sparsely inconspicu-
ously punctulate. Head 0.84 and 0.86 width
prothorax. Prothorax: width length 1.24
and 1.29; base /apex 1.14 and 1.11; margins
moderate, with basal angles well defined, c.
right or obtuse (not acute). Elytra long;
width elytra prothorax 1.65 and 1.69; striae
moderately impressed, not distinctly punc-
tulate. Mea.suremcnts: length c. 4.7-5.3;
width c. 1.9-2.1 mm.
Type. From Simla(h), northern India;
now in Oberthiir Coll., Paris Mus. (not
seen ) .
Occurrence in New Guinea. Doubtful;
see following Notes.
Measured specimens. A i from Coim-
batore. South India, and 9 from Lawa,
Malita, Davao Prov., Mindanao, Philippine
Islands (both specimens M.C'.Z. ).
Notes. This species is now recorded
over a very wide area, from SE. Asia
including Ceylon and Japan to Timor and
New Caledonia, but apiiarentb not .\us-
tralia. New Cluinea is included in the
species' range by Andrewes and Csiki, but
I have found no detailed record of its oc-
currence there. I suspect its supposed oc-
cuiicnce is erroneous, b;ised on the old
specimens in the British Museum described
below ;is ualhuci. I h;i\(- not seen true
sliii,ninla Irom New (iuiiiea.
Anomofarus wallacei n. sp.
Dcscripliou. With c h;ir;icteis ol genus;
lonn shorter than usual in genus; brownish
pieeous, el\'tr;i e;uli willi laint p;iler AVCd
behind Immeius (corresponding to post-
The Carabid Beetles of New Guinea
Darlington
189
humeral spot of stigmula), a small testa-
ceous spot on intervals 2 and 3 just before
apex, and lateral margin narrowly brownish
testaceous; appendages brownish testa-
ceous; surface (in part) moderately shining,
with reticulate microsculpture lightly im-
pressed and punctulation rather sparse.
Head 0.84 and 0.89 width prothorax. Pro-
thorax: width/length 1.23 and 1.22; base/
apex 1.00 and 1.05. Elytra shorter and more
narrowed basally than usual in genus; width
elytra/prothorax 1.69 and 1.74; striae mod-
erately impressed, not distinctly punctulate.
Inner wings apparently fully developed in
spite of narrowing of humeri. Measure-
ments: length 4.3-4.9; width 1.8-2.0 mm.
Types. Holotype 5 (British Mus.) and
1 9 paratype (M.C.Z., Type No. 31,492)
both from Dory, West N. G. (presumably
collected by Wallace); c^ unknown.
Notes. This species is very close to
stigmuhi (above) but has elytra shorter,
more narrowed at base, and with markings
reduced, and the prothorax coaptively nar-
rowed at base, as the ratio base apex shows.
It may prove to be a geographic subspecies
of stigmida. It may prove not to be from
New Guinea (because the "Dory" locality
is always dubious), but it seems not to be
known anywhere else.
Anomofarus plagifer n. sp.
Description. With characters of genus;
form as in Figure 115; brownish black,
elytra with conspicuous, common, square
spot just before apex pale and lateral
margins slightly pale translucent; append-
ages testaceous; surface rather shining but
lightly microreticulate and faintly sparsely
punctulate. Head 0.88 width prothorax.
Prothorax strongly cordate (more so than
in stigmida); width/length 1.35; base/apex
1.02; posterior angles abruptly right-acute.
Elytra of moderate length, slightly nar-
rowed toward base ( less than in walktcei ) ;
width elytra prothorax 1.57; striae well im-
pressed, not distinctly punctulate. Measure-
ments: length c. 4.9; width c. 2.0 mm.
Type. Flolotype S (Bishop Mus.) from
Port Moresby, Papua, May 20, 1956
(Gressitt), taken in light trap; the type is
unique.
Notes. As compared with stigmida,
plagifer is slightly broader, with more
cordate prothorax, and is darker in color,
without basal but with more conspicuous
subapical elytral marks.
Anomofarus ocellafus n. sp.
Description. With characters of genus;
form as in Figure 114; black (bluish black
in some lights), each elytron with trans-
verse-oval pale spot before middle between
striae 1 and 7; appendages brownish testa-
ceous, femora darker brown; shining, reticu-
late microsculpture light ( faint on part of
pronotal disc) and punctulation very fine,
faint, sparse. Head 0.85 width prothorax.
Prothorax cordate; width/length 1.31; base/
apex 0.96; side margins narrower than
usual. Elytra: width elytra prothorax 1.59;
striae moderately impressed. Measure-
ments: length 4.4; width 1.8 mm.
Type. Holotype 9 (Louwerens Coll.,
eventually to Leiden Mus.) from Sorong
"Kpg. Roefci," West N. G., July 8-Aug. 14,
1948 (M. A. Lieftinck); the type is unique.
Notes. The small size, relatively shining
surface, and unique markings distinguish
this species.
Anomofarus fransversus n. sp.
Description. With characters of genus;
form as in Figiue 116; large, with wide-
cordate prothorax; dull aeneous black,
elytra more alutaceous, with c. entire cross-
band before middle and incomplete trans-
verse mark before apex pale; elytral margins
bicolored, pale at transverse fascia, dusky
elsewhere; femora bicolored, dark with
pale apices; appendages otherwise brown-
ish testaceous, antennae slightly darker
distally; surface closely microreticulate,
sparsely punctulate. Head 0.72 and 0.76
width prothorax. Prothorax: width length
1.46 and 1.43; base apex 1.08 and 1.03;
posterior angles abruptly acute. Elytra:
190 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
width elytra/prothorax 1.46 and 1.44; striae
fine, lightly impressed, not punctulate; in-
tervals almost flat. Measurements: length
c. 5.5-6.0; width 2.1-2.3 mm.
Types. Holotype S (M.C.Z., Type No.
31,493) and 1 9 paratype both from Do-
bodura, Papua, Mar.-July 1944 (Darling-
ton); 3 paratypes, Popondetta, Papua, 60
m, Aug. 30-31, Sept. 1-4, 1963 (Sedlacek),
light trap.
Notes. The large size, dull partly aluta-
eeous surface, and color pattern, especially
the evenly transverse anterior elytral fascia,
characterize this species.
Anomofarus ornafus Louwerens
Louwerens 1956, Treubia 23, p. 237.
Description. With characters of genus;
form c. as of following species (fuscipcs);
piceous black, elytra each with broad pale
mark before middle, this mark being ir-
regularly transverse-oblique with outer-an-
terior corner extended toward and some-
times reaching humerus, and elytra also
with subapical-sutural spot pale and lateral
margins entirely pale; legs entirely testa-
ceous; antennae brownish testaceous; sur-
face shining, reticulate microsculpture light
on head, faint on disc of pronotum, more
distinct on elytra. Head 0.cS6, 0.(S5, and
0.83 width prothorax. ProtJiora.x cordate;
width length 1.43, 1.36, and 1.33; base/apex
1.08, 1.07, and 1.08; posterior angles
abruptly right or acute. Elytra: width
elytra prothorax 1.72, 1.77, and 1.77;
striae moderately impressed, not punctu-
late. Measurements: length 5.1-5.8; width
2.2-2.4 mm.
Types. From Obi Is. (Laiwui, 0-200 m.
Sept.-Oct. 1953, A. M. R. Wegner); holo-
type in Leiden Mus. (not seen).
Occurrence in Neic Guinea. West N. {,.:
8, Cyclops Mts., Sabron, Camp 2, 2000 ft.
(610 m), July 1936 (Cheesman); 4, vie.
Ilollandia (various collectors); 1, Kebar
Vy. W. of Manokwari, 550 m, Jan. 4-31,
1962 (S. & L. Quale, Bishop Mus.), in light
trap; 1, Sansapor, Aug. 1944 (Darlington).
Measured .specimens. A pair (69) from
Cyclops Mts. and a 9 paratype from Obi
Is.; figures given in this order.
Notes. See under the following species
( fuscipes ) .
Anomofarus fuscipes n. sp.
Description. With characters of genus;
form average; black ( slightly brownish or
aeneous), elytra each with slightly oblique
transverse fascia just before middle and
subapical-sutural spot pale and lateral
margins entirely pale brownish translucent;
femora brown (not distinctly bicolored),
tibiae and tarsi paler; antennae brown;
upper surface rather shining, reticulate
microsculpture light especially on disc of
pronotum. Head 0.76 and 0.77 width pro-
thorax. Prothorax cordate; width length
1.38 and 1.41; base apex 1.17 and 1.09;
basal angles abrupth' right or acute.
Elytra: width elytra prothorax 1.66 and
1.59; striae moderately impressed, not punc-
tulate. Measurements: length 4.1-5.5;
width 1.9-2.3 mm.
Types. Holotype $ (M.C.Z., Type No.
31,494) from Dobodura, Papua, Mar.-July
1944 (Darlington); and paratypes as fol-
lows. Papua: 1, Karema, Brown R., Mar.
8-11, 1955 (E. O. Wilson, M.C.Z.), taken
in lowland rain forest; 2, Ml. Riu, Sudest
Is., 250-350 m, "No. 10," Sept. 3, 5, 1956
(L. J. Brass, A.M.N.H.); 1, Abaleti, Rossel
Is., 0-50 m, "No. 12," Oct. 9, 1956 (Brass,
A.M.N.H.). N-E. N. G.: 1, vie. Nadzab,
Julv 1944 (Darlington); 1, same localitv,
May 20-22, 1955 (E. O. \Ailson, M.C.Z.'),
in dry e\'ergr{>en forest; 1, Eriina, Astrolabe
Bav, 1896 (Biro); 1, Stephansort, Astrolabe
Ba'v, 1898 (Bin')). West N. G.: 4, Ilol-
landia, Nov. 21, 1944; Ma\ 1945; May 4,
1947 (Iloogstraal, M.C.Z. ); 1, same localitv,
May 1945 ( Malkin, U.S.N.M.).
Measured si)ecinu'ns. The • holot\pe and
1 9 parat\pe from Ilollandia.
Notes. This species is very close to the
[M'eeeding (ornatus), but is distinguished
by brown lather than testaceous femoia
and 1)\ more transverse anterior eKtral
The Carabid Beetles of New Guinea • Darlinaton 191
fascia which approaches the humeri less
closely, although this mark varies somewhat
in both species. A. fuscipcs occupies ap-
proximately the eastern half of New Guinea,
oniaftis the western half and the Moluccas,
but their ranges are not strictly allopatric,
since both occur in the vicinity of Hol-
landia.
Anomotorus wau n. sp.
Description. With characters of genus;
form as in Figure 117; black, elytra each
with oblique fascia before middle and with
common subapical-sutural spot testaceous
and margins brownish translucent; append-
ages brownish testaceous with femora
darker; shining, reticulate microsculpture
and fine sparse punctulation present but
light especially on pronotum. Head 0.84
width prothorax. ProtJiorax narrow-cordate,
with relatively narrow margins; width
length 1.27; base apex 1.05; sides more
oblique posteriorly (less broadly rounded)
than in fmcipes, with posterior angles
abruptly e. right. Elytra: width elytra
prothorax 1.67; striae well impressed, not
punctulate. Measurements: length 5.1;
width 2.0 mm.
Type. Holotype 9 (Bishop Mus.) from
Wau, Morobe Dist., N-E. N. G., 1200 m,
July 5, 1961 (Sedlaceks), taken in light
trap; the type is unique.
Notes. Although this may be only a form
of the preceding species (fiiscipes), it has
a relatively narrow prothorax and reduced
elytral fasciae and will probably prove to
be worth distinguishing. More material
from more localities is needed to show
whether it is a species or a geographic
subspecies.
Tribe PENTAGONICINI
Fentugonicidac Auct. including Jeannel 1949,
Coleop. Carabiques de la Region Malgache,
Part 3, p. 767.
Jeannel 1942, Faune de Fianee, Coleop. Carabiques,
Part 2, p. 1017, footnote.
Penta^onicinae Basilewsky 1953, Exploration Pare
National TUpemba, Fasc. 10, p. 183.
Scopodmi Csiki 1932, Coleop. Cat., Carabidae,
Harpalinae 7, p. 1500 (see for synonymy and
additional references).
Jedlicka 1963, Ent. Abhandlungen 28, p. 505.
The beetles of this tribe resemble Lebiini
but (according to Jeannel 1949) are not
related to them. Pentagonicini can usually
be recognized at a glance by form, and
the tribe is defined by technical characters
including obliteration of the suture that,
in most Carabidae, separates the mentum
from the base of the head posteriorly.
However, this suture is still indicated in
Parascopodes.
This tribe consists of four genera. One,
Pentagonica., occurs in all the warm regions
of the world. One, Parascopodes (de-
scribed below), consists of a single species
that occurs in both eastern New Guinea
and northeastern Australia. One, Actenonyx
(1 species), is confined to New Zealand.
And the fourth genus, Scopodes, is best
represented in Australia and extends to
New Zealand and to mountains on New
Guinea and on Java. Ecologically, Pen-
tagonica alone of these 4 genera is pri-
marily arboreal, occurring especially in
masses of vines and other vegetation near
the ground, although some species are
found among dead leaves on the ground.
Parascopodes cyaneus occurs, in my limited
experience, in grass or on the ground under
grass (I am not sure which). The New
Zealand genus (Actenonyx) is probably
ground-living. And Scopodes is ground-
living but some species occur on logs or
tree trunks.
The distribution of genera of Penta-
gonicini suggests two possible geographic
histories. The tribe may once have been
better represented in other parts of the
world and may have withdrawn (or may
be withdrawing) into the Australian Region.
Or the tribe may have originated in Aus-
tralia and diversified there, and Pentagonica
may have spread from there over the rest
of the world, its spread perhaps facilitated
by its invasion of arboreal habitats in
which flight and dispersal may have been
192 BiiUctin Miiscinu of Comparative Zoology, Vol. 137, No. 1
favored. The very wide distributions of
some species of Pento^onica show that the
insects do disperse readily. Of course, there
is a third possil^ihty, that the geographic
liistory of the tribe has been more complex
than can be guessed from present distribu-
tions of genera and cannot now be deci-
phered at all. Nevertheless, the history of
the tribe is worth guessing about. Its
distribution may become more significant
if other Carabidae or other animals are
found to have similar geographic patterns.
There is, for example, a suggestive general
similarity between the distributions of this
tribe of carabid beetles and of the parrots,
which are most diverse in the Australian
Region with one of the several Australian
subfamilies spread over the warmer parts
of the world (Darlington, Zooiieoiiraplnj,
Wiley, 1957, pp. 271-272, ;30()-;301, fig. .34).
Key to Genera of Pentagoxicim of
New Guinea
1. Form Lt'/;;'a-like, with eyes only iioniuilly
prominent (Fig. 118) (p. 192) _.._ Pcntd^onica
— Fomi more eompaet, with eyes larger and
more abruptly pronnnent (Figs. 120, 121) _ 2
2. Ligula normal, not miieh swollen, mueli
shorter than paraglossae; i front tarsi 2-
seriately squanmlose; elytral striae and in-
tervals regular, without eonspieuous foveae
(p. 19.5) Parascopodes
- Ligula swollen, club-like, as long as or longer
than paraglossae; i front tarsi with soles of
densely packed slender squamae; elytral
striae and intervals usually more or less ir-
regular, .^rd intervals usually with eon-
sj^iiciinns lovcae (p. 197) _ Scopixlcs
Genus PENTAGONICA Schmidt-Goebel
Schmidt-( ioclx'l I S4(), I'auniila (]olco|). Hiiiiianiae,
p. 47.
Csiki 19.32, Clolcop. Gat., ("arahidae, llarpaiiiiac 7,
\i. 1.500 ( see lor additional r(i<icnees, s\iion\niy,
and list of species ).
Di(iu.n()si.s. Immediately recogni/able b\-
lorm ( I'ig. 11<S) and tribal characters.
l)c.scrij)liou. None reciuired here.
Tijjx' species. Peiitagonica ntficollis
Schmidt-Cioebel ( below ) .
Generic distribution. All warm regions
ol the world.
Notes. The members of this genus are
winged, active, and apparently diurnal.
They usually live in dense vegetation within
a few feet of the ground, or sometimes in
leaf litter on the ground. Specific char-
acters in the genus arv few, principally
slight differences of form, microsculpture,
and color pattern. There is some individual
variation, including apparent dimorphism
of color in some cases. And understanding
of the species is made more difficult by the
very wide distributions of some of them.
Because the prothorax has no distinct an-
terior angles, the ratio base apex is omitted
in the following specific descriptions.
Key to Species of Pentagonica ok New Guinea
1. Prothorax strongly pedunculate; ( bicolored,
dark with prothorax red; elytral striae punc-
tulate; size small, length .3-.3.8 mm) (p.
192) pallipcs
- Prothorax less strongly pedunculate; size usu-
ally larger 2
2. Pronotum with lateral margins usually con-
nected posteriorly by a weak transverse
ridge that is not ((uite basal; reticulate
microsculpture of elytra often somewhat
transverse; (color above unitormK d;irk or
with prothorax red; length c. 3.5-4.5 mm)
(p. 193) l^himla
- Proncjtum with lateral margins connected
posteriorly b>' a line ridge that reaches
extreme base; elytral microreticulation usu-
ally c. isodiametric; size larger .3
3. Prothorax relatively narrower ( width length
1.57 and 1.65); (color dark, with lateral
margins of prothorax and elytra strikingh'
pale) (p. 194) crichsoui
- Prothorax relatively wider (width length
1.72-1.79) [ 4
4. Bicolored, d;uk with red prothorax (p.
194) nificollis
- Not bicolored, entirely dark 5
5. Flytral striae clearly indicated (hut scarcely
impressed); antennae rilativeb dark ;uid
femora relatixcK pale (p. 194) papiia
- FKtral striae \irtu:dl\ obsolete; antennae
testaceous, Icmoia rclatixcK (!;nk (p. 19.5)
_ cstriaia
Pentagonica pallipes (Nietner)
Nielncr 18.5(i, 1. Asiatic Soc. Bengal 25, p. .525
(Klliolia).
(.'siki 1932, (^oK'op. i'.ni.. Gar:ibida<', Ihupalinae 7,
p. 1.502 (see for sNiionvinx and additional refer-
ences ).
The Carabid Beetles of New Guinea • Darlington
193
Jedlicka 1963, Ent. Abliandlungen 28, pp. 505,
507.
Description ( for recognition only ) . Head
and elytra dark, prothorax red. Head 0.91
and 0.93 width prothorax. Prothorax pedun-
culate; width length 1.71 and 1.59. Elytra:
width elytra/ prothorax 1.62 and 1.79; striae
punctate; reticulate microsculpture slightly
irregular but scarcely transverse. Measure-
ments: length 3.0-3.8; width c. 1.5-1.9 mm.
Type. From Ceylon; should be in Stettin
Mus. (not seen).
Occurrence in Neiv Guinea. Nineteen
specimens from all 3 political divisions of
New Guinea; most from low altitudes (in-
cluding Oro Bay near Dobodura) but 2 at
1200 m at Wau. One specimen ( from Wau )
is labeled as taken at light.
Measured specimens. A 6 from Hollandia
and 9 from Port Moresby.
Notes. The known range of this rela-
tively distinct species is from Ceylon, the
Malay Pen., etc., to the Philippines, New
Guinea, New Britain, and mid-peninsular
Cape York, Australia (collected hv me
in 1958).
Pentagonica blanda Andrewes
Andrewes 1929, Tijdschrift voor Ent. 72, pp. 315,
339.
?htzocn.'iis Jedlicka 1934, Sbornik Ent. Mus. Prague
12, p. 123.
?})hili]>pmeims Jedlicka 19.34, Sbornik Ent. Mus.
Prague 12, p. 124.
?hottclwri Jedlicka 1935, Acta Soc. Ent. Czecho-
slovakia 32, p. 140.
?eurijodes Andrewes 1938, Ann. Mag. Nat. Hist.
(l"l) l,p. 207.
?quadratipennis Louwerens 1956, Treubia 23, p.
236.
Description ( for recognition only ) . Form
(Fig. 118) broad; color above eitlier en-
tirely dark or dark with red or reddish
prothorax; antennae brown or testaceous;
legs pale, often with darker femora. Head
0.77 and 0.74 width prothorax. Prothorax:
width/length 1.83 and 1.80; lateral margins
usually connected by a poorly defined
(sometimes vague) transverse prebasal
ridge. Elytra: width elytra prothorax 1.51
and 1.54; sutural angles usually blunted or
narrowly rounded but sometimes denticu-
late; striae lightly impressed, usually finely
punctulate ( but variable ) ; reticulate micro-
sculpture often ± transverse. Measure-
ments: length c. 3.5-4.5; width c. 1.6-1.8
mm.
Types. Of blanda, from Sumatra, in
British Mus. (seen); of Jedlicka's species,
from the Philippines, types of luzoensis
and philippinensis in Jedlicka Coll. (not
seen), of bottcheri, in British Mus. (seen);
of euryodes, from Java, in British Mus.
(seen); of quadratipennis, from Halmahera,
Molueeas, in Leiden Mus. ( not seen ) .
Occurrence in Neic Guinea. Very com-
mon throughout New Guinea: c. 130
specimens (about half from Dobodura),
most at low altitudes but a few up to 2500
m (at 1200 m at Wau).
Measured specimens. A pair (69) from
Dobodura.
Notes. I am not ready to synonymize the
names listed above under blanda, but I
suggest that they may all prove to apply
to a single variable species or to members
of a group of very closely interrelated
species that ranges from SE. Asia across
the Malay Archipelago to NE. Australia.
This species or species group varies strik-
ingly in color pattern (individuals uni-
colored or bicolored) and to some extent
in proportions, size, degree of paleness of
margins and appendages, and distinctness
of punctures of elytral striae. Most in-
dividuals from New Guinea are entirely
dark above, but 3 from Finschhafen, N-E.
N. C, and 8 from Hollandia, West N.
C, have the prothorax red. These super-
ficially resemble ruficollis (second follow-
ing species ) , but have different prothoracic
bases, usually rounded sutural angles, and ±
transverse elytral microsculpture. Except
for the red prothorax, these individuals do
not seem to differ from specimens with
dark prothorax taken at the same localities.
At these localities the species is apparently
dimorphic in color of prothorax. However,
at some other localities intermediates oc-
194 Bulletin Musctmi of Comparative Zoology, Vol. 137, No. 1
cur: 4 examples that I haxe from Sansapor
( Vogelkop), West N. G., have the prothorax
paler than head and elytra but reddish
brown rather than clear red. Obviously
this species or group ot species requires
further study, of material from outside as
well as inside New Guinea, before its varia-
tion can be understood.
Most of the many specimens that I col-
lectt^d in New Guinea were taken by day,
by sweeping low vegetation. However, a
few individuals, including both color forms
at Finschhafen, are from light-trap material
and evidently flew to light at night.
I collected numerous dark (not bicolored)
specimens apparently of this species at
several localities in North Queensland in
1957-195(S. The species seems not to have
received a name in Australia.
Penfagonica erichsoni Schmidt-Goebel
Schmidt-Coel^el 1846, Faunula CoKop. Biniianiat',
p. 48.
Csiki 1932, Coleop. Cat., Carabidat', llaipalinae 7,
p. 1501 (see for synonymy and additional refer-
ences ) .
Jedlieka 1963, Knt. Ahliandlnntjen 28, pp. 506.
.511.
Description ( for recognition only), h^orm
of large Fcntuiionica with rather narrow
prothorax; dull black, reflexed margins of
prothorax and elytra very pale or translu-
cent, legs testaceous, antennae brown.
Head 0.85 and 0.84 width protliorax. Pro-
thorax: width length 1.57 and 1.65; margins
posteriorly connected b\ a fine curved
ridge across extreme base. Elytra: width
elytra/ prothorax 1.82 and 1.78; sutural
angles usually denticulate; striae sHghtl\
impressed, vaguely or not distinctl\ punc-
tate; elytral microreticulation r. isodiamet-
ric. Measurements: length e. 5.0-5.5;
width e. 2.0-2.2 mm.
Type. l<'rom liurma; in Prague Mus. (not
seen ) .
Occurrence in Netc Guinea. Tapua: 1,
Dobodura, Mar.-lul\ 1944 (Darlington);
1, Woodlark Is. (Murn;i), Kulumadau Hill,
Apr. 20-30. 19.57 (W. W . Brandt, l^isliop
Mus.). N-E. N. G.: 1, Simbang, Huon
Gulf, 1898 (Biro); 1, Wau, 1200 m, Sept.
15-30, 1962 (Sedlacek); 1, Wum, Upper
Jimmi Vy., 840 m, July 16, 1955 (Gressitt).
Measured specimens. Two $ 9 , from
Dobodura and Wum.
Notes. P. erichsoni ranges from Ceylon
and SE. Asia to New Guinea and mid-
peninsular Cape York, Australia ( 1 speci-
men. Rocky R., 1958, taken by myself).
Penfagonica ruficollis Schmidt-Goebel
Sehmidt-Coebel 1846, Fannnla Coleop. Birmaniae,
p. 48.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1503.
Jedlieka 1963, Ent. Abhandlunsen 28, p. 509.
Description (for recognition only ). Form
of large Pentagonica; black, prothorax red,
appendages usually black or brown. Head
0.81 and 0.77 width prothorax. Prothorax:
width length 1.79 and 1.78. Elytra: width
elytra prothorax 1.59 and 1.61; sutural
angles usually denticulate; striae very light
or c. absent; microsculpture c. isodiametric.
Measurements: length c. 5.0-5.7; width e.
2.0-2.3 mm.
Type. From Burma, should be in l^rague
Mus. (not seen).
Occurrence in New Guinea. Twt'nt\-six
specimens from Papua, N-E. N. (i.. anil
West N. (i., at low altitudes and up to
1950 m.
M easurrd sjx'cimens. A i")air ( ci 9 ) b-om
Dobodura, Papua.
Notes. The known range of ruficollis is
Irom SE. Asia to .Australia.
Penfagonica papua n. sp.
Description. Form (Fig. 119) of large,
rather slender Pentaiionica: entireK' black
or piecous except suture sonictiuies reddish,
margins sometimes reddish hut not con-
trastinglx pale; h-mora brown, tibiae and
tarsi testaceous; antennae brown with 1st
segments darker biowii; rt'tieulate micro-
seulpture isodiametric on head and pro-
notum. nioic incgnlai- or slightK tiansvcrse
on eKtra. Head 0.78 and 0.78 width pro-
tliorax; labium transxcrse. 6-setose ante-
The Carabid Beetles of New Guinea • Darlington
195
riorl) , the 4 inner setae mueh smaller than
outer ones; mentum without tooth; ligula
not much swollen, 2-setose; paraglossae
slightly shorter than ligula and apparently
attached to it. Prothorax: width length
1.73 and 1.72; margins posteriorly connected
1)\ basal loop. Elytra: width elytra pro-
thorax 1.63 and 1.63; sutural angles usually
narrowly rounded, not denticulate; striae
very finely and lightly indicated, scarcely
impressed, irregularly punctulate; 3rd in-
tervals apparently usually with 3 minute,
well spaced punctures but latter small, dif-
ficult to find, perhaps sometimes absent.
Secondary sexual characters: £ front tarsi
scarcely dilated but with slender squamae
in 2 slightly irregular rows; c^ with 1, 9
2 setae each side last ventral segment.
Measurements: length 4.8-5.7; width 1.8-
2.2 mm.
Types. Holotype $ (M.C.Z., Type No.
31,495) and 10 paratypes all from Dobo-
dura, Papua, Mar.-July 1944 (Darlington).
Measured s})ecimei}s. The 6 holotype and
1 9 paratype.
Notes. The preceding Description gives
all the characters that seem to me worth
mentioning in this genus, in which the
species are so similar to each other. The
particular characters that separate this
species from others in New Guinea are
indicated in the Key to Species. The present
new species is larger than blandii, relatively
narrower, with different pronotal base and
slightly different elytral microsculpture, as
well as more lightly impressed elytral striae.
As compared with erichsoni, papua has a
wider prothorax, shallower elytral striae,
and darker prothoracic and elytral margins.
The closest relative of papua may be
ruficollis ( above ) , but the latter is bicolored
and always (in the specimens before me)
has the sutural angles subdenticulate or at
least angulate, while papua is dark, with
sutural angles usually blunted, rarely sub-
denticulate. The difference in sutural
angles is not absolute but is enough to sug-
gest that the color difference is specific.
Penfagonica esfriafa n. sp.
Description. Form of large Penta^onica;
black or piceous, legs dark, antennae in-
cluding basal segments pale; entire upper
surface with c. isodiametric microsculpture,
at most slightly transverse on elytra. Head
0.83 and 0.82 width prothorax; details of
mouthparts as in papua. Prothorax: width,
length 1.76 and 1.78; margins posteriorly
connected by a basal loop. Elytra: width
elytra prothorax 1.71 and 1.74; sutural
angles usually finely denticulate; striae ef-
faced or at most faintly indicated; dorsal
punctures, if present, minute, almost un-
detectable. Secondary sexual characters as
in preceding species (papua). Measure-
ments: length 4.9-6.2; width 2.1-2.6 mm.
Types. Holotype c^ (Bishop Mus.) and
1 9 paratype from Eliptamin Vv., N-E.
N. G., 1200-1350 m, June 19-30, 1959
(W. W. Brandt); and additional paratypes
as follows. N-E. N. G.: 1, Eliptamin Vy.,
1350-1665 m, June 23-30, 1959 {\Y. W.
Brandt, Bishop Mus.); 1, Wau, Morobe
Dist., 1200 m, Jan. 3-4, 1963 (Sedlacek);
1, "No. 14," Umi R., Markham Vy., 480 m,
Nov. 11, 1959 (L. J. Brass, A.M.N.H.).
West N. G.: 1, Hollandia, Dec. 15, 1944
(Hoogstraal, M.C.Z.); 1, Star Mts., Sibil
Vy., 1245 m, Oct. 18-Nov. 8, 1961 ( S. & L.
Quate, Bishop Mus.), taken in Malaise
trap; 3, Japen Is., Mt. Baduri, 1000 ft. (c.
300 m), Aug. 1938 (Cheesman). Some
paratypes in M.C.Z., Type No. 31,496.
Measured specimens. The 6 holotype and
1 9 paratype from Eliptamin Vy.
Notes. P. c.striata is the same size and
nearly the same form as papua but slightly
broader, with differently colored append-
ages, usually denticulate rather than blunted
sutural angles, and virtually no elytral
striae. The latter character, as well as
color, differentiat(\s estriafa from ruficollis.
PARASCOPODES n. gen.
Diagnosis. Form Scopodes-Mke. Head:
eyes enormous; labrum transverse-quadrate,
6-setose; mentum \\'ith basal suture in-
196 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
dicated, and with broad, short, more or less
emarginate tooth; hgula not swollen, 2-
setose; paraglossae much longer than ligula,
with apices narrowly rounded. Prothoia-x
weakly subpedunculate, angulate at sides,
2-setose each side. Elytra with entire, regu-
lar striae; 3rd intervals very inconspicuously
3-punctate. Secondary sexual characters: 6
front tarsi narrow, 2-seriately squamulose
below; i copulatory organs as figured
(Fig. 178); most other characters as in
Scopodes (below).
Description. See Diaiinosis (above) and
Description of the single species (below).
Type species. Scopodes cyaneus Sloane.
Generic distribution. As of the single
known species: eastern New Guinea and
North Queensland, Australia.
Notes. Although the Sro/>or/c.s-like form
and enormous eyes are striking specializa-
tions, the labrum, mentum, ligula, elytra,
and perhaps the S tarsi of Farascopodes
cyaneus seem generalized within the tribe
and suggest that the insect may be a
superficially specialized relict of a primitive
stock from which both Fentaiionica and
Scopodes may have evolved.
Farascopodes cyaneus (Sloane)
Scopodes cyancii.s Sloane 1907, Proc. Linncan Soc.
New South Wales 32, p. 380.
Description. With characters of genus;
form as in Figure 120; color above blue, more
or less purplish on elytra, below piceous;
appendages testaceous, mouthparts in part
browner; shining, reticulate microsculpture
virtually absent on head, light, irregular,
usually slightly transverse on pronotum and
elytra. Head 1.36, 1.32, 1.31 width pro-
thorax; 2 setae over each eye; Iront smooth
at middle, with a deep groo\'e each side.
Prothorax: width length 1.19, 1.16, 1.23;
margins narrow, each with 2 setae on small
projections; disc strongly convex; middle
line and trans\-erse impressions sharply im-
pressed, latter irreguhuK subpunctatc; sur-
face of disc smooth excei^t \\ illi laiiit liaiis-
verse strigulation toward sides. I'Jyhd:
width elytra prothorax 1.72, 1.72, 1.65;
striae entire, regular, coarsely punctate;
intervals regular, flat or slightly convex,
3rd with 3 very inconspicuous dorsal punc-
tures (see Notes, below). Inner icings
dimoiphic: short in 1 i from Dobodura,
full in all other specimens. Lower sui^ace
sparsely pubescent; base of abdomen on
each side ( under bases of femora ) with c.
6 parallel grooves that may form a stridu-
lating organ. Legs slender; 4th hind-tarsal
segments shallowly emarginate; 5th seg-
ments with accessory setae; claws simple.
Secondary sexual characters: i tarsi as de-
scribed under genus; 6 with 1, 9 2 setae
each side last ventral segment. Measure-
nients: length 3.7-4.0; width 1.4-1.6 mm.
Type. From Kuranda, North Queens-
land, Australia; should be in Sloane Coll.,
Canberra, but I was not able to find it
there in 1957.
Occurrence ifi New Guinea. Papua: 3,
Dobodura, Nhir.-July 1944 (Darlington); 1,
Misima Is. ("St. Aignan") (ex coll. F. C.
Morrell, British Mus.). Also 1, "New
Cuinea, Sayer" (Sharp Coll., British Mus.).
Measured .specimens. A pair ( c^ 9 ) from
Dobodma and a 9 from N. of Mareeba,
Australia (see below); figures listed in this
order.
Notes. Besides the specimens from New
Cuinea, I have 1 9 taken by myself N. of
Mareeba (about 20 miles SW. of the t>'pc
locality). North Queensland, Australia, Feb.
1958, in flooded grassland. 1 can lind no
significant difference between the New
Cuinean and Australian specimens, but oi
course I have not been able to compare 6
characters.
\u a footnote to the original description.
Sloane sa\s that he eould liiid onl\' 1
( fine ) dorsal puncture on the el\tra oi the
t\pe. However, under a good stereoscopic
inieioseope with good light, 1 lind what
appear to be .3 punctures neaiK e\('iil\
spaced along the length of each 3rd inter\al
in both New (iuinean and Australian in-
(li\ idnals.
The Carabid Beetles of Ne\\' Guinea
Darlington
197
The habitat of this species, in or under
grass at low altitudes, is different from that
of any Scopodes in New Guinea. If this
insect is a relict of an ancestral stock from
which Pcntap.onica and Sco))odcs have
evolved, as I ha\e suggested under the
genus, it may still be in an ancestral habitat
from which Pentag,onica may have invaded
low vegetation, and Scopodes, habitats on
the ground.
Genus SCOPODES Erichson
Krichson 1842, Archiv fiir Naturgeschichte 8,
Band 1, p. 123.
Sloane 1903, Proc. Linnean Soc. New South Wales
28, pp. 637-638 (key to Australian species).
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1504 (see for additional references, synonymy,
and list of species ) .
DiaiJ^nosis. Form (Fig. 121) character-
istic, with eyes very large and sides of
prothorax usually angulate; mentum not
toothed; ligula swollen, club-like, longer
than paraglossae, 2-setose; i front tarsi
with soles of numerous close-packed slender
squamae not in 2 series. For additional
diagnostic characters see under the tribe
and Key to Genera of Pcntaiionicini.
Description. None required here, but
note wings full or reduced or dimorphic in
different Australian species, full in all New
Guinean species except altus, in which
reduced.
Ty})e species. Scopodes hoops Erichson,
of southern Australia.
Generic distribution. Australia, New
Zealand, New Guinea, Java; and 1 species
(not seen by me) described from New
Caledonia.
Notes. This genus divides into 2 groups.
The typical group includes all the Austra-
lian and New Zealand species, 1 species
(altus) at very high altitudes on the Snow
Mts. in New Guinea, and 1 ( irregidaris
Andrewes) on mountains in Java. The
other New Guinean members of the genus
form a distinct, endemic group of relatively
compact, often brightly colored species oc-
curring mostly at moderate altitudes in the
mountains. These species can be arranged
in what seems to be an evolutionary se-
quence involving change of form, loss of
elytral striae, simplification and reduction
of sculpture, and intensification of color,
but the lines of evolution have probably
been complex. In the last species of the
Key (adonis. Fig. 123), which is at the end
of the apparent evolutionary sequence, the
Scopodes form is much modified (the pro-
thorax being long and narrow and without
lateral angulations, and the elytra being
oval ) ; the elytral striae and sculpture have
almost disappeared, and the color is deep
purple.
These beetles live in the ground or in
logs. They are probably diurnal: most of
them are winged, but they apparently do
not come to light at night. Evelyn Ghees-
man, in The Land of the Red Bird ( London,
Herbert Joseph Ltd., no date, p. 134), says
apparently of Scopodes cheesmani (which
I describe below), "Other insects were
very tiny violet beetles, which appeared
from inside the log, out of one of its many
passage ways, and ran about on the sur-
face." This was between 3400 and 4500 ft.
(c. 1380 m) altitude in the rain-forested
Cyclops Mts.
In the following descriptions, the width
of prothorax includes the tubercles. The
frontal sulci are counted between the eyes
just behind the anterior supraocular setae.
The sculpture of the front of the head, espe-
cially of the clypeus, is variable, heavily
impressed or almost obsolete in different
individuals of single species from single
localities, and I have therefore not de-
scribed it in detail in most cases.
Key to Species of Scopodes of New Guinea
1. Prothorax with 2 setae each side, the pos-
terior on conspicuous dentiform processes;
(depressed; elytra with c. complete striation
indicated) ( p' 198) altufi
- Prothorax with only 1 seta each side, c. %
of prothoracic length from apex, without
posterior-lateral setae or processes - 2
198 Bulletin Museum of Comparative Zoology, Vol. 137. No. 1
2. Elytra with conspicuous irregular sericeous
pattern aiul extensively but irregularly
striate 3
- Elytra without conspicuous sericeous pat-
tern 4
3. Color bronze, elytral foveae conspicuously
bine, legs pale (p. 199) tafa
- Color dark bronze or greenish, eh tral foveae
not conspicuoush' contrasting, legs dark (p.
199) ..--' cJihithti
4. Color usually bronze, never primarily bright
blue or purple 5
- Color primarily blue or purple 7
5. Elytra with striae indicated at least anteri-
orly 6
- Elytra without striae (p. 201) sim))lcx
6. Extensively snbstriate (p. 200) icilsoiii
- Striae indicated only at base of elytra (p.
200) Inisdlis
7. Prothorax with lateral margins (p. 201 )
checsiiuini
- Prothorax without lateral margins (p. 201)
adoni.s
Scopodes alius n. sp.
Description. With characters of genus;
form (Fig. 121) relatively depressed, with
humeri narrowed; black, appendages dark;
rather shining, but complexly sculptured
as described below. Head 1.26 and 1.23
width prothorax; labrum long, strongly
rounded, with 4 decurved setae anteriorly,
the inner setae much shorter than the outer
(setae sometimes broken off); front with
c. 20 or more fine sulci, latter subparallel
at sides and base but connected and partU
transverse at middle ol head; front anteri-
orly and clypeus and labrum more finel\
and closely longitudinally sculptured in
S 6 , sculpturing reduced in 9 (see Notes).
Prothora.x: width length 1.24 and 1.27;
sides margined, each with 2 setae, at an-
gulation c. ':'. from apex and on conspicuous
triangular process b(4()re base; disc with
weak median longitudinal imiiression and
strong subbasal and subapical transverse
imprt\ssions; surface of disc covered vvith
rather line, complex, more or less anas-
tomosing sculptuic. Elytra: width elytra/'
prothorax 1.69 and — (elytra too spread to
measure in 9 ); striae indicated ])\' imdula-
tions of the surlace bul not sliarpK dclincd.
slightly irregular; intervals slightly convex,
3rd with 3 moderate foveae; elytral surface
with rather light irregular reticulate micro-
sculpture. Inner tt/ng.s- reduced to c. Vs
length of elytra. Measurements: length c.
3.3-3.5; width c. 1.3 mm.
Types. Holotype S (Leiden Mus. ) from
Scree Vy. Camp, Snow Mts., West N. G.,
3600 m, Sept. 19, 1938; 1 i paratype
(M.C.Z., Type No. 31.497) from same
locality, 3800 m, Sept. 1938; 1 9 paratype
(Leiden Mus.), from Lake Habbema, Snow
Mts., 3250-3300 m, late July-Aug. 1938 (all
specimens collected by Toxopeus ) .
Measured specimens. The 6 holotype and
9 paratype.
Notes. This species differs from all other
Scopodes known from New Guinea in its
depressed form and in possessing 2 pairs
of lateral prothoracic setae. In these ways it
resembles some Australian species including
the type of the genus, Scopodes hoops
Erichson, but (dttis is more shining than
Jjoops, with better defined pronotal sculp-
ture, and other minor differences of detail.
S. altus resembles also irre^idaris Andrewes
of Java but is black rather than aeneous,
with elytra less narrowed anteriorly and
more distinctly striate, as shown b\ com-
parison with a specimen ol irreu^ularis from
"G. Tengger, Java, l^rescher." Of the 3
specic^s just discussed, the Australian hoops
has dimorphic wings; the New Guinean
and Javan specie's, reduced ones; but they
all are probabK derived Irom a winged
ancestor that dispersed i)\ I light. The
distribution of this group ol Scopodes in
the Malax' Archipelago is comparable to
that of Meeyclothorax. which also occurs
primariK' in Australia but is represented at
verv high altitudes on the Snow Mts. ol
New Guinea and on the mountains ol Java
(Darlington, Hull. M.C.Z. 126. 1962, pp.
505-507). S(H' also Mieroferouia, [vige 18
ol my i^rescut pajier.
Whether the dillcrence in sculpture of
clypeus and labrum of the 6 S (Scree Vy.)
and the V (Lake llabbema) is sexual.
The Carabid Beetles ok New Guinea • DarUnglon
199
geographic, or individual cannot be decided
without more material.
Scopodes fafo n. sp.
Description. With characters of genus;
form compact, convex; aeneous, labrum
dark, dorsal and lateral foveae of elytra
blue, appendages pale, antennae browner
distally; head and pronotum shining, sculp-
tured as described (below), elytra irregu-
larlv sericeous with irregular reticulate
microsculpture. Head 1.19 and 1.20 width
prothorax; labrum rounded, 6-setose; front
with c. 8 longitudinal sulci abbreviated an-
teriorly and running into coarse rugosity
posteriorly; labrum more finely longitudi-
nally rugose. ProtJwrax: width length
1.26 and 1.27; sides margined, each with
seta-bearing puncture on triangular process
at angulation c. % from apex; disc with
coarse rugosity transverse posteriorly but
more confused anteriorly. Elytra subquad-
rate; width elytra prothorax 1.83 and 1.77;
several striae indicated on disc but striation
obsolete externally and apically; 3rd in-
terval conspicuously 3-foveate. Measure-
ments: length 3.5-4.0; width 1.4-1.7 mm.
Ti/pes. Holotype 9 (British Mus.) and
4 paratypes (2 in M.C.Z., Type No. 31,498)
all from Mt. Tafa, Papua, 8500 ft. (c.
2600 m), Feb. 1934 (Cheesman).
Additional material. One $ , Wau, Bull-
dog Rd., N-E. N. G., 2400 m, May 31, 1962
( Sedlaceks ) .
Measured specimens. The c^ holotype and
1 9 paratype.
Notes. See preceding Key for characters
distinguishing this from related species.
The specimen from Wau is slightly larger
than the types, with narrower humeri (but
still with large folded inner wings ) and less
distinct elytral striation. Additional ma-
terial may show it to be a distinguishable
form.
Scopodes chimbu n. sp.
Description. With characters of genus;
form compact; color dark, subaeneous or
greenish, elytral foveae not contrasting,
appendages dark except antennae paler
basally; head and pronotum sculptured but
with irregular reticulate microsculpture.
Head 1.15 and 1.22 width prothorax; labrum
rounded, 6-setose; front with c. 7 longitu-
dinal sometimes slightly sinuous sulci
shining, elytra irregularly sericeous and
running into coarse rugosity posteriorly and
sometimes anteriorly. Prothorax: width/
length 1.25 and 1.21; sides margined, each
with 1 seta-bearing puncture, on tubercle
at angulation c. % from apex; disc coarsely
rugose, the rugosity in general transverse
but somewhat confused especially anteri-
orly. Elytra subquadrate; width elytra/
prothorax — and 1.83 ( elytra of c5 too spread
to measure); several striae indicated on
disc, outer striae fainter or obsolete; 3 con-
spicuous foveae on each 3rd interval. Mea-
surements: length 3.5-4.4; width 1.4-1.7
mm.
Types. Holotype $ (M.C.Z., Type No.
31,499) and 2 (99) paratypes from
Chimbu Vy., Bismarck Rge., N-E. N. G.,
5000-7000 ft. (c. 1500-2135 m), Oct. 1944
( Darlington ) .
Additional material. N-E. N. G.: 2, Mt.
Mis(s)im, Morobe Dist. (1 specimen at
6400 ft. = 1950 m) (Stevens, M.C.Z.); 1,
Joangang, 500 m, Apr. 7-8, and 1, Tumnang,
1400-1600 m, Apr. 14-15, both on Mongi
Watershed, Huon Pen. ( 1955, E. O. Wilson,
M.C.Z.); 1, Saruwaged (Salawaket) Rge.,
upper Bunbok Vy.. 1800-2000 m. May 1955
(E. O. Wilson, M.C.Z.); 1, Sepalakembang,
Salawaket Rge., 1920 m, Sept. 12, 1956 (E.
J. Ford, Jr., Bishop Mus.); 1, Kepilam,
2420-2540 m, June 21, 1963 (Sedlacek); 1,
Finisterre Rge., Saidor, Matoko, Aug. 29-
Sept. 5, 1958 (W. W. Brandt, Bishop Mus.).
Measured specimens. The 6 holotype and
1 9 paratype.
Notes. This species varies somewhat from
locality to locality, but I do not have enough
material to distinguish geographic forms.
Except that my specimens were taken on
the ground in the more open part of the
200
Bulletin Miiscinu of Comparative Zoology, Vol. 137, No. 1
Chimbu Valley, I can say nothing about
their habits.
Scopodes wilsoni n. sp.
Description. With characters of genus;
compact, convex; color dark, subaeneous,
elytral foveae not contrasting, appendages
dark, antennae paler basally; shining, head
and pronotum coarsely sculptured, elytra
irregularly microreticulate but without con-
spicuous sericeous pattern. Head 1.25 and
1.24 width prothorax; labrum narro\\'ly
rounded, 6-setose; front with c. 8 longitu-
dinal sulci running into coarse rugosity pos-
teriorly and usually anteriorly. Frothorax:
width/length 1.23 and 1.18; sides margined,
each with seta-bearing puncture on tuber-
cle at angulation c. % from apex; disc
coarsely rugose, the rugosity transverse but
somewhat irregular and varying in depth,
and disc also variably punctulate. Elytra
quadrate; width elytra/prothorax 1.91 and —
(elytra too spread to measure in second
specimen); all or several striae indicated
for much of length; 3 conspicuous foveae on
or near each 3rd interval. Measurements:
length 3.4-4.1; width 1.4-1.7 mm.
Types. Holotype $ (M.C.Z., Type No.
31,500) from Nganduo to Yunzain, Mongi
Watershed, Huon Pen., N-E. N. G., 1000-
1500 m, Apr. 6, 1955 (E. O. Wilson); and
following paratvpes (all i 6 ) from N-E.
N. C: 1, Nad/ab, Markham Vy., July 13,
1944 (K. V. Krombein, U.S.N. M.), this
specimen further labeled "E. fork Ngafir
Cr. 100()-3()00 ft. nativc> trail"; 1, Mt.
Mis(s)im, Morobe l^i.st., 6400 Ft. (1950 m)
(Stevens, M.C.Z.); 1, Wau, 1300 m, Dec.
10, and 1, same locality, Nami Ck., 1750 m,
Aug. 12 (both 1961, Sedlaceks).
Addilional nuiterial. Fapua: 1, Owen
Stanley Rge., Goilala, Bome, 1950 m, Apr.
30-Mav 2, 1958 (W. \V. Brandt, Bishop
Mus.).' West IS. (;.: J, Battau Camp, 1150
m, Feb.-Mar., and 1, Sigi (lamp, 1500 in,
Feb. 26, both 1959 (Neth. Ind. -American
[Snow Mts.] Exp.; Toxopeus).
Measured specimens. The A holotype and
9 paratype from Nad/ab.
Notes. As in the case of chimbu (pre-
ceding), this species appears to vary geo-
graphically, but my material is too limited
to justify describing geographic forms.
Scopodes basalis n. sp.
Description. With characters of genus;
very compact, convex; dark green or aene-
ous, elytral foveae not contrasting, labrum
and legs dark, antennae yellowish brown;
shining, reticulate microsculpture faint or
absent e\en on elytra. Head 1.22 and 1.20
width prothorax; labrum narrowly rounded.
6-setose; front with c. 7 longitudinal sulci
sometimes abbreviated anteriorly and pos-
teriorly. Trothorax: width length 1.21 and
1.25; sides margined, each with seta-bearing
puncture on tubercle at angulation c. V:i
from apex; disc coarsely transversely rugose
(rugosity sometimes only lightK" impressed)
and rather sparsely fineh' punctulate.
Elytra quadrate; width elytra/prothorax
1.79 and 1.82; striae absent or \irtuall\- so
except deeph' but variably impressed at
base, sometimes only on basal declivit\-; 3
conspicuous punctiform foveae on positions
of 3rd interxals. Measurements: length c.
3.3; width c. 1.7 mm.
Types. Holotype A (M.C.Z., Type No.
31,501) from Joangang, Mongi Watershed,
Huon Pen., N-E. N. G., 500 m, Apr. 7-8,
1955 (E. (). \\'ilson); and following para-
types from N-E. N. (,.: 1, Saruwageil
(Salawaket) Rge., upper Bunbok \'v., 2300-
3200 m. May 29-31 (E. (). Wilson, M.C.Z.),
"mossy forest"; 1, Mt. Mis(s)im, Morobe
Dist. (Stevens, M.C.Z.); 1, TorricelH Mts..
Siaute, sea level, Nov. 9-17, 1958 (\\. W.
Brandt, Bishop Mus. ).
Measured sjx'cimens. The s holot\ jie and
9 paratyjie Irom Saruwaged Bge.
Notes. S. basalis diHers from iriisinii not
oiiK in reduction ol cKtral striation bnt
also in more pnnelilorm cKtral loxcae and
\irtnal absence ol icticnlatc inicrosculi')turc>
of eKtia. 'Ilic oeiui rcnce ol an indixidnal
near sea IcncI is uiinsnal in this genns in
New (iuinea.
The Carabid Beetles of New Guinea • Darlington
201
Scopodes simplex n. sp.
Description. With characters of genus;
compact, convex; aeneous black, elytra
sometimes faintly purplish, front anteriorly,
clypeus, and labrum ])right aeneous, ap-
pendages dark, base of antennae paler;
shining, elytra lightly irregularly microreticu-
late. Head 1.25 and 1.25 width prothorax;
front with 7 sulci running into coarse
rugosity posteriorly and sometimes anteri-
orly. Prothorax: width/length 1.19 and
1.22; sides margined, each with seta on
tubercle c. Mi from apex; disc lightly trans-
versely wrinkled, ± punctulate ( variable ) .
Elytra subquadrate; width elytra prothorax
1.90 and 1.92; disc virtually estriate, with 3
conspicuous punctiform foveae near posi-
tion of each 3rd interval. Measurements:
length 3.8-4.2; width 1.5-1.7 mm.
Types. Holotype c^ (M.C.Z., Type No.
31,502) and 2 paratypes from Nganduo to
Yunzain, Mongi Watershed, Huon Pen.,
N-E. N. G., 1000-1500 m, Apr. 6, 1955 ( E.
O. Wilson); and additional paratypes as
follows. N-E. N. G.: 1, Gemeheng, Mongi
\\'atershed, Huon Pen., 1300 m, Apr. 11-13,
1955 (E. O. Wilson, M.C.Z.); 1, vie. Xad-
zab, July 1944 (Darlington); 2, Sattelberg
(British Mus., ex Coll. Hauser). West
N. G.: 8, Wissel Lakes, Moanemani, Kamo
Vy., 1500 m, Aug. 14, 1962 (Sedlacek); 2,
Wissel Lakes, Enarotadi, 1500 m, Aug. 14,
1962 (Sedlacek).
Measured specimens. The i holotype and
1 9 paratype from Nganduo to Yunzain.
Notes. This species, like the preceding
( hasalis ) , occurs at relatively low altitudes,
sometimes down almost to sea level (at
Nadzab ) .
Scopodes cheesmoni n. sp.
Description. With characters of genus;
form (Fig. 122) compact, convex; blue-
purple (in general blue with elytra puiple
with foveae bluer); shining, reticulate
microsculpture faint or obsolete even on
elytra. Head 1.31 and 1.35 width prothorax;
front with 7 sulci running into coarsely
rugose areas posteriorly and usually anteri-
orly. Frothorax narrower and less angulate
than usual; width/length 1.13 and 1.18;
sides margined, each with seta on small
projection c. M', from apex; disc coarsely
transversely sulcate or wrinkled, ± punctu-
late especially anteriorly. Elytra subquad-
rate, slightly narrowed toward base; width
elytra/prothorax 1.98 and 2.09; disc estriate,
with 3 conspicuous punctiform foveae on
position of each 3rd interval. Measure-
ments: length 3.9-4.4; width 1.6-1.8 mm.
Types. Holotype $ (British Mus.) and
2 paratypes from Mt. Lina, Cyclops Mts.,
West N. G., 3500 ft. (1067 m). Mar. 1936
( Cheesman ) ; and additional paratypes as
follows. West N. G.: 4, Cyclops Mts.,
3400-4500 ft. (c. 1040-1370 m). Mar. 1936
(Cheesman); 3, Rattan Camp, 1150 m,
Feb.-Mar., and 1, Sigi Camp, 1350 m. Snow
Mts., Feb. 28, 1939 (Toxopeus); 1, Bivak
36, 1220 m, July 30, and 1, Bivak 39, 1300
m, June 30, 1959, Star Rge. (Neth. New
Guinea Exp., Leiden Mus.). N-E. N. G.:
1, Chimbu Vv., Bismarck Rge., 5000-7500
ft. (c. 1500-2300 m), Oct. 1944 (Darling-
ton); 1, Eliptamin Vy., 1665-2530 m, June
23-30, 1959 (W. W. Brandt, Bishop Mus.).
Some paratypes in M.C.Z., Type No. 31,503.
Measured specimens. The i holotype and
1 9 paratype from Mt. Lina.
Notes. Although not very different in
form and basic structure from the several
preceding compact species, cheesmani does
suggest transition toward the following
(adonis).
The habits of this species are suggested in
Notes under the genus.
Scopodes adonis n. sp.
Description. With characters of genus;
form ( Fig. 123 ) less compact, with longer
prothorax and more oval elytra, than in
other members of the "New Guinean group"
of Sco})odes; purple; front anteriorly,
clypeus, and labrum cupreous; legs dark;
antennae with basal and outer segments
brown, segments 2-5 paler; shining, upper
202 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
surface without reticulate microsculpture,
sparsely punctulate. Head 1.37 and 1.32
width prothorax; front with c. 7 slightly
unequal sulci behind level of anterior
supraocular punctures, the sulci abbreviated
anteriorly and cur\'ing toward sides pos-
teriorly; neck irregularly rugose. Prothorax
oval long; width length 0.93 and 0.96;
sides not margined, not angulate, each with
seta-bearing puncture c. Va from apex; disc
strongly convex, with fine but distinct mid-
dle line, weakly transversely wrinkled.
Elytra suboval; width elytra prothorax 2.13
and 2.17; humeri narrower and more
broadly rounded than in other species of
group; outer-apical angles obtusely angu-
late, sutural angles acute; striae absent; 3
minute, inconspicuous seta-bearing punc-
tures on position of each 3rd interval. Mea-
siirement.s: length 4.7-4.9; width 1.7-1.9
mm.
T\j])es. Ilolotype i (Bishop Mus.) and
8 paratypes (some in M.C.Z., Type No.
31,504) from Torricelli Mts., Mokai Village,
N-E. N. G., 750 m, Dec. 8, 16 (holotype,
Dec. 8), 1958 (W. W. Brandt); and 2 para-
types, Torricelli Mts., Mobitei, 750 m, Feb.
28-Mar. 4 and Apr. 1-15, 1959 (W. W.
Brandt, Bishop Mus.).
Measured speeimcn.s. The i holotype and
1 9 paratype from Mokai Village.
Notes. This very distinct species is dis-
tinguished from all others known to me by
form, prothorax long with margins obsolete,
elytra oval with acute sutural angles, elytral
foveae reduced, and color. However, as
noted in discussion under the genus, it prob-
ably does not represent a separate stock in
New Guinea but seems to be at the end of
an evolutionary sequence which includes
other New Guinean species.
Tribe HEXAGONIINI
Csiki 19.'32, Colcop. Cat., (;aial)itl;K', Ilaipalinac 7,
p. 1506 (sec tor s>n()n\niy and additional rclcr-
enccs).
flcxofioniitue Jcanncl 1948, (lolrop. C'aiahiqiics dc
la i^ogion Malgadic, I'ait 2, p. 7.59.
This is a small tribe (4 genera) of char-
acteristically formed ( Fig. 124 ) , subparallel,
often flattened carabids at least some of
which are specialized to live under the
leaf sheaths of plants. In this tribe the
inner lobe of the maxilla has a slender,
Diocahle apical segment that is diagnostic,
occurring in no other Carabidae except the
Cicindelinae.
The tribe is confined to the Old-World
tropics. The genus Hexa^onia (below) is
widely distributed there; 1 additional genus
occurs in the Oriental Region; 2 more, in
Madagascar.
Genus HEXAGON/A Kirby
Kirby 1825, Trans. Linnean Soc. London 14, p.
563.
Csiki 1932, Coleop. Cat., Cavabidat', Ilaipalinae 7,
p. 1506 (see for synonymy and additional refer-
ences ) .
Jeannel 1948, Coleop. Carahiques de la Region
Malgache, Part 2, p. 759.
Basilewsky 1948, Bull. Mus. Hist. \at. Belgian
Congo 24, p. 3 (African species).
Diaiinosis. See under tribe, of which this
is the only genus represented in New
Guinea.
Deseri))t}on. None required here.
Ty))e s))eeies. II. terniiiuifa Kirby, of SE.
Asia.
Generie distribution. Tropical .\sia and
islands to New Guinea and northeastern
Australia; Africa, Madagascar.
Notes. Members of this genus are rather
diverse in the Oriental Region including
the Philii)pines, but ()nl\ I spc^cies group
extends to New (Guinea and Australia.
ProbabK' bee;uise the\- oceup) an unusual
niche and perhaps also beeausi' the\' are
diurnal and nia\ not ll\ to light, these
insects are rarel\ eolleeled. The 1 New
Guinean and the 1 ( nnd(\scrib(Kl ) .Austra-
lian s|ieeies are eaeli known Ironi a single
collection made b\ nusell, by breaking
down tall grass and other a([uatic \-egeta-
tion into water and picking np the beetles
;is the\ came to the surhice.
The Carabid Beetles of New Guinea • Darlington 203
Hexagonia popua n. sp.
Description. With characters of tribe;
form as in Figure 124; head black; pro-
notum piceous, reddish at base and apex;
elytra red in anterior 1-2 or more, black
posteriorly, the black area extending farther
forward on inner than on outer part of
elytra; lower surface irregularly reddish and
piceous; appendages brown; shining, reticu-
late microsculpture faint or absent on head
and pronotum, imperfect on elytra; head
and pronotum irregularly punctate. Head
1.05 and 1.05 width prothorax; front semi-
circularly impressed each side; neck deeply
transversely impressed. Prothorax subcor-
date; width length 1.06 and 1.09; base/apex
not calculated (because prothorax rounded
into neck without distinct anterior angles);
margins narrow, each with seta c. % from
apex, without posterior seta; disc with mid-
dle groove deeply impressed and subpunc-
tate, other impressions irregular and weak.
Elytra: width elytra/prothorax 1.66 and
1.67; striae well impressed, punctate; inter-
vals slightly convex, 3rd with 3 or 4 con-
spicuous seta-bearing punctures, 5th with 1
such puncture c. ^.i from apex on outer edge.
Lower surface: head below transversely
rugulose; prothorax below subrugosely
punctate. Inner wings full. Legs moderate;
tarsi wide; 4th tarsal segments very deeply
emarginate, long-lobed; claws simple, not
toothed. Secondary sexual characters: S
tarsi not or not much modified, without
special squamules; i with 2, 9 3 short
setae near apex each side last ventral seg-
ment. Measurements: length 7.2-7.8; width
2.2-2.4 mm.
Types. Holotype S (M.C.Z., Type No.
31,505) and 12 paratypes all from Aitape,
N-E. N. G., Aug. 1944 (Darhngton).
Measured specimens. The c^ holotype and
1 ? paratype (sexes of these specimens
determined by dissection).
Notes. This new Hexagonia is probably
related to lucasseni van der Poll of Java but
is slightly more slender and much darker,
with pronotum piceous (red in lucasseni)
and elytra more extensively black posteri-
orly. One or more similar but apparently
undescribed species occur in the Philip-
pines.
The habitat of the species is noted under
the genus.
Tribe ODACANTHINI
Sloane 1917, Proc. Linnean Soc. New South Wales
42, p. 413.
1923, Proe. Linnean Soe. New South
Wales 50, p. 30.
Jedlieka 1963, Ent. Abhandlungen 28, p. 488.
Habu 1967, Fauna Japonica, Carabidae, Tnui-
eatipennes Group, p. 13.
Odacantliidae Jeannel 1948, Coleop. Carabiques
de la Region Malgaehe, Part 2, p. 745.
Odacanthinac Basilewsky 1953, Exploration Pare
National I'Upemba, Fase. 10, Carabidae, p. 108.
CoIUuriui Csiki 1932, Coleop. Cat., Carabidae,
Ilaipalinae 7, p. 1517 (see for .synonymy and
additional references ) .
Liebke 1938, Festschrift Einbrik Strand 4, pp. 37-
141.
Most members of this tribe have a char-
acteristic form (Figs. 125-131), with pro-
thorax very long and narrow, usually much
narrower than head. Technical characters
of the tribe are given by authors cited
above. The tribe is well represented in
the tropics of both hemispheres, less well
represented in most temperate regions, but
several remarkable endemic genera occur
in Australia.
Liebke ( 1938 ) has published a useful
generic classification of this tribe for the
world. His classification is, however, artifi-
cial, as shown by the failure of some of my
new New Guinean species to fit into it.
The form of the 4th hind-tarsal segments,
used by Liebke in the first couplet of his
key, is a particularly unsatisfactory generic
character. Form of the 4th hind-tarsal seg-
ments does characterize some genera, but
it is extremely variable in others. See, for
example. Notes under Dicraspeda in the
following pages. However, I cannot under-
take to revise the generic classification in
dealing with the few members of the tribe
that occur in New Guinea, except in the
case of primarily New Guinean genera.
204 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Ecologically, Odacaiithini are active,
winged carabids some of which live in
foliage and some on the ground. Of the
New Guinean forms, Dicraspeda ( includ-
ing Philcmonia and Macrocentra) lives in
understory foliage of rain forest; Clarcncia
and some CoUiuris, in or under low vegeta-
tion or dead leaves in wet places; Casnoideo,
among reeds and in other vegetation in
water; Eudalia (in Australia), Dohodura,
and I think also Lachnothorax, in gravel
and among stones bv running water; and
Basisticu.s ( in Australia ) , on the ground
in relatively dry places including open
Eucahjptu.s woodland.
Eight genera, 19 species of Odacanthini,
are known from New Guinea. Two or 3
stocks of CoIUuris, 1 species of Casnoidca,
and Lachnothorax have probably reached
New Guinea rather rec(uitly from the
Oriental Region. A second species of
Casnoidea and also Basi.sticus micans (which
may be related to Coditiris) are shared
with Australia. Cdarcncia and Eudalia are
shared with Australia and may be of Aus-
tralian origin. Monotypic Dohodura is con-
fined to New Guinea but may be derived
from Eudalia. And Dicra.s))eda {.sensu lato)
has radiated chiefly in the rain forests of
New Guinea, where 6 diverse species now
exist.
Key to Genera of Odac:antiiini of
New Guinea
1. Lateral niaiyins of protlinrax iDtomplcte
(i)i(l 4tli liiiul-tarsal sciiiiK'iils \ci>' lonij;-
lobc'd; strikinjfly bicokiied, bkick (or jiiccous)
and red ( p. 207 ) (.Uisnoidca
— Not as above in one oi' more ways; nsually
not blaek-and-red l)ieoIore(l (olisenrely so
in lidsislicns ) 2
2. Antennae with 3rd se,unients \er_\ h)njf, c.
% longer than 4th segments (p. 209) _..
Cldicncid
— Antennae with ■ivd segments shorter, ('{jual
to or not more tlian 'A longer tlian 4t]i
segments 3
3. Head with fine eosta eaeh side above eye
ami pronotum elianneled at sides and mid-
(Ue; ( l)ase ot elytra not i(un\clti pnnetate-
striate) (yi. 210) l^icraspcdd
— Not as above in one oi more ways 4
4. Body pnbeseent ([i. 214) I .(kIiiioIIidiiix
- Not pubescent 5
5. Eyes not margined on inner edges by costae
and front smooth 6
- Eyes usually margined by costae, or if
(rarely) not margined, front so coarsely
pimetate that supraocular costae are indis-
tinct 7
6. Elytra not spined (p. 214) EiidaUd
- Elytra spined (see also Notes under this
genus) (p. 215) ... Dohodura
7. Side margins of prothorax absent or incom-
plete or ( if margins nearly complete ) pro-
notum not channeled at middle; elytra not
\erv differentlv sculptured at base and apex
(p.' 205) _._! Colliuris
- Side margins of prothorax complete and
pronotum channeled at middle; elytra
coarsely punctate-striate in anterior %,
smooth posteriori)' ( p. 20(S ) Basisticiis
Genus COLLIURIS Degeer
Degeer 1774, Mem. Hist. Insectes 4, p. 79.
Gsiki 19.32, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1.518 (see for synonymy and additional refer-
ences ) .
Liebke 1938, Eestschrift Endirik Strand 4, p. 45.
ledlicka 1963, Ent. Abhandlungen 28, pp. 489.
490.
Diuiinosis. See preceding Keij to Genera.
Deseription. None required here.
Tijpe speeies. Attelahus surinameusis
Linnaeus, of South America.
Generic distribtition. All wami regions
of the worlfl. In the Asiatic- Australian area,
species are moderately numerous and di-
verse in southern Asia and the western
Malay Archipelago, but only 4 species
( representing 3 indc^pcMident stocks ) have
been foiuid in New (Guinea. According to
Liebke, the genus does not reach Australia,
but I found 3 species of it ( representing 2
ol the same stocks that are in New Guinea)
in North Oueensland in I95(S.
Notes. I cannot fit the New Guinean
species into T-iebkes subg(Mi(Ma satisfac-
torily. ('. ro.s.si (below) and also the Aus-
tralian C. ohscura Gastelnau ha\e \irtualK
complete lateral prothoracic margins, which
rules these species out of Colliuris entirely,
according to Li(>bke (who in fact incor-
rectly put ohscura in Dicraspeda). Never-
theless, in loiiu and other characters these
The Carabiu Beetles of New Guinea • Darlington
205
species seem to me to be closely related to
Colliuris subgenus EticoUiuris of Africa and
the Oriental Region.
Key to Species of Colliuris of New Guinea
1. Prothorax with lateral margins; intervals 3,
5, 7 with seta-bearing punctures; (pronotum
and elytral striae coarsely punctate, head
smooth) (p. 205) rossi
- Prothorax without lateral margins, or margins
reduced to sutures; only 3rd intervals with
seta-bearing punctures 2
2. Not maculate; subcylindric, elytra not more
than 2x width prothorax; entire upper sur-
face coarsely punctate (p. 205) fii.svipcnnis
- Quadrimaculate; broader, elytra more than
2X width prothorax; upper surface c. im-
punctate except for finely punctate elytral
striae 3
3. Less slender, prothoracic width/length 0.60
and 0.63; elytra not or not much impressed
(p. 206) papiiu
- More slender, prothoracic width/length 0.46
and 0.46; elytra transversely impressed near
base (see also Description) (p. 206) par
Colliuris rossi n. sp.
Description. Form (Fig. 125) of Colliuris
of Oriental fiiscipennis group; black, legs
brown with dark knees, antennae brown,
paler at base; shining, without reticulate
microsculpture. Head 1.13 width prothorax;
right mandible with acute tooth on inner
edge; a fine costa over each eye separated
from eye by groove; anterior and posterior
supraocular seta-bearing punctures present
but no other setae posteriorly; front convex,
slightly impressed at middle and anteri-
orly, impunctate; mentum with acute tooth;
ligula broad, 2-setose; palpi slender, acumi-
nate, not pubescent. Prothorax long-oval;
width/length 0.71; base apex 1.34; side
margins entire except confused by puncta-
tion at extreme base, narrow, each with 1
seta before middle; middle line light; disc
strongly convex, irregularly coarsely punc-
tate. Elytra subparallel; width elytra/pro-
thorax 2.00; apices obliquely sinuate-trun-
cate with outer and sutural angles blunt;
striae coarsely punctate, the punctures be-
coming finer posteriorly; intervals 3, 5, 7
each with 4 to 6 well spaced seta-bearing
punctures. Inner wings full. Lower surface
not pubescent, coarsely punctate anteriorly
including sides of metasternum, smooth
posteriorly. Legs normal; tarsi not pubes-
cent and not sulcate above; 4th hind-tarsal
segments shallowly emarginate. Secondary
sexual characters: 6 front tarsi narrow, 3
segments narrowly 2-seriately squamulose;
c5 last ventral segment c. semicircularly
emarginate at apex, with 1 seta each side; $
copulatory organs as in Figure 179; 9 un-
known. Measurements: length c. 6.5; width
2.0 mm.
Type. Holotype £ (Gal. Acad.) from
Finschhafen, N-E. N. G., May 7, 1944 (E.
S. Ross); the type is unique.
Notes. This new species seems closely
allied only to C. obscura (Castelnau) of
NE. Australia, but ohscura has only the 3rd
elytral intervals with seta-bearing punc-
tures. Otherwise the two species agree in
most characters including presence of
virtually entire prothoracic margins, distri-
bution of coarse punctation (head impunc-
tate, pronotum contrastingly coarsely punc-
tate), i secondary sexual characters, and
presence of an acute tooth on the right
mandible in some individuals. However,
presence of the mandibular tooth does seem
to be an individual character in ohscura:
in my series from Cairns, the tooth is well
developed in some and almost absent in
other specimens.
The generic assignment of ohscura and
rossi is doubtful. Sloane (1923, Froc.
Linnean Soc. New South Wales 48, p. 31 )
thought ohscura might go in Arame, which
is doubtful, and Liebke (1938, Festschrift
Embrik Strand 4, p. 89) put it in Dicraspeda,
which is certainly wrong. Only the pres-
ence of nearly entire prothoracic margins
prevents placing both ohscura and rossi in
Colliuris in Liebke's classification, and I
doubt if the prothoracic margins are of
generic value in this case.
Colliuris fuscipennis (Chaudoir)
Chaudoir 1850, Bull. Soc. Nat. Moscow 23, Part 1,
p. 26 {Casnoiiid) .
206
Bulletin Miiscuni of Comparative Zoology, Vol. 137, No. 1
Andrewes 1927, Ann. Mag. Nat. Hist. (9) 19, p.
106 (Odacantha).
Csiki 1932, Colfop. Cat., Carabidae, Harpalinae 7,
p. 1527 (see for synonymy, '■varieties," and addi-
tional references).
Lie1)ke 1938, Festschrift Enilirik Strand 4, p. 65,
fi,y. 27.
Tedlicka 1963, Ent. Abhandkingen 28, p. 494, figs.
190-193.
Description. None required here. Note
form rather slender; eolor bhiek with apex
of elytra reddish, legs testaceous; upper
surface including head coarsely punctate;
length c. 5% mm.
Type. From China ("Chine, Tchusan?");
in Oberthiir Coll., Paris Mus. (not seen).
Occurrence in Ncic CAiinea. Papua:
1, Lake Daviumbu, Fly R., Aug. 19-30,
1936 (Archbold Exp., A.M.N.H.).
Notes. Fiiscipennis is the oldest name
for a very common species (n- group of
closely related species previously known
from SE. Asia to Celebes and the Philip-
pines. Andrewes (1927) says of it that
fuscijx'nnis Chaudoir, punciata Nietner,
haeniorrhoidalis Motscliulsky, and jlavi-
cauda Bates "appear to dilfer very little
from each other; they may all prove to
belong to the same species, but at present
I have not the means of dc-ciding this."
(isikis, Liebke's, and Jedlicka's treatment
of some of the doubtful forms as subspecies
or varieties is not acceptable. Under these
circumstances I can only refer the New
Cuinean individual to fuscijx'nuis scnsn
l(il(K pending revision ol all related tornis.
Colliuris papua n. sp.
Description. With character's of genus;
lonn { I'ig. 126) ol ('olliuri.s, with mod-
erately broad elytra scarcely impressed be-
lore middle; black, each elytron with 2 c.
round yellow spots, centered on tth and 5th
intervals, belore middle and before apex;
apjiendages brown, antennae paler al bas(.\
moderatcK shining, reticulate microsculp-
ture hunt and r. isodiametric on front, trans-
verse on head posteriori)' and on pronofum,
indistinct on elytra. Head 1.44 and 1.11
width prothorax; 2 setae ()\cr each e\c bnt
no other setae posteriorly; front scarcely
impressed, impunctate. Prothorax long,
swollen at sides behind middle, strongly
narrowed anteriorly; width length 0.60 and
0.63; base apex 1.54 and 1.56; lateral mar-
gins reduced to sutures, each with 1 seta
before middle; disc strongly convex, base
scarcely impressed, middle line fine, surface
faintly transversely strigulose, punctate
across base. Elytra: width elytra prothorax
2.24 and 2.22; apices obliquely truncate
( slightly emarginate ) with outer angles
rounded, sutural angles scarcely blunted;
striae formed In rows of small punctures
which become minute posteriorly; 3rd in-
tervals with c. 4 seta-bearing punctures, in-
tervals 5 and 7 without punctures. Inner
winfis full. Lower surface punctate only
around front coxae and at front of meso-
sternum. Lci^s normal; tarsi not pubescent
and not sulcate above; 4th hind-tarsal s(>g-
ments shallowly emarginate. Secondary
sexual characters: A front tarsi narrow,
narrowly 2-seriately squamulose below;
last ventral segment slightK' emarginate at
apex in S , variably impressed in 9 , with
1 seta each side in c5 , 2 in 9 . Measure-
ments: length e. 5.5-6.0; width 1.6-1.8 mm.
Types. Holotype i (M.C.Z., Type No.
31,506 ) and 35 paratypes all from Dobo-
dura, Papua, Mar.-July 1944 ( Darlington ) .
Measured s])eeiniens. The ' holot\pe and
1 9 paratype.
Notes. See Notes imder the following
species, par.
Colliuris par n. sp.
Description. Similar in forTU and most
characters to preceding t papua ) but more
slender, with elytra impressed before mid-
dle; anterior eUtral spots longer antl almost
eonlined to 5th inlei\als, legs pale at base;
reticulate microsculptnic not distinct on
pronotuni. Head 1.65 and 1. 58 width \nn-
thorax. Prothorax: width length 0.46 and
0.46; base apex 1.55 and 1.56; disc trans-
\('rsel\ iinprc^ssed and constricted before
base. Elytra: width ebtra prothorax 2.60
and 2.62. ]A)irer surface as in pa))ua excei")t
The Carabid Beetles of New Guinea • Darlington 207
with row of coarse punctures each side
prosternum before coxae. Secondary sexual
characters: as for papiia. Measurements:
length c. 6.0-7.0; width 1.7-2.0 mm.
Types. Holotype $ (M.C.Z., Type No.
31,507 ) and 5 paratypes all from Hollandia,
West N. G., July-Sept. 1944 ( Darlington ) .
Additional material. N-E. N. G.: 7,
Aitape, Aug. 1944 (Darlington).
Measured specimens. The $ holotype and
1 9 paratype.
Notes. C. papua and par seem to be
allopatric representatives of one ancestral
stock, but they differ too much to be con-
sidered subspecies. C. par is the more
widely distributed: I found it at Cape
Gloucester, New Britain, in 1944, and
(1 9 ) at Lockerbie, near the tip of Cape
York, Queensland, Australia, in 1958. A
second 9 from Lockerbie is superficially
similar but much more compact and differs
in other details. I think it probably rep-
resents a 3rd, distinct species of this group
of Colliuris.
Genus CASNOIDEA Castelnau
Castelnau 1834, Etudes Ent. 1, p. 40.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1534 (see for synonymy, additional refer-
ences, and list of species).
Jedlicka 1963, Ent. Abhandlungen 28, pp. 489,
498.
Ophinnca Eschscholtz 1829, Zool. Atlas 2, p. 5
(not Ophionea Klug 1821).
Liebke 1938, Festschrift Embrik Strand 4, p. 79.
Diagnosis. See preceding Key to Genera.
Description. None required here.
Type species. Of both Casnoidea and
Ophionea Eschscholtz, Attelahus indica
Thunberg.
Generic distribution. SE. Asia including
Ceylon and Japan to Australia; a species
recorded also from the Seychelles Is.
Notes. This genus of slender, usually
strikingly bicolored (red and black) carabids
includes several species widely distributed
in SE. Asia and the Malay Archipelago.
They are usually found in grass, reeds, and
other vegetation growing in water.
Key to Species of Casnoidea of New Guinea
1. Color dark with basal Mi or -/-, of elytra red
(p. 207) ^estroi
- Color red with head, post-median elytral
fascia, and sometimes bases of elytra dark
(the post-median fascia with a pale spot on
each elytron) 2
2. Pronotum conspicuously punctate (p. 207)
ptiHcticollis
- Pronotum not conspicuously punctate 3
3. Elytra not dark at l)ase (p. 208) -.
( nigrofasciaia )
- Elytra dark at base (p. 208) _.... (imJira)
Casnoidea gesfroi (Maindron)
Maindron 1910, Bull. Soc. Ent. France for 1910,
p. 34 ( Oi)J}ionc(i) .
Dupuis 1913, Ann. Soc. Ent. Belgium 57, p. 270.
Liebke 1938, Festschrift Embrik Strand 4, p. 79,
fig. 60 {Ophionea).
gestiunis Seidlitz 1912, Archiv fiir Naturgeschichte
77, Part 3, p. 155 (error for gestroi).
Description. None required here; see
preceding Key to Species; length c. 7.0-
7.5 mm.
Type. From Dilo, Papua, July 1890 (D.
Loria); presumably in Paris Mas. (not
seen ) .
Occurrence in New Guinea. Papua: 7,
Dobodura, Mar.-July 1944 (Darlington); 2,
Kiunga, Fly R., July 15-21, Aug. 1-3, 1957
(W. W. Brandt, Bishop Mus.). West N. G.:
1, Waris, S. of Hollandia, 450-500 m, Aug.
8-15, 1959 (T. C. Maa, Bishop Mus.); 1,
Wasian, Vogelkop, Sept. 1939 (Wind,
M.C.Z.).
Notes. Tliis distinct species is evidently
widely distributed in New Guinea and is
represented also on New Britain (an un-
described subspecies from Cape Glouces-
ter) but is unknown elsewhere.
Casnoidea puncficollis (Sloane)
Sloane 1923, Proc. Linnean Soc. New South \\'ales
48, p. 31 (Ophionea).
Liebke 1938, Festschrift Embrik Strand 4, p. 80
(Ophionea).
Description ( for recognition only ) . Form
as in Figure 127; red, head black, elytra
with broad transverse fascia ( bluish ) black,
the fascia with an elongate pale fleck on
each 5th interval, legs bicolored; shining.
20S Bulletin Miiscmu of Comparative Zoology, Vol. 137, No. 1
without reticulate microsculpture. Head
short, rounded posteriorly; front wrinkled
anteriorly, impunctate. Prothorax long-
oval; side margins irregularly indicated
anteriorly; disc conspicuously punctate.
Eh/fra punctate-striate. Measurements (New
Guinean specimen): length c. 7.5; width
2.1 mm.
Type. From Burdekin R., Queensland,
Australia; in Sloane Coll., C.S.I.R.O., Can-
berra ( seen ) .
Occurrence in New Guinea. Papua: 1,
Kiunga, Fly R., Aug. 1-3, 1957 (W. W.
Brandt, Bishop Mus.).
Notes. I do not have puncticoUis from
Australia and have identified the New
Guinean individual from description. Note
that C. ^estroi (preceding species) as well
as puncticoUis occurred at Kiunga.
iCasnoidea nigrofasciafa (Schmidt-Goebel))
Schmidt-Goebel 1846, Faumila Coleop. Bir-
iiianiae, p. 21 (Opiiionea) .
Aiidrewes 1930, Treubia 7, Suppk-ineut, p. 3'34
( Ophionea ) .
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1535 ( see for synonymy and additional refer-
ences ) .
I.iebki' 1938, Festschrift Embrik Strand 4, p. 80,
fiji. 57 (Opiiiouca ).
Description. None required here; see
preceding Key to Sj)ecies.
Type. From Buruia; in Prague Mus.
(not seen).
Occurrence in New Guinea. Probably
does not occur.
Notes. C. ni<irofasciaia ranges from SE.
Asia to Java and Borneo. It is apparently
not recorded bom Celebes or the Moluccas.
New Guinea is included in the species'
range by Csiki, but I can find no authority
for this. 1 suspect that a too-hasty com-
piler, not noticing the negative, picked
"New Guinea" out of Andrewes' (1930)
statement that "I havi' seen no examples
cithci' hoin Jaj^an or New (iuinea."
(Casno/c/ea indica (Thunberg))
i'lmnbc'ru 1784, Novas Inseclorinn Species .3, p.
()8, flu. 81 (Altclalms).
Andrewes 1930, Cat. Indian Insects, Part 18,
Carabidae, p. 241 {Ophionea).
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1534 ( see for synonymy and additional refer-
ences ) .
Licbke 1938, Festschrift Embrik Strand 4, p. 79,
ti^. 55 ( Ophionea).
Louwerens 1958, Treubia 24, p. 249 (Moluccas)
{Oi)]uonea).
Description. None required here; see
preceding Key to Species.
Type. From "India orientali"; presumed
lost (not seen).
Occurrence in New Guinea. Doubtful.
Notes. This common Oriental carabid
ranges from SE. Asia including Ceylon
and Japan to Celel>es and the Moluccas.
A specimen in the British Museum is labeled
"Dory, New Guinea" but may be from
Celebes or the Moluccas ( see Part I of m\
"Carabid Beetles of New Guinea," p. 331).
Other collectors have failed to find the
species in New Guinea. Andrewes' state-
ment that indica occurs south to New
Cruinea is probabh' based on the doubtful
"Dory specimen.
Genus BASISTICUS Sloane
Sloane 1917, Proc. Linnean Soc. New South Wales
42, p. 415.
1923, Proc. I.innean Soc. New South
Wales 48, p. 30.
Liebke 19.38, Festschrift Emlirik Strand 4, p. 81.
Diapiosis. See preceding Key to Genera.
Description. None required here.
Type species. Odacant}ia micans Macleay
( below ) .
Generic distribution. As of the single
known species.
Notes. This genus is close to CU)lliuris
(sensu Jafo ), from which it differs in luuing
the lateral margins of the prothora.x entire.
Basisficus micans (Macleay)
MacleaN 18()4, 'I'rans. Kwi. Soc. New South Wales
1, p. 107 (Oducanlha).
CsiVi 1932, Coleop. ('at., (Carabidae, Harpalinae 7,
p. 15.3.5 (see for additional references),
l.iebke 1938, Festschrift Fnibrik Strand 4, p. 81.
Description ( lor recognition ouK). Form
The Carabid Beetles of New Guinea
Darlinfitoii
209
of ColUuris; head and prothorax red, base of
elytra dark reddish, smooth part of elytra
piceous, antennae red, legs dark with pale
bases; elytra very coarsely punctate-striate
in anterior Vs, smooth with striae of minute
punctules in posterior %; length c. 6.5 mm.
Type. From Port Denison, northern Aus-
tralia ( presumably near Bowen, Queens-
land); probably in Macleay Mus., Sydney
(not seen).
Occurrence in New Guinea. Papua: 1,
Rouku, Morehead R., March 1962 (W. W.
Brandt, C.S.I.R.O.).
Notes. This is an Australian species, well
known in North Queensland. I have speci-
mens from the vicinity of Cairns, Mareeba,
and Townsville. The single individual from
New Guinea matches Australian ones well.
In Australia, this insect is found on the
ground in open woodland; the type was
"found under dried cow dung."
Genus CLARENCIA Sloane
Sloane 1917, Proc. Linnean Soc. New South Wales
42, p. 415.
1923, Proc. Linnean Soc. New South
Wales 48, p. 30.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1535.
Liebke 1938, Festschrift Embrik Strand 4, p. 81.
Diagnosis. See preceding Key to Genera;
note form of large CoUiuris; antennae with
very long 3rd segments (and see Notes,
below ) ; £ front tarsi with 3 segments with
numerous narrow squamae loosely ar-
ranged (not 2-seriate as in CoUiuris); last
ventral segment slightly emarginate at apex
in both sexes, with 1 seta each side in <^ , 2
in 9 .
Description. None required here.
Type .species. Casnonia aliena Pascoe, of
Australia.
Generic distribution. Eastern Australia,
New Guinea.
Notes. Although only 1 Clarencia is cur-
rently recognized (Csiki, 1932), 4 species
are represented in Australian material col-
lected by me in 1957-1958. One of these
species (described below as quadridens)
occurs also in New Guinea, and 1 additional
species of the genus is endemic in New
Guinea.
The antennae of some Australian Clar-
encia not only have very long 3rd seg-
ments but also have the 4th segments
uniquely modified: expanded and obliquely
truncate at apex so that the 5th segments
hinge forward, and with the pubescence
of the 4th segments restricted to the seg-
ments' anterior edges. This modification
of the 4th segments is only slightly indicated
in the New Guinean species, more clearly
in papua than in quadridens.
The species of Clarencia are usually
found in wet places, often by standing
water, either among wet leaves or in or
under low vegetation.
Key to Species of Clarencia of New Guinea
1. Elytra toothed at sutural and outer-apical
angles (p. 209) (iiiadridens
- Elytra not toothed (p. 210) \ni\>iia
Clarencia quadridens n. sp.
Description. With characters of genus;
form as in Figure 128, with elytra trans-
versely impressed near base; black, elytra ±
yellowish at apex but not spotted, epipleuri
pale, femora pale at base, dark at apex,
tibiae dark banded with pale, tarsi pale,
antennae brown darker basally; shining,
without distinct reticulate microsculpture.
Head 1.37 and 1.27 width prothorax; front
with conspicuous V-shaped impression an-
teriorly, impunctate. Prothorax long, with
sides swollen behind middle; width length
0.59 and 0.61; base apex 1.38 and 1.40;
disc very convex, with fine middle line,
punctate across base with a few punctures
along lateral margins and across apex but
otherwise impunctate. Elytra: width elytra/
prothorax 2.30 and 2.47; apices obliquely
sinuate-truncate with outer and sutural
angles acutely dentate; striae formed by
lines of punctures anteriorly, obsolete pos-
teriorly; 3rd intervals with c. 6 and 5th
intervals with c. 4 seta-bearing punc-
tures. Legs slender; tarsi above not pubes-
cent and not sulcate; 4th hind-tarsal seg-
210 Bulletin Museum of Comparative Zoology. Vol. 137, No. 1
ments shallovvly emarginate. Measurements:
length 10.0-11.5; width 2.8-3.1 mm.
Types. Holotype c^ (M.C.Z., Type No.
31,508) and 14 paratypes from Hollandia,
West N. G., Jiily-Sept. 1944 (Darhiigton);
and additional paratypes from West N. G.
as follows: 4, Hollandia, Apr., May 1945
(B. Malkin, U.S.N.M.); 2, "Neth. N. G."
without further locality (T. Aarons, Cal.
Acad.).
Additional material. West N. G.: 1
teneral S , Maffin Bay, Aug. 1944 ( Dar-
lington). Papua: 1 $, Normanby Is.,
Wakaiuna, Sewa Bay, Jan. 1-8, 1957 (W.
W. Brandt, Bishop Mus.).
Measured specimens. The S holotype and
1 9 para type from Hollandia.
Notes. The denticulate elytra distinguish
this species from all other known Clarencia
in Australia as well as New Guinea. This
species occurs in Australia: I have one
9 from near Cairns, N. Queensland (Dar-
lington). This Australian specimen and the
one from Normanby Is. have more yellow
at apex of elytra than Hollandia speci-
mens do.
Clarencia papua n. sp.
Description. With characters of genus;
form c. as in quadridens (above) except
elytra relatively narrower and not dentate;
])lack, elytra scarcely paler at apex, legs
bicolored as in quadridens, antennae brown;
shining, without distinct reticulate micro-
sculpture. Head 1.32 and 1.22 width pro-
thorax; front convex except impressed an-
teriorly as in quadridens, impunctate.
Frotliora.x formed as in (juadridens but
slightly shorter; width lengtli 0.62 and 0.68;
base/apex 1.37 and 1.41; disc very convex,
middle line fine, surface closely wrinkled-
punctate across base, variabK punctate
across apex, and more extensively punctate
at sides than in quadridens. Elytra: width
elytra prothorax 2.21 and 2.10; ajiices
oblicjuely sinuate-truncate with outer and
sutural angl(\s narrowK' rounded; striae
nearly entire (longer than in (piadridens),
punctate, the punctures becoming minute
posteriorly; 3rd and 5th intervals with a
few seta-bearing punctures. Legs c. as in
(juadridens. Measurements: length c. 9.0-
10.0; width 2.5-2.9 mm.
Types. Holotype $ (U.S.N.M.) and 7
paratypes (some in M.C.Z., Type No. 31,509)
from Hollandia, West N. G., Apr., May,
June (holotype, Apr.) 1945 (B. Malkin); 1
paratype, same locality, "11/5/44" (W. T.
Nailon, Fenton Coll.); 1 paratvpe, Yent-
chan. Main R., Sepik, N-E. N.' G., Feb.
1965 (R. Hornabrook).
Additional material. Papua: 2, Lake
Daviumbu, Fly R., Aug. 19-30, Sept. 11-20,
1936 (Archbold Exp., A.M.N.H.).
Measured specimens. The c5 holotype and
1 9 paratype.
Notes. See preceding Description for
details distinguishing papua from quad-
ridens. Actually, papua may be more closely
related to undescribed Australian species.
Genus DICRASPEDA Chaudoir
Chaudoir 1S62, Bull. Soc. Nat. Moscow 35, Part
2, p. 300.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1536 (see for partial s>n()n\in\ and addi-
tional references).
Liebke 1938, Festschrift Embrik Strand 4, pp. 43,
88.
Mdcroccntra Chaudoir 1869, Revue et Ma.uasin
Zool. (2) 21, p. 205 (new synonymy).
Liebke 1938, Festschrift Embrik Strand 4, pp. 39,
100.
Loxocdia Sloanc 1907, Dcutsihc Ent. Zeitscluitt
for 1907, pp. 179, 474.
Pliilcmonki Liebke 1938, Festschrift Enilirik
Strand 4, pp. 39, 83 (new s>n()n> ni\- ).
Diagnosis. Form as in Figiu'e 129 but
somewhat xariable. color black or metallic,
not maculate. Head: a line costa over each
eye; antennae with 3rd segments not or
not much longer than 4th segments. Pro-
tliora.x moderatelx long; pronotum chan-
neled at sides and with deep median
gr()()\('. Elytra: apici's \ariable ( sei' fol-
lowing Key to Species). Legs: tarsi not
jKibescent above (Liebke's statenunt that
tarsi ol Macroccntrd aic pubescent above
is erroneous); lib hiiid-tarsal segments
\ariab1y emarginate or lobed (see Notes,
The Carabid Beetles of New Guinea
Darlington
211
below). Secondary sexual characters: $
front tarsi narrow, narrowly 2-seriately
sqiiamulose; 6 with 1, 9 2 setae each side
last ventral segment.
Description. None required here.
Type species. Of Dicraspeda, D. brunnea
Chaudoir (see below). Of Macrocentra, M.
(juadrispinosa Chaudoir, of New Guinea.
Of Loxocara, L. quadrispinosa Sloane { =
M. quadrispinosa Chaudoir). Of PhiJemonia,
P. longiloba Liebke, of New Guinea.
Generic distribtition. Most diverse in
New Guinea; several New Guinean species
reach New Britain, etc.; species of Di-
craspeda sensu stricto (small forms with
unanned elytra) occur in Australia, and 1
(brunnea Chaudoir, below) extends to
Timor, Java, and the Philippines.
Notes. The 6 New Guinean species here
brought together in Dicraspeda are super-
ficially diverse, differing in form, presence
or absence of elytral spines, and form of 4th
hind-tarsal segments. But the differences
are all gradational (see following para-
graphs ) , different characters vary indepen-
dently, and the 6 species all share characters
given in the preceding Diagnosis. More-
over, they all inhabit understory foliage
of rain forest, and I think that they are all
probably derived from one ancestral stock
that has diversified in this habitat. Five
of the species are lowland forms and are
sympatric, occurring together at Dobodura.
The sixth species, D. ("Macrocentra")
violacea (Sloane), occurs at moderate alti-
tudes in the mountains.
The elytral apices are obliquely truncate
with sutural angles usually slightly blunted
and outer angles obtuse in D. (sensu
stricto) brunnea. In the 3 species of
"Philemonia" the sutural angles are either
slightly blunted (most individuals of longi-
Joba), variably denticulate (dubia and
some individuals of other species), or
spined (typical individuals of bispinosa);
the outer-apical angles are well formed in
these species and usually acute in longiloba,
but not spined. And in the 2 species of
"Macrocentra' (quadrispinosa and violacea),
outer-apical as well as sutural angles are
spined.
Variation of the 4th hind-tarsal segments
is noteworthy and is not correlated with
the insects' size or with form of elytral
apices. The 4th hind-tarsal segments are
shallowly emarginate (Fig. 163) in D.
(sensu stricto) brunnea; very deeply emar-
ginate with extremely long lobes (Fig. 165)
in D. ("PJiilemonia" ) longiloba, which re-
sembles brunnea in size and elytral apices;
and intermediate but variable in the other
species (other "Philemonia" and "Macro-
centra" ) .
Key to Species of Dicraspeda of New Guinea
1. Elytra without spines or with spines only at
sutural angles 2
- Elytra with spines at sutural and outer-apieal
angles 5
2. Fourth hind-tarsal segments emarginate for
c. Mi segments' length; elytra with sutural
angles slightly blunted, outer-apical angles
obtuse; length c. 5.5-6.0 mm (p. 211) - brunnea
- Fourth hind-tarsal segments more deeply
emarginate; elytra with sutural angles den-
ticulate or spined ( except in most longiloba ) ;
size larger 3
3. Fourth hind-tarsal segments very long-
lobed (Fig. 165); sutural angles usually
blunted; length c. 6.5-7.5 mm (p. 212) ..
longiloba
- Fourth hind-tarsal segments with shorter
lobes; sutural angles denticulate or spined;
size usually larger -— 4
4. Fourth hind-tarsal segments with lobes c.
^2 segments' length; sutural angles denticu-
late; length 6.5-8.0 mm (p. 212) ilulna
- Fourth hind-tarsal segments with longer
lobes; sutural angles spined or denticulate;
length 8.0-9.5 mm (p. 212) bi.s-pinosa
5. Color black; tarsi sulcate-carinate above;
length c. 11-13 mm (p. 213) quadrispinosa
- Color green-purple; tarsi not sulcate-carinate
above; length c. 11-12 mm (p. 213) -..
violacea
Dicraspeda brunnea Chaudoir
Chaudoir 1862, Bull. Soc. Nat. Moscow 35, Part 2,
p. 300.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1536 ( see for synonymy and additional refer-
ences ) .
Liebke 1938, Festschrift Embrik Strand 4, p. 89.
Description. None required here; note
212 BiiUctin Museum of Comparative Zoology. Vol. 137, No. 1
size small; pronotum punctate across base
and apex and in lateral and median grooves
but widely smooth at middle; elytral apices
unarmed; 4th hind-tarsal segments not
deeply emarginate (Fig. 163); length c.
5.5-6.0 mm.
Type. From Celebes; in Oberthiir Coll.,
Paris Mus. (not seen).
Occurrence in New Guinea. Papua: 9,
Dobodura, Mar.-July 1944 (Darlington);
1, Nonnanby Is., Wakaiuna, Sewa Bay, Jan.
1-8, 1957 (W. W. Brandt, Bishop Mus.).
N-E. N. G.: 1, Wareo, Finschhafen (Rev.
L. Wagner, S. Australian Mus.). West
N. G.: 3, Hollandia and vicinity (various
dates and collectors ) .
Notes. D. bninnea is recorded from
Australia (Queensland), Timor, Celebes,
Java, and Mindanao, and I have specimens
from Leyte and Luzon and New Britain
as well as from New Guinea.
Dicraspeda longiloba (Liebke)
Liehke 1938, Festschrift Einbrik Strand 4, p. 83
( F/i ilemonia ) .
Description. None required here; note
elytra with sutural angles blunt or at most
minutely denticulate; 4th hind-tarsal seg-
ments very long-lobed ( Fig. 165 ) ; length c.
6.5-7.5 mm.
Type. From N-E. N. G. ("Deutsch-Neu-
Guinea"); in Liebke Coll., present location
unknown (not seen).
Occurrence in New Guinea. N-E. N. G.:
the type. Papua: 5, Dobodura, Mar.-July
1944 (Darlington).
Notes. I have a specimen also from Cape
Gloucester, New Britain.
Dicraspeda dubia (Gestro)
(Jestro 1879, Ann. Mus. Civ. C.cnoa 14, p. 558
(Odacanlha) .
Liebke 1938, Festsebrift i;ii.l)rik Strand 4, p. 83,
fJLC. f:)7 {Fluh'Dionid) .
Descri])tion. None required here; note
elytra with sutural angles variably denticu-
late but not si)ined; 4th hiud-taisal seg-
ments rather short-lobcd but somewhat
varia])le; length c. 6.5-8.0 mm.
Type. From Fly R., presumably Papua;
in Genoa Mus. (not seen).
Occurrence in New Guinea. Papua: 3,
Dobodura, Mar.-Julv 1944 (Darlington);
5, Kokoda, 1200, 1300 ft.. May, Aug., Sept.,
Oct. 1933 (Cheesman). N-E. N. G.: 1,
Aitape, Aug. 1944 (Darlington); 1, Wau,
Morobe Dist., 1200 m, Aug. 18, 1961 ( Sed-
lacek), in light trap. West N. G.: 1, Waris,
S. of Hollandia, 450-500 m, Aug. 16-23,
1959 (T. C. Maa, Bishop Mus.); 1, Xabire,
S. Geelvink Bay, 0-30 m, July 2-9, 1962
(Gressitt); 1, Waigeu Is., Camp 1, Mt. Nok,
2500 ft. ( c. 760 m ), May 1938 ( Cheesman ) .
Notes. I found this or a closely related
species also at Bamaga, near the tip of
Cape York, Australia.
Dicraspeda bispinosa n. sp.
Description. With characters of genus;
form as in Figure 129; brownish black, legs
dark, antennae and mouthparts paler
brown; moderately shining, reticulate mi-
crosculpture indistinct on head and pro-
notum, isodiametric or slightK' transverse
on elytra. Head 1.17 and 1.17 width pro-
thorax; front irregular!)' impressed anteri-
orly, impunctate; mentum w ith long narrow
tooth; ligula 4-setose. Prothorax elongate-
subquadrate with sides swollen below mar-
gins; width length 0.99 and 0.98; base '
apex 1.19 and 1.12; pronotum strongly con-
vex, narrowly channeled each side near
margin and with well impressed middle
groove; surface punctate chiefly across
base, slightly at apex. Elytra: width elytra/
prothorax 2.08 and 2.05; apices obliquely
sinuate-truncate with sutural angles spiued
or denticulate, outer-apical angles obtuse,
and apical margin in part minuteh' d(>nticu-
late; striae entire, punctate; 3rd inter\als
3-punctate, the posterior puncture near
apex. Inner n'i)iij.s full. /.ri,'.v normal; tarsi
not suleate and not pubescent aboM'; 4th
hind-taisal segments long-lobed (Fig. 164).
Secondary sexual cJiaracters as of genus;
last Ncntral segment with small notch at
apex in both sexes. Mcdsiircinciils: length
The Carabid Beetles of New Guinea • Darlington 213
(including spines) c. 8.0-9.5; width 2.8-
3.3 mm.
Types. Holotype $ (M.C.Z., Type No.
31,510) and 15 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
additional paratypes as follows. Papua: 2,
Kokoda (Cheesman); 2, Mt. Lamington,
1300-1500 ft. (c. 400-460 m) (C. T.
McNamara, S. Australian Mus.). N-E.
N. G.: 4, Wau, Morobe Dist., 1150, 1200,
1300 m, dates in Jan., Feb., Oct., 1961, 1963
(Sedlacek); 1, Finschhafen, Huon Pen., 80-
200 m, Apr. 13, 1963 (Sedlacek).
Additional material. N-E. N. G.: 4,
Finschhafen, 80 m, Apr. 16, 1963 (Sedlacek);
1, same locality, 80-200 m, Apr. 13, 1963
(Sedlacek); 3, Pindiu, Huon Pen., dates
in Apr. 1963 (Sedlacek).
Mea.sured specimens. The 6 paratype and
9 holotype from Dobodura.
Notes. This new species would go in
Philemonia in Liebke's classification. The
specimens listed under Additional material
have the sutural angles of the elytra den-
ticulate rather than spined, but I think they
are referable to bispinosa. Note that both
spined and denticulate forms have been
found at Finschhafen.
Dicraspeda quadrispinosa (Chaudoir)
Chaudoir 1869, Revue et Magasin Zool. (2) 21, p.
206 ( Macrocentra ) .
Sloane 1907, Deutsche Ent. Zeitschrift lor 1907, p.
474 (Macrocentra).
Liebke 1938, Festschrift Embrik Strand 4, p. 100
( Macrocentra ) .
Louwerens 1956, Trculiia 23, p. 223 (Moluccas)
(Macrocentra).
Loxflcara quadrispinosa Sloane 1907, Deutsche Ent.
Zeitschrift for 1907, p. 180.
Description. None required here; known
among New Guinean Colliurini by size
large; color plain black; elytra spined at
sutural and outer-apical angles; tarsi sul-
cate-carinate above; length (including
spines) c. 11-13 mm (rarely shghtly smaller
or larger).
Types. Of quadrispinosa Chaudoir, from
Dorey, West N. G. ( \\'allace ) ; in Oberthiir
Coll., Paris Mus. Of quadri.spinosa Sloane,
from Simbang, N-E. N. G.; "returned to
Dr. Horn (for Bennigsen's collection)"
(not seen).
Occurrence in New Guinea. Common:
227 specimens from localities on New
Guinea and Normanby, Ferguson, Wood-
lark, Rossel, Sudest, and Waigeu Is.; ap-
parently confined to low altitudes, up to
700, 750, 800, and 975 m at different
localities, but none found higher; cnmmnn
at Dobodura.
Notes. This characteristic New Guinean
carabid has been foimd also in the Moluc-
cas, New Britain, and the Solomons, but
not Australia.
Dicraspeda violacea (Sloane)
Sloane 1907, Deutsche Ent. Zeitschrift for 1907,
pp. 181, 474 (Macrocentra).
Liebke 1938, Festschrift Embrik Strand 4, p. 100
(Macrocentra).
hahiJis Sloane 1907, Deutsche Ent. Zeitschrift for
1907, p. 181 (name used in error for violacea).
Description. None required here; similar
to preceding (iptadrispinosa) but head and
pronotum greenish or bluish, elytra purple;
tarsi not sulcate-carinate above; length c.
11-12 mm.
Type. From Sattelberg, N-E. N. G.; "re-
turned to Dr. Horn (for Bennigsen's col-
lection)" (not seen).
Occurrence in New Guinea. N-E. N. G.:
8, Wau, Morobe Dist., 1200, 1300 m, Jan.,
Mar., Apr., June, Sept., Nov., 1961-1963
(Sedlaceks); 1, Eliptamin Vy., 1665-2530
m, June 23-30, 1959 (W. W. Brandt, Bishop
Mus.); 1, Finisterre Rge., Saidor, Kiambavi
Village, July 22-29, 1958 (\^^ W. Brandt,
Bishop Mus.); 1, W^areo, Finschhafen (Rev.
L. Wagner, S. Australian Mus.); 1, Goroka,
E. Highlands, 5200 ft. (c. 1600 m), J. H.
Barrett, Dept. Agr. Port Moresby). West
N. G.: 3, Rattan Camp, Snow Mts., 1150,
1200 m, Feb.-Mar. 19.39 (Toxopeus); 5,
Fac Fac, Vogelkop, 100-700 m, June 9,
1959 (Gressitt and T. C. Maa, Bishop
Mus.), in light trap; 1, Mt. Baduri, Japen
Is., 1000 ft., Aug. 1938 (Cheesman).
Notes. D. violacea apparently replaces
quadrispinosa above 1000 m altitude in
214
Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
New Guinea, Init the two species overlap
below 1000 m. D. violacea occurs also in
New Britain (Gaulim, Gazelle Pen., 130 m,
Nov. 28, 1962, Sedlacek).
Genus LACHNOTHORAX Motschulsky
Motsthulsky 1862, fitude Ent. 11, p. 48.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1542 (see for additional references).
Liebke 1938, Festschrift Embrik Strand 4, p. 103.
Jeannel 1948, Coleop. Carabiques de la Region
Malgache, Part 2, p. 7.56.
Diaii,nos-is. See Key to Genera of Col-
liurini of New Guinea; this is the only con-
spicuously pubescent colliurine in New
Guinea.
Description. None required here.
Type species. L. higuttatus Motschulsky,
of India.
Generic disfrihtition. Africa and Mada-
gascar; SE. Asia to New Guinea.
Notes. The few species of Lachnothorax
are all very much alike. They are probably
ground-living, and I suspect that they occur
by running water.
Lachnothorax. tokkia Gestro
Gestro 1875, Ann. Mns. Civ. Cenoa 7, p. 856.
Csiki 1932, Coleop. ('at., Carabidae, Harpalinae 7,
p. 1542 (see for synonymy and additional refer-
ences ) .
Liebke 1938, Festschrift Embrik Strand 4, p. 101.
Jedlicka 1963, Ent. Abliandlnngen 28, p. 504.
Description. None required here; note
form, color black with pale spot before
apex each elytron, and consjiicuous pubes-
cence; length c. 5.0-5.5 mm.
Type. From Kandari, SE. Celebes; in
Genoa Mus. (not seen).
Occurrence in New Guinea. N-E. N. (i.:
7, Stephansort, Astrolabe Bay, 1900 (Biro).
Notes. Lachnothorax tokkia has been r(^-
corded previously from the Malay Pen..
Sumatra, Java, and Celebes, and a ver\
closely relat(>d species {hi<iuttata Motschul-
sky) occurs in India and Geylon.
Genus EUDALIA Castelnau
Caslelnan 1867, Notes on Australian Coleop., p. 16.
Sloane 1917, Proc. Linnean Soc. New South Wales
42, pp. 415, 417-422 (with key to Australian
species ) .
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1542 (see for synonymy and additional refer-
ences ) .
Liebke 1938, Festschrift Embrik Strand 4, pp. 44,
105.
Diapiosis. See Key to Genera.
Description. None required here.
Type species. Odacantha latipennis Mac-
leay, of Australia.
Generic distribution. Australia (c. 6
species ) and New Guinea ( 1 species,
doubtfully assigned to this genus).
Notes. The species described below as
Eudalia anomala has entire lateral pro-
thoracic margins and therefore runs to
Eudalia in Liebke's key, but if the margins
were obsolete, it would run to Andrewesia,
to which it may be more closely related.
{Andrewesia ohcsa (Andrewes) ranges
from the Malay Pen. to the Moluccas.)
Generic characters and limits in this group
(as in so many others!) need revision.
Eudalia anomala n. sp.
Description. Form as in Figure 130;
black, elytra faintly aeneous and tipped
with yellow, legs testaceous, antennae
brown paler at base; h(>ad and pronotum
shining and without reticulate microsculp-
ture, elytra duller with isodiametric meshes.
Head 1.18 and 1.17 width prothorax;
strongly constricted at neck; antennae with
segments 3 and 4 sube(jual, pubescent from
4th segmcMits; mandibles moderate^ in knigth
and curvature; front eon\ex, irregularly
impressed anteriorK', impunetate; mentuin
with moderate tooth; liguJa broad, with 2
long and 2 shorter setae; palpi slender, not
pubescent. Protliorax suborbicular except
parallel at base; width length 0.89 and
0.90; base apex 1.25 and 1.26 (sides of pro-
thorax curve into condyle of neck at apex);
lateral margins narrow but (Mitire, with a
seta-bearing pimcture inside margin (on
disc) before middle; disc \('ry convex,
strongly transverseh- impressed at base;
middle line slightly impressed; surface
The Carabid Beetles of New Guinea • Daiiinaiott
215
punctate across base, impunctate or nearly
so elsewhere. Elytra ample; width elytra,
prothorax 2.15 and 2.16; apices obliquely
truncate with outer angles obtuse and
inner angles acute-blunted; striae entire,
punctate; 3rd intervals with 4 or 5 seta-
bearing punctures including 1 near base.
Inner winp,s full. Legs moderate; tarsi not
pubescent above and not sulcate; 4th hind-
tarsal segments emarginate but not lobed.
Secondary sexual characters: i front tarsi
narrow, 2-seriately squamulose; last ventral
slightly emarginate at apex in 6 , not in $ ,
with 1 seta each side in i , 2 in 9 . Mea-
surements: length c. 7.0; width 2.5 mm.
Types. Holotype $ (A.M.N.H.) and 1
i paratype (M.C.Z., Type No. 31,511)
from Menapi, Cape Vogel Pen., Papua,
0-30 m, "No. 1," Aug. 8-11, 1953 (Geoffrey
M. Tate); and 1 9 i^aratype from Wasian,
Vogelkop, West N. G., Sept. 1939 (Wind,
M.C.Z.).
Measured specimens. The i holotype and
9 paratype.
Notes. For possible relationships of this
species, see under genus. This species is
smaller and much less punctate and less
roughened above than any typical (Aus-
tralian) Eudalia known to me, and the
New Guinean species has the outer-apical
elytral angles more angulate.
DOBODURA n. gen.
Diagnosis. See Key to Genera of Col-
liurini of Netv Guinea.
Description. Form ( Fig. 131 ) c. as in
some CoUiuris. Head without supraocular
costae; mandibles long, slender, weakly
arcuate; antennae very long, 3rd segments
c. V4 longer than 4th segments, 1st segments
with 1 long seta near apex; mentum with
triangular tooth; ligula rounded, with 2 long
setae at apex and 2 shorter setae laterally;
paraglossae small, membranous; palpi slen-
der, not pubescent. Prothorax: lateral mar-
gins entire; median impressed line fine;
base deeply transversely channeled with
transverse ridge behind channel. Elytra
spined. Inner icings full. Legs slender;
tarsi not pubescent above, not sulcate
above; 4th hind-tarsal segments moderately
emarginate, emargination c. '/•; length of
segment; claws not toothed. Secondary
sexual characters: 6 front tarsi scarcely
dilated, 3 segments narrowly 2-seriately
squamulose; last ventral segment of $
weakly, of 9 subcircularly emarginate,
with 1 seta each side in i , 2 in 9 .
Type species. D. armata (below).
Generic distribution. The single species
is known only from New Guinea.
Notes. This striking new genus may be
related to Eudalia but differs in form,
longer and less arcuate mandibles, much
longer antennae with relatively longer 3rd
segments, and presence of elytral spines.
The position of the principal spines, c.
opposite the ends of the 4th intervals
rather than at the sutural or outer-apical
angles, is unusual in this tribe.
Dobodura armata n. sp.
Description. With characters of genus;
form as in Figure 131; black above and
below, appendages testaceous except femora
dark on inner sides; shining, reticulate
microsculpture absent or indistinct on most
of upper surface, present and c. isodiametric
on elytra posteriorly. Head 1.07 and 1.06
width prothorax; front evenly convex except
slightly impressed anteriorly, impunctate;
neck slightly constricted. Prothorax suboval,
swollen at sides below margins; width
length 0.92 and 0.91; base/apex 1.14 and
1.15; margins each with seta-bearing punc-
ture c. -o from apex; disc c. evenly convex,
impunctate. Elytra: width elytra prothorax
2.00 and 2.09; apices with sutural and outer
angles denticulate or short-spined and with
long spines c. opposite ends 4th intervals;
striae entire, formed by lines of fine punc-
tures; 3rd intervals with 3 well spaced
seta-bearing punctures. Secondary sexual
characters as of genus; S copulatory organs
as in Figure 180. Measurements (types);
length (including spines) c. 10.5-11.5;
width 3.3-3.5 mm.
216 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Types. Holotype S (M.C.Z., Type No.
31,512) and 5 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
2 paratvpes, Mt. Hansemann, Astrolal^e
Bay, N-E. N. G. (Biro).
Additional material. West N. G.: 1 $ ,
mountain slope above Bernhard Camp, 100
m, Apr. 8, 19.39 (Toxopeus).
Measured specimens. The i holotype and
1 9 paratype from Dobodura.
Notes. My specimens were taken among
spray-drenched stones beside small torrents
in rain forest.
The specimen from Bernhard Camp has
the strial punctures of the elytra coarser
than in the types and the tip of the aedeagus
slightly different. Additional material may
sho\\' it to represent a distinguishable sub-
species.
Tribe DRYPTiNI
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1548 (see for synonymy and additional refer-
ences ) .
Jeannel 1949, Coleop. Carabicjues de le Region
Malgache, Part 3, p. 1063.
Jedlicka 1963, Ent. Abhandlunyen 2S, p. 4S1.
Halm 1967, Fanna Jap<"iica, Caral)idae, Trun-
eatipennes Group, p. 266.
Drypfiddc Jeannel 1942, Faune de France, Coleop.
Carabicjues, Part 2, p. 1098.
Dryptimie Basilewsky 19.53, Exploration Pare Na-
tional rUpemba, Ease. 10, Carabidae, p. 228.
Members of this small but widely distrib-
uted tribe (represented in New Cuinea
by only 2 genera) are easily recognized by
characteristic form (P'ig. 132); pubescent
surface; antennae with very long 1st and
very short 2nd segments; and elytra with-
out raised outer margins. The New Guinean
species live chiefly in grass, 1 think. They
are winged and probably diurnal.
Key to (;kni:ha of Dkyi'tim ok New Guinea
I. CHaws simple (p. 216) _ /);■(//)/«
- Claws pectinate (p. 218) Dcscra
Genus DRYPTA Latreille
Latreille 1796, Precis Caracteres Generiqnes In-
sectcs, p. 75.
Csiki 1932, Coleop. (]at., Carabidae, Harpalinae 7,
p. 1.548 (see lor additional reterences).
Andrewes 1936, Proc. R. Ent. Soc. London (B)
5, p. 134 (key to "Indian" species).
See also references under tribe.
Diaiinosis. See characters given for tribe
and in preceding Key to Genera.
Description. None required here.
Type species. Carahus dentatus Rossi, of
Europe, etc.
Generic distribution. Tropical and warm
temperate regions of the Old World; 1
species listed from Brazil.
Notes. The Oriental-Australian species of
Drypta are much alike, differing chiefly in
proportions and color.
Key to Species of Drypta of New Guinea
1. Head, prothorax, and elytra blue-green (p.
216 ) fkipua
- Head and prothorax red; elytra brown,
black, blue-black, or striped 2
2. Elytra broadly longitudinally striped with
red (p. 217) _ inaatcrsi
- Elytra not striped 3
3. Less slender ( prothoraeic width dength r.
0.78 or more); femora dark (p. 217) fumi^ata
- More slender ( prothoraeic width length c.
0.7.5 or less); femora pale (p. 217) siilcicollis
Drypta popua n. sp.
Description. With characters of genus;
form as in Figure 132, c. average in genus;
greenish blue, appendages yellow with
apices of femora and of 1st antennal seg-
ments narrowly darker; entire upper sur-
face closely punctate. Head 1.09 width
prothorax; eyes moderate, genae convex.
Protliora.x subcylindric; width length 0.76;
base apex 1.08; lateral margins indistinct;
middle line poorly defined. Elytra: width
elytra prothorax 2.09; apices obliciueK sub-
truncate with outer angles obtuse-blunted
and sutmal angles c. right; striae impressed,
coarsely ver\ closely punctate; inten^als
more finely, less closely punctate. Le^s:
tarsi not sulcate abo\(>; 4th hiud-tarsal seg-
ments long-lobed; claws cm\(>d, not pecti-
nate, each w ith obtuse angulation of innc^r
edge near base but with no trace of teeth.
Secondary sexual characters not determined
{a unknown). Measurements: length 8.5;
w idth 2.7 mm.
Type. Holotype 9 (M.C.Z., Type No.
The Carabid Beetles of New Guinea
Darlington
217
31,513) from Lae, N-E. N. G., Oct. 1944
(Darlington); the type is unique.
Notes. I am not sure of the relationships
of this unexpected species. In Andrewes'
( 1936 ) key to "Indian" species of Drijpta
(see reference under genus), papua runs
to couplet LS ( 19 ) but fits neither species
there named, having a relatively narrower
head and broader prothorax than aeiheria
Andrewes (of Assam) and more closely
punctate elytral intervals than cijanopa An-
drewes (of Bengal). Drypta papua does
not resemble any Australian species of the
genus. It does superficially resemble
Desera cleg,ans Sloane (below) but is
smaller, with relatively broader prothorax
and obtuse rather than acute outer-apical
elytral angles, and of course with simple
rather than pectinate tarsal claws.
Drypta mastersi Macleay
Macleay 1871, Trans. Ent. Soc. New South Wales
2, p. 82.
Chaudoir 1877, Bull. Soc. Nat. Moscow 52, Part 1,
p. 257.
Description. None rec^uired here; note
elytra striped with red; length (of New
Guinean specimen) c. S.5 mm.
Type. From Gayndah, South Queensland,
Australia; presumably in Macleay Mus.,
Sydney (not seen).
Occurrence in New Guinea. Papua: 1,
Rouku, Morehead R., W. Papua (opposite
the tip of Cape York), Apr. 1962 (W. W.
Brandt, C.S.I.R.O.).
Notes. In Australia, mastersi ranges north
at least to mid-peninsular Cape York. I
do not know whether it is really distinct
from Drypta australis Dejean of more-south-
ern Australia.
Drypta fumigata Putzeys
Putzeys 1875, Ann. Mus. Civ. Genoa 7, p. 720.
Chaudoir 1877, Bull. Soc. Nat. Moscow 52, Part
1, p. 258.
Description. None required here; length
13.5-15.0 mm.
Type. From Andai, Papua, Aug. 1872
(Beccari and D'Albertis); in Genoa Mus.
(not seen).
Occurrence in New Guinea. Papua: 3,
Dobodura, Mar.-July 1944 (Darlington); 5,
Kiunga, Fly R., dates from Julv 23 to
Sept. 25, 1957 (W. W. Brandt,' Bishop
Mus.); 1, Owen Stanley Rge., Goilala,
Rome, 1950 m, Apr. 1-15, 1958 (W. W.
Brandt, Bishop Mus.); 1, Popondetta, N.
Dist., Jan. 29, 1965 (R. Hornabrook). N-E.
N. G.: 1, Erima, Astrolabe Bay, 1896
(Biro); 1, Chimbu Vv., Bismarck Rge.,
5000-7500 ft. (c. 1500-2300 m), Oct. 1944
(Darlington); 1, Aiyura, E. Highlands,
5600 ft. (c. 1700 m), "9.10.1960" (J. H.
Barrett, Dept. Agr. Port Moresby), at light;
2, Okapa, June 23, 1965 (R. Hornabrook).
West N. G.: 2, Hollandia, Apr., May 1945
(R. Malkin, U.S.N.M.); 32, Sansapor, Aug.
1944 (Darlington).
Notes. This species is presumably of
Oriental origin, but I cannot determine to
which Oriental species it is most closely
related.
Drypta suicicollis Putzeys
Putzeys 1875, Ann. Mus. Civ. Genoa 7, p. 721.
Chaudoir 1877, Bull. Soc. Nat. Moscow 52, Part 1,
p. 258.
Description. None required here; length
c. 10.5-11.5 mm.
Type. From Andai, Papua, Aug. 1872
( Reccari and D'Albertis ) ( note locality
same as for type of fumigata); in Genoa
Mus. (not seen).
Occurrence in New Guinea. Papua: 1,
Dobodura, Mar.-Julv 1944 (Darlington);
3, Kiunga, Fly R., Aug. 1-3, 14-17, 1957
(W. W. Rrandt, Bishop Mus.). N-E. N. G.:
1, Erima, Astrolabe Bay, 1897 (Biro); 1,
Stephansort, Astrolabe Bay, 1897 (Biro).
West N. G.: 1, Hollandia-Binnen, 25 m,
Oct. 16, 1957 (Gressitt); 1, Humboldt Bay
Dist., 1934 (British Mus.); 3, Tor R.
(mouth), 4 km E. of Hoi Maffen, July 2,
1959 (T. C. Maa, Rishop Mus.). at light;
2, Wasian, Vogelkop, Sept. 1939 (Wind,
M.C.Z.).
Notes. This species too is probably de-
rived from an Oriental (not Australian)
stock, but I do not know its exact relation-
ships.
218 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Genus DESERA Hope
Hope 1831, Zoological Mistellany, p. 21.
Csiki 1932, Coleop. Cat., Carabidae, Haipalinae 7,
p. 1553 (see for additional references).
Andrewes 1936, Proc. R. Ent. Soc. London ( B )
5, p. 136 ( kev to "Indian" species).
1939, Ann. Mag. Nat. Hist. (11) 3, p. 133.
Dendrocelhis Schmidt-Goebel 1846, Faunula
Coleop. Birmaniae, p. 24.
Diafinosis. Characters as of Drijpta, ex-
cept cla\\'s pectinate.
Description. None required here.
Ty))e .species. Dcsera nepalen.si.s Hope,
of SE. Asia (see following Note.s).
Generic distribution. SE. Asia to Aus-
tralia; Africa.
Note.s. Dcsera differs from Drypto ap-
parently only in having pectinate tarsal
claws. A modern revision of the species is
needed to show whether both genera are
really nionophyletic and distinct.
Andrewes ( 1939 ) outlines the history of
the name Dcsera. It was used by Hope
( 1S31 ) in combination with the valid de-
scription of a new species ( )icpalcnsis,
which is therefore the type species), and
the combined description iricludes refer-
ence to the pectinate tarsal claws. This
use validates Dcsera Hope 1831 under
Article 16(a) (VI) of the 1964 edition of
the International Code of Zoological No-
menclature.
A single, common species ol this genus
occurs in New Cuinea.
Desera elegans (Sloane)
Sloanc 1907, Dcutsilic Knt. Zcitsclirilt lor 1907,
p. 473 ( Dcii(lrocclliis).
Andrewes 1927, Ann. Mag. Nat. Hist. (9) 19, p.
110.
Description ( for recognition only). With
characters of tribe and genus; form slender;
green (sometimes l^lue-green or l)roiv/e-
green), antennae red with 1st segment dark
at apex, legs red with knees usualK' darker
(legs and antennae sometimes more ex-
tensively dark); knigth c. 9.5-10.5 mm.
Ty])C. Froiu Haining Berge, Cazelle Pen.,
New Itritaiii; in Deutsche Ent. Instituti',
Berhn-Dahlem (Andrewes 1927) (not seen).
Occurrence in New Guinea. Common:
197 specimens from many localities widely
scattered over New Guinea, from sea level
to c. 1700 m altitude; occurs at Dobodura
and \\'au.
Notes. Desera clciians of New Guinea,
New Britain, and New Ireland is similar
to genicidata King (SE. Asia to the Moluc-
cas) on one side and to smaraiidula
Chaudoir (Australia) on the other. In fact
a single individual from Rouku, Morehead
R., West Papua, Apr. 1962 ( W. W. Brandt,
C.S.I.R.O. ) looks more like the Australian
smarag,dula than like the New Guinean
elegans. Relationships (or identities?) of
these and other similar species in the
whole Asiatic-Australian area need further
study.
Tribe ZUPHIINI
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1562 (see for synonymy and additional refer-
ences ) .
Habn 1967, Fanna Japonica, Carabidae, Tnm-
catipennes Gronp, p. 253.
Ztiphiidae Jeannel 1942, Fanne de France, Coleop.
Carabiqnes Part 2, p. 1091.
ZitpJiiitac Jeannel 1949, Coleop. Carabiciues de la
Region Malgaelie, Part 3, p. 1047.
Ziiphiiuae Basilewsky 1953, Fxploration Pare Na-
tional I'l'pcniba I'^ase. 10, p. 224.
This is another small but wideK distrib-
uted tribe. Its characters and taxonomic
liiuits need not be discussed here. It is
represented in New Cuinea by 2 easily
recognized genera and a total ol 6 known
species.
The members of the tribe \\\r in wt't
places, often among dead lea\cs on tlu'
ground {Yjipliiuni) or in gniss and vegeta-
tion growing in water (Planctes). Most
sjieeies, including all those in New Cluinea,
are winged.
K\:\ ro Ckmvuv oi /.ii'iium oi Niw Guinea
1. Head Mil)triangiilar, \cr\ wide at base: eKtra
not eostate (p. 219) Zujiliitnii
- Head normal; elytra wilii Mian> line (.ostac
(p. 220) '- - I'laiictcs
The Carabid Beetles of New Guinea • Darlington 219
Genus ZUPHIUM Latreille
Latreille 1806, Genera Crustaceorum et Iiisectonim
1, p. 198.
Csiki 1932, Coleop. Cat., Caiabidae, Harpalinae 7,
p. 1562 (see for synonymy and additional refer-
ences ) .
Jedlicka 1963, Ent. Abhandlungen 28, p. 477.
See also references under tribe.
Diagnosis. Form (Fig. 133) diagnostic;
and see preceding Key to Genera.
Description. None required here.
Type species. Carahus oJens Rossi, of
Europe, etc.
Generic distribution. All tropical and
some temperate regions of the world.
Notes. The various Oriental and Austra-
lian species of Zuphium are not well under-
stood. For example, the 7 listed Australian
species were all described between 1867
and ISScS, most of them from single speci-
mens or single localities, and they have
never been revised. The real relationships
of the 2 New Guinean species are therefore
doubtful, although I have made some com-
parisons.
Besides the 2 species recorded from New
Guinea below, I have seen (British Mus.)
1 specimen of Zuphium celehense Chaudoir
labeled as from Dory, presumably collected
by Wallace. I think this specimen is prob-
ably really from Celebes ( see Part I of my
"Garabid Beetles of New Guinea," pp. 330-
331 ) , and I see no reason to list the species
from New Guinea even tentatively.
Key to Species of Zuphium of Ne\v Guinea
1. Large (c. 8.3 mm); color piceous (p.
219 ) thouzeti
- Small (c. 3.5 mm); color brown (p. 219)
simiinu
Zuphium thouzeti Castelnau
Castelnau 1867, Notes on Australian Coleop., p. 17.
1868, Trans. R. Soc. Victoria 8, p. 103.
Description. None required here. Note
size large; color dark, not spotted; length
(of New Guinean specimen) c. 8.3 mm.
Types. From Rockhampton, Queensland,
Australia; present location of type un-
known (not seen).
Occurrence in New Guinea. Papua: 1
9, Port Moresby, Feb.-May 1943 {W. B.
Jones, U.S.N.M.).
Notes. Besides the types from Rock-
hampton, Gastelnau had a specimen from
Port Denison (near Bowen) farther north,
and I have specimens (identified from de-
scription) from W. of Ravenshoe and N. of
Mareeba still farther north in Queensland.
The Port Moresby specimen agrees rea-
sonably well with my Australian ones ex-
cept that the color of the legs varies.
Zuphium sinuum n. sp.
Description. Form as in Figure 133, very
small; brown, head slightly darker, append-
ages and lower surface paler; dull, entire
upper surface densely microreticulate or
roughened. Head 0.92 and 0.94 width pro-
thorax; antennae short, middle segments c.
2x long as wide; surface densely micro-
reticulate, moderately punctulate. Pro-
thorax: width/length 1.13 and 1.10; base/
apex 0.88 and 0.81; posterior angles right-
acute and not quite basal ( base very briefly
subpedunculate); surface closely roughened-
punctate. Elytra: width elytra prothorax
1.65 and 1.70; apices sinuate at middle of
width, lobed between sinuations and suture;
surface roughened, striae indicated but not
well defined. Secondary sexual characters:
i front tarsi slightly dilated, 3 segments
with soles of dense short squamae; c^ with
1, 9 2 setae each side last ventral segment.
Measurements: length c. 3.5; width 1.3-
1.4 mm.
Types. Holotype 9 (M.G.Z., Type No.
31,514) from Aitape, N-E. N. G., Aug. 1944
(Darlington); and 1 6 paratype, Kota
Nika, Res. Hollandia, West N. G., Jan. 31,
1956 (R. T. Simon Thomas, Louwerens
GolL).
Measured specimens. The S paratype and
9 holotype, in this order.
Notes. This species or a close relative
occurs also at Cape Gloucester, New Brit-
ain. Small size, color, dull surface, and
sinuate elytral apices distinguish sinuum
from other comparable species including
220 BuUc'tin Mu.scin)i of Comparative Zoolog,!), Vol. 137, No. 1
cclc'])cnsc Chaudoir (see under genus), in
whieh the elytral apices are scarcely sinuate.
Z. inconspicuum Schmidt-Goebel of Burma,
etc., has strongly sinuate elytral apices but
is much more shining than .sintiinn.
Genus PLANETES Macleay
Macleay 1825, Aniuilosa Javanica, p. 28.
Csiki 1932, Coleop. Cat., Caiahidae, Harpalinat- 7,
p. 1567 (see for syiuiiiyiiiy and aclclitional refer-
ences).
Jcdiieka 1963, iMit. Ahhandlnnuen 28, p. 464.
Di(ii!,n()si.s. Form c. as in Figure 134;
elytra each with more than 20 line longitu-
dinal costae.
Description. None required here.
Type species. P. himacuJatus Macleay,
of Java, etc.
Generic distrihution. SE. Asia including
Oylon and Japan to northern Australia;
Africa.
Notes. Most and most diverse species of
Planetes occin- in the Oriental Region. Four
species, all rather small and imspotted, are
known in New Guinea. And only I or 2
species, the same as or close to New
Guinean lorms, ha\t' l)een found in Aus-
tralia. This geographic pattern suggests
dispersal from Asia to Australia.
The species of this genus that I have
collected live in swamps and beside stand-
ing water. The\' are \\inged and often fly to
light.
In the present work I have not distin-
guished Uetero^Jossa Nietner from Planetes,
although the two probably are distinct (see
Ihibu, 1967, reference cited under tribe).
Ki;v K) SiM'CiKs OF Planetes ok Nkw CIuinka
1. Smaller, lenijtli 6.0—7.5 mm; see also l^c-
■srrii)li()u (p. 220) scccnuiutus
— Usually larger; // leniilli under 8 mm, 19th
( posthiimeral ) elytral intervals specially eon-
sjiieuous 2
2. Klytra with an interval (the 19th, near
humeri) more conspieuons than others at
base; see also Description (p. 220) Iniincidlis
— Elytra with no single interval more eoii-
spieuous than others at base 3
.3. Prothorax subeordate, with sides not oi'
weakh' sinuate; pronolum more evcuK pmie-
tate, the punctures rather coarse and of c.
uniform size (p. 221) . aiistidlis
- I'rothora.x strongly cordate, with sides stronglj'
sinuate; pronotum less evenly punctate, with
coarsi' and fine punctures mixed (p. 221)
cordens
Planetes secernendus OberthUr
Oberthur 1883, Notes Leyden Mus. 5, p. 217.
Description ( for recognition only ) . Form
small; sparsely inconspicuously pubescent;
piceous, not spotted, appendages brownish
testaceous; prothorax cordate or subeordate,
pronotum unevenly punctate, the punctures
varying in size and usually sparser near
middle of pronotum; elytra each with more
than 20 fine costae, the costae subequal
except 1st, 4th, 7th, etc. usualh' sliii^lithj
wider or more prominent toward base and
apex, but 1 9th costa not specially con-
spicuous at base; length r. 6.0-7.5 mm.
Types. From Sumatra; in Oberthiir
Coll., Paris Mus. (not seen).
Occurrence in New Guinea. Sixty-three
specimens from localities (including Dobo-
dura) scattered over most of the length of
New Guinea; at low altitudes only, none
above 500 in.
Notes. P. secernendus is now known
from the Malay Pen. (British Mus.),
Sumatra, Java (British Mus.), Borneo.
Leyte and Luzon in the Philippines
(M.C.Z.), New Guinea, and New Britain
(M.C.Z. ). Geographic \;uiation probably
occurs but is confused 1)\ iiidi\idual \aria-
tion especialK (in New (Guinea) in form
and pimctation ol prothorax. See also Addi-
tional material and Notes under /'. humer-
al is. below.
Plonefes humeralis n. sp.
nescri))lion. With characters ol genus;
ionn as in preceding species (seeenioidus),
reddish biown, sometimes darkcM', append-
ages slightK i^aler; head and pronotum
shining betwcHMi piuictures, eKlia duller.
lle(ul 0.77 and 0.7(S w idth prothorax, \\ eakly
impressed across base; I rout comex, slightly
impressed anleriorly, slightK irregularly
The Carabid Beetles of New Guinea • Darlington
221
finely punctate. Protlwrax narrowly sub-
cordate; width length 1.25 and 1.26; base
apex 1.06 and 1.05; sides broadly arcuate
anteriorly, moderately sinuate posteriorly,
with moderate margins, each with usual 2
setae; disc slightly convex, with middle line
well impressed but lateral longitudinal im-
pressions vague; baso-lateral impressions
moderate, roughened; surface of disc mod-
erately punctate with punctures of mixed
sizes, more closely punctate across base and
apex. Elytra subparallel; width elytra pro-
thorax 1.35 and 1.37; apices obliciuely trun-
cate, outer angles broadly rounded, sutural
angles scarcely blunted; each elytron with
more than 20 fine costae, the 1st, 4th, 7th,
etc., slightly more prominent than others
and the 19th specially prominent (but still
fine) at base. Secomkinj sexual characters:
5 front tarsi slightly dilated, 3 segments 2-
seriately squamulose; 1 principal seta each
side last ventral segment in both sexes.
Measurements (of types): length 7.3-8.3;
width 2.5-2.9 mm.
Types. Holotype c^ (Bishop Mus.) and
1 £ paratype (M.C.Z., Type No. 31,515)
from Eliptamin Vy., N-E. N. G., 1200-1350
m, July 16-31, 1959 (W. W. Brandt); 1
6 paratype, Torricelli Mts., Mokai Village,
N-E. N. G., 750 m, Jan. 1-23, 1959 (W. W.
Brandt, Bishop Mus.); 1 $ paratype, Mt.
Dayman, Maneau Rge., Papua, 700 m, "N.
Slope No. 6," July 13-20, 1953 (Geoffrey
M. Tate, A.M.N.H.).
Additiorml material. N-E. N. G.: 3,
Krisa, Vanimo, Apr. 1939 (Cheesman).
West N. G.: 1, Dojo, 2nd Strip, Res. Hol-
landia, Apr. 15, 1957 (R. T. Simon Thomas,
Louwerens Coll.).
Measured specimens. The 6 holotype and
i paratype from Eliptamin Vy.
Notes. The diagnostic character of this
species is the relative conspicuousness of
one costa (the 19th) at base of each elytron.
Form, color, punctation, and size are also
characteristic of the types. However, the
individuals listed under Additional material
are darker and much smaller than the types,
c. 6.5 mm long. They have the 19th costae
relatively conspicuous, as in the types, but
otherwise are more like secernendus. More
material from more localities is needed to
show wliether these specimens are referable
to humeralis or to secernendus, or whether
they represent a separate species. One
possibility is that humeralis occurs prin-
cipally at higher altitudes than secernendus
and that intermediates occur where their
ranges overlap.
Planeies ausfralis (Macleay)
Macleay 1871, Trans. Ent. Soc. New South Wales
2, p. 82 (PoUsticiis).
Description. None required here; note
size, prothorax with sides weakly or not
sinuate; pronotum c. evenly rather coarsely
punctate; length (in New Guinea) 7.7-9.5
mm.
Type(s). From Rockhampton, Queens-
land, Australia; presumably in Macleay
Mus., Sydney (not seen).
Occurrence in New Guinea. Papua: 1,
L. Daviumbu, Fly R., Sept. 11-20, 1936
(Archbold Exp., A.M.N.H.). West N. G.:
1, Kota Nika, Res. Hollandia, Jan. 25, 1956
(R. T. Simon Thomas, Louwerens Coll.),
in light trap; 1, Hoi Maffin, near Sarmi, July
18, 1959 (T. C. Maa, Bishop Mus.).
Notes. The New Guinean specimens
agree reasonably well with specimens from
Cairns, North Queensland, identified as
australis from description.
Planetes cordens n. sp.
Description. With characters of genus;
form as in Figure 134, depressed, with
wide-cordate prothorax; reddish piceous,
appendages paler; surface inconspicuously
pubescent (as usual); head and pronotum
shining between punctures, elytra dull.
Head 0.76 and 0.79 width prothorax, ir-
regularly impressed across base; front con-
vex except irregularly impressed anteriorly,
with a little irregular fine punctation. Pro-
thorax: width length 1.42 and 1.44; base
apex 0.97 and 0.96; sides broadly rounded
anteriorly, strongly sinuate posteriorly, mod-
222 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
erately margined, with usual 2 setae; pos-
terior angles well defined, right or slightly
obtuse; pronotum weakly convex, with well
impressed middle line and less distinct lon-
gitudinal impressions each side nearer mar-
gin than middle; baso-lateral impressions
shallow, closely microreticulate; surface of
disc otherwise rather closely punctate with
mixture of moderate and minute punctures.
Eli/fra subparallel; width elytra prothorax
1.34 and 1.30; apices obliquely truncate
with outer angles broadly rounded and in-
ner angles scarcely blunted ( as usual ) ;
each elytron with more than 20 fine costae,
the 1st, 4th, 7th, etc. slightly more promi-
nent than others, but no costa specially con-
spicuous at base. Secondary sexual char-
acters as for humcralis (2nd species above).
Measurements: length c. 9.0-9.5; width
3.2-3.3 mm.
Types. Holotype $, (Hungarian National
Mus.) and 3 paratypes (2 in M.C.Z., Type
No. 31,516) from Madang ("Friedrich-
W'ilh.-hafen"), N-E. N. G., 1896 (Biro);
and additional paratypes as follows, all
from N-E. N. G. ( Biro ) : 1, Stephansort,
Astrolabe Bay, 1900; 1, Simbang, Huon
Gulf, 1899; 1, Erima, Astrolabe Bay, 1897.
Meastired specimens. The S holotype and
1 9 paratype from Stephansort.
Notes. Among New Guinean Planetes,
this should be immediately recognized by
rather large size and wide-cordate prothorax
with disc flatter than usual and punctate as
described. Why Biro should have found
this species at four localities while no one
else has found it anywhere is a mystery.
Perhaps he obtained his specimens in a
special habitat b\' sjx'cial collecting methods,
perhaps by sitting leaf-debris from the
ground in rain forest.
Tribe HELLUODINI
Csiki 1932, Coleop. Cat., Carabidae, Harpaliiiac 7,
p. 1571 (sec for synoiiyiiiy and additional iclcr-
cnces ) .
This is a small tribe, confined to thv
tropical Asiatic-Australian area. ()iil\- 3
genera are recognized, of which only 1 is
represented in New Guinea and ( northern )
Australia. However, Iloloponerus fiodef-
froyi (Fairmaire) (1881, Le Naturaliste 3,
p. 381; 1883, Ann. Soc. Ent. Belgium 27, p.
2) of New Britain, although considered a
lebiine by Fairmaire and listed as one in
the Coleopterorum Catalogus (Csiki 1932,
p. 1361 ), may belong in this tribe. I do not
know this insect, but the description is of a
large carabid (perhaps the largest member
of the family in New Britain), 28 mm long,
with long mandibles, prothorax expanded
at sides, elytra sinuate-truncate and not
spined, and head at base with a strong
spine on each side. This description sug-
gests a very large Poi:,onoglossus-\ike carabid
with genae, which are prominently angu-
late or tuberculate in some Pogonoglossus,
produced as spines.
Genus POGONOGLOSSUS Chaudoir
Chaudoir 1862, Bull. Soc. Nat. Moscow 35, Part
2, p. 304.
Csiki 1932, Coleop. Cat., Carabidae, Harpaliiiae 7,
p. 1571 (see for synonymy and additional refer-
ences ) .
Andrewes 1937, Bull. Soc. Ent. France for 1937, pp.
152// (with key to species of Ja\a and Sumatra).
Diaiinosis. Form including form of eyes
characteristic; upper surface at least partly
pubescent; antennae not geniculate, with
moderate 2nd segments; see description of
ligula and paraglossae, below.
Description (characters common to New
Guinean species). Form as in Figures 135-
140; variably pubescent. Head: eyes ±
abruptly prominent, genae rounded or an-
gulate-tubereulate behind eyes; 2 setae over
each eye; antennae not gtmiculate, 2nd seg-
ments moderate (not \('r\ short), segments
1-4 variably setulose, outer segments mor(>
densely pub(>seent; mandibles long, weakly
arcuate; neck deeply transxcrseK con-
stricted; front 2-impressed anterior!) ;
clypeus irregularly truncate, apparenth
2- or 4-setose anteriorU- (setae difficult
to distinguish Ironi other jMibescence );
labium \ ariable. 6-sc>tose; ineiituin with
The Carabid Beetles of New Guinea • Darlington 223
triangular tooth; ligiila short, rounded, with
c. 4 setae at apex and 2 more near middle
of length; paraglossae very slender, much
longer than and free from ligula; palpi
with apical segments narrowly subtruncate.
Prothorax cordate or subcordate; apex
slightly or moderately (not deeply, in New
Guinean species) emarginate, with anterior
angles usually rounded ( ± pointed in some
g,lobncoUis) and not or not much advanced
beyond arc of emargination; base subtrun-
cate or emarginate at middle, ± oblique at
sides; posterior angles or sides of prothorax
just before angles usually minutely emar-
ginate; margins moderate or wide, reflexed,
each with seta at base and before middle;
disc usually only weakly convex, with mid-
dle line and transverse impressions distinct,
baso-lateral impressions present Init not
sharply defined. Elytra quadrate; margins
usually faintly subserrate; apices obliquely
emarginate-truncate with membranous mar-
gins, with outer angles usually rounded
(obtuse in popiio), inner angles c. acute or
blunted, not armed; striae entire ( ± obso-
lete in unicolor and globricoUis); intervals
variably punctate, 3rd with up to 3 or 4
special dorsal punctures (often not distin-
guishable especially in species with exten-
sive general punctation). Inner uiniis full.
Lower siu-jace variably punctate-pubescent.
Legs moderately slender; tarsi pubescent
above, not sulcate above; 4th hind-tarsal
segments shallowly emarginate; 5th seg-
ments setulose above and below; claws
simple. Secondary sexual characters: S
front tarsi not or not much widened, 3 seg-
ments narrowly 2-seriately squamulose; S
usuallv with 2 or 3, 9 3 or 4 setae each
side last ventral segment, but these setae
and their punctures sometimes difficult to
distinguish.
Type species. P. validicornis Chaudoir,
of Java.
Generic distribution. SE. Asia to north-
ern Australia.
Notes. Species of this genus are probably
moderatelv numerous and diverse from the
SE. corner of Asia to New Guinea (fewer
in northern Australia), but individuals are
rarely collected. Of 9 Javan and Sumatran
species, Andrewes ( 1937 ) saw only single
specimens of 6; and of 9 (or 10, with
unicolor (Macleay) ) New Guinean species,
I have seen a satisfactory series of only 1.
All the New Guinean species appear to be
endemic. I compared some of them with
the Andrewes Collection in 1948 ( see Notes
under several species, below); none fits
the description of P. horni Sloane ( 1907,
Deutsche Ent. Zeitschrift for 1907, p. 184)
of New Britain; and the 2 Australian species
that I have seen are different from any New
Guinean species.
Key to Species of Pogonoglossus of
New Guinea
1. Elytra] striae distinct, impressed 2
- Elytral striae faint or absent 9
2. Genae rounded or irregular behind eyes but
not conspicuously angulate or tuberculate
(see Description of taylori) 3
- Genae conspicuously angulate or tuberculate . 7
3. Entire upper surface including much of
head rather closely punctate or (on elytra)
roughened 4
- Part or all of upper surface sparsely punctate
or impunctate 6
4. Elytra with outer-apical angles obtuse but
distinct; size medium (length 9.0-11.5 mm);
(found at low altitudes) (p. 224) painia
- Elytra witli outer-apical angles rounded;
size either larger or smaller; (often at higher
altitudes ) 5
5. Larger, length 12.3-13.0 mm (see also De-
scription) (p. 224) taylori
- Smaller, length 7.6-8.7 mm (see also De-
scription) (p. 225) ininor
6. Prothorax less wide (width/length 1.49),
with moderate margins (p. 225) major
- Prothorax very wide (width/length 1.88 and
1.97), with very wide margins (p. 225) latior
7. Sides of prothorax oblique but scarcely sinu-
ate posteriorly; length c. 13 mm (p. 226) -
obliquus
- Sides of prothorax sinuate posteriorly;
smaller °
8. Length 9.6-11.0 mm (p. 226) grossuliis
- Length 7.0-9.0 mm (p. 227) parvus
9. Pronotum densely minutely punctate and
pubescent (p. 227) unicolor
- Pronotum virtually impunctate and glabrous
(p. 227) glahricollis
224 BuUetin Museum of Comparative Zoology, Vol. 137, No. 1
Pogonoglossus papua n. sp.
Description. With characters of genus;
form as in Figure 135; brownish black, ap-
pendages dark; entire upper surface mod-
erately pubescent, punctate, with reticulate
microsculpture indistinct or (on elytra) ir-
regular. Head 0.84 and O.Sl width pro-
thorax; genae rounded, not strongly angu-
late; front moderately punctate, shining be-
tween punctures. Prothorax strongly cor-
date; width/length 1.45 and 1.48; base/apex
1.14 and 1.19; base broadly slightly emar-
ginate, slightly oblique at sides; sides
strongly sinuate well before base; basal
angles shaiply formed, c. right; margins
rather wide, moderately reflexed; disc
weakly convex, surface moderately closely
punctate-pubescent, less shining than head
but more shining than elytra. Elytra:
width elytra prothorax 1.46 and 1.46; outer-
apical angles obtuse but more distinct than
usual in genus, striae moderately impressed,
indistinctly punctulate; intervals slightly
convex, closely punctate-pubescent. Sec-
ondary sexual characters as for genus; S
with 2, 9 3 special seta-bearing punctures
each side last ventral segment. Mca.siire-
ments: length c. 9.0-11.5; width 3.4-4.1
mm.
Types. Holotype $ (M.C.Z., Type No.
31,517) and 10 paratypes from Dobodura,
Papua, Mar.-July 1944 (Darlington); and
additional paratypes as follows. Papua: 1,
without precise locality (Hungarian Na-
tional Mus.). N-E. N. G.r 1, Lae, sea level,
July 24, 1955 (Gressitt), in light trap; 1,
Busu H., E. of Lae, 100 m, Sept. 13, 1955
(Gressitt); 7, Aitape, Aug. 1944 (DarHng-
ton). WestN. G.: 1, Hollandia, July-Sept.
1944 (13arlingt(m); 2, same locality. May
1945 (II. Iloogstraal, M.C.Z.); 1, same lo-
cality, June 1945 ( B. Malkin, U.S.N.M.); 3,
same locality, dates in Nov., Dec, Jan.
1944-1945 (W. T. Nailon, Fenton Coll.);
1, Sentani, 90+ m, June 22, 1959 (Gressitt
and T. G. Maa, Bishop Mus.), in light trap;
1, Hoi Maffin, near Sarmi, July 18, 1959 (T.
G. Maa, Bishop Mus.); 1, Neth. N. G. with-
out further locality, Oct. 10, 1944 (T.
Aarons, Gal. Acad.). Also 1, "Sinimi" { =
Senimi R., Papua?), "Vr, 1943" (T. Niimura,
Ueno Goll.).
Measured specimens. The S holotype and
1 9 paratype from Dobodura.
Notes. In Andrewes' key ( 1937, see refer-
ence under genus), this species runs to
latus Andrewes of Sumatra but has the pro-
thorax probably narrower at base and less
emarginate at apex and the pronotum cer-
tainly more closely punctate. Of Australian
species, papua is closest to porosus Sloane
( I have specimens, identified from descrip-
tion, from Rocky R., mid-peninsular Gape
York ) but has a more strongly cordate pro-
thorax and better defined outer-apical
elytral angles.
Since papua is the common Poii,ono'jJos-
sus in New Guinea, I shall take it as a
standard for comparison of sexeral of the
following species.
Most of my Dobodiua specimens were
taken in piles of dead leaves on the ground
in rain forest.
Pogonoglossus taylori n. sp.
Description. With characters of genus;
form and characters c. as preceding species
{papua) except as follows. Head 0.15 and
0.78 width prothorax; genae more promi-
nent than in papua, nearly wide as eyes,
minutely tuberculate and e. subangulate be-
hind eyes. Prothorax: width length 1.44
and 1.58; base/apex 1.18 and 1.18; apex
slightly more emarginate than in ))apua
and sides slightly more broadl\ and e\enl>
reflexed. Elytra: width elytra prothorax
1.43 and 1.43; outcM-apical angles more
rounded than in papua. Measurenwnis:
length 12.3-13.0; 4.3-4.9 mm.
Types. Holotype S (M.G.Z., l\pe No.
31,518) from Aiyura, N-E. N. G.. 'l900 m
July 1962 (R. W. Taylor, #2147), in rain
forest; 1 9 paratvpe, Eliptanu'ii \\'., N-E.
N. G., 1665-2530 m, June 23-30. 1959 (W.
W. Brandt, Bishop Mus.); 1 s paratvpe,
Okapa, N-E. N. G., Aug. 29, 1965 (R.
Hornabrook).
The Carabid Beetles of New Guinea • Darlington
225
Measured specimens. The i holotype and
9 paratype from Eliptamin Vy.
Notes. This is apparently a mountain-
Hving species probably related to the low-
land papiia but differing from it as indi-
cated in the Description above.
Pogonoglossus minor n. sp.
Description. With characters of genus
(except i unknown); form c. as in papua;
characters c. as in papna except as follows.
Color browner (less black), surface slightly
more shining. Head 0.88 and 0.86 width
prothorax; eyes slightly smaller and genae
more evenly rounded than in papiia. Pro-
thorax: width length 1.47 and 1.49; base
apex 1.08 and 1.07. Elytra: width elytra
prothorax 1.50 and 1.51; outer-apical angles
more rounded than in papiia; intervals less
roughened. Measurements: length 7.6-8.7;
width 3.0-3.3 mm.
Types. Holotype 9 (M.C.Z., Type No.
31,519) from lower Busu R., Huon Pen.,
N-E, N. G., May 12, 1955 (E. O. Wilson),
in lowland rain forest; 1 9 paratype, W'au,
Morobe Dist., N-E. N. G., 1200 m, June 22,
1961 (Sedlaceks); 1 9 paratype, Hollandia,
West N. G., Jan. 20, 1945 (W. T. Nailon,
Fenton Coll.); 1 9 paratype, Njau-limon,
S. of Mt. Bougainville, West N. G., 300 ft.,
Feb. 1936 (Cheesman).
Measured specimens. The 9 holotype and
9 paratype from Njau-limon.
Notes. P. minor differs from papua as
indicated in the preceding Description.
The 2 species are sympatric but minor is
apparently the less widely distributed,
being known only from a comparatively
small part of east-central New Guinea.
P. minor is similar also to porosus Sloane
of North Queensland, Australia, but has the
head more punctate and the prothorax more
strongly cordate.
Pogonoglossus major n. sp.
Description. With characters of genus;
form as in Figure 136; irregular brownish
piceous, appendages dark brown; rather
shining, reticulate microsculpture absent or
indistinct on head and pronotum, light,
irregular, moderately transverse on elytra;
surface punctate as described below. Head
0.82 width prothorax; genae prominently
rounded but not angulate; front sparsely
punctate-pubescent. Prothorax weakly cor-
date; width/length 1.49; base/apex 1.05;
sides broadly sinuate before obtuse except
minutely subdenticulate posterior angles;
surface irregularly rather sparsely punctate-
pubescent. Elytra: width elytra/prothorax
1.44; outer-apical angles broadly rounded,
sutural angles blunted; striae deep, entire,
finely punctulate; intervals convex, sparsely
punctate, 3rd with apparently 3 or 4 special
dorsal punctures difficult to distinguish
from other punctures. Secondary sexual
characters: S front tarsi as genus; 6 with
apparently 3 principal setae on left, 4 on
right side last ventral segment; 9 unknown.
Measurements: length 17.5; width 6.0 mm.
Type. Holotype S (M.C.Z., Type No.
31,520) from vie. Nadzab, N-E. N. G., July
1944 (Darlington); the type is uniciue.
Notes. This is the largest New Guinean
Pogonoglossus. It is about the size of P.
Jwrni Sloane (Deutsche Ent. Zeitschrift for
1907, p. 184) of New Britain but has the
prothorax more narrowed behind with more
obtuse posterior angles, the outer elytral
striae not fainter, and the elytral intervals
sparsely rather than closely setose-punctate.
Pogonoglossus lafior n. sp.
Description. With characters of genus;
form as in Figure 137, very broad; brownish
black, appendages dark; moderately shin-
ing, reticulate microsculpture indistinct on
head and pronotum, light, fine, rather
strongly transverse on elytra; punctation as
described below. Head 0.74 and 0.67 width
prothorax; genae oblique for most of length,
slightly rounded or very obtusely subangu-
late behind eyes; front c. impunctate ex-
cept sparsely punctate laterally and pos-
teriorly. Prothorax very wide, cordate;
width length 1.88 and 1.97; base/apex 1.24
226 Btilletin Museum of Comparative Zoology, Vol. 137, No. 1
and 1.12; sides strongly but variably sinuate
well before c. right or obtuse posterior
angles; margins widely reflexed; disc more
convex than usual, sparsely inconspicuously
punctate-pubescent. Elytra: width elytra
prothorax 1.45 and 1.30; outer-apical angles
rounded, sutural angles blunted; striae
entire, well impressed, slightly irregular
but scarcely punctulate; intervals convex,
sparsely inconspicuously punctulate, 3rd
with 3 or 4 dorsal punctures difficult to
distinguish. Sccundary sexual characters: 6
front tarsi as for genus; 6 with 3, 9 4 setae
each side last ventral segment. Measure-
ments: length 14.5-15.5; width 5.7-6.0 mm.
Types. Holotype c5 ( Leiden Mus. ) and
1 9'paratype (M.C.Z., Type No. 31,521)
from Lower Mist Camp, Snow Mts., West
N. G., 1550 m, Jan. 31, 1939 (Toxopeus).
Notes. See Key to Species for distinguish-
ing characters of this well defined species.
The 9 has a wider prothorax with more
obtuse angles than the S , but this is prob-
ably individual rather than sexual variation.
I have no doubt the 2 specimens are con-
specific.
Pogonoglossus obliquus n. sp.
Description. With characters of genus;
form as in Figure 138; black, appendages
dark; shining, reticulate microsculpture
absent or indistinct even on elytra; puncta-
tion as described l)elow. Head 0.77 width
prothorax; genae prominently angulate-
tuberculate behind eyes; front almost
smooth, very sparsely punctulate-pubescent.
Prothorax very wide; width/length 2.00;
base/apex 1.22; sides oblicjue and converg-
ing and scarcely sinuate before obtuse pos-
terior angles; margins widely reflexed; disc
moderately convex, sparsely punctate-jm-
bescent, more closely so across base and
apex. Ehjtra: width not measured (speci-
men broken); humeral margins wider ihan
usual; outer-apical angles roimded, sutural
angles acute, scarcely blunted; striae entire,
impressed, irregular but seareel\- punctu-
late; interxals con\-ex, finely s])arsel\- punc-
tulate, 3rd with c. 3 special dorsal punc-
tures difficult to distinguish. Secondary
sexual cJiaracters: 6 front tarsi as for
genus; i with 3 setae each side last
ventral segment; 9 unknown. Measure-
ments: length c. 13 mm; width not mea-
sured.
Ty])e. Holotvpe 6 (Bishop Mus.) from
Eliptamin Vy.,'N-E. N. G., 1665-2530 m,
June 23-30, 1959 (W. W. Brandt); the type
is unique.
Notes. The single specimen was received
in bad condition and remounted in pieces
on a card, but it shows the essential char-
acters of this very distinct species. See Key
to Species for its differential characters.
Pogonoglossus grossulus n. sp.
Description. With characters of genus;
form average; black or brownish black, ap-
pendages dark; shining, reticulate micro-
sculpture indistinct even on elytra. Head
0.80 and 0.79 width prothorax; genae promi-
nently angulate-tuberculate behind eyes;
front virtually smooth at middle, very
sparsely punctulate-setose at sides. Pro-
thorax wide-cordate; width length 1.95 and
1.94; base/apex 1.23 and 1.19; sides broadly
sinuate before c. right or obtuse posterior
angles; margins wide, widely reflexed; disc
weakly convex, very sparsely punctulate-
pubescent. Eh/tra: width elytra prothorax
1.27 and 1.36; outer-apical angles rounded,
sutural angles acute (except for membranous
margins); striac> deep, scarcely punctulate;
intervals convex, very sparsely punctulate,
3rd with up to 3 special dorsal punctures
difficult to distinguish. Secondary sexual
characters as for genus; S with 2, 9 with 3
setae each side last ventral segment. Mea-
surcmoils: length 9.6-11.0; width 3.7-4.1
mm.
Types. Holotype 9 (M.C.Z., Type No.
31,522) Irom \ic. ZcMigarn, \'\-. of Kua H.,
Mongi Watershed, Ihion Pen., I\-K. N. (i..
800 m, Apr. 14, 1955 (E. O. Wilson); 1
9 paratype, Lae, N-E. N. (i.. JuK 1941
(F. E. Skinner, Purdue V. Coll., borrowed
fr. Bisho]-) Mus.); 1 ^ paratope. Kokoda.
The Carabid Beetles of New Guinea • Daiiington 227
Papua, 1200 ft. (366 m), Sept. 1933 are distinet in both specimens and may
(Cheesman). prove to be characteristic of the species,
Measured specimens. The ^ paratype and although similar marks are indicated in
9 holotype, in this order. some individuals of some other species.
Notes: Although the 3 individuals listed
above vary somewhat, they agree in form Pogonoglossus unicolor (Macleay)
of genae, wide-cordate prothorax, shining Macleay 1886, Proc. Linnean Soc. New South
sparsely punctate surface, and moderate Wales (2) 1, p. 137 {Planetes).
size. I think they probably represent a ^'"''"'^ ^^^'^^ Deutsche Ent. Zeitschrift for 1907,
single, variable species.
Description ( significant details only, from
Pogonoglossus parvus n.sp. Macleay's description). Color brownish
Description. With characters of genus; hlack, legs dark; head shining, pronotum
form as in Figure 139; brownish black, head and elytra dull and densely minutely punc-
with 2 narrow oblic^ue red marks posteriorly tate; prothorax a little wider than long, with
(see following Notes), appendages dark titles narrowed to posterior angles which
brown; moderately shining, reticulate micro- ^re "rather obtusely rectangular"; elytra
sculpture indistinct on head, irregular or "with 8 or 9 almost invisible striae"; length
transverse and light on pronotum and elytra, c. 10 mm.
Head 0.85 and 0.85 width prothorax; genae Type. From Fly R. (probably Papua);
angularly prominent behind eyes; much of should be in Macleay Mus., Sydney (not
front virtually impunctate. Prothorax cor- seen).
date; width/length 1.48 and 1.56; base apex Notes. Sloane (1907) adds nothing to
1.21 and 1.23; sides broadly sinuate before Macleay's description of unicolor except
obtuse or nearly right posterior angles; ^^'^^ the insect is a Pogonogjo.ssus. The
margins rather narrow and not strongly re- ^^^Y lightly striate elytra distinguish it from
flexed; disc moderately convex, finely ^H known New Guinean species of this
punctate-pubescent ( pubescence rubbed off genus except glabricoUis Van Emden ( be-
in part in holotype ) . Elytra: width elytra low), from which it differs in having the
prothorax 1.34 and 1.35; outer-apical angles pionotum densely punctate and pubescent
narrowly rounded, sutural angles acute or rather than smooth and virtually glabrous
slightly blunted; striae well impressed, ir- '^■'^ i" glabricoUis.
regular but scarcely punctulate; intervals „ ; ; , • //■ v, .- ■
r.r^r.,.^v. c. 1 1^ 1 4. 1 i. o 1 Pogonoglossus glabncollis Van Emden
convex, sparsely punctulate-pubescent, 3rd ^ ^ »
^^'ith special dorsal punctures not surely ^''" ^'"''™ ^•^•^"' ^^^""^^■•" E"^- ^^^^""'^ 9^' P- ^^■
distinguishable. Secofu/f/rj/ .sexjva/ c/i<3/rtcfers Description. With characters of genus;
as for genus; c^ with 2 or 3 (unsymmetric), form as in Figure 140 (but somewhat vari-
9 3 setae each side last ventral segment, able); irregular dark brown or brownish
Measurements: length 7.0-9.0; width 2.5- black, head with 2 reddish marks posteri-
3.2 mm (the 6 is the larger). orly, appendages brown; moderately shining,
Types. Holotype $ (M.C.Z., Type No. elytra duller, reticulate microsculpture in-
31,523) from vie. Hollandia, West N. G., distinct on head and pronotum, light and
July-Sept. 1944 (Darlington); and 1 9 irregular on elytra. Head 0.85 and 0.83
paratype, Dobodura, Papua, Mar.-July 1944 width prothorax; genae rounded; front
(Darlington). almost impunctate. Prothorax cordate, vari-
Notes. The small size, angulate genae, able; width/length 1.39 and 1.60 {sic);
and form and microsculpture distinguish base/apex 0.98 and 1.07; sides broadly sinu-
this species. The red marks on the head ate before obtuse or c. right posterior
228 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
angles; margins rather narrow l:)iit variable;
anterior angles rounded or bluntly pointed;
disc almost flat, scarcely punctulate. Elytra:
width elytra/prothorax 1.35 and 1.37; outer-
apical angles broadly rounded, sutural
angles acute or blunted; striae faintly in-
dicated or virtually obsolete; surface closely
punctulate; up to 3 apparent dorsal punc-
tures sometimes visible on position of 3rd
intervals. Secondary sexual characters un-
determined ( i unknown); 9 with 3 or 4
(sometimes unsymmetric) setae each side
last ventral segment. Measurements: length
12.5-16.0; width 4.2-5.3 mm.
Type. A 9 from N-E. N. G.; in Van
Emden Coll., British Mus. (seen).
Occurrence in New Guinea. Papua: 1
9, Kokoda-Pitoki, 400 m. Mar. 23, 1956
(Gressitt); 1 9, Fiume Purare, Jan. 1894
(Loria, borrowed from Straneo). N-E.
N. G.: the holotype; 1 9, Motae, Kuku
Kuku, E. Highlands, 6000 ft. (c. 1830 m),
"1/3/64" (R. Hornabrook). West N. G.: 1
9 , Geelvink Bay, 1878 ( Raffray and Main-
dron, Paris Mus.).
Measured specimens. The 9 9 from
Kokoda-Pitoki and Motae.
Notes. Although the 4 individuals listed
above vary in several characters ( e.g. form
of prothorax, degree of obliteration of
elytra! striae), the variations are not obvi-
ously concordant, and I think only one very
distinct species is involved. It is uniquely
characterized by form, elytral striae faint or
obsolete, and combination of virtually im-
punctate jironotum and densely punctulate
elytra.
Tribe HELLUONINI
Sloanc 191 I, I'loc. Liiiiican Soc. New Sontli Wales
39, p. 568.
Csiki 1932, Colcop. Cat., Carabidac, liarpalinac 7,
p. 1572 (see for synonymy and additional rcfcr-
fnccs).
Jeanne! 1949, Colcop. Carahiciucs dc la Hejjion
MaljJiaclie, Part 3, p. 1041.
Jrdlicka 1963, Knt. Abliandlnn«en 2S, p. 167.
llclliioiiinac Basilfwsky 1953, Exploration Pare
National I'Upemba, Fasc. 10. p. 219.
This is still another small but widely
distributed tribe. The members of it are
medium-sized or large carabids, usually of
characteristic form, usually with sparse or
short pubescence, and usually with mouth-
parts including the labrum strikingly modi-
fied. Three genera are confined to the
Americas; 6, to Africa and/or the Oriental
Region (except that a species of Crea^.ris
extends to Australia); 13, to the Australian
Region. (A supposed helluonine on Ne\v-
Caledonia has been shown not to be one
by Britton, 1937, Ent. Monthly Magazine
73, p. 127. ) The Australian genera form a
distinct group of the tribe, characterized by
Sloane (1914, p. 570). Five genera (1 of
them new) and 8 species of Australian-
group Helluonini occur in New Guinea,
where the only other member of the tribe
is CreaiS,ris lahrosa, \\'hich ranges frotn
Ceylon and India to Australia.
In spite of Sloane's ( 1914 ) careful stud\
of the Australian genera, I have had trouble
with the generic classification of the New
Guinean forms. This is partK' because m\'
material is inadequate: 2 obviously distinct
new species are represented by unique fe-
males which I have assigned to Ilelhionidius
with some doubt, and I have been forced to
base an apparent new genus on a single
male. I myself found no Helluonini during
11 months in New Guinea and I can sa\
nothing about their habitats or habits there
except that all the New Guinean species
are winged and that some of them fl\' to
light. In Australia, different hc41uonines
lixc on the ground and on tree trunks, usu-
alK in open or openly-wooded places rather
than in rain forest.
K'i;v lo (Iknkha ok IIki.ia omm or Xi:w Cvinka
1. Front iemora not anuniate-protnlicrant he-
low: size smaller, lenutli r. 9 mm (p.
229 ) _ - - Creofiris
- Front femora tliickened and lilnntly angn-
late or protnl)eranl hi'low near base; size
larjier — - 2
2. Prothorax inoderateK' narrowed posteriorly,
witii si(.les moilerately siimate (p. 2'v3 ) _,
IIcIIikkIciiki
The Carabid Beetles of New Guinea • Darlington
229
- Piothoiax strongly narrowed posteriorly, with
sides strongly sinuate and base subpeduncu-
late 3
3. Ligula subtriangular, narrowed anteriorly,
with apex narrowly rounded ( p. 233 )
Helhiosoma
- Ligula very wide, with apex broadly rounded
or eniarginate 4
4. Labrum with 2 principal setae; elytra with
Sth intervals much wider tlian 7th and closely
punctate; length ( in New Ciuinea ) c. 30 mm
( p. 233 ) Gigadema
- Lal)rum with 4 or more principal setae; elytra
with 8th intervals not much wider than 7th
and less closely punctate; length c. 20 mm
or less 5
5. Tarsal segments unusually widened or par-
allel-sided, the 4th hind-tarsal segments eniar-
ginate for more than V2 the segments' length;
labrum usually with 4 principal setae (p.
229 ) Helluoniduis
- Tarsal segments not thus widened and not
parallel-sided, the 4th hind-tarsal segments
shallowly emarginate; laJMum with c. 10
principal setae (see also Description) (p.
232 ) HcUuopapud
Genus CREAGRIS Nietner
Nietncr 1857, J. Asiatic Soc. Bengal 26, p. 139.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1575 (see for synonymy and additional refer-
ences ) .
Diagnosis. See preceding Key to Genera.
Description. None required here.
Type species. Creagris lahrosa Nietner,
below.
Generic distribution. Six species in the
Oriental Region, 1 of them extending to
New Guinea and Queensland, Australia;
possibly an additional species in Queens-
land.
Notes. The listing of C. wiJsoni Castelnau
(the supposed endemic Queensland species)
also from Java by Csiki (p. 1576) is ap-
parently a compiler's error based on a mis-
reading of Sloane 1914 ( see reference under
following species ) .
Creagris labrosa Nietner
Nietner 1857, J. Asiatic Soc. Bengal 26, p. 139.
Sloane 1914, Proc. Linnean Soc. New South Wales
39, p. 570.
1920, Proc. Linnean Soc. New South Wales
45, p. 322.
Csiki 1932, Coleop. Clat., Carabidae, Harpalinae 7,
p. 1575 (see for synonymy and additional refer-
ences ) .
Description. None required here; note size
small; color dark brown; labrum expanded,
r. circular, shallowly channeled each side;
mentum with lateral lobes and median
tooth all produced as long very slender
processes; front femora not angulate below;
length c. 9 mm.
Types. From Ceylon; in Berlin U. Zool.
Mus. and Stettin Mus. (not seen).
Occurrence in New Guinea. Papua: 2,
Mt. Lamington, 1300-1500 ft. (c. 400-460
m) (C. T. McNamara, S. Australian Mus.).
Notes. C. lahrosa is now known from
Ceylon, India, Burma, etc., Java, New
Guinea, and Queensland, Australia (a
specimen from Mackay, recorded by
Sloane, 1914).
Genus HELLUONIDIUS Chaudoir
Chaudoir 1872, Revue et Magasin Zool. (2) 23,
p. 216.
Sloane 1914, Proc. Linnean Soc. New South Wales
39, pp. 571, 582.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1580 (see for synonymy and additional refer-
ences ) .
Diagnosis. Form c. as in Figure 142;
pubescent (as usual in tribe); genae vari-
able; labrum variable, produced or angu-
late at apex, usually with 4 principal setae;
mentum toothed; ligula rounded; prothorax
strongly constricted at base; elytra with Sth
intervals not much wider than 7th, irregu-
larly punctate; 4th hind-tarsal segments ±
wide, deeply emarginate; see also preceding
Key to Genera of UeUuonini of New Guinea.
Description. None attempted here; ma-
terial inadequate.
Type species. Aenigma cyanipcnne Hope,
of Australia.
Generic distribution. Eastern and north-
ern Australia, New Guinea.
Notes. Of the 4 New Guinean species
now placed in this genus, only chrysocomes
Maindron is a typical Helluonidius. The
other 3 species, 2 of them based on unique
230 Bulletin Museum of Coniptudfive Zoology, Vol. 137, No. 1
females (sex determined by dissection),
differ in form of labrum and differ among
themselves in form of genae, form of tarsi,
and in other ways. They are obviously
distinct species, but males are needed to
determine generic assignments.
Key to Species of Helluonidius of
New Guinea
1. Lalnuni loiijier than wide, narrowly pro-
duced at apex (p. 230) chrysocoitics
- Labrum wider than long, not narrowly pro-
duced at apex 2
2. Labrum witli median setae close to margin;
cKtra microreticulate (p. 230) laevifrons
- I^abrum with median setae c. V:\ lalirum's
length behind margin; elytra not micro-
reticulate 3
3. Genae abruptly truncate, c. straight from
posterior edges of eyes to neck; tarsi \'er>'
wide (Fig. 184) (p. 231) latipes
- Genae moderately convex; tarsi less wide (p.
231) politus
Helluonidius chrysocomes Maindron
Mainchon 1908, Ko\a (Guinea 5, Li\raisf)n 2, p.
299.
Description. None required here; form
as in Figure 142; form of labrum ( longer
than u'ide, and narrowK' produced at apex )
unique in tribe Ilelluonini in New Guinea;
genae prominent; antenna and hind tarsus,
Figure 182; color black or dark brown; sur-
face without reticulate microsculpturc;
lengtli c. 16.()-1(S.5 mm.
Type. From "Sentani" (near llollandia ),
West N. G.; probably in Paris Mus. (not
seen ) .
Occurrence in Neic Cuineo. Twenty-two
speeimens from 11 localities scattered oxer
most ol the length of New (Guinea, from
I^)rt Moresby and the Fly R. to the N'ogel-
kop; at low iiltitudes, none labeled higluM"
than SOO ni ( Araucaria Camp).
Notes. This is a t\'pical IJelluoniditis,
very close to ;ind perhajis the same as one
of the Australian species, which are all \'(m\'
similar and which n(H>d revision.
Most spec iiiK'iis ol elinjsocomes haxc the
surface especially ol the elytra w ith a jiearly
luster which is apparently due to a surface
film of some sort, but fi individuals w ideU'
scattered within the range of the species are
shining black without luster.
Helluonidius laevifrons n. sp.
Description. With characters of genus;
form as in Figure 143; antenna and hind
tarsus. Figure 183; black, appendages dark;
head and pronotum shining (but punctate
as described below), elytra duller, with c.
isodiametric microsculpturc. Head 0.84
width prothorax; genae oblique, not at all
prominent; chpeus subtrimcate, 1 -setose
each side; labrum wider than long, obtuseK'
subangulate, sinuate each side of angula-
tion, 4-setose with inner setae very close to
anterior margin (see Notes, below); front
deeply impressed each side, the impressions
punctate but front otherwise broadh' im-
punctate; neck moderately impressed and
punctate; mentum with large triangular
tooth; ligula wide, irregularly rounded, de-
pressed at middle, with 3 pairs seta-bearing
punctures one behind the other; inner lobe
of maxillae with inner edge irregular near
middle and with strong hook at c. right
angles from inner-apical angle; palj^i rather
short and thick. Protliora.x: width length
1.19; base/apex 1.02; base head 0.72; mar-
gins narrow, scarcely interrupted; disc ir-
regularly convex, irregularly coarsely punc-
tate with punctures tcmding to form longi-
tudinal rows between n;urow smooth sp;ices
at middle. Elyfrd: width ebtra prothorax
1.22; striae well impressed, punctulate; in-
tervals slight 1\ convex, most int(M\als
slightly irrt\gularly 2-seriately punctate,
si)ccial dorsal punctures of 3rd not sureK
distinguish;ibl(\ 8th slightK' wider than 7th
and irregularly (not 2-seriately ) punctukite.
Lciis: tarsal segments more nearl\- j^arallel
than usual (Fig. 183); 4th hind-tarsal seg-
ments emarginate for slightK more* than ^-i
length. Measurcnicnls: length 18.0; width
6.5 mm.
Type. Il()l()t\pe V (Bishop Mus.) Irom
Torrieelli Mts.,'M()kai Village, N-E. N. G.,
750 m. j;ui. 1-23, 1959 (W. \\ . Brandt);
the type is unique.
Notes. The lionl maiuiii ol ihc ];i1)rum
The Carabid Beetles of New Guinea • Dadingion
231
is slightly sinuate on each side, with the
inner seta-bearing puncture near margin at
the sinuation. This form of lahrum is inter-
mediate between that of //. cJirysocomcs
and those of the following two species. But
in some other ways ( form of genae and
especially form of tarsi ) the present species
is strongly characterized, not intermediate.
Helluonidius lafipes n. sp.
Description. With characters of genus;
form as in Figure 144; antenna and hind
tarsus. Figure 184; dark brown, appendages
dark; shining, without reticulate microsculp-
ture, but punctate as described below.
Head 0.82 width prothorax; genae oblique,
not at all prominent; clypeus subtruncate,
2-setose each side; labrum wider than long,
bluntly obtusely angulate, not sinuate at
sides of angulation, 4-setose with inner setae
almost % length of labrum behind anterior
margin; front irregularly convex, deeply im-
pressed each side anteriorly, punctate at
sides and base but c. impunctate at middle;
mentum with strong bluntly triangular
tooth; ligula wide, rounded, scarcely im-
pressed at middle, probably 6-setose as in
laevifrons (above) but anterior setae
covered or broken; inner lobe of maxillae c.
as in laevifrom-, palpi stout. Frothorox:
width length 1.40; base apex 1.01; base '
head 0.72; margins narrow, much inter-
rupted anteriorly; disc irregularly convex,
irregularly punctate. Elytra: width elytra/
prothorax 1.52; striae impressed, scarcely
punctulate; intervals slightly convex, ir-
regularly 2-seriately punctate, 3rd with
special dorsal punctures not surely distin-
guishable, 8th not wider than 7th, irregu-
larly, rather sparsely in part 2-seriately
punctate. Le^s: tarsi exceptionally wide
(Fig. 184); 4th hind-tarsal segments much
wider than long, deeply and widely emar-
ginate; 5th segments wide and flattened.
Measurements: length 19.8; width 6.8 mm.
Type. Holotype 5 (Leiden Mus.) from
Rattan Camp, West N. G., 1200 m, Feb.-
Mar. 1939 (Toxopeus); the type is unique.
Notes. This species is assigned to
Helluonidius with doubt. The genae are
formed as in the preceding species (laevi-
frons) and the labrum is almost the same
in shape, but the 2 inner setae of the
labrum are much farther back and the tarsi
are strikingly different.
Helluonidius polifus n. sp.
Description. With characters of genus;
form as in Figiue 145; antenna and hind
tarsus. Figure 185; brownish black, append-
ages dark; shining, without reticulate micro-
sculpture but punctate as described below.
Head 0.84 width prothorax; genae moder-
ately convex, subprominent; clypeus broadly
slightly emarginate, 2-setose each side;
labrum wider than long, bluntly angulate,
with apparently 2 principal setae on right
and 4 on left, the inner seta on each side
almost V:'. length of labrum behind anterior
margin; front weakly convex, deeply im-
pressed each side anteriorly, punctate at
sides and across base and with a few widely
scattered punctures near middle; mentum
with strong blunt tooth; ligula wide,
rounded, scarcely impressed, 6-setose as in
laevifrons: inner lobe of maxillae c. as in
laevifrons and latipes, with hook from inner-
apical angle; palpi less thick than in pre-
ceding species. Prothorax: width length
1.33; base/apex 0.97; liase/head 0.68; mar-
gins narrow, much interrupted anteriorly;
disc weakly irregularly convex, irregularly
coarsely punctate. Elytra: width elytra/
prothorax 1.44; striae impressed, punctulate;
intervals convex, irregularly 2-seriately
punctate, 3rd with special dorsal punc-
tures not surely identifiable, 8th not much
wider than 7th, irregularly in part 2-seri-
ately sparsely punctate. Legs: tarsi mod-
erately wide and flattened, but less so than
in preceding species (latipes); 4th hind-
tarsal segments wide, very deeply emargi-
nate. Measurements: length 16.7; width
5.3 mm.
Type. Holotype $ (M.C.Z., Type No.
31,524), from Maba Vy., Menyama, Mo-
232
BuUetiu Museum of Comparative Zoology, Vol. 137, No. 1
robe Dist., N-E. N. G. (L. Hastings); 1 i
paratype (Bishop Mus.), Oriomo River, 6
m, Feb. 13, 1964 ("H.C."), light trap; 1
9 paratype (M.C.Z.), Maprik, N-E. N. (;..
Oct. 14, 1957 (Gressitt), hght trap.
Notes. This species is simihir to the pre-
ceding {hiti])cs) l)ut has more prominent
genae and narrower tarsi. The genae are
less prominent than in dinjsocomes but lead
toward that species, while the labrum sug-
gests a relationship \\ ith Jatipcs.
HELLUOPAPUA n. gen.
Dia^no.sis. Form of lleUuonidius; labrum
wide, multisetose; ligula wide, slightly
emarginate, setose as in llcUuonid\ns\ inner
lobe maxillae strongly hooked on inner side
before apex; Sth elytral intervals e. wide as
7th, 2-seriately punctate with few or no
scattered punctures; tarsi slender, 4th hind-
tarsal segments shallowly emarginate; front
femora obtusely pronu'nent below near base;
i front tarsi without squamae; i copula-
tory organs as in Figure LSI.
De.seription. See description of only
known species, below.
Ti/})e speeies: H. toxopei, below.
Gencrie distri])ution. Known from a
shigle locality in West N. (i.
Notes. This new genus differs from Ilel-
hionidiiis in labrum multisetose; hook of
inner lobe of the maxillae subapieal (not
apical), and tarsi much more slend(>r with
4th hind-tarsal segments shallowly (not
deeply) emarginate. It differs from Cwi,^/-
dema in labrum multisetose and (Sth el\ tial
intervals much narrow(M- and 2-seriatel\
(not densely) ptinctatc. It fits no other
genus in Sloane's (1914, pp. 571-572) key.
And it differs from all previously known
Ilelluonini of Sloane's ( 1914, p. 570) "Aus-
tralian (irouj) in lacking sexual s((namae
on i front tarsi.
Helluopapua toxopei n. sp.
Deseiij)li()iL With characters of genus;
form as in I'igure 146; slend(>r, subparallel.
depressed; antennae and hind tarsus, Figure
186; black, appendages dark brown, 2
minute red spots on head posteriorly; sur-
face sparsely pubescent, moderately shining,
microsculpture faint and irregular on head
and pronotum, more distinct and e. isodia-
metric on elytra, and surface punctate as
described below. Head 0.90 width pro-
thorax; genae rounded, moderately promi-
nent; clypeus slightly sinuate-truncate, with
several setae each side; labrum wider than
long, wide in front, broadly sinuate each
side in front with apex obtusely angulate,
with several principal setae each side but
n(jne near middle; front irregularly convex,
deeply impressed each side and transversely
impressed anteriorly (individual character?),
irregularly punctate at sides and posteriorly;
mentum with strong triangular tooth and
side lobes long and pointed; ligula as de-
scribed for genus; palpi rather slender.
ProtJiorax: width length 1.38; base apex
0.78; base head 0.69; margins narrow, not
interrupted; disc weakly irregularly convex,
surface irregularly pvmctate. Elytra: width
elytra prothorax 1.52; striae impressed, not
punctulate; intervals slightK' comex. rather
sparsely 2-seriately punctate. Measure-
ments: length 22.5; w idth 6.7 mm.
Type, llolotype S (sex determined by
dissection) (Leiden Mus.) from Rattan
Camp, West N. G., 1200 m, Feb.-Mar.
1939 (Toxopeus); the t\'pe is uniciue.
Notes. Although this ,j is from the same
localit)' as the 9 type of lleUuo)\idius
Idlipes. the two specimens differ in so man\'
ways that the\' cannot be one spt>cii\s but
ha\(' to be assigned to different genera:
the two specimens differ in lonn. in genae.
in sha])e and setae ol labrum, in position
of hook of inner lobe of maxillae, and in
form of tarsi, and this is just a beginning of
the list of differences. A r(>\ision ol all Ntn\
(iuinean and Australian members of the
tribe, with mucli additional material, will
l^robabK' be nec-essary to decide the real
relationships of this new genus as well as of
the new species ol I lelluonidiits described
above.
The Carabid Beetles of New Guinea • Darlington 233
Genus HELLUOSOMA Castelnau
Castelnau liSttT, Notes on Australian Coleop., p.
20.
Sloane 1914, Proc. Linnean Soc. New South Wales
39, pp. 571, 585.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1581 (see for synonymy and additional refer-
ences ) .
Diopiosis. See preceding Key to Genera.
Description. None required here.
Type species. H. atrum Castelnau (be-
low ) .
Generic distribution. Tropical Australia
and New Guinea.
Notes. Only one species of this genus is
adequately known, although a second
species may exist in Australia ( Sloane 1914,
p. 586).
Helluosoma afrum Castelnau
Castelnau 1867, Notes on Australian Coleop., p. 21.
Sloane 1914, Proe. Linnean Soc. New South Wales
39, p. 586.
Csiki 1932, Coleop. Cat., Caraliidae, Harpalinae 7,
p. 1581 (see for synonymy and additional refer-
ences ) .
Description. None required here; note
form c. as in Helluoniditis; color black or
dark brown; genae prominent; labrum
wider than long, obtusely angulate, 4-setose;
ligula narrower than usual in tribe, nar-
rowed anteriorly, narrowly rounded at apex;
length c. 12.5-15.0 mm.
Type. From Rockhampton, Queensland,
Australia; present location unknown (not
found at Melbourne in 1958).
Occurrence in New Guinea. Papua: 3,
Rouku, Morehead R., Apr. 1962 {\\\ W.
Brandt, C.S.I.R.O.); 7, Port Moresby and
vicinity, various dates and collectors ( Dept.
Agr. Port Moresby; A.M.N.H.); 1, Bisia-
numu, Sogeri Subdistrict, c. 1600 ft. (485
m). Mar. 1955 (J. J. H. Szent-Ivany and J.
McAdam, Dept. Agr. Port Moresby).
Notes. The relatively narrow ligula is
diagnostic of this species in this tribe in
New Guinea. I find no significant dif-
ferences between specimens from Australia
and from New Guinea, although individual
variation occurs in both places.
Genus HELLUODEMA Castelnau
Castelnau 1867, Notes on Australian Coleop., p. 19.
Sloane 1914, Proc. Linnean Soc. New South Wales
39, pp. .571, 586.
Csiki 1932, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1581 (see for .synonymy and additional refer-
ences ) .
Diapiosis. See preceding Key to Genera.
Description. None required here.
Type species. HeUuomorpha hatesi Thom-
son {= tinicolor Hope), of Australia (see
below ) .
Generic distribution. Eastern and north-
ern Australia, New Guinea.
Notes. Two species of this genus occur
in Australia, one of them extending to New
Guinea.
Helluodema un/co/or (Hope)
Hope 1842, Proc. Ent. Soc. London for 1842, p. 47
( Aeni<itna).
Sloane 1914, Proc. Linnean Soc. New South Wales
39, p. 587.
Csiki 1932, Coleop. Cat., Caral)idae, Harpalinae 7,
p. 1581 (see for synonymy and additional refer-
ences ) .
Description. None required here; note
fonn (Fig. 141) slender; genae prominently
rounded; prothorax only moderately (not
strongly) constricted before base; labrum
wider than long, obtusely angulate, 4-setose;
ligula broadly rounded, obtusely emargi-
nate at apex; length c. 13-15 mm.
Type. From Australia; present location
unknown.
Occurrence in New Guinea. Papua: 3,
Rouku, Morehead R., Apr. 1962 {W. W.
Brandt, C.S.I.R.O.). West N. G.: 2,
Merauke, sea level. Mar. 24, 28, 1955 ( L. D.
Brongersma, Leiden Mus.), evidently taken
in light trap.
Notes. This species occurs in eastern
Australia at least from northern New South
Wales to Cooktown. The New Guinean
specimens agree well with Australian ones.
Genus GIGADEMA Thomson
Thomson 1859, Arcana Naturae, p. 93.
Sloane 1914, Proc. Linnean Soc. New South Wales
39, pp. 572, 593.
234 Bulletin Museum of Comparative Zoology. Vol. 137, No. 1
Csiki 1932, Coleop. Cat., Caiabidae, Harpalinae 7,
p. 1582 (see for synonymy and additional refer-
ences ) .
Diapwsis. See preceding Kcij to Genera.
Deseription. None required here.
Type .species. G. titanum Thomson ( =
nocte Newman), of Australia.
Generic distribution. Australia; south-
ern New Guinea.
Notes. This is the principal genus of the
tribe in Australia. One of the 12 or more
Australian species is now recorded from a
single locality in southern New Guinea,
almost opposite the tip of the Cape York
Peninsula.
Gigadema maxiilare Sloane
Sloane 1914, Proc. Linnean Soc. New South Wales
39, pp. 595, 599.
Description. None required here; note
large size; prothorax constricted at base;
color dark; surface short-pubescent; labrum
c. long as wide, rounded, 2-setose; length
of Australian specimens 27-35, of New
Guinean specimen 32 mm.
Types. From tropical Queensland, Aus-
tralia: Townsville, Kuranda, Cooktown,
Princess Charlotte Bay. I here designate
as lectotype a specimen labeled "Cktn., Q.,
Olive, i " and "Giiiadcmo nuixillare SI., Id.
by T. G. Sloane"; in Sloane Coll., C.S.I.R.O.,
Canberra ( seen ) .
Occurrence in New Guinea. Papua: 1
9 , Rouku, Morehead R., Apr. 1962 (W. W.
Brandt, C.S.I.R.O.).
Notes. The Papuan 9 agrees with Aus-
tralian examples of maxillare in nonsexual
characters, but a ^ is needed to confirm
the identification. Another specimen ap-
parently of nuixillare, but also a 9 , is
before me from Mona Is., Torres Straits
(J. VV. Schomberg, S. Australian Mus.).
Tribe BRACHININI
Brachijitiiii CIsiki 1932, (Coleop. Cat., (;aral)ida(\
Harpalinae 7, p. 1593 (sec for synonymy and
additional references ) .
Jedlicka 1963, Ent. .AbhanilliniHcn 28, p. 524.
Bnichinidac Jearniel 1942, Faime de France,
Coleop. Carabiques, I'art 2. p. 1102.
1949, Coleop. Carabiques de la Region
Malgache, Part 3, p. 1079.
BrachUdnac Basilewsky 1953, Exploration Pare
National I'Upemba, Fasc. 10, Carabidae, p. 235.
Habu 1967, Fauna Japonica, Carabidae, Trun-
catipennes Group, p. 280.
The beetles of this tribe are bombardiers
(but are not the only Carabidae that "shoot"
repellents) and are well known to most
entomologists in most parts of the world.
Their form is characteristic, and identifica-
tion is confirmed by presence of S visible
ventral abdominal segments.
Two genera of the tribe are very widely
distributed, and these are the only genera
that reach New Guinea. Fhcropsophus,
which occurs (discontinuously ) in all prin-
cipal tropical regions, has 6 New Guinean
species of which 5 are endemic and 1 shared
with Australia. Brachinus, which is almost
worldwide except that it does not reach
Australia, has 1 New Guinean species which
is endemic but which is the easternmost
member of an Oriental species group.
Key to Geneh.a of Br.\chixixi of New Guixe.\
1. Elytra with costae strong, distinct, and
separate to apex (p. 234) PJwropsopiuts
- Elytra with costae weaker and ])econnng
faint and in part vaguely connected before
ape-x (p. 239) Brarhiitus
Genus PHEROPSOPHUS Soiier
Solicr 1833, Ann. Soc. Ent. France 2, p. 161.
Csiki 19.32, Coleop. Cat., Carabidae, Harpalinae 7,
p. 1595 (see for subgenera and ad(h'tional rcler-
ences ).
l\ira})hcropn(>]>h\is lliilxMitiuil 1911, Diiitsche \\\\[.
Zeitschritt for 1914. pp. 140. 442 (new s>-n()n-
yniy).
Csiki 1933, Coleop. Cat.. Carabidai'. Harpalinae 8,
p. 1604 (as subgenus of /'/icro/wo/^/n/.s).
Dia^^nosis. See preceding Key to Genera.
Description. None required here; see
Figures 147-151.
7'(//)<" species. Ol Pli(ropso))hu.s. Brachi-
nus senciialensis Dejean, oi Africa. Of
Paraplieropsoplius, P. intermedins Huben-
tlial {— verticalis Dejean) b\ present des-
ignation.
Generic distribution. All principal trop-
ical and s()m(- wann-teinperate regions of
The Carabid Beetles of New Guinea • Darlington 235
the world. In the Asia tic- Aiistrahan area
species are numerous in southeastern Asia
and the western Malay Archipelago, fewer
eastward, and only I variable species
reaches Australia.
Notes. The supposed "subgenus" Para-
pheropsophus was based on trivial char-
acters, primarily on the shape of the dark
mark between the eyes ( which is varialile )
supported by a supposedly characteristic
habitus which, according to Hubenthal, is
'ieichter zu erkennenden als zu beschrei-
benden." In my opinion all the 4 "species"
and 5 additional "varieties" listed in this
subgenus by Csiki (following Hubenthal)
are forms of a single species {verticals
Dejean) which is not worth subgeneric
separation from PJieropsophus.
The 6 species of Fheropsophus that occur
in New Guinea are ecologically differenti-
ated. P. verficalis is very common in a
variety of wet places and is winged. P.
amnicola has been found only on the banks
of large rivers (Markham and Sepik) and
is always winged. The other 4 species are
rare, local, flightless forms; 1 (cafiilus) is
known to occur in leaf-litter on the ground
in rain forest, and this is probably the habi-
tat of the others too.
Besides the 6 species of Phcropsophus
treated below, I have seen a single speci-
men of javanus Dejean (agnatus Chaudoir)
labeled as from New Guinea ("New Guinea,
Mimika R., A. F. R. Wollaston. 1911-229."
(British Mus.)). This conspicuous species
is common in the western Malay Archipel-
ago but has not been found in New Guinea
by recent collectors and has apparently not
been found in the Moluccas. I think the
specimen in question is probably wrongly
labeled. New Guinea should be deleted
from the range of the species as given by
Csiki (p. 1601) and others.
Key to Species of Fheropsophus of New Guinea
1. Elytra each with 8 costae c. equally promi-
nent 2
- Elytra with odd costae more prominent than
even ones at least at base ,5
2. Strictly bicolored: head and prothorax yellow
or reddish yellow without dark marks, elytra
dark without pale marks (p. 235) _ amnicola
- Not thus bicolored: all or part of pronotum
and part of head dark, elytra dark with or
without pale marks 3
3. Front yellow with isolated usually V-shaped
dark mark between eyes; inner wings large
and folded (p. 236) verticalis
- Posterior half or more of head dark; inner
wings vestigial 4
4. Head bicolored, yellow anteriorly, dark pos-
teriorly; prothorax wider than long at middle
(by measurement); femora not or only
minutely black-tipped; length 8.5-12.5 mm
(p. 237) aptim^iuorpJuis
- Head dark with small V-shaped reddish mark
on front; prothorax as long ( at middle ) as
wide and appearing longer; femora con-
spicuously black-tipped; length c. 18 mm (p.
237) pedes
5. Pronotum (sparsely) pimctate, much rough-
ened at base and apex; length c. 15-16 mm
(p. 238) catuht.s
- Pronotum \irtually impmictate, scarcely
roughened; length 20.5 mm (p. 238) __ cants
Fheropsophus amnicola n. sp.
Description. With characters of genus;
form (Fig. 147) c. as of verticalis; head
and pronotum yellow without dark marks,
elytra black without pale marks, append-
ages yellow. Head 0.92 and 0.93 width
prothorax. Prothorax subcordate; width/
length 1.14 and 1.13; base/apex 0.94 and
1.01; sides broadly arcuate anteriorly,
broadly sinuate before c. right posterior
angles; margins very narrow; disc convex,
irregularly subpunctate and punctulate.
Elytra moderately narrowed anteriorly;
width elytra/ prothorax 1.64 and 1.66; each
elytron with 8 well defined costae (in-
cluding raised suture), the costae equally
elevated and equally prominent at base;
surface of costae finely microreticulate, in-
tercostal intervals longitudinally roughened.
Inner icings full. Measurements: length c.
8.5-15.0; width 3.2-5.3 mm.
Types. Holotype c^ (M.C.Z., Type No.
31,525) and 34 paratvpes from vie. Nadzab,
N-E. N. G., July 1944 (Darhngton); and
1 paratype. Main R., Sepik, N-E. N. G.,
Feb. 1965 (R. Hornabrook).
Measured specimens. The S holotype and
1 9 paratype from Nadzab.
236 Bulletin Museum of Comparative Zoolop^y, Vol. 137, No. 1
Notes. The preceding description con-
tains all the characters that now seem worth
specifying for this new species, which is
distinguished from ceiiicaJis primarily by
color. (Jolor, pioperly understood and with
allowance for variation, is in fact specific
in this genus. The geographic and ecologic
restriction of amtiicola. as compared with
verticalis\ is another indication that the two
species are fully distinct. My specimens
were all taken on thi' banks of the Markham
R., and none was found in anv other situa-
tion.
Pheropsophus verticalis Dejean
Dfjfan 1825, Spitit-s (ieneral Coleop. 1, p. 302.
Csiki 19.33, Coleop. Cat., Caral)idae, Harpalinae
8, p. 1604 (see for Australian "varieties" and
additional references ) .
atistrali.s Castelnau 1867, Notes on Australian
Coleop., p. 23 (new synonymy).
papucmis Maeleay 1876, Proc. Linnean Soc. New
South Wales 1, p. 166 (new synonymy).
Heller 1910, AbhandhuiKen und Berichte Zool.
VIus. Dresden 13, No. 3, p. 7.
luaclcayi Sloane 1894, Proe. Linnean Soe. New
Soutli Wales (2) 9, p. 4.53 (new synonymy).
haliothonix Heller 1910, Abhandlunuen und
Berichte Zool. Mus. Dresden 13, No. 3, p. 6
( new synonymy ) .
intermt'dius Iluhentiial 1914, Deutsclit' Ent.
Zeitschrift for 1914, p. 440 (new synonymy).
Deseription. None required here. This is
the only New Ciuinean Plwropsophiis with
an isolated black (or brown) frontal spot.
See also under genus (above) and Notes
(below); length 8.5-16.5 mm.
Types. Of verticalis Dejean, from
"Nouvelle Tlollande" (= Australia ); in
Oberthiir Coll., Paris Mus. Of (tiislralis
Castelnau, from Hockhampton, Queensland,
Australia; present location unknown. Of
ixipnemis Maeleay, from Katow , Papua;
presumably in Maeleay Mus., Sydney. Of
nuirlediji Sloane, from King's Sound, NW.
Australia; lectotype not designated. Of
haliolliorax Heller, from l''iiischhafen, N-K.
N. (i.; in Dresden Mus. Of intermedins
Ilubenthal, from Neu Itrilain; in Hcrlin
Zool. Mus. (Of all these types, I have seen
only some cotxpes of niaeleai/i. )
Occurrence in New Cui)\ea. Coinmou
throughout New Guinea: 215 specimens;
most from low altitudes, only 2 individuals
from above 1000 m; common at Dobodura.
Notes. The synonymy proposed abo\'e is
based not on comparison of types but on
examination of much material from many
localities in Australia as well as New
Ckiinea, New Britain, and some other islands.
As a result of it I have concluded that, in
the area in question, all the Fhcropsophus
with an isolated dark frontal spot belong
to one variable species, verticalis Dejean,
which ranges over the whole of Australia
and New Guinea and extends to New
Britain, New Ireland, the Solomons, and
perhaps other islands.
In reaching this conclusion, I first con-
sidered the supposed separate northern
Australian species, maclcayi Sloane. Of it,
Sloane had only 3 specimens, which hap-
pened to be rather small ( 11.5-13 mm) and
to share some minor peculiarities of form
and markings including presence of yellow
shoulder spots and elytral fasciae of con-
stant shape. My material from subtropical
and tropical northern Australia shows that
these characters are in fact indixidual rather
than si)ecific. For example, 6 specimens
from the Blackall Range, in subtropical
South Queensland, \ary in size from 11.5
to 14.5 mm ( to apex of {>K'tra ) and vary
also considerabl) in lorm and somewhat in
markings, although none has shoulder
spots; and 4 specimens from mid-j:)eninsular
C>ape York (Co(mi and Silver Plains) are
large (e. 14.5-16.5 mm) and \ary in exact
form of prothorax, in prominence of humeri,
and in markings: e.g., 2 have and 2 have
not Ncllow shoulder spots.
The New Guinean individuals that I in-
clude in verticalis vary in details of form,
especially in prominiMu-e of humeri. 4'he\-
\ary in si/.r Irom e. S.5 to 16.5 mm. And
tlu'v N'ary in markings: the trans\(M-se usu-
all\ \'-shaiied dark mark between the eyes
is relatixcK constant but is soiiictimes
slighth extentled posteriori) ; the pronotum
\aries Irom wholly dark to broadl\- reddish
Ncllow with ()nl\- the margins dark (inter-
The Carabid Beetles of New Guinea • Darlington 237
mediates are common ) ; and the elytra vary
from \\'holly dark to conspicuously marked,
with median fascia often present but vari-
able (but rarely large), shoulder spots
sometimes present (distinct in only 2 in-
dividuals, from S. Highlands and Popon-
detta, and vestigial in a few other individ-
uals), and apices sometimes with (variable)
yellow margins. Although much of this
variation is surely individual, some of it is
or may be geographic. For example. New
Guinean specimens usually have the elytra
less heavily spotted than Australian speci-
mens, although extremes overlap. But I
think nothing is to be gained by recognizing
subspecies now. The variations of this
species should first be analyzed statistically,
in detail, using series of specimens from
exact localities, not just the New Guinean
against Australian specimens. This will be
third stage taxonomy (see Part I of my
■'Garabid Beetles of New Guinea," p. 329),
far beyond what I can attempt now.
The wings of vcrticalis are fully de-
veloped, or at least large enough to be
strongly folded at apex, in all my Australian
specimens and all New Guinean specimens
that I now assign to this species. However,
occasional short-winged individuals occur
that may prove to be mutants of veiticalis
although I am tentatively treating them as
a separate species, ciptinomorphtis Heller
( below ) .
P. vcrticalis is common in a variety of
wet places. Although all individuals are
winged, they may not often fly and are
not often taken in light traps. Observations
on their flight would be interesting.
The great variation in size of adults sug-
gests that the larvae may be parasitoid, per-
haps feeding on pupae of other beetles, as
some other members of the tribe are known
to do.
Pheropsophus apfinomorphus Heller
baliothorax var. apfiiH>iuori)}ui.s Heller 1910,
Abhandlun^en unci Berichte Zool. Mus. Dresden
13, No. 3, p. 7.
Description. Form as in Figure 148;
similar to vcrticalis (above) but elytra
more narrowed to base; head bicolored,
yellow anteriorly, dark posteriorly; elytra
not marked (in the few specimens seen);
inner wings vestigial, reduced to thin strips
less than Vj long as elytra; length (to apex
of elytra) c. 8.5-12.5 mm.
Type. From New Guinea, exact locality
not given, but altitude stated as 120 m; in
Dresden Mus. (not seen).
Occurrence in Netc Gtiinca. N-E. N. G.:
1, Aitape, Aug. 1944 (Darlington). West
N. G.: 1, Waris, S. of Hollandia, 450-500
m, Aug. 16-23, 1959 (T. G. Maa, Bishop
Mus.); 1, Maffin Bay, Aug. 1944 (Darling-
ton); 1, same locality, June 15, 1944 (E. S.
Ross, Galifornia Acad. ) .
Notes. Heller does not say whether the
type is winged, but the 4 specimens listed
above answer his description in color of
head (which is diagnostic), and their form,
with narrowed humeri, does recall A))tinus.
If it were not for the different head
marking, I would consider my specimens of
aptinomorphus to be short- winged mutants
of vcrticalis. The distribution of the short-
winged indi\iduals, widely scattered and
occurring with vcrticalis (which I have
from Waris and Maffin Bay and which
probably occurs at Aitape too), would be
consistent with their being mutants. But
if they are mutants, then the mutant gene
apparently must modify color of head as
well as length of wings and form of elytra,
and I do not dare assume that this is the
case without further evidence.
Pheropsophus pedes n. sp.
Description. With characters of genus;
form as in Figure 149, with relatively long
narrow prothorax and elytra ample but
strongly narrowed to base; black, head with
irregular red marks including U-shaped
mark between eyes, appendages reddish,
femora broadly tipped with black. Head
0.96 width prothorax; eyes normal; 1st
antennal segments swollen; labrum semi-
circular; front 2-impressed anteriorly, with
surface finelv c. isodiametricallv reticulate.
238
BuUciin Museum of Comparotivc Zoolomj, Vol. 137, No. 1
Profhomx long; width/length 1.01; base '
apex 1.06; margins narrow, each with 1
principal seta behind middle of length; disc
moderately conxex, ^^'ith scattered punc-
tures (the punctures with setae as usual in
genus), strongly roughened at base, less so
apically, and with disc finely irregularly
microreticulate. Elytra: width elytra/pro-
thorax 1.95; each elytron with 8 costae (in-
cluding raised suture) c. equalh' dexeloped;
siuface of costae finely irregularh' micro-
reticulate, intercostal intervals longitu-
dinally roughened. Inner aings evidently
atrophied. Measurements: length LS; width
6.5 mm.
Ti/pe. Ilolotype 9 (Bishop Mus.) from
Bomberi, Vogelkop, West N. G., 700-900
m, June 7, 1959 (T. C. Maa); the type is
imique.
Notes. I do not know the relationships
of this obviously distinct species. Char-
acters distinguishing it from other New
Ouinean Plier(>})so})1ius are given in th(> pre-
ceding Key tu Speeies.
Pheropsophus cafulus n. sp.
Deseription. With characters of genus;
form as in Figure 150, with elytra strongly
narrowed to base; brownish black, head
variably red-marked anteriorly, appendages
reddish'. Head 0.91 and 0.90 width pro-
thorax; eyes moderate; 1st antennal seg-
ments slightly swollen; labrmn transverse,
slightly prominent at middle; front 2-im-
pressed anteriorly, with surface finely ir-
regularly microreticulate, roughened jios-
teriorh. Prothorax (juadrate-subcordate;
width length l.OI and 1.07; ba.se/apex 1.11
and 1.13; sides weakly arcuate anteriorly,
broadly sinuate before basal angles; latter
acute except bhmted, slightK produced
posteriorly; margins moderate, each with
one principal seta behind middle of k'ugth;
disc irregularly convex, irregularh rough-
ened especially posteriorly and anteriorly,
and surface also with scatteri'd punctures
(with hairs as usual) and irregularly faintly
microreticulate. Elytra: width elytra pro-
thorax 1.72 and 1.74; each cKtron with S
costae (including raised suture), even
costae stronger than odd ones and reaching
base; surface of costae faintly finely micro-
reticulate, intercostal intervals finely longi-
tudinally roughened, and alternate intervals
each with row of widely spaced setae.
Inner wings reduced to vestiges that hardly
extend beyond edge of metathorax. Mea-
surements: length e. 15-16; width 5.0-5.6
mm.
Types. Holotype i (M.C.Z., Type No.
31,526) and 2 9 9 paratypes all from
Dobodura, Papua, Mar.-July 1944 (Dar-
lington ) .
Measured specimens. The i holotype and
1 9 paratype.
Notes. I do not know the relationships of
this very distinct species, which is quite
different from any of the preceding ones
although closely related to the following
(canis). The beetles were taken among
dead leaves on the ground in rain forest.
Pheropsophus canis n. sp.
Description. With characters of genus;
form as in Figure 151, with elytra strongly
narrowed to base; brownish black, head red
anteriorly dark posteriori)-, pronotum with
faint reddish marks, legs reddish yellow,
antennae brown. Head 0.S7 width pro-
thorax; eyes moderate; 1st antennal seg-
ments slightly swollen; labrum transverse,
with margin broadh' rounded; front weakh'
2-impressed, closely irregularly microreticu-
late, roughened posteriorly. Prothorax nar-
rowK' subcordale; width length 1.07; base '
apex 1.06; sides broadU' arcuate anteriorh,
broadly sinuate before slightly obtuse,
slightly blunted, posterior angles; latter not
produced posteriorly; margins moderate,
each with 1 principal s(>ta behind middle;
disc moderately conxcx. weakh roughened
posteriorly but otherwise uearK' smooth,
surlace w ith Wwc irregular reticulate" micro-
sculpture. Elytra: width eKtra i^rothorax
l.(S(); each eKtron with 5 conspicuous costae
(nos. 1, 3, 5, 7, (S), the intermediate costae
(nos. 2, 4, 6) weak or obsolete; surface^ of
costae lineK' irreirularK microreticulate. iu-
The Carabid Beetles of New Guinea • Darlington 239
tercostal intervals finely roughened; even
intervals probably with widely spaced setae,
but latter in part broken oft or missing.
Inner wings evidently vestigial. Measure-
ments: length c. 20.5; width 7.6 mm.
Type. Holotype 9 (Hawaiian Sugar
Planters Association) from Koitaki, Papua,
1500 ft. (c. 460 m), Nov.-Dec. 192(S ( Pem-
berton Coll.); the type is unique.
Notes. This flightless species is probably
a geographic representative of the pre-
ceding (eatuhis) but is larger, slightly dif-
ferent in proportions, with antennae darker,
pronotum much less roughened, and even
intervals of elytra much more reduced.
Other, related forms of this flightless group
are to be expected elsewhere in New
Guinea.
Genus BRACHINUS Weber
Weljer 1801, Observationes Entomologicae, p. 22.
Bidclujnus Auct. including Csiki 1933, Coleop.
Cat., Carabidae, Haipaliiuu' (S, p. 1606 (see for
additional references ) .
Diagnosis. See preceding Key to Genera.
Description. None required here.
Tyi)e species. Carahus crepitans Lin-
naeus, of Europe.
Generic distribution. Most of the world,
except Australia. In the Indo-Australian
area, many diverse species occur in India,
etc.; fewer, in the western part of the Malay
Archipelago; and a single species occurs
in New Guinea.
Brachinus papua n. sp.
Description. Form as in Figure 152; dark
brown or brownish black, head red anteri-
orly or with 2 red spots between eyes,
pronotum sometimes vaguely reddish, ap-
pendages reddish with tibiae, tarsi, and
apices of femora darker brown; dull, entire
upper surface with fine, c. isodiametric but
irregular reticulate microsculpture, and
much of upper surface with inconspicuous
fine pubescence, often in part rubbed away,
and perhaps missing on front, part of pro-
thoracic disc, and disc of elytra. Head 1.01
and 0.99 width prothorax; eyes moderate,
genae oblique, setose; front longitudinally
impressed each side, nearly smooth or
slightly roughened and punctulate ( vari-
able), with head more roughened posteri-
orly. Prothorax subcordate (exact form
variable); width/length 1.07 and 1.07; base/
apex 1.03 and 1.01; margins moderate, each
with apparently 1 principal seta near middle
of length; disc weakly convex, slightly finely
transversely wrinkled, irregularly punctu-
late, slightly longitudinally roughened at
base and apex. Elytra ample; width elytra/
prothorax 2.34 and 2.33; intervals slightly
raised but not costate. Inner icings full.
Measurements: length 17.5-19.0; width
7.3-8.0 mm.
Types. Holotype c5 (M.C.Z., Type No.
31,527) from Hollandia, West N. G., 250
ft.. May 1945 (H. Hoogstraal); 1 paratype,
same locality, Apr. 1945 (B. Malkin,
U.S.N.M.); 2 paratypes, Tanahmerah, Res.
Boven Digoel, West N. G., Feb. 1958
( R. T. Simon Thomas ) ; and 1 paratype,
Fenichel (Hungarian National Mus.).
Measured specimens. The S holotype and
1 9 paratype from Tanahmerah.
Notes. This, the first known New
Guinean Brachinus, may represent higut-
ticeps Chaudoir of Java, etc., but papua is
larger than my specimens of bigutticeps,
with less sharply bicolored legs and less
roughened pronotum. Further study is
needed to clarify the relationships of these
and related species in the Malay Archipel-
ago.
Tribe PSEUDOMORPHINI
Csiki 1933, Coleop. Cat., Carabidae, Harpalinae 8,
p. 1634 (see for synonymy and additional refer-
ences ) .
Pseiidomorphidae Auct. including Notman 1925,
Proc. United States National Mus. 67, Art. 14,
p. 1.
Pseudomorphini (Figs. 153-159) do not
look like Carabidae but superficially re-
semble dytiscids or scolytids or Crypto-
cep]}alus-\ike chrysomelids. They are nu-
merous onlv in Australia; a few small
species occur in New Guinea; a species of
240 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
the Australian and New Guinean genus
Adelotopus has lieen found m Java; one
genus, Cnjptocephalomorpha, ranges from
New Guinea aeross the Malay Archipelago
to the SE. corner of Asia; and a supposed
endemic genus is localized on New Cale-
donia. Outside this area the tribe contains
only a single genus, Fseudomorpha, con-
fined to the Americas and ranging from
-southern United States to Brazil and
Argentina/
Although I have only 15 specimens of
this tribe from New Guinea, they include 3
genera and 7 species, and all the species are
different from the single pseudomorphine
{Adelotopus papuanus) previously known
from the island. ( An ^^ Adelotopus sp.^' listed
from New (Guinea by Heller, in Abhand-
lungen und Berichte Zool. Mus. Dresden
13, 1910, No. 3, p. 4, has not been identi-
fied.) Because my material is very limited
and because the species are well defined
by easily seen characters, I shall treat the
members of this tribe rather superficially,
leaving dissection of the mouthparts, etc.,
to the next reviser of the tribe as a whole.
Notman's ( 1925 ) keys to the Australian
species of this tribe are very useful but,
because they are based largely on old de-
scrij^jtions rather than on specimens, the\'
should be used with caution.
Most Australian Pseudomorphini live on
the tnuiks of trees, especially on the shaggy
trunks of Euealyplus. They are v(My active,
winged beetles. I do not know tiie habits
ol the New Guinean h)rms.
' The lollowiiiji; notes arc necessary to justify
my siininiar\ of [\\c distrihiition ot this zoot^t'o-
.Uraphically iiilcrcstinu tribe. The African Ihj-
ilroporoDiorplui has only 1 entire \entral abdominal
sesnients; it is not a pscucloniorphine but probabK
a harpaline. Paus.sotwpus is Australian and jirob-
ably does not occur on Batchian Is. ( \otnian
1925, p. 5, footnote; conlirnied 1)\ nie at the
British Mus. in 1948). And SilplioiiioiplKi
cntiaroidcs (Newman), listed from Oceania 1)\
Csiki (1933, p. 1639), is probably really Aus-
tralian; Newman kIvcs no localit\' except "Its
habitat is 3753," but lie refers to the inseit as
''this jircttx' antipodean.
Key to Gener.\ of Pseudomorphini of
New Guinea
1. Eyes superior in position, not interrupting
lateral margins of heatl (p. 240) _ _ Adelotopus
- Eyes lateral in position, broadly interrupting
lateral margins of head __ 2
2. Form wider, more depressed; head not
strongK deflexed, labrum and mandibles
\isible from in front (p. 242) __ SpliaJIoiuorpha
- Form subc\lindric; head sti"ongl\ deflexed,
labrum and mantlibles ( except sometimes
their tips ) not \ isible from in front ( p.
242 ) Cnjptocephdiontorplia
Genus ADELOTOPUS Hope
Hope 1834, Trans. Ent. Soc. London 1, p. 11.
Notman 1925, Proc. United States National Mus.
67, Art. 14, pp. 5, 6.
Csiki 1933, Coleop. Cat., Carabidae, Harpalinae 8,
p. 1634 (see for additional references).
Diciii^nosis. See preceding Key to Genera.
Description. None required here.
Type species. A. '^yrinoidcs Hope, of
Australia.
Generic distribution. Australia including
Tasniania (many species). New Guinea
( 4 species ) , and Java ( 1 species ) .
Notes. A. jaeohsoni Ritsema ( 1909, Notes
Leiden Mus. 31, p. 255) of Java evidently
realK' is an Adelotopus, not a Cryptoeephalo-
morpha, for Ritsema knew both genera.
Key to SrEf:]Ks of Adei.otoi'I s of New GnNF..\
!. Piceous, with narrow reddish tianslucent
margins i)ut otherwise unmarki'd; length 6.6—
7.0 nun (p. 240) exactor
- Black, elytra marked or tipped with red or
\('llow ; smaller 2
2. Ehtra with red apices (p. 241) debitor
- Ehtra with xellow or reddish marks near
1 )ase 3
3. i'^K tra w itli wide basal fascia rufo-teslaceous;
sfutelluni \ cllow-niartiined (p. 241) pa]>uami:<
- f'Ktra eacli witii large sni)l)asa! ncHow spot;
stiitelhini not \ ellow -inartiined (p. 211) -
bijtigu.s
Adelotopus exacfor n. sp.
Deseripiioii. With characters of genus;
lorni ;is in I'^igine 15'3, inodiTately convex;
piceous. unmarked except i^rothorax and
el\ tra narrow 1\- reddish translucent at sides,
lower surlace red; no dorsal pubescence or
The Carabid Beetles of New Guinea • Darlinaton
241
setae except on humeral margins; not
shining, entire upper surface with tine c.
isodiametric to sHghtly transverse micro-
sculpture and very fine well spaced punc-
tulation. Head 0.54 width prothorax. Pvo-
thorax: width length 1.98; base/apex 1.84;
margins moderate, moderately reflexed.
Elytra: width elytra prothorax 1.07; basal
marginal line sharply impressed, nearly
entire, not punctate; posthumeral impres-
sions weak; margins moderate; striae not
indicated. Measurements: length 6.6-7.0;
width 3.5-3.7 mm.
Types. Holotype, sex not determined (S.
Australian Mus.) and 1 paratype (M.C.Z.,
Type No. 31,528) both from Wareo,
Finschhafen ("Finsch Haven"), N-E. N. G.
(Rev. L. Wagner).
Measured specimen. The holotype. The
paratype has the pronotum broken and can-
not be satisfactorily measured.
Notes. In Notman's ( 1925 ) key to Aus-
tralian Adclotopus this runs to hydrohioides
Westwood, but comparison with Australian
specimens identified as hydrohioides shows
that exactor is narrower, with narrower
prothoracic and elytral margins.
Adelotopus debitor n. sp.
Description. With characters of genus;
form as in Figure 154, very convex espe-
cially elytra; black, apical % of elytra red,
lower surface black anteriorly, red posteri-
orly, appendages red; no dorsal pubescence
or setae except on humeral margins; shining,
fine isodiametric microsculpture distinct on
head, indistinct on pronotum, absent on
elytra, but whole upper surface with fine
moderately spaced punctulation. Head 0.54
width prothorax. Prothorax: width length
2.18; base apex 1.76; margins moderate,
poorly defined, moderately reflexed espe-
cially anteriorly. Ehjtra: width elytra/pro-
thorax 1.04; basal margin irregular or obso-
lete on c. inner V2 of width of elytron; sides
scarcely impressed behind humeri; margins
moderate; striae not indicated. Measure-
ments: length 5.5; width 2.8 mm.
Types. Holotype, sex not determined
(Bishop Mus.), from Wau, Morobe Dist.,
N-E. N. (;., 1200 m, Jan. 16, 1961 ( Sedla-
ceks); 1 paratype (M.C.Z., Type No.
31,590), Kokoda-Pitoki, Papua, 450 m,
Mar. 24, 1956 (Gressitt).
Notes. In Notman's (1925) key to the
Australian species of Adelotopus, this runs
to apicahs Macleay or possibly haemor-
rhoidalis Erichson, but comparison with
Australian specimens of these species shows
that debitor is broader anteriorly and more
narrowed posteriorly.
Adelofopus papuanus Gestro
Gestro 1893, Ann. Mns. Civ. Genoa 33, p. 287.
Notman 1925, Proc. United States National Mils.
67, Art. 14, p. 8.
Description ( significant details from
Gestro). Form perhaps similar to other
New Guinean species of genus; black, pro-
thorax with reddish margins, elytra with
wide basal fascia reddish testaceous and
scutellum with testaceous margin; shining,
very finely punctulate; length 4.75 mm.
Type. From "Ighibirei, lungo il Kemp
Welch"; in Genoa Mus. (not seen).
Notes. This species seems surely different
from any New Guinean Adelotopus I have
seen. Gestro compared it only with A.
himacuhitus Macleay of Australia.
Adelotopus bijugus n. sp.
Description. With characters of genus;
form as in Figure 155, very convex; black,
elytra each with large yellowish plagia
before middle, lower surface mainly dark,
some ventral segments narrowly yellow at
apex, appendages rather dark; no dorsal
pubescence or setae except on humeral
margins; shining, reticulate microsculpture
faint or absent but whole upper surface
rather closely conspicuously punctulate.
Head 0.55 and 0.57 width prothorax. Pro-
thorax: width length 1.72 and 1.84; base/
apex 1.71 and 1.60; margins moderate, well
reflexed, not sharply defined inwardly.
Ehjtra: width elytra prothorax 1.01 and
1.02; basal margin irregular or obsolete in
inner V2 of width of elytron; sides slightly
242 Bulletin Museum of Comparative Zoolofiij. Vol. 137, No. 1
impressed behind humeri; margins mod-
erate anteriorly, obsolete posteriorly; striae
not indicated on elytral surface but vaguely
suggested under the surface in the pale
areas. Measurements: length 4.4-5.2; width
2.2-2.5 mm.
Types. Holotype, sex not detennined
(Bishop Mus.), from Wau, Morobe Dist.,
N-E. N. G., 1200 m, May 25, 1962 ( Sed-
lacek), and paratvpes as follows: 1, Wau,
1200 m, Dec. 1, 1961; 1, Wau, 1200 m, Dec.
IS, 1961; 1, Wau, 1250 m. Mar. 27, 1964;
1, Mt. Missim (near Wau), 980-1100 m,
Aug. 14, 1964; all specimens collected by
the Sedlaceks; 2 paratvpes now in M.C.Z.
(Type No. 31,529).
Notes. In Notman's (1925) key to the
Australian species of Adelotopus, this runs
to ])ini(ieul(itus Macleay but has wider pro-
thoracic and elytral margins than my Aus-
tralian himaculatus.
Genus CRYPTOCEPHALOMORPHA Ritsema
Ritsema 1875, Tijdschrift voor Eiit. 18, Verslaj^, p.
XCII.
Notman 1925, Pioc. Ihiited .States National Mus.
(i7. Art. 14, pp. 5, 12.
Diafinosis. See preceding Key to Genera.
Description. None re(|uired here.
Type s])ecies. C. g.averei Ritsema, of
Java, etc.
Generic distribution. Prexiouslv known
from ThailaiHl, Sumatra, Java, Borneo
(British Mus.), and Phili|>|>iMe8 (Luzon,
in M.C.Z. ); now recorded bom New
(iuinea.
Notes. This genus of very small scoKtid-
likc I^seudomorphini is the only genus of
the tribe known to be w idel\ distiibuted in
the Mahi\' Archijic lago and the onl) genus
known to reach the Asiatic mainland.
Crypfocephalomorpha papua n. sp.
Description. With characters ol genus;
lonn as in Figure 156, subcylindric; entirely
slightly irregular reddisli brown; no dorsal
pubescence or setae; reticulate microsculp-
tiuc indistinct or absent but whole upper
surface with fine moderately spaced punc-
tulation. Head 0.72 and 0.71 width pro-
thorax. Profhorax: width length 1.41 and
1.49; base apex 1..37 and 1.41; margins very
narrow. Elytra: width elytra prothorax
1.00 and 1.00; basal margin apparently
absent ( possibly hidden by base of pro-
thorax); side margins very narrow; striae
not indicated. Measurements: length 3.0-
3.3; width 1.4-1.5 mm.
Types. Holotype, sex not determined
(British Mus.) and 1 paratype (M.C.Z.,
Type No. 31,530) both from Kokoda,
Papua, 1200 ft. (c. 370 m), Sept. (holotype)
and June (paratype) 1933 (Cheesman),
the paratype taken at light.
Notes. The plain, unspotted coloration
distinguishes this from the 2 previously
known species of the genus, gaverei Ritsema
and collaris Waterhouse, both of which
occur farther west in the Malay Archipel-
ago.
Genus SPHALLOMORPHA Westwood
Westwood 1841, Trans. l,innt>an Soc. London 18,
p. 414.
Notman 1925, Proc. United States National Mus.
67, Art. 14, pp. 6, 25.
Csiki 1933, Coleop. Cat., Carabidae, Harpalinae 8,
p. 1641 (sec for additional references).
Diagnosis. See preceding Key to Genera.
Description. None required here.
Ty)>e sjx'cic.s. S. decipiens Westwood. of
Australia.
Generic distri])ution. Australia (many
species) and New (Guinea (3 species).
Notes. The follow ing 3 species, like other
members ol the tribe in New (^.uinea, are
probably not directK relati'd among them-
s(4\'c>s but are all independentU related to
Australian sjiecitvs.
Ki;v ro Si'kciks of Spum.lomohi'U i ok
\i \\ (".riM \
1. C^olor patli'ni l-niacuhitf ( el> tra eacii w itii
2 larjie yellow spots) (p. 243) titiadnta
- Color iialtcrn not 4-niaeuIate 2
2. I'lKtra each with a lar^e, irregular yellow
plaiiia (p. 243) _, _ dtipla
I'-Ktra with a counnon heart-shaped >(4I()w
plauia (p. 243) iniita
The Carabid Beetles ok New Guinea • Darlin^,ton 243
Sphallomorpha quadrua n. sp.
Description. With characters of genus;
form as in Figure 157, moderately convex;
brownish black, margins of prothorax and
elytra paler, elytra each with 2 large pale
yellow spots as indicated ( Fig. 157 ) , below
reddish with head darker, appendages red-
dish; no dorsal pubescence or setae except
setae at humeral angles and at posterior
angles of head; dull, entire upper surface
isodiametrically to irregularly microreticu-
late and finely punctulate. Head 0.63 width
prothorax. Trothorax: width length 2.50;
base/apex 1.51; margins narrowly slightly
reflexed. Elytra: width elytra prothorax
1.10; base not margined; sides scarcely
impressed behind humeri; margins not
strongly reflexed; striae not indicated. Mea-
surements: length 5.7; width 3.0 mm.
Type. Holotype, sex not determined
(Chicago Mus.), from Gadaisu, Papua,
Nov. 15, 1917 (J. T. Zimmer); the type is
unique.
^otes. In Notman's (1925) key to Aus-
tralian members of this genus, quadrua
runs to quadrimaculata Macleay, but
quadrua is dull, while the description of
quadrimacuJata calls for a "brilliant shining
l)lack" insect.
The unique type of quadrua was prob-
abl)' taken in a light trap, for many insect
scales are stuck to its surface.
Sphallomorpha dupla n. sp.
Description. With characters of genus;
form as in Figure 158, moderately convex;
black, lateral and basal margins of pro-
thorax and lateral margins of elytra slightly
rufescent, elytra each with large yellow
plagia as indicated ( Fig. 158 ) , lower surface
and appendages reddish or yellow; no
dorsal pubescence or setae except setae at
humeral angles and posterior angles of
head; head and pronotum dull, elytra more
shining, entire upper surface with fine c.
isodiametric microsculpture and faint fine
punctulation. Head 0.58 width prothorax.
Prothorax: width length 2.32; margins
poorly defined, weakly reflexed. Elytra:
width elytra'prothorax 1.10; base not mar-
gined; sides scarcely impressed behind
humeri; margins narrowly reflexed; striae
not indicated on surface. Measurements:
length 5.8; width 3.2 mm.
Type. Holotype, sex not determined
(Bishop Mus.), from Wau, Morobe Dist.,
N-E. N. G., 1500 m, June 15, 1962 ( Sedla-
ceks); the type is unique.
Notes. The color pattern of dupla is
somewhat like that of eohjtnhetoides West-
wood and himaculata Castelnau, but dupla
is much smaller than either of these Aus-
tralian species.
The type of this species too has insect
scales stuck to it and was probably taken
in a light trap.
Sphallomorpha unita n. sp.
Description. With characters of genus;
fonn as in Figure 159, moderately convex;
without pubescence or setae above, except
setae behind eyes; piceous black, margins
faintly paler, elytra with large common
plagia as indicated (Fig. 159), lower sur-
face and appendages irregular dark red-
dish; no pubescence or setae above except
setae at posterior angles of head (ap-
parently not at humeral angles ) ; mod-
erately shining, but whole upper surface
isodiametrically to irregularly reticulate
(finely) and just visibly punctulate. Head
0.54 width prothorax. Prothorax: width/
length 2.47; base/apex 1.72; sides narrowly
moderately reflexed. Elytra: width elytra/
prothorax 1.08; base not margined; sides
slightly impressed behind humeri; margins
narrowly reflexed; striae faintly indicated
especially in pale area. Measurements:
length 5.1; width 3.0 mm.
Type. Holotype, sex not determined
(British Mus.), from Mafulu, Papua, 4000
ft. (1220 m), Jan. 1934 (Cheesman); the
type is unique.
Notes. The marking of U7iita is probably
like that of cordifer Blackbum, of North
Queensland, Australia, but cordifer is c. 8
mm long, against c. 5 mm for unita.
(Received 23 March 1967.)
244 Bulletin Miiscinn of Cowparotivc ZooIofi,ij. Vol. 137, No. 1
Fig, 1, Per/gono rex n. sp., r$ holotype; 2, P. rossi' n. sp., ho'o.; 3, P. dentifer n. sp., holo.; 4, Physo/aefhus covicpps
Andrewes, rj Enarotadi; 5, Omesfes (orfo Andrewes, 9 Hollandia; 6, Dicrochile acuta n. sp., j holo.; I, D. o/fernans
n. sp., c$ paratype, Chimbu Vy.; 8, Microferon/o bare n. sp., i holo.; 9, Chlaenius pan n. sp., 9 poro., Koto Nika;
10, C. olthofi n. sp., 9 pare, Bernhord Camp; II, Oodes nil n. sp., 9 holo.; 12, O. ross/' n. sp,, d holo.; 13, O.
wilsoni n. sp., 9 holo.; 14, O. por n. sp., 6 holo.; 15, O. longior n, sp., i holo.; 16, Diaphoromerus papuellus n.
sp., 9 pare, Kokoda; 17, Lecanomerus angusfi'or n. sp., 9 para., Hollandia; 18, L. latior n. sp., S holo.; 19, Chy-
doeus papua n. sp., 9 para., Mt. Wilhelm; 20, Trichofichnus sfroneoi (Louwerens), 9, Hollandia; 21, T. modus n. sp.,
S holo.; 22, 7. mongi n. sp., 9 holo.; 23, T. delicatus n. sp., 9 para., I. Deslccs; 24, Harpaloxenus fortis n. sp., 9
para., Doboduro; 2S, Lyfer glober n. gen. & sp., c5 para., Finschhafen; 26, Co/eo/issus papua n. sp., 9 para., Hollandia
area; 27, C. ongulofus n. sp., 9 para., Doboduro; 28, H yphaereon levn n. sp., 9 para., Sibil,
The Carabid Beetles of New Guinea • Dorlinatou
245
Fig. 29, Hyphoereon timidus n. sp., i paratype, Dobodura; 29A, same, wing fo same scale, 9 para., Dobodura; 29B,
some, wing to same scale, another 9 para., Dobodura; 30, Egadroma cyc/ops n. sp., 9 holotype; 31, Acupalpus exactus
n. sp., 9 holo.; 32, A. papua n. sp., 9 para., Dobodura; 33, Odonfomosoreus humero/is n. gen. & sp., 9 para., Dobo-
dura; 34, Anaulacus s/omensis Chaudoir, A Geelvink Bay; 35, Sarothrocrepis papua n. sp., i para., Dobodura; 36,
Somofr/chus elevotus (Fabricius), 9 Peleliu Is.; 37, Aristolebia papua n. sp., 9 para., Wau; 38, Lefaio endynomena n.
sp., £ holo.; 39, L. externa n. sp., S holo.; 40, L cordifer n. sp., i holo.; 41, L insularum n. sp., £ holo.; 42,
Lachnodermo ioveolatum Sloone, 9 Goilalo; 43, Sinurus opocus Chaudoir, c5 Woigeu Is.; 44, Stenofe/us spinosus n. sp.,
9 para., Dobodura; 45, Misce/us sibling n. sp., 9 para., Dobodura; 46, Holcoderus elongatus (Saunders), 9 Wau; 47,
M/nuf/iodes mefa///ca n. sp., 6 holo.; 48, M. papuana (Sloane) , i Dobodura; 48A, same, another c5 , Dobodura; 49, M.
lineella (Chaudoir), 9 Morotai Is.; 50, M. queenslandica (Sloane), i Rocky Scrub; 51, M. rossi n. sp., c5 holo.; 52,
M. sedlacekorum n. sp., i holo.; 53, M. subnitens n. sp., S holo.; 54, M. simplex n. sp., 9 holo.
246 Bulletin Museum of Comparative Zoo/ogy, Vol. 137, No. 1
Fig. 55, Mmuthodes sexuolis n. sp., 9 holotype; 56, M. s. s/gnofo n. subsp., .', paratype, Sambeang; 56A, same, $ ,
Wau; 57, M. regularis n. sp., $ para., Dobodura; 58, M. irregularis n. sp., 6 para., Hollandlo; 59, Cofoscopus brun-
neus n. sp., i holo.; 60, C. /ofus n. sp., 9 hole.; 61, C. sidus n. sp., i , Sibil; 62, C. dobodura n. sp., 9, Kiungo; 63,
C. taylori n. sp., 9 para., Mt. Missim; 64, C. rex n. sp., 6 para., Kiunga; 65, Pencalus liguratus Chaudoir, S , Dobo-
dura; 66, Coptodera grossa n. sp., 9 holo.; 67, C. ornatipennis Louwerens, 6 , Dobodura; 68, C. Imeolata Bates, 9 ,
Ore Bay; 69, C. papuella n. sp., i holo.; 70, C. wou n. sp., i para., Wau; 71, Minuphloeus mixtus n. gen. & sp., 9
pare, Wau; 72, Agonochi/o minuthoides n. sp., $ holo.; 73, A. duplicata n. sp., j holo.; 74, A. variabilis n. sp., 9
para., Wagete; 74A, same, 6 hole.; 74B, same, c^ para., Wau; 75, A. expansa n. sp., 9 holo.; 75A, same, c? para.,
Mt. Hagen; 76, A. dorsofo n. sp., ^^ holo.; 76A, same, 9 , Edie Ck.; 77, Oxyodonfus (ripunctafus Chaudoir, 9 , Dobo-
dura; 78, Mochlherus obscurus ISIoane), 9 , Dobodura; 79, Mochfheroides niger Jedlicka, 6 , Cape Gloucester.
The Carabid Beetles of New Guinea • Darlinaton 2A1
Fig. 80, Dolichocfis microdero Andrewes, $ , Dobodura; 81, D. d/sforfa n. sp., $ paratype, Kokodo; 82, D. denfofa n.
sp., 9 para., Dobodura; 83, D. subrofundo n. sp., c5 holotype; 84, D. subquodrofa n. sp., 6 holo.; 85, D. divisa n. sp.,
c5 holo.; 86, Stricklandia lata n. sp., 9 para., Wagete; 87, Pe//ocypas papuo n. sp., S para., Modong; 88, Ce/ae-
nep/ies parallelus Schmidt-Goebel, 9 , Dobodura; 89, Synfomus quodripunctatus (Schmidt-Goebe!), c5 Wau; 90, M/cro/esfes
curfotus n. sp., 9 para., Luzon; 91, M. c/nctus n. sp., S holo.; 92, Apristus biroi n. sp., 9 para., Madang; 93, Plochi-
onus pollens (Fabricius), 9 , Java; 94, Pareno fasciata (Chaudoir), 9 , Mumeng; 95, Anchista binotata (Dejean), 9 , Hogita;
96, Endynomeno pradieri (Fairmaire), 9 , Samoa; 97, Demetrida goroka n. sp., 9 holo.; 98, D. tesselata n. sp., 9 holo.;
99, D. subfenuis n. sp., c^ holo.; 100, D. tenuis n. sp., 9 holo.; 101, D. genicula n. sp., 9 para., Wau; 102, D.
latangula n. sp., 9 para.. Brown R.; 103, D. kokoda n. sp., 9 para., Kokoda; 104, D. forma n. sp., 9 pare, Pindiu;
105, D. rex n. sp., i para., Pindiu; 106, D. nigripes n. sp., 6 holo.; 107, D. vigil n. sp., c5 holo.; 108, D. n/gr/ceps
n. sp., i holo.; 109, D. seticollis n. sp., 9 para., Enarotadi.
248 Bulletin Museum of Comparative Zoology, Vol. 137, No. 1
Fig. 110, Phloeocarabus eup/enes n. sp., S paratype, Kiunga; 111, Trigonothops lateralis n. sp., 9 holotype; 112, No
foforus papua n. sp., 9 para., Dobodura; 113, Anomotarus gress/ff/ n. sp., 9 holo.: 114, A. ocellatus n. sp., 9 holo.;
115, A. plagiler n. sp., 6 holo.; 116, A. fronsversus n. sp., c5 holo.; 117, A. wau n. sp., 9 holo.; 118, Pentagonica
blonde Andrewes, c5 Dobodura; 119, P. papua n. sp., 9 para., Dobodura; 120, Paroscopodes cyoneus (Sloane), c^ Do
bodura; 121, Scopodes altus n. sp., <^ holo.; 122, S. cheesmani n. sp., c5 para., Cyclops Mts.; 123, S. adonis n. sp.,
$ para., Mobitei; 124, Hexogon/o papua n. sp., c5 para., Aitape; 125, Colliuris rossi n. sp., i holo.; 126, C. popuo
n. sp., 9 para., Dobodura; 127, Cosnoidea puncticollis (Sloane), 9, Kiungo; 128, Clarencia quadndens n. sp., 6 para.,
Hollandia; 129, Dicrospedo bispinosa n. sp., 9 holo.; 130, Eudalia anomala n. sp., 9 para., Wasian; 131, Doboduro
armata n. gen. & sp., i para., Dobodura; 132, Drypfo papua n. sp., 9 holo.; 133, Zuphium sinuum n. sp., 9 holo.;
134, Planete: cordons n. sp.,9para., Stephansort; 135, Pogonog/ossus papua n. sp., 9 para., Dobodura; 136, P. mo(or
n. sp., 6 holo.; 137, P. /odor n. sp., 9 para.. Lower Mist Camp; 138, P. obliquus n. sp., 6 holo.
The Carabid Beetles ok New Guinea • Darlington 249
Fig. 139, Pogonoglossus parvus n. sp., $ paratype, Doboduro; 140, P. glabricollis Van Emden, 9, Mofae; 141, Hellu-
odema un/co/or (Hope), c^ , Rouku; 142, Helluonidius chrysocomes Maindron, 9 , Nabire; 143, H. laevifrons n. sp., 9
holotype; 144, H. latipes n. sp., 9 hole.; 145, H. politus n. sp., 9 holo.; 146, Helluopapua toxopei n. gen. & sp., $
holo.; 147, Pheropsophus amnicola n. sp.; 148, P. aptinomorphus Heller, S Maffin Bay; 149, P. pedes n. sp., 9 holo.;
150, P. catulus n. sp., $ holo.; 151, P. cam's n. sp., 9 holo.; 152, Brachinus papua n. sp., 9 para., Tanahmerah;
153, Adelotopus exactor n. sp., holo.; 154, A. debitor n. sp., holo.; 155, A. bijugus n. sp., holo.; 156, Crypfocepfio/o-
morpha papua n. sp., holo.; 157, Spha/Zomorpho quadrua n. sp., holo.; 158, S. dupla n. sp., holo.; 159, S. unita n.
sp., holo.; 160, Demetrida moda n. sp., S middle tibia, para., Dobodura; 161, D. brunnea n. sp., same, holo.; 162,
D. reversa n. sp., same, holo.; 163, Dicraspeda brunnea Chaudolr, 4th segment right hind tarsus (without setae), S , Do-
boduro; 164, D. bisp'mosa n. sp., same, c^ holo.; 165, D. longiloba (Liebke), same, S , Dobodura; 166, Arhtolebia
papua n. sp., 4th segments middle & hind tarsi (without setae), 9 para., Wau; 167, A. capitis n. sp., same, S holo.;
168, Miscelus sibling n. sp., right eye & supraocular setae, i holo.; 169, M. un/co/or Pufzeys, some, 9, Wau.
250 Bulletin Museum of Comparative Zoology. Vol. 137, No. 1
182
83
84
86
Fig. 170, Mi<:roleronia boro n. sp., i copulatory organs, holotype; 171, Chlaenius pan n. sp., same, holo.; 172, ^ric'io-
(ichnus o/fus n. sp., same, paratype, Wau 1200 m; 173, T. dux n. sp., same (apex of middle lobe only), para., Wau 1700
m; 174, iyler glaber n. gen. & sp., same, pore, Finschhafen; 175, Odonfomosoreus humeralii n. gen. & sp., same,
para., Doboduro; 176, Copfodero grossa n. sp., same, para., Wau; 177, A/lmuph/oeus mix/us n. gen. & sp., same, para.,
Wau; 178, Paroscopodes cyaneus (Sloane), same, Dobodura; 179, Colliuris rossi n. sp., same, holo.; 180, Doboduro
armata n. gen. & sp., same, para., Dobodura; 181, Hel/uopopuo toxopel n. gen. & sp., same, holo.; 182, Helluonldius
chrysocomes Maindron, right antenna and right hind tarsus (without setae), 9 , Nabire; 183, H. laevifrons n. sp., same,
9 holo.; 184, H. latlpes n. sp., same, 9 holo.; 185, H. politus n. sp., same, 9 hole.; 186, Helluopapua toxopei n.
gen. & sp., same, i holo.
The Carabid Beetles of New Guinea • Darlington 251
Q.
O
Ot
o
o
15
C
Q.
O
a) c
D
a
_o
6
o
3
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Geographic Variation in Anolis disfichus
Cope [Lacertilia, Iguanidae) in the
Bahama Islands and Hispaniola
ALBERT SCHWARTZ
HARVARD UNIVERSITY VOLUME 137, NUMBER 2
CAMBRIDGE, MASSACHUSETTS, U.S.A. SEPTEMBER 27, 1968
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GEOGRAPHIC VARIATION IN ANOLIS DISTICHUS COPE (LACERTILIA,
IGUANIDAE) IN THE BAHAMA ISLANDS AND HISPANIOLA
ALBERT SCHWARTZ^
ABSTRACT
Anal is cUsfichus is widely distributed in
the Bahama Islands and Hispaniola, includ-
ing the Hispaniolan satellite islands of Ile-a-
Vache, Grande and Petite Cayemite, Isla
Catalina, and Isla Saona. Analysis of vari-
ation in head scutellation, body color and
chromatic repertory, and dewlap pigmenta-
tion shows that A. distichus is divisible into
the following subspecies: A. d. distichus —
Bahama Islands: New Providence, Exuma
Cays, Long Island, Ragged Islands; A. d.
distichoidcs — Bahama Islands: Andros; A.
d. J)iminiensis — Bahama Islands: South
Bimini; A. d. dapsilis new subsp. — Bahama
Islands: Eleuthera; A. d. ocior new subsp. —
Bahama Islands: Rum Cay and San Sal-
vador. The status of the Cat Island pop-
ulations is questionable. On Hispaniola,
A. disticJiiis has been divided into the
following subspecies: A. d. dominicen-
sis — most of Haiti and the northern half
of the Republica Dominicana; A. d. igni-
gidaris — central southeastern Republica
Dominicana; A. d. propenis new subsp. —
extreme eastern Republica Dominicana; A.
d. sejunctus new subsp. — Isla Saona; A. d.
tostus new subsp. — Isla Catalina; A. d.
ravitergum new subsp. — south central Re-
publica Dominicana; A. d. favillarum new
subsp. — Sierra de Baoruco in southwestern
^ Dept. of Biology, Miami-Dade Junior College,
Miami, Florida 33167.
Bull. Mus. Comp. Zool
Republica Dominicana; A. d. aurifcr new
subsp. — central portion of Tiburon Penin-
sula in southwestern Haiti; A. d. vinosus
new subsp. — southwestern portion of Tibu-
ron Peninsula; A. d. siippar new subsp. —
extreme tip of Tiburon Peninsula; A. d.
patruelis new subsp. — He Grande Cayemite.
A. d. floridamis has been re-established as a
valid name for the continental Florida
populations which do not agree with their
Bahaman relatives; it is suggested that
floridamis is in actuality a Bahaman form
from the western portion of Andros Island.
An extensive history of A. distichus is pre-
sented to account for the distribution and
variation in the species.
INTRODUCTION AND
ACKNOWLEDGMENTS
Anolis distichus Cope is a rather small
and stocky anoline lizard which occurs
throughout the islands of the Great Bahama
Bank, Rum Cay and San Salvador, on
Hispaniola and some of its satellite islands,
and in Florida. The species was first de-
scribed in 1861 from New Providence Is-
land in the Bahamas. In 1863, Reinhardt
and Liitken named Anolis dominicensis
from Hispaniola and, although recognizing
the similarities between the two species,
considered dominicensis specifically distinct
from distichus. Barbour (1937) apparently
first combined the t\\'0 species (as A. d.
distichus and A. d. dominicensis). This
., 137(2): 255-310, September, 1968 255
256 Bulletin Museum of Comparative Zoology, Vol 137, No. 2
combination was followed by Mertens
(1939) and Cochran (1941); Mertens gave
a thorough review of the subspecies of A.
distichus which had been described in
the 76 years between the naming of A.
dominicensis and 1939. These subspecies
include altavelensis Noble and Hassler
( Isla Alto Velo off Cabo Beata, RepubHca
Dominicana), caudaJis Cochran (He de la
Conave, Haiti), juliae Cochran (Ile-a-
Vache, Haiti), and wetmorei Cochran (Isla
Beata, Republica Dominicana), as well as
distichoidcs Rosen (Andros Island, Bahama
Islands). Mertens himself named two new
subspecies from the Repuljlica Dominicana
(ignifiukiris and aJhido'gidaris) and resur-
rected brcvirostris Bocourt as applicable to
specimens from the vicinity of Barahona in
the southwestern Republica Dominicana.
Finally, Ohver (1948) described A. d.
biminicnsis from South Bimini Island in the
western Bahamas and Smith and McCauley
(1948) named A. d. floridanus from south-
em Florida. Thus, as presently understood,
there are 12 subspecies of A. distichus rec-
ognized, of which three are Bahaman, eight
occur on Hispaniola and its associated
islets, and one is on the North American
continent.
The present paper is a result of collections
made by myself and parties in both the
Bahama Islands and Hispaniola and of
specimens and information gathered by Dr.
Ernest E. Williams at the Museum of Com-
parative Zoology at Harvard University
under grants from the National Science
Foundation, B-16066 and GB-2444, and
from the American Philosophical Society.
Dr. Williams, who has for some time been
involved with Hispaniolan anoles, recog-
nized that some of the forms associated
with A. distichus in actuality pertain to an-
()th( r (and similar) species, A. brcvirostris.
He suggested that he and I joiiitK' work
out the variation in A. di.sticJius on His-
paniola as part of a rather extensive paper
dealing with new data which have ac-
cumulated concerning these lizards on that
island. But because of other duties. Dr.
Williams has agreed to a partition of the
larger work and has also suggested that I
summarize the new information on A.
distichus by myself. His collections of
anoles made in Haiti and my own collec-
tions from the Republica Dominicana sup-
plement one another very nicely, so that a
more or less complete picture of the situa-
tion of A. distichus on the entire island of
Hispaniola is now much more possible than
heretofore. There are still certain gaps in
our knowledge, and these will be pointed
out in their proper places.
Most specimens which I have examined
are in the Albert Schwartz Field Series
(ASFS); a more limited amount of material
has been borrowed from the American
Museum of Natural History (AMNH),
Carnegie Museum (CM), Field Museum of
Natural History (FMNH), Museum of
Comparative Zoology (MCZ), Richard
Thomas (RT), University of Florida, Flor-
ida State Museum (UF/FSM), Museum of
Zoology, University of Michigan (UMMZ),
and United States National Museum
(USNM). I am grateful to the following
curators and their assistants for the loan
of this supplemental material: Charles M.
Bogert and George W. Foley, Neil D. Rich-
mond, Robert W^ Inger and Hymen Marx,
Ernest E. Williams, Wayne King, Charles
F. Walker, Doris M. Cochran and James A.
Peters. Paratypes of new subspecies have
also been deposited in tlie Academy of Nat-
ural Sciences of Pliiladelphia JANSP),
Museum of Natural History, University of
Kansas (KU), and the University of Illinois
Museum of Natural History (UIMNH).
Since coloration and pattern play such a
major role in differentiating the \arious
subspeci(^s of A. distichus, I have not con-
sidered it worlhwhik to borrow all the
available specimens of the species which
exist in collections. Maiu' ot these older
specimens arc long preserved and now
much faded. 1 have attempted to examine
all material which might be- assignable to
new taxa proposed herein, and have ex-
amined all specimens w Inch are designated
Anolis distichus • Schwartz
257
as paratypes. Lists of referred specimens in
several cases include localities and museum
numbers (MCZ) which I assign to certain
taxa on the basis of provenance; specimens
so listed have not been examined by myself.
ASFS specimens have of course been
studied in detail. The probability is high
that almost all lizards listed as referred
specimens are correctly designated sub-
specifically, since they have come from
areas whose borders are delimited by fresh
material which I have examined. Excep-
tional instances or uncertain allocations are
noted in the text.
In the field I have had the capable as-
sistance of Patricia A. Heinlein, Ronald F.
Klinikowski, David C. Leber, Dennis R.
Paulson, and Richard Thomas. Mr. Thomas
succeeded in securing two distinctive sub-
species of A. disticlnis on Isla Saona and
He Grande Cayemite for me, and Messrs.
Paulson and C. Rhea Warren made especial
efforts to secure these lizards when they
visited Cat Lsland, San Salvador, and Long
Island on my behalf. Mr. Warren has also
donated specimens collected by himself on
South Bimini Island and in southern Flor-
ida. Carefully taken color notes from living
specimens have been indispensable, and
frankly, without them, the variational pic-
ture of A. distichus throughout its range
would be impossible to inteipret; I there-
fore wish to commend the efforts of others
in this particular matter of information on
fresh material, without which parts of the
present paper would be in doubt.
I am particularly in the debt of David
C. Leber, whose water color portraits of
the various subspecies of Anolis distichus
aid greatly in the visualization of the color
differences in these lizards. Plates I and
II are the result of Mr. Leber's work.- Of
the 16 portraits, ten were executed in the
field, often under trying circumstances; the
remaining six were rendered from freshly
preserved specimens and extensive color
- Publication of these plates has been made
possible by N.S.F. grant (;B-6944 to Ernest E.
Williams.
notes, at times additionally accompanied by
Kodachrome transparencies. These latter
portraits, completed under the critical eyes
of myself and Richard Thomas ( whose field
notes are herewith gratefully acknowl-
edged), are as accurate as those done in
the field.
HISTORICAL SUMMARY
As noted in the introduction, there are
12 subspecies of A. distichus. However,
these subspecies in actuality represent two
species, whose prior names are A. distichus
Cope and A. brevirostris Bocourt. Varia-
tion in the latter species, as well as its eco-
logical interrelationships with A. distichus,
are presently under study by Dr. Williams
and need not concern us further here. In
general, the two species are allotopic but
broadly sympatric, although A. disticlnis is
much more widely spread on Hispaniola
than is A. brevirostris. In certain regions,
however, the two species are precisely
syntopic; in the most general terms, A.
brevirostris inhabits xeric regions and A.
disticlnis more mesic situations, but there
are obvious and bold exceptions to this
statement (for example, A. distichus on ex-
tremely hot and dry Isla Catalina off the
southern Dominican coast).
The named fonns which are correctly
associated with A. brevirostris are caudalis
Cochran and wetmorei Cochran, whereas
the balance of the subspecies (domiuicensis,
ignii^ukiris, albidogularis, juliae, distichoides,
biminicnsis, jloridanus) are correctly as-
sociated with A. distichus. The most tren-
chant scale difference between the two
species is the absence of a "preoccipital"
scale in A. brevirostris and its presence in
A. distichus. Even this character is not
constant in either species, since most speci-
mens of A. distichus from South Bimini and
many from Andros lack the "preoccipital"
(primarily by fusion with the interparietal),
and occasional specimens from other Baha-
man Islands (most commonly from Eleu-
thera) lack the "preoccipital" either by
fusion with the interparietal or by frag-
258 Bulletin Museum of Comparative Zoology, Vol. 137, No. 2
mentation. Twenty-three A. distichns of a
total of 1001 examined from Hispaniola
and its satellites lack the "preoccipital,"
primarily hy fragmentation (thus the area
usually occupied by the "preoccipital' is
crowded by a number of small scales ) or
by fusion with the interparietal — the latter
being the less common condition. Of these
23 aberrant Hispaniolan A. dlstichus, none
is from regions where A. di.stichiis and A.
brevirostris are sympatric, but three are
from areas where A. brevirostris might be
expected to occur ( Llanos de Azua ) .
I have made no attempt to examine large
series of A. brevirostris but have studied 46
specimens of this species from the Departe-
ment de lOuest in Haiti (localities include
the northern shore of the Golfe de la
Gonave, the Cul de Sac Plain and the
southern coast in the Jacmel area) and the
vicinities of Barahona and San Juan in the
Republica Dominicana. In this lot of ma-
terial, I find that the "preoccipital" is very
variable in occurrence and shows an
amount of variation equal to that in A.
distiehns. The scale is most often absent
(fused or fragmented) in lizards from the
Barahoila area in the Republica Domini-
cana, but in Haitian material it is more often
present, although at times tiny or small in
size. The amount of overlap in size of the
"preoccipital" in A. distichus and A. bre-
virostris is fairly broad, and there are many
specimens of the latter that have a "preoc-
cipital" as large as that of many specimens
of the former. I do not interpret this con-
dition as intergradation or hybridization,
but as part of the variation of each species.
There are pattern differences between the
two species, since A. brevirostris has a jxiir
of black nuchal spots, which is absent in
A. distielius; no A. brevirostris ever assumes
a green color, as do main' subspecies of
Hispaniolan A. distieJnis. As far as my
observations are concerned, A. brevirostris
is the smaller lizard, reaching a maximum
snout-vent length in males of 47 mm,
whereas A. distichiis is generally larger,
with the largest males of all races repre-
sented by large numbers having snout-vent
lengths between 48 and 58 mm.
One name, altavelensis, has not been
associated with either A. distiehus or A.
brevirostris. This foiTn resembles A. dis-
tiehus in having a "preoccipital," but, be-
cause of other differences, Dr. Williams
suggests that it not be associated with this
species and that it be considered as a
species distinct from either A. distiehus or
A. brevirostris. The fauna of Isla Alto Velo
presents consistent peculiarities when com-
pared with that of adjacent Isla Beata and
the Peninsula de Barahona, and specific
status for A. altavelensis is no exception,
since both Isla Beata and the Peninsula de
Barahona south of the Sierra de Baoruco
are inhabited solely by A. brevirostris. Thus
altavelensis, with its "preoccipital," is un-
expectedly like A. disticlius (which occurs
in this region exclusively in the Sierra de
Baoruco and the eastern Massif de la Selle,
and not in the lowlands or along the coast )
rather than like A. brevirostris. Doubtless
A. cdtavelensis has had a long independent
history from the balance of A. distielius;
a similar situation occurs in the Alto Velo
Leioeejdialus (which I have regarded as a
peculiarly disjunct subspecies of the geo-
graphicallv removed L. vinculum; Sch-
wartz, 1967).
The material on which the name A.
dominicensis was based had as its prove-
nance merely "Haiti"; Dr. Williams has ex-
amined the syntypes and assures me that
they are indeed identical \\ith those lizards
which are currently called A. d. domini-
censis, and not with A. brevirostris. With
the description of several ne\\ mainland
Hispaniolan subspecies of A. distiehus, it
is ajipropriate to restrict the type locality
of A. d. dominicensis in order to clarify
m\' concepts of that subspecies. I herein'
designate Port-au-Prince, Departcment de
rOuest, Haiti, as the t\ pe localit)' of A. d.
dominicensis. It is not uiilikel\' tliat the
original specimens, collected by A. II. Riise,
did indeed come from the \icinit\' of the
capital of Haiti; Port-au-Prince has long
Anolis distichus • Schwartz
259
Figure 1. Partial dorsal views of heads of Anolis distichus showing modifications of head scales. Symbols: interparietal,
widely spaced horizontal lines; preoccipital," widely spaced vertical lines; supraorbitals in contact with interparietal,
narrow vertical lines; median azygous head scales, dense stipple; postfrontals, open stipple; scales in lateral contact with
postfrontals, heavy stipple.
A) ASFS 10283, Andros Island, Bahama Islands; 2/2 scales in lateral contact with postfrontals; supraorbital semicircles
completely separated by a series of 10 median azygous head scales; "preoccipital" absent by fragmentation; 0/0 supra-
orbitals in contact with interparietal.
B) ASFS X4709, South Bimini Island, Bahama Islands; 3 scales in lateral contact with postfrontal on right side, left
side abnormal; supraorbital semicircles in contact; 6 median azygous head scales; "preoccipital" absent by fusion with
interparietal (denoted by overlap of symbols); 2 2 supraorbitals in contact with interparietal.
C) KU 93369, Carrefour Canon, Haiti; 3- 3 scales in lateral contact with postfrontals; supraorbital semicircles in con-
tact; 4 median azygous head scales, including "preoccipital" (denoted by shading); "preoccipital" present; 0/1 supra-
orbitals in contact with interparietal.
D) USNM 157924, 10 km W Bani, Republica Dominicana; 2,2 scales in lateral contact with postfrontals; supraorbital
semicircles in contact; one (the "preoccipital, denoted by shading) median azygous head scale; "preoccipital" present;
1 1 supraorbitals in contact with interparietal.
been a prominent Caribbean seaport. An-
other possibility might be Cap-Haitien, and
assumption of this city as the source of the
original dominicensis material would not
alter my taxonomic conclusions, since I
regard the populations of A. distichus at
Cap-Haitien as identical with those at
Port-au-Prince. In favor of Port-au-Prince
as the type locality of dominicensis is the
(admittedly oblique) association of Riise
with the type specimen of Sphaerodactijhis
copei Steindachner, a lizard which does
occur in the environs of Port-au-Prince but
not at Cap-Haitien (see Schwartz and
Thomas, 1965:317, for discussion of S.
copei ) .
METHODS
When he described A. d. biminiensis,
Oliver (1948) analyzed some Bahaman
populations of A. disticluis on the basis of
various scale counts and relationships.
Hoping that an application of his counts
to non-Bahaman A. distichus might reveal
differences other than coloration and pat-
tern between various subspecies, I have
followed his techniques and applied them
to the material I have examined. Repre-
sentations of several of the variant condi-
tions are shown in Figure 1. The scale
counts employed are:
1 ) Number of scales across the snout at
the level of the second canthal scale. I
follow Williams (1962:2) in making this
count, in that the second canthal is reckoned
from the anterior border of the orbit.
2) Number of loreal rows.
3) Scales between the supraorbital semi-
circles.
260
Bulletin Museum of Compomtive Zoology, Vol. 137, No. 2
Table 1. Sixteen subspecies of Anolis distichus, showing statistically significant differences
IN means of number of median azygous head scales. Size of sample in first column, means
and two standard errors of means in second column, a plus in tables indicates that the two
subspecies involved differ significantly (non-overlap of two standard errors of mean); a
minus indicates no statistical differences. Two subspecies (sejunctus, tosius) are not included
because of very small sample size.
N
M
s
2
o
o
1
,60
13
.2
so
s
5
.2
.5)
.5)
60
3
a.
c
;^
a.
1
2
a
3
.2
'^
^
^
1
distichus
126
6.0
±
.23
X
+
—
—
—
+
+
+
+
+
+
+
+
+
+
+
distichoides
160
8.7
-±^
.16
X
+
+
+
+
+
+
+
+
+
+
+
+
+
+
biminiensis
42
5.5
±
.78
X
—
+
+
+
+
+
+
+
+
+
—
+
dapsilis
101
6.2
-+-
.36
X
—
+
+
+
+
+
+
+
+
+
+
+
actor
55
5.8
-±^
.49
X
+
+
+
+
+
+
+
+
+
+
+
dominicensis
235
3.9
-+-
.20
X
+
+
+
—
—
+
—
+
—
+
i^ni^idaris
106
3.5
±
.08
X
+
+
—
—
—
—
—
+
+
))IOl)CIUS
58
2.8
±
.16
X
—
+
+
+
—
+
+
+
ravitergurn
49
2.6
-+-
.33
X
+
+
+
+
+
+
+
favillanim
26
3.8
-±^
.48
X
—
—
—
—
—
+
uurifcr
64
3.7
±
.32
X
—
—
—
+
+
vinosus
100
3.4
-+-
.26
X
. —
—
+
+
juliae
27
3.4
-±^
.46
X
• —
+
+
suppar
174
3.4
±
.19
X
+
+
patruelis
25
4.6
•+^
.62
X
+
floridanus
90
7.9
±
.30
X
4) NiimlxT of rows of .scales between
supraorbital semicircles and interparietal
scale. This figure is written as a fraction
(i.e., 1/1, meaning that the interparietal is
separated from the semicircles by one scale
on each side). A count of 0/0 means that
there are no scales between the semicircles
and the interparietal, and that the semi-
circles and interparietal are thus in contact.
5) Numlier of subdigital lamellae on
phalanges II and III of the fourth toe.
6) Number of scales bordering the post-
frontal laterally (see Oliver, 1948:16, for
drawings showing these scales in A.
distichus). In a small number of speci-
mens of A. (lislicliiis, the postfrontal may
abnormally extend so far laterally as to
make contact with one of the canthals. In
such instances, I have not included the
canthal as a scale in contact with the post-
frontal, since the condition is obviously
anomalous.
7) Number of median (usually azygous)
scales posterior to the postc^iormost para-
median pair of snout scales, usually re-
stricted to the midline from the anterior
border of tlie prefrontals posteriorly, and
including the "preoccipital" as an unpaired
median scale. Thus a count of 1, for ex-
ample, means that between the anterior
border of the postfrontals and the inter-
parietal, there is only one scale ( usually the
"preoccipital"). A count of 0 occurs when
the "preoccipital" is fused with the inter-
parietal, provided that there are no addi-
tional median azygous scales. If some or
all of the scales between the postfrontals
are i)aired, but extremely irregularly so,
the count includes these irregularly paired
scales as median azygous scales, since they
do not present the r(>gular conformation of
the paired paramedian snout scales of A.
(list id HIS. l\d)U 1 shows the statistical
significance of differences between sub-
species in this character.
8) Number of supraorbital semicircle
scales in contact with interparietal. This
count is ixutK' correlated with (4), the
Angus distichus • Schwartz
261
Table 2. Sixteen subspecies of Anolis distichus, showing statistically significant differences
IN means of number of postmental scales. See Table 1 for details.
N
M
•5
2
"3
•2
■i
s
IS
1
u
e
1
o
■I
e
a.
g
1
1
.-1
5
Co
.2
s
disticluis
127
5.5
-+-
.22
X
+
+
—
+
+
+
+
+
+
+
+
+
+
+
distichoides
159
4.6
^2
.14
X
—
+
+
+
+
+
+
+
+
+
+
+
+
])iiniiiicnsis
42
4.8
±
.20
X
+
+
+
+
+
+
+
+
+
+
+
+
dapsiUs-
105
5.2
±
.21
X
+
+
+
+
—
+
+
+
+
+
+
+
ocior
56
6.7
-±^
.28
X
—
+
— •
+
—
+
—
+
+
+
dot)nnicensis
240
6.6
±
.13
X
+
—
+
+
+
+
+
+
+
ignigidaris
105
6.0
-±^
.29
X
+
—
+
+
+
+
+
+
propcrus
58
6.7
-+-
.38
X
+
—
+
+
+
+
+
niv iter gum
56
5.6
-±^
.34
X
+
+
+
+
+
+
favilhiium
28
6.1
-±^
.41
X
+
+
+
+
+
+
aurifer
62
7.2
+
.28
X
—
—
• —
—
+
viuosus
98
7.4
-+^
.27
X
—
—
—
+
juliae
31
7.2
±
.47
X
+
—
+
sui))nir
171
7.9
±
.22
X
—
+
patrucUs
25
7.8
-+-
.48
X
+
floridanus
89
4.4
-+:
.23
X
number of rows of scales between the
semicircles and the interparietal; for in-
stance, if the latter count is 1/1, the num-
ber of supraorbital scales in contact with
the interparietal will of necessity be 0/0.
However, if the count of (4) is 0/0 (i.e.,
there are no scales between the semicircles
and the interparietals), then (8) may have
a fairly wide fluctuation.
9) Number of postmental scales. Table
2 shows the statistical significance of differ-
ences between subspecies in this character.
10) Presence or absence of a "preoc-
cipital."
The above counts have been taken on
1588 specimens from Florida, the Bahamas,
and Hispaniola. Some of them have proved
to be useful, primarily on a modal rather
than an absolute level, in defining the sub-
species. Very small samples often show
such a wide diversity in some counts that
it is impossible to state with certainty what
the modal condition is, but with increas-
ingly large samples, in most cases a distinct
mode can be easily determined for each of
the counts. The degree of overlap between
the various counts for the different samples
is often great, so that it is difficult to
identify a particular lizard to subspecies on
the basis of any single count. Reliance
must be placed on such features as dewlap
pattern and coloration, and coloration and
pattern of the head and body.
Of the counts taken, those of scales across
the snout, number of loreal rows, and
lamellae overlap so broadly between the
samples and are so variable intra se that
they serve no useful purpose insofar as
diagnosing the subspecies is concerned.
The data for these counts are presented in
each case, but merely for the sake of com-
pleteness.
BAHAMAN VERSUS HISPANIOLAN
POPULATIONS AS A WHOLE
The only statement contrasting the dif-
ferences (if any) between all the Bahaman
populations of A. distichus versus all the
Hispaniolan populations is that of Cochran
(1941:146) who noted that "In adult ex-
amples of disticluis there are distinct keels
on the enlarged scales of the femur, while in
dominicensis these scales are always
smooth. . . ." Examination of large num-
262 Bulletin Museum of Comparative Zoology, Vol. 137, No. 2
bers of A. distichus indicates that no such
dichotomy exists, and the two major geo-
graphic subdivisions cannot be distin-
guished on the basis of presence or absence
of keeled scales on the anterior femoral face.
Additionally, no other scale character will
separate the two segments of A. distichus
absolutely, but there are a few characters
which generally differentiate the two sec-
tions.
1 ) There is a tendency for Bahaman
populations to have the supraocular semi-
circles completely separated by a single
median row of azygous head scales. This
character reaches its greatest development
in the populations from South Bimini and
Andros in the Bahamas, but occurs casually
in all other Bahaman samples. No His-
paniolan specimen shows this character.
2) All Bahaman populations but one
have 0/0 scales between the semicircles and
the interparietal as the modal condition,
whereas in Hispaniolan samples there are
either 0/0 or 1,1 scales modally between
the semicircles and the interparietal, with
1/1 having the higher incidence by pop-
ulation.
3) Median head scales in the Bahamas
vary in mean from 5.5 to 8.7, whereas in
Hispaniola the means vary from 2.6 to 5.0 —
the highest mean being probably higher
than it is in reality, since the sample is com-
posed of only six lizards. Table 1 shows
the data on head scales.
4) The absence of the "preoccipital'
scale is most frequently encountered in
Bahaman populations and occurs only very
rarely in Hispaniolan A. disticJius, as pre-
viously pointed out. All Bahaman popula-
tions from which I have examined samples
have at least one or a few specimens which
lack the "preoccipital" scale, whereas onl\-
a very few Hispaniolan A. distichtis lack
this feature.
Although none of the above is com-
pletely diagnostic of Bahaman versus His-
paniolan A. di.stic]nis, it does suggest that
there has been a greater divergence be-
tween the two major segments of A.
distichus than between intra-Bahaman and
intra-Hispaniolan populations.
There is also one suggestive color dif-
ference between Bahaman and Hispaniolan
A. disticJnis. With one exception, all Baha-
man populations are incapable of a true
green phase. The general coloration of
Bahaman lizards is a pale ashy gray to
sandy tan, capable of becoming dark wood
brown, although this latter condition is
rather rarely observed. Very occasionally
Bahaman lizards are observed to be a very
pale ashy green, but bright or dark green
lizards, such as occur in several Hispaniolan
populations, are unknown from the Baha-
mas. The one Bahaman exception is lizards
from Rum Cay and San Salvador; on these
two isolated islands, A. disticluis is distinc-
tively colored ( in reference both to other
Bahaman and to Hispaniolan populations)
in that it is regularly a pea-green or yellow-
green. In fact, the yellow component of the
dorsal pigmentation may be more striking
than the green hues. The Rum Cay-San
Salvador lizards are the onh' populations in
the Bahamas where A. disticlius is known to
be greenish rather than gray or tan.
Many Hispaniolan subspecies of A.
disticluis, on the other hand, do indeed have
a green phase, the greens varying from
bright to a pale ashy (which is much more
distinctly green than any green observed in
Bahaman lizards other than those on Rum
Cay and San Salvador). Even this color
repertory distinction between the two seg-
ments of A. disticlius is not absolute, since
some Hispaniolan subspecies are not known
to be able to assume the green phase, and
thus resemble the Bahaman populations.
As far as dewlap coloration and pattern
are concerned, the Bahaman A. distichus
are verv likt> some ol their Hispaniolan
relatives. Although I lia\e no quantitati\e
data, the dewlaps of l^ahaman A. distichus
appear smaller than do those ol the His-
paniolan lizards, but this ma\ be merely
an artilact of observation or preservation
techni(iues. The dewlap pattern and colora-
tion ol Hahanian .A. J/.s//r7///.v resemble those
Anolis distichus • Schwartz
263
of lizards from various Hispaniolan lo-
calities; the most aberrant dewlap pattern
and colors occur in specimens from the
extreme southwestern portion of the Tiburon
Peninsula of Haiti and on its adjacent Ile-a-
Vache.
SIZE AND NATURAL HISTORY
The largest specimens of A. disticlius are
from the southeastern uplands of Haiti. On
the Montague Noire in the vicinity of
Peneau and Furcy, males reach a snout-
vent length of 58 mm and females 48 mm.
In general, in all populations, females reach
a maximum size of about one centimeter
less than males. The smallest of the
maximally sized males (46 mm snout-vent
length) are from Isla Catalina off the
southern coast of the Repiiblica Domini-
cana, and the smallest maximally sized fe-
male (38 mm) is from Isla Saona. How-
ever, the samples from both islands are
very small (five males and one female
from Saona; three males from Catalina),
so that these comments are equivocal.
Etheridge (1966:351) stated that the
largest Bahaman A. distichus he had ex-
amined had snout-vent lengths of 48 mm
(New Providence, Andros, Cat) to 53 mm
(Eleuthera). On the other hand, he noted
that Hispaniolan specimens reached a maxi-
mum snout-vent length of about 50 mm.
My own Bahaman data, based on 385 speci-
mens in contrast to Etheridge's data for
126 specimens, do not agree with his
Bahaman figures, since the maximally sized
Eleuthera male (of 107 Eleuthera speci-
mens) I have measured has a snout-vent
length of 50 mm, slightly smaller than
Etheridge's maximum for that island. The
largest Bahaman males I have seen are
from San Salvador and Rum Cay, and have
snout-vent lengths of 53 mm, precisely the
same as the largest male (from Eleuthera)
examined by Etheridge. These discrep-
ancies have little significance, but they in-
dicate that populations on various Baha-
man Islands do differ in maximum adult
size.
A. distichus has a broad distribution on
Hispaniola and is rivalled in this respect
only by Anolis ricordi Dumeril and Bibron,
Anolis cybotes Cope, and Anolis scmilincatus
Cope. It occupies situations varying from
mesic oases in otherwise extremely xeric
regions (Cul de Sac-Valle de Neiba plain)
to rain forest at high elevations; it even
occurs in only slightly more shady areas
within xeric areas themselves (vicinity of
Monte Cristi, Republica Dominicana).
Typically, A. distichus prefers shady and
mesic forested or pseudo-forested situa-
tions, such as hardwood forests, coffee and
cacao groves, mango-breadfruit-royal palm
associations, overgrown and shady fence-
rows along abandoned fields, etc. In some
areas it literally swarms, whereas in other
and apparently quite similar areas it is
extremely uncommon. A. distichus, in dense
forest, often prefers large trees which ex-
tend above the lower canopy, and in cacao
groves (where A. disticlius and A. cijhotcs
occur syntopically on the same trees), A.
distichus in general seems to prefer the
more exposed — and thus slightly more
sunny — branches, although a mature cacao
grove is inherently very deeply shaded and
cool and often canopied by much larger
forest trees. Sleeping A. distichus are not
easily observed, as Rand (1962:11) pointed
out. I saw none in Haiti in two months
fieldwork, and encountered the first
sleeping individual near Miches in the Re-
piiblica Dominicana; this lizard was on the
upper surface of an herb leaf within two
feet of the ground. In northwestern Re-
publica Dominicana, near Palo Verde, in an
extensive patch of flood plain hardwoods
along the Rio Yaque del Norte, Thomas
and I encountered many A. disticlius sleep-
ing in company with A. cybotes and A.
chlorocyanus Dumeril and Bibron. Here
A. distichus customarily slept on the leaves
and twigs of small herbs and shrubs, within
three feet of the ground, whereas both
A. cybotes and A. cldorocyonus slept on the
tips of small branches of saplings or on the
tips of long and slender lianas and vines.
264 Bidletin Museum of Comparative Zoohg.ii, Vol 137, No. 2
A. chlorocyanus slept distinctly higher in
the canopy than A. cijhotcs, since no A.
chlorocyanus was enconntered below eight
feet above the ground and most \\'ere above
ten feet and inaccessible. Considering the
occurrence at this locality of the vine-
inhabiting and climbing snakes Epicrates
gracilis Fischer and Uromaccr oxyrhynclius
Dumeril and Bibron, the use of the tips of
branches and pendant vines by A. cyhofcs
and A. chlorocyanus is most suggestive; the
distinctly lower and non-tree or vine as-
sociated sleeping sites for A. distichus may
well have a distinct positive survival value
in an area where these two primarily
arboreal snakes are abundant.
A. distichus occurs in Hispaniola at eleva-
tions from below or near sea level (Valle
de Neiba) to at least 6000 feet (1830
meters), in the Sierra de Baoruco, Massif
de la Selle, and Cordillera Central.
In the Bahamas, A. distichus occurs with
some frequency in hammock woods or
coppice (South Bimini, New Providence),
but also occupies (as Rand, 1962:4, noted
for Hispaniolan A. distichus) isolated large
and often gray-barked trees, such as Ficus,
with whose bark coloration the Bahaman
races blend excellently, and which addi-
tionally offer sanctuary among adventitious
roots and buttresses. Other trees with which
A. disticlius is customarily associated in the
Bahamas are Coccoloha, Lysilonui, and
TerminaJia; all have pale bark which ren-
ders the lizards inconspicuous. In Nassau,
A. disticlius occurs commonly on crannied
limestone walls and street cutbanks, and on
San Salvador the species was abundant
about tlie ruins of Sandy Point House
(=Watling's Castle), both on the sur-
rounding trees and saplings and on the
buikling itself.
On some Bahaman islands, A. distichus
is (juite common. Thus, it is abundant on
New Providence and Eleuthera, for in-
stance, and Oliver (1948:32) noted that
C. M. Breder, Jr., secured a series of 164
A. distichus from native boys on Andros;
my own observations on Andros do not
indicate such a present abundance of A.
distichus, however. On South Bimini, A.
distichus is only moderately common;
Oliver (o)). cit.:22) secured 20 specimens
from Ficus and CoccotJirinax, but recent
collectors have not secured these lizards so
abundantly on South Bimini. At the other
extreme of abundance lies Rum Cay, where
A. distichus is distinctlv uncommon; here
the lizards were observed and collected
primarily on Cocos palms and other trees
in the settlement of Port Nelson, and only
occasional individuals were observed away
from human habitations. Only two individ-
uals, both on Cat Island, have been noted
sleeping in the Bahama Islands. Richard
Thomas observed these lizards sleeping on
small limbs, between 6 and 7 feet ( ± 2
meters) above the ground; one sleeping
lizard was in a Sa1)al grove and the other
in an open group of large trees surrounded
by thorn scrub. Occasional individuals have
been collected diurnally beneath rocks both
inland and near the strand, so it is possible
that Bahaman A. distichus resort also to
such situations for nocturnal retreats.
Rand ( 1962 ) has summarized his ob-
servations on three Hispaniolan anoles (A.
distichus, A. cyhotcs, A. cldorocyanus) both
in the field in the Republica Dominicana
and in the laboratory. My observations on
A. disticJius differ somewhat from his; for
instance, he regarded this species as living
"primarily on isolated trees and fence posts
and along the edges of woods and trails
in open woods." The abundance of A.
(listiclius in Dominican cacao gro\es (admit-
tedly an artificial situation) and in dense
mesic woods high in the Cordillera Central
is in contrast to Rand's statement. Such
difference s ma\- well r(41(>ct different habits
in difierent regions, and suggest that one
species of anole may occup\' \arying habi-
tats in dif brent areas, and that extreme
caution should bc^ used in generalizing
about the habitat preferences of geographi-
cally \\idel\' distributed anol(\s. It is also
pertinent in this connection that Mertens
(1939:15) rcportcxl th(> occurrence of A.
Angus distichus • Schwartz
265
distichus (along with A. cijhotes) in pine
forest at Paso Bajito in the Cordillera
Central. In the higher pine woods near
Constanza, at elevations between about
40()() and 6000 feet ( 1220 and 1830 meters),
A. distichus is at best rather rare, preferring
in this region residual stands of rainforest.
It has not been taken or observed in the
vieinity of Valle Nuevo (about (SOOO feet;
2440 meters) where A. shrcvci Cochran is
the commonest (and perhaps only) anole
of the cool and open pine-forested slopes.
SYSTEMATIC ACCOUNT
Anolis distichus distichus Cope
Anolis distichus Cope, 1861, Proc. Acad. Nat. Sci.
Philadelphia: 208.
Ty))e locahty. New Providence Island,
Bahama Islands.
Definition: A subspecies of A. distichus
characterized by small size (males to 49
mm, females to 44 mm snout- vent length),
dorsum pale ashy gray to sandy tan and
without a green phase, dewlap pale yellow,
rarely with a vague basal to more extensive
orange blush, modally 0/0 scales between
the supraorbital semicircles and the inter-
parietal, 0/0 supraorbitals in contact with
the interparietal, 2/2 scales in contact lat-
erally with the postfrontals, and high mean
number (6.0) of median azygous head
scales.
Distribution: The Bahama Islands:
known definitely from New Providence,
the Exuma Cays (Warderick Wells Cay,
Staniel [= Stanyard] Cay, Darby Cay),
Great Exuma, Little Exuma, Long Island,
and Great Ragged Island ( Fig. 2 ) .
Comments: A. d. distichus is widely
distributed on the islands to the east of the
Tongue of the Ocean on the Great Bahama
Bank and presumably on the Ragged Is-
lands. Specimens from Cat Island will be
discussed later.
In life, A. d. distichus is normally a gray
lizard, but some specimens are sandy tan in
life. Occasional specimens demonstrate a
boldly contrasting pattern of brownish black
ground color with black crossbands; in this
phase the snout is smudged with sooty black
and the eyeskin is also sooty. Rarely some
lizards show a very pale greenish gray
phase. Although I have not so recorded it,
I assume that A. d. distichus can become
rich dark brown as can several of the other
Bahaman subspecies. The dorsum is at best
only very weakly longitudinally striate with
darker, and there may be a single vague
scapular chevron, its apex pointed pos-
teriorly. The interocular dark bar is vari-
able, but even when best expressed, is not
especially prominent; other head markings
are vague and ill defined. The venter is
cream to very pale yellowish, and the un-
derside of the tail is very pale yellow also.
The dewlap is regularly pale yellow (Pi.
I). Rarely is there a basal orange blush;
if present, the orange is extremely faint
and only barely discernible. Very occa-
sional specimens ( Long Island ) have the
pale orange more extensive.
The islands to which I have attributed
the nominate subspecies may be con-
veniently divided for further discussion
into four areas: 1) New Providence, 2) the
Exuma Cays, including Great and Little
Exuma, 3 ) Long Island, and 4 ) Great Rag-
ged Island. The samples from these four
areas are alike in dorsal coloration and pat-
tern and presumably in dewlap color (I
have not seen live Ragged Island speci-
mens), and on these bases I group them
together. In scale characters, there are
some differences which may be pertinent,
but I have chosen not to emphasize them.
The following data are from a series of 49
New Providence specimens, 16 from the
Exumas, 57 from Long Island, and 10 from
Great Ragged Island. I have seen living
specimens from New Providence and Long
Island, and freshly preserved material from
the Exumas.
Long Island and Great Ragged Island
specimens modally have 0/0 scales between
the semicircles and the intei-parietal, and
0/0 is one of two bimodes (each with 20
specimens) on New Providence. In the
266
Bulletin Museum of Comparative Zoology, Vol. 137, No. 2
78°
1^
74'
\j>^:
0 20 , «0 60
j)\
26*-
-23'
23'
^^^=^.
O I
^^
78'
74°
''O
Figure 2. Map of the Bahama Islands, showing distribution of the subspecies of Anoiis disfichus; diagonal lines, upper
left to lower right, A. d. d/stichus; diagonal lines, upper right to lower left, A. d. daps/7is; open stippling, A. d. distichoides;
close stippling, A. d. oc/or; F, A. d. Iloridanus; B, A. d. bimmiens/s; overlap of symbols for A. d. distichus and A. d.
dapsilis suggests area of intergradation between these two subspecies.
specimens from the Exuma Cays, there are
modally 1/1 scales lietween the semicircles
and the interparietal. In number of supra-
orbitals in contact with the interparietal,
0/0 is the mode in all samples except that
from Great Ragged, which has 1 /I modally
(although 0/0 has a freciuency of two
lizards and 1/1 a frecpiency of three lizards).
All samples modally have 2^2 scales in
lateral contact with the postfrontals. The
highest incidence of complete median sep-
aration of the supraorbital semicircles oc-
curs on Long Island (four of 53 lizards),
whereas New Providence has three of 46,
the Exumas one of 16, and none occurs on
Great Ragged. Scales across the snout
range from 4 to 6 (New Proxidence), 8
(Exumas), or 7 (Long), and are either 5
or 6 on Great Ragged; modes 4 (New
Providence), 5 (Great Ragged) and 6
(Exuma (]ays. Long). Loreal rows \ar\'
from 3 to 5 on New Providence (mode 4),
and 4 to 6 on the Exumas (mode 4) and
Long (mode 5); loreal rows on Great
Ragged are 4 or 5 (mode 4). Fourth toe
lamellae vary from 15 to 20 (New Provi-
dence, the Exumas), 14 to 19 (Long), and
15 to 19 (Cireat Ragged), with modes of
18 in the former two samjiles and in the
Great Ragged lizards, and 17 on Long
Island. Median azygous head scales vary
between 3 and 13 (mode 5, mean 6.2) on
Anolis DiSTicHus ' Schwartz
267
New Providence, 3 and 9 (mode 5, mean
5.6) on the Exumas, 2 and 10 (mode 6,
mean 5.9 ) on Long, and 3 and 10 ( mode
6, mean 5.9 ) on Great Ragged. Postmentals
vary from a low of 4 in all samples to 7 on
New Providence, the Exnmas, and Great
Ragged, and 10 on Long. The mode is 4
on New Providence and the Exmnas, and
6 on Long and Great Ragged. The means
of postmentals are 5.0 ( New Providence ) ,
5.1 (Exnmas), 5.4 (Long), and 5.3 (Great
Ragged). The "preoccipital" is regularly
present; one specimen from New Provi-
dence, one from the Exumas, one from
Great Ragged, and four from Long lack
this scale.
From the above data, it appears that a
certain amount of divergence has taken
place in the four areas which are inhabited
by A. d. distichus. In general, the Exuma
Cays material is closer in most counts to
the lizards from New Providence ( although
the scales between the semicircles and the
interparietal are a notable exception). The
Long Island lizards, on the other hand,
differ somewhat more. The sample from
Great Ragged Island is too small for de-
tailed comment. In the alxsence of any
established chromatic or pattern differences,
I place all four populations in the nominate
subspecies, although I acknowledge the
modal differences mentioned.
As noted above, A. d. distichus is com-
mon on Ne\\' Providence, where it was ob-
served abundantly in Nassau ( especially on
rock walls and exposed limestone street
cuts), and on and about the limestone
bluffs near the coast at Cave Junction. At
the latter l^^cality, the lizards occurred also
on saplings and large Ficus about the
bluffs, and on Coccoloba on the coast. In
high coppice near Nassau East, A. distichus
was extremely abundant, both on the trees
and saplings and on an old rock wall which
extended for some distance through the
woods. The species is only moderately
common in coppice on Great and Little
Exuma.
Specimens examined: BAHAMA IS-
LANDS. New Providence (localities not
mapped): Nassau, 23 (AMNH 76348-54 +
16); Cave Point, 3 (ASFS 10301-03); Cave
Junction, 10 (ASFS V7206-15); 0.9 mi.
(1.4 km) W Cave Junction, 2 (ASFS V7226-
27); hills south of Lake Cunningham on
Gladstone Road, 4 (ASFS V2092-95); Pros-
pect Ridge, 2 (ASFS V2102-03); The
Grove, 1 (ASFS V2104); Windsor Field, 1
(ASFS V2110); 0.6 mi. (1.0 km) NW
Yamacraw Beach, 1 (ASFS V7242); 0.3
mi. (0.5 km) E Nassau East, 2 (ASFS
V1063S-39). Exuma Cai/s: Warderick
Wells Cay, 3 (AMNH 76326-28); Staniel
(=Stanyard) Cay, 5 (AMNH 76329-33);
Darby Cay, 2 (AMNH 76334-35). Great
Exmna: 3.2 mi. (5.1 km) NW George
Town, 5 (ASFS V7033-36, ASFS V7053).
Little Exuma: 5.7 mi. (9.1 km) SE The
Ferry, 1 (ASFS V7043). Long Island:
Simm's, 2 ( MCZ 42282-83); Cray's Settle-
ment, 5 (ASFS V8562-64, ASFS V8567-68);
2 mi. (3.2 km) E Gray's Settlement (not
mapped), 3 (ASFS V8579-81); Deadman's
Cay Settlement, 5 (UMMZ 115596); Clar-
ence Town, 37 (MCZ 37986-95, MCZ
86931-53, UMMZ 80510-2 specimens,
FMNH 25372-73); 3.6 mi. (5.8 km) SE
Clarence Town (not mapped), 4 (ASFS
V10835-38); Roses, 1 (FMNH 22750).
Great Ragged Island: Duncan Town, 10
(UMMZ 118008-6 specimens; UMMZ
1 18009-4 specimens ) .
Anolis distichus disfichoides Rosen
Anolis distichoides Rosen, 1911, Liinds Univ.
Arrskr. N.F., Afd. 2, 7(5):29.
Type locality: Stanniard Creek, Andros
Island, Bahama Islands.
Definition: A subspecies of A. disticlius
characterized by moderate size (males to
51 mm, females to 43 mm snout-vent
length), dorsum grayish tan to gray and
without a green phase, entire dewlap
orange to yellowish-orange, modally 1/1
scales between the supraorbital semicircles
and interparietal, 0/0 supraorbitals in con-
tact with the interparietal, 2/2 scales in con-
tact laterally with the postfrontals, and very
268 BuUetin Museum of Comparative Zoology, Vol. 137, No. 2
high mean number ( 8.7 ) of median azygous
head scales correlated with the high in-
cidence (about 50 per cent) of complete
separation of supraorbital semicircles medi-
ally.
Di.strihution: The Bahama Islands:
known from Andros Island ( including
Mangrove Cay) and the Berry Islands
(known definitely from Frazer's Hog Cav)
(Fig. 2).
Comments: The status of A. d. disti-
chuidcs has been disputed in the past.
The main claim for its recognition has
been the orange dewlap (PL I), in contrast
to the yellow dewlap of topotypical A. d.
distichus. Although I have collected very
few dlstichoidcs ( as pointed out previously,
I have observed it rarely on Andros ) , those
males which I have seen in life have had an
orange dewlap consistently. Scale data
from 161 A. d. dlstichoidcs show the fol-
lowing: snout scales 4 to 8 (mode 6),
loreal rows 4 to 6 (mode 5); supraorbital
semicircles in contact in 108 specimens and
completely separated by median azygous
head scales in 51 lizards; modallv 1/1 scales
between semicircles and interparietal and
0/0 supraorbitals in contact with inter-
parietal; 2/2 scales ui lateral contact with
postfrontals; fourth toe lamellae 15 to 21
(mode 18); median azygous head scales 3
to 14 ( mode 9, mean 8.7 ) ; "preoccipital"
more often present ( 84 lizards ) than absent
( 74 lizards ) ; postmentals 2 to 8 ( mode 4,
mean 4.6). The almost equal incidence of
presence or absence of the "preoccipital"
is noteworthy, although more dislichoidcs
have this scale than lack it. The high mean
of median head scales is correlated with
the high Irequency of complete separation
of the semicircles. No other subspecies of
A. disticliKs, either Bahaman or Ilispaniolan,
has so high a mean, although it is ap-
proached most closely (7.9) by the main-
land populations of A. d. floridaiuis. South
Bimini A. distichus likewise have a high
incidence of complete semicircle separation,
but the mean number of median head
scales is much lower (5.5).
Tlie above scale features, especially the
high number of median head scales and the
high incidence of absence of the "preoc-
cipital," as well as the frequent separation
of the supraorbital semicircles, all dif-
ferentiate disticlioidcs from the nominate
subspecies. Adult male A. d. distichus are
also slightly smaller and have a yellow
rather than orange or yellow-orange dew-
lap. Both subspecies resemble each other
in dorsal color, although I have not noted
dlstichoidcs being tan in life. As in the
nominate subspecies, head markings are
suppressed or absent in dlstichoidcs; the
interocular bar is not prominent when
present and is often absent. There may be
a series of four dorsal chevrons, but these
are often obscure or absent, and the degree
of dorsal dark striation is likewise variable,
with a strong tendency for the lizards to
lack striae.
A. d. dlstichoidcs is the only Bahaman
subspecies which modally has 1 1 scales
between the semicircles and the inter-
parietal. Sixty-five lizards fall into this
category. On the other hand, 61 lizards
have 0/0 scales between the semicircles and
the interparietal, so the modalit) is not
strong. The virtualh' bimodal ceMidition in
this scale character is not obviously due to
the samples invoh ed; since Andros is a ver\'
large island (nearly 100 miles long and up
to 40 miles wide) and is much dissected by
bights and minor \\ater\\a\ s. it was con-
ceivable that the two modalities were due
to the pooling of data from two ]")opulations
which are divergent in this character.
This is not the case, since most of the
disticlioidcs sample under study ari' from
Mangrox'e Caw and w ithin this lot ol lizards
th(> bimodality is clearly shown.
The size of Andros and \hc inace(\ssibility
of its west coast is possibK sigiiilieant in
another matter. There is but a single A.
di.slichus axailable lioni llic entire west
coast of Andros. This is a iciuale ( UF/FSM
18005); its sex precludes knowledge of dew-
lap color and its geographic uniqueness
prcNcnts an assessment ol the characters of
Angus nisiiciivs • Schwartz
269
the populations whence it was taken. The
specimen is mentioned here and Hsted be-
low as A. (I. distichoidcs\ but for several
reasons I suspect that the population
whence it was derixed in actuality repre-
sents A. (I. floiidanus. Further comment
upon this lizard will be made in the discus-
sion of the history of the latter subspecies.
The occurrence of A. d. dlstichoides on
the Berry Islands has not been previously
reported. Two specimens from Frazer's
Hog Cay collected by l^ichard Thomas are
clearly referable to this subspecies; one is a
male with an orange dewlap and the other
a female. The male has the semicircles
completely separated by a median row of
S scales, and both lizards lack the "preoc-
cipital." Possibly these two lizards might
be better associated with the subspecies on
South Bimini, but I consider them dis-
ficlwides on the basis of provenance and the
affinities of the Berry Islands fauna.
Specimens- examined: BAHAMA IS-
LANDS. Andws Island: no further lo-
cality, 22 (UMMZ 80369-4 specimens,
UMMZ 80377-11 specimens, 80381-6 speci-
mens, UMMZ 80384); Morgan's Bluff (not
mapped), 7 (UF/FSM 17626, UF/FSM
17628, UF FSM 17630-32, UF/FSM 17634,
UF FSM 17637); ca. 0.5 mi. (0.8 km) N
Nicholl's Town, 1 (ASFS V6972); NicholFs
Town (not mapped), 1 (UF/FSM 18013);
Coakley Town, 4 (MCZ 41986-89); south
side, mouth of Fresh Creek, 10 (ASFS
10280-86, UMMZ 115598-3 specimens);
Mangrove Cay, 103 (MCZ 42013 + 15 un-
tagged specimens, AMNH 63073-19 speci-
mens, UMMZ 260210-4 specimens, UMMZ
10922.3-5 specimens, UMMZ 115597-34
specimens, plus 25 untagged specimens
from AMNH 63067); south side. South
Bight, 1 (MCZ 42001); Little Creek, 5
(UMMZ 118006); Pure Gold (not mapped),
15 (MCZ 42026-29 + 11 specimens); west
coast, 2 mi. (3.2 km) at 55° from mouth
of Deep Creek (not mapped), 1 (UF/FSM
18005). Berry Islands: Frazer's Hog Cay,
2 (ASFS \T0667-68).
Anolis distichus biminiensis Oliver
Anoli.s distichus- l)iminiensis Oliver, 1948, Amer.
iMiis. Novitates, No. 1383:16.
Tijpe locality: Western end of South
Bimini Island, Bahama Islands.
Definition: A subspecies of A. distichus
characterized by small size (males to 50
mm, females to 44 mm snout- vent length),
dorsum pale gray and without a green
phase, dewlap orange, modally 0/0 scales
between the supraorbital semicircles and
the interparietal, 2/2 supraorbitals in con-
tact with the interparietal, 3 3 scales in
contact laterally with the postfrontals, and
high mean number (5.5) of median azygous
head scales.
Distribution: The Bahama Islands:
known only from South Bimini (Fig. 2).
Comments: At the time of the descrip-
tion of A. d. biminiensis, Oliver had twenty
specimens of this subspecies. Additional
lizards taken since that time confirm his
diagnosis of the race. Most striking, in
comparison with all other subspecies, is the
postfrontal contact with 3/3 scales laterally
and the modal 2/ 2 supraorbitals in contact
with the inteiparietal. Data for the series
of 44 specimens are: snout scales 4 to 6
(mode 4), loreal rows 4 to 6 (mode 5);
supraorbital semicircles in contact in 30
specimens and completely separated by
median azygous head scales in 13 lizards;
modally 0/0 scales between semicircles and
interparietal and 2/2 supraorbitals in con-
tact with interparietal; fourth toe lamellae
14 to 19 (mode 16); median azygous head
scales 1 to 10 (mode 5, mean 5.5); "preoc-
ciptal" usually absent (41 of 44 lizards;
see comments below); postmentals 4 to 6
(mode 5, mean 4.8). Of the three lizards
which have the "preoccipital" present, in
one (AMNH 68638) the scale is veiy tiny
and in the second (AMNH 68637) the scale
which I consider the "preoccipital" may in
actuality be a fragment of the interparietal.
Only in one lizard (CM 32552) is there
an unequivocal "preoccipital" present. In
having such a high percentage of absence
270 Bulletin Museum of Coniparafivc Zoology, Vol. 137, No. 2
(by fusion) of the "preoccipital," biminiensis
stands alone among all subspecies of A.
disticJiiis.
Aside from the scale characters noted
aboN'c, A. (/. Inminicnsis differs from A. d.
distichus in the color of the dewlap — orange
in the former (PI. I) and yellow in the
latter. In this feature biminiensis resembles
distichoidcs; it seems very likely that the
population on South Bimini is a direct
derivative of disticJioidcs on Andros, with
resulting intensification by isolation of some
of the characters of the Andros subspecies.
A. d. biminiensis usually is a gray lizard,
but it is capable of turning a rich velvety
brown. The shade of the orange dewlap
is that of plate 9 I 10 and plate 10 L 9; all
color designations are from Maerz and
Paul, 1950. The venter is creamy to whitish
or grayish, and the underside of the tail
and hindlimbs has been noted as pale
yellow (pi. 17 J 1). Head markings and
dark body striae are usually obsolete, but
the interocular bar is at least often in-
dicated, and young lizards show both the
interocular bar and an occipital dark V.
Oliver (1948:22) noted that A. d. ])imini-
ensis was encountered at low heights on
light gray colored trees such as Fictis and
Coccothrinax. More recently ])iminicnsis
has been collected on trees in hammock
woods (high coppice) as well as on isolated
Ficiis. The absence of A. distichus from
North Bimini is puzzling. Sutcliffe (1952)
did not report the species from North Cat
Cay south of South Bimini in the chain,
but Wayne King advises me that he has
collected the species in this chain but the
specimens have been lost. Presumably the
absence ol A. d. biniinicnsis from North
Bimini (paralleled b\' that of S))Ii(i('rodacli/-
lus dccoratiis fhivicdiidiis Harbour, which
also occurs, among the Biminis, onl\ on
South Bimini) is due to a fluke of coloniza-
tion from Andros, and the li/ards haxc been
unable to cross even ihe narrow water
gap between South and North J^imini.
Specimens exdttiincd: liAIIAMA IS-
LANDS. South Bimini: no other locality,
3 (MCZ 80132-34); western end, 12 (ASFS
X4709-15, ASFS X4721-24, ASFS X4932);
western part, 2 (ASFS V10750-51); west
end, 27 (AMNII 68637-38 + 6 specimens,
AMNH 68639 + 8 specimens, MCZ 49739-
40, UMMZ 118303, CM 34118-20, CM
32549-52).
Ano/;s distichus dapsilis' subsp. n.
Holotype: MCZ 81139, an adult male,
from ocean side, opposite Hatchet Bay,
Eleuthera Island, Bahama Islands, one of a
series taken 15 June 1966 by Richard
Thomas. Original number V10385.
Faratypes (all from Eleuthera Island,
Bahama Islands ) : ASFS V10386-405, same
data as holotype; ASFS 17144-49, AHce-
town, 23 October 1961, native collector;
ASFS 17167-74, Alicetown, 24 October
1961, native collector; ASFS 17176-82,
Alicetown, 25 October 1961, native col-
lector; AMNH 96509-15, ANSP 27163-69,
CM 40623-29, KU 93380-86, MCZ 92001-08,
UIMNII 61696-700, UF FSM 21526-33,
USNM 160692-99, Alicetown, 26 October
1961, nati\e collector; ASFS 17498-500,
Alicetown, 30 October 1961, native col-
lector; ASFS 17151, Hatchet Bay (not
mapped), 24 October 1961, A. Schwartz;
ASFS V6799-800, 4 mi. (6.4 km) N Rock
Sound, 2 October 1965, R. Thomas; ASFS
V6864, 4 mi. (6.4 km) NW, thence ca. 2
mi. (3.2 km) E Rock Sound, 5 October
1965, R. Thomas; ASFS \'6811, Southeast
Point, 4 Octobc>r 1965, R. Thomas.
Definition: A subspecies of A. disticlnis
characterized b\ small si/e (males to 50
mm, females to 45 mm snout-xcnt length),
dorsum pal(> ashy gra)' with a yellowish
cast and without a green i)hase, dewlap
orange with occasionalK a \cry narrow
yellow bolder, modalK 0 0 scales between
the supraorbital semicircles and the inter-
parietal, 0 0 and 1 2 supraorbitals in con-
tact with interparietal. 2 2 scales in eon-
tact lateralK with tlu' iiostfrontals, and
I'lom Latin, r/c;/)s///.s. iilcntilnl.
Anolis DisriCHus • Schwartz
271
high mean number ( 6.2 ) of median azygous
liead scales.
Distribution: The Bahama Islands:
known only from Eleuthera Island (Fig. 2).
Comments: The holotype has the fol-
lowing measurements and scale counts:
snout-vent length 50 mm, tail 42 mm, distal
half regenerated; 4 scales across snout, 4
loreal rows, semicircles in contact, 0 0
scales between supraorbital semicircles and
interparietal, 2 2 supraorbitals in contact
with interparietal, 2 2 scales in lateral con-
tact with postfrontals, 15 fourth toe
lamellae, 4 median azygous head scales,
"preoccipital" present, 4 postmentals.
Scale counts for the series of 107 A. d.
dapsiJis are: snout scales 4 to 7 (mode
6), loreal rows 3 to 5 (mode 4); supra-
orbital semicircles in contact in 99 speci-
mens and completely separated by median
azygous head scales in six lizards; modally
0 0 scales between semicircles and inter-
parietal and 0/0 and 1/2 (both \\\\h 24
lizards ) supraorbitals in contact with inter-
parietal; 2/2 scales in lateral contact with
postfrontals; fourth toe lamellae 14 to 20
(mode 17); median azygous head scales 2
to 11 (mode 6, mean 6.2); "preoccipital"
usually present (93 of 107 lizards); post-
mentals 4 to 9 (mode 6, mean 5.2).
The dorsum of A. d. dapsiJis is usually
pale ashy gray with a yellowish cast, and
the head in adults regularly lacks any
darker markings, including the interocular
dark bar. In subadults and juveniles, the
interocular bar and occipital V are some-
what more obvious. The dorsum lacks
longitudinal dark striae but may be vaguely
streaked with darker gray. The dewlap is
completely orange, or orange with a very
narrow yellow edge; hues noted for the
dewlap are those of plate 11 C 10 and
plate 10 E 12, and the yellow border has
been noted as that of plate 10 H 3. The
eye ring is white and the eye skin gray or
tan, the latter in contrast to the gray head
and dorsum. There is no evidence that
dapsiJis has a dark brown phase, but I
assume that this color occurs. One lizard
was recorded as being pale gray with a
very faint greenish cast when caught.
A. d. dapsiJis differs from A. d. disticJms
in dewlap color (orange versus pale yel-
low) and in reaching a very slightly larger
size; in this latter context, Etheridge (1966:
351 ) reported 48 mm as the maximum size
for New Providence specimens and 53 mm
as a maximum on Eleuthera. Although
none of the 107 A. d. dapsiJis examined by
me is so long as that reported by Etheridge,
his data indicate that dapsiJis is even larger
than A. d. disticJms.
The Eleuthera subspecies resembles
distichoidcs and J)iminiensis in dewlap
color. It differs from these two more west-
ern subspecies in several ways: the 2 2
lateral postfrontal contact separates dapsiJis
from himiniensis with 3 3, and the higher
mean number of median head scales (8.7)
in disticJioidcs differentiates that form from
dapsiJis (with 6.2). A. d. Jjiminiensis and
A. d. disticJioidc's both have the supraorbital
semicircles more often separated than does
dapsiJis, and both the western subspecies
more regularly lack the "preoccipital."
The holotype and paratopotypes from
the ocean side of Eleuthera at Hatchet Bay
were taken from saplings around the edges
of an abandoned and overgrown Cocos
grove. The specimens from Alicetown were
from an edificarian situation. A. d. dapsiJis
is common on Eleuthera; I observed many
at Hatchet Bay Plantation on isolated
LysiJoma trees on the lawns and in high
coppice between Hatchet Bay and The
Glass Window. Considering the quantity
of specimens examined by me, as well as
many more in collections which I have not
studied, A. d. dapsiJis must be the com-
monest subspecies of A. disticlius in the
Bahamas.
Anolis distichus ocior^ subsp. n.
HoJotype: MCZ 81140, an adult male,
from Port Nelson, Rum Cay, Bahama Is-
lands, one of a series taken 20 June 1966
* From Latin, ocior, more rapid.
272 Bulletin Museum of Comparative Zoology, Vol. 137, No. 2
bv Albert Schwartz and Richard Thomas.
Original number V10488.
Parafi/pcs (all from Rum Cay, Bahama
Islands ) : ASFS Vl()489-90, ASFS V10493-
94, MCZ 81147-48, same data as holotvpe;
ASFS V10418-21, Summer Point, 17 June
1966, R. Thomas; ASFS V10446, Port Nel-
son, 17 June 1966, R. Thomas; ASFS
V10473, Summer Point, 18 June 1966, R.
Thomas.
Associated specimens: BAHAMA IS-
LANDS. San Salvador (localities not
mapped): no further locality, 7 (MCZ
36729-31, FMNH 222, FMNH 225-26,
FMNH 263); Cockbmn Town, 19 (ASFS
V2277, ASFS V2279-81, ASFS V2297-302,
ASFS V2355-60, ASFS V2285); 4.2 mi. (6.7
km) N Cockburn Town, 1 (ASFS V10572);
9.9 mi. (14.9 km) by road N Cockburn
Town, 1 (ASFS V10539); 7.1 mi. (11.4 km)
N Cockburn Towai, 1 (ASFS V2292); 1.2
mi. (1.9 km) N Dixon Hill, 1 (ASFS
V2278); Dixon Hill, 3 (ASFS V2286-88);
Sandy Point House, 7.6 mi. S Cockburn
Town, 10 (ASFS V10559-67, RT 1464);
2.3 mi. (3.7 km) E Watlings Castle ( =
Sandy Point House), 1 (ASFS V2339);
Green Cay, 1 (ASFS V10625); Man Head
Cay, 1 (ASFS V2337).
Definition: A subspecies of A. di.sficljus
characterized by moderate size (males to
53 mm, females to 48 mm snout-vent
length), dorsum yellow-gray to brown with
a prominent pale flank stripe between the
fore- and hindlimbs bordered above and
below by dark brown or gray and with a
pea-green phase, dewlap pale yellow,
modally 0/0 scales between the supraorbital
semicircles and the interparietal, 1/1 supra-
orbitals in contact with the interiiarietal,
2/2 scales in contact laterally with the
postfrontals, and In'gh mean number (5.8)
ol median azygous head scales.
Distribution: The Bahama Islands:
known Iroiu Rum (lay and San SaKador.
including its satellites Creen Cay and Man
Head Cay (Fig. 2).
Comments: The holotype has the follow-
ing measurements and scak> counts: snout-
vent length 53 mm, tail ca. 90 mm; 6 scales
across snout, 4 loreal rows, semicircles in
contact, 1/1 scales between supraorbital
semicircles and interparietal, 0/0 supra-
orbitals in contact with interparietal, 3/3
scales in lateral contact with postfrontals,
19 fourth toe lamellae, 7 median azygous
head scales, "preoccipital" present but
somewhat fragmented, 7 postmentals.
Scale counts for the series of 59 A. d.
ocior are: snout scales 4 to 8 (mode 6),
loreal rows 3 or 4 ( mode 4 ) ; supraorbital
semicircles in contact in 49 specimens and
completely separated by median azygous
head scales in nine lizards; modally 0/0
scales between semicircles and interparietal
and IT supraorbitals in contact with inter-
parietal; 2 2 scales in lateral contact with
postfrontals; fourth toe lamellae 15 to 19
( mode 17 ) ; median azygous head scales 2
to 10 (mode 6, mean 5.8); "preoccipitar
usually present ( 55 of 57 specimens ) ; post-
mentals 5 to 9 ( mode 7, mean 6.7 ) .
Dorsally, Rum Cay A. d. ocior varies
from unstriate gray to brown, but most
specimens observed were some shade of
green, from a grayish pastel green to a rich
pea-green. There is a complete cream
labial stripe ^^'hich extends aboxe the
shoulder and continues down th(> flank be-
tween the fore- and hindlimbs and is
bordered both above and below by dark
gray (dark gray-green in the green phase)
or brown. The green phase of ocii^r is
fairly bright, but not so bright a green as,
for example, Anolis earoli)t(')isis. In the
green phase tht>re arc no lu>ad markings,
but an occipital \' is often prcvsent in the
gray phase. The dewlap is ncIIow (PL I).
The venter is a rich >ellowish tan in all
phases, slightK' brighter (more yellow)
under the tail and along tlu^ 1ow(M' lips.
Si)ecinicns from San Salvador ditk'r Irom
those from lUmi Cay described above in
that th(\' do not sliow the gic en phase so
consistently nor ((uite so brightK'. The
cream subocular inaik is conspicuous, and
the flank stripe is jiresent but not so distinct
noi" so reuularb' bordered \\ ith darkcM" as in
Anolis distichus • Schtvartz
273
Rum Cay lizards. The dewlap is yellow
on San Salvador. In scutellation, Rum Cay
and San Salvador specimens are completely
comparable in both modes and means in
all counts taken; the largest female (48
mm) is from the small series from Rum
Cay, whereas the largest female from the
much longer San Salvador series is smaller
(44 mm). I group the lizards from these
two islands together, since it is apparent
that they are derivative populations which
together are more divergent from the
balance of the Bahaman populations than
they are from one another. Isolation on San
Salvador and Rum Cav has resulted in
some differentiation in situ, but not suf-
ficient for nomenclatorial recognition.
A. cl. ocior differs from all other Baha-
man subspecies in having a green phase.
From himiniensis, distichoidcs and dopsilis,
ocior differs in having a yellow rather than
an orange dewlap. The Rum Cay-San Sal-
vador subspecies resembles A. d. distichus
in dewlap color, but has 1/1 supraorbitals
in contact with the interparietal in contrast
to 0 0 in the nominate race, and also is
larger and has a green phase, which A. d.
distichus lacks. A. d. ocior is the only
Bahaman subspecies with 1/1 supraorbitals
in contact with the interparietal, and has
the highest mean number of postmentals
(6.7) of any Bahaman subspecies; the
highest postmental mean other than that of
ocior is that of disticJuis (5.5) among the
Bahaman subspecies.
On Rum Cay, A. d. ocior is uncommon;
all of our specimens were taken in edifi-
carian situations, especially on Cocos,
Lysiloma, and TcnninaJia in Port Nelson
and on a Lysiloma near a cottage at Sum-
mer Point. Lizards were also observed on
Thrincix palms near the beach, but not
commonlv. On San Salvador, A. d. ocior
is more abundant, but is still not so com-
mon as is A. d. distichus on New Providence
or A. d. dapsilis on Eleuthera, for instance.
Specimens were collected on Ficus and
Terminalia in Cockburn Town and were
observed on exposed fence posts in com-
pany with Anolis sagrei; the latter species
was more common in such situations. At
Sandy Point House, A. d. ocior was ex-
tremely abimdant on saplings about the
ruins and on the walh of the ruins them-
selves. On Green Cay, A. d. ocior is
moderately common on Coccoloha tangles,
and the single lizard from Man Head Cay
was taken under a flat rock among strand
plants.
Rum Cay and San Salvador stand isolated
from the Great Bahama Bank on t\\'o
separate banks of their own. Rum Cay
lies closest to Long Island (which is in-
habited by A. d. distichus), whereas San
Salvador is about equidistant from Long
Island and Cat Island (but is closer to
Rum Cay than to either of these). Aside
from Cychira rileyi Stejneger and Lepto-
typhlops columhi Klauber which are en-
demic to San Salvador, and SphaerodactyJus
corticohi Carman which occurs on both
islands, the heipetofauna of Rum Cay and
San Salvador is depauperate. Doubtless
A. d. ocior has been a long resident of these
two islands; it has diverged strikingly from
the balance of the Bahaman subspecies.
Cat Island
Cat Island, located on its own bank along
with Little San Salvador, lies southeast of
Eleuthera (which is inhabited by A. d.
dapsilis), east of the Exuma Cays (which
are inhabited by A. d. distichus) and north-
west and west of Rum Cay and San Sal-
vador (which are inhabited by A. d. ocior).
I have examined 27 A. distichus from Cat
Island, of which 14 were freshly taken by
Dennis R. Paulson. These lizards I leave un-
assigned subspecifically, although I doubt
that they merit nomenclatorial separation
from the balance of the Bahaman sub-
species.
In dorsal color and pattern, the Cat
Island lizards resemble New Providence
A. d. disticlius. They do not have a green
phase and thus are unlike ocior, but like
distichus and dapsilis. The dewlap colora-
tion is variable — more so than in any other
274 BiiUetin Museum of Comparative Zoology. Vol. 137, No. 2
Bahaman race; in a single series, Paulson
noted that three had pale yellow dewlaps,
one had a yellow dewlap with an orange
center, and the fifth had an orange dewlap
with a narrow yellow edge. Thus, in dew-
lap color, the Cat Island lizards combine
(are intermediate in?) the characters of
both disiicluis and ocior, on the one hand,
and dapsilis, on the other.
In the relationships between the inter-
parietal and supraocular semicircles, the
Cat Island lizards are not distinctive and
resemble both distichus and dapsilis but
not ocior (which modally has 1/1 supra-
orbitals in contact with the interparietal,
in contrast to 0/0 or 1/2 in disHchus and
dapsilis). The "preoccipital" is absent in
six of 24 lizards; this is a higher proportion
than dapsilis, distichus or ocior. The mean
of median head scales is 4.8, in strong con-
trast to 6.2 in dapsiJis, 6.0 in distichus, or
5.8 in ocior. The postmental mean is 6.0,
higher than both disticJitis and dapsilis, but
lower than ocior. The postfrontal contact is
bimodal, with both 2/2 and 3 3 having
equal frequencies of eight lizards; there is
also a strong tendency (as intimated by
the bimode of 3/3) for Cat Island lizards
to have 3/4 and 4/4 scales in contact lat-
erally with the postfrontals ( 18 of 26 lizards
have three scales in contact unilaterally),
whereas counts above 2/3 are relatively
uncommon in distichus (22 of 128 lizards),
dapsilis (five of 102 lizards) and ocior
(six of 59 lizards). Such high lateral post-
frontal contact counts are more usually
encountered in ])iminiensis (23 of 42
lizards). It should be recalled that 3/3 is
the modal condition in himinicnsis.
On the basis of dewlap coloi', it would
seem appropriate to consider the Cat Is-
land lizards intermediate betw(>en distichus
and da))silis, and the geographic position ol
Cat Island is in accord with a possibly
double "invasion" of lizards from the is-
lands to the n()rth\\'est and west. 1 can
see no ocior inlluencc in the (-at Island
lizards. In contrast to the situation \\'ith
the dewlap color, the scale counts pres(^nt
a peculiar melange of characters which
cannot reasonably be attributed to inter-
action of the two adjacent races. It is
probable that Cat Island has been colonized
at \'arious times by both distichus and
dapsilis, but that there has been imposed
upon these two parent stocks other local
differentiation on Cat Island, so that the
Cat Island lizards resemble their parent
stocks in some characters but have diverged
considerably in others.
Specimens examined: BAHAMA IS-
LANDS. Cat Island: Orange Creek, 7
(ASFS V2145-51); Arthur's Town, 7 ( MCZ
39580-83, UMMZ 79449); Bennett's Har-
bour, 4 (AMNH 76337-40); Tea Bay, 2
(ASFS V2159-60); The Bight, 5 (ASFS
V2188-91, CM 20444); hills above The
Bight, 1 (ASFS V2123); 1 mi. (1.6 km) S
McQueen, 1 (AMNH 76336).
Anolis distichus c/om/n/cens/s Reinhardt and
Lijtken
Auolis ilominicen.sis Reinhardt and Liitken, 1863,
Vid. Mc'dd. Nat. Foren. Kjobenhavn: 261.
Anolis disticlius (lU)i(Iogitlaris Mertens, 1939, Abh.
St'nckenl)er,ij;. Naturf. Ges., 449:59.
Tij])e localitij: Haiti; restricted to Port-
au-Prince, Dept. de lOuest, Haiti.
Definition: A subspecies of A. distichus
charactt^rized by very large size (males to
58 mm, females to 48 mm snout-vent
length), dorsum varying between all green
and all dark brown with darker longitudinal
striae in all phases, dewlap pale yellow
(occasional!)' \\hitc or almost so) to yel-
low with a hunt orange basal blush,
modally 11 scales between the supra-
orbital semicircles and the interparietal, 0 0
supraorbitals in contact with th{> inter-
parietal, 2/2 scales in contact laterally with
the postfrontals and moderate mean num-
ber (3.9) ol median azNgons h(>ad scales.
Distribution: All of Ilaiti with the ex-
ception ol tlu' Tiburon Peninsula W(\st of
Miragoane (precise limits along the south-
ern coast ol the Tiburon Peninsula at the
longitnde of Miragoane unknown); the
Hepi'ibliea Dominicana in extrcMue western
Angus distic.iivs • Schwart:
275
70"
-tor
-•^
P
I
•5 't^.^ • ^^»
t
J
70-
n to so 40
Figure 3. Map of Hispaniola, showing distribution of tfie subspecies of Ano//s disf;chus; fine diagonal lines, A. d.
dommicensis; horizontal lines, A. d. ignigularn; diagonal lines, upper right to lower left, A. d. properus; open stippling,
A. d. ravitergjm; crosshatching, A. d. favillarum- close stippling, A. d. aurifer; vertical lines, A. d. vinosus; diagonal
lines, upper left to lower right, A. d. suppor; P, A. d. patruelis; J, A. d. juliae; T, A. d. tostus; S, A. d. se|unctus.
Pedernales Province on the south, through
extreme western Independencia Province,
thence east through San Juan Proxince to
northern La Vega Province (Jarabacoa),
Sanchez Ramirez Province (Cotui), San
Cristolial Province (Gonzalo), and Samana
Province (mouth of the Rio Yuna), and
north to the northern coast in Maria Trini-
dad Sanchez Province (Cabrera), but ex-
cluding the Peninsula de Samana; possibly
the He de la Tortue off the northern coast
of Haiti (Fig. 3).
Comments: A. d. dominicensis has the
widest distribution of any of the His-
paniolan subspecies. Throughout this wide
range, it is remarkably constant in dewlap
and dorsal colorations. The dewlap is most
often pale yellow (PI. I), but at times (and
not segregated geographically) there is a
vague and pale orange basal blush on the
otherwise yellow dewlap. Occasional speci-
mens (for example, in the Sierra de
Baoruco in Pedernales Province and at
Cap-Haitien in northern Haiti) have the
dewlap very pale yellow to practically
white.
In the green phase, the ground color is
fairly bright and marbled and/or streaked
with green, brown, black, or yellow. The
head is pale green, and the ventral color
varies from pale green to gray or even
black. The underside of the tail ranges
from bright yellow to yellow-orange. In
the brown phase, the back is a rich choco-
late or wood brown; some specimens seem
incapable of achieving a uniform brown
and have a marbled or mottled pattern of
darker and lighter browns. There is an
intermediate color phase ( greenish tan or
grayish brown), which presumably is as-
sumed between the definitive green or
solid brown conditions.
Scale counts of the series of 245 A. d.
dominicensis are: snout scales 4 to 10
(mode 4), loreal rows 3 to 7 (mode 4);
supraorbital semicircles always in contact;
modally 1 1 scales between the semicircles
and the interparietal and 0 0 supraorbitals
in contact with the interparietal; 2/2 scales
in lateral contact with postf rentals; fourth
toe lamellae 14 to 24 (mode 19); median
azygous head scales 2 to 11 (mode 3,
mean 3.9); "preoccipital" almost always
present (235 of 251 lizards); postmentals
276 Bulletin Museum of Comparative Zoology, Vol. 137, No. 2
2 to 12 (modes 6 and 7, mean 6.6). The
largest males are from the higher elevations
in the Montagne Noire. These elevations
(5000 and 5600 feet— 1525 and 1708 meters)
are among the very highest at whieh A.
disticJius has been collected. A. d. domini-
censis also occurs at sea level in many
coastal situations, and below sea level in
the extreme eastern Cul de Sac Plain.
The specimens from the eastern Cul de
Sac Plain are of special interest. In this
region (Thomazeau, Manneville), the dor-
sal coloration is like that of specimens from
the uplands on the northern slopes of the
Morne 1 Hopital ( Petionville, Morne Cal-
vaire, for instance ) , but the dewlap color
is a deep orange (pi. 4 C 11, pi. 4 G 10
and pi. 4 G 11), at times with a faintly
brown cast. This is one of the regions
where A. distichus is sympatric (but not
syntopic) with A. brevirostris, which in this
same area has an orange dewlap. Of the
two species in the Thomazeau-Manneville
area, A. ])icviiostris is distinctlv the in-
habitant of the xeric scrub and A. disticlius
the inhabitant of more mesic situations,
oases, and cultivated areas. In this region,
A. hrevirostris is the widespread lizard of
open areas, whereas A. distichus is re-
stricted to certain less rigorous habitats and
is in effect surrounded by A. hrcvirustiis.
The orange dewlap of A. disticJius may well
be the result of partial or complete isola-
tion of the A. disticJius populations from
the balance of the species. To the east, in
the Valle de Neiba in the Republica
Dominicana, A. d. doniiiiiccnsis remains un-
known, but is replaced to the east of Lago
Enriquillo by the western extreme of an-
other subspecies. Doubtless A. d. domini-
censis will ultimately be collected between
the Dominico-Jlaitian liorder and the west-
ern end oi Lago liinric^uilJo in the Re-
publica Dominicana.
Considering tlu^ fact that A. d. domini-
censis occurs at a great \ariety of eleva-
tions, it is obvious thai it also occupies a
great variety of habitats, from the hot
oases in the Cul de Sac Plain to upland
mesic cacao groves and rain forest. Cul-
tivated lands are quite suitable, and it is
often the dominant anole of shady fence
rows and the interior of humid coffee plant-
ings and woods. Along the coast it occurs
in mangroves (Trou Forban), mesic and
open banana-breadfruit-royal palm associa-
tions, on large trees in open cultivated
semi-arid regions, and it is common almost
everywhere, at least where a minimal patch
of shady woods occurs. In the hot and dry
Valle de Cibao, A. d. dominicensis was en-
countered in thorn and tree-cactus scrub
but in the more shady situations. In short,
throughout its broad Hispaniolan range,
A. d. dominicensis is encountered in almost
any situation which offers shade and refuge.
I have noted in the introduction the
sleeping habits of A. d. dominicensis at
Miches and Palo Verde in the Republica
Dominicana. One other observation has
been made; two A. d. dominicensis were
taken sleeping exposed on a large, wet log
lying adjacent to a rushing stream, in a
deep and cool montane ravine at 2200 feet
(671 meters), near Puesto Grande in the
Cordillera Septentrional in northern Re-
publica Dominicana.
I do not regard A. d. aJJjidoiiularis
Mertens as a valid subspecies. Mertens
was misled into the description of (dJ)ido-
ii.uJ(iris by the material which he regarded
as dominicensis from Haiti; his ^'domini-
censis' were three males and a female
from Gonaives, and two males and a fcMiiale
from St. Marc. His comments (1939:56)
on the distinguishing characters of A. d.
dominicen,sis (as based upon these seven
specimens) do not apply to dominicensis —
i.e., that dominicensis is ne\cr green but
rather is gray to gray-brown, has a bright
and clear supralabial streak, a pair of dark-
scapular spots and a lined dorsum, and a
chrome-orange-yellow dewlap with citron-
yellow scales. These are precisely the
characters — esp(>eially the alw ays gray color
and the pair of scapular spots and a lined
dorsum — which distinguish the species A.
l>r('rirosiris Irom .A. dislicJms. It is ajiiiarcMit
Anolis Di^riciius • Schwartz
277
that Mertens, when describing albidogularis,
did not have for comparison specimens of
A. d. dominicensis, as he presumed, but
rather A. hrevirosirls. The characters of
aJhidoii,ul(iris are those of dominicensis,
and specimens from the vicinity of the type
locaHty (Monte Cristi, RepubHca Domini-
cana) do not differ significantly in any
feature from topotypical Port-au-Prince ma-
terial. The pale dewlap coloration which
is ascribed to albidogularis is not consistent
in the Valle de Cibao population and oc-
curs only sporadically elsewhere; specimens
which I have collected near Monte Cristi
and in the Valle de Cibao have the dewlap
color pale yellow, as do specimens from
elsewhere within the range of A. d. domini-
censis.
A. d. dominicensis differs from all the
Bahaman subspecies except ocior in having
a green phase; the green of ocior is a much
more yellow-green than the green of
dominicensis. Of the Bahaman subspecies,
all are smaller than dominicensis; ocior
most closely approaches dominicensis in
size. The median head scale mean of
dominicensis (3.9) is lower than that of any
Bahaman race (5.5 to 8.7). Only distichoides
in the Bahamas has the 11 scales between
the semicircles and the interparietal as does
dominicensis. Other head scale differences
( such as the regular presence of the "preoc-
cipital" and the regular contact between the
semicircles in dominicensis) are also sig-
nificantly different in comparison with the
Bahaman subspecies.
A. d. dominicensis presumably comes into
contact with four other subspecies. In one
of these instances (Sierra de Baoruco) no
intergradient specimens are known, since
there is an hiatus between the closest rec-
ords of dominicensis and this next adjacent
form to the east. In three instances, how-
ever {ignigularis; the subspecies to the
west on the Tiburon Peninsula; and the
subspecies to the east in the Valle de
Neiba), there are samples which I interpret
as intermediates. In the case of ignigularis,
the material from the higher elevations in
the eastern portion of the Cordillera Central
( vicinity of Constanza, Paso Bajito, etc. )
shows the dewlap rather intermediate be-
tween the yellow or yellow-with-orange-
blush dominicensis condition and the solid
orange dewlap with a narrow yellow border
of ignigularis, although the dewlap in gen-
eral is much closer to that of ignigularis
than to that of dominicensis. I have in-
cluded these Cordillera specimens with
ignigularis for that reason.
A small series from Padre las Casas, Azua
Province, Republica Dominicana, I interpret
as intergradient between dominicensis and
the Valle de Neiba-Llanos de Azua sub-
species. This lot is closer to the latter race,
and I have discussed it in detail there.
Finally, lizards from the vicinity of Saint
Michel du Sud on the Tiburon Peninsula
are intermediate in dewlap color between
dominicensis and the next adjacent race
to the west on the Peninsula (which has a
deep orange dewlap with a narrow yellow
edge ) , but they are closer to the latter sub-
species, and I have included them in the
discussion of that race rather than with
dominicensis.
I have seen no fresh material from He
de la Tortue and only three old specimens
which are distinctive in neither scutellation
nor what is discernible of pattern or pig-
mentation. I include Tortue in the range
of A. d. dominicensis only provisionally,
since on all other satellite islands where
A. distichus is found, it is racially distinct.
Thus there is a good likelihood that fresh
specimens from Tortue will demonstrate
that there is a different subspecies present
there.
Referred specimens: HAITI. Dept. du
Slid: Miragoane, 30 (MCZ 25489-98 + 20
untagged specimens); Butete, nr. Miragoane
(not mapped), 7 (MCZ 6613.3-39); Etang
Miragoane, 7 (MCZ 66140-46). Dept. de
TOuest: 7.1 mi. (11.4 km) E Miragoane,
1 (ASPS X3850); 3 mi. (4.8 km) W Grand
Goave, 300 feet (92 meters), 1 (ASFS
X3856); 1.1 mi. (1.8 km) NE Fauche, 2
(ASFS X2045-46); 5 km S Dufort, 4 (MCZ
278 Bulletin Museum of Comparative Zoology. Vol 137, No. 2
63099-102); 4 mi. (6.4 km) SE Leogane, 4
(ASFS V8463-66); Leogane, 2 (MCZ
13779-80); ga Ira, 9 (MCZ 63898-906);
bridge over Riviere Momance on road to
Leogane, 1 (MCZ 63103); Mariani, 7 mi.
(11.2 km) E Gressier, 7 (ASFS V8446-52);
Diquini, 17 (MCZ 59430-32, MCZ 8696-
700, MCZ 8703, MCZ 8705, MCZ 8710,
MCZ 8712, MCZ 8714-18); Port-au-Prince,
1 (MCZ 51427); Boutillier Road, S of Port-
au-Prince, 17 (MCZ 59413-29); SVV of
Port-au-Prince (not mapped), 1 (MCZ
51258); 2.8 km S Peticmville, 1700 feet
(519 meters), 2 (ASFS V81 17-18); 5 mi.
(8.0 km) NE Petionville, ca. 160 meters,
3 (ASFS V9405-07); 3 km (airline) W
Petionville, Morne I'Hopital, 920 meters, 11
(ASFS V843.5-45); Morne Calvaire, 1 mi.
(1.6 km) SW Petionville, 2300 feet (702
meters), 44 (ASFS X1237-80); Kenscoff, 2
(MCZ 45745, MCZ 59401); Morne Bourette
(not mapped), 2 (MCZ 47546 -f one un-
tagged specimen ) ; Peneau, 5000 feet ( 1525
meters), 4 (ASFS X1350-51, ASFS X1574-
75); Furcy, 5600 feet (1708 meters), 45
(ASFS X1591-95, MCZ 63535-39, MCZ
59393-97, MCZ 59433-41); Peneau and
Furcy, ca. 4000-5000 feet (1220-1527
meters), 4 (ASFS V4821-44); Hatte Lathan
(not mapped), 2 (MCZ 51421-22); Thom-
azeau, 4 (MCZ 13771-72, MCZ 37455,
USNM 59191); near Thomazeau, 2 (MCZ
37495-96); Tete Source, 1.4 km NNE
Thomazeau, 3 (ASFS V8173-75); Manne-
ville, 9 (ASFS V8194, CM 38881, MCZ
59390-92, MCZ 63107-10); Ste. Philomene
(not mapped), 1 (MCZ 51428); 3.9 mi.
(6.2 km) NW Ganthier, 1 (ASFS X2171);
Gormand, nr. Saltrou (not mapped), 2
(MCZ 68614-15); Colomhier, iir. Saltrou,
4 (MCZ 68616-19); Lan Hanane, nr.
Saltrou, 5 (MCZ 68620-24); Tete a n-:au.
nr. Saltrou, 6 (MCZ 68625-30); Thiotte,
nr. Saltrou, 9 (MCZ 69631-39); Caroye, nr.
Saltrou (not mapped ), 31 ( MCZ 69315-45);
Londry, nr. Saltrou (not mapped), 4 (MCZ
69346-49); Citadelle, m-. Saltrou (not
mapped), 15 (MCZ 69350-64); Maviet(>,
nr. Saltrou (not maiijied), 15 (M('Z
69365-79); Mapou, in-. Saltrou, 7 (MCZ
69380-86); ca. 3.5 mi. (5.6 km) NE Trouin,
800 feet (244 meters), 1 (ASFS V9664);
5 mi. (8.0 km) S Trouin, 700 feet (214
meters), 3 (ASFS V9668-70); Jacmel, 1
(ASFS V9825); ca. 5.5 mi. (8.8 km) NW
Jacmel, 600 feet (183 meters), 1 (ASFS
V9784); 3 mi. (4.8 km) E Jacmel, 2
(ASFS V9757-58); ca. 1 mi. (1.6 km) W
Cayes Jacmel, 4 (ASFS V9700-03); 10 mi.
(16.0 km) NNE Marigot, 3200 feet (976
meters), 1 (ASFS V9737); Trou Forban, 1
(ASFS V8216); 1.6 km SW Trianon, 1100
feet (336 meters), 3 (ASFS V8278-80); 1.6
km NE Trianon, 6 (ASFS V8282-87); 7
mi. (11.2 km) N Mirebalais, 1 (ASFS
X2234); La Tombe, nr. Mirebalais (not
mapped), 21 (MCZ 68204-24); Fer-a-
Cheval, nr. Mirebalais, 5 (MCZ 68225-29);
Boudou, nr. Mirebalais (not mapped), 13
(MCZ 68230-42); Ledie, nr. Mirebalais
(not mapped), 4 (MCZ 68243-46); Dubui-
son, nr. Mirebalais (not mapped), 3 (MCZ
68247-49). Dept. dc rArtibonitc: south
end, Etang Bois Neuf, 1 (MCZ 59942);
Pierre Payen, 8 (MCZ 59402-03, MCZ
59107-12); bridge over Riviere de FArti-
bonite, St. Marc road, 2 (MCZ 59404-05);
Passe Peine, 3 (MCZ 63055-57); 8 to 9
km W Marmelade, 3500 feet ( 1068 meters),
2 (ASFS V9913-14); 5 mi. (8.0 km) NW
St. Michel de TAtalaye, 4 (ASFS V10030-33);
2 mi. (3.2 km) NW St. Michel de I'Atalave,
2 (ASFS V10034-35); Hinche, 5 (MCZ
25499-503); C;r()s Morne, 8 (MCZ 63075-82).
Dcpt. du Noid: 3 mi. (4.8 km) NW
Terrier Rouge, 1 (ASFS \T()163); Dondon,
10 (MCZ 6306:3-72); Dondon, southeastern
outskirts, 4 (ASFS V10017-20); ca. 2 km
S Dondon, 2 (ASFS \'10038-39); Grande
Riviere du Nord, 13 ( MC:Z 66655-67); Cap-
llaitien, 94 (MCZ 37483-92 + 69 untagged
specimens, MCZ 63058-62, ASFS \T0194-
204); Ti Guinin, near Cap-IIaitien (not
mapp(>d), 8 ( MC:Z 66668-75); Citadelle
Laferriere, 7 (MCZ 33370, MCZ 6307:3-74,
MCZ 66651-54); m-. Citadelle Laferriere,
2 (MCZ 25487-88); 4 mi. (6.4 km) SSW
Limbe. 200 feet (61 meters), 1 (ASFS
Anolis DiSTiCHUs ' Schwartz
279
V9964); 4 mi. (6.4 km) N Port Margot,
east side of Riviere de Port Margot, 1
(ASFS V9971); ca. 2 km inUmd from Anse
a Margot, 1 (ASFS V10277); Chouchou, 10
mi. (16.0 km) NW Port Margot, 6 (ASFS
V9978-83); 1 mi. (1.6 km) SW Le Borgne,
west side Riviere du Borgne, 2 (ASFS
VlOOOl-02); Dept. du Nord Quest, Jean
Rabel, 1 (MCZ 63098); Bombardopolis, 15
(MCZ 63083-97). Ik dc la Toiiue: 3
(MCZ 37493-94, USNM 95121). RE-
PUBLICA DOMINICANA. Pedcmales
Prov.: 19 km N Pedernales, 1000 feet (305
meters), 1 (ASFS V2702); Las Mercedes,
ca. 1400 feet (427 meters), 1 (ASFS
V2659); 1 km S Los Arrovos, 4100 feet
(1251 meters), 1 (ASFS V2605); 27 km
N Puerto de Alcoa, 1 (ASFS X9765). In-
dependencia Prov.: Aguacate, 3 (MCZ
58467-69); 8 km E Aguacate, 1600 feet
(488 meters), 1 (ASFS X9945); 7.6 km
NW La Descubierta, ca. 2000 feet (610
meters), 2 (ASFS V4375-76); Guayabal,
6 km N Postrer Rio, 4 (MCZ 58470-73).
San Rafael Prov.: 18 km SW Hondo Valle,
6000 feet (1830 meters), 1 (ASFS V360);
9.0 mi. (14.4 km) NW Elias Pina, 1 (ASFS
V330); Rancho La Guardia, 13 (MCZ
58441-53); Pedro Santana, 1 ( MCZ 58440);
Banica, 1 (MCZ 58438); 3 km E Banica,
1 (MCZ 58439); 3 km NE Banica, 13
(MCZ 58454-66). San Juan Prov.: San
Juan, western edge, 6 (ASFS V499-504);
15 km SE San Juan, 4 (ASFS V487-90);
3 km E Las Matas, 4 (ASFS V305-08);
Rio Arriba del Norte, 1950 feet (595 meters),
3 (ASFS V521-23); 7 km N Carpintero,
9 (MCZ 58500-08); 7 km NW Vallejuelo,
2600 feet (793 meters), 3 (ASFS V302,
ASFS V394-95). La Vega Prov.: Jarabacoa,
2 (MCZ 58480-81); 3 km NE Jarabacoa, 1
(ASFS V1948). Sanchez Ramirez Prov.:
12.3 km E Cotui, 5 (ASFS V611-15). San
Cristobal Prov.: 10 km NE Gonzalo, 600
feet (183 meters), 2 (ASFS V3131-32).
Sarnand Prov.: south side of Rio Yuna,
approximately 1 km upstream from mouth,
7 (ASFS V2961-67). Maria Trinidad
Sanchez Prov.: 11.2 km S Cabrera, 3 (ASFS
V4244-46). Duarte Prov.: 1 km NW
Arensoso, 3 (ASFS V1841-43). EspaiUat
Prov.: 2 km N Puesto Grande, 2200 feet
(671 meters), 2 (ASFS V1962-63). Puerto
Plata Prov.: Puerto Plata, 2 (MCZ 5442,
MCZ 43670); Sosua, 8 (ASFS V1631-32,
MCZ 13754-59); 6 km E Imbert, 700 feet
(214 meters), 2 (ASFS V1691-92). Santi-
ago Prov.: Santiago, 1 (MCZ 58482);
Licey al Medio, 4 (MCZ 58317-20);
Ceboruco (not mapped), 12 (MCZ
58483-94); 3 km S Pena (not mapped),
5 (MCZ 58495-99); 6 km E El Rubio, 1000
feet (305 meters), 2 (ASFS V2922-23); 7
km SE El Rubio, 2300 feet (702 meters),
1 (ASFS V2924). Valve rde Prov.: 7 km
E Valverde, 2 (ASFS V2954-55). Monte
Cristi P) w.: 24 km E Monte Ciisti, 1
(MCZ 43681); 4 km E Pepillo Salcedo, 1
(ASFS V1167); 2 km NE Palo Verde, 10
(ASFS V130.3-12); 1 km S Palo Verde, 4
(ASFS V1357-60). Dajabon Prov.: 6 km
S Copey, 1 (ASFS VI 170); 1 km S Loma
de Cabrera, 900 feet (275 meters), 1 (ASFS
V1171).
Anolis distichus ignigularis Mertens
Anoli'i distichwi i^ni<iiilaris Mertens, 1939, Abh.
Senckenbers. Naturf. Ges., 449:58.
Type locality: San Pedro de Macoris,
San Pedro de Macoris Province, Republica
Dominicana.
Definition: A subspecies of A. distichus
characterized by moderate size (males to
55 mm, females to 44 mm snout-vent
length), dorsum usually green anteriorly
and rich and translucent reddish tan pos-
teriorly ( but capable of turning completely
brown), dewlap vivid orange centrally
with a narrow yellow margin, modally 0/0
scales between the supraorbital semicircles
and the interparietal, 1 2 supraorbitals in
contact with the interparietal, 2/2 scales
in contact laterally with the postfrontals,
and low mean number (3.5) of median
azygous head scales.
Distribtition: The Republica Dominicana
from eastern San Cristobal Province in the
west, east along the coast to the type
280
Bulletin Museum of Comparative Zoology, Vol. 137, No. 2
locality, thence inland to the vicinity of
Higiiey and to the north coast (east of
Miches ) in La Romana Province; along the
north coast to the Bahia de San Lorenzo
in El Seibo Province, south into eastern San
Cristobal Province ( Bayaguana ) , and west
into the Cordillera Central; Peninsula de
Samana, west to the vicinity of Yayales
(Fig. 3).
Comments: The dewlap and dorsal
colors of A. (1. ig.ni fibular is are very constant
throughout the range of the subspecies.
The vivid and extensive orange center and
narrow yellow margin of the dewlap (PI.
I ) are diagnostic features of iii,nifi,uloris
in the eastern and central portions of the
Rcpublica Dominicana. The dorsum is usu-
ally a rather dark green anteriorly, grading
rapidly into a translucent reddish tan pos-
teriorly. The lizards can become completely
brown, although this brown is of a more
reddish shade (cinnamon) than that of
A. (1. (lominiccnsis. The extent of the orange
center of the dewlap is slightly variable,
but the bright pigment is never restricted
to a small and indistinct orange blush, as
it is occasionally in dominicensis.
Scale data on the series of 103 A. d.
i^nii^idaris are: snout scales 4 to 8 (mode
4 ) , loieal rows 3 to 6 ( mode 5 ) ; supraorbital
semicircles always in contact; modally 0/0
scales between the semicircles and the
interparietal and 1/2 supraorbitals in con-
tact with the interparietal; 2/2 scales in
lateral contact with postfrontals; fourth
toe lamellae 14 to 22 (mode 20); median
azygous head scales 1 to 9 (mode 3, but
4 scales are almost equally as common,
mean 3.5); "preoccipital" usually present
(100 of 103 lizards); posimc>ntals 4 to 10
( mode 5 or 6, mean 6.0 ) . The asymmetrical
mode of 1/2 supra()rl)itals in contact with
the interparietal is pecuhar, but the mode
is fairly strong (34 individuals; next highest
category is 0/0 with 26 lizards). Judging
from the high incidence (26) of 1/1
supraorbitals in contact with the inter-
parietal, I suspect that iii,ui^ularis is a pop-
ulation which is in the process of exolxing
from a mode of 11 to 2/2 but has not
completed the transition.
The largest males have snout-vent lengths
of 55 mm, and both are from the vicinity
of Higiiey; the status of that particular
population is probably intergradient be-
tween igni(i.ularis and the race next to the
southeast, but these two large males are
clearly much more like ifiuigularis than the
drab southern form. The largest female
ignigidaris has a snout-vent length of 44
mm; this individual, from the Valle de
Culata, at an elevation of 5000 feet (1525
meters) in the Cordillera Central, is from
an area of extreme intergradation with
dominicensis.
A. d. igniguhiris is readily separable
from all previously discussed subspecies;
the combination of orange dewlap and
bicolor dorsum occurs in no other form.
Comparison with the three orange-dew-
lapped Bahaman subspecies, himinicnsis,
distichoidcs, and dapsiUs, is easily made.
Aside from these three races lacking the
bicolored dorsum, all are smaller, have
much higher median head scale means ( 5.5
to 8.7 in contrast to 3.5 in ignigularis),
lower means of postmentals (4.6 to 5.2 in
contrast to 6.0), and have a high percentage
of specimens which lack the "preoccipital."
Of the three Bahaman subspecies, only
dapsilis modally has 0 0 scales between
the supraorbitals and the interparietal and
12 supraorbitals in contact with the inter-
parietal as doc\s ignigularis. In d(i})silis,
however, 1 2 is one of two bimodes. Ex-
tended comparisons witli dominicensis are
not necessarv'; th(> dewlap and dorsal colora-
tions arc sufficient to distinguish the two
races. The 1/1 scales between the semi-
circl(\s and the interparietal and 0 0 supra-
orbitals in contact with the interparietal in
dominicensis diller from the conditions of
()() and I 2 in iis.)iiii.ul(iris.
The api^arcntU' disjunct range of igni-
gtdaris is oi especial interest. Were it
not lor the series (seven specimens) from
the mouth of (h(^ l^io Yuna, I would con-
sider that iiiiiiiLuhiris has a continuous dis-
Anolis distichus • Schwartz
281
tribution about the western end of the
Bahia de Samana. However, the Rio Yuna
lizards are clearly dominicensis and have
the pale yellow dewlaps of that subspecies.
Although there is evidence (Cochran,
1941:2) that the Peninsula de Samana was
in historic time an island separated from
the mainland, this seems hardly likely when
the isthmus is visited today, since, although
it is low-lying and swampy and is bisected
b\' the canos de Gran Esfcro, it is also
heavily forested, and it seems doubtful that
the Samana has been completely severed
from the mainland so recently. Doubtless
the Peninsula has been completely insular
at various times in the past. It seems
possible that ignigularis invaded the
Samana across the Bahia de Samana from
the south, while the former was cut off
from the balance of Hispaniola, and be-
came established there, rather than having
reached the Peninsula around the ^^'estern
end of the Bahia. Another possibility is
that dominicensis has followed down the
Rio Yuna from the interior and has invaded
the area at the head of the bay, thereby
severing the two components of the igni-
gularis population. Larger numbers of
specimens from this immediate area may
demonstrate intergradation; the series at
hand from the mouth of the Rio Yuna,
however, does not show it.
Intergradation between ignigulaiis and
dominicensis occurs in the eastern Cordil-
lera Central, although specimens from the
foot of the eastern escarpment of the
Cordillera (vicinity of Monseiior Nouel)
are clearly ignigularis. Specimens from the
area about Constanza and Paso Bajito are
much like ignigularis, except that the dew-
lap orange is somewhat paler (although
usually very extensive), and the dorsum
is more regularly all green rather than
sharply bicolor. These specimens I regard
as closer to ignigularis and have so listed
them below.
A. d. ignigularis comes into contact with
two other subspecies, that to the south and
east in the La Romana to Cabo Engaiio
region, and that to the southwest in the
Llanos de Azua. No intergrades are known
for the latter contact, and the break be-
tween the two subspecies must be rather
sharp (see comments below). The four
lots of fresh material from the vicinity of
Higiiey and Bejucal are much closer to
ignigularis than to the subspecies to the
south, although the series from 2 miles south
of Higiiey has one male with a yellow
dewlap without any orange. Taken as a
whole, the Higiiey and Bejucal material
is close to ignigularis. In two other areas,
the intergrades between these two forms
are closer to the unnamed subspecies and
will be discussed belo\\'.
Like A. d. dominicensis, ignigularis has
a wide altitudinal range, from sea level to
elevations of at least 6000 feet (1830
meters) in the Cordillera Central. In the
lowlands, it is a customary denizen of moist,
shady cacao groves and other wooded
situations. In the Cordillera it is encoun-
tered most frequently in heavily wooded
ravines and local stands of rain forest, al-
though at Valle de Culata it was found on
a rail fence in an exposed and abandoned
pasture. It does not occur commonly in
the pine woods in the highlands.
Specimens examined: REPlJBLICA DO-
MINICANA. San Cristobal Prov.: 15.5
km SE El Cacao, 1400 feet (427 meters),
1 (ASFS V2463); El Tablazo, nr. Rio
Nigua, 15 km NW San Cristobal, 7 ( MCZ
58714-20); La Cabirma de la Loma, north-
west of San Cristobal, 4 (MCZ 79269-72);
Colonia Ramfis ( = La Cabirma de la
Loma ) , 5 ( MCZ 58721-22, MCZ 58566-68 ) ;
1 km NW Colonia Ramfis ( not mapped ) , 5
(MCZ 58561-65); 3 km SE Colonia Ramfis
(not mapped), 7 (MCZ 58569-75); 6 km
SE Colonia Ramfis (not mapped), 4 (MCZ
58576-79); 9 km SE Colonia Ramfis (not
mapped), 5 (MCZ 58580-84); 12 km SE
Colonia Ramfis, 5 (MCZ 58585-89); 15 km
SE Colonia Ramfis ( not mapped ) , 3 ( MCZ
58590-92); 7 km N San Cristobal, 6 (MCZ
58593-98); Mt. Calabozo, near San Cristobal
(not mapped), 3 (MCZ 58599-601); 2 mi.
282 Bulletin Museum of Comporative Zoology, Vol. 137, No. 2
(3.2 km) SE San Cristobal, 2 (ASFS
X7774-75); 3 km W Bayaguana, 4 (ASFS
V602-06); 10 km NE Bayaguana, 1 (ASFS
V3141); Comate, Municipio Bayaguana, 5
(MCZ 79286-90); Monte Plata, 1 (MCZ
16441). Distrito Nacional: 9.8 mi. (15.7
km) E Santo Domingo, 5 (ASFS X7735-39);
Santo Domingo, 8 (MCZ 53945, MCZ 58655,
MCZ 58708, MCZ 75185-86. MCZ 79266-68).
San Pedro de Macoris Prov.: 6 km N San
Pedro de Macoris, 2 (ASFS X7832-33).
La Romana Prov.: Bejucal, 5 (MCZ
58602-06); 1 mi. (1.6 km) NE Higiiey, 5
(ASFS V771-75); 2 mi. (3.2 km) S Higiiey,
4 (ASFS V747-50); 6.6 km W. Higiiey, 1
(ASFS V1013); 24.8 mi. (39.7 km) ESE
Miches, 2 (ASFS X7891-92). EJ Seibo
Prov.: 1.4 mi. (2.2 km) SE Miches, 1
(ASFS X9349); 14 km SW Miches, 8 (MCZ
75187-94); 6.6 mi. (10.6 km) NW Hato
Mayor, 2 (ASFS X7871-72); San Francisco,
6 km SE Hato Mayor, 1 (MCZ 58614); 2.1
mi. (3.4 km) N El Valle, 2 (ASFS X7866-67);
Sabana de la Mar, 42 (ASFS V3081-98,
MCZ 58615-38); 3.5 mi. (5.6 km) S Sabana
de la Mar, 7 (ASFS X7841-44, ASFS
X7930-32); 20 km S Sabana de la Mar, 11
(MCZ 58639-49); Cueva de Cano Hondo,
5 (ASFS X9284-88); Bahia de San Lorenzo
( small beach west of railway bed ) , 2 ( ASFS
V3150-51). La Vcm Prov'.: Paso Bajito, 1
(ASFS X8787); 7 km E Paso Bajito, Casa
de los Michelenas, 3 (ASFS X8781-83); El
Rio, 6 (MCZ 64371-76); 7.1 mi. (11.4 km)
E El Rio, 3500 feet ( 1068 meters), 1 (ASFS
X8112); Constan/.a, 9 (MCZ 44387, MCZ
58652-54, MCZ 58709-13); 9.1 mi. (14.6
km) N Constanza, 6000 feet ( 1830 meters),
3 (ASFS X8487, ASFS X8700-01); 9 km N
Constanza, 1 (ASFS X8699); 5.1 mi. (8.2
km) N Constanza, Valle de Culata, 5000
feet (1525 meters), 11 (ASFS X8488-98);
6 km W Constanza, 4250 feet ( 1296 meters),
2 (ASFS X88.32-.33); Loma Vieja, 1 (MCZ
44383); Paraje La Palma, Municipio Con-
stanza (not mapped), 41 (MCZ 75153-79,
MCZ 79273-85); El Convento, Municipio
Constanza (not mapped), 5 (MCZ 79291-95);
El Montazo, Municipio C^onstanza (not
mapped), 1 (MCZ 79296); Seccion La
Culata, Paraje La Cienaga, 1 (MCZ 75180);
Municipio Jarabacoa, Seccion Manabao,
Paraje la Cienaga, 3 (MCZ 75181-83); be-
tween Constanza and Jarabacoa (not
mapped), 5 (MCZ 64383-87); Loma
Rucilla, 3 (MCZ 44384-86); Monsenor
Nouel, 1 (MCZ 64370); 1.2 mi. (1.9 km)
SE Monseiior Nouel, 700 feet (214 meters),
1 (ASFS X8125); Piedra Blanca, 6 (MCZ
64377-82); 2 km NW La Cumbre, 2 (MCZ
56850-51). Samand Prov.: 8 km SE
Yayales, 1 (ASFS V1918); 6 km E Sanchez,
2 (ASFS V1908-09); Sanchez, 11 (MCZ
37497-506 + one untagged specimen ) ; 5 km
W Samana, 1 (ASFS V1983); Samana, 2
(MCZ 5448, MCZ 43699); Puerto Escondido,
4 (ASFS V2974-77).
Anolis distichus properus' subsp. n.
Holoti/pe: MCZ 81130, an adult male,
from 0.5 mi. (0.8 km) NW Boca de Yuma,
La Romana Province, Repiiblica Domini-
cana, taken 31 August 1963 by Ronald F.
Klinikowski. Original number V920.
Paratijpcs ( all from La Romana Province,
Repiiblica Dominicana): ASFS V921, same
data as holotype; ASFS X8235, Rio
Cumavasa, 17 km W La Romana, 28 June
1963, D. C. Leber; MCZ 16443-51, La
Romana, 1922, E. Lieder; MCZ 75203, MCZ
7520.5-06, La Romana, 27 March 1963, C.
E. Ray, R. Allen; MCZ 75184, MCZ
75195-97, 5 km E La Romana, 27 March
1963, C. E. Rav, R. Allen; USNM 157917,
8 km E La Romana, 19 Jul\- 1963, R.
Thomas; ASFS X9316, 2 km E La Romana,
19 Jul)' 1963, R. Thomas; ASFS Vl()62-63,
mouth of Rio Chavon, west side, 4 Septem-
ber 1963, R. F. Klinikowski; MCZ 58607.
MCZ 58609-13, Sanate, 12 km S Higiiey,
26 August 1958, C. E. Ray and A. S. Rand;
AMNll 96472-75, 0.3 mi. (0.5 km) NW
Boca de Yuma, 29 August 1963, A. Sch-
wartz, R. Thomas; ASFS VI 135, 2.5 km
NW Boca de Yuma, 4 September 1963,
nati\ e collector; RT 807, 2.5 km NW Boca
Latin. ])r<^)crus, (\n
ick.
zcnbaA ,;(»a
miS rttuo2 ,bn9 moh-^ 212A) won bnossS
i
Plate I
First row: left, A. disfichus disiichus (ASFS 10301), Cawe Point, New Providence Island, Bahama Islands, snout-vent length
46 mm; right, Anolis d. d'ntichoides (ASFS 10280), Fresh Creek, Andros Island, Bahama Islands, snouf-vent length 46 mm.
Second row: left, A. d. biminiensls (ASFS X4932), western end. South Bimini island, Bahama Islands, snout-vent length 47
mm; right, Anolis d. ocior (MCZ 81140), Port Nelson, Rum Cay, Bahama Islands, snout-vent length 53 mm.
Third row: left, A. d. dominicensis (ASFS XI 237), Morne Calvoire, 1 mi. SW Petionville, 2300 feet, Dept. de I'Ouest,
Haiti, snout-vent length 54 mm; right, A. d. ignigularis (ASFS X7735), 9.8 mi. E Santo Domingo, Distrito Nacional, Re-
publica Dominicano, snout-vent length 48 mm.
Fourth row: left, A. d. properus (MCZ 81130], 0.5 mi. NW Boca de Yuma, La Romano Province, Republica Dominicano,
snout-vent length 48 mm; right, A. d. sejunctus (MCZ 81131), environs of Mono Juan, Isia Saona, Republica Dominicano,
snout-vent length 50 mm.
Plate II
First row: left, A. d. tostus (MCZ 81134), western end, Isia Catalina, Republica Donfiinlcana, snouf-vent length 49 mm;
right, A. d. ravifergum (MCZ 81132), 16.5 mi. S San Jose de Ocoa, Peravia Province, Republica Dominicana, snout-vent
length 53 mm.
Second row: left, A. d. favillarum (MCZ 81133), 3 km NE Las Auyamas, 3300 feet, Barahona Province, Republica Do-
minicana, snout-vent length 50 mm; right, A. d. auriler (MCZ 81135), 11 km N Cavailion, 1300 feet, Dept. du Sud,
Haiti, snout-vent length 52 mm.
Third row: left, A. d. v'mosus (MCZ 81136), Camp Perrin, Dept. du Sud, Haiti, snout-vent length 53 mm; right, A. d.
juliae (ASFS X3548), western end, lle-a-Vache, Haiti, snout-vent length 48 mm.
Fourth row: left, A. d. suppar (MCZ 81137), Dame-Marie, south side of tov/n along coast, Dept. du Sud, Haiti, snout-
vent length 52 mm; right, A. d. patruelis (MCZ 81138), vicinity of Pointe Sable, He Grande Cayemite, Haiti, snout-
vent length 49 mm.
n woi«i
;^ q,r.n
»lflifoS lo xliniDiv »(8£fiiS i.l;M| ivbuitcq
.t: ,1^*!' ■ ?*-!*^ "'t»H
ii;i'l "'
Angus distichus • Schwartz
283
de Yuma, 6 August 1963, native collector;
MCZ 75198-202, Boca de Yuma, 28 March
1963, C. E. Ray, R. Allen; MCZ 75207-21,
Juanillo, 29 March 1963, C. E. Rav, R.
Allen; UIMNH 61681-83, 16.5 km SE El
Macao, 31 August 1963, R. F. Klinikowski,
R. Thomas.
Intergrades between A. d. ignigularis
and A. d. properus (but closer to the latter):
Republica Dominicana, La Romana Prov-
ince: 12 km NE La Romana, 2 (ASFS
X9319-20); 0.7 mi. (1.1 km) SE El Macao,
3 (ASFS X7879-81).
Definition: A subspecies of A. distichus
characterized by moderate size (males to
54 mm, females to 45 mm snout- vent
length), dorsum plain ashy to very pale
green (rarely) and without any distinct
dark markings on the head, dewlap very
pale yellow with at times a pale central
orange blush, modally 0 0 scales between
the supraorbital semicircles and the inter-
parietal and 0/0 supraorbitals in contact
with the interparietal, 2/2 scales in contact
laterally with the postfrontals, and very
low mean number ( 2.8 ) of median azygous
head scales.
Distribution: La Romana Province, Re-
publica Dominicana, from the Rio Cumayasa
on the west, east and north around Cabo
Engano to the vicinity of El Macao; inter-
grades with A. d. ignigularis northeast of
La Romana, south of Higiiey, and at El
Macao (Fig. 3).
Comments: The pale and drab A. d.
properus stands in strong contrast to its
1)rightly colored relative ignigularis to the
north and west. The two are readily sepa-
rable on the basis of body color, since
properus is always faded and pale and
usually ashy gray in the field, although it
is capable of a very pale green phase. No
specimens were observed to become solid
dark brown, one of the phases in the
repertory of ignigularis. The pale yellow
dewlap of properus is in harmony with the
balance of its faded coloration (PI. I);
occasional specimens have a pale orange
central blush on the dewlap. In dewlap
color, properus resembles dominicensis
( from whose range it is separated by
ignigularis and an undescribed subspecies),
but it can be differentiated from domini-
censis by the lack of bright green and
dark brown phases. The head is virtually
patternless, and this character will dif-
ferentiate properus from the unnamed sub-
species to the west in the Valle de Neiba
and Llanos de Azua. The hindlimb band-
ing, which is a fairly constant feature of
dominicensis and ignigularis, is much re-
duced or absent in properus.
The holotype has the following measure-
ments and counts: snout-vent length 48
mm, tail ca. 57 mm, tail regenerated; 5
scales across snout, 4 loreal rows, semi-
circles in contact, 0/0 scales between
supraorbital semicircles and interparietal,
1/1 supraorbitals in contact with inter-
parietal, 3/3 scales in lateral contact with
postfrontals, 19 fourth toe lamellae, 3
median azygous head scales, "preoccipital"
present, 7 postmentals.
Scale counts for the series of 58 A. d.
properus are: snout scales 4 to 9 (mode
4 but 6 scales is almost equally modal),
loreal rows 4 to 6 (mode 5); supraorbital
semicircles in contact in all specimens;
modally 0/0 scales between supraorbital
semicircles and interparietal, and 0/0
supraorbitals in contact with interparietal;
2/2 scales in lateral contact with prefrontals;
fourth toe lamellae 15 to 21 (mode 17);
median azygous head scales 1 to 6 (mode
3, mean 2.8); "preoccipital" usually present
(55 of 58 specimens); postmentals 4 to 11
(mode 6, mean 6.7). The largest male (54
mm snout-vent length) is from La Romana,
and the largest female (45 mm) is from
2.5 km N\\' Boca de Yuma.
Intergradient specimens between pro-
perus and ignigularis from south of Higiiey
and Bejucal have already been noted under
the discussion of the latter subspecies. The
Higiiey material was collected on fence
posts in a shady pasture; in life these
lizards were olive to gray with more or less
longitudinal dark striae (which properus
284 Bulletin Museum of Comparaiivc Zoology. Vol. 137, No. 2
lacks). One specimen had a dirty yellow
dewlap, whereas in the remainder of the
series the dewlaps had variable amounts of
orange centrally and pale yellow edges.
These lizards are appropriate both geo-
graphically and in characteristics as inter-
mediates between propcrus and ignigularis,
but as a whole they are closer to the latter.
The Bejucal series, which I did not see
in life, at least shows dorsal pattern features
which are likewise more like those of
ignigularis than propcrus.
The specimen from 0.7 mi. (1.1 km) SE
El Macao has the dorsum tan and striated.
The dewlap has a restricted patch of dull
orange basally and a wide pale yellow
margin. This specimen, by virtue of its
striate dorsum and more prominent orange
dewlap blotch, seems intermediate between
propcrus and ignigularis, although much
closer to the former. Specimens of igni-
gularis were taken about 30 kilometers
northwest of El Macao.
The remaining locality for ignigularis X
properus intergrades is 12 km NE La
Romana. Tlie situation at this locality is
most peculiar, since the two specimens
were collected in a mesic and forested
ravine which presently cuts deeply through
cane fields. The dewlap of the single male
was dull orange centrally with a narrow
dull yellow edge — an ignigularis character.
The dorsal color was dull grayish to dull
greenish; the back was not bicolor and the
hues were much subdued and faded, but
not so pale as those of propcrus. These
specimens combine the characters of pro-
pcrus and ignigularis.
Comparisons of propcrus with both
(lominiccnsis and ignigularis have been
made above. Of the Bahaman su])species,
propcrus most closely resembles (lisficlius;
bom bitninioisis, disticlioiilcs and dapsilis,
propcrus differs in having a yellow rather
than an orange dewlap, as well as in se\(>ral
scale characters, thc> most striking of which
is the extremely low mean for median head
scal(\s (2.8 in contrast to 5.5 to 8.7 in the
al)ove listed Bahaman subspecies). A. d.
ocior in its non-green phase is somewhat
like propcrus. but the longitudinally lined
flanks will distinguish the former from the
latter, as will also the median head scales
(5.8 in ocior). From the nominate race,
which propcrus most closely resembles in
general aspect and dewlap color, propcrus
differs in lower mean median head scales
(2.8 versus 5.9), greater number of post-
mentals (6.7 versus 5.4), larger size (54
mm versus 49 mm ) , and in never having
the supraorbital semicircles separated by a
row of scales. A. d. disiichus and A. d.
propcrus both have 0/0 scales between the
semicircles and the interparietal and 0/0
supraorbitals in contact with the inter-
parietal as the modal conditions.
The range of A. d. propcrus embraces
the arid portion of extreme eastern His-
paniola. Lizards were taken on fences and
in xeric woods and on rocks with which
their color blends exceptionally well. Al-
though I have no data on elevation, there
are no high mountains in this southeastern
region, and thus no specimens come from
elevations of any consequence. The type
locality lies on the forested limestone ridge
which parallels the coast at Boca de Yuma.
At Boca de Chavon, A. d. propcrus was
collected in coastal stands of Coccoloha.
Nowhere does the subspecies appear to be
so abundant as ignigularis and domini-
ccnsis.
Anolis distichus sejuncfus' subsp. n.
Ilolofypc: MCZ 81131, an adult male,
from en\irons of Mano Juan, Isla Saona,
Republica Dominicana, taken 19 JuK- 1964
by Richard Thomas. Original number
V3064.
Parafijpcs: ASFS V3061-63, AMNIT
96476, USNM 157918, sam(> data as holo-
type.
Dcfinilion: A sul^spccies of A. dislicJius
characterized by small size (males to 50
mm snout-vent Icmgtli; only one female
known, with snout-Ncnt Icngtli ol 38 mm).
I'^roni I,aliii, sriunffcrr, to so\(-r.
Anolis distichus • Schwartz
285
dorsum light gray with darker spots and
flecks and suffused with greenish yellow,
head without any distinct dark markings,
dewlap uniform pale yellow, modally 1/1
scales between the supraorbital semicircles
and the interparietal and 0/0 supraorbitals
in contact with the interparietal, 2/2 scales
in contact laterally with the postfrontals,
and moderate mean number (4.3) of median
azygous head scales.
Distribution: Isla Saona, Republica Do-
minicana (Fig. 3).
Comments: The five males and one fe-
male taken on Isla Saona are the only
specimens of A. disticlius known from that
island. The dorsum is light gray and (in
large males) is suffused with greenish
yellow. Thus, sejtinctus seems to have both
a gray and a greenish color phase in its
repertory. In both phases, the dorsum is
marked with a scattering of dark spots and
flecks, although the head lacks any defini-
tive pattern. The dewlap is regularly faint
yellow (PI. I ) . The holotype has the fol-
lowing measurements and scale counts:
snout-vent length 50 mm, tail 37 mm, two
thirds regenerated; 7 scales across snout, 5
loreal rows, semicircles in contact, 0/0
scales between semicircles and interparietal,
1/1 supraorbitals in contact with inter-
parietal, 2/3 scales in lateral contact with
postfrontals, 18 fourth toe lamellae, 3
median azygous head scales, "preoccipital"
present, 5 postmentals. The series of six
A. d. sejunctiis has the following scale
counts: snout scales 5 to 7 (mode 5),
loreal rows 4 to 6 (mode 5); supraorbital
semicircles in contact in all specimens;
modally 1/1 scales between supraorbital
semicircles and interparietal and 0/0 supra-
orbitals in contact with interparietal; 2/2
and 3/3 (each with two specimens) scales
in lateral contact with postfrontals; fourth
toe lamellae 15 to 18 (mode 18); median
azygous head scales 3 to 6 (mode 3 or 5,
both with two lizards, mean 4.3); "preoc-
cipital" always present; postmentals 5 to 7
(mode 5, mean 5.7).
A. d. sejunctiis resembles A. d. propenis
on the adjacent mainland. Two features
distinguish them: the absence in properns
of any dorsal markings, with a consequently
plain back, and the greenish tints which
sejunctiis is apparently able to assume
regularly. A. d. propenis only very rarely
has any green hues in its repertory. The
two races resemble each other in lacking
any dark head pattern, and in having a
uniform pale yellow dewlap (although
propenis may have a pale basal orange
blush). Scalewise, propenis modally has
0/0 scales between the semicircles and the
interparietal, whereas sejunctiis has a for-
mula of 1/1. In mean of median head
scales, propenis (2.8) is lower than
sejunctiis (4.3).
From the balance of the subspecies,
sejunctiis differs from distichoides, bimini-
ensis, dapsilis, and ignigiilaris in having a
yellow rather than an orange dewlap. From
the yellow-dewlapped forms distichus and
ocior, sejunctiis differs in having 1/1 scales
between the semicircles and the inter-
parietal (0/0 in the two Bahaman sub-
species) and 0/0 supraorbitals in contact
with the interparietal (0/0 in distichus,
1/1 in ocior). A. d. distichus is incapable
of a green phase, and the lineate sides and
unpatterned back of ocior contrast with the
patterned back and plain sides of sejunctiis.
The usual differences in presence of the
"preoccipital" and higher frequency of
complete separation of semicircles in Baha-
man versus Hispaniolan races apply as well.
Compared with the yellow dewlapped A.
d. dominicensis to the west, sejunctiis dif-
fers in smaller size, flecked and spotted in
contrast to striate dorsum, and lower mean
number of postmentals (5.7 versus 6.6).
The aspect of these two subspecies is quite
different.
The area about Mano Juan is generally
shady woody scrub, and the lizards were
taken in this habitat as well as in the
settlement of Mano Juan.
The fauna of Isla Saona is becoming in-
286 Bulletin Mtiseum of Comparative Zoology, Vol. 137, No. 2
creasingly well known, and all species
which occur there which have been studied
(Leiocephahis hinatii.s, Ameiva chrysolaema,
Ameiva tacniiua, D) amicus parvifrons) are
represented by distinctive subspecies, which
show expected affinities with their rela-
tives on the adjacent mainland or in ex-
treme eastern Hispaniola. A. disticlius
follows this pattern. Presumably A. d.
■scjunctus is widespread on Saona, despite
the fact that it is known only from the
area about Mano Juan.
Anoiis distichus fosfus' subsp. n.
Holotypc: MCZ 81134, an adult male,
from Isla Catalina, western end, Republica
Dominicana, taken 20 August 1963 by
Richard Thomas. Original number V558.
Paiatypes: ASFS V559-60, same data
as holotype.
Definition: A subspecies of A. disticlius
characterized by (presumably) small size
(males to 46 mm snout-vent length; fe-
males imknown ) , dorsum yellow-tan with
little or no flecking or striations and no
head pattern, dewlap deep orange centrally
with a yellow border, modally 0/0 scales
between the supraorbital semicircles and
the interparietal, and moderate mean num-
ber (5.0) of median azygous head scales.
Distribution: Isla Catalina, Republica
Dominicana ( Fig. 3 ) .
Comments: The three male specimens of
A. d. to.-itus are so distinctive that I have no
hesitancy in describing them as a subspecies
which differs both from propcrus on the
adjacent coast and sejunctus on Isla Saona
to the east. The yellow-tan dorsum (pi. 12
J 3) is like that of no other subspecies of
A. distichus; the patternless head resembles
that of both sejunctus and properus, but the
extensively orange-centered dewlap (Pi.
II) is more like that of i'^niiiularis (which
usually has th(> orange center largcT and
the yellow edge much narrower) than the
pale yellow dewlaps of both propcrus and
sejunctus. There is no evidence (but the
~ Kroin Latin, Inrrere, to parch.
number of specimens both collected and
observed was few) that to.stus has a green
phase or has green pigment in its repertory.
The holotype has the following measure-
ments and counts: snout- vent length 46
mm, tail broken; 5 scales across snout, 5
loreal rows, semicircles in contact, 0/0
scales between semicircles and interparietal,
2 2 supraorbitals in contact with inter-
parietal, 2/2 scales in lateral contact with
postfrontals, 16 fourth toe lamellae, 5
median azygous head scales, "preoccipital"
present but tiny, 4 postmentals. The series
of three A. d. tostus has the following scale
counts: snout scales 5 and 6 (mode 6),
loreal rows 4 and 5 (mode 4); supraorbital
semicircles in contact in all specimens;
modally 0 0 scales between supraorbital
semicircles and interparietal; no mode for
number of supraorbitals in contact with
interparietal — counts of 1/1, 2/0, 2/2; no
mode for number of scales in lateral con-
tact with postfrontals — counts of 2/2, 2/3,
3/3; fourth toe lamellae 16 to 19 (mode
16 ) ; median azygous head scales 4 to 6
(no mode; mean 5.0); "preoccipital" always
present; postmentals 3 and 4 (mode 4,
mean 3.7). The mean of median head scales
is the highest for any Ilispaniolan popula-
tion; the small sample of tostus renders the
significance of this high figure dubious.
From the subspecies disticlius, ocior,
dominicensis, propcrus, and sejunctus, A. d.
tostus differs in having a yellow-tan dorsum
and a dewlap with a d(>ep orange center
and a broad yellow edge. It resembles the
races ])imi)iiensis, distichoidcs, dapsilis, and
iiiniiiidaris in having an orange dewlap, but
differs from thest> races in dorsal color and
pattern. Scale counts are not profitably
compared.
On Isla Catalina, A. d. lost us was col-
lected exclusively in dry hammock woods
( = low^ coppice), and I'ven there was un-
common. Since Isla ('atalina is \er\' dry
and much of it is sun-baked scrub and
grassland, presumably foslus is restricted
to the shadier situations in xcM'ie woods.
Anolis distichus • Schwartz
287
Anolis distichus ravitergum^ subsp. n.
Holotype: MCZ 81132, an adult male,
from 16.5 mi. (26.4 km) S San Jose de
Ocoa, 500 feet (122 meters), Peravia
Province, Repiiblica Dominicana, one of a
series taken 24 August 1963 by Ronald F.
Klinikowski, Albert Schwartz, and Richard
Thomas. Original number V728.
Paratypes (all from the Republica Do-
minicana): ASFS V729-35, same data as
holotype; ASFS X7988, 1.8 mi. (2.9 km) W,
thence 1.1 mi. (1.8 km) N Azua, Azua
Province, 24 June 1963, R. Thomas; AMNH
96477-80, CM 40604-08, 1.8 mi. (2.9 km)
W, thence 2.7 mi. (4.3 km) N Azua, Azua
Province, 24 June 1963, R. F. Klinikowski,
D. C. Leber; MCZ 58422-23, 12 km N
Azua, Azua Province, 11 August 1958, C. E.
Ray, A. S. Rand; ASFS V3169-77, 2 km
W Puerto Viejo, Azua Province, 27 July
1964, D. C. Leber, R. Thomas; UIMNH
61684-85, 15.2 mi. (24.3 km) S San Jose
de Ocoa, Peravia Province, 24 August 1963,
A. Schwartz; UF FSM 21514-15, 1.8 mi.
(2.9 km) S San Jose de Ocoa, 1300 feet
( 397 meters ) , Peravia Province, 24 August
1963, R. F. Klinikowski, R. Thomas; USNM
157919-25, 10 km W Bani, Peravia Province,
27 July 1964, D. C. Leber, R. Thomas; MCZ
58421, 13 km NW Bani, Peravia Province,
6 August 1958, C. E. Ray, A. S. Rand; KU
93359-64, 4.2 mi. (6.7 km) NE Sabana
Grande de Palenque, San Cristobal Prov-
ince, 27 June 1963, A. Schwartz.
Referred specimens: REPUBLICA DO-
MINICANA. Independencia Prov.: 6.3 mi.
(10.1 km) SW Neiba, 3 (ASFS V269-71).
Baoruco Prov.: 3.9 mi. (6.2 km) ENE
Neiba, 4 (ASFS V221-24); 3.4 mi. (5.4
km) ENE Neiba, 1 (ASFS V246); 0.8 mi.
(1.3 km) SW Neiba, 4 (ASFS V248-50,
RT774).
Intergrades between A. d. ravitergum
and A. d. dominicensis: REPUBLICA DO-
MINICANA. Azua Prov.: Padre las Casas,
3 (MCZ 58477-79).
Definition: A subspecies of A. distichus
** From Latin, nivuni, gray, and tergum, back.
characterized by large size (males to 56
mm, females to 45 mm snout-vent length),
dorsum ashy gray to tan or pale greenish,
head usually with a distinct interocular
dark brown bar and a dark U extending
from the eyes across the occiput, dewlap
pale yellow, at times with a faintly orange
center, modally 0/0 scales between the
supraorbital semicircles and the inter-
parietal and 1/1 supraorbitals in contact
with the interparietal, 2 2 scales in contact
laterally with the postfrontals, and a very
low mean number ( 2.6 ) of median azygous
head scales.
Distribution: The Valle de Neiba and
the Llanos de Azua, from east of Lago
Enriquillo east to the vicinity of Sabana
Grande de Palenque in San Cristobal
Province, Republica Dominicana (Fig. 3).
Comments: A. d. ravitergum is typically
an ashy gray to drab tan lizard \\'ith a
fairly prominent brown head pattern. Some
individuals show a greenish phase, but the
green is neither bright nor vivid. The
dewlap is pale yellow and occasionally has
a pale orange center (PI. II). Specimens
with orange-centered dewlaps are com-
moner in the Valle de Neiba and may be
demonstrating in this area the residual
genetic influence of the subspecies in the
uplands of the adjacent Sierra de Baoruco.
The back is usually moderately marked
with vague longitudinal striae, but some
specimens are plain above. A few lizards
(as preserved) lack the head markings
described for the subspecies, but in general
the markings are a consistent feature of the
entire series. The venters are whitish and
the midersides of the tails vary from pale
yellow to orange or yellowish green. In
general, the entire coloration is faded and
subdued.
The holotype has the following measure-
ments and counts: snout-vent length 53
mm, tail 56 mm, incomplete; 4 scales across
snout, 5 loreal rows, semicircles in contact,
0 0 scales between supraorbital semicircles
and inteiparietal, 1/2 supraorbitals in con-
tact with interparietal, 2/2 scales in lateral
288
Bulletin Mtiseuyn of Comparaiive Zoology, Vol. 137, No. 2
contact with postfrontals, 19 fourth toe
lamellae, 2 median head scales, "preoc-
ciptal" present but divided longitudinally,
6 postmentaLs.
Scale counts for the series of 57 raviter-
gum are: snout scales 4 to 8 (mode 6),
loreal rows 3 to 6 (mode 4); supraorbital
semicircles in contact in all specimens;
modally 0/0 scales between supraorbital
semicircles and interparietal and 11 supra-
orbitals in contact with interparietal; 2/2
scales in lateral contact with postfrontals;
fourth toe lamellae 14 to 23 (mode 17);
median azygous head scales 0 to 5 (mode
3, mean 2.6); "preoccipital" usually present
(49 of 52 specimens); postmentals 4 to 9
(mode 5, mean 5.6). The largest males
(56 mm) are from 3.9 mi. (6.2 km) ENE
Neiba, Baoruco Province, and the largest
female (45 mm) is from 12 km E Azua,
Azua Province.
In having a yellow dewlap, ravifcrgiim
differs from the orange-dewlapped sub-
species Jyiminicnsis, distichoides, dap.silis,
ignigidari'i and to.stii.s. A. d. ravitergum in
its drab coloration is most like properus
and sejunctiis. The presence of a head
pattern and of at least vague striae on the
dorsum will distinguish ravitcrg,um from
these races. From dominiccn.sis, ravitcrfs.um
differs in dorsal coloration ( lacking either
a bright green or a dark brown phase),
in having 0/0 scales between the semicircles
and the interparietals and 1/1 supraorbitals
in contact with the interparietal (1/1 and
0/0, respectively, in dominiccnsis), and
lower mean number of median head scales
(2.6 versus 3.9). A. d. nwitcrgum resembles
A. d. ocior in general dorsal color, but the
latter race has a prominent lateral pale
streak, a brighter green phase, and much
higher mean of median head scales (2.6
versus 5.(S). A. d. lavUcrgiun is the second
largest subspecies, being exceeded in snout-
vent length only by A. d. dom'micemis. It
is most closely approached by ig,nig,idaiis
and properus in si/e.
Presumably A. d. ramterfi^iim comes into
contact with three other subspecies of A.
distichus. To the west in the Valle de
Neiba it must meet dominicensis some-
where between Neiba, on the one hand,
and Aguacate and the mountains above
La Descubierta, on the other. No specimens
are available from this hiatus, and it is
interesting that the lizards from both the
latter localities are from the ascending
slopes of the Sierra de Neiba (2000 feet —
610 meters) and the Sierra de Baoruco
(1600 feet — 488 meters). Since ravitcriium
is in essence an inhabitant of the floor of
the Valle de Neiba in the western portion
of its range, the zone of intergradation may
well be narrow and restricted to the lower
slopes of the ranges. The nearest localities
in the valley floor whence I have seen
A. d. dominicensis to the west in Haiti are
Manneville and Thomazeau; comment on
the orange-dewlapped populations of do-
minicensis in the Thomazeau-Manneville
region has already been made. It is perti-
nent to note again that the highest fre-
quency of orange dewlap centers in raviter-
gum is in the Valle de Neiba east of the
Thomazeau-Manneville area.
A short series of three specimens is from
Padre las Casas, Azua Province, on the
southern dry slopes of the Cordillera Cen-
tral. These lizards, even though preserved
for some time, still are noticeably green,
especially about the head; the larger female
has a snout-vent length of 47 mm, which
is near tlie upper extreme of female
dominicensis but l)elo\\ that of nwiicrii^um.
The general area about Padre las Casas is
transitional between the lower arid Llanos
de Azua and the more mesic interior up-
lands, but its aspect and launa (i.e., Ameiva
lineohita) are closer to those of tlie hot
lowlands. I would expect on ecological
grounds that the /\. distichus at Padre las
C^asas would \)v r(nilcriS.ii})i; geographically,
however, it is an almost ideal situation for
intergradation between a lowland and (in
tills area) highland subspecies.
A. d. raviteris.ii7n comes into contact in
the east with A. d. iiS.niii,id(iris, in south-
western San Ciisl(')bal Pioxince. Here, the
Anolis distichus • Schwartz
289
line of demarcation between the two sub-
species is extremely sharp, since ravitergnm
is known from 4.2 mi. ( 6.7 km ) NE Sabana
Grande de Palenque and i<i,niii,tihiris from
2 mi. (3.2 km) SE San Cristobal and 15.5
km SE El Cacao at 1400 feet, as well as
from a series of specimens from various
measured localities along the road from
San Cristobal to El Cacao. The ignigiilaris
localities are distinctly upland and mesic,
although the locality southeast of San Cris-
tobal is in the mesic lowlands. The distance
between the San Cristobal and Sabana
Grande localities is about 16 kilometers
airline, but the situation near San Cristobal
(a shaded fence row adjacent to pasture
in a generally mesic region) is in contrast
to the drier coastal region near Sabana
Grande. In general, this area in the vicin-
ity of Bani is becoming increasingly well
known as either a place where there is
fairly rapid shift in subspecies or as the
extreme limit of distribution of species,
since on the west are the xeric Llanos de
Azua and on the east the more mesic re-
gions which extend toward Santo Domingo.
The specimens which I have examined
from this general region are referable to
either ravitergnm or ignigularis, and I do
not regard any of them as intergradient.
There is presumably also a zone of con-
tact between ravitergnm and the unde-
scribed subspecies in the Sierra de Bao-
ruco, but there are no specimens from
lower intermediate elevations, and all ma-
terial at hand from the eastern Baoruco
is clearly the race indigenous to that mas-
sif and shows no tendencies toward ravi-
tergum (see however the discussion below
concerning the material from southwest of
Barahona in the Sierra de Baoruco).
Although A. d. ravitergnm is essentially
a lowland subspecies in the Valle de Neiba
and the Llanos de Azua, it does ascend
the southern rolling piedmont of the Cor-
dillera Central in the vicinity of San Jose
de Ocoa and also occurs in the Sierra de
Ocoa. But in both these regions, condi-
tions are xeric and merely continuations
of the same habitat in the lower plains.
The highest elevation for A. d. ravitergnm
is 1300 feet (397 meters); presumably it
also occurs below sea level at the eastern
end of Lago Enriquillo.
The relationships of A. d. ravitergnm
and A. hrevirostris in the Valle de Neiba
will be discussed later in detail by Dr.
Williams. It is pertinent at this time to
point out that in this low and arid valley,
A. d. ravitergnm is more or less confined
to shady palm oases and other less rigor-
ous situations, whereas A. hrevirostris is
the lizard of the open scrub. On the as-
cending slopes of the Sierra de Baoruco,
A. distichns and A. hrevirostris are pre-
cisely syntopic; in this area of syntopy, the
vegetational cover is intermediate between
that of the rain forest above and the arid
plains below.
Anolis distichus favillarum' subsp. n.
HoJoti/pc: MCZ 81133, an adult male,
from 3 km N Las Auyamas, 3300 feet ( 1007
meters) Barahona Province, Republica Do-
minicana, taken 24 July 1963 by David C.
Leber. Original number X9593.
Paratypes (all from Barahona Province,
Republica Dominicana): ASFS X9592,
same data as holotype; ASFS X983S-41,
7.0 mi. (11.2 km) S Cabral, 2300 feet (702
meters), 27 July 1963, R. Thomas; CM
40609-12, 7.1 mi. (11.4 km) S Cabral, 2300
feet (702 meters), 27 July 1963, D. C.
Leber, R. Thomas; ASFS X9832-33, 8.8
mi. (14.1 km) S Cabral, 2700 feet (824
meters), 27 July 1963, D. C. Leber, R.
Thomas; MCZ 58424, MCZ 58426-28, MCZ
58430-31, MCZ 5843.3-35, MCZ 58437, La
Cueva, 11 km SW Cabral, 17 August 1963,
C. E. Ray, A. S. Rand; UF/FSM 21516, 8
km NE Las Auyamas, 2600 feet (793 me-
ters), 28 July 1963, native collector; UF/
FSM 21517, 24 km SW Barahona, 3700 feet
(1129 meters), 2 August 1963, D. C. Leber;
AMNH 96481-83, 24 km SW Barahona,
3700 feet ( 1129 meters), 6 July 1964, D. C.
^ From Latin, faiilla, glowing ashes.
290 Bulletin Museum of Comparative Zoology, Vol. 137, No. 2
Leber, R. Thomas; MCZ 65353, Hermann's
finca, near Paraiso, 2400 feet (732 meters),
26 August 1932, W. G. Hassler.
Definition: A sul^jspecies of A. distichtis
charaeterized by moderate size ( males to 54
mm, females to 47 mm snout-vent length ) ,
dorsum bright dark green and heavily stri-
ate with darker green or brownish, head
with rusty temples and interparietal scale
yellow-green and sharply set off from re-
mainder of green head coloration, dewlap
vivid orange centrally with a narrow pale
yellow edge, modally 1/1 scales between
the supraorbital semicircles and the inter-
parietal and O/'O supraorbitals in contact
with the interparietal, 2 2 scales in contact
laterally with the postfrontals, and a mod-
erate mean number (3.8) of median azy-
gous head scales.
Distribution: Known only from inter-
mediate and higher elevations in the east-
ern portion of the Sierra de Baoruco in the
Republica Dominicana (Fig. 3).
Comments: Perhaps the most gaudy of
the Ilispaniolan mainland races of A clis-
tichiis is favillarum. This is especially true
when it is compared with its neighbors
(lominiccnsis in the west and raviterii,tim
in the north. The latter is essentially a
drab brownish lizard with a pale yellow
dewlap and the former a green lizard with
a pale yellow dewlap. Neither has the rusty
temples and sliarply distinct yellow-green
interparietal scale nor the vivid orange fa-
vAllariim dewlap (Pi. II).
The holotype of A. d. favillarum has the
following measurements and counts: snout-
vent length 50 mm, tail broken; 4 scales
across snout, 4 loreal rows, semicircles in
contact, 0/0 scales between supraorbital
s(Miiicircles and interpari(>tal, 2/2 supra-
orbitals in contact with interparietal, scales
in lateral contact with postfrontals indeter-
minate, 21 fourth toe lamellae, 1 median
head scale, "preoccipital" absent, 5 post-
mentals.
The series of 28 A. d. favillarum has the
following counts: snout scales 3 to 6 (mt)de
4), loreal rows 4 to 6 (mode 4); su]')raor-
bital semicircles in contact in all specimens;
modally 1/1 scales between supraorbital
semicircles and interparietal and 0 0 supra-
orbitals in contact with interparietal; 2/2
scales in lateral contact with postfrontals;
fourth toe lamellae 16 to 22 (mode 19);
median azygous head scales 1 to 6 (mode
4, mean 3.8); "preoccipital usually present
( 26 of 28 specimens ) ; postmentals 4 to 8
(mode 7, mean 6.1).
In having an orange dewlap, favillarum
differs from the subspecies distichus, ocior,
dominicensis, properus, sejunctus, and
raviterg,um. Although in dewlap color A.
d. favillarum resembles himiniensis, disti-
ehoides, dapsilis, ignigularis, and tostus,
none of these races is deep bright green
above with rusty temples and a distinct yel-
low-green parietal. Despite the dewlap simi-
larities, for instance, it is hard to visualize
two subspecies more distinct in general ap-
pearance than favillarum and tostus, or fa-
villarum and biminensis. In having 1/1
scales between the semicircles and the inter-
parietal and 0/0 supraorbitals in contact
with the interparietal, favillarum resembles
distichoides, dominicensis, and sejunctus.
The moderate mean of median head scales
(3.8) in favillarum is lower than those of
the other orange-dewlapped races (5.0 to
8.7) with the exception of i<j,nigidaris (3.5).
In some ways favillarum most closely re-
sembles ig,niiiularis, but these two sub-
species can be differentiated in that favil-
larum lacks the bicolor dorsum of i'j^niiiu-
laris, and i'j,niii,ularus lacks the rusty tem-
ples of favillarum. The ranges of the two
are separated by some 62 kilometers at their
nearest points (and the distance is longer
if one considers the intervening coastal em-
bayments and irregularities) as well as by
the intervening lowland subspecies raviter-
il.um in the Llanos de Azua. A. d. favil-
larum is so distinctive in color and pattern
that it really requires little detailed com-
parison with an\ other subspecies.
A. d. favillarum pr('suiiial)ly comes into
contact with raviterii,um to the north at the
base of th(> SicMia de Baoruco and with
Angus distichus • Schwartz
291
dominicensis to the west in the western por-
tion of the Sierra de Baoruco. The lack of
faviJIannn X lovitcr^tim intergrades has
])een explained in the discussion of the lat-
ter subspecies. The absence of faviUarum
X dominicenses intergrades is doubtless due
to the fact that there is no material avail-
able from the central (and virtually inac-
cessible) portion of the Sierra de Baoruco.
A. distichus from the Sierra de Baoruco
along the Dominico-Haitian border are
dominicensis. One of the paratypes of A.
d. fuviUarum is of possible interest insofar
as the problem of intergradation between
this subspecies and ravitergum is con-
cerned. This adult lizard, from 24 km SW
Barahona at an elevation of 3700 feet (1129
meters), was noted as having a plain yel-
low dewlap. The specimen might be inter-
preted as showing tendencies toward the
r(iviter<ium dewlap condition; on the other
hand, this seems unlikely, especially in view
of the extreme elevation of the locality. I
consider it more likely that it is simply a
faviUarum with an aberrantly colored dew-
lap.
A. d. faviUarum is esentially a denizen
of mesic woods and cafctah's at higher ele-
vations in the Sierra de Baoruco; the known
altitudinal limits for the subspecies are
from 2300 feet (702 meters) to 3700 feet
(1129 meters), although the subspecies
must occur at both higher and somewhat
lower elevations in this mountain range.
In the area of syntopy with A. hrcvirostris
(the lower altitudinal limits noted above),
the vegetational cover is transitional be-
tween that of the very mesic uplands and
that of the Valle de Neiba below.
Anolis distichus ourifer^'* subsp. n.
Holotype: MCZ 81135, an adult male,
from 11 km N Cavaillon, 1300 feet (397
meters), Dept. du Sud, Haiti, one of a
series taken 6 August 1962 by Dennis R.
Paulson, David C. Leber, and native col-
lectors. Original number X3717.
^" From Latin, aiirifer, gold bearing.
Parati/pes ( all from Dept. du Sud, Haiti ) :
ASFS X3658-63, ASFS X3680-84, ASFS
X3718-23, AMNH 96484-87, KU 93365-68,
CM 40613-16, UIMNH 61686-89, same data
as holotype; MCZ 74838-64, Pourcine,
Massif de la Hotte, 31 December 1962—
2 January 1963, F. Vuilleumier; MCZ
74833-37, Trou Bois on Jeremie Road, 30
December 1962, D. Hill.
Referred specimens: HAITI. Dept. du
Sud: Tosia, 1 (MCZ 69756); nr. Massif de
la Hotte (=Pic Macaya), 3 (MCZ 38254-
56); Petit Trou de Nippes, 8 (USNM
80801-08).
Definition: A subspecies of A. distichus
characterized by moderate size (males to
54 mm, females to 46 mm snout-vent
length), dorsum heavily marbled with
varying shades of greens and browns, dew-
lap vivid orange with a narrow yellow
border, modally 1/T scales between the
supraorbital semicircles and the inter-
parietal and 0/0 supraorbitals in contact
with the interparietal, 2/2 scales in contact
laterally with the postfrontals, and a moder-
ate mean number ( 3.7 ) of median azygous
head scales.
Distribution: Known definitely from
only three localities (the type locality,
Pourcine, and Trou Bois ) on the north and
south flanks of the Massif de la Hotte on
the Tiburon Peninsula in southwestern
Haiti; by inference and observation (see
below) assumed to occur from southeast
of Jeremie east to the vicinity of Saint
Michel du Sud, where aurifer intergrades
with dominicensis (Fig. 3).
Comments: The Tiburon Peninsula of
Haiti, west of about the longitude of
Miragoane on the north coast and a pres-
ently unknown locality on the south coast,
is inhabited by a complex of (at least)
three subspecies of A. distichus. In addi-
tion to these three mainland races, there
are additional subspecies on Ile-a-Vache
off the south coast and He Grande Cayemite
off the north coast. The three mainland
subspecies are very different in dewlap
color in life, but the dewlap colors and
292 Bulletin Museum of Comparative Zoology, Vol. 137, No. 2
pattern are of course fugitive in preserved there are no fresh specimens from the
material. Consequently, the precise bound- northern coast of the Tiburon Peninsula in
aries of the various races can be defined this region.
only in terms of freshly collected specimens. The measurements and counts of the
and many older specimens from this region holotype of A. d. aurifcr are: snout-vent
may be placed with a particular subspecies length 52 mm, tail ca. 70 mm; 6 scales
only if there are adequate field data on across snout, 5 loreal rows, semicircles in
color in life — which in some critical ma- contact, 11 scales between supraorbital
terial there are not. Questionable sub- semicircles and interparietal, 0/0 supra-
specific assignments will be noted in ap- orbitals in contact with interparietal, 2/2
propriate discussions. scales in lateral contact with postfrontals,
The series of A. d. aurifcr from the type 20 fourth toe lamellae, 4 median head scales,
locality was examined by me in life. These "preoccipital" present, 6 postmentals.
lizards were heavily mottled and streaked The series of 67 A. d. aurifcr has the fol-
above with varying shades of greens and lowing counts: snout scales 4 to 7 (mode
browns, but lacked any bright colors (i.e., 4), loreal rows 3 to 6 (mode 4); supra-
rusty temples) on the head or body. The orbital semicircles in contact in all speci-
dewlap was vivid orange with a narrow mens; modally 1/1 scales between supra-
yellow margin (PI. II); some males had the orbital semicircles and inteiparietal and
dewlap orange-red, a still more distinctive 0/0 supraorbitals in contact with inter-
and vivid color. The series from Pourcine parietal; 2/2 scales in lateral contact with
in the Museum of Comparative Zoology postfrontals; fourth toe lamellae 16 to 23
was noted by the collector to have the (mode 19, but 20 has almost the same
dewlaps orange "with yellow spots in the frequency); median azygous head scales
orange" — this latter a feature not seen 1 to 7 (mode 3, mean 3.7); "preoccipital"
in the topotypical series. The Trou Bois always present; postmentals 4 to 10 (mode
lizards were likewise noted to have "bril- 7, mean 7.2).
liant orange-red" throats. Richard Thomas The orange dewlap of A. d. aurifcr dif-
collected a single male A. disticJui.s- about ferentiates the subspecies from the yellow-
7.5 km ( airline ) south-southeast of Roseaux dewlapped races disticlius, ocior, domini-
which also had an orange dewlap, but the cen.^is, propcrus, scjunctus, and ravitcrg.uin.
lizard escaped. Tliese localities summarize In addition to dewlap and dorsal color
the known distribution of orange-dewlap- and pattern (none of the above subspecies
ped A. disticlius in this region. I have in- has a lieavily mottled green-and-brown
eluded the single lizard from Tosia, three back), aurifcr differs in the high mean
from Pic Macaya, and eight from Petit number of postmentals (7.2 in aurifcr, 5.4
Trou d(! Nippes with aurifcr on the basis to 6.7 in the above races, with ocior ap-
of provenance. Tosia is on the Les Cayes- proaching aurifcr most closely). The other
Jeremie road on the north side of the Massif orange-dewlapped subspecies are ])imini-
de la Ilotte, and the lizard might lie as- oisis, distichoidcs, dapsilis, iiiniiiularis,
signed to th(> Les Cayes-Camp Perriii sul)- tostus, and faviltaruni, of wliicli aurifcr is
species d(\scribed below. However, there closest geographicalK to faiillaruni, but
are no known specimens of the more south- from which it is separated by the interven-
ern race from the north slope of the La ing range of dominiccnsis. vMl these sub-
Hotte, and it seems likc>ly that the speci- species differ in dorsal pattern and color
mcu is an aurifcr. Tlie same comments fiom aurifcr (in hut, oiiK faiillaruni has a
apply equally well to the Pie Macaya green phase); the rusl\ lemi:)l(\s of favil-
lizards. The series from Petit 'I'rou de larun\ additionally distinguish it from
Nippes falls into the same category, since aurifcr. The m(>an postmentals ol aurifcr
Anolis DiSTiCHUS • Schtvcirtz
293
(7.2) aid in separating it from the other
orange-dewlapped subspecies (3.7 to 6.1,
with faviUamm approaching aurifer most
closely ) .
A. d. aurifer is presumed to intergrade
with A. d. dominiccnsis in the vicinity
of Saint Michel du Sud, southwest of
Miragoane. A series of 18 specimens (ASFS
X3830-47) from 3.5 mi. SW Saint Michel
du Sud, 1000 feet (305 meters), was noted
as having the de\\'laps pale orange with a
yellow edge — precisely the condition ex-
pected at the place of intergradation of an
orange-dewlapped and a yellow-dewlaj^ped
race. Purely on the basis of provenance, I
consider two other lots of specimens from
this same region (MCZ 66113-32, Fond des
Negres, and MCZ 25504-08, 10 mi. [16.0
km] SW Miragoane) aurifer X dominicensis.
The latter lot may be assignable to A. d.
dominicensis, but the Fond des Negres
series is close to Saint Michel du Sud, the
known locality for aurifer X dominicensis
intergradation. Other than these inter-
grades, the eastern limits of aurifer are un-
known; specimens from the north coast in
the Miragoane region were clearly domini-
censis in life.
In the northwest aurifer intergrades with
the yellow-dewlapped population on the
tip of the Tiburon Peninsula in the area
about Roseaux, and in the south aurifer
intergrades with another subspecies in the
vicinity of Cavaillon. In both cases, these
intergrades will be discussed with their
respective subspecies.
The distribution herein attributed to A.
d. aurifer is indeed most peculiar, since it
is assumed to occur on both sides of at
least the eastern portion of the Massif de
la Hotte, and along a portion of the north
coast as well. Much of the upland range
assigned to aurifer is extremely difficult to
penetrate, and it may be some time before
the details of the distribution of aurifer are
clarified. On the basis of the few annotated
series presently available, there is no choice
but to regard all these specimens as one
subspecies.
The type locality of A. d. aurifer is a
rocky shaded hillside on the southern slopes
of the Massif de la Hotte.
Anolis distichus v/nosus'^ subsp. n.
Holotype: MCZ 81136, an adult male,
from Camp Perrin, Dept. du Sud, Haiti,
one of a series taken 22 July 1962 by
native collectors. Original number X2711.
Parafiipes (all from Dept. du Sud, Haiti):
ASFS X2533-49, ASFS X2560-70, AMNH
96488-97, UIMNH 61690-95, CM 40617-22,
UF/FSM 21518-23, same data as holotype;
MCZ 63125-31, Camp Perrin, 5 August
1960, A. S. Rand and J. D. Lazell, Jr.; ASFS
X3361-62, Les Cayes, 2 August 1962, D. R.
Paulson; MCZ 63111-17, Les Cayes, 3 Au-
gust 1960, A. S. Rand and J. D. Lazell, Jr.;
ASFS X3353-55, 9.9 km ENE Port-Salut,
650 feet (198 meters), 3 August 1962, D.
C. Leber, D. R. Paulson; ANSP 27156-62,
KU 93369-75, USNM 157926-27, Carrefour
Canon, 500 feet (153 meters), 1 August
1962, R. F. Klinikowski, A. Schwartz; MCZ
63118-21, Carrefour Canon, 4-5 August
1960, A. S. Rand and J. D. Lazell, Jr.; MCZ
63122-24, Les Platons, above Carrefour
Canon, 5 August 1960, A. S. Rand and J.
D. Lazell, Jr.
Referred specimens: HAITI. Dept. du
Sud: Tombeau Cheval, 3 (MCZ 63132-34).
Definition: A subspecies of A. distichus
characterized by moderate size (males to
54 mm, females to 45 mm snout- vent
length), dorsum marbled with greens and
browns of varying shades, dewlap with
a rather restricted basal maroon (wine
colored) blotch or spot and a broad pale
yellow margin, modally 1/1 scales between
the supraorbital semicircles and the inter-
parietal and 0/0 supraorbitals in contact
^^'ith interparietal, 2/2 scales in contact
laterally with the postfrontals, and a low
mean number (3.4) of median azygous
head scales.
Distrd)ution: The southern slopes of the
Massif de la Hotte from Camp Perrin (and
^^ From Latin, viiio.siis, full of wine.
294 Bulletin Museum of Comparative Zoology, Vol. 137, No. 2
Tombeau Cheval?) and Les Platons, south
to Les Cayes, and west onto the Presqii ile
du Port-Sakit; intergrades with A. d. aurifcr
at Cavaillon (Fig. 3).
Comments: The holotype of A. d. vinosus-
has the following measurements and eounts:
snout-vent length 53 mm, tail 35 mm, bro-
ken; 5 scales across snout, 5 loreal rows,
semicircles in contact, 1/1 scales between
supraorbital semicircles and interparietal,
0/0 supraorbitals in contact with inter-
parietal, 2/2 scales in lateral contact with
postfrontals, 20 fourth toe lamellae, 4
median head scales, "preoccipital" present,
6 postmentals.
The series of 102 A. d. vinosus has the
following counts: snout scales 4 to 8
(mode 4), loreal rows 4 to 6 (mode 5);
supraocular semicircles in contact in all
specimens; modally 1/1 scales between
supraorbital semicircles and interparietal
and 0/0 supraorbitals in contact with inter-
parietal; 2/2 scales in lateral contact with
postfrontals; fourth toe lamellae 16 to 24
(mode 20); median azygous head scales
2 to 8 ( mode 3, mean 3.4 ) ; "preoccipital"
always present; postmentals 4 to 11 (mode
7, mean 7.4 ) .
Compared with all other subspecies of
A. distichiis, from both the Hispaniolan
mainland and the Bahamas, none is so
easily differentiable as vinosns. The com-
bination of maroon or wine-red centered
dewlap with a broad yellow margin (Pi.
II) and heavily mottled brown and green
dorsum will distinguish it from any other
subspecies. Only A. d. jidiac on Ile-a-Vache
resembles A. d. vinosus in dewlap color
and pattcMii; jidiuc will be discussed lurther
below. The amount o\ maroon in the basal
spot ol the vinosus dewlap is \ariable, and
the illustrated indixidual (wliicli is the
holotype) resembles the maximal condi-
tion. The range of vinosus is bordered on
the east by the orange-dewlapped aurifcr
and on the northwest by a ycllow-dewlap-
ped subspecies. In both cases, the contrast
between the vinosus dewla]:) and that of
its neighbors is striking, and the races are
easily separable. In dorsal coloration,
vinosus is most like aurifcr, with a marbled
or mottled pattern of browns and greens.
As far as scales are concerned, there is
nothing distinctive about vinosus; along
with the Hispaniolan subspecies domini-
censis, sejunctus, favillarum, and aurifer,
vino.sus has 1/1 scales between the supra-
orbitals and the interparietals and 0/0
supraorbitals touching the interparietal.
With a mean of 3.4 median head scales,
vinosus ranks low among all subspecies, and
with a mean of 7.4 postmentals, it ranks
among the highest.
A. d. vinosus is extremely common
throughout its range and especially so at
Camp Perrin, where it was observed and
taken on trees and hedgerows along dirt
roads. At Carrefour Canon, these lizards
were abundant in a cafctal with cacao,
shaded by a high canopy. In Les Cayes,
A. d. vinosus was abundant about ^^'alls and
buildings, trees and gardens, etc.
Intergrades between vinosus and the
race to the northwest will be discussed
later. Intergrades between vinosus and
aurifcr are represented by a series of three
specimens from Cavaillon (ASFS X3729-31).
The two males in this short series had
dewlaps which had the basal maroon spot
paler (more reddish-orange) than in
vinosus, and the broad margin of the dew-
lap distinctly more orange — a combination
which I interpret as demonstrating inter-
gradation between the two subspecies.
These Cavaillon speciminis were collected
on the same day as the topot\pical series
of aurifcr, and direct comparisons of the
intensities of the dewlap colors in both
lots were made directh with on(> another.
I ha\(- associated the three specimens
from Tombeau Cheval ( MCZ 63132-34)
with vinosus rallier than aurifcr or the race
to the northw(\st on the basis of provenance.
Tombeau (/h('\al lies on about the high
point ol the load between Les Ca\'es and
Jeremie, and just north of (lamp Perrin.
Since Tonibeaii C!he\'al is closer to Camp
Penin than to an) other locality whence
Angus distichus • Schwartz
295
A. distichus is known in this region, I have
considered the specimens from that locaHty
as vinosus, although I admit the possibiHty
of error in such an assignment in this
particular region.
Anolis disfichus juliae Cochran
Anolis doniinicensis juliue Cochran, 1934, Occ.
Papers Boston Soc. Nat. Hist., 8:169.
Type localify. Ile-a-Vache, Haiti.
Definition: A subspecies of A. distichus
characterized by moderate size (males to
53 mm, females to 44 mm snout-vent
length), dorsum brownish-gray to green,
somewhat marbled with darker browns and
greens, dewlap almost completely dark
wine-red with a pale yellow margin, mod-
ally 0 0 scales between the supraorbital
semicircles and the interparietal and 0/0
supraorbitals in contact with the inter-
parietal, 2 '2 scales in contact laterally
with the postfrontals, and a low mean
number (3.4) of median azygous head
scales.
Distribution: Ile-a-Vache, Haiti ( Fig. 3).
Comments: A. d. juliae is obviously an
insular derivative of the mainland A. d.
vinosus, which it resembles in general dew-
lap pigmentation. Four characters separate
the two subspecies: 1) the wine-red pig-
ment in the dewlap of juliae is brighter
(more red) than that of vinosus, 2) the
extent of the wine-red spot is greater in
juliae than in vinosus (PI. H), 3) the
dorsum of juliae is generally paler and less
marbled and dark than that of vinosus,
and 4) the modal condition of 0/0 scales
between the semicircles and the inter-
parietal and 0 0 supraorbitals in contact
with the interparietal differ from the 1/1
and 0 0 (respectively) modes in vinosus.
Comparisons with the remaining races are
unnecessary, since no subspecies, other than
vinosus, has the red-blotched dewlap of
juliae.
Measurements and scale counts of the
holotype (a male) of A. d. juliae are:
snout-vent length 47 mm, tail ca. 49 mm,
broken; 4 scales across snout, 5 loreal rows,
semicircles in contact, 1 0 scales between
supraorbital semicircles and interparietal,
0/1 supraorbitals in contact with inter-
parietal, 3 2 scales in lateral contact with
postfrontals, 21 fourth toe lamellae, 4
median azygous head scales, "preoccipital"
present, 8 postmentals.
Scale counts on the series of 31 A. d.
juliae are: snout scales 4 to 8 (mode 4),
loreal rows 4 to 6 (mode 5); supraorbital
semicircles always in contact; modally 0/0
scales between the semicircles and the in-
terparietal and 0/0 supraorbitals in contact
with the interparietal; fourth toe lamellae
16 to 22 (mode 20); median azygous head
scales 2 to 6 (mode 4, mean 3.4); "preoc-
cipital" usually present (29 of 30 lizards);
postmentals 5 to 9 (mode 8, but 6 has
almost as high a frequency, mean 7.2).
Where we collected on the western end
of Ile-a-Vache, A. d. juliae was moderately
common, occurring about houses and on
trees in cultivated areas, as well as on
Cocos trunks in old coconut plantings.
Specimens examined: HAITI. Ile-d-
Vache: no other locality, 9 (MCZ 37517—
holotype, MCZ 37518-19— paratypes, MCZ
6171, MCZ 86767-71); western end, 22
(ASFS X3516-36, ASFS X3548).
Anolis distichus suppar^- subsp. n.
Holotype: MCZ 81137, an adult male,
from Dame-Marie, south side of town along
coast, Dept. du Sud, Haiti, taken 13 March
1966 bv Richard Thomas. Original number
V9236.'
Paratypes ( all from Dept. du Sud, Haiti ) :
ASFS V9237, same data as holotype; ASFS
V9268, ca. 5 km (airline) S Dame-Marie,
13 March 1966, R. Thomas; ASFS V9269,
ca. 10 km ( airline ) WSW Moron, 13 March
1966, R. Thomas; ASFS V9192-94, ASFS
V9213, ca. 7.5 km (airline) WSW Moron,
13 March 1966, E. Cyphale, R. Thomas;
MCZ 74766, MCZ 74768-810, MCZ
74812-25, Marfranc, 26-27 December 1962,
D. Hill and F. Vuillemier; USNM 160682-86,
^-' From Latin, sitfipar, almost equal.
296
BuUetin Museum of Comparaiwc Zoology, Vol. 137, No. 2
Jeremie, 9-10 March 1966, R. Thomas, na-
tive collectors; UF/FSM 21524-25, Jeremie,
11 March 1966, R. Thomas; AMNH
96501-04, Jeremie, 11 March 1966, R.
Thomas, native collector; MCZ 63106,
Jeremie, 31 July 1960, A. S. Rand and J. D.
Lazell, Jr.; MCZ 3346, MCZ 86772-77,
Jeremie, no date, D. F. Weinland; KU
93376-79, 2 km NW Jeremie, 14 March
1966, native collector; MCZ 69766-79, Car-
refour Sanon, nr. Jeremie, December 1962,
G. Whiteman; MCZ 69780-91, Place Negre,
nr. Jeremie, December 1962, G. Whiteman;
CM 37811 + 10 untagged specimens. Place
Negre, nr. Jeremie, 10-11 December 1961,
L. Whiteman; MCZ 69792-809, Mayette,
nr. Jeremie, December 1962, G. Whiteman;
MCZ 64630-37, Tiga, nr. Jeremie, 15 De-
cember 1960, G. and L. Whiteman; MCZ
69751, Lancenise, nr. Jeremie (not mapped),
December 1962, G. Whiteman; MCZ
69757-65, La Source, nr. Jeremie (not
mapped), December 1962, G. Whiteman;
MCZ 69754-55, Perine, nr. Jeremie (not
mapped), December 1962, G. Whiteman;
MCZ 65627-28, nr. Jeremie, 1960, L. and
G. Whiteman; MCZ 69752-53, Bozo, nr.
Jeremie ( not mapped ) , December 1962,
G. Whiteman; ASFS V9359-60, ca. 8 km
(airline) S Marche Leon, 3000 feet (915
meters), 15 March 1966, native collector.
Referred specimens: HAITI. Dcpt du
Slid: Tiburon, 6 (MCZ 6170, MCZ 86778-
82); Paroty, nr. Jeremie (not mapped), 1
(MCZ 64638); Place Negre, nr. Jeremie,
39 (MCZ 64675-713); nr. Jeremie, 7 (MCZ
3346).
Definiiion: A sul)speci{\s of A. distichiis
characterized by moderate size ( males to
54 mm, females to 44 mm snout-xcnt
length), dorsum pale green, somewhat
marbled with gray and yellow, dewlap
pale yellow to yellow-green or grayish
yellow and at times with a dull yellow-
orange basal smudge, modally 11 scales
between the supraorbital semicircles and
the interparietal and 0 0 supraorbitals in
contact with the interparietal, 2 2 scales in
contact laterally with the postfrontals, and
a low mean number (3.4) of median
azygous head scales.
Distrd)iition: The extreme western tip of
the Tiburon Peninsula in Haiti, from Dame-
Marie east to Jeremie, and south on the
northern slopes of the Massif de la Hotte
in the vicinity of Marche Leon; occurrence
at Tiburon problematical (see below)
(Fig. 3).
Comments: The terminal subspecies on
the western tip of the Tiburon Peninsula
is remarkably different from its neighbors
to the east [ourifer) and south (vinosus)
and in fact resembles its relative domini-
censis far to the east, both in dorsal color
and dewlap color. In having a yellow
dewlap (PI. II), suppar is readily distin-
guishable from aiirifer (orange dewlap)
and vinosus (maroon-centered dewlap).
The resemblances to dominiccnsis are
strong, including a dorsal green color, a
pale yellow dewlap, comparable means of
median head scales (3.4 and 3.9), and 1/1
scales between semicircles and interparietal
and 0/0 supraorbitals in contact with inter-
parietal. The major differences are the
higher mean number of postmentals (7.9
in suppar — the highest mean of any sub-
species— and 6.6 in dominicensis) and the
more pastel or paler green dorsum.
The measurements and counts for the
holotype of A. d. suppar are: snout-vent
length 52 mm, tail 65 mm; 5 scales across
snout, 5 loreal rows, semicircles in contact,
1/1 scales between supraorbital semicircles
and interparietal, 0 0 supraorbitals in con-
tact with interparietal, 2 2 scales in lateral
contact with postlrontals, 22 fourth toe
lamellae, 4 median head scales, "preoc-
cipifal" present, 7 postmentals.
Scale counts for the series oi 176 /\. d.
suppar are: snout scales 4 to 8 (mode 4),
loreal rows 3 to 6 (mode 5); modally 11
scales between supraorbital semicircles and
interparietal and 0 0 supraorbitals in con-
tact with interparietal; 2 2 scales in lateral
contact w ith postfrontals; fourth toe lamel-
lae 16 to 25 (mode 18 and 19); median
azygous head scales 1 to 7 (mode 3, mean
Anolis distichus ' Schwartz
297
3.4 ) ; "pieoccipital" usually present ( 170 of
176 lizards); postmentals 4 to 13 (mode 7,
mean 7.9 ) .
The dewlaps of A. d. suppar have been
noted in life as pale yellow-green (Jeremie,
Dame-Marie, and west-southwest of Moron)
and pale yellow (Marche Leon; pi. 17 E
1, west-southwest of Moron). A male from
the Moron region also has a dull yellow-
orange (pi. 9 J 10) basal smudge. The
dorsum is usually pale or pastel green,
somewhat overlaid with grayish marbling
and/or streaking, and commonly there are
yellow or paler green middorsal blotches,
especially on the anterior trunk and neck.
Preserved specimens, regardless of fresh-
ness of preservation, very regularly show
both a broad dark (black) V-shaped collar
which arises from about the angle of the
jaws and extends across the neck, and a
large dark (black) area on the upper side
of the head, separated from the collar by
a narrow paler (gray) V-shaped band.
Since no note of these markings was made
in life, they must not be conspicuous in the
living animal, but they are remarkably
consistent in the preserved lizards. I do
not know if suppar is capable of a brown
phase.
Comparisons of suppar with the adjacent
aurifer and vinosus were made above. From
the orange-dewlapped subspecies bimini-
cnsis, distichoides, dapsilis, igni^ularis,
tostus, and favilJarum, suppar differs in
having a yellow dewlap. From juJiac, sup-
par also differs in having a yellow dewlap
instead of a dewlap with an extensive
wine-colored basal blotch. From the yel-
low-dewlapped races (distichus, ocior, pro-
pcrus, scjunctus, and ravitergum; compari-
son with dominicensis was made above)
suppar differs in being (always?) green
(in contrast to distichus, properus, sejunctiis,
and ravitergum) and in lacking the lateral
pale line of ocior (although many suppar
have the flank stripe fairly well developed,
it is not clearly outlined above and below
by darker). Other differences from ocior
include a much lower mean number of
median head scales (3.4 versus 5.8), 1/1
scales between the semicircles and the
interparietal (0/0 in ocior), and 0/0 supra-
orbitals in contact with the interparietal
(1/1 in ocior).
The specimens from Tiburon were col-
lected by Garman and thus are quite old
and faded, and there are no color data on
them. I consider them suppar only pro-
visionally; Tiburon is 28 kilometers airline
south of Dame-Marie, but it may be
precisely in this intervening region that
suppar intergrades with vinosus. The
Tiburon lizards may be vinosus; there are
no specimens from any locality between
Dame-Marie and Tiburon, on the one hand,
or between Tiburon and Port-Salut, on the
other.
Although there is no evidence of inter-
gradation between suppar and vinosus,
there is evidence of intergradation between
suppar and aurifer. A series (MCZ
74826-32) from Roseaux was noted as
having the dewlap with a "deep orange
rust spot at base." It may be recalled
that there is a sight record of an aurifer
from 7.5 km (airline) south-southeast of
Roseaux. The zone of intergradation be-
tween suppar and aurifer appears to be
very narrow, centering in the region about
Roseaux.
A. d. suppar is quite common throughout
most of its range, occurring from sea level
to elevations of 3000 feet (915 meters)
above Marche Leon on the northern slopes
of the Massif de la Hotte. In habitat, it
does not differ from other altitudinally
wide-ranging races, in that it was taken in
edificarian situations, along the southern
slopes of the Monts Cartaches, and in both
natural and artificial wooded situations
which the species inhabits throughout its
range.
Anolis distichus patruelis^' subsp. n.
Holotypc: MCZ 81138, an adult male,
from vicinity of Pointe Sable, He Grande
^ • From Latin, patritelis, relating to a cousin.
298
Bulletin Museum of Compurative Zoologij, Vol. 137, No. 2
Cayemite, one of a series taken 18 March
1966 by Richard Thomas and native col-
lectors. Original number V9409.
Paratypes: ASFS V9410-14, ASFS V9423-
26, MCZ 81142-46, USNM 160687-91,
AMNH 96505-08, same data as holotype;
MCZ 25519, Grande Cayemite, 3 August
1927, W. J. Eyerdam.
Definition: A subspecies of A. distichus
characterized by small size (males to 50
mm, females to 42 mm snout-vent length),
dorsum green to gray, usually not promi-
nently striate, dewlap solid dark reddish
to mustard orange, modally 1/1 scales be-
tween the supraorbital semicircles and the
interparietals and 0/0 supraorbitals in con-
tact with the interparietal, 2/2 scales in
contact laterally with the postfrontals, and
a moderate mean number (4.6) of median
azygous head scales.
Distribution: He Crande Cayemite, Haiti
(Fig. 3).
Comments: The measurements and scale
counts for the holotype of A. d. patnielis
are: snout-vent length 49 mm, tail ca. 60
mm; 4 scales across snout, 4 loreal rows,
semicircles in contact, 1/1 scales between
supraorbital semicircles and interparietal,
0/0 supraorbitals in contact with inter-
parietal, 2/2 scales in lateral contact with
postfrontals, 19 fourth toe lamellae, 2
median head scales, "preoccipital" present,
8 postmentals.
The series of 25 A. d. patnielis has the
following counts: snout scales 4 to 8
(mode 4), loreal rows 4 to 6 (mode 4);
modally 1/1 scales between supraorbital
semicircles and interparietal and 0/0 supra-
orbitals in contact with interparietal; 2/2
scales in lateral contact with postfrontals;
iourth toe lamellae 16 to 22 (mode 20);
median azygous head scales 2 to 7 (mode
4, mean 4.6); "preoccipital" always present;
postmentals 6 to 11 (mode 8, mean 7.8).
The dorsum of A. d. pafruelis varies be-
tween green and gray; most specimens
show little or no striae, but others ha\(' a
lineate dorsum. The dewlap varies in life
from dark reddish to mustard orange (pi.
6 K 9, pi. 5 L 11), and has an orange
( rather than pale yellow ) margin ( PI. II ) .
He Grande Cayemite is adjacent to the
northern section of the presumed mainland
range of A. d. aurifer, and A. d. patruelis
resembles the former subspecies in dewlap
color. A major difference is the absence in
patnielis of the narrow yellow dewlap
margin which occurs in aiiiifer: the richer
and deeper hues of the patnielis dewlap
are likewise different than the brighter
pigments of aurifer. The back of aiirifer
is heavily marbled and mottled with greens
and browns, whereas that of patnielis is
generally much plainer, lacking pronounced
mottling, and is rarely clearly striate.
A. d. patnielis, with its deep orange
dewlap, differs from the subspecies which
have yellow dewlaps [distichus, ocior,
dominicensis, propenis, sejunctus, raviter-
gum, and suppar) and those which have
a maroon or wine-red basal spot ( vinosus,
juliae). The other orange-dewlapped sub-
species are himiniensis, distichoides, dapsilis,
ignigularis, tostus, favillanim (and aurifer,
with which patnielis was compared above).
Aside from the differences in dorsal pig-
mentation and pattern, the deeper hue of
the dewlap color, and the absence of a
yellow dewlap margin in patruelis, the
Grande Cayemite subspecies differs from
all other orange-dewlapped forms in having
a very high mean of postmentals (7.8 in
patruelis. 3.7 to 7.2 in other orange-throated
subspecies, with aurifer luuing the highest
mean ). In fact, other than suppar, patruelis
has a higher postmental mean than all
other subspecies.
Most of the i)arat\'pic series were native
collected; the lizards came from dry
scrubby woods growing on almost bare
limestone and from about the \illage at
Pointe Sable.
There is a short series (USNM 80814-18)
of A. distiehtis from He Petite ('ayemite,
just to the west of Grande Cayemite. These
specimens have long bcH'n in preservati\e,
and consequent 1\ no details of coloration
or pattern are discernible. They may be
Anolis distichus • Schwartz
299
fxitnielis, although, as pointed out for
Ameiva taeniura Cope, which is known
from lioth the Cayemites, there is a pos-
sibiHty that each island has its own sub-
species (Schwartz, 1967a). In this short
Petite Cayemite series of five specimens,
the postmentals range between 4 and 7
(two specimens have counts of 4 and 5,
and are thus lower than the much longer
series from Grande Cayemite). One lizard
(USNM 80818) has only a single median
azygous head scale, the "preoccipital," a
condition not observed in the Grande
Cayemite series. I consider the Petite
Cayemite lizards A. d. patniclis only pro-
visionally.
The Florida Populations
Anolis- distichus was first reported from
the continental United States by Smith and
McCauley (1948), who described A. d.
floridanits on the basis of a short series of
six specimens from Brickell Park in down-
town Miami, Florida. The status of the
mainland lizards was later discussed by
Duellman and Schwartz (1958:279-281),
who regarded floridanus as a synonym of
A. d. distichus. Of the four scale characters
and one pigmental trait, these authors noted
that "floridanus' (of which form they ex-
amined 77 specimens in detail ) agreed with
topotypical distichus in number of in-
fraorbital scales, number of scales bordering
the median suture ( = median azygous head
scales), and in having the throat unpig-
mented, but disagreed with the nominate
subspecies in having a higher percentage
(63.6 per cent versus 14.0 per cent) of
specimens with the supraocular semicircles
separated and in the modal number of
scales separating the prefrontal from the
anterior supraocular (mode 1 and 2 with
almost equal frequencies in "floridanus,"
mode 2 in topotypical distichus). With
increased knowledge of Anolis distichus in
its insular range, it is appropriate to reas-
sess the status not only of "floridanus" but
also that of another mainland Floridian
population.
Through the efforts of C. Rhea Warren,
I have been able to examine a short series
of seven lizards ( RT 1478-84 ) from North-
west South River Drive in Miami, Florida.
These lizards were green in life and cap-
able of becoming solid brown; they repre-
sent a small sample from a large and very
successful colony centering near the junc-
tion of the Miami Canal ( the northwestern
extension of the Miami River) and the
artificial Tamiami Canal. The specimens
are typical in all ways of A. d. dominicemis,
with the possible exception of three of the
seven specimens having 3/3 scales in lat-
eral contact with the postfrontals. This
high frequency is doubtless due to the
small sample size; in the series of 245 A. d.
dominicensis from Hispaniola, 32 have 3/3
scales in lateral contact with the post-
frontals, whereas 162 have 2/2 scales in this
position. The yellow dewlaps with oc-
casional orange basal blush and the green
dorsa agree in detail with my concepts of
A. d. dominicensis, and I assume that these
lizards have been recently introduced into
this region through some fluke of inter-
national shipping. King and Krakauer
(1966:146) have reported this population
as A. d. dominicensis at my suggestion.
AnoWs disfichus floridanus Smith and
McCauley
Anolis distichus floridanus Smith and McCauley,
1948, Proc. Biol. Soc. Washington, 61:160.
Type locality: Brickell Park, Miami,
Dade County, Florida.
Definition: A subspecies of A. distichus
characterized by small size (males to 50
mm, females to 45 mm snout-vent length),
dorsum gray to dark brown and without a
green phase, dewlap pale yellow ( occasion-
ally pale orange), modally 1/1 scales be-
tween the supraorbital semicircles and in-
terparietal, 0 0 supraorbitals in contact
with the interparietal, 2/3 scales in contact
laterally with the postfrontal, and very high
mean number (7.9) of median azygous head
scales correlated with the very high in-
cidence (about 60 per cent) of complete
300
Bulletin Mtisciiin of Comparative Zoolop,!/, Vol. 137, No. 2
separation of supraocular semicircles medi-
ally.
Distribution: Known only from the ex-
treme eastern coastal and near-coastal mar-
gin of Dade County, Florida ( Fig. 2 ) .
Comments: I have examined 90 A.
clistichus (aside from the A. d. dominiccnsis
noted above) from southern Florida. Al-
though Duellman and Schwartz (1958:
279-281 ) considered A. d. floridanus syn-
onymous with A. d. disticJius from New
Providence, the above definition clearly
shows that floridanus differs from distichus
in several characters which elsewhere in
the Bahamas and Ilispaniola I regard as
indicative of subspecifity. The use of the
name A. d. floridanus for the continental
lizards mainly involves the philosophical
problem of its appropriateness if the main-
land A. distic])us have been introduced
only recently by man. This question is
discussed below.
The two major samples which I have
studied come from two localities ( Brickell
Park and its vicinity in downtown Miami,
and Fairchild Tropical Garden). Mr. War-
ren advises me that A. disticJius occurs
elsewhere in Miami and in Coral Gables,
Florida, and Wayne King ( in litt., 28
September 1966) reported its occurrence
at one additional locality in Miami, four in
Coral Gables, three in Coconut Grove, and
one in Kendall. I have not examined ma-
terial from any of these localities. Dr.
King suggests that the Brickell Park-Fair-
child Garden population is continuous (the
Coconut Grove localities and a locality at
the junction of Brickell Avenue and the
Kickenbacker Causeway in downtown Mi-
ami I ill in fairly well the hiatus between
the two presumed terminal stations for
A. d. floridanus), and I concur. King and
Krakauer (1966:146) stated that all other
localities are the result of secondary in-
troductions by reptile fanciers; a second
method for dispersal may be that Fairchild
Garden supplies plants for ornamental pur-
poses to Dade County and to jirixate
persons for decorative planting, with re-
sultant accidental distribution of A. flori-
danus throughout the county.
The two terminal localities are distant
about 8.5 miles ( 1.3.6 km ) from one another.
Both are more or less coastal, and Fairchild
Tropical Garden has for many years been
a center to which plants from outside the
United States have been introduced for
purposes of culture and exhibit. The
Brickell Park locality lies in downtown
Miami on the coast on the south side of the
Miami River. The lizards are extremely
abundant at both localities. In most char-
acters the two samples are alike, and if
they represent two different "introductions,"
their later convergence has been along re-
markably similar lines.
The largest mainland male and female
are both from Brickell Park (snout- vent
length 50 mm in the male, 45 mm in the
female), whereas the largest of each sex
from Fairchild Garden are 47 mm and 39
mm. The scale characters of the two pop-
ulations are: snout scales 3 to 6 (mode 6)
at Brickell Park, 3 to 7 (mode 6) at Fair-
child Garden; loreal rows 4 and 5 (mode
4) at Brickell Park, 4 to 6 (mode 4) at
Fairchild Garden; semicircles usually not
in contact (27 of 42 lizards from Brickell
Park, 25 of 48 lizards from Fairchild Gar-
den ) ; modally 1/1 scales between the supra-
orbital semicircles and the interparietal and
0 0 supraorbitals in contact with the in-
terparietal in both samples; fourth toe
lamellae 15 to 19 (both localities), modes
17 or 18 (Brickell Park) and 18 or 19
(Fairchild Garden); modally 3/3 scales in
lateral contact with postfrontals at Brickell
Park, liut almost an (^((ual frequenc)' of
2 2 at this localit\ ; inodalh' 2 3 scales in
lateral contact with postfrontals at Fair-
child Garden; median azygous luad scales
5 to 12 (mode 8, mean 8.0) at Brickell
Park, 4 to fO (mode 8, mean 7.8) at Fair-
child (harden; "preoccij^ital" usualK" present
(40 of 42 h/ards from Brickell Park, 42
of 48 lizards from Faiichild Garden); post-
mentals 3 to 6 (mode 4, mean 4.3) at
lirickcl! Park, 3 to 8 (mode 5 or 6, mean
Angus Disricnus • Schwartz
301
4.4) at Fairchild Garden. The dewlap is
pale yellow to yellow with an extensive
pale orange blush; dorsally the lizards are
gray, incapable of a green phase but cap-
able of becoming dark brown.
The mainland A. disticlius obviously are
related to the Bahaman populations of the
species rather than to the Hispaniolan
forms. Such features as the high incidence
of complete separation of the semicircles,
the low number of postmentals, and the
lack of a green phase all point to the
Bahamas as the place of origin of A. d.
fJoridanus. It has generally been assumed
tliat the continental A. disticlius are the
result of a very recent introduction, either
fortuitous or intentional, by man from the
Bahamas. If such were the case, it should
be a simple matter to determine from which
of the five Bahaman subspecies floridoniis
has been drawn. This is not the case; A.
(I. floridaiius presents a suite of characters
which distinguishes it from all Bahaman,
as well as Hispaniolan, populations. If the
forerunners of floridanus were only recently
introduced by man, then differentiation in
Florida of floridanus must have been ex-
tremely rapid. If, on the other hand, A. d.
floridanus has had a history other than that
generally accepted — i.e., it has been in
Florida for a longer period or A. d. flori-
danus has been introduced only recently
luit evolved its peculiar characteristics else-
where— its differences from any other sub-
species could be accounted for more readily.
Evidence for the relationship and a sug-
gested history of the continental popula-
tions are offered below.
Turning first to dorsal and dewlap colors,
floridanus resembles all the Bahaman sub-
species except ocior in the former (since
floridanus lacks a green phase) and only
distichus in the latter. The scale char-
acters, on the other hand, are distinctive.
The very high incidence of complete separa-
tion of the semicircles (57.8 per cent if
both samples are combined; 64.2 per cent
in the Brickell Park sample alone, 52.1
per cent in the Fairchild Garden sample
alone) is much greater than that of any
Bahaman subspecies, being approached
most closely by distichoides (32.1 per
cent) and biminiensis (30.2 per cent). In
modally having 1/1 scales between the
semicircles and the interparietal, floridanus
differs from all Bahaman populations ex-
cept distichoides. New Providence and
Exuma Cays A. d. distichus do have 1/1
as the modal condition (or as a bimode in
the former case), however. Although the
modal condition is 2/3 scales in lateral
contact with the postfrontals in floridanus
(30 individuals), 29 lizards have 2/2 scales
in lateral contact and 25 have 3/3 (of
which 15 are from Brickell Park, where
3/3 is the mode). Such a high incidence
of 3/3 scales in lateral contact with the
postfrontals is unequalled in any Bahaman
population except biminiensis, where 3/3
is the mode.
The mean of 7.9 median azygous head
scales in floridanus is higher than that of
any Bahaman subspecies with the excep-
tion of 8.7 in distichoides. In having 0/0
supraorbitals in contact with the inter-
parietal, floridanus is like disticlius, disti-
choides, and dapsilis, but unlike bimini-
ensis and ocior. The regular occurrence of
the "preoccipital" in floridanus resembles
the condition in all Bahaman subspecies
except distichoides and biminiensis which
more often lack the "preoccipital." Finally,
the mean of 4.4 postmentals in floridanus
is lower than those of all Bahaman pop-
ulations, being most closely approached by
distichoides (4.6) and biminiensis (4.8).
From the above resume, it is apparent
that, although floridanus agrees with nomi-
nate distichus in dorsal and dewlap colors,
it differs markedly from it in scale char-
acters. The two populations which bear
the closest resemblance in scalation to
floridanus are distichoides and biminiensis,
and, not unexpectedly, these two races are
those most geographically adjacent to
floridanus. The occasional occurrence in
floridanus of a pale orange dewlap also
suggests that one or the other of these
302
BiiUetin Museum of Comparative Zoology, Vol. 137, No. 2
orange-dewlapped subspecies may have
been the source of floridanu.s.
There are three possible histories for the
continental population of A. distichus:
1 ) The populations were indeed ac-
cidentally or purposely introduced by man
in the relatively recent past (shortly prior
to their discovery and description in 194(S)
from somewhere in the Bahamas.
2) The populations are the result of a
natural overseas introduction from some-
where in the Bahamas whose population
as yet remains unsampled; the Bimini chain
or Andros seem likely candidates, but A. d.
floridanu.s does not agree with either
himinicn.si.s or distichoide.s in all details of
color or scalation.
3) The populations reached Florida dur-
ing the Pleistocene from the Bahamas but
remained undetected there until 1948, by
which time their precise place of origin in
the Bahamas had become obscure because
of in situ evolution to A. d. floridanu.s on
the mainland.
Several facts should be taken into con-
sid(Mation before proceeding. 1 ) A. d. flo-
ridanu.s is closest in characteristics to those
Bahaman lizards which are geographically
most nearly adjacent to the mainland — J)i-
minicnsis and di.stichoide.s. 2) The distri-
bution of A. d. floridanu.s is primarily coastal
(as one might expect of a natural invader)
and is not now (and perhaps was not ever)
disjunct, as prc>viously supposed. 3) Al-
though there is much overseas boat traffic
between the Biminis and Miami, there is
less between Andros and Florida; Dr. King
has pointed out that "in the early nineteen
hundri'ds there were large foiu--, five- and
six-masted barks tliat sailed freight all over
the Bahamas and between th(> Bahamas and
Florida. Any of tliese could have been a
vehicle for introducing di.sficJiu.s into l^'lor-
ida." 4) That A. distichus arrived in Flor-
ida in the Pleistocene but remained unde-
tected until the jiresent century may tak(>
credulity. It is not impossible that such a
chain of events took place, since the Miami
region has only in the present century been
a large urban center, and small or restricted
coastal populations of this fast-moving and
inconspicuous lizard could have been easily
overlooked by earlier collectors and visiting
scientists, who likely were more concerned
with protecting themselves from mosqui-
toes in the coastal regions where A. d. flori-
danus occurs. 5) Along these lines it is in-
teresting to note that A. d. distichus was
first described from New Providence in
1861, whereas Nassau was the home of the
First Royal Governor of the Bahamas in
1718, and through the following century
and a half became a veritable Bahaman
metropolis, far more visited by travelers
and scientists than the Miami region. No
one suggests that A. d. di.stichus owes its
tenancy of New Providence to the years
just prior to the year it was described, yet
this has always been the assumption of the
status of A. d. floridanus.
Considering all of the above facts and
suggestions, I adhere to a combined se-
quence of events as regards A. d. floridanus
as noted above in postulations ( 1 ) and
( 2 ) . I think it most likely that A. d. florid-
anus was introduced, either by natural
overseas transport or by man long before
its discoxery in 1948 but in historical times.
The source of this introduction remains a
mystery, but one region may be mentioned.
The west coast of Andros is ver\' poorly
known zoologically, and this coast is the
one closest to the Florida mainland. The
fragmentation of Andros by cross-island
waterways and bights and its large size
(Andros is as large as Puerto Rico but less
physiographiealK' di\'erse) suggest that in-
tra-island dilfercMitiation in somi- reptiles
may ha\(' taken place there. This is indeed
the ease with Andros Amciva auJycri (per-
sonal eoninumicatioii, (Clarence J. McCoy,
Jr.), and there* is no reason to doubt tliat it
occurred in other rei)til(\s. \'erification of
this supposition can be had only by collec-
tion oi series ol specimcMis from this inacces-
sible west coast. Dr. King is th(> onh' scien-
tist who has crossed the Big Mud to western
Andros in recent times, and he secured but
Anolis distichus • Schwartz
303
a single female A. distichus there. Western
Andros remains the only extensive area in
the Bahamas whose fauna is extremely
poorly known.
The geographical juxtaposition of this
coast to Florida makes it even more attrac-
tive as a source of introduction of A. d.
floridanus. The assumed derivation of A.
d. floridanus from A. d. distichoidcs is logi-
cal, considering the resemblances between
the two in some scale features. Dr. King
{in lift., 30 September 1966) suggested
that the Bahaman sponge fleet used to ply
the waters of the large banks in the Ba-
hamas (including the Big Mud west of
Andros). Boats from the fleet put into
Miami for sale of cargo until the Baha-
man sponge industry was destroyed by the
sponge blight in 1938 and 1939. Inten-
tional or accidental transportation of A. d.
floridanus from the west coast of Andros
to the Miami area might well have been
effected by ships in the sponge trade. If
such is the case, then, A. d. floridanus,
despite its name, is in reality a Bahaman
subspecies from the western section of
Andros which has been introduced into
Florida in the relatively recent past.
Specimens examined: FLORIDA. Dade
County: Brickell Park, 35 (UMMZ
106189-31 specimens, UMMZ 108100,
UMMZ 108372-3 specimens); south of
Miami River on Brickell Ave., Miami, 7
(UMMZ 109232); Fairehild Tropical Gar-
den, Miami, 48 (RT 1485-97, UMMZ
108189-2 specimens, UMMZ 108190-6 speci-
mens, UMMZ 108371-25 specimens, UMMZ
109231-2 specimens).
DISCUSSION
Anolis distichus is one of the most widely
distributed species of amphibian or reptile
in the Antillean region; its occurrence on
Hispaniola, the Bahamas, and Florida is
exceeded by that of another anole {Anolis
sa^rei Dumeril and Bibron on the Bahama
Islands north of the Crooked Island Passage,
Cuba and the Isla de Pinos, Jamaica, Cay-
man Islands, Peninsula de Yucatan, Swan
Islands, Florida Keys), of TypMops htm-
hricalis Linnaeus ( Bahama Islands north
of the Crooked Island Passage, Cuba and
the Isla de Pinos, Hispaniola), and of the
boa Epicrates aufiulifer (Great Bahama
Bank, Sheep Cay off Great Inagua, Cuba
and the Isla de Pinos, Hispaniola). Through-
out its range, A. distichus varies in success;
in more mesic situations and on some is-
lands, it may be very abundant, but in
xeric regions it is less so or absent.
As far as the Bahaman herpetofauna is
concerned, A. distichus holds a unique posi-
tion. It is generally conceded that the
Bahaman herpetofauna has been derived
mainly from that of Cuba, with a smaller
Hispaniolan element. There are 13 Cuban
forms in the Bahamas ( Ilyla septentrionalis,
EleutJierodactylus planirostris, Sphaero-
dactylus decoratus, SphaerodactyJus notatus,
Tarcntola americana, Anolis angusticcps,
Anolis caroUnensis, Anolis sagrei, Leio-
cephalus carina! us, Leiocephalus loxogram-
mus, Ameiva auberi, Typhlops biminiensis
and Typhlops lumbricalis). These species
all have Bahaman and Cuban populations
which are identical or only racially dif-
ferentiated, with the exception of L.
loxogrammus (which is endemic to the
Bahamas but closely related to the Cuban
L. raviceps) and Typhlops lumbricalis (in
which the nominate form occurs both in
the Bahamas and in part of eastern Cuba ) .
The latter species likewise is presumed
( Thomas, in press ) to have originated on
Hispaniola rather than Cuba, but it is in-
cluded here as a Cuban element in the
Bahaman fauna, since the Bahaman pop-
ulations have been derived directly from
Cuba rather than from Hispaniola. In gen-
eral, the Cuban species are limited in the
Bahamas to the islands north of the Crooked
Island Passage (i.e., the Great and Little
Bahama banks), although there are excep-
tions (L. loxogrammus), and some forms
may not occur on both banks (A. caro-
Unensis) or may occur only, but not be
widespread, on the Great Bahama Bank
( T. americana). However, compared to
304 BuUetin Museum of Comparative Zoology, Vol. 137, No. 2
the Hispaniolan faunal element in the Ba-
hamas, the species with Cuban affinities
are widely distributed.
The Bahaman species with Hispaniolan
affinities include eight species (Aristelliger
cochranae, Spliaerodoctyhis inag,uae, A7ioli.s
distichus, Leiocephalus inoguoe, Leiocepha-
Iii.s arcmirius, Ameiva mcnjnardi, Epicrates
cxsul, and Epicrates angidifcr). I include
E. an^idifcr in this series rather than
with the Cuban element (the species
occurs on both islands), since the more
widely ranging Bahaman subspecies is
more closely related to the Hispaniolan than
the Cuban race. Of these eight species,
five are limited to the islands between the
Crooked Island Passage and Hispaniola,
one {E. exsid) to the Little Bahama Bank,
and the remaining two (A. distichus and
E. anguJifcr) occur on the Great Bahama
Bank (or at least primarily on this bank).
Spluierodactyhis ''anthrocinus'' on New
Providence and Andros is without doubt
introduced from Hispaniola, since the Baha-
man form {"anthracinus') is identical with
one of the Hispaniolan subspecies of S.
copci. Thus of the eight Bahaman species
with Hispaniolan affinities, only two (A.
distichus and E. angulifer) are widespread
in the Bahamas, whereas 12 of the 13
Cuban species are widespread on the Great
Bank. (There are 19 other species of
reptiles — including seven nominal species of
the genus Cyclura — in the Bahamas, but
these are not readily classifiable as to origin;
none is obviously or certainly related to
either Hispaniolan or C'uban congeners,
and need not concern us furtlier in this
context. )
Froiu the above brief summary of the
affinities of the liahaman herpetofauna, it
apjx'ars that A. distichus has a rather
uni({ue position therein, since it is one of
the two Bahaman reptiles which are wid(>-
spread in the Ikihamas and haxc had an
Hispaniolan origin. Noteworthy is the fact
that A. distichus is absent from the Baha-
mas south of the Crooked Island Passag(\
and from the Turks and Caicos islands.
This is peculiar, since these islands would
seem to have been likely and handy
stepping-stones from the Hispaniolan main-
land to the Great Bahama Bank. Not only
is A. disticJius absent from these islands,
but they have only a single anolis (A.
scriptus), which is not closely related to
A. distichus (scriptus, although associated
with disticlius as an Eastern Island Alpha
anole, belongs to the cristatellus rather
than the bimocukitus group; fide Etheridge,
in litt.). There is no distichus relative in
Cuba, the more logical place for invasion
of the Great Bahama Bank. It seems then
that A. disticluis arrived on the Great Ba-
hama Bank directly from Hispaniola, with-
out using either Cuba or the southern Ba-
hamas south of the Crooked Island Passage
as way stations.
I have already commented on the general,
although not absolute differences between
the Bahaman and Hispaniolan segments of
A. distichus. These differences are not
especially striking, either in dewlap color
and pattern or in dorsal color. The com-
plete absence of separation of the semi-
circles in any Hispaniolan A. distichus, the
higher mean numbers of median head
scales in the Bahamas, and the tendency
toward absence of the "preoccipital' in the
Bahamas do indicate, however, that the
Bahaman lizards have diverged as a unit
in some scale characters. Most Bahaman
subspecies have lost the ability to become
green, and this phenomenon has taken place
also in some Hispaniolan populations. In
general, the loss of the green phase is
correlated with more arid, in contrast to
distinctly mesic, situations. lIowcNcr, the
range of A. d. ocior in the Bahamas is not
(specially more mesic than that of the
other Bahaman su])species, and ocior re-
tains a green phase. It is suggestive that
those islands (Rum Cay and San Salvador)
inhabited by ocior are two ol tlie three
Bahaman islands occupied by A. distichus
which are not on the Great Bank. The
other (\\ception. Cat Island, is presently
censis) in ha\ing a xcllow d(>wlap and
Anolis distichus ' Schwartz
305
biminien sis distichoides dapsllis
florid anus
ocior
dominicensis
1
supper favillarum I ravitergum ignigularis
^ \ I ^ ^
vinosus aurifer ' tostus properus
juliae patruelis , sejunctus
Figure 4. Dendrogram of the relationships between the subspecies of Ano//s distichus. Bohaman subspecies above solid
horizontal line, Hispaniolan subspecies below solid horizontal line; subspecies on the south island of Hispaniola and its
satellite islands enclosed within dashed line in lower left of diagram.
cut off from the Great Bank, but in contrast
to Rum and San Salvador, Cat is not far
removed from the Great Bank and is still
connected to Eleuthera by a narrow sub-
marine strip (Clench, 1938:536). Its his-
tory has been at least partially associated
with that of the Great Bank.
Not only does A. d. disticJius have the
broadest distribution of any subspecies in
the Bahamas, but it also resembles the most
widespread Hispaniolan race ( A. d. domini-
also is the Bahaman subspecies which
occurs closest geographically to Hispaniola
on the Ragged Islands and Long Island.
A. d. distiduis seems appropriate, both
geographically and in dewlap color, as a
direct Bahaman derivative from Hispaniola
(Fig. 4). In the Bahamas, various sub-
species have differentiated from A. d.
distichus on more or less peripheral islands
or island groups. Thus, biminiensis, disti-
choides, and dapsilis are all essentially
orange-dewlapped A. distichus with head
scutellation features differing from those
of A. d. distichus but occurring in the
latter subspecies as casual variants.
A. d. ocior, in contrast to the balance of
the Bahaman races, retains a green phase
and a yellow dewlap — characters which
ally it directly with dominicensis. It does,
however, have some "Bahaman" charac-
teristics, such as the high number of median
head scales. Possibly ocior reached Rum
Cay and San Salvador from the Great
Bahama Bank prior to the loss of the green
phase by the parental stock there, but at a
time when some head scale modifications
were already established or becoming so.
I do not consider it likely that ocior has
had a direct and separate connection with
dominicensis, but rather that it has had a
long history independent of that of the
balance of the Bahaman races.
On Hispaniola, the situation is more
306
Bulletin Museum of Comparative Zoology, Vol. 137, No. 2
complex. In essence there is a single, wide-
spread, yellow-dewlapped subspecies (do-
niinicensis) with a series of "peripheral"
races on the mainland {i<i,m<i,ularis, propcrus,
ravitcrii,um, fa villa rum, aurifer, vino.sus,
■suppar) and a series of subspecies from the
satellite islands (sejunctus, to.stus, jtiliae,
patruelis). Of the mainland subspecies,
those with plain or drab dorsa and yellow
dewlaps in general inhabit the more arid
regions {propcrus, ravitc'rfi,tim) and the
orange-dewlapped and brightly colored sub-
species inhabit both lowland and highland
mesic areas {ipiigulari.s, favilkirum, aurifer).
A. d. suppar is a distinct exception to this
statement, since the extreme tip of the
Tiburon Peninsula is mesic, and .suppar is
a green lizard with a yellow dewlap, much
like dominicensis. The most strikingly dif-
ferent mainland subspecies in dewlap color
is vinosus.
Of the satellite island races, all but one
resemble their relatives on the immediately
adjacent mainland in dewlap color and in
color repertory. The major exception is
A. d. to.stu.s on Isla Catalina. The adjacent
mainland is inhabited by propcrus, which
is yellow-dewlapped, whereas tostus has
essentially an orange dewlap, more like
that of ii!,nii[i,ularis to the west. The faunal
history of Isla Catalina is peculiar, in that
it includes an endemic and relict sub-
species of Amciva lincolata and a popula-
tion of A. clirijsolacma, a species which is
not known from the adjacent coast. Both
these lizards, as well as tostus, show dis-
tinctly more western than adjacent or east-
em affinities. Under these circumstances
it is likely that tostus represents an iiiniiiu-
laris, rather than a ])ropcrus, derivative,
and that through changing conditions on
the adjacent mainland, there has been a
shift to the westward of subspecic\s along
the coast, with propcrus replacing iii,niii,u-
laris. As far as the l)ahuice of the satc>lHte
island races is concerned, none pres(>nts
any problem. It is noteworthy that jidiac
shares with vinosus on the adjacent main-
land the strikingly different style of dew-
lap pigmentation and pattern.
Although not now a satellite island, the
Peninsula de Samana presumably was so at
one time. The occurrence there of a disjunct
population of ignigularis has been discussed
in detail in the text.
There is no \^'ay of determining \\'hether
A. distichus was primarily and primitively
an inhabitant of the historical north or south
island {.scnsu Williams, 1961) of Hispaniola.
The occurrence of dominicensis on most
of the north island and on the basal half
of the Tiburon Peninsula suggests in some
ways that the species was primarily north
island, and invaded the south island sec-
ondarily. This thesis presents the problem
of the very distinct and apparently isolated
favilkirum in the Sierra de Baoruco, and
the exceptionally distinctive vino.sus and
juliae on the distal Tiburon Peninsula and
Ile-a-Vache. The three subspecies on the
tip of the Tiburon Peninsula {suppar,
vino.sus, aurifer), of \\'hich two are quite
different in dewlap color from dominicensis,
may indicate that this region was colonized
directly from the north island across the
Golfe de la Gonave and not seriallv along
the peninsula itself. If so, then these ter-
minal populations may ha\e di\erged in-
dependently and remained isolated from
other A. di.sticJius populations until the
arrival of dominicensis across the much
narrower inter-island strait and subseciuent
contact along the Tiburon Peninsula. A. d.
favilkirum in the Sierra de Baoruco ma\'
represent still another isolated derixative
from a north island stock. I cannot suggest
that favillarum was historically derived
from the adjacent lowland and drab
ravitcrgum, howe\(>r. Another possibilit)'
is that favillarum is an njiland offshoot
irom dominicensis (just as, to the east,
ignigularis is an orange-dewlapped dotnini-
ccnsis deri\ati\t' ) in the Sierra de Baoruco,
and thus has exoKcd rather iccently. The
ai^i^arent absence ol /\. distichus from tht>
southern side ol the \'alle de Neiba in this
Angus distichus • Schwartz
307
particular region seems to enforce the latter
suggested derivation of favillorum.
Although A. (lisficluts is very widespread
in Hispaniola, its absence in two regions
is remarkable. The species is unknown
from He de la Gonave, which is inhabited
by the A. distichus cognate, A. hrevirostris.
Although the coast of Gonave is hot and
arid, the interior is less hostile and more
shaded. It is strange that A. distichus and
A. hrevirostris do not share Gonave as they
do similar portions of the mainland.
The other major region whence A.
distichus is absent is the Peninsula de
Barahona. The lowlands of this peninsula,
south of the Sierra de Baoruco, are in-
habited exclusively by A. hrevirostris, again,
as on Gonave, despite the ample avail-
ability of apparently suitable habitat for
A. distichus. The species likewise is un-
known from the coastal lowlands on the
Peninsula from the city of Barahona south
(A. hrevirostris is the exclusive species of
the pair in Barahona itself and its environs),
and also apparently along the southern
Haitian coast between the Dominican vil-
lage of Pedernales east to the area near
Jacmel. (There is some doubt in this latter
case because of the confused and unlocat-
able records for A. distichus labeled as
coming from localities "near Saltrou." At
least, all locatable stations where A.
distichus has been taken "near Saltrou" are
upland, and this is nicely correlated with
the occurrence of the species on the Domini-
can side of the border north of Pedernales
in the extreme eastern Sierra de Baoruco.)
The Peninsula de Barahona south of the
Sierra de Baoruco is emerging in His-
paniolan herpetology as a most distinctive
area. The high mountain massifs of the
Baoruco to the north and the La Selle to
the west, coupled with the narrow and steep
coastal "plain" at the eastern end of the
Sierra de Baoruco, effectively trap lowland
xerophiles to the south. Included in the
list of such disjunct or practically disjunct
forms are Typhlops sijntherus and Lepto-
typhlops pyrites as endemic species; Ameiva
chrysolaema ficta and A. c. leheri, Ameiva
lineolata privigna, Leiocephalus harahonen-
sis oxygastcr and aureus, DipJogJossus cur-
tissi aporus, Amphishaena gonavensis hy-
porissor and A. g. leheri as endemic
subspecies. Although A. hrevirostris is by
no means restricted to this region, it is of
interest that A. distichus has been unable
to penetrate it either along the steep eastern
coastal "plain," or from the uplands of the
eastern Massif de la Selle or from the
Sierra de Baoruco. A. hrevirostris is the
conspicuous and common member of the
pair in the lowlands south of the moun-
tains.
Although the Peninsula de Barahona is
in general arid, there seem to be ample
areas which would be quite suitable for
A. disticlius; such regions are invariably
inhabited by A. hrevirostris. Correlated
with the absence of A. distichus from the
Peninsula de Barahona is its absence from
Isla Beata (which has the endemic sub-
species A. hrevirostris ivetmorei). The
Beata fauna is easily derivable from that
on the adjacent Peninsula de Barahona, and
the lack of A. distichus on Beata is not
noteworthy. The relationships of AnoUs
altavelensis on Isla Alto Velo have already
been noted.
Although I am reluctant to attribute the
absence of a species from a particular region
to the catch-all phenomenon of competi-
tion, an explanation which may be glibly
invoked without precise data, the situation
between A. distichus and A. hrevirostris
suggests very strongly that competition may
indeed be the reason for the absence of
the former in some regions occupied by the
latter. The He de la Gonave and the
Peninsula de Barahona are both arid re-
gions. In the Cul de Sac Plain, where mesic
oases occur within otherwise arid scrub,
A. distichus is confined to the former habi-
tat, whereas the latter is occupied by A.
hrevirostris. Wherever the two species oc-
cur sympatrically in an arid situation, A.
hrevirostris is regularly the more "success-
ful" and A. distichus the species whose
308 Bulletin Museum of Comparative Zoology, Vol. 137, No. 2
distribution is limited to favoral)le pockets
within the area occupied by A. hrevirostris.
Such encounters ( Thomazeau-Manneville;
Haitian coast in the Jacmel area; arid
coast on the north shore of the Golte de la
Gonave ) invariably are "unfavorable" for
A. cUsticlius. If the He de la Gonave and
the Peninsula de Barahona were originally
colonized by A. hrevirostris, it seems likely
that A. distichus may simply not have been
able to penetrate into these regions to
reach ecologically suitable habitats (shaded
woods, oases, etc. ) because of the previous
presence there of A. hrevirostris. The situa-
tion on Gonave may be less complex, since
as an off-shore island, Gonave may never
have been reached by A. distichus. On the
other hand, it seems plausible, in the light
of evidence from areas of contact between
A. hrevirostris and A. distichus elsewhere
and the absence of A. distichus from the
Peninsula de Barahona, that A. distichus
may not be able to compete with A.
hrevirostris on Gonave where the latter
species is already well established.
In summary, A. distichus has a wide
distribution in the Bahama Islands and on
Hispaniola, having arrived in the Bahamas
directly from Hispaniola without employing
either Guba or the southern Bahamas as
way stations. These two major segments of
A. distichus have been isolated from one
another for a sufficiently long period for
some differentiation to have taken place
between them, but in general they are
similar. In the Bahamas, A. d. distichus is
considered the basic stock whence have
been derived four peripheral subspecies,
of which one (ocior) was isolated on Rum
Cay and San Salvador prior to the separa-
tion of the remaining three Bahaman races
from A. d. distichus. On Hispaniola, A. d.
dominicensis is suggested as a north island
parent stock (whence the Bahaman races
also were derived) which has invaded the
south island. Previously, the terminal portion
of the Tiburon Peninsula has received A.
distichus across the Golfe de la Gojiave,
and three subspecies had diflerc>ntiated
there. With the invasion of A. d. domini-
censis across the inter-island strait, this
subspecies came in contact with the eastern-
most (aurifcr) of the Tiburon races. The
subspecies in the Sierra de Baoruco
(favillarum) is considered a relatively re-
cent derivative from dominicensis. A. d.
tostus on Isla Catalina suggests that there
has been a westward shift in A. distichus
populations along the southeastern coast,
with the result that to.stus is, alone of the
four satellite island subspecies, unlike its
neighbor on the adjacent mainland. The
absence of A. disticlius from He de la
Gonave and the Peninsula de Barahona is
attributed to the inability of A. di.stichus to
compete with A. hrevirostris in arid areas
where the latter is well established.
LITERATURE CITED
Barbour, Thomas. 1937. Third list of Antil-
lean reptiles and amphibians. Bull. Mus.
Comp. Zool., 82(2): 77-166.
Clench, William J. 1938. Origin of the land
and freshwater mollusk fauna of the Bahamas,
with a hst of the species occurring on Cat
and Little San Salvador Islands. Bull. Mus.
Comp. Zool., 80(14): 481-541, 2 figs.. 3 pis.
Cochran, Doris M. 1941. The herpetology of
Hispaniola. Bull. U. S. Natl. Mus., 177:
i-vii, 1-398, 120 figs., 12 pis.
DuELLMAN, William E., and Alrert Schwartz.
1958. Amphibians and reptiles of southern
Florida. Bull. Florida State Mus., 3(5):
181-324, 28 figs.
Etheridge, Richard. 1966. Pleistocene lizards
from New Providence. Quart, [our. Florida
Acad. Sci., 1965, 28(4): 349-358.
King, Wayne, and Thomas Krakauer. 1966.
The exotic herpetofauna of southeast Florida.
Quart. Tour. Florida Acad. Sci., 29(2):
144-154.
Maerz, A., AND M. Rea Paul. 1950. A dic-
tionary of color. New York, McGraw-Hill
Book 'Co., pp. i-vii, 1-23, 137-208, 56 pis.
Mertens, Rohert. 1939. Herpetologische Erge-
bnisse einer Reise nach der Insel Hispaniola,
Westindien. Abh. Senckenberg. Naturf. Ces.,
449: 1-84, 10 pis.
Oliver, James A. 1948. The anoline lizards of
liimini, Bahamas. Amer. Mus. Novit., No.
i:i83: l-,36, 3 figs.
bA\D, A. Stanley. 1962. Notes on Hispaniolan
herpetology. 5. The natural histor>' of tlirec
sympatric species of Aiioli.s. Breviora, Mus.
Cuwp. Zool. No. 154: 1-15.
Anolis distichus ' Schwartz
309
Schwartz, Albert. 1967a. The Ameiva (La-
certilia, Teiidae) of Hispaniola. III. Ameiva
taeniitra Cope. Bull. Mus. Comp. Zool.,
135(6): 345-375, 2 figs.
. 1967b. The Leiocephahis ( Lacertilia,
Iguanidae) of Hispaniola. II. The Leio-
cephalus personatus complex. Tulane Stud.
Zool., 14(1): 1-53, 12 figs.
Schwartz, Albert, and Richard Thomas.
196.5. Subspeciation in Sphaerodactijhis copei.
Quart. Jour. Florida Acad. Sci., 1964, 27(4):
316-332, 1 pi.
Smith, Hobart iM., and Robert H. McCauley,
Jr. 1948. Another new anole from south
Florida. Proc. Biol. Soc. Washington, 61:
159-166.
SuTCLiFFE, Robert. 1952. Results of the Cat-
herwood-Chaplin West Indies Expedition,
1948. Amphibia and Reptilia. Notul. Nat.,
Acad. Nat. Sci. Philadelphia, No. 243: 1-8.
Thomas, Richard. In Press. The lumbricalis
group of West Indian Typhlops. Bull. Mus.
Comp. Zool.
Williams, Ernest E. 1961. Notes on His-
paniolan heriDctology. 3. The evolution and
relationships of the Anolis semilineatus group.
Breviora, Mus. Comp. Zool., No. 136: 1-8,
1 pi.
. 1962. Notes on Hispaniolan herpetology.
6. The giant anoles. Breviora, Mus. Comp.
Zool, No. 155: 1-15, 1 fig., 1 pi.
(Received 19 June 1967.)
Wmmm:m-
Muiletin OF THE
Museum of
Comparative
Zoology
Ammonoids of the Late Scythian
(Lower Triassic)
BERNHARD KUMMEL
HARVARD UNIVERSITY VOLUME 137, NUMBER 3
CAMBRIDGE, MASSACHUSETTS, U.S.A. APRIL 28, 1969
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Memoirs 1864-1938
JoHNSONLV, Department of Mollusks, 1941-
OccASioNAL Papers on Mollusks, 1945-
Other Publications.
Bigelow, H. B. and W. C. Schroeder, 1953. Fishes of the Gulf of Maine.
Reprint, $6.50 cloth.
Brues, C. T., A. L. Melander, and F. M. Carpenter, 1954. Classification of In-
sects. $9.00 cloth.
Creighton, W. S., 1950. The Ants of North America. Reprint, $10.00 cloth.
Lyman, C. P. and A. R. Dawe (eds.), 1960. Symposium on Natural Mam-
malian Hibernation. $3.00 paper, $4.50 cloth.
Peters' Check-list of Birds of the World, vols. 2-7, 9, 10, 12, 15. (Price Hst on
request. )
Turner, R. D., 1966. A Survey and Illustrated Catalogue of the Teredinidae
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Whittington, H. B. and W. D. I. Rolfe (eds.), 1963. Phylogeny and Evolution
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© The President and Fellows of Harvard College 1969.
AMMONOIDS OF THE LATE SCYTHIAN (LOWER TRIASSIC)
BERNHARD KUMMEL
CONTENTS
Introduction 312
Plan of Study 313
Acknowledgment 316
Lower Triassic Chronology 316
Summary of Prolitingaritcs Zone Ammonoids _ 324
Paleogeographic Implications of ? rohungarites
Zone Ammonoids 336
Summary of CohmiI)ites Zone Ammonoids — - 336
Stratigraphy and Faunas of the Late Scythian . 337
Albania 338
Chios .___._ 339
Werfen Formation 342
Mangyshlak Peninsula 345
Afghanistan 347
Salt Range, West Pakistan 347
India 348
Timor 349
New Zealand 351
China 351
Japan 353
Primorye Region .___ 353
Northeastern Siberia 355
Spitsbergen 357
Ellesmere Island 357
British Columbia 358
Tobin Formation, Nevada 358
Confusion Range, Utah 358
Thaynes Formation, Southeast Idaho 359
Systematic Paleontology 360
Sageceratidae 360
Pseudosagcceras 360
Cordillcrites 364
Dieneroceratidae 367
Dieneroceras ____ 367
Subvishniiites — - 373
Hemilecanites 374
Xenoceltitidae 375
Xenoceltites 375
Prefloiianites 379
Paranoritidae .._. 383
Pseudaspidites 383
Proptychitidae 384
Proptijchitoides 384
Procamites 391
Paranannitidae 397
Arnautoceltites 397
Prosphingitcs 403
VickuJdcrites 408
Zenoites 410
Isctditoidcs 411
Chiutitcs 419
Czekaiiowskites 420
Popovites 421
Monocanthites 422
Tunglanites 422
Cohindntes 424
Sid)cohiiuhites 427
ParadiuarUes 437
Pscuduceltites 437
Procohaubites 441
Pienkites 441
Protropites 444
Chioceras 445
Arianites 446
Meropella 447
Epiceltites 447
Ussuriidae 448
Panissuria 448
Hedenstroemiidae 448
Metahedemtroemia 448
Beatites 449
Lanccolites 450
Meekoceratidae 450
Svalbardiccras 450
Stacheites 455
Dagnoceras 457
Metadagnoceras 460
Balkanites 465
NordopJuceras 465
Psetidokijnwtites 475
Arctomeekoceras 476
Boreomeekoceras 476
Arctotirolites 477
Noritidae 477
Albmiites 477
Prionitidae 482
Hemiprionites 482
Bull. Mus. Conip. Zool., 137(3): 311-702, April, 1969 311
312 BuUetin Museum of Comparative Zoology, Vol. 137, No. 3
Sibiritidae 483
Sibirites - 483
Keyserlingites 485
Olenckites 488
Eukashmirites 490
Anakashmirites 490
Tirolitidae 49 1
TiroUtes 49 1
Diaplococeras 503
Bittnerites 504
Doricranites _- 505
Dinaritidae 506
Dinarites 506
HoJolobus 511
Pseudodinorites - 511
Hellenitidae 511
Hellenites 511
Beyrichitidae 516
Beyrichites 516
Gymnitidae 517
Eogymnites 517
Hungaritidae 517
Prohiingcirites 517
Dolmatites 522
Ussuritidae 524
Eophyllites — - 524
Palaeophyllites 527
Uss'urites - 528
Leiophyllites 531
References 536
Index 542
ABSTRACT
An evaluation of all taxa of ammonoids of
the Late Scythian Frohun^aritcs Zone shows
the fauna to consist of 65 genera with 154
species. Of the total number of genera, 42
are confined to this zone, 20 genera are
known to range up from the preceding
Cohimhites and Oucnitcs Zones, one genus
is known from both older and younger
horizons, and two genera are also present
in the overlying Anisian. There are 24
genera endemic to the Tethyan region.
The western Pacific, eastern Pacific, and
Arctic faunas have only two endemic genera
each. Indexes of faunal similarity between
the major faunas of the Prolnin^a rites Zone
are presented. The largest number of
genera of this zone is found in the Tethyan
region. The Arctic region has only approxi-
mately 30 percent as many and the western
and eastern areas an intermediate number.
It is suggested that this is a true faunal
gradient and could well reflect a climatic
pattern.
The Cohimhites fauna lying immediately
below that of the Prohuniiarites Zone is well
known from southeast Idaho and Siberia,
and appears to be present in Arctic Canada.
This fauna is, as yet, not known in the
Tethyan region. The Cohimhites fauna is
closely related to that of the Prohun^aritc.s
Zone.
INTRODUCTION
We are well along in the second century
of the systematic study of invertebrate
paleontology and stratigraphy. During the
early phases of this period, published con-
tributions consisted mainly of monographs
describing the whole range of fossil forms,
then known, for wide geographic regions.
This initial pattern of publication was fol-
lowed by a phase consisting mainly of
monographs of faunas of specific strati-
graphic units. This second phase gradually
merged into a third where specific biologi-
cal groups, with or without time restric-
tions, became the focal point of study. We
are now in a fourth phase in the history of
invertebrate paleontology where it is pos-
sible to attempt a total synthesis of animal
groups on a world scale for relatixely short
miits of time. The development of this
latest phase in paleontology comes at a
most appropriate time. Whereas there are
still many areas that have received little
or no intensive geological study, we do
have a considerable fund of data from all
parts of the world. In recent years there
has developed a more intense interest in
the problems of animal e\-olution and
zoogeography. However, towards these
goals invertebrate paleontology can make
significant contributions onb if a sound
taxonomic base is present.
We are now in the midst of a great
resurgence of interest in the question of
permanence or non-permanence of the con-
tinents and oceanic basins, brought on pri-
mariU b>' tiie application of paleomagnetic
stndies. Faleontologv must and can jilay
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 313
an important role in this fundamental re- first attempt to establish a series, stage,
evalnation of the earth's evokition. The and zonal scheme for the marine Triassic
importance of paleontological data to these was by Mojsisovics, Waagen, and Diener
problems is commensurate with the quality ( 1895 ) . It was in this paper that the term
and quantity of data available. The vast Scythian^ was first introduced for the
majority of the paleontological data ac- Lower Triassic, and the sequence of the
cumulated over the past century consists Ceratite beds in the Salt Range of West
of faunal studies in which genera and Pakistan was selected as the type. At this
species are treated as segregated elements time there had already appeared or were
with little or no relevance to other con- in an advanced stage of preparation Moj-
temporaneous faunas. Syntheses which at- sisovics' monumental works on the Alpine
tempt to evaluate and re-appraise all Triassic (Mojsisovics, 1873-1902, 1882),
generic and specific taxa on a global scale Waagen's monograph of the Salt Range
are not many. It is such syntheses, how- Ceratite bed fauna (Waagen, 1895), the
ever, that will contribute most to the stoiy faunas of the Lower Triassic of the Hima-
of the earth's biological and physical his- layas (Diener, 1897), the fauna of the
tory. L'ssuri Bay (Primorye) region, and the
In an attempt to achieve a limited goal, Olenek fauna of northern Siberia ( Moj-
I have set up a program to develop a sisovics, 1886, 1888). Thus, by the turn of
three-dimensional picture of a stratigraphic the century, it was fairly well established
stage based primarily on ammonites. For that marine formations of Lower Triassic
this purpose I have selected the Scythian age existed in the Tethyan realm, in the
Stage ( Lower Triassic ) . The first require- circum-Pacific region and in the Arctic
ment of this program is the establishment region.
of a uniform taxonomic procedure based Throughout the early period of study of
on consideration of all available data. This Lower Triassic ammonoids, the primary
contribution is devoted to consideration of guiding philosophy in taxonomy was typol-
the two upper zones of the Scythian. In ogy. In fact, at that time there was no
the chapter on systematic paleontology all alternative — each fauna was new, small,
taxa of Upper Scythian ammonoids are con- and unique; the correlation of faunas was
sidered. These data provide the basis for tenuous at best; and, finally, nomenclatural
the summary chapters concerning popula- and species concepts were primitive or
tion structures, geographic distribution of absent. The typological approach did
species and genera, and other problems. facilitate description and comparison, and
it led, in time, to a more complete docu-
PLAN OF STUDY mentation. One needs only to examine
The first described species of Lower many original collections upon which the
Triassic (Scythian) ammonites was Am- early classic studies were made, to have
monites hogdoanus von Buch (1831). The data on their collecting, and to know the
first centurv of study of Scvthian am- prevailing geological and zoological phi-
monoids was focused primarilv on docu- losophies, to appreciate why the faunas
mentation of each newly discovered fauna ^ere treated as they were. Whereas one
and on attempts to construct a zonal frame- can understand and appreciate the guiding
work. It was recognized very early in the philosophies behind these early studies, it
development of our geologic time scale that is q"ite clear that the resulting conclusions
central and northern Europe were not ap- are not satisfactory for many purposes.
propriate areas to establish a chronological n^i c^ m ■ ■ ^ j .. i
). ^ 1 J T™ Scythians are an ancient nomadic tribe
framework for the Triassic system based ^j^^^ occupied the area north of the Caspian Sea,
on a succession of marine faunas. The the region which yielded Ammonites hogdoanus.
314 Bulletin Museum of Comparative Zoologij, Vol. 137, No. 3
Table 1. Summary of numbers of species and specimens described in sixteen major publica-
tions ON Lower Triassic (Scythian) ammonoids.
Percent
Total species described
Species described on basis
Species described on basis
Species described on basis
Species described on basis
Species descri]:)ed on basis
Species described on basis
Species described on basis
Species described on basis
of 1 specimen
of 2 specimens
of 3 specimens
of 4 specimens
of 5 specimens
of 6-10 specimens
of 11-20 specimens
of >20 specimens
1194
548
237
89
63
38
104
57
55
46
20
7.5
5
3
8.9
5
4.6
An analysis of the taxonomic treatment in
16 major publications on Lower Triassic
( Scythian ) ammonites is summarized in
Table 1. These publications contain the
description or documentation of 1,194
species of which .548 (46 percent) were
based on a single specimen. In fact, only
216 species ( 18.5 percent ) were based on
6 or more specimens. When one looks at
only the new species described in these
16 monographs one gets a clearer insight
into early taxonomic procedures. These
data are summarized in Table 2. There
are 668 new species of which 308 (46
percent) are based on a single specimen.
Only 100 of these new species ( 15 percent)
were based on 6 or more specimens.
Although at any one period there were
usually only from one to four active workers
on Lower Triassic ammonoids, a vast
amount of geologic and paleontologic data
was accumulated during a century or so of
study. I believe we now are in a position
to approach the problem as a whole and
not in terms of isolated parts, as has been
the pattern in the past. The first step in
such a synthesis is a thorough restudy of
all available type collections of the early
classic studies. I have had the opportunity
of studying the following faunas: the
Werfen fauna described ])y Kittl (1903),
deposited in the Natural History Museum,
Vienna; the Albanian Suhcohtmhitc.y fauna
described by Arthaber (1908, 1911), de-
posited in the Paleontological Institute, Uni-
versity of Vienna; the Suhcoiinnhilc.s fauna
of Chios described by Renz and Renz
(1948), deposited in the Natural History
Museum, Basel; the Lower Triassic faunas
of Timor described by Welter ( 1922 ) , de-
posited in the Paleontological Institute,
Bonn, and at Delft; various collections from
the U.S.S.R. described by Mojsisovics (1882,
1886), Popov (1961), and Kiparisova
(1961); the Kashmir fauna, described by
Diener (1913); the Himalayan fauna de-
scribed by Diener ( 1897 ) and Krafft and
Diener (1909), deposited in the Geologi-
cal Survey of India, Calcutta; the fauna of
the Lower Triassic of Spitsbergen deposited
in the Paleontological Institute, Stockholm,
and the Arctic Institute, Norway; the fauna
of Arctic islands of Canada and British
Columbia described by Tozer (1961a, 1965a,
b ) and deposited in the Geological Survey
of Canada, Ottawa; the faunas of the west-
ern United States described by J. P. Smith
(1932), deposited in the U.S. National
Museum, Washington; and finally the large
collections in the British Museum (Natural
History). In addition I have had before
me new collections made by myself in
Nevada, Utah, Idaho, Madagascar, Afghani-
stan, and the Salt Range of West Pakistan.
The only large fauna of Lower Triassic
ammonites that I ha\e not personalh- ex-
amined is that from south China, described
byChao (1950, 1959).
The examination of these many faunas al-
lowed a direct comparison of related taxa.
Many siH'cimeus, several of them t\pes,
had been inade(iuatc4y, and in sonK> cases
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 315
Table 2. Summary data on numbers of specimens on which new species were established in
the same sixteen major publications on Lower Triassic ( Scythian ) ammonoids as tabulated in
Table 1.
Percent
Total new species
Species estalilished on basis of 1 specimen
Species established on basis of 2 specimens
Species established on basis of 3 specimens
Species established on basis of 4 specimens
Species established on basis of 5 specimens
Species established on basis of 6-10 specimens
Species established on basis of 11-20 specimens
Species established on basis of >20 specimens
668
308
137
55
42
25
48
28
24
46
20
8
6
4
7
4
4
misleadingly described or illustrated. The on these late Scythian ammonoids gave
numbers of specimens in these collections more than token tribute to the range of
vary greatly. In some, e.g. the Salt Range morphological variation in their species,
and Himalayan collections, only the figured Even though many of the faunas from single
specimens of the original monographs are horizons and localities had yielded an
available, whereas the Subcolumbitcs fauna abundance of certain taxa, emphasis was
of Chios, described by Renz and Renz placed on "differences" rather than simi-
( 1948 ) and preserved in the Natural His- larities. In many of these cases it can be
tory Museum, Basel, is nearly twice as large clearly demonstrated that within the fauna,
as indicated in the original monograph. one is dealing with a highly variable single
In approaching the problem of evaluation species complex rather than with a complex
and synthesis of all these faunas, I elected of several species of a genus. In most such
to study them zone by zone. In this paper cases species were differentiated on the
I am treating the two upper zones of the
Scythian, the Proluing^a rites Zone and the
Cohimhites Zone. Rather than focus my
studies on a fauna by fauna analysis, I
basis of differing shell parameters, that is,
degree of whorl compression, involution,
suture, etc. Plots of measurements of large
numbers of specimens often show the
chose to make the genus my unit of study, particular distinguishing parameters to be
For each genus of late Scythian ammonoids, part of a gradational series,
an analysis was made of all species and The total numbers of specimens in nine
specimens that had been or should be as- faunas of the Frohungaritcs Zone and the
signed to it. For many of the genera there numbers of specimens on which the species
were usually one or more faunas which are based are summarized in Table 3. The
contained an appreciable number of speci- numbers of species for each of these faunas
mens of a species, thereby yielding data are based on the results presented in the
on intraspecific variation. The insight systematic portion of this paper. The num-
gained from studies of species represented ber of species based on 20 or more speci-
by large numbers of specimens was of mens is relatively small. At the same time,
great help in the analysis of species repre- these relatively few species include any-
sented by one or very few specimens from where from 36 to 90 percent of the total
other localities. In essence, the analysis of number of specimens in these faunas,
a population of many specimens provided It is thus quite apparent that only about
the framework within \\'hich isolated speci-
mens could be more logically interpreted.
Essentially, none of the previous studies
25 percent of the total number of species
for the ProJiungarites Zone have been es-
tablished on the basis of samples of reason-
316 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Table 3. Summary data on numbers of specimens and the numbers of specimens per species in
NINE major faunas OF THE PrOHUNGARITES ZoNE.
Albania Chios
China Primor\e B. C. Tobiu Timor
Elles- Ham-
mere mond
Is. Cr.
Total specimens in collection
No. of species in fauna
No. of species with 50 specimens
Percentage of total specimens
No. of species with 20-49 specimens
Percentage of total specimens
No. of species with 10-19 specimens
Percentage of total specimens
No. of species with 5-9 specimens
Percentage of total specimens
No. of species with 1—4 specimens
Percentage of total specimens
730
1900
97
97
41
186
24
23
296
32
41
21
17
11
7
11
5
11
5
9
0
0
0
2
0
0
1
53.4
49.0
0
0
0
53.7
0
0
67.5
6
9
0
1
0
2
0
0
2
27.8
12.5
0
36.2
0
34.7
0
0
23.6
7
3
2
0
1
1
0
1
0
15.0
2.4
26.7
0
36.5
5.4
0
47.8
0
1
7
6
6
2
1
1
1
2
0.7
2.6
43.3
41.2
31.7
3.7
25
26.1
6.0
13
13
13
10
8
1
10
3
6
3
1.5
30
22.6
31.7
2.2
75
26.1
2.7
able size. Note the difference in these
figures from those given in Tables 1 and 2.
Species based on few specimens are diffi-
cult to evaluate. If two or more species are
based on few specimens from the same
horizon and locality and are differentiated
on criteria known to be highly variable in
related groups, I have tended to synonymize
them. The prime assumption is that a
larger sample would "fill in" the morpho-
logical gap with gradational forms. How-
ever, species based on few specimens from
widely separated localities are generally
kept separate even though there is a strong
indication that they may be identical with
other named species.
ACKNOWLEDGMENTS
It is a great pleasure to acknowledge the
hospitality and help of the following in-
dividuals who allowed me to study speci-
mens under their charge: Dr. E. Gasche,
Natural History Museum, Basel; Dr. F.
Steininger, Paleontological Institute, Vienna;
Professor O. II. Schindewolf, Tiibingen
University; Professor H. K. Erben, Bonn
University; Professor 11. J. MacCillavry,
Geological Institute, Amsterdam; Mr. M.
V. A. Sastry, Geological Svu'vey of India;
Dr. M. K. Howarth, British Museum (Nat-
ural History), London; Dr. E. T. Tozer,
Geological Survey of Canada; Dr. N. J.
Silberling, Stanford University; Dr. L. D.
Kiparisova, Geological Institute, Lenin-
grad; Dr. Yu. N. Popov, Arctic Institute,
Leningrad; Dr. Yuji Bando, Kagwawa Uni-
versity, Japan; Dr. Keiji Nakazawa, Uni-
versity of Kyoto, Japan. I have benefited
greatly from fossil exchanges with Dr. Yu.
D. Zakharov, Far East Geological Institute,
Vladivostok, Dr. Yu. N. Popov, Dr. E. T.
Tozer, Dr. N. J. Silberling, Dr. H. K. Erben,
and Dr. E. Gasche.
Throughout the laboratory phases of this
study, I was assisted by Miss Victoria
Kohler and I am deeply indebted for the
excellence and competence of her assis-
tance. It is a pleasure to acknowledge the
support of the following research grants
from the National Science Foundation
which made the project possible: NSF-
G19066, GB-2354, GB-5109X. My field
work in Afghanistan and Japan was sup-
ported by a grant from the Shaler Fund of
Harvard University.
LOWER TRIASSIC CHRONOLOGY
The name Scythian was lirst introduced
as a series name for the Lower Triassic by
Waagen and Diencr in Mojsisovics, Waa-
gen, and Diener (LS95); the sequence for
thc> Salt Range of West Pakistan was se-
lected as the type, with the addition of the
Otoccras wooduardi Zone of the Himalayas
at the base of th(> sequence (Table 4).
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 317
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318 Bidkiiu Museum of Comparative Zoology, Vol. 137, No. 3
Table 5. Zonal scheme for the Lower Triassic (Scythian) proposed by Spath (1930'
Ages
Zones (India)
Some Equivalents
Stephanitan
Columbitan
Upper-
I. Owenitan
superbiis?
EO-TRIAS
flemingiam
volutu.^
Flemingitan
fallax
Lower^
Gyronitan
rotundatus
radiosiis
Otoceratan
woodwardi
Arctoceras beds, Spitsbergen; OJenekites beds, N.
Siberia.
Columbites beds, Albania; Anasibirites beds, Spits-
bergen, Utah, Timor; Tirolifes beds, Werfen.
Meekocenis beds, California.
Meekoceras beds, Timor;
Hedenstroemia beds, Himalayas (lower part).
"Ophiceras" beds, Timor.
"Meekoceras" beds, Himalayas.
Proptycliites beds, Primorye.
ProptycJiites beds, Greenland.
Otoceras beds, Greenland.
The Scythian was thus proposed as a series
division which included seven ammonoid
zones, but the zonal context of the Scythian
has since undergone considerable change.
What is so puzzling is that Waagen and
Diener separated from the Scythian the
fauna of the Upper Ceratite limestone
(zone of Steplioiiitcs superbus) of the Salt
Range, establishing a new stage (Hy-
daspian) of the Middle Triassic. This was
done on the basis of what they interpreted
as a striking change between the faunas of
the Upper Ceratite limestone and the un-
derlying members of the Ceratite beds.
The basis of Waagen and Diener's correla-
tion of these divisions was summarized as
follows: "Es kann wohl nicht in Zweifel
gestellt werden, dass die tieferen Abtei-
lungen der Ceratiten-Schichten als zeitliche
Aquivalente jener Bildungen anzusehen
seien, welche in Mittel-Europa den Namen
Buntsandstein' tragen. Anderseits jedoch
haben wir gesehen, dass angefangen von
den tiefsten Ablagerungen des Lower
Ceratite Limestone bis hinauf zur oberen
Grenze der Ceratite Sandstones eine
kontinuierliche Serie der Cephalopoden-
Faunen angetroffen wird, dass aber vom
Ceratite Sandstone zum Upper Ceratite
Limestone ein betrachtlicher Wechsel sich
einstellt. Hier muss also eine Formations-
grenze durchgezogen werden, und diese
Grenze kann nur jene zwischen der Sky-
thischen und Dinarischen Serie sein.
Wir glauben daher die Oberen Ceratiten-
Kalke an die Basis der Dinarischen Serie
stellen zu sollen und betrachten sie in-
nerhalb der letzteren als den Typus einer
besonderen Stufe. Die Bezeichnung der
letzteren als Hydaspische Stufe ist dem
alten Namen des die Salt Range an ihrer
Ostseite umfliessenden Ihelum ( Hydaspes )
entnommen" ( Mojsisovics, Waagen, and
Diener, 1895: 1291 ) . Tliis scheme of classifi-
cation was completely accepted bv Smith
(1896, 1901, 1904).
Noetling (1901, in Freeh 1905) interpreted
the fossiliferous Triassic beds of the Salt
Range (through the zone of Stcpluinifes
superbus of the Upper Ceratite limestone)
as including a complete succession of faunal
zones for the Scythian. Tliis view was held
by Diener (1912:256), Welter (1922:92),
and others.
This view on the age span of the Ceratite
beds and the scope of the Scythian stage
generally prevailed until Spath (1930:76)
published a preliminary scheme for the
subdivisions of the Scythian (Table 5). This
is an extremely interesting modification of
the earlier attempt to establish subdivisions
of the Scythian. This scheme differed in
detail from the proposals and conclusions
of previous authors, but it agreed with
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 319
Table 6. Zonal scheme for the upper half of the Lower Triassic (Scythian) proposed by
Spath (1934).
Divisions
Zones
Equivalents
.2
!-l
H
I
o
W
i-i
a
Q
Prohimgaritan
(Olenikitan ?)
Columbitan
Owenitan
I
f Columbites
\ Tirolites — .
' Anasihirites
Owenites
Pseudosageceras
Upper Arctoceras beds, Spitsbergen.
Olenck beds, Siberia ( partim ) .
P. middlemissi l^eds, Kashmir.
Suhcohimbites beds, Albania, Timor.
Cohivdrites beds, Idaho.
Tirolites beds, Alps, etc., Idaho.
Amisibirites beds, Timor, Utah.
Chocolate Limestone, Byans?
Upper Ceratite Limestone, Salt Range.
Kasliiniiites beds, Kashmir, Timor.
Meekoceras beds, Timor, Idaho, California.
Timor, Himalayas, W. America.
several of these in having a single zone
(Stephanites siiperbtis) to represent most
of the upper Scythian. At the same time,
within the late Scythian, Spath (1930:76)
I'ecognized three ages: the Owenitan,
Cohimbitan, and Stephanitan.
Spath's proposal was admittedly a tenta-
tive scheme in need of further analysis.
What was critically lacking, especially for
the upper part of the scale, were sufficient
stratigraphic sections with these faunal
zones in sequence. Spath was cognizant
that Smith ( 1904 ) , Hyatt and Smith ( 1905 ) ,
and Smith (1914) had reported Scythian
ammonoid faunas in sequence in south-
eastern Idaho, but published data were
very limited. Smith's ( 1932 ) monograph
on the Lower Triassic ammonoids of North
America provided a comprehensive treat-
ment of these ammonoid faunas. In south-
eastern Idaho, Smith encountered within
the Thaynes Formation a sequence of three
ammonoid zones which formed the basis
for the upper part of his chronologic
scheme for the Lower Triassic:
Columbites Zone
Tirolites Zone
Meekoceras Zone<
' Anasihirites Subzone
Owenites Subzone
Pseudosageceras muJti-
lobatwn Subzone
As my own monograph is devoted to the
ammonoid faunas of the upper Scythian,
no further discussion is needed here in re-
gard to the zones of lower Scythian. A
comprehensive discussion of the Meekoceras
Zone has been published by Kummel and
Steele (1962), and Kummel and Erben
(1968).
Smith considered his Anasihirites Sub-
zone to be equivalent to the fauna of the
Upper Ceratite limestone of the Salt Range.
Smith furthenuore considered the specimen
( presumably from the topmost limestone of
the Dolomite beds) that Waagen (1895:
130, pi. 21, figs, la-c) described as Pseiid-
harpoceras spiniger as allied to his own
species, Tseiidharpoceras idahoense, from
the Cohimhites fauna of southeast Idaho,
and thus of Columbites Zone age. This
scheme enlarged the scope of the Scythian.
Smith's zonal scheme and analysis of the
Scythian came under the searching pen of
L. F. Spath (1933, 1934). Though Spath
was rather caustic in his remarks on Smith's
zonal scheme, the differences between
these two authorities were not that exten-
sive. Spath presented a new scheme of
zones and correlations in 1934; that for the
upper half of the Scythian is shown on
Table 6. It can readily be seen that this
zonal scheme differs from that proposed by
Smith ( 1932 ) mainly in the introduction of
a division of one or more zones above the
Columbitan. In this conclusion Spath was
guided mainly by "intuition," which was
320
Bulletin Museum of Comparative Zoology, Vol. 137, No.
supported by the biologic character of the
faunas. Stratigraphic data on the faunas
he assigned to his Prohungaritan division
were either completely lacking or very
ambiguous. In his conclusions, explaining
the significance of the Prohungaritan divi-
sion, Spath (1934:34) states: "I am merely
relying on the obvious differences between
the lowest Anisian and the highest Scythian
faunas so far known, and the only diffi-
culty is to find a name for this time inter-
val that will prove sufficiently accurate to
serve for a label, even if it is not the best
that could ultimately be proposed." Spath's
conclusions regarding additional zones
above the Columbites Zone were verified
with the discovery of a Frohun^aritcs fauna
in the Thaynes Formation of southeast
Idaho, 1,000 feet above the Columbites
fauna ( Kummel, 1954).
In recent years a number of additional
localities have been studied which have
yielded sequences of ammonoid faunas in
upper Scythian formations. Aside from
southeastern Idaho, good stratigraphic sec-
tions with fossil faunas are now known for
Ellesmere Island (Tozer, 1961a, 1965a),
the Primorye Region around Vladivostok
(Kiparisova, 1961; Zakharov, 1966), Kwangsi,
China (Chao, 1959), the Salt Range of
West Pakistan (Kummel, 1966), and Afghani-
stan (Kummel, 1968). In southeast Idaho,
Utah, Ellesmere Island, Primorye Region,
and northern Siberia, the upper Scythian
comprises a sequence of three distinctive
faunas: the Oiccnitcs (or Mcckocems)
fauna, the Columbites fauna, and at top
the Prohungarites fauna or its equivalents.
The Columbites fauna is not known from
south China, West Pakistan, or Afghanistan.
Other areas that have yielded faunas of
ProJiuugarifcs Zone age but are isolated in
having no other Scythian ammonoid faunas
above or below are those of the Werfcn
Formation of southeast Europe, Albania,
Chios, Mangyshlak Peninsula, Kashmir,
Timor, New Zealand, Japan, and Nevada
( Tobin Formation ) .
Even though there has been a vast in-
crease in the amount of data on Scythian
strata and faunas over the past decade,
considerable differences of interpretation
on the correlation of late Scythian zones
persist. Kiparisova and Popov ( 1956 ) re-
jected Spath's (1934) Prohungaritan divi-
sion and accepted Columbitan as the latest
Scythian biostratigraphic unit. They con-
sidered this division as including the
Olenekites fauna of northern Siberia, the
SuJ)colum])ites fauna of the Primorye Re-
gion, Albania, and Chios, and the Arctoceras
fauna of Spitsbergen. At a later date these
two authors (Kiparisova and Popov, 1961)
modified their late Scythian chronology
and accepted a Prohungarites Zone as the
latest Scythian zone, overlying a Columbites
Zone. Their Prohungarites Zone was then
stated to include the following faunas:
Prohungarites middlemissii of Kashmir, the
ProJiungaritcs fauna of southeast Idaho, the
Olenekites fauna of northern Siberia, and
Subcolumbites fauna of the Primorye Re-
gion, Albania and Chios, the Procarnites-
Leiophijllites fauna of Kwangsi, China, and
the Stocheites Zone of Astakhova (1962)
of the Mangyshlak Peninsula. Later, these
authors introduced a further revision
(Kiparisova and Popov, 1964) in the cor-
relation of late Scythian faunas. They now
considered the Olenekites fauna of northern
Siberia, as well as the Subcolumbites fauna
of the Primorye Region, as being of Co-
lumbites Zone age. The correlations of the
ProJiungaritcs Zone included a new but
undescribcd ProJtungarites element from
northern Siberia, a questionable Prohun-
garites from the Primorye Region, the
Kashmir Prohungarites middlemissii, the
Prohungarites fauna of southeast Idaho,
and the Procarnites'Lciophijllitcs faunas of
K\\'angsi, China.
Our knowledge of the Scythian of China
greatly increased with the appearance of a
large monograph by Chao ( 1959 ) on faunas
from Kwangsi. Chao followed the lead of
Kiparisova and Popov (1956) in using the
Columbitan division to encompass all of
the late Scythian. P"or Kwangsi Province,
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 321
Chao recognized three zones within his
Cokimbitan division, namely:
Trocarnites-LeiophijUitcs Zone
Cohnnhifes costofus Zone
TiroJites darwini Zone
The paleontology and stratigraphy of these
Kwangsi faunas are discussed in detail on
page 351. It will suffice here to summarize
my conclusions. The Tirolitcs darwini
Zone was recognized from a single, frag-
mentary specimen from an isolated horizon.
This specimen is not well enough preserved
to be identified. The name-giving species
of the second zone, Cohimhitcs costahis,
is a synonym of Prenkites timorensis. The
fauna listed as comprising the Procarnites-
LeiophyUites Zone was derived from loose
blocks. The assemblage of species and
genera which were assigned to the Co-
himhifes cosfafiis Zone and the Procarniics-
LeiophijUites Zone are such that one can
only conclude they are part of a single
zone. Recent contributions by Tozer (1961a,
1965a, b ) have sho\^'n that British Columbia
and the Arctic islands have faunal se-
quences comparable to those of southeast
Idaho.
Late Scythian faunas lying in strati-
graphic position above the mid-Scythian
Owenites Zone are now knov^m from nine
locahties. In Afghanistan a Subcolumhitcs
fauna occurs immediately above an Oucnites
fauna ( Kummel, 1968 ) and the same situa-
tion is present in the Salt Range and
Surghar Range of West Pakistan ( Kummel,
1966) and in Kwangsi Province of south
China (Chao, 1959). In the Primorye Re-
gion the Sub cohimhitcs fauna is strati-
graphically above a Cohimhitcs fauna
which in turn lies above an Owenites fauna.
In northern Siberia the Olcnekitcs fauna
occurs above a Cohimhitcs fauna which,
in places, lies above an Owenites fauna.
The same is tiiie for Ellesmere Island. In
southeastern Idaho the uppermost Scythian
is marked by a Prohungarites fauna, which,
in turn, is underlain by a Cohimhitcs fauna
and this by an Owenites fauna. The
Tirolitcs Zone of Smith (1932) is of only
local importance and not equivalent to the
fauna of the Werfen Formation (see p.
342).
We thus have a sequence of two faunas
(Cohimhitcs and Prohungarites) above the
mid-Scythian Owenites Zone in southeast
Idaho, Ellesmere Island, and the Primorye
Region and in northern Siberia. However,
in China, West Pakistan, and Afghanistan,
wherever we have stratigraphic control, the
late Scythian Sitbcolumbites or Prohun-
garites fauna lies directly above Owenites
Zone faunas. This raises a question as to
the relationships of the Cohimhites fauna
and its independence as a zonal entity.
The Cohimhitcs fauna and its equivalents
are known from southeast Idaho, the
Primorye Region and northern Siberia.
There are 21 genera of ammonoids in one
or another of these three main faunas. The
largest number of genera (15) comprise
the Cohimhitcs fauna of southeast Idaho.
The Primorye Region contains eight genera
and northern Siberia six genera at this
horizon. Of the total of 21 genera, only
two are restricted to this horizon. Two
genera are also present in earlier horizons,
seven genera are present in both older
and younger horizons, and ten genera are
also present in younger horizons. Tliere
are no species in common in the faunas
assigned to the Cohimhites Zone and those
assigned to the Prohungarites Zone, but
the ten genera in common include such
late Scythian members as Procohimhites,
Pseudoccltitcs, Svalhardiceras, Metadagno-
ceras, Nordophiceras, Keijserhngites, Olcne-
kitcs, HcUcnites, DaJmatites, and Ussuritcs.
The faunas assigned to the Cohimhitcs
Zone clearly are intimately related to those
assigned to the Prohungarites Zone and are
quite distinct from the faunas of the
Owenites Zone. Zakharov (1966) expressed
this relationship by treating the two faunas
as subzones of a single late Scythian Zone.
There is no question but that this sugges-
tion has merit. However, at a species level,
these two successive faunas are very dis-
322 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
tinct. The bothersome factor is the absence
of the Columhites Zone in Tethys. This,
however, may be due more to preservation
than to anything else. Only eight areas
within Tethys have yielded late Scythian
faunas. The Suhcolumbites faunas of
Albania and Chios are isolated, without
other Scythian faunas above or below. The
Werfen fauna and that from the Mangyshlak
Peninsula are from semi-isolated embayed
regions along the northern margin of
Tethys and contain a large percentage of
endemic genera and species and are the
youngest Scythian ammonoid faunas in
their respective areas. The Kashmir record
is based on specimens found as float. There
are thus only two areas within Tethys
where a sequence of two ammonoid zones,
Proluingaritcs over Oicenites, is known.
These occur at Kotal-e-Tera, Afghanistan,
and in the Salt Range and Surghar Range
of West Pakistan. At Kotal-e-Tera the
Scythian comprises approximately 90 feet
of liinestone and dolomite of which 75 feet
includes an Owenites fauna and the upper
15 feet contains a Prohungarites Zone fauna.
Ammonoids are abundant and fairly well
preserved in the Owenites Zone, but the
fossils from the Frohuno^aritcs Zone are
neither common nor \\e\\ preserved. In the
Salt Range and Surghar Range of West
Pakistan the Prohungarites Zone is thought
to comprise all of the Narmia Member of
the Mianwali Formation ( Kummel, 1966).
Fossils in the Narmia Member are very
scarce and not well preserved. The lowest
units of the Narmia Member, which in-
cludes the Bivalve Limestone of Waagen
(1895), contain Nordophiceras planorbis
(Waagen) and Xenoceltites sinuatus (Waa-
gen). These two species could possibly
be of Columhites Zone age, but because
of scarcity and generally poor preservation
of the Narmia Member fossils one cannot
be sure.
A new area of Permian and Triassic out-
crops has been reported by Sokolov and
Shah (1965) around Ghazaband Pass,
Quetta District, West Pakistan. The pres-
ence of the Scythian in this region is based
on a single specimen identified by Mr. A.
N. Fatmi of the Geological Survey of
Pakistan as Columhites sp. The fomiation
\\'hich yielded this specimen comprises
300-500 meters of shale with thin interbeds
of limestone. It is unfortunate that this
specimen has neither been described nor
illustrated.
The absence of a Columhites Zone fauna
in Kwangsi, China, may be due to facies
differences or possibly to insufficient field
studies. There is obviously a great need
for more data, but on the basis of the
picture developed here, I advocate con-
sidering the Columhites and Prohungarites
Zones as closely related but distinct; the
absence of the Columhites fauna in Tethys
and south China is most likely due to col-
lection failure and or adverse facies for
preservation.
The nomenclatin-e of these late Scythian
zones presents problems. The Columhites
Zone seems fairly well established. The
name-giving species, Columhites parisianus,
is present at the type locality in Paris
Canyon, southeast Idaho, in the Primorye
Region, and possibly in Arctic Canada.
The presence of Columhites parisianus in
the Primorye Region makes it difficult to
understand why Zakharov (1966) selected
a different species to identify this zone.
Columhites is not present in northern
Siberia, but the genera and species of
Popov's Dieneroceras Zone are very close
to those in the Columhites fauna of south-
east Idaho.
The naming of the Scythian post-Co-
lumhites Zone has not been settled to
everyone's satisfaction, mainly because of
erroneous views concerning composition of
the faunas and their correlation. Spath
( 1934 ) was the first to clearly recognize
the need for a unit (or zone) above that
of Columhites. For this upper unit he
introduced the Prohungaritan Division, with
the following comment: "On the other
hand, the uppermost beds of the Eo-Trias
are as yet very incompletely known, and
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 323
the use of the term Prohungaiitan (or in which it is not present relatively easy.
Olenikitan) is provisional, for the OZen//c/tes Wright (in Ager and Nichols, 1963) has
especially may yet be found to belong suggested an interesting base upon which
to the Columbitan, together with Keyser- to develop a logical picture of the spatial
lingites, although its derivative Dtirgaites relationships of the many late Scythian
is a characteristic element of the Lower faunas now known. Evidence is mounting
Anisian" (Spath, 1934:32, 33). We now that ammonites were gregarious and per-
have a much larger fund of data than was haps had distinct paths of migration, pos-
available to Spath. There is no longer any sibly as part of the breeding process. The
question as to the need for an additional modification and superimposing of gre-
zone above that of Coliimbitcs, but at the garious swarms could lead to the kinds of
same time I conclude that only one such assemblages that are encountered in these
zone is present world wide. To be sure, there late Scythian faunas. The faunas of the
are areas where several local zones are Werfen Formation and the Mangyshlak
present, e.g. Mangyshlak Peninsula, but Peninsula, with their large component of
these are only parts of a global zone. endemic genera and species, are clearly
There is only one species of ammonite geographic isolates developed in embay-
in the late Scythian that is world wide in dis- ments off the margin of Tethys.
tribution — Pseudosageceras muUilohaium. Few precise radiometric dates are avail-
However, a number of genera and even able for the Triassic ( Harland, et al., 1964 ) .
species have very widespread distribution. Holmes ( 1959 ) assigned a period of 45
Such forms and their patterns of over- million years, and Kulp ( 1961 ) 49 million
lapping associations provide the key for years to the Triassic; thus it seems reason-
definition of a global late Scythian Zone, able to conclude that the Scythian had a
Within Tethys there is a fairly homogeneous duration of possibly 7 to 10 million years,
fauna characterized by such forms as Suh- Within the Scythian there exist most prob-
coJumhites, Albonites, and Prohungorites. ably only five or six world-wide zones.
These Tethyan faunas have a high degree This suggests a duration of approximately
of similarity with faunas from the western 1 to 2 million years for each zone,
and eastern Pacific regions. There is much A few comments are appropriate regard-
less similarity with the faunas of the circum- ing the use of Scythian as the stage name
Arctic region. However, the presence of for the Lower Triassic in light of recent
several genera which are particularly char- attempts to introduce other names. The
acteristic of the circum-Arctic region within history of the name Scythian has already
faunas of the western and eastern Pacific been discussed (p. 313). The name has
and in Tethys offers the opportunity to had general acceptance as the stage name
make a correlation. For this latest zone for the Lower Triassic ever since the tenn
of the Scythian I feel we should retain the was introduced in 1895. The term is well
name Prohungorites as originally, though entrenched in the paleontological literature,
tentatively, suggested by Spath (1934). works on regional geology, text-books, etc.
There is surely much that remains to be In recent years proposals have been made
known about late Scythian ammonites, but to abandon the term Scythian and replace
on the basis of the available data, I see it with two or four new stage names. Some
no need to introduce changes in Spath's of these proposals were clearly intended
nomenclature. The genus Prohungarites is for use within a particular country, others
now known to be widely distributed in the were hopefully proposed for wider applica-
Tethyan, Pacific and Arctic realms. The tion. For instance, in New Zealand there
large number of forms associated with has been a general abandonment of Euro-
Prohungarites makes correlation of faunas pean stage names, and a new classification
324 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
specifically adapted to the New Zealand
stratigraphic panorama is used. Tlie local
stages are then correlated with the Euro-
pean stages. Ammonites are not common
in Triassic formations of New Zealand;
pelecypods and brachiopods are more com-
mon and play a greater role in the defini-
tion of the local stages. It should be noted
that the Triassic ammonites, known to date,
are genera common to faunas of Tethys
and the circum-Pacific region. A some-
what similar approach to Triassic chronol-
ogy has been proposed by Ichikawa ( 1950,
1956) for Japan. This system of classifica-
tion has not been generally adopted in
Japan.
Kiparisova and Popov (1956, 1961, 1964)
in a series of papers have proposed aban-
donment of Scythian as the stage name for
the Lower Triassic and the substitution of
two new stage names, the Indian and the
Olenekian. The lower of these stages, the
Indian, derives its name from the Indus
River, and the upper, the Olenekian, from
the Olenek beds of the Olenek River region,
northern Siberia. Tliese two stages are
equivalent to the Lower Eo-Trias and Up-
per Eo-Trias of Spath ( 1934 ) ; the boundary
of the stages was placed between the
Flemingitan and Owenitan divisions of
Spath. In a later contribution on this sub-
ject Kiparisova and Popov ( 1964 ) altered
the spelling of their stage names to Indus
and Olenek and lowered the boundary be-
tween the zones. Tliese authors concluded
that the ammonites of the Flcming,itcs
flemingiamis Zone are really equivalent to
the Owenites Zone; thus the upper zone
for the Indus stage at its type locality in
the Salt Range is the zone of Koninckites
volutu.s which these authors placed within
the Gyronitc.s- Zone.
The most recent contribution of new
stage names for the Lower Triassic of
Canada is by E. T. Tozer (1965b). This
author proposes for use in Canada four
stages: Griesbachian, Diencrian, Smithian,
Spathian. Tozer had the cooperation of the
Canadian Permanent Committee on Geo-
graphical Names in establishing names of
creeks in Arctic Canada after these promi-
nent Triassic paleontologists. The type
areas of these new stages are based on
sequences in Ellesmere and Axel Heiberg
Islands.
The presumed justification for establish-
ing these new stage names is to clarify com-
munication. Local stage names under cer-
tain circumstances are useful. However,
they tend to be based on incomplete data
and approached from a provincial point
of view. We need only look back at the
history of development of the geological
time scale to see the bad effects of the
proliferation of stage names for certain sys-
tems and the resultant confusion. One
would think we had reached a stage of
maturity in the science where repetition of
this sort of thing would not take place.
SUMMARY OF PROHUNGARITES ZONE
AMMONOIDS
The ammonoids of the Prohungarites
Zone, as here understood, comprise 65
genera with 154 species (Table 7). Of the
total number of genera, 42 are confined
to this zone, 20 genera are known to range
up from the preceding Columljifcs and
Owenites zones, one genus is knowm from
both older and younger horizons, and 2
genera are also present in the overlying
Anisian. The geographic distribution of
the genera is summarized in Table 8.
Twenty-four genera, or 36.9 percent of the
fauna, are confined to a single locality or
region; 10 genera, or 15.4 percent of the
fauna are known from two localities. Thus
52.2 percent of the total fauna of 65 genera
are known from only one or tw o localities.
On the other hand, only 10 genera, or 15.4
percent, are known from 6 to 11 localities
or regions. The degree of geographic re-
striction of genera is of particular interest.
There are 24 genera endemic to the Tethyan
rt>gion, that is, known only from one or two
localities within Tetlns. However, the
western Pacific, eastern Pacific, and Arctic
faunas have only two endemic genera each.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 325
Table 7. Summary table of genera and species of ammonites and their geographic distribu-
tion IN THE Prohungarites Zone. Symbols used in right hand column (range of genera and
species) are as follows: X = present only at this horizon, — = present in both younger
AND older horizons, e = present also in earlier horizons, 1 = PRESENT ALSO IN LATER HORIZONS.
c
o.S ^
•St: J'
P DC .^
cm H
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Oh
bij
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tlj (J oS
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60
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■12 bc-ls o "-^
3 5
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3
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-§ rt-2 MS c5 -Si ^>. *
S— ~ iK— "i: ■- >; S "^ ^^ re— -m-.2^ "Cos:
^^c/^ ^:/: -Cc/^ ;^0 aUi =« -CH 2 « 3^ CO; O 0) go, cs^
«^ ^^ 0^^ O^ ^^ K^ n^ PnU KZ HZ U;^ c/3^«c8
Sageceratidae
Pseudosageceras
inultilobatum
drincnse
albanicum
pasquayi
simplex
Cordillerites
angulatus
X
X
X
X
X
X
X
X X
X
X X
X
X
X X
X
X
e
e
X
X
X
X
e
e
Dieneroceratidae
Dieneroceras
mediterranea
skutarensis
karazini
Stibvishnuites
enveris
Hcmilecanites
discus
paradisctts
Xenoceltidae
Xcnoceltites
sinuatus
crenoventrosus
spitshergensis
Prcflorianites
sulioticus
garbinus
multiplicatus
intermedius
X
X
X
X
X
X X
X
X
X X
X
X
X
X
X
X
X
X
e
X
X
X
e
X
X
X
X
e
X
X
e
e
X
X
X
X
Proptychitidae
Proptychitoidcs
decipiens
trigonalis
arthaberi
tunglaneiwu
kummeli
Procarnites
kokeni
immaturus
lolouensis
X
X
X
X
X X X X X
Paranannitidae
Arnautoceltites
mediterraneus X X
bajarunasi
involutus
gracilis
teicherti
Prosphingites
czekanowskii
alt X
lolouensis
suhglobosus
globosus
insularis
coombsi
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
e
X
X
X
X
X
X
X
326 Bulletin Museum of Comparotive Zoology, Vol. 137, No. 3
Table 7. Continued
^^ ^^ ^^ ^ ^-^ ^^ ^^
|S .^ ^ ^ Us I 1^1-^ .§-a.^§i 111^:5 I i ^ g fj £
^^ I JslJc^l.S S ^: "l |*|«| N i:Sao sis I 5 |l^-3
^Z: <;r2. u- S <^ c;5> t^- H- 5i ui- 4^ £-i O- (^--^ W3. psC ^u K2 HZ Ulz c;5- rt 5
Vickohleritcs X X
stiudaicus XX X
Zenoites X
heleiuie X X
vonderschmiui X X
arcticus X X
Isculitoides XX X
orighus XXX X
vllipticus X X
siiboviformis X X
minor X X
icasserbergi X X
liammondi X X
Chiotites X
glolndaris X X
Czekanowskites X X
decipiens X X
Popovites X
occidentalis X X
borealis X ..X
Monocanthitcs X
monoceras X X
Ttinglanites X
Icuficularis X X
o/e.vi XX X
Suhcohimbites X
pcrrhiismifhi XXX XX ■ X
dii.smani XX X
rohtisttis X X
midtiformis X X
americanus X X
Paradinarites X
siini X X
Pseudoceltites e
dohmpaensis X X
nevadi X X
Procolumbites X
karataticiktis X X
Prenkites X X
malsorcnsis XX X
hclenac X X
timorensis XXX X
Protropite.i X
hj7nit X X
Chioceras X
mitzopotdoi X X
nodosum X X
Arianifes X
nnusacchi X X
MeropeUa X X
plejanac X X
Epiceltites X
gentii XX XX
stibgracilis X X
Ussiiriidae
Parussnria e
lalilohata X X
Hedenstroemiidae
Metahedenstroemia X
kastriotae XX X
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 327
Table 7. Continued
(-;
r^
tz
i~^
tc
2
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<-^
<-*-
pc;
^
C^
a =5
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^
cc ^
*-
s
^
^
a
s
2
2
w:
=
5;^
c
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c
o
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rv..
c
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■y^
■^
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B
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6
7:
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bio
rat:
C>4-,
0; C
Sit;;-
0^ ^
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= T.-~ .-ti '-^ a^
Si
Si
-^ "w ^ O -C^ ;:; '^
■•Co ^IS §>S>
ffi" ci;U KZ HZ
o-
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UZ
a;
c
5? <*--Fl
;(X, s
K C3
berthae X
Lanceo/ifes
discoidalis X
Meekoceratidae
Sra/fcnrdtcera.s
«btrici<ni
denlosus
freboldi
chowadei
Stacheites
prionoides X
flowerl
Dagnoceras
nopcsanum X
zappancnse X
latilobatiim
eUipticum
Metadaguoceras
pidcher
tobini
frecmani
terbunicum X X
Balkanites
tabulatus X
Nordophiccras
pseudosimplex
planorbis
coinpressttm
Pseudokymatites
svilajamts X
Arctomeekoceras
rottindatum
Boreomeckoceras
keyscrlingi
Arctotirolitcs
menensis
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Noritidae
Albanites
triadicus
Sibiritidae
Sibirites
eichwaldi
renzi
Keyserlingites
sitbrobustus
middendorffi
bearlakensis
bearriverensis
Olenekites
spiniplicatns
mangy sJdaken^is
canadensis
Euka^hmirites
subdimorphus
contortits
Anakashmirites
X X X X
X
X
X
X X X X
X
X
X
X
X
X
X
X
X
X
X
X
e
X
e
X
X
X
X
X
X
X X
X
X
X
X
X
X
X
e
X
X
X
X
X
X
e
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
e
X
X
X
X
e
X
X
X
X
X
X
e
328 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Table 7. Continued
S.2
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CO
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£-3
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Tirolitidae
TiroUtes
idricnuis
cassiauiis
cingulatus
rossicus
impolitus
morpheas
Diaplococeras
liccanum
connectens
BUtnerites
bittneri
Doricranitcs
bogdoantis
acuttis
Dinaritidae
Dinarites
dalmcitijius
carnioliciis
liatsikasi
nndatus
Hololohus
monoptychiis
Psciidodinarites
mohamedaniis
Hellenitidae
Hellenites
praemaUtrus
radiatus
Beyrichitidae
Betjrichites
latirae
Gymnitidae
Eogymrntes
arthoberi
Hungaritidae
Prohungarites
crassepUcatus
ttibcrculatus
middleniissii
carinatus
mckelcci
gutstadti
Dcdmatites
morlaccus
Ussuritidae
Eophyllites
dieneri
orieutalis
amurensis
PalaeophtjlUtes
steiruTKmni
Vssurites
nieveri
hoesi
1 .ciophyllites
variabilis
radians
serpentimts
admaris
maritinius
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X X
X
X X
X
X
X
X
X
X
X
X
X
e
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
e
X
X
1
X
X
X
X
X
X
X
X
X
X
e
X
X
X
X
X
X
X
1
X
X
1
X
X
X
X
X
Ammonoids of the Late Scythian (Lower Triassic) • Ktimmel
329
Table 8. Geographic distribution of ammo-
NOID GENERA OF PrOHUNGARITES ZoNE IN TERMS
OF NUMBERS OF LOCALITIES OR REGIONS IN WHICH
EACH GENUS IS PRESENT.
Number of
Number of
Percentage of
Localities
Genera
Total Fauna
1
24
36.9
2
10
15.3
3
6
9.2
4
8
12.3
5
7
10.7
6
5
7.7
7
2
3.1
8
0
0
9
1
1.6
10
1
1.6
11
1
1.6
Within the Tethyan region the highest
degree of endemism is found in the fauna
of the Werfen Formation, with 6 genera
out of a total of 11 restricted to that fomia-
tion. A seventh genus, Dalmatites, is in the
Proliungaritcs Zone, restricted to the Werfen
Formation, but another species of this
genus occurs in the underlying Columbites
Zone in southeast Idaho. Diaplococcras is
represented by two species, but the other
endemic genera by only a single species
each. Two of the species are known from
only a single specimen each and the others
from very few specimens.
The Sub columbites fauna of Albania in-
cludes four endemic genera — Protropites,
Arianites, Beatites, and Eogymnites; Pro-
tropites is a fairly common member of that
fauna but the other three genera are known
only from a single specimen each. The
Subcolumbites fauna of Chios also contains
four endemic genera — Chiotites, Chioceras,
Meropella, and Beyrichites. Among these
four genera, Chiotites is abundantly repre-
sented in the Chios fauna, but the other
three genera are known from only three
or four specimens each. In the Mangyshlak
Peninsula the late Scythian fauna includes
three endemic genera — Procolumbites, Eii-
kashmirites, and Doricranites; of these,
Doricranites appears to be by far the most
common form. In the Salt Range the
Prohungarites Zone contains fragmentary
specimens of Anakashmirites, a genus not
recorded from any other locality in this
zone.
Within the western Pacific realm there
are two endemic genera — Paradinarites and
Parussuria — from the Subcolumbites faunas
of Kwangsi, China. In the eastern Pacific
realm there are only two endemic genera
in the Prohungarites Zone faunas. Mono-
canthites is known only from late Scythian
strata in British Columbia, and Ussiirites
only from Nevada and Utah. Ussurites is a
fairly common Anisian genus, but the
species recorded here are the first from
the late Scythian. In the Arctic realm
there are two endemic genera, Boreomee-
koceras and Arctotirolites, both known
from the Olenek fauna.
In evaluating the degree of similarity of
these late Scythian faunas of the Prohun-
garites Zone, Simpson's index of faunal
resemblance is useful (Simpson, 1943, 1947,
1953). This index is symboHzed as 100
C/N, in which C stands for the number of
taxonomic units common to two faunas,
and N is the total number of genera in the
smaller of the two. The number of genera
in common and the index of correlations,
on the basis of genera of these faunas at-
tributed to the Proliungaritcs Zone, are
given in Tables 9 and 10.
It is recognized that the composition of
any of these faunas is influenced by factors
of preservation, collection techniques, and
facies. Evaluation of the relative signifi-
cance of these factors in regard to each of
the faunas is most difficult. The Sub-
columbites faunas of Albania and Chios ap-
pear to have been thoroughly collected.
This applies also to the fauna of the Werfen
Formation; however, in this case more
stratigraphic data are desirable. It is my
impression from the literature that the
Mangyshlak fauna contains more genera
and species than those described to date.
The Afghanistan and Salt Range faunas are
known from small collections, mainly of
poorly preserved specimens. The chance
330 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Table 9. Summary chart of numbers of genera in common between faunas of the Prohun-
GARITES Zone from nineteen localities or regions.
1
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y.
a
c
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C/3
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c
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15
33
si
II
0
Werfen Fm.
11
1
1
3
0
2
0
0
0
0
0
1
0
0
0
1
1
1
Albania
1
28
19
7
8
6
0
10
13
1
11
4
0
0
6
8
2
3
Chios
I
19
27
7
7
3
0
9
10
1
9
4
0
1
5
8
1
3
Mangyshlak
3
7
7
15
4
5
1
4
3
0
4
3
0
1
1
4
3
4
Afghanistan
0
8
7
4
11
5
0
5
5
1
6
3
1
1
3
4
1
3
Salt & Smghai
Ranges
2
6
3
5
5
13
11
5
5
0
4
5
2
1
3
3
2
4
Kashmir
0
0
0
1
0
1
1
1
0
0
0
0
0
0
0
0
0
1
Timor
0
10
9
4
5
5
1
12
4
0
4
2
0
0
2
1
2
New Zealand
0
1
0
0
0
0
0
0
1
0
1
1
0
0
1
0
0
0
Kwangsi, Chin.
I 0
13
10
3
5
5
0
4
16
1
8
4
0
0
4
6
0
2
Japan
0
1
1
0
1
0
0
0
1
1
1
{)
0
0
0
1
0
0
Primorye
Region
0
11
9
4
6
4
0
4
8
1
11
0
0
3
5
1
2
Olenek
1
4
4
3
3
5
0
2
4
0
3
15
2
2
4
1
.")
5
Spitsbergen
n
()
0
0
1
2
0
0
0
0
0
2
2
2
2
0
0
2
Ellesmere
Island
0
0
1
1
1
1
0
0
0
0
0
2
2
4
3
0
0
2
British
Columbia
0
6
5
1
3
3
0
2
4
0
3
4
2
3
11
2
0
3
Tobin Range
I
8
8
4
4
3
0
6
1
5
1
0
0
2
10
2
O
Confusion
Range
1
2
1
3
1
2
0
1
0
0
1
2
0
0
0
2
4
1
Southeast
Idaho
I
3
3
4
3
4
1
2
2
0
2
5
2
2
3
3
1
9
Total Genera
11
28
27
15
11
13
I
12
16
1
11
15
2
4
11
10
4
9
of ever uncovering larger and better pre-
served faunas from these localities is not
good. The Timor fauna is known from
isolated blocks mainly from a single locality.
It is apparent that Welter's ( 1922 ) mono-
graph has by no means dealt exhaustively
with this fauna. New collecting on Timor
is badly needed. The Pioliungarites Zone
faunas of New Zealand and Japan consist
of a single species each. In these two areas
facies and preservations have been the
most severe limiting factors. The Sub-
columhites fauna of Kwangsi, China, ap-
pears to have been adequately collected.
The faunas of the Primorye Region and
northern Siberia have been collected exten-
sively but there is urgent lu^ed of more
detailed stratigraphic and geographic data.
I have the impression from the literature
that ammonites are not abundant in these
regions. Neither are ammonites of the
Frohuniiaritcs Zone abundant in Ellesmere
Island or in British C'olumbia. In these
two areas logistical difficulties further com-
plicate the problem and offer perhaps an
explanation for the small size of the known
faunas. The hunia of the Tobin Formation
has been well collected, but I am sme that
more search will yield additional specimens.
The same applies to the fauna from Ham-
mond Creek in southeast Idaho.
Considering all these factors, one should
use some r(\straint in evaluating the faunal
indices of Table 10; this of course applies
more to ihc smaller faunas. If one assumes
that contiguous faunas \\'ithin a geologic
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 331
Table 10. Index of faunal similarity at a generic level for nineteen ammonoid faunas of
THE PROHUNGARITES ZoNE.
c
^
2
C
^
CA
rt
'—
■a
c
a
N
1/
1-'
^^2
1-1
i5
1;_
pi
11
bc
S 1/
1*
I2
0
^
<
u
S
<
C>3
t^
H
2
u:
lA
CU
5
Cn
s
ffl
t2
u
tXl
Werfen Fm.
100
9
9
27
0
18
0
0
0
0
0
0
9
0
0
0
11
25
11
Albania
9
100
70
46
72
46
0
83
100
81
100
100
26
0
0
54
80
50
33
Chios
9
70
100
46
63
23
0
75
0
62
100
81
26
0
25
45
80
25
33
Mangyshlak
27
46
46
100
36
38
100
33
0
20
0
36
20
0
25
9
40
75
44
Afghanistan
0
72
63
36
100
45
0
45
0
45
100
54
27
50
25
27
40
25
33
Salt & Surghar
Ranges
IS
46
23
38
45
100
100
41
0
38
0
36
38
100
25
27
30
50
44
Kashmir
0
0
0
100
0
100
100
100
0
0
0
0
0
0
0
0
0
0
100
Timor
0
83
75
33
45
41
100
100
0
33
0
36
16
0
0
18
30
25
22
New Zealand
0
100
0
0
0
0
0
0
100
100
0
100
100
0
0
100
0
0
0
Kwangsi, Chin
a 0
81
62
20
45
38
0
33
100
100
100
72
26
0
0
26
60
0
22
Japan
0
100
100
0
100
0
0
0
0
100
100
100
0
0
0
0
100
0
0
Priniorye
Region
0
100
81
36
54
36
0
36
100
72
100
100
27
0
0
27
50
25
22
Olenek
9
26
26
20
27
38
0
16
100
26
0
27
100
100
50
36
10
50
55
Spitsbergen
0
0
0
0
50
100
0
0
0
0
0
0
100
100
100
100
0
0
100
Ellesmere
Island
0
0
25
25
25
25
0
0
0
0
0
0
50
100
100
75
0
0
50
British
Columbia
0
54
45
9
27
27
0
18
100
26
0
27
36
100
75
100
20
0
33
Tobin Range
11
80
80
40
40
30
0
30
0
60
100
50
10
0
0
20
100
50
33
Confusion
Range
25
50
25
75
25
50
0
25
0
0
0
25
50
0
0
0
50
100
25
Southeast
Idaho
11
33
33
44
33
44
100
22
0
22
0
22
55
100
50
33
33
25
100
Total Genera
11
28
27
15
11
13
1
12
1
16
1
11
15
2
4
11
10
4
9
province reflect better the actual com-
position of the fauna, a computation of
faunal resemblance between geologic prov-
inces could be more meaningful. Scythian
paleogeography is characterized by Tethys
and by marginal geosynclines on the con-
tinents surrounding the Pacific and Arctic
oceans. The Spitsbergen, Ellesmere Island,
and northern Siberian (Olenek) faunas thus
fall within an Arctic province. The eastern
Pacific province includes the faunas from
British Columbia, Nevada, Utah, and south-
east Idaho. The western Pacific province
includes the faunas from the Primorye Re-
gion, China, Japan, Timor, and New Zea-
land. The Tethyan province includes the
Kashmir, West Pakistan, Afghanistan, Man-
gyshlak, Chios, Albania, and Werfen For-
mation faunas. The index of correlation of
the faunas between these geologic provinces
is given in Table 11. The Tethyan province
has the largest fauna in total numbers of
genera, and the largest number of genera
are endemic to that province. In contrast,
the Arctic province has the smallest total
number of genera, while the eastern and
western Pacific provinces are intermediate
and have the same number of genera.
There is a high degree of correlation of the
western and eastern faunas with those of
Tethys; the Arctic faunas show a lesser
degree of correlation. It is of interest to
note the relatively low degree of correla-
tion between the Arctic province and the
western Pacific; the correlation between
the Arctic and the eastern Pacific provinces
332 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Table 11. Index of faunal similarity at a
generic level for ammonoid faunas of the
Prohungarites Zone between the major
paleogeographic provinces.
Table 12. Geographic distribution of ammo-
NOiD species of Prohungarites Zone in terms
OF NUMBER OF LOCALITIES OR REGIONS IN WHICH
A SPECIES IS PRESENT.
Tethvs W. Pacific E. Pacific Arctic
Number of
Localities
Tethys
Number of
Species
Percentage of
Total Fauna
100.0 92.0 84.0 68.7
W. Pacific
92.0
100.00
48.0
31.2
E. Pacific
84.0
48.0
100.00
56.3
Arctic
68.7
31.2
56.3
100.0
Total genera
57
25
25
16
Number of
endemic genera
24
2
2
2
Percentage of
endemic genera
42.1
8.0
8.0
12.5
(56 percent) is approximately twice as
good. At the same time the correlation be-
tween the eastern and western Pacific prov-
inces is only 48 percent.
Only two of the 65 genera of the Pro-
Juin^a rites Zone, Pseudosoiieceras and Fro-
S])hin0tcs, are present in all four of the
main geologic provinces. There are, how-
ever, ten additional genera which occur in
the Tethyan, western Pacific, and eastern
Pacific provinces:
Cordillerites
Hemilecanites
Procarnitcs
Arnautoceltites
Isciilitoides
Siibcoltimbites
Metadagnoceras
HeJlenites
Prohungarites
LeiophtjUites
Of this list of 10 genera, 7 are confined to
the Proliungaritcs Zone, 2 are known from
older horizons, and one is known from
younger horizons.
Discounting for the moment the endemic
genera within each province, the above
widely distributed genera constitute 33 per-
cent of the Tethyan fauna and 48 percent
of the western and eastern Pacific faunas.
The faunas of the Arctic regions are more
distinctive than the index of faunal simi-
larities tends to suggest. The Arctic prov-
ince includes 16 genera, of which only two
(Boreomeekoceras and Arctotirolites) are
endemic. Whereas all but two of the
genera in the Arctic province are present
in one or more of the other three major
geologic provinces, there are significant dif-
1
2
3
4
5
7
10
117
23
8
2
2
1
1
75.9
14.9
5.2
1.3
1.3
0.6
0.6
ferences in the relative representation of
genera within this and the other provinces.
The most characteristic and abundant genera
in the Arctic fauna are Svalhardiceras,
Sibirites, Keijserlingites, and Olenekites.
Each of these genera is represented outside
of the Arctic province by few species and
specimens. Svalhardiceras is also known
from a few fragmentary specimens in the
Salt Range and in southeast Idaho. Sibirites
is also known from only a few small speci-
mens in the Subcolumbites fauna of Chios.
Outside the Arctic region, Keyserlingitcs
is known in the Tobin Formation of Ne-
vada, the Thaynes Formation of southeast
Idaho, and at Kotal-e-Tera, Afghanistan.
Finally, Olenekites is known from only a
few specimens in southeast Idaho and on
the Mangyshlak Peninsula. In contrast to
the sparse representation of these genera,
in terms of numbers of localities represented
and numbers of specimens outside of the
Arctic province, they are widespread and
common elements of the Arctic fauna.
T^enoites was first recognized in the Sub-
eoliimbites fauna of Chios and is now
known also from Ellesmere Island. Like-
wise, Pro))tyehitoides is a very common
Tethyan form, now known to be repre-
sented by one specimen in northern Siberia.
In contrast, Prosphingites is quite common
in the eastern and western Pacific realms
but is relatixely rare or uncommon in
Tetliys and the Arctic province.
An analysis of the Prohungarites Zone
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel
333
Table 13. Summary chart of numbers of species in common between the faunas of the Pro-
hungarites zone from nineteen localities or regions.
£
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Southeast
Idaho
Werfen Fm.
15
1
1
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Albania
1
34
24
3
4
2
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6
0
5
1
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1
0
0
1
1
1
2
Chios
1
24
40
3
4
2
0
6
0
7
1
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1
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0
1
2
1
2
Mangyshlak
2
3
3
19
3
2
0
3
0
1
0
1
1
0
0
0
1
1
0
Afghanistan
0
4
4
3
4
2
0
3
0
2
1
1
1
0
0
0
1
1
0
Salt & Surghar
Ranges
0
2
2
2
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4
0
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0
0
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Kashmir
0
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0
0
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0
0
0
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0
0
0
0
0
0
0
0
Timor
0
6
6
3
3
2
0
13
0
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0
1
1
0
0
0
1
1
0
New Zealand
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
Kwangsi, China 0
5
7
1
2
1
0
2
0
22
1
0
0
0
0
1
1
0
0
Japan
0
1
1
0
1
0
0
0
0
1
1
0
0
0
0
0
0
0
0
Primorye
Region
0
2
2
1
1
1
0
1
0
0
0
14
1
0
0
1
1
1
0
Olenek
0
1
1
1
1
1
0
1
0
0
0
1
18
2
1
2
1
1
0
Spitsbergen
0
0
0
0
0
0
0
0
0
0
0
0
2
3
1
1
0
0
0
Ellesmere
Island
0
0
n
0
0
0
0
0
0
n
0
0
1
1
4
1
0
0
0
British
Columbia
0
1
1
0
0
0
0
0
0
1
0
1
2
1
1
11
0
0
0
Tobin Range
0
1
2
1
1
1
0
1
0
1
0
1
1
0
0
0
9
1
0
Confusion
Range
0
1
1
1
1
1
0
1
0
0
0
1
1
0
0
0
1
3
0
Southeast
Idaho
0
2
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
7
Total Species
15
34
40
19
4
4
1
13
1
22
1
14
18
3
4
11
9
3
7
fauna at the generic level shows consider-
able similarity within and between the
major geologic provinces. At the species
level, as one would expect, there is a greater
degree of endemism, that is, species known
from a single region. Table 12 summarizes
the geographic distribution of the species
known from the Prohungaritcs Zone. It is
impressive to note that approximately 75
percent of the species are known from
only a single locality or region and only
about 8 percent are known from 3 to 8
localities. The numbers of species in com-
mon between the various localities and re-
gions are tabulated in Table 13, and the
index of correlation (percent) in Table 14.
In general there are very low or zero
indices of correlation. It is impressive, how-
ever, to see the very high correlation be-
tween the Siihcohimhites faunas of Albania
and Chios; lesser but still significantly high
correlations exist between the Albanian and
Chios faunas and those of Timor and
China. The main aspect of the chart is
that there is a higher degree of correlation
between localities within a geologic prov-
ince than there is between different prov-
inces.
In collecting from most ammonitiferous
deposits it is a common experience that a
few genera and species are overwhelmingly
predominant, a few species are represented
by modest numbers, and a group of species
is represented by only one or two specimens
each, even after extensive collecting. Even
though there are multiple factors which
334 BiiUefin Museum of Comparative Zoology, Vol. 137, No. 3
Table 14. I
NDEX
OF
FAUNAL SIMILARITY
AT
THE
SPECIES 1
LEN-EL FOB
L NINETEEN AMMONOID
FAUNAS OF
THE
Prohvngarites Zone
a
^
5
C3
^
.3
.2
<
V5
U
>•
c
"5
<
en
N
>
z
II
It
r-
^1
.sec
Si
0
1/
0;
IX
c
"3
c
II
U
C/2
Werfen Fm.
100
6
6
13
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Albania
6
100
70
15
100
50
0
46
0
22
100
14
5
0
0
9
11
33
28
Chios
6
70
100
15
100
50
0
46
0
31
ino
14
5
0
0
9
22
33
28
Mangyshlak
13
15
15
100
75
50
0
23
0
5
0
7
5
0
0
0
11
33
0
Afghanistan
0
100
100
75
100
50
0
25
0
50
100
25
25
0
0
0
25
25
0
Salt & Surghar
Ranges
0
50
50
50
50
100
0
50
0
25
0
25
25
0
0
0
25
33
0
Kashmir
0
0
0
0
0
0
100
0
0
0
0
0
0
0
0
0
0
0
0
Timor
0
46
46
23
75
50
0
100
0
15
0
7
7
0
0
0
11
33
0
New Zealand
0
0
0
0
0
0
0
ion
0
0
0
0
0
0
0
0
0
0
Kwangsi, China 0
22
31
5
50
25
0
15
0
100
100
0
0
0
0
9
11
0
0
Japan
0
100
100
0
100
0
0
0
0
100
100
0
0
0
0
0
0
0
0
Primorye
Region
0
14
14
7
25
25
0
7
0
0
0
100
7
0
0
9
11
33
0
Olenek
0
5
5
5
25
25
0
7
0
0
0
7
100
66
25
18
11
33
0
Spitsbergen
0
0
0
0
0
0
0
0
0
0
0
0
66
100
33
33
0
0
0
Ellesmere
Island
0
0
0
0
0
0
0
0
0
0
0
0
25
33
100
25
0
0
0
British
Columbia
0
9
9
0
0
0
0
0
0
9
0
9
IS
33
25
100
0
0
0
Tobin Range
0
11
22
11
25
25
0
11
0
11
0
11
11
0
0
0
100
3:3
0
Confusion
Range
0
33
33
33
25
33
0
33
0
0
0
33
33
0
0
0
33
100
0
Southeast
Idaho
0
28
28
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
100
Total Species
15
34
40
19
4
4
1
13
1
22
1
14
18
3
4
11
9
3
7
can and do influence preservation, I be-
lieve it reasonable to assume that the num-
ber of specimens of a species represented
in a well collected fauna does reflect the
relative abundance of the species. If one
accepts this assumption, it makes possible
a fiulher evaluation of the composition of
the various faunas of the Prohung,aritcs
Zone and changes of abundance within
the various populations.
Data on the total number of specimens
and number of specimens per species for
nine faunas are given on Table 3. The
collections from the Suhcolumhitc.s- faunas
of Albania and Chios are by far the largest.
The five commonest species in the Suh-
columhitcs fauna of Albania in order of
abundance are:
Siihc(>luml>ifcs iicniuisiiiitJii
Procdiiiitc.s kokciii
Isculito ides origin is
Protropifcs liihiii
Frrtikitrs nuilsDrcnsis
The 9 species in the Chios fauna that are
represented b\' 50 or moic specimens in
order of abundance are:
Isculitoidcs ()iiiJ.itiis
Frocariiitcs kokciii
Lciopliyllitc.s varidhili.s
Chioccid.s i)iifz()i)i)ul(>i
Alhauitcs triadicu.s
}'](ypU[lUltcs divncri
I Irllcnitcs praciiuitiinis
I'.svudosvgc'ceras alhanicum
Sid)((>lui)d)itcs pcninisniithi
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 335
Three of these species {Isciilitoides
originis, Procarnites kokeni, StibcoJumhites
pcrrinismifhi) are predominant members in
iioth faunas. Protropites hilmi is endemic
to Albania, as is Chioccras mitzopouloi
to the Chios fauna. Whereas Prcnkitcs
malsorcnsis is predominant in the Albanian
fauna, it is represented by only two speci-
mens in the Chios collection; HcUenitcs
pracmoturus is very common in Chios but
represented by only two specimens in the
Albanian collection. Even though Leiophijl-
lites variabilis is not on the list from Albania,
it still is quite abundant, with 43 specimens
in the collection; the same applies to
EophyUites dieneri with 40 specimens.
Subcohinihitcs is predominant in the
Albania, Chios, Primorye, and Tobin For-
mation faunas. In Albania and Chios the
species is perrinismithi, in the Primorye
Region multiformis, and in the Tobin For-
mation amcricanus. Subcolumbites is also
known from the Afghanistan and K\\'angsi
faunas, though in both collections by very
few specimens only.
Hellenitcs is a predominant element only
in the Chios fauna; in the Albanian, Kwangsi,
and Nevada faunas it is represented by
very few specimens. Procarnites kokeni
occupies a position of predominance in the
Albanian, Chios, and Kwangsi faunas. It
is known from very few specimens in the
Afghanistan, Salt Range, and Timor faunas.
Isciditoides is a predominant element in
the Albanian, Chios, and southeast Idaho
faunas but has only minor representation
in the faunas of Afghanistan, Salt Range,
China, Primorye, Nevada, and British
Columbia.
Olenekitcs is a predominant element in
the faunas of northern Siberia and Elles-
mere Island, modestly represented in the
Tobin Formation of Nevada (S.W. Muller,
personal communication), and very minor
in the faunas of the Mangyshlak Peninsula
and southeast Idaho. Prohungarites is over-
whelmingly the predominant element of
the fauna from the Upper Thaynes Forma-
tion of southeast Idaho but is apparently
Table 15. Number of species per genus among
GENERA OF AMMONOIDS IN THE PrOHUNGARITES
ZONTE.
Niim
ber of
Number of
Percentage of
Species
per
Genus
Genera
Total Genera
1
29
44.6
2
13
20.0
3
8
12.3
4
5
7.7
5
5
7.7
6
4
6.1
7
1
1.5
only a minor element of the faunas of
Timor, Kashmir, and the Mangyshlak
Peninsula.
A look at the 11 genera which are repre-
sented in the Tethyan, western Pacific and
eastern Pacific provinces yields some inter-
esting data. Four of these genera {Cordd-
lerites, Uemilecanitcs, Prosphingites, and
Metadagnoceras) are not a common ele-
ment in any of the faunas from which they
are recorded. Procarnites and Hellenites
are very common in one or more localities
within Tethys but nowhere else. Sub-
cohimbites, Isciditoides, and Arnautoceltites
are predominant elements in one or more
faunas in the Tethyan, western Pacific and
eastern Pacific provinces. Prohungarites is
common only in the eastern Pacific and
LeiophijUites in the western Pacific.
Most genera from the Prohungarites
Zone, that is, approximately 45 percent, are
known from only a single species (Table
15). This for the most part reflects the
high number of monotypic genera for this
zone. At the same time some genera, for
example Cordillerites, are widely distri-
buted in three of the major paleogeographic
provinces, though represented by a single
species only. Only 15 genera, or 23 per-
cent of the total fauna, are represented by
four or more species. Eight of these genera
with four or more species are among those
which are known from at least three of the
major paleogeographic provinces. Most of
the other genera of this group are con-
spicuous elements of the faunas of one or
two of the major paleogeographic provinces.
336 BuUeiin Museum of Comparative Zoology, Vol. 137, No. 3
PALEOGEOGRAPHIC IMPLICATIONS OF
PROHUNGARITES ZONE AMMONOIDS
Interest in paleoclimates has greatly
heightened since the introduction of paleo-
magnetic techniques. In the excellent re-
view on paleomagnetism by Cox and Doell
(I960), it was encouraging to read in their
summary a statement of the need for more
and better paleontological data as a test of
the geophysical conclusions now being ad-
vocated by many researchers. In the past
decade there has been a definite increase
in paleontological contributions to the prob-
lem of paleoclimates. The question rests
on the assumption that temperature is the
most significant of the factors that controls
the distribution of many organisms. Clear-
cut temperature gradients are recognized
in the distribution patterns of many modem
faunas and floras. The general problem of
temperature gradients has been given an
excellent review by A. G. Fischer (1960).
The most active student of paleontology
who has been applying the techniques of
faunal gradients to the problem of paleo-
climates has been F. G. Stehli. Ample
testimony to the complexity of the problem
can be seen in the number of enthusi-
astic rebuttals to many of the interpreta-
tions of Stehli and others. Craig (1961)
has amply reviewed the techniques and
limitations in using fossils for paleoclimatic
interpretations. He concludes that (a) "we
have insufficient knowledge of, and there-
fore control over, the variables that can
affect the form and distribution of fossils,"
and (b) "stratigraphic correlation is not
yet sufficiently accurate to be certain
that we are interpreting paleoclimates of
the same geological age" (Craig, 1961:224).
Craig emphasizes that these somewhat
"pessimistic conclusions" are meant to
"sound a cautionary note."
Interpretation of the significance of the
geographic distribution of the ammonoids
of the Frohiing,aritcs Zone is handicapped
by the sparscness of data from the South-
ern Hemisphere*. There are no late Scythian
marine deposits nor faunas as yet identified
from Central and South America. The
same applies to all of Africa, Madagascar,
Australia, and Antarctica. As stated several
times above, the basic paleogeographic
pattern of the Scythian is that of Tethys,
with circum-Atlantic and circum-Arctic
marginal seas. There are no marine late
Scythian deposits in the Atlantic basin south
of Spitsbergen. This pattern of distribu-
tion of the known data precludes any direct
comment on the problem of large scale
horizontal displacements of continents. It
is, however, worthwhile to consider the
spatial distribution of the ammonoids of
the Proluin^arites Zone. We are first of all
dealing with faunas considered to be part
of a single zone that probably had a time
duration of one to two million years.
Working on a world scale I do not believe
we shall be able to resolve our correlations
much finer than this. The fact that the
Tethyan region has the largest numbers
of genera and species, the circum-Arctic
region only approximately 30 percent as
many, and the western and eastern Pacific
areas an intermediate amount could ver\
well reflect a climatic pattern. In addition,
carbonate rocks are common in the Tethyan
and circum-Pacific regions but are essen-
tially absent in the circum-Arctic region for
this segment of time. If one accepts the
assumption that the known record to date
is a reasonably valid reflection of the extent
and diversity of late Scythian faunas, I
find it most plausible to see here a true
faunal gradient which most prol)ably re-
flects climate. The data inteipreted in this
vein are more plausible when compared to
the pattern of world continents that exists
today than to any of the many and varied
alternative maps which ha\e been drawn.
SUMMARY OF COLUMBITES ZONE
AMMONOIDS
The CohimJ)ites Zone is now known from
four localities, southeast Idalio, Arctic
Canada, Primorve Region, and northern
Siberia. A suininar\' list of the spc^cies and
iienera of this zone is tabulated in Table 16.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 337
Table 16. Summary table of genera and species of ammonites and their geographic distribu-
tion IN the Columbites Zone. S-i^MiBOLS used in right hant) column (range of genera and species)
ARE AS follows: X = present only at this horizon, — PRESENT IN BOTH YOUNGER AND OLDER
horizons, e =: PRESENT ALSO IN EARLIER HORIZONS, 1 = PRESENT ALSO IN LATER HORIZONS.
5 o
^
i-.
a
5 o
1)
C.2
r.
c — -
N '-
^ -S
— ' .—
N ^
=o .S
m
1 :.
U 1/
2"^
^ be
•■S CD
0 2
^ 4-
~ " aj
S 3 ?
"o o
Is-
'^ ^ r~
in 5
O Oj
5'2
UcAi
5z
fe: £
rv- ;;:
Uc/3
Sz
a; P-,
rt S
Sageceratidae
Metadagnoceras
1
Pseudosageceras
- —
unicum
X
X
midtilohatum
X
X
—
Nordopliiccras
1
Coidillerites
—
euomphalus
X
X
angidatus
X
—
alexeevae
X
X
Dieneroceratidae
jacksoni
X
X
Dieneroceras
—
pilaUitn
X
X
demokidovi
X
X
Prionitidae
apostolicus
X
X
X
Hemipiionites
e
Subvishnuites
—
costatus
X
X
eiekitensis
X
X
Sibiritidae
Xenoceltitidae
Keyseiiingitcs
X
1
Xenoceltites
—
stephensoni
X
X
spencei
X
X
Olenekites
X
1
Preflorianites
montpelierensis
X
X
Tirolitidae
Tiwlites
Paranoritidae
Pseudaspidiies
e
harti
smithi
X
X
X
X
popovi
X
X
astakhovi
X
X
posteriiis
X
X
Paranannitidae
Cohtnibitcs
Hellenitidae
X
Hellenites
idahocnse
X
1
X
X
parisianus
X
X
X
Procolumhites
X
1
inopinatiis
X
X
Neocohimbites
X
X
Hungaritidae
Pseiidoceltites
e
Dahnotites
—
cheneyi
X
X
kittli
X
X
Meekoceratidae
Ussuriidae
Svalbardiceras
X
1
Ussiirites
1
sheldoni
X
X
mansfieldi
X
X
The limited areas from which this fauna
is known do not allow the broad synthesis
that was possible for the ammonoids of
the Frohungarites Zone. Most of the perti-
nent data regarding the ammonoids of this
zone and their relationship to the under-
lying faunas of the Owenites Zone and the
overlying faunas of the Prohunga rites Zone
have already been discussed in the section
on chronology. The relationships of the
ammonoid fauna of the Columbites Zone to
those of the underlying Owenites Zone
and the overlying Frohungarites Zone are
summarized at a generic level on Table 17.
STRATIGRAPHY AND
LATE SCYTHIAN
FAUNAS OF THE
Ammonoid faunas of late Scythian age
are now known from 17 localities or regions
( Fig. 1 ) . In this chapter are summarized
the stratigraphic and faunal data on each
of these occurrences. The discussion first
338 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Table 17. Total genera and their ranges in the three lhpper zones of the Late Scythian.
Symbols used in the four right hand columns are as follows: X = present only at this hori-
zon, = PRESENT IN BOTH YOUNGER AND OLDER HORIZONS, e = PRESENT ALSO IN EARLIER HORIZONS,
1 = PRESENT ALSO IN LATER HORIZONS.
Total
Genera
Tethys W. Pacific E. Pacific Arctic
X
Trohungarites
65
57
25
25
16
42
20
1
2
Coliimbites
21
8
15
6
2
2
7
16
Owenites
58
34
43
32
14
38
6
1
13
centers on the Tethyan faunas proceeding
from west to east; then the areas in the
western Pacific belt are discussed; this
leads to the circum-Arctic region, and
finally to the east Pacific localities. For
each locality that has previously been mono-
graphed the original faunal list is given
plus a summary list of the species I recog-
nize based on the discussion in the sys-
tematic portion of this report.
Albania
One of the most abundant and diverse of
late Scythian faunas is that from Kcira,
Albania. This fauna, originally collected
by Dr. Franz Baron Nopcsa and studied
by G. Arthaber (1908, 1909, 1911), occurs
in a one meter bed of red nodular lime-
stone. There are no other Scythian faunas
known in the sequence of strata at Kcira.
Arthaber recognized in this fauna 59
species, placed in 30 genera; a list of these
species is as follows:
Pseud osageccms drincusc Arthaber
Sageceras alhaniciim Arthaber
Pronoritcs triadiciis Arthal)er
Pronoritcs osmanictis Artliaber
Pronorite.s arhamis Artliaber
HcdcnstrocDiid kastriutae Arthalier
Hcdenstroeiu ia .■■iki])ciar('nsis Arthaber
Bcatites hcrthac Artlial)er
Procarnitcs kokciii Arthaber
Procarnitcs skundcrhcgis Artliaber
Paranannitcs nieditcrrciiiciis Arthaber
Proptychites latifituhriatus de Koninck
Proptychitt'.s krafffi Arthaber
Proptyciiitcs liigoiidlis Arthaber
Proptychilcs hertisci Arthaber
Proptychites ohliqucplicutus Waafien
Xenodiscus suliotictts Arthaber
Xenaspvi mcditerranca Arthaber
J(i])onitcs sugriva Diener
Monophyllites dieneri Arthaber
MonophylUtes pitamaha Diener
Monophyllites kingi Diener
Monophyllites hara Diener
MonopJiyllitcs nopcsai Arthaber
Lecanites skutarensis Arthaber
Lecanites fishtae Arthaber
Lecanites niazi Arthaber
Lecanites discus Arthaber
Ophiceras sakuntala Diener
Ophiceras efr. nangaensis Waa^en
Dagnoceras nopcsanum Arthaber
Dagnoceras zappanense Arthaber
Dagnoceras terbunicum Arthaber
Dagnoceras konianuni Arthaber
Dagnoceras lejanutn Arthaber
Meekoceras radiosuin Waasen
Meekoceras skodrensc Arthaber
Meekoceras hakki Arthaber
Meekoceras nialioinedis Arthaber
Aspidites hasserti Artliaber
Aspidites marginalis Arthaber
Tirolites illyricus Mojsisovics
Tirolites rectangtdaris Mojsisovies
Prosphingites alt Arthaber
Pseudosihirites efr. dichotonius Waagen
Protropites hihni Arthaber
Prenkites malsorcnsis Arthaber
Isculites originis Arthaber
Styrites lilangensis Diener
Columhites europaeus Arthaber
CoUunhites perrinisniithi Arthaber
Cohunhites chisniani Arthaber
Cuhnnhites niirditensis Arthalu'r
Arianites musacchi Arthaber
Paragoceras dtikagini Arthaber
Celtites aniauticus Arthaber
Celtites kcirensis Arthaber
l\piceltites gentii Arthaber
(?) Tropiceltites praeniaturus Arthaber
What specimens remain of the Kcira col-
lection studied by Arthaber are deposited
in the Paleontological Institute, University
of Vienna. This collection now consists
mainly of the illustrated specimens. The
fate of the nuun- unfigured parat\'pes is
Ammonoids of the Late Scythian (Lower Triassic) • Kummel
339
Figure 1. Index map of localities where faunas of Prohungarifes Zone age fiave been reported. (1) Upper Thaynes For-
mation, southeastern Idaho; (2) Upper Thaynes Formation, west-central Utah; (3) Tobin Formation, Tobin Range, Ne-
vada; (4) Subco/umbifes fauna. Providence Range, southeastern California; (5) Humboldt Range, Nevada; (6) Toad-
Grayling Formation, northeastern British Columbia; (7) Upper Scythian of Ellesmere Island and Axel Heiberg Island; (8)
Spitsbergen; (9) Olenek-Lena River Basin, Siberia; (10) Okhotsk-Kolyma Land, Siberia; (11) Primorye Region around Vlad-
ivostok; (12) Osawa Formation near Sendai, Japan; (13) south Ofago, South Island, New Zealand; (14) Prohungarifes fauna,
Nifoekoko, Timor; (15) Subco/umbifes fauna, Kwangsi, China; (16) Prohungarifes fauna, Kashmir, Himalayas; (17) Narmia
Member, Mianwali Formation, Salt Range and Surghar Range, West Pakistan; (18) Subco/umbifes fauna, Kotal-e-Tera, Afghani-
stan; (19) Tyur-Upa Suite, Mangyshlak Peninsula, Caspian region; (20) Subco/umbifes fauna of Chios; (21) Subco/umbifes
fauna of Albania; (22) Tiro/ifes fauna of Campil Member of Werfen Formation.
unknown, though part or all of them may
be in the collection obtained by the British
Museum (Natural History) in 1922, by
purchase. All of the existing specimens
from Albania in these two institutions have
been thoroughly studied and are discussed
in the systematic part of this paper.
Arthaber's illustrations of his species are
highly retouched photographs and in many
cases quite misleading. Likewise, many of
his suture drawings were not as successful
as one would like. Unretouched photo-
graphs of the primary types and new draw-
ings of many sutures are presented in the
systematic chapter.
My own studies of this fauna lead me to
conclude that it consists of 32 species,
placed in 27 genera. A summary list of
the species recognized is given on Table
18. I was unable to evaluate four of Artha-
ber's species: Lecanites fishtae, Lccanites
niazi, Meekoceras skodrense, Aspidites mar-
ginalis. I consider these as unrecognizable
species, and thus they are not treated in
the taxonomic summary of the fauna.
Chios
The largest and most diverse of any late
Scythian ammonite fauna is that from the
island of Chios collected by Dr. C. Renz
and associates. The collection consists of
nearly 2,000 specimens and is deposited
in the Natural Historv Museum, Basel. The
ammonites come from hard, red, siliceous
limestone; there are no other Scythian
faunas known from Chios. The fauna is
almost identical to that from Kcira, Albania,
which likewise occurs in a hard, red,
siliceous limestone. Renz and Renz ( 1947,
1948) recognized 116 species, subspecies,
340
Bulletin Muscidu of Cotnparativc Zoology, Vol. 137, No. 3
Table 18. Summary list of species recognized in this report from the Subcolumbites faunas
OF Albania and Chios.
Alba-
nia Chios
Alba-
ma
Chios
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Fseudosagcceras mult iJobat urn Noetling
Pseiidosageceias drincnse Arthaber
Pseudosageceras alhanicum ( Arthaber )
Tseudosageccras pasquayi Renz and Renz
Cordilleritcs angulatus Hyatt and Smith
Dieneroceras mcditcrranea (Arthaber)
Diencroceras skutarensis (Arthaber)
Siihvi.'ihnuitcs cnveris (Arthaber)
Heniilecanites discus (Arthaber)
Preflorianitcs suliotictis (Arthaber)
Prcflorianitcs garbintis (Renz and Renz)
Proptycliitoidcs dccipiens Spath
Propfychitoides trigonalis (Arthaber)
Procarnitcs kokcni (Arthaber)
AniaiitoccJtitcs ineditcnaueiis (Arthaber)
Prosphingites «// Arthaber
Zcnoiics helcnac Renz and Renz
Zenoites vondcrsclunitti
(Renz and Renz)
Iscidifoides originis (Arthaber)
Cliiotites glohidaris Renz and Renz
Tunglanitcs alexi n. sp.
Subcohnnbites perrinismithi (Arthaber)
Subcohiinhitcs diismani (Artliaber)
Vickoldc'iitcs sundaicus (Welter)
Prenkitcs inalsorensis (Arthaber)
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Prenkites timoreusis Spath
Prcnkites helenae Renz and Renz
Proiropites hihni Arthaber
Chioceras mitzopouloi Renz and Renz
Chioceras nodosum Renz and Renz
Arianitcs musacchi Arthaber
McropcUa plcjanac Renz and Renz
Epicclfites genti Arthaber
MctaJicdenstroemia kastriotac (Arthaber)
Beatites hcrthae Arthaber
Dagnoccras nopcsaniim Arthaber
Dagnoceras zappancusc Arthaber
Metadagnoccras terbunicum (Arthaber)
Albanitcs triadicus (Arthaber)
Sibiritcs rcnzi n. sp.
Tirolites idrianus (Hauer)
Dinarites dahnafinus (Hauer)
Dinaritcs hatsikasi Renz and Renz
Hcllcniies pracmaturus (Arthaber)
Hellenites radiatus Renz and Renz
Beyrichites laurae Renz and Renz
Eogymnitcs arthabcri (Diener)
Eophyllitcs dicncri (Artliaber)
Palacophyllites steinmanni Welter
Leiophyllites variabilis Spath
and varieties placed in 38 genera and sub-
genera; a list of these taxa is as follows:
CoJumbitcs curopacus Arthaber
Cohnnbitcs peninismitJii Arthaber
Columbites europacusperrinisniithi Renz and Renz
Columbitcs niiiditcnsis Arthalser
CoIu))}bit('s dianae Renz and Renz
Columbitcs dianae var. Renz and Renz
Coluudjitcs parisianus Hyatt and Smith
Columbites .opened Smith var. cliiotica Renz and
Renz
Columbites ex aff. plicatuli Smith
Columbitcs malayamis C. Renz
Columbitcs utahujanus var. C. Renz
Columbites malayanus C. Renz var. crassa Renz
and Wvnx
Columbitcs bubulinae Renz and Renz
Columbites graecoamericcmus Renz and Renz
Colut)d)itcs Icvaulinjis Renz and Renz
Columbitcs aithaliac Renz and Renz
Columbites hellenicus Renz and Renz
Prcnkites ))udsorensis Arthaber var. Renz and Hcnz
Prcnkites .sundaicus Welter
Prenkites helenae Renz and Renz
ef. Styrites lilangensis Diener
Iscultitcs originis Arthaber
Iscultitcs globulus Renz and Renz
Iscultites globulus var. Renz and Renz
Lscidtitcs antiglobulus Renz and Renz
Iscultitcs antiglobulus var. Renz and Renz
Iscultitcs globidusoriginis Renz and Renz
Iscultites globulusant iglobidus Renz and Renz
Anasibirites aff. anguloso (Waagen)
Chioceras mitzopouloi Renz and Renz
Chioceras mitzopouloi \ar. mcridionalis Renz and
Renz
Chioceras nodosum Renz and Renz
Prosphingites ex aff. czekanowskii Mojsisovics
Prosphingites rondcrschmifti Renz and Renz
Prosphingites (Cliiotites) globularis Renz and Renz
Prosphingites (Chiotites) supcrglobosus Renz and
Renz
Prosphingites (Zenoites) helenae Renz and Renz
Prosphingites (Zenoites) Julcnac \ ar. maradovunen-
sis Renz and Renz
Ccltitcs kcirensis Artliaber
I'lpiccltifcs gcntii Arthaber
llcllcnitcs i>racnuiturus (Arthabcn)
Ilcllcnitcs pracnuiturus \ar. aegaeiea Renz and
Renz
Ilcllcnitcs trikkiiliiu>i Ren/ and Renz
Ammonoids of the Late Scythian (Lower Triassic) • Knmmel 341
UeUcnitcs trikkalinoi var. Renz and Renz
Hellenites trikkalinoi var. ^raeca Renz and Renz
Hcllenites ( Pallasiics ) radiatus Renz and Renz
HcUcnites (Palhsites) striatus Renz and Renz
Hellenites (Pallasites) striatus var. densicosiata
Renz and Renz
Dinarites ntidus Mojsisovics
Dinaritcs evohitior Kittl
Dinarites liatsikasi Renz and Renz
Stacheites dionysi Renz and Renz
Stacheites dionysi var. Renz and Renz
Dagnoeeras terbuniciim Arthaber
Dagnoceras nopcsanuni var. involute Renz and
Renz
Meekoceras cf. gracilitatis White
Inyoites garhinus Renz and Renz
Ophieeras cf. demissum Oppel
Flemingites pseudorusselli Renz and Renz
Lecanites skutarensis Arthaber
Lecanites discus Arthaber
Xenodiscus sulioticus Arthaber
Koninckites bernoulii Renz and Renz
Koninckites bernoidlii var. Renz and Renz
Koninckites timorensis Renz and Renz
Beyrichites praematurus Renz and Renz
Beyrichites latirae Renz and Renz
Proptychites mohamedis (Arthaber) var. applanata
Renz and Renz
Proptychites ktenasi Renz and Renz
Proptychites arthaberi Welter
Pro])tychites balcanicus Renz and Renz
Proptychites mistardisi Renz and Renz
Proptychites lawrencianus (de Koninck) mut.
postindica Renz and Renz
Proptychites buxtorfi Renz and Renz
cf. Nannites hii^dostanus Diener
cf. Nannites medius Krafft and Diener
Paranannites aspenensis Hyatt and Smith var.
europaea Renz and Renz
Paranannites mediterraneus Arthaber
Paranannites mediterraneus var. media Renz and
Renz
Paranannites chionensis Renz and Renz
Paranannites compresstts Renz and Renz
Monophyllites {Leiophyllites) praeconfucii Renz
and Renz
Monophyllites {Leiophyllites) georgalasi Renz and
Renz
Monophyllites (Leiophyllites) rosae Renz and
Renz
Monophyllites (Leiophyllites) dieneri Arthaber
var. involuta Renz and Renz
Monophyllites (Leiophyllites) palaeotriadicus Renz
and Renz
Monophyllites (Leiophyllites) aff. pitamaha Diener
Monophyllites (Schizophyllites) betilloni Renz and
Renz
Monophyllites (Schizophyllites) betilloni var.
evoluta Renz and Renz
Monophyllites (?Schizophyllites) pseudohara Renz
and Renz
Monophyllites (Palacophyllites) thalmanni Renz
and Renz
Monophyllites ( Palacophyllites) praekieperti Renz
and Renz
Procarnites kokeni Arthaber
Procarnites kokeni var. Renz and Renz
Procarnites kokeni var. evoluta Renz and Renz
Procarnites kokeni var. panteleimonensis Renz and
Renz
Procarnites skanderbegis Arthaber
Hedenstroemia pityaussae Renz and Renz
Pronorites triadicus Arthaber var. Renz and Renz
Pronorites arbanus Arthaber
Pronorites arbanus Arthaber var. Renz and Renz
Pronorites arbanus Arthaber var. mediterranea
Renz and Renz
Pronorites orientalis Renz and Renz
Pronorites cf. ostnanicus Arthaber
Pronorites scluiubi Renz and Renz
Pronorites schaubi var. kephalovunensis Renz and
Renz
Pronorites reicheli Renz and Renz
cf. Cordillerites angidatus Hyatt and Smith
Pseudosageceras cf. claviscllatum Diener
Pseudosageceras intermontanum Hyatt and Smith
Pseudosageceras drinense Arthaber
Pseudosageceras drinense var. incentrolata Renz
and Renz
Pseudosageceras (Metasageceras) pasquayi Renz
and Renz
Sageceras albanicum Arthaber var. Renz and Renz
Arianites (Meropellu) plejanae Renz and Renz
cf. Paragoceras dukagini Arthaber
My own study of the Renz collection in
the Natural History Museum, Basel, leads
me to conclude that the fauna consists of
41 species placed in 27 genera. A summary
of my taxonomic conclusion is given on
Table 18. The illustrations of this fauna in
the Renz and Renz ( 1948 ) monograph are
excellent and thus need not be reproduced
here.
The Subcolumbites faunas of Chios and
Albania contain a few specimens that are
"nonnal" Anisian species. In the Chios
fauna there is Beyrichites laurae, recognized
on the basis of three specimens, and in the
Albanian fauna there is Eogymnites artha-
beri. The preservation in red limestone and
the distribution of the fossils in lenticular
"pockets," at least as far as the Chios fauna
is concerned, does raise a question of pos-
sible mixing. Renz and Renz (1948:61)
342 Bulletin Museum of Comparaiivc Zoology, Vol 137, No. 3
recognized the unusual aspect of Beijrichites
in their Chios fauna but came to the con-
ckision that the faunas were not mixed.
The fact that Beijrichites i^ represented by
only three specimens tends to support their
thesis. On the basis of the data available
I have elected to accept Beyrichites as a
valid member of the late Scythian am-
monoid faunas.
Werfen Formation
Tlie ammonite fauna of the Werfen For-
mation of the Alps and associated regions,
characterized by the great abundance of
Tirolitcs, has been an enigma among
Scythian faunas. It is a unique fauna quite
unlike that of any other of the Scythian.
This uniqueness is the cause of an array
of conflicting opinions as to its age. A re-
view of the conclusions on the age of this
fauna by such pioneer students as Moj-
sisovics, Waagen, Diener, etc. cannot serve
any useful puipose here. It was J. P. Smith
( 1932 ) who provided the basis for the
prevalent current interpretation of the age
relations of the Tiwlitcs fauna. Within the
Thaynes Fonnation cropping out in Paris
Canyon, southeast Idaho, Smith encoun-
tered a poorly preserved fauna 225 feet
above the Meelcoceras fauna and 30 feet
below his Columhites fauna. Within this
fauna, Smith (1932:11) recognized four
species of ammonites: Dalmatites at-
tenuottis, Tirolites harti, T. knia^liti, and
T. peali. On the age and correlation of this
small fauna Smith (19.32:10) concluded:
"Tirolites- constitutes the most abundant
and characteristic element of the fauna,
with species closely allied with those of
the Campil beds of the Tyrol, giving a
definite correlation and marking the first
appearance of Mediterranean types in the
American Lower Triassic." Smith thus rec-
ognized a Tirolites Zone stratigraphically
between his Meekoceras Zone below and
the Cohimhites Zone above. This scheme
was also adopted by Spath (19.34:27), and
in the Treatise (Kummel in Arkell, et al.,
1957). The age relations of the so-called
Tirolites fauna thus hinged on the strati-
graphic position of a small lot of poorly
preserved ammonites from a single locality
and horizon in southeastern Idaho. Need-
less to say, this was a most unsatisfactory
situation.
The discovery and study of a number of
new upper Scythian faunas in the past
couple of decades has contributed many
new faunal and stratigraphic data that war-
rant a new evaluation of the Werfen fauna.
The most comprehensive previous analysis
of the Werfen fauna was that by Kittl
( 1903 ) who recognized 59 species, listed
below.
Diiiaiites Jacvis Tommasi
Dinaritcs mticliiamis (Hauei)
Dinarites evolutior Kittl
I^inaritcs hianpulatus Kittl
Dinaritcs niidiis Kittl
Dinaritcs dalmatimis (Hauei)
Dinaritcs nuiJticosiatus Kittl
Dinaritcs tirolitoidcs Kittl
Dinaritcs ?anp.ulatus Kittl
Dinaritcs (Hercegovitcs) mohamcdantis Mojsisovics
Dinaritcs (Hcrccp.ovitcs) dioclctiani Kittl
Dinaritcs (Liccaitcs) circmnplicatiis Mojsisovics
Dinaritcs (Liccaites) connectcns Mojsisovics
Dinaritcs (Liccaitcs) Jiccanus (Hauer)
Dinaritcs (Liccaitcs) pro<ircsstis Kittl
StacJwites prionoidcs Kittl
Ccratitcs (Paraccrafitcs) prior Kittl
Tirolites (HoIoIoJjus) monoptychns Kittl
Tirolites carniolicus Mojsisovics
Tirolites serratelohatiis Kittl
Tirolites idrianus (llaiicr)
Tirolites hetcrophanus Kittl
Tirolites mcrcurii Mojsiso\ics
Tirolites paucispinatns Kittl
Tirolites scminiidtis Mojsisovics
Tirolites disians Kittl
Tirolites (jucnstcdti Mojsisovics
Tirolites rohtistiis Kittl
Tirolites diinidiatns Kittl
Tirolites stachci Kittl
Tirolites dinanis Mojsisinics
Tirolites Jujhridus Kittl
Tirolites angnsfns Kittl
Tirolites sid}ilhjrieti.s Kittl
Tirolites illyrieits Mojsiso\'ics
'Tirolites rc))idstis Kittl
Tirolites rotiformis Kittl
Tirolites rccfanfitdaris Mojsisox ics
Tirolites uudidalus Kittl
Tirolites (mgustilolxilns Kittl
Tirolites cassiantis (Quciistedt)
Tirolites xpinosus Mojsisovics
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 343
Tirolites haueri Mojsisovics
Tirolitcs inultispinattis Kittl
Tirolites percostatus Kittl
Tirolites tiirgidus Mojsisovics
Tirolites daruini Mojsisovics
Tirolites spinosior Kittl
Tirolites smiriagini ( Auerbach )
Tirolites kerneri Kittl
Tirolites toulai Kittl
Tirolites {Svilajites) cingulatus Kittl
Tirolites { Svilajites) tietzei Kittl
Tirolites {Bittnerites) malici Kittl
Tirolites {Bittnerites) hittneri Kittl
Tirolites (Bittnerites) ?telleri Kittl
Kyniatites svilajanus Kittl
Meckoceras caprilense Mojsisovics
Diilmatites morlaccns Kittl
I have had the opportunity of examining
the very hirge collections of Werfen am-
monites that constituted the basis for Kittl's
monograph. All of Kittl's taxa are treated
in the systematic portion of this report.
My own analysis of this fauna leads me to
believe it consists of only 13 species in-
cluded in 9 genera. A summary list of these
species is as follows:
Dahnatites morlaccns Kittl
Stdcheites prionoides Kittl
Dinarites dalmatinns (Hauer)
Dinarites carniolicus (Mojsisovics)
Diuplococeras liccannm (Hauer)
Diaplococeras connectens (Mojsisovics)
Psendokymatites svilajanns (Kittl)
Psendodinarites mohamedantts ( Mojsisovics)
Bittnerites hittneri Kittl
Hololohns monoptychns (Kittl)
Tirolites idrianus (Hauer)
Tirolites cassianiis ( Quenstedt )
Tirolites cingnlatns Kittl
The species of Tirolites completely domi-
nate the fauna, being represented by several
hundred specimens. Dinarites is also repre-
sented by many specimens but still con-
siderably less than specimens of Tirolites.
The other species are represented by very
few specimens. Stacheites prionoides, Pseti-
dokijmatites svilajanus Kittl, and Hololohm
monoptijchus are represented only by a
single specimen each. Diaplococeras lic-
canum is represented by 2 specimens, Bitt-
nerites hittneri by 5 specimens, Psendo-
dinarites mohamedaniis by 6 specimens,
Dahnatites morlaccus by 10 specimens, and
Diaplococeras connectens by 11 specimens.
Species of Tirolites and Dinarites include
approximately 95 percent of all the speci-
mens in this collection.
Recently, Ganev ( 1966 ) has described
a small fauna from the Campil beds of
Bulgaria, recognizing the following species:
Lanceolites discoidalis Ganev
Balkanites tahulatus Ganev
Tirolites hispinatns Ganev
Dinarites niuchianns (Hauer)
Dinarites progressus Kittl
Through the courtesy of Dr. Ganev, I
have photographs and plaster casts of the
specimens illustrated in his paper. This is
the first record of Lanceolites outside of
North America where the type species of
the genus is a prominent member of the
Meckoceras fauna. Balkanites is a new
genus known only from Bulgaria. The
specimens assigned to Tirolites hispinatus
I believe to be Tirolites cassianus, those
assigned to Dinarites progressus belong in
Diaplococeras connectens; and those as-
signed to Dinarites muchianus belong in
Dinarites dalmatinus. It appears from
Ganev's (1966) paper that he had only 10
specimens in his fauna.
As stated above, the age assignment of
the Werfen fauna arrived at by Smith rests
almost entirely on the position of the
Tirolites fauna in the Thaynes Formation
exposed in Paris Canyon, southeast Idaho.
An evaluation of all genera and species in
the Werfen fauna suggests a quite different
age assignment. The Werfen fauna in-
cludes only 11 genera of ammonoids, but
6 of these ( Diaplococeras, Hololohns, Bitt-
nerites, Psendodinarites, Balkanites, and
Psendokymatites) are endemic to the
Werfen Formation. The species of these
endemic genera are represented by only
27 specimens. The morphological charac-
teristics of these genera are not of any
particular help in establishing an age for
the fauna. Dalmatites is represented in
the Tirolites fauna of southeast Idaho (D.
attenuatus) and in the overlying Columhites
fauna (D. kittU n. sp.). Smith (1932:81)
described Dalmatites richardsi from the
344 Bulletin Museum of Comparative Zoology. Vol. 137, No. 3
Meekoceras fauna of southeast Idaho.
These are the only species of this genus
recorded to date. I agree with Spath ( 1951:
20) that Dalmatites ropini Diener (1907:
93, pi. 9, figs. 5, 6) of Anisian age is
generically distinct from the Scythian
species mentioned above. Lanceolites was
previously known only from two species in
the Alcckoccras limestone of western United
States. Tlie genus Stacheites is now known
from four additional localities. Astakhova
( 1960b ) records Stacheites prionoicles from
the uppermost of her faunal horizons in the
Scythian formations on the Mangyshlak
Peninsula. In fact, she named her upper-
most zone the Stacheites Zone but did not
describe or illustrate her specimens. Kum-
mel ( 1966 ) recorded a poorly preserved
specimen as Stacheites sp. indet. from the
uppermost fossiliferous horizon of the
Scythian strata in the Surghar Range of
West Pakistan in a rock unit also containing
Prohuniiarites sp., Procamitcs kokeni,
Dag^nuceras cf. zappanensc. In the syste-
matic portion of this report Stacheites
floweri n. sp. is described from the Tobin
Formation of Nevada where it is associated
with Suhcolumhites, Hemilecaniies, Isculi-
toidcs, etc. An indeterminate species of
Staclieitcs from the Prohun^arites fauna of
the upper Thaynes Formation of southeast
Idaho is described in the systematic portion
of this paper. These four additional records
of Stacheites are clearly from horizons
assignable to the late Scythian Prohtin<^ai-
ites Zone.
Species of Dinarites are the second most
abundant form represented in the Werfen
fauna. Fortunately, species of this genus
are now known from the Suhcolumhites
fauna of Chios and from the Scythian l"or-
mation of the Mangyshlak Peninsula. The
Chios specimens that Renz and Renz
(1948) assigned to Dinarites nudus Moj-
sisovics and Dinarites evohitior Kittl are
here considered more properly assigned to
Dinarites dahnatinus. A third dinaritid in
the Chios fauna, Dinarites liatsikasi Renz
and Renz, is unique because of its close
similarity to Dinarites undatus Astakliova
from the Mangyshlak Peninsula. Dinarites
undatus is recorded from the Tirolites Zone
of Astakhova, lying above beds containing
Procamitcs kokeni and Prohungarites cari-
natus and beneath horizons containing
species of Alhanites, Olenekites, Epiceltites,
etc.
Finally, there is the genus Tirolites, the
dominant element in the Werfen fauna
and the genus Smith ( 1932 ) relied upon,
almost entirely, in correlation of his Tiro-
lites Zone. In concluding that Tirolites had
a very narrow stratigraphic range. Smith
( 1932 ) ignored the species that Krafft and
Diener (1909) described from the Heden-
stroemia fauna of the Himalayas (T. in-
jucundus), the specimens described by
Arthaber (1911) from the Suhcolumhites
fauna of Albania, or his own record ( Smith,
1932) of a species of Tirolites from his
Columhifes fauna. In addition, Popov
( 1961 ) has described tirolitids from late
Scythian strata in northern Siberia, and
Silberling (in Hose and Repenning, 1959)
records tirolitids from a late Scythian
horizon in western Utah. The genus
Tirolites is the dominant element only in
the Werfen Formation fauna; in all other
recorded occurrences the genus is repre-
sented by relatively few specimens. The
age span of the genus now clearly encom-
passes all of the upper half of the Scythian.
In summary, the Werfen Formation
fauna contains 11 genera, 6 of which are
endemic. One genus (Dalmatites) is known
from the Meekoceras and Columhites
faunas, another (Lanceolites) is known
from tlie Meekoceras fauna. Two genera
(Stacheites and Dinarites) are known from
outside of the Mediterranean region by
species from late Scythian faunas of Pro-
hungarites Zone age. It thus appears to be
imsound to rely on Tirolites to establish
the age of the Werfen favma; for this, much
greater reliance can be put on Stacheites
and Diiuirites. This leads to the conclusion
that the Werlen fauna is of late Sevthian,
Prohuniiarites Zone age.
Ammonoids of the Late Scythian (Lower Triassic) • Ktimmel 345
Mangyshlak Peninsula
The presence of Scythian fonnations on
the Mangyshlak Peninsnla has been known
for the better part of this centuiy. In
spite of several investigations of tliese strata
there prevails considerable controversy
over the stratigraphy and correlation of
most of the units. M. V. Bajarunas (1936)
was the first to present a stratigraphic
sequence with a list of the ammonites pres-
ent, which unfortunateh' included many
nomina nuda. According to Bajarunas, the
lowest 15 m of the Triassic section consist
of limestones and marls which contain
Doricranites bogdoanus v. Buch, D. rossicus
Mojsisovics, D. acutus Mojsisovics, and
Siibdoricranitcs discoidcs (nomina nuda
for both genus and species ) . It is of interest
to note Bajarunas's comment that this fauna
was characterized by numerous individuals
but few species. Above this lower fossilif-
erous unit are 100 m of unfossiliferous
siliceous shales. These are followed by
200 m of marly shales with seams of marly
limestone and calcareous marly concretions,
rich in ammonites. This series of strata is
divisible into four units; the lowest part,
of 23 m, contains Ophiceras cf. detnissum,
Xenodiscus sp., Pscudosagcceras multi-
lobatiwi Noetling, Neotoceras niokrinskii
Bajarunas (both genus and species nomina
nuda). The next 30 m are chiefly charac-
terized by Pseud osageceras muUilobatum
Noetling, Frocolumbites karatauciki Bajar-
unas (both genus and species nomina nuda),
Procamitcs andnisovi Bajarunas (nomina
nuda), Thcnnalites n. sp. and other species.
The third division, of 80 m, contains Co-
himbites cf. parisianus Hyatt and Smith,
C. asiaticus Bajarunas (nomina nuda), C.
dohmpcnsis Bajarunas (nomina nuda), C.
adai Bajarunas (nomimi nuda), C. ligati-
formis Bajarunas (nomina nuda), C. turur-
pensis Bajarunas (nomina nuda), C. gracilis
Bajamnas (nomina nuda), Tirolitcs n. sp.
and others (unnamed). The uppermost
division, of 65 m, is characterized by several
forms of Tirolitcs and Dinarites.
On the basis of this faunal sequence.
Bajarimas (1936) came to the conclusion
that since Doricranites lay below beds con-
taining Pscudosagcceras and Ophiceras, the
Doricranites strata must be correlative with
the Otoccras beds of the Himalayas, that
is, at the base of the Scythian. However,
the lack of any detailed discussion of these
faunas or any illustrations, plus the large
number of nomina nuda, are severe handi-
caps in fonnulating any judgment on the
basis of Bajarunas' conclusions.
Kiparisova (1947) briefly discussed tlie
Mangyshlak section and described some of
the ammonites, making specific note of the
fact that detailed stratigraphic data were
lacking for much of the material available
to her. However, she did describe from the
upper unit ( the 80 m unit below the upper-
most division of 65 m with Tirolitcs and
Dinarites) of Bajarunas' section Columbites
doJ)iapacnsis Kiparisova ( = C. dohuipaen-
sis Bajarunas MS), Kashmirites subdimor-
phus Kiparisova, Anasibirites gracilis Ki-
parisova ( = Columbites gracilis Bajarunas
MS ), and Tirolitcs rossicus Kiparisova.
In the volume on the stratigraphy of the
USSR, Kiparisova (1958a) contributed
some additional data on the Mangyshlak
sequence and the faunal associations. She
states that the sequence begins with up to
250 m of calcareous shales of which the
lower 10 m consist of sandy shale and coarse
sandstone with lenses of conglomerate.
Above the lower sandy bed, the shales con-
tain beds of limestone with abundant am-
monoids (Doricranites bogdoanus v. Buch,
Tirolites cassianus Quenstedt, Procarnites
andrusovi Kiparisova, Pscudosagcceras mul-
tilobatum Noetling, etc.) and pelecypods.
Above this are up to 400 m of argillaceous
and sandy shales with interbedded lime-
stone and sandstone. The limestones con-
tain ammonoids (Columbites cf. parisi-
anus Hyatt and Smith, Tirolites cassianus
Quenstedt, Procarnites andrusovi Kipari-
sova, Anasibirites gracilis Kiparisova, and
others) plus pelecypods, gastropods, and
brachiopods. Kiparisova placed these hori-
zons in the upper half of the Scythian.
346 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Table 19. Stratigraphy and faunas of Scythian strata on the Mangyshlak Peninsula after
AsTAKHOVA (1960 a, b; 1964). The species marked with an asterisk were only listed by Asta-
KHOVA; THEY WERE NEITHER DESCRIBED NOR ILLUSTRATED.
c
o
a
Stacheites
Zone
Columbites
Zone
u
0)
^
a
i
§
s
0)
^
-^
w5
,^
r-
(n
•— '
S
Tirolites
1— (
Zone
C/3
<
PL|
P
Pscudosageceras
>-l
Zone
H
S3
1
a
Doricranites
§
Zone
o
*J
S
Leiophijllites radians Astakhova
Nannites ])a jam nasi Astakhova
*'Paranannites aspenen.si.s Hyatt and Smith
'^Stacheites prionoides Kittl
Alhanites danispanensis Astakhova
Anasihirites suhgracilis Astakhova
Anasihirites gracilis Kipaiisova
*Olenekitcs tururpensis Astakhova
Olenekites niangyshlakensi^ Astakhova
Procuhnnhitcs karataiicikiis Astakhova
Columbites constrictilis Astakhova
*Columhites parisianus Hyatt and Smith
Columbites dolnapaensis Kiparisova
Dinarites undatus Astakhova
Kashmirites subdiniorphus Kipaiisova
Kashmirites contortus Astakhova
Tirolites impolitus Astakhova
Tirolites clegans Astakhova
* Tirolites smiriagini (Auerbach)
* Tirolites cassianus (Quenstedt)
* Tirolites spinosus Mojsisovics
'■'Tirolites rossicus Kiparisova
Procarnites andrusovi Kiparisova
"'Pscudosageceras multilobatum Noetling
Doricranites tu)nulosus Astakhova
Doricranites Innceolatus Astakhova
''Doricranites rarecostatus Astaklio\a
'^Doricranites discus Astakhoxa
Doricranites scharicus Astakhova
"'Doricranites ovatus Astakhova
Doricranites rossicus (Mojsisovics)
Doricranites bogdoanus (v. Buch)
Doricranites acutus (Mojsisovics)
Subdoricranites discoides Astakliova
Subdoricranites orbiculatus Astakhova
Kiparisovites carinatus Astakhova
Related to Bajarunas' interpretation of
the age of the Doricranites beds was the
conchision that tlie Perniian-Triassic forma-
tions on tlie Mangyslilak Peninsula were
gradational. In both these eonclnsions,
Bajarunas received support from Shevyrev
and Shle/inger (1960). These authors state
that throughout the entire extent of the
Kara-Tau Hange there are ru) signs, e\ en
the most indirect, of an interruption in
sedimentation or of a basal conglomerate.
They likewise are emphatic that the lower
horizon with the Duiicnniites fauna does
not contain Tirolites, as had been claimed
by some authors. The short paper by
Shevyrev and Shlezinger ( 1960 ) suffers
from the same vague and incomplete data
that characteri/(Hl the contributions of
HajariMias (1936) and Kijiarisoxa (1947,
1958a).
For the first comprehensive discussion of
the stratigraphy and faunas of the Scythian
formations on the Mang\'shlak Peninsula
we are indebted to Astakhoxa ( 1960a, b.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 347
Table 20. Summary List of Species Recog-
nized IX This Report from Each of the Zones
Established by Astakhova (1960 a, b) in the
Scythian of the Mangyshlak Peninsula.
Stacheites Zone
Columbites Zone
Tirolites Zone
Lciophyllitcs radians-
Arnautoceltites hajarunasi
Alhanites triadiciis
Epiceltites sid?pracilis
Olenekitcsmanp.yshhken.sis
Procohnnbites karataticikus
Pseudoceltites dolnapacnsis
Dinarites undatus
Eukashmirites
subdimorpJius
Eukashmirites contortiis
Tirolites rossicus
Tirolites impolitus
Pseudosaaeceras Zone Procarnites kokeni
Dorikranifes Zone
Prohungarites carinatus
Dorikranites hogdoaniis
Dorikranites acutus
1964 ) . This author considered the Scythian
strata, which she named the Tyur-Upa
Suite, to be transgressive on the underlying
Permian (Dohiapa Suite) fonnations. The
basal 6-10 m of strata are stated to contain
lenticular beds of conglomerate which in-
clude pebbles of the underlying red Per-
mian Dolnapa Suite. Astakhova recognized
a sequence of three lithologic members and
five faunal zones within her Tyur-Upa
Suite. These data are summarized on Table
19. The Doricranifcs Zone was correlated
with the "Meekoccras' beds of the Hima-
layas, the Primorye Region, and Timor.
The Pseuclosofieceras Zone was correlated
with the Hedenstrocmia beds of the Hima-
layas, the Fk'min<s,ifes beds of Timor, the
Flerningifes beds of the Primoiye Region,
and the Pseudosageceras beds of the west-
ern United States. The Tirolites Zone was
correlated with the Tirolites Zone of the
eastern Alps and of southeast Idaho. The
Columbites Zone was correlated with the
SuhcoJumhites fauna of the Primorye Re-
gion, the Olenekites fauna of northern
Siberia, and the Columbites fauna of south-
east Idaho. The Stacheites Zone was cor-
related with the Prohungarites Zone of
Spath (1934).
Astakhova thus concluded that her Tyur-
LTpa Suite included all of the Scythian ex-
cept for the lowest zone (Otoceras). Care-
ful analysis of the described and illustrated
species leads me to a quite different con-
clusion: that all of the Mangyshlak faunas
belong to a single zone, that of Prohun-
garites. In the systematic portion of this
paper each of the species from the Tyur-
Upa Suite is discussed. A summary of the
species that I recognize in each of Astak-
hova's zones is listed on Table 20.
Afghanistan
Excellent exposures of Lower Triassic
Scythian strata occur at Kotal-e-Tera, 90
kilometers southeast of Kabul. A well pre-
served and diverse Owenites fauna has been
described from this locality by Kummel
and Erben ( 1968 ) . A unique feature of
this Owenites fauna is its complete mixing
with a typical Anasibirites fauna. The beds
containing the Oicenites fauna are 75 feet
thick and are overlain by 14 feet of strata
that have yielded a poorly preserved Sub-
columbitcs fauna. Kummel (1968) has
recorded the following species from these
beds:
Pseudosageceras inultilobatum Noetling
Subvishnuites sp. indet.
Subvishnuites cf. cnveris Arthaber
Xenoceltites sp. indet.
Procarnites kokeni (Arthaber)
Isciditoides cf. originis (Arthaber)
Subcolumbites perrinismithi (Arthaber)
Vickohlerites cf. sundaicus (Welter)
MeropcUa cf. plcjanae Renz and Renz
Albanites triadicus (Arthaber)
Keyserlingites sp. indet.
Leiophyllites sp. indet.
These strata are overlain by black mud-
stones containing an Anisian fauna.
Salt Range, West Pakistan
The Ceratite beds of the Salt Range,
West Pakistan, occupy an important place
348 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
in the development of Lower Triassic
(Scythian) chronology. The geology of
the Salt Range and Trans-Indus ranges was
first monographed by Wynne ( 1878, 1880 ) ,
and the ammonites monographed by Waa-
gen ( 1895 ) . In establishing a standard
chronologic scheme for Triassic marine
facies, Mojsisovics, Waagen, and Diener
( 1895 ) proposed the sequence of zones of
the Salt Range as the type for the Lower
Triassic. At that time the Ceratite beds
were thought to comprise the entire Lower
Triassic. It has long been recognized that,
with a very few exceptions, the ammonoids
from the Salt Range described by Waagen
( 1895 ) came from zones representing only
parts of the middle and lower part of the
Scythian.
A complete restudy of the Triassic forma-
tions in the Salt Range and Trans-Indus
ranges has been published by Kummel
( 1966 ) . The Ceratite beds of Wynne and
Waagen have been named the Mianwali
Formation, including three members: Kath-
wai, Mittiwali, and Nannia. Nearly all of
Waagen's ( 1895 ) ammonoids came from
the Mittiwali Member. The Namiia Mem-
ber is approximately equivalent to the
Dolomite group of Waagen ( 1895 ) . From
this stratigraphic horizon, Waagen ( 1895 )
described a single ammonoid specimen as
Psciidharpoceras spinigcr. Waagen con-
sidered this species to be closely related to
the genus Tropites and concluded that it
indicated a late Triassic (Keuper) age. It
should be pointed out that Psetidharpoceras
spinigcr is based on a single specimen from
an unknown horizon, but thought by Waa-
gen to be from the topmost limestone unit
of his Dolomite group in the Sheik-Budin
Hills in the Trans-Indus Region. In addi-
tion, Waagen described Dinarites simiatus,
Lecanites Jaquetis, and Lecanites pkmorhis
from his Bivalve beds, the basal unit of the
Narmia Member.
Kummel's (1966) extensive field studies
of the Triassic formation in the Salt Range
and Trans-Indus ranges yielded a small,
and generally poorly preserved fauna in-
cluding the following species:
Pseiidosageceras miiltilohatiim Noetling
Suhvishniiitcs sp. indet.
Xenoceltitcs .siniiatiis (Waagen)
Xenoceltites sp. indet.
Procarnitcs kokeni (Arthaber)
I.sculitoidcs sp. indet.
Anakashmirites sp. indet.
SvaUiardiccras sp. indet.
Stachcitcs sp. indet.
Dagnoceras sp. indet.
Nordophiceras planorhis (Waagen)
Nordophiccras cf. planorhis (Waagen)
Arctoincckoccras sp. indet.
Tirolites sp. indet.
Prohungariics cf. crasscpJicatiis (Welter)
Indi
la
Scythian strata have been recognized in
the Himalayas for a century and have oc-
cupied an important role in the develop-
ment of Scythian paleontology and stratig-
raphy. However, most of the abundant
Scythian faunas known from Spiti to Kash-
mir belong to the lower half of that stage.
The youngest Scythian horizon known is
that of Sibiritcs spinigcr from Byans which
is equivalent to the Anasibiritcs Subzone
of the Owcnites Zone, approximately mid-
Scythian in age.
A late Scythian horizon is probably pres-
ent in Kashmir as shown by the presence of
Prohiingarifcs middlemissii (Diener, 1913).
This species is quite similar to Prohiin-
go rites mckclvei n. sp. from the upper
Thaynes Formation of southeast Idaho. The
genus Prohungarites is known from the
Mangyshlak Peninsula, the Salt Range,
Timor, Olenek region, Nevada, and south-
east Idaho in horizons here considered late
Scythian in age. The Kashmir species, how-
ever, was collected from loose blocks and
no stratigraphic data are available.
Recently, Tozer (1965a) has suggested
that the horizon of Kcyscrlingitcs dicneri in
the Himalayas may be upper Scythian in
age rather than lower Anisian, as concluded
by Diener, Spath, and odiers. The age
assignment of these beds has a sufficient
number of ambiguities to warrant a thorough
re-analysis.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel
349
Krafft and Hayden measured the follow-
ing section near Lilang in Spiti (Diener,
1912:56):
4. Dark grey limestone, often concretion-
ary with shaly partings. Upper
Muschelkalk 22 ft.
.3f. Grey limestone with Ceratites ravana 16 in.
3e. Grey concretionary limestone 6 in.
3d. Shales with Spiriferina stracheiji 4 in.
3c. Grey limestone 3 in.
3b. Hard, grey limestone with Kcijscr-
lingites dieneri 4 in.
3a. Thin layers of grey limestone and
shale 3 ft.
2. Nodular limestone ( Niti limestone of
Noetling) 60 ft.
1. Shaly limestone with RJn/iiclioiiclla
gricsbachi 3 ft.
Krafft measured the following section in
1900 at the Bambanag Cliff (Diener,
1912:57):
4. Upper Muschelkalk with numerous
specimens of Pti/rhitc.s, IloUandites,
Beijrichitcs khanikuffi, Gym nites
vasantascna 20 ft.
3i. Shales with many concretions, con-
taining Spiriferina stracheiji 2 ft.
3h. Dark grey limestone with Spiriferina
stracheiji and Spirigera stoliczkai 1 ft.
3g. Black shales 5 in.
3f. Dark grey limestone with Keijser-
lingites dieneri, MonophijUites hara,
M. kingi, Spiriferina stracluiji.
Spirigera stoliczkai 5 in.
3e. Black shale 2 in.
3d. Limestone as 3f containing Gijninites
sp. 7 in.
3c. Black shales 5 in.
.3b. Limestone as 3h with MonophijUites
sp. and Dahnatites ropini 6 in.
3a. Black shales with KeyserJingites sp. 5-6 in.
2. Nodular limestone ( Niti limestone of
Noetling) unfossiliferous 50 ft.
1. Earthy limestone with Rhynchonella
gricsbachi and Retzia himaica 3 ft.
Thus the basic pattern of this strati-
graphic interval is a thin limestone unit
with RJjyncJioncUa griesbadii, followed by
a thick nodular, essentially unfossiliferous,
limestone, then about six feet of limestone
and shale, with KeyserUngites dieneri, etc.,
and at the top limestones with an abundant
Anisian fauna. Diener (1912:55) empha-
sizes the homogeneity of this sequence of
facies between Spiti and Painkhanda. In
regards to the cephalopod faunas of the
Lower Muschelkalk, the sections at Spiti
have yielded a considerably larger number
of species than those of Painkhanda. A
tabulation of the species Diener (1907)
described from these Lower Muschelkalk
sections is given on Table 21. There are
only f(nu- species which are common to
the two districts. For some reason, Tozer
(1965a: 11) based his argument mainly on
the fauna from Bambanag Cliff; however,
if we accept the correlation of strata as
proposed by Diener, the large number of
unquestionable Anisian species from the
Spiti sections does not support a suggestion
that these strata and faunas could be late
Scythian in age. There is an additional bit
of evidence that lends support to Diener's
conclusions, and this is the faima from the
Middlemiss Crag near Chitichun. This is
a fauna from "exotic" blocks that has six
species in common with that of the Lower
Muschelkalk of Spiti, and two species in
common with the Lower Muschelkalk at
Painkhanda (Table 21). The presence of
such genera as Psilosttiriu, Procladisites,
etc., clearly indicates that this is Anisian in
age; however, KeyserJingites is not present
in this fauna.
Timor
Among the several beautifully preserved
faunas of Scythian age from Timor, only
one, that of "Block E bei Nifoekoko," is of
late Scythian age. Welter (1922) described
the following species from this fauna:
Cohimbites sp. ind.
Monophyllites sp. ind. ex aff. dieneri Arthaber
Palaeophyllites steinnianni Welter
ProptycJiites aiiJiaberi Welter
Hungarites cf. middlemissi Diener
Hiingarites crasseplicatus Welter
Hungarites tuberculatiis Welter
Pronorites arbanus Arthaber
Pronorites sp. ind. ex aff. arbani Arthaber
Welter ( 1922 ) noted that five of his nine
species were related to species of the Suh-
columhites fauna of Albania described by
Arthaber (1908, 1911). He, however, placed
the horizon of this fauna at about that of
350 Bulletin Miiscimi of Comparative Zoolofnj, Vol 137, No. 3
Table 21.
StrMMARY List of Species from the Lower Muschelkalk of the Himalayas, Tibet
AND Timor. Data from Diener (1895, 1907) and Welter (1915).
Gyundi Lilang, Po,
R., Spiti Spiti Spiti
Bam-
ShaLshal banag
Cliff Cliff
Middle- Timor Timor
miss (Welter (Welter
Crag Bed 2) Bed 3)
Hollandites vyasa Diener
Donithites kansa Diener
Danubitcs cunbika Diener
Daniihifes alternecostatiis ( Welter )
DaniiJntcs conipressus (Welter)
Keyscrlin^itcs dicncri Mojsisovics
Key.scrlin^ites pahari Diener
Key.serlinp.ife.s pagoda Diener
Kcyscrliufiites angiistecosiafus Welter
Japonitcs iigia Diener
Japonites meridianus Welter
Japonitc.s raphacli.s zojac Tonnnasi
Stacheitcs wcl)hianits Diener
Dahnatites ropini Diener
Sibil ites prahlada Diener
Sihirites pandya Diener
Gymnites depauperatus Diener
Gyrnnites volzi Welter
Procladiscites yasoda Diener
Psilosturia moiigolica (Diener)
Megaphyllifcs evolutm Welter
LeiophyUitcs ronfucii (Diener)
LeiopliyUites ))radytimna (Diener)
Leiophyllites iiiiddlemissii (Diener)
LciophyUites pitamaha (Diener)
Leiophyllites laevi.s (Welter)
Leiophyllites indoaustralica (Welter)
Ussurites hara (Diener)
Usstirites kingi (Diener)
Rommanites cf. simionescui Kittl
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
his Owenites limestone and approximately
mid-Scythian in age.
The British Museum (Natural History)
has large collections of Timor Scythian am-
monoids. In the l^ritisli Museum catalogue
of Triassic ammonoids, Spath ( 1934 ) de-
scribed a few new species from these blocks
at Nifoekoko which are characterized by
the manganese coating on the specimens.
Kummel (1968) has lik(>wise described new
species from this fauna. In this report the
species of ammonites recognized in the
Nifoekoko fauna (with manganese coated
specimens) are as follows:
Proptychitoidc.s arthuberi { Welter)
Procarnites kokeni ( yVrthalx'r )
Iseulitoidcs originis ( Arthab(M- )
Prenkites timorcnsis Spath
Dagnoceras zappaucn.se Arthaber
M etadagnoceras freemani Knninirl
Albanite.s triadicus (Arthaber)
Prohungarites cra.sseplicatu.s ( Welter)
Prohioxgarites tubereulatiis (Welter)
Eophyllites orientalis Spath
Pataeophyllitcs .stciniiianni Welti'r
Included as a member of the Frohun-
i^a rites Zone fauna of Timor is Prcnkitcs
.sinidaiciis- Welter. The one specimen on
which tills species is based came from Noel
Niti, Timor, but no identifiable associated
forms are known. This specimen has been
selected as the type of a new genus,
VickoJilcritcs Kummel (1968a).
Tozer (1965a: 12) has suggested that the
beds on Timor with Kci/.s-erliiu^ilci anp^iis-
tccostaitis Welter are possibly late Scythian
Ammonoids of the Late Scythian (Lower Triassic) • Kitmmel 351
in age. Tlie fact that the Triassic of Timor
is represented only by isolated blocks and
that extremely condensed sections (within
the blocks) are common has complicated
interpretation of many of the Timor faunas.
Most of the species that Welter (1915)
assigned to the Anisian came from a single
block of limestone within which three
distinct layers were recognized. One of
these layers, 60 cm thick, contained a
typical Anisian fauna of 8 species, including
Acrochordiceras (Pamcrochordiccras) an-
odosum Welter, Gymnitcs sp., etc. Adjacent
to this was a 30 cm thick unit with 14
species of ammonites which are listed in
Table 21 (Welter's bed 2). Four of these
species were first described from the Lower
Muschelkalk of the Middlemiss Crag of
Tibet, namely Japonites tigra Diener, Pro-
cladiscites yasoda Diener, Psilosturia mon-
p,oUca (Diener), and Us.siiritcs hara (Die-
ner). Two of these species were first
described by Diener from the Lower
Muschelkalk at Lilang, Spiti (beds which
contain KeyserUn<i,ites dicneri), namely
Japonites ii'^ra Diener and Ussurites hara.
Tozer (1965a: 12) agrees that this 30 cm
unit is "undoubtedly Anisian" in age.
Adjacent to this unit is a third, of 100 cm
in thickness from which Welter recognized
only three species, KeyserUngites angtisteco-
stattis, Ussurites hara (Diener) and Leio-
phyUites indoausirahca (Welter). It is this
fauna that suggests to Tozer the possibility
of an upper Scythian age on the basis that
the two associated species "do not establish
an Anisian age." However, Ussurites hara
(Diener) and LeiophyUitcs indoaustraJica
occur also in Welter's unit 2 and Ussurites
hara occurs in the Himalayas at Spiti,
Painkhanda, and at the Middlemiss Crag,
all units with an abimdant Anisian fauna.
Finally, among the specimens of Keyser-
Ungites angustecostatus Welter in the
British Museum, three fragments are from
a "large block with Gymnites and Leiophyl-
lites, etc., that also yielded Parasageceras"
Spath (1934:359). The facts regarding
these Timor faunas do not support the
suggestion that the 100 cm layer with
KeyserUngites angustecostatus may be late
Scythian in age.
New Zealand
Scythian ammonoids are extremely rare
in New Zealand where only two small
faunas have been discovered. The first of
these faunas consisted of only 9 specimens
placed in 4 species of Owcnites Zone age
(Kummel, 1959). The second fauna con-
sisted of 24 specimens of a single species,
Prosphingitcs coomhsi Kummel (1965).
The moiphological characters of this species
and its genetic relationships suggest that it
is of late Scythian {Prohungaritcs Zone)
age. Prosphingites coomhsi is extremely
close to Prosphingites insidaris Kiparisova
from the Suhcolumbites fauna of the
Primorye Region.
China
Our knowledge of the upper Scythian of
China is derived mainly from a monograph
by Chao ( 1959 ) on ammonite faunas from
Kwangsi Province. Chao adopted the strati-
graphic divisions of Spath (1934), and for
the Columbitan division he recognized
three zones:
Procarnites-LeiophylUtes Zone
Columbitan division Cohimbitcs- costatiis Zone
Tirolites dancini Zone
An analysis of all of the taxa recognized
by Chao in these upper Scythian zones is
included in the taxonomic portion of this
paper. The stratigraphic data are some-
what ambiguous but do not tend to support
Chaos interpretation.
The Tirolites Zone was recognized on the
basis of a single, poorly presei-ved frag-
ment of body chamber collected from an
isolated horizon 3 km southwest of Pachuan
in the Fengshan district, Kwangsi. The
identification of this ammonoid fragment
is highly dubious. This, coupled with the
complete lack of any associated fauna or
stratigraphic data, leads me to reject this
as a valid record of the so-called TiroUtes
Zone.
352 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
The remaining portion of Chao's Co-
lumbitan fauna is represented in three
sections in western Kwangsi — tlie NaHHng,
the Yali, and the Chashanao.
The Nahhng section is developed about
one kilometer northeast of Lolou village
and was divided by Chao (1959:158) into
three divisions. The lower unit is composed
of 15 m of black, thin bedded limestone.
The middle unit is composed of grey, thin
bedded limestone and calcareous shales,
about 40 m thick. The upper unit consists
of black, well bedded limestone 15-20 m
thick, but the upper part of this unit is cut
off by a fault. The lower unit contains an
Owcnites fauna. From the calcareous shale
beds of the middle unit, Chao (1959)
identified the following species ( collections
541a, b of Chao ) :
Prosphingites loloucnsis Chao
Meekoceras sp. indet.
Cohnuhitcs osymmetricus Chao
Columbitcs ? sp.
Isculitoides sp.
Di^it(>))]uiUitcs loloucn.sis Chao
Auuka-'iliniiiitcs ? sp.
From the upper unit, below the fault,
Chao (1959) recognized the following
species ( collections 542a of Chao ) :
Paianannites stibglobosiis Chao
Prospliiiigitcs invoJiitits Chao
C(ilui)i})it('s asynuuctriciis Chao
Prenkites kwangsicnsis Chao
Dognoceras cllipticiim Chao
Hcllcnitcs pracmatiirus (Arthabei)
Celtites sp.
Within the village of Lolou, Chao (1959)
uncovered an isolated limestone block from
which he identified the following species of
ammonites ( collection 542b of Chao ) :
Procariiilcs oxynostiis Chao
Procarniies acutus Spath
Cordillcritcs orienlalis Chao
Proptycliilvidc.s coDiprcssus (^hao
Tunglanites lenticularis Chao
ParaudnuUc.s siibglohosiis Cliao
Isculiluidas cUipticus CJhao
Isculitoides aff. ortginis ( Artliahcr)
Xcnoccltitcs crenovciitrosus C'hao
Xenoceltites comprcssiis Cliao
Leiophyllites oxynostiis Chao^
Leiophyllitcs lolouensis Chao-^
Leiophyllites serpcntintts Chao
Digitophyllites lolouensis Chao
Subnieckoceras compressinu Chao
Siibtueekoceras loloiiense Chao
Siibmeekoceras longiseptatum Cliao
Partissuria lafiJobafa Cliao
Anakashmiritcs aff. nivalis Dieiier
Lecanites sp.
The Yali section, in the Fenghan district,
is stated by Chao (1959:173) not to have
been well studied. It apparently repre-
sents a collection of ammonites (horizon
546 of Chao) from an unmeasured section.
Chao identified the following ammonites
from this section:
Proptychitoides ? simjdex Chao
Columbites yaliensis Chao
Coliimbites hiiangi Chao
Colinnbitcs eostattis Chao
Paianannites inculutiis Chao
Paranannites minutiis Chao
Prenkites kivangsianiis Chao
Fcngshanitcs robusttis Chao
Dagnoceras latilobatuni Chao
Hellenites pracniatitnis (Arthaber)
The Chashanao section lies on the border
of the Hochih and Tunglan districts. The
Scythian strata here consist of only 17 m of
strata resting unconformably on Lower
Permian limestones. Near the top of this
sequence a 0.6 m bed of black limestone
yielded the following ammonites, identified
by Chao ( 1959, his horizon 610) :
Subcolumbites kwangsianus Chao
C(>hi>u])ites hiiangi Chao
Isculitoides globosus Chao
Tunglanites lenticularis Chao
Paradinaritcs suni Chao
Anakashmirites sp.
Proptychitoides luughmcnsis Chao
Henilecanites discus Arthaber
Chao concludes that the fauna of his
collections 541a, b, 546, and 610 is es-
^ In the list of species iioin this eollection Chao
(1959:160) does not include these two species.
Instead, he has Leiophylliics ktc(ing.sien.sis Chao sp.
nov. and L. vermifonni.s Chao sp. nov.: however,
neither of these species is described in the taxo-
nomic portion of his nionoyraph. Tlic two species
listed aboNC arc from collection 5121).
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 353
Table 22. Stjmmary List of Species Recog-
nized IN This Report from the Late Scythian
OF KwANGSi, China, in Terms of the Five
Distinct Collections Studied by Chao (1959).
.a
j:
_0
<- ^
r- ^
Sc-l
0 ^H
5 CI
CD
^2
O lO
Oio
J— •
c»o
^co
0
0;
., 0
•2 c
"^ c
0 ^
1— o
bt-H
St-S
o
<-' 'X
O 1/
o *^
rt -J-"
.£ tl
.5 tj
:;; m
c/^ 9
-c i^
CS
— _a/
ffl s
V
2 —
'a'B
"^ '*'
v'"^
-— "7^
i:'c
ZU
:zu
J>
>^u
UU
Cordillcritcs angulatus
X
Xenoccltites crenoventrosus
X
Hemilecanites discus
X
Proptych itoidcs tunglanensis
X
X
X
Procarnitcs kokeni
X
Procaniites lolouensis
X
X
Amatitocchites involutus
X
Prosphinpitcs lolouensis
X
X
Prosp]}ingitcs subglobosus X X
Isctditoides ellipticus X
Tuuglanites lenticularis X X
Suhcohnubites perrinismithi XX X
Subcolumbites robustus X
Paradinarites suni X
Prenkites timorensis X XX
Parussuria lotilobata X
Dagnoceras eUipticinn X
Dagnoceras latilobatum X
Nordophiceras compressurn X
Uellcnitcs pracmaturus X X
LciopJujllites scipeutinus X
sentially the same and comprises his Zone
of Columhites costatus. Because a specimen
identified as Procarnites kokeni was found
from the top part of the Lower Triassic
Hmestone sequence east of Lolou, Chao
(1959:160) conchides that his collection
542b (the loose block) "may represent the
highest horizon of the Columhites stage"
and refers this fauna to his Procarnites-
LeiophyUites Zone.
All of the taxa described by Chao from
these sections are discussed in the taxonomic
portion of this paper. This study leads
me to conclude that there are only 21
species of ammonites in these four collec-
tions. Tliese are Hsted on Table 22 along
with their geographic occurrence. The
limestone block, collection 542b, which
Chao assigned to his Pwcarnites-Leiophyl-
lites Zone, comprises 11 species. Four of
these species occur in one or more of the
remaining three collections. Of the seven
remaining species, C o rd ill e rites angulatus
ranges throughout the upper half of the
Scythian; the genus Xenoceltites ranges
throughout the upper half of the Scythian
and is not common in late Scythian fonna-
tions; Procarnites kokeni is a common ele-
ment of the Subcolumbites fauna of Albania
and Chios, and it is also known from the
Salt Range and from the Prohungarites
fauna of Timor; the genus Isculitoides is
apparently confined to tlie late Scythian
and occurs in most localities where such
faunas are known; Parussuria latilobata is
the only species of this genus from a late
Scythian horizon; Nordophiceras is known
from the Olenckitcs fauna and Dieneroccras
fauna of northern Siberia, the Columhites
fauna of southeastern Idaho, and from the
Salt Range of West Pakistan; fomis like
Leiophyllites serpentinus are common in
the Suhcolumhites fauna of Albania, Chios,
and the Primorye Region.
I can see no actual difference in the
faunal composition of collection 542b and
collections 610, 542a, and 541a, b. Taking
into account the factors of preservation,
collection failure, and the composition of
other late Scythian faunas, as those at
Albania, Chios, Timor, Primoiye, etc., plus
the fact that there are no stratigraphic data
on collection 542b, these four collections
appear more likely to represent one single
zone.
Japan
The present data on Triassic stratigraphy
and ammonite faunas of Japan have been
very ably summarized by Bando ( 1964a,
1966). The upper Scythian is represented
by two specimens of Suhcolumhites per-
rinismithi from the Osawa Formation in
the Kitakami Massif,
Primorye Region
This is another of the classic areas that
have occupied an important place in Scyth-
354 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
ian studies going back to 1895 when Diener Suhcolumhites solitus Kiparisova
, ,. , , M r- . .-u ^„ ,,^ Suhcohimljitcs anonialu.s Kiparisova
published the fn-st monograph on am- p,,,„„„„„,, g,,,,-,,-, Kiparisova
monites from this region. There have been Pamnannites suhovijormis Kipariso\a
a number of studies on the ammonites and Pmanannites minor Kiparisova
iDelecvpods of these faunas; however, it is Lciophyllites pracmoturus Kiparisova
1 { 1.1 1 ti ■4-,-„^., ,.f i^;.^o,-,-c-^,,o EopiniUitcs amurensis Kiparisova
larcelv through the wntmgs ot Kipaiisova ,, ; ., / r-> i -^ \ j ■ i-- • ,.„
icii-^Kiy iiiiwugi. f3 1 l)(iiuiJ)>tcs (Daniihites) odmans K]pav\sova
(1947, 1961) that a picture of the stratlg- Oanuhitcs (Danuhitcs) hwcrUis Kiparisova
raphy and paleontology can be assessed. Dcmuhites (Preflorianites) inflaius Kiparisova
Korzh (1957 1959) has contributed to our Dcmuhites {Preflorianites) nmritimus Kiparisova
understanding of the petrography and M^'^c,phyUites inunaturus Kiparis.na
paleogeography of the Primoiye Scythian The above species are apparently in
formations. Regional stratigraphic data as direct association with Suhcolumhites. In
yet leave much to be desired. The Scythian addition, there are four species from the
fonnations of the Primorye Region repre- upper Scythian formations of the Primorye
sent 350-700 m of strata (Kiparisova, 1961: Region whose relations with the Suhco-
191). Four assemblages of ammonites are Uimhifes assemblage are not clear. First,
recognized within these formations. Kipari- there is Hellcnites (?) inopinatus Kiparisova
sova ( 1961 ) named the lowest assemblage which occurs at Cape Zhitkov, a locality
the Proptychitcs Zone, which she correlated ^ith a well developed Suhcolumhites fauna,
with the Otoceratan and Gyronitan ages Next, there is Prohung^arites (?) popovi
of Spath ( 1934 ) . This assemblage is con- Kiparisova from the strata on the west coast
fined to 100-200 m of strata. In the over- of Amur Bay which Kiparisova considers
lying 50 m of strata a Fleminoites Zone to be latest Scythian or earliest Anisian in
was recognized and correlated with the ^g^. Finally, there are two species — Co-
Flemingitan age of Spath (1934). The third himhitcs sp. indet. and Daii,noccras ?
faunal assemblage Kiparisova (1961) named unicum Kiparisova — that Kiparisova lists
the Prosphingitcs Zone. This assemblage is separately as beneath the main Suhco-
clearly identical to the Meekoceros fauna Jumhites fauna.
of the western United States. Recently, Recent studies by Zakharov have greatly
Kiparisova and Popov ( 1964 ) have pre- increased our knowledge of the stratigraphy
sented evidence to the effect that the .^j^d paleontology of the Primorxe Scythian
Flcmin^ites Zone and Vrosphiiv^itcs Zone cleposits. Zakharov ( 1966; additional per-
of the Primorye Region are equivalent, and sonal communication, 1967 ) recognizes a
they recommend excluding the Flemingites single Scythian zone above that of Owenites
Zone from the Scythian time scale. kocneni to which he gives the name Co-
The fourth and uppemiost faunal as- luuihifcs parisiauus Zone, ^^'ithin this zone
semblage in the Primorye Scythian forma- j^^. recognizes two subzones, a lower Neo-
tions was named the Suhcolumhites Zone columhites imignis Subzone and an upper
by Kiparisova. This upper zone encom- Suhcolumhites multiformis Subzone. Zak-
passes 100-150 m of strata. It is the fauna harov's zonal scheme with the key species
of this zcme that is i^ertinent to this paper, fgj. each segment is as follows:
Kiparisova ( 1961 ) recognized tlu' following Prosphin^ites insularis KiparisoN a,
species from her Suhcolumhites Zone: Suhco- P. filohosus Kiparisova, Sulx-o-
lumhitcs hinihites nitiUiforniis Kiparisova,
Pseudosdiiecerus simplex Kiparisova mtilti- Prenkites aff. timoren.sis Spatli.
Pseudosageeeras lon^ilolxitum Kiparisoxa forwis Paranannites ^meilis Kipariso\a,
Dieneroceras dieneri (Hyatt and Smith) Me'^aptmllites inunaturus Kipari-
Xetioeellites spitsher'^ensis Spaih Colum- sova.
Prusphin^ites ^lohosus Kiparisova bites .
Prosphingites insularis Kiparisova parisi- Svalhardieeras pansense Zak-
Suheolumbites multiformis Kiparisova anus Ikuon, Metadatinoeeras unieum
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 355
Neoco- (Kiparisova), Cohimhites parisi-
lumhitcs anus Hyatt and Smith, Pro-
insigui.s coltiiiihites sp., Neocoluinhites
insignis Zakharo\-, Keyserlingitcs
mcridianu.s Zakharo\', Olenckites
soiiticiis Zakharov, Hellcnites
inopinatus Kiparisova, H. tcher-
mjschewiensis Zakharov.
Zakharov's complete monograph on the
Scythian faunas of the Primorye Region is
not available to me at this moment. On the
basis of the data available, the ammonoids
of the Siibcohtmbites fauna of the Primorye
Region include the following species:
Pseudosageceras muUilohatum Noetling
Pseudosageceras simplex Kipariso\a
Dieneroccras kaiazini n. sp.
Xenoceltites spitshergensis Spath
Procaniites itiiDiafiirtis (Kiparisova)
Anuuttoccltitcs gracilis (Kiparisova)
Prosphingites glohosus Kiparisova
Prosphingitcs insidaris Kiparisoxa
Isculitoides suhmiforiiiis Kiparisova
Subcoliimbites intdtiformis Kiparisova
Prenkites aff. tiiiiorcnsis Spath
Leiopliijllites adinaris (Kiparisova)
Leiophyllitcs maritimiis (Kiparisova)
LeiopJnjlliics variabilis (Spath)
As mentioned above, Kiparisova ( 1961 )
was uncertain as to the precise horizon of
Frohiin'^oritcs (?) ])opoci Kiparisova. Zak-
harov (personal commimication ) informs
me that he has found ArctoJiiingaritcs
primoriensis Zakharov, Megaphijllites at-
losoviensis Zakharov, and LeioplujUifes
pruematunts Kiparisova on the west coast
of Amur Ray at a horizon 30 m below beds
with PioJuingorites (?) popovi, and con-
cludes that it is an Anisian form. In this
I agree.
One of the more important contributions
that Zakharov has made toward our under-
standing of the Scythian of the Primorye
Region is the recognition of the Columbites
Zone which he assigned to his Neocolum-
hites insignis Subzone. In the monograph
by Kiparisova (1961), it could only be in-
ferred, primarily on the basis of strati-
graphic position, that Columbites sp. indet.
and Dagnocems uniciim most probably rep-
resented the Cohunbites Zone. Zakharov
(personal communication) lists the fol-
lowing species from his Neocolumbites
insignis Subzone:
Svalbardiceras parisense Zakharov
Metadagnoceras tinicinu (Kiparisova)
Columbites parisianus Hyatt and Smith
Procolumbites sp.
Neocohimbites insignis Zakharov
Ketjscrlingitcs meridianus Zakliarov
Olenckites sonticus Zakharov
Hellcnites inopinatus Kiparisova
Hellcnites tchcrnyscheiciensis Zakharov
Cohimbites parisianus is the name giver
and primary member of the Columbites
fauna of southeastern Idaho. In addition,
the Idaho fauna also includes species of
Svalbardiceras, Keyscrlingites, and Hel-
lcnites. The relationship of the Primorye
Columbites fauna with that of the Die-
neroccras Zone of northern Siberia, which
I consider equivalent to the Columbites
fauna of southeast Idaho, is less direct;
there is not a single genus or species in
common. However, there is a close tie
between the fauna of the Dieneroccras
Zone of northern Siberia and the Colum-
bites fauna of Idaho, especially in species
of Dieneroceras, Pseudaspidites, and Nor-
dophiceras.
Northeastern Siberia
Until a decade ago our knowledge of the
Lower Triassic of northeastern Siberia
rested almost entirely on two contributions
of Mojsisovics ( 1886, 1888 ) . In recent years
our knowledge of this region has greatly
increased, largely due to the writings of
Yu. N. Popov. A summary of the stratig-
raphy of the major outcrop areas in this
area of Siberia can be found in Popov
(1958, 1960). Data on the sedimentology
and paleogeography of the Permian and
Lower Triassic formations in the Verk-
hoyansk Range can be found in Shutov
( 1958 ) . The principal recent discussion of
the ammonites of this region is in a mono-
graph by Popov (1961).
In the general area of the Olenek River
region, the Scythian comprises two facies
and stratigraphic units. The lower unit is a
clastic facies with Estheria and plant re-
356 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
mains; this is overlain by a marine clastic
unit containing two ammonoid faunas — the
Dicneroceras fauna of Popov, overlain by
the Olcnekites fauna. Further to the east,
lower Scythian ammonoid horizons occur,
but these are not the concern of this report.
The age and correlation of the Olenekites
fauna have been under discussion ever since
its monographic treatment by Mojsisovics
(1886, 1888). This author recognized the
following species in this fauna :
Dinarites spiniplicatus Mojsisovics
Diuarites vohittis Mojsisovics
Diiiaiitcs densipUcatus Mojsisovics
Dinarites altu.s Mojsisovics
Dinarites intermedins Mojsisovics
Dinarites ^hiciaJis Mojsisovics
Dinarites laevis Mojsisovics
Dinarites tolli Mojsisovics
Ceratites si<j.matoideus Mojsisovics
Ceratites nmltiplieatns Mojsisovics
Ceratites hyperhorens Mojsisovics
Ceratites fissiplieatns Mojsisovics
Ceratites discretus Mojsisovics
Ceratites niiddendorffi Keyserling
Ceratites sehrencki Mojsisovics
Ceratites subrohnstns Mojsisovics
Ceratites nikitini Mojsisovics
Ceratites biingei Mojsisovics
Ceratites deei)>iens Mojsisovics
Ceratites inustranzeffi Mojsisovics
Sil)irites eichwaldi (Keyserling)
Sil)irites pretiosus Mojsisovics
Xenudiseus enonipliahis ( Keyserling )
Xenodiscus schmidti Mojsisovics
Xenodisens deniosns Mojsisovics
Meekoceras karpinskii Mojsisovics
Meekoceras rotnndatum Mojsisovics
Meekoceras sihiricum Mojsisovics
Prosphingites czekanowskii Mojsisovics
In his recent monographic treatment of
the Olenek fauna, Popov (1961 and 1962a)
recognized the following species:
Fseudosageceras longilohalnm Kiparisova
Colnmhites (?) aff. ornatus Smith
Colmnhiles morplieo.s Popov
TiroUtes ex gr. eassianus (Qucnstedt)
Tirolites gerhaetisis Popov
Sibirites eiehxiuddi (Keyserling)
Sihirites pretiusns Mojsisovics
Sibirites suhpreliosus Popov
Parasibirites grarnbergi ( Popov)
Purasibiriles rariacnlealns Popov
Parasibirites niixtns Popov
Olenekites spiniplicalus Mojsisovics
Olenekites glacialis Mojsisovics
Olenekites altus Mojsisovics
Nordophiceras schmidti ( Mojsisovics )
Boreomeekoccras keyserlingi ( Mojsisovics )
Keyserlingites middendoiifi (Keyserling)
Keyserlingites subrobnsfus ( Mojsisovics)
Keyserlingites nikitini ( Mojsisovics)
Prophingites czekanowskii Mojsisovics
Arctoceras sim))lex (Mojsisovics)
Procarnites kunimeli Popov
Hemiprionites sibiricus (Mojsisovics)
Arctomeekoceras rotundatum (Mojsisovics)
Anasibirites raricostatus Popov
Pseudotirolites menensis Popov
My own analysis of this fauna leads me to
believe it consists of the following species:
Psendosageceras njultilobatuin Noetling
Preflorianites mnltiplicatus (Mojsisovics)
Proptyehitoides kunimeli (Popov)
Prosphingites ezekanouskii Mojsisovics
Czekanowskitcs dccipiens ( Mojsisovics )
Scalbardieeras schmidti ( Mojsisovics)
Svalbardiceras dentosns ( Mojsisovics)
Svalbardiceras sibiriciim ( Mojsisovics)
Nordophiceras pseudosimplex n. sp.
Arctomeekoceras rotundatum ( Mojsisovics )
Boreomeekocera.s keyserlingi ( Mojsisovics )
Sibirites pretiosus ( Mojsisovics )
Keyserlingites niiddendorffi ( Keyserling)
Keyserlingites subrobustus ( Mojsisovics)
Olenekites spiniplicatus ( Mojsisovics)
Tirolites mor))heos (Popov)
Arctotiroliti's menensis Popov
Smith (1932) considered the Olenekites
fauna as correlative with the Colnmhites
fauna of southeast Idaho and the youngest
zone of the Lower Triassic. Spath (1934)
expressed considerable concern over cor-
relation of the Olenek fauna, clearly rec-
ognizing the problem as it then existed. He
concluded, however, that the Olenek fauna
was latest Scythian in age and even sug-
gested that his latest division of the
Scythian could just as well be named
Olenekitan. The time relationship of the
Olenekites fauna and the Vwhungarites
fauna was clarified by the discovery of
elements of these faunas associated together
at a horizon approximately 1000 feet abo\e
the Colnmhites fauna in soutlieastern Idaho
(Kummel, 1954). Popov (1961) concluded
that the Olenekites Zone was correlative
\\ itii the Colnmhites Zone. Kiparisova and
Popov (1956) had previously arrived at a
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 357
similar conclusion. At a later date, Kipari-
sova and Popov (1961) accepted the
Frohiinp,arites Zone as being younger than
the CoJumbitcs Zone and correlated the
Olenekites fauna with it.
In a recent contribution, Kiparisova and
Popov ( 1964 ) correlated the Olenekites
fauna with the Colufnbites and Tirolites
faunas of southeast Idaho, and recorded an
additional younger zone, that of Frohun-
garites tubercidatus Welter. Descriptions of
the fossils and stratigraphic sequence of
this new assemblage are apparently in
press. It is reasonable to expect that in
this northern Siberian region two local
zones could be useful for detailed strati-
graphic analysis. On the other hand, on
the basis of the data available, it appears
more plausible that these two local zones
are correlative with the Frohungaritcs Zone
as interpreted here. The correlation of the
Olenekites fauna with the Columbites
fauna of southeast Idaho has no basis
whatsoever.
This brings us to the problem of cor-
relation of the Dieneroceras Zone of Popov.
That author (Popov, 1961, 1962a) recog-
nized the following species in this zone:
Pseudosageceras longilobatum Kiparisova
Dieneroceras deiuukidovi Popov
Dieneroceras apostolicus ( Smith )
Dieneroceras khelaliensis Popov
Dieneroceras nikabitensis Popov
Nordophiceras karpinskii ( Mojsisovics)
Nordophiccras alexeevae Popov
Nordopliiceras olenekensis Popov
Nordophiceras contrarius Popov
Koninckites posteriiis Popo\'
Imjoites eiekitensis Popov
Hemiprionites costatus Popov
My own analysis of this fauna leads me
to believe it contains the following species:
Pseudosageceras multilohatuni Noetling
Dieneroceras dcmokidovi Popo\-
Dieneroceras apostolicus ( Smith )
Subvishnuites eiekitensis (Popov)
Pseudaspidites posterius (Popov)
Nordophiceras euomphahis (Keyseding)
Nordophiceras alexeevae Popov
Hemiprionites costatus Popov
Popov ( 1962a ) concluded that the fauna
of his Dieneroceras Zone was correlative
with the Owcnites Zone of the circum-
Pacific region. In a personal communica-
tion, he further stated that he believed his
fauna to be correlative to the Anasibirites
Zone (upper Owenites). This correlation
was followed by Kiparisova and Popov
( 1964 ) . The conclusion arrived at here
that the Dieneroceras Zone fauna of Popov
is of Columbites Zone age is based pri-
marily on the close relations and possible
identity of the species of Dieneroceras,
SiibvisJinuites, Fseiidaspidites, and Nor-
dophiceras with the forms in the Coltimbites
fauna of southeast Idaho.
Spitsbergen
There is a considerable literature on
Triassic stratigraphy and ammonoids of
Spitsbergen. An extensive review of the
older literature of the Triassic stratigraphy
and paleontology with much new data has
recently been published by Buchan, et al.
(1965). The late Scythian is represented
by only three species of ammonoids:
Keyscrlingites sidjrobustus and Svalbardi-
ceras spitzbergensis Frebold, and Svalbardi-
ceras schmidti (Mojsisovics). These are
typical representatives of the late Scythian
{Frohungaritcs Zone) fauna of the circum-
Arctic region.
Ellesmere Island
A number of localities in the Blaa Moun-
tain and Blind Ford formations of Ellesmere
Island have yielded a small but highly
interesting late Scythian fauna. Tozer
( 1961a, 1965a ) has recognized the follow-
ing species in these faunas:
Olenekites canadensis Tozer
Svalbardiceras freboldi Tozer
Keyscrlingites subrobustus ( Mojsisovics )
Popov it es borealis Tozer
Zenoites arcticus Tozer
The above assemblage of species includes
several very typical late Scythian forms of
the circum-Arctic region. Underlying the
horizon \\'hich vields the above fauna, Tozer
(1965a) records Nordophiceras pilatum
(Hyatt and Smith) "associated with small
358 Bulletin Museum of Comparative Zoology, Vol 137, No. 3
ammonoids, probably Columhites sp." This
appears to identify the presence of the
Columhites Zone.
British Columbia
Late Scythian ammonoids are apparently
rare in British Columbia. Recently Tozer
( 1965a ) has described a small fauna from
three localities in northeastern British Co-
lumbia in the "Toad-Grayling Formation."
All the specimens come from a 20-30 foot
bed within the formation. From these three
localities Tozer ( 1965a ) has recognized the
following species:
Piocarnites modestus Tozer
Kcyserliufiites stibrohustus ( Mojsisovics)
Popovites occidcntaUs Tozer
P.sc'udosa^cceras bicarinatinn Tozer
Leiophyllites sp. iiidet.
Pro.sphiufiites ci. P. czekanoicskii Mojsisovics
Picfloriuiiites intermedins Tozer
Monacauthites monoceras Tozer
Mrtada^nocerci.s pidchcr Tozer
Siall)anlicei(is ciiowadei Tozer
I.sciditoides minor Tozer
My own analysis of this small fauna,
consisting of 41 specimens, leads me to
conclude that Procarnites modestus is a
synonym of Proearnites immaturus (Kipari-
sova) from the Primorye Region and that
Pseudosageeeras hicarinatum is a synonym
of Cord tile fit es angulatus Hyatt and Smith.
Tobin Formation, Nevada
The geology of the Tobin Formation, as
it is developed in the Mount Tobin quad-
rangle, Nevada, has been ably discussed by
Muller, et al. (1951). These authors rec-
ognized the Scythian age of the formation
and recorded the following fossils from
it: Claraiu cf. C. aurita, Mijopliaria sp.,
Lingulci sp., SuJ)cohimhites sp., Ilungarites
sp. In addition, Muller has shown the
author specimens of Olcnekites from the
Tobin Formation; the precise locality and
horizon within the formation of these
Olenckites is not known to the writer.
In 1959, in company with N.J. Silberling,
then of the U.S. Geological Surve\\ we
discovered a fossiliferous bed with well
preserved specimens a few tens of feet
above the base of the Tobin Fonnation.
This site has become USGS Mesozoic lo-
cality M2562 and is described geographi-
cally as follows (Silberling, written com-
munication): "Pershing County, Nevada.
South tip of Tobin Range, Cain Mountain
1:62,500 quad. Center NW V4 Sec. 9, T. 26
N., R. 39 E. 5,500 feet south, 27.5 west from
elevation point 5088 on range crest." The
fauna we collected from this site yielded
the following species :
Subcolumbites americanus n. sp.
Arnautoceltites teicherii n. sp.
StacJieitc's flowcri n. sp.
Isculitoidcs tca.sscrijergi n. sp.
Ussurites sieveri n. sp.
Metcida^noceras tobini n. sp.
HemilcciDiites ixiradiscu.s n. sp.
In the general vicinity of USGS locality
M2562 talus blocks of unknown strati-
graphic position within the Tobin Forma-
tion have yielded the following species:
Pseudosageeeras tntdtdobatitm Noetling
Keyserlingites sp. indet.
Ilellenifes radiattis Renz and Renz
Proltun^arites mckelvei n. sp.
Prohungarites sp. indet.
Confusion Range, Utah
The Thavnes Formation, including a
series of fossiliferous horizons, is well de-
veloped in the Confusion Range of west-
central Utah (Hose and Repenning, 1959).
The lower beds of the Thaynes Formation,
containing an abtmdant Meekoceras fauna,
rest on the Gerster Limestone of Permian
age; there is thus a significant hiatus be-
tween these formations. A second horizon
with ammonoids occurs from 1,100 to 1,120
feet above the base of the Thaynes Fonna-
tion. N. J. Silberling (in Hose and Repen-
ning, 1959:2188) has identified "OpJiiccras"?
sprncei Hyatt and Smith and "(^/^///c't'/Y/.s"?
i(U-ksoni Hyatt and Smith from this fauna.
These are species occurring in the Co-
]und)iles fauna of southeast Idaho and a
correlation with that fauna is suggested.
Abimdant ammonites, of poor to lair pres-
ervation, were found from 1,420 to 1,530
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel
359
feet above the base of the Thaynes ( USGS
Collection Mill). SilberHng (in Hose and
Repenning, 1959:2188) identified in this
collection Froptijchitoides mahomedis (Ar-
thaber), Xenoceltites cf. X. spitsbcrgensis
Spath, Tirolites cf. T. spinosus Mojsisovics,
T. aff. hoiieri Mojsisovics, and Pseudosage-
ceras sp. Through the courtesy of Dr. N. J.
Silberling I have had the opportunity of
studying USGS collection Mill and have
identified the following species:
Ussurites hoesi n. sp.
Pscttdoccltites neuadi n. sp.
Tirolites cf. cassianus (Quenstedt)
Pseudo.sageceras multilohatum Noetling
This fauna is of particular interest, as
none of the genera are present in the
Prohungarites fauna of the upper Thaynes
Formation in southeast Idaho. Tlie Suh-
columbites fauna from the Tobin Forma-
tion of Nevada contains a species of Us-
surites which, however, is quite distinct
from the Confusion Range species. Pseiido-
ccltifcs occurs in the Owenites Zone, Co-
himhites Zone, and in the latest Scythian
Prohungarites Zone. One species, Pseiido-
celtites doJnapaensis Kiparisova, is recorded
from the Cohimhitcs Zone of Astakhova
( 1960b) on the Mangyshlak Peninsula. The
Columhites fauna of southeast Idaho con-
tains Pscudoceltitcs cheneyi n. sp. which
is quite similar to P. nevadi. Tirolites like-
wise occurs through the upper half of the
Scythian but is of no assistance in die dating
of this fauna. Finally, Pscudosageceras
rtudtilohatum is present in all the Scythian
faunal zones.
There are thus few direct data for precise
age assignment of this fauna. It represents
a distinctive assemblage of species not
directly comparable to any other fauna.
The stratigraphic position, lying above a
horizon with species of Cohimbites Zone
age, suggests that we can include this fauna
within the late Scythian Prohungarites Zone.
Thaynes Formation, Southeast Idaho
The Thaynes Formation of southeast
Idaho includes one of the most complete
sequences of ammonoid faunas for the up-
per half of the Scythian. Extensive data
on the stratigraphy and facies relations of
the formation have been presented by
Kummel ( 1954, 1957 ) . In the general area
of Bear Lake, in southeast Idaho, the
Thaynes Formation contains five distinct
ammonoid horizons. The basal unit of the
formation is the lower limestone member
of Kummel ( 1954 ) and contains an abun-
dant Meekoccras fauna. Immediately over-
lying the lower limestone member is the
lower shale member which contains a well
developed Anasihirites fauna. There has
been considerable debate as to whether
these two faunas represent distinct zones
or are part of a single zone. Recently,
Kummel and Erben ( 1968 ) have presented
new data that favor considering these two
faunal horizons as subzones of the Owenites
Zone of mid-Scythian age, as originally sug-
gested by Smith ( 1932 ) . Approximately
620 feet above the lower limestone unit
with Meekoccras, occurs a small fauna of
two species of ammonites: Tirolites harti
and Dalmafites attcnuattis. Smith (1932)
correlated this fauna with that of the
Werfen Formation of the Alps and ad-
joining regions and established a Tirolites
Zone. The correlation of this fauna has
been discussed in detail on page 342 in
conjunction with the discussion of the
Werfen fauna. The conclusion arrived at
here is that this Idaho fauna is not correla-
tive with the Werfen fauna but is more
related to the overlying Cohimbites fauna,
and should probably be considered as
merely a local subzone of the Cohimbites
Zone.
One thousand feet above the lower lime-
stone member with Meekoccras is the
middle shale member which contains the
richly fossiliferous Cohimbites fauna. Smith
(1932) described the following species of
ammonites in this fauna from outcrops in
Paris Canyon:
Ophiceras jacksoni Hyatt and Smith
Ophiceras spcncei Hyatt and Smith
Meekoceras ciirticostatuin Smith
360 BuUctin Museum of Comparative Zoology, Vol. 137, No. 3
Meekoceras micromphahis Smith
Mcckoceras pilatum Hyatt and Smith
Meekoceras sanctorum Smith
Vscudharpoceras idahoense Smith
Tirolites cf. iUyrictis Mojsisovics
Fseudosageceras multiJohotum Noethng
Ccltites aposiolicus Smith
Ccltites plaiiovolvls Smith
Ccltites iirsensis Smith
Coluinbites consanguineus Smith
Columbites ligatus Smith
Cohiiu])ites niiniintis Smith
Coluiid)ites onuitus Smith
Columbites parisianus Hyatt and Smith
Columbites spencei Smith
Several new outcrops of the middle shale
member have been encountered in the area
around Bear Lake in southeastern Idaho,
which have yielded an abundance of addi-
tional specimens. Examination of all of
Smith's types and my own large collections
yields the following species as comprising
the Columbites fauna:
Fseudosageceras multilobatum Noetling
CordiUerites angulatus Hyatt and Smith
Dieneroceras aposiolicus ( Smith )
Subvishnuites sp. indet.
Xeuoccltitcs spencei (Hyatt and Smith)
Preflorianites montpelierensis n. sp.
Pseudaspidites popovi n. sp.
Columbites f)arisianus Hyatt and Smith
Pscudoceltites chcneyl n. sp.
Svalbardiceras sheldoni n. sp.
Nordophiceras pilatum (Hyatt and Smith)
Nordophiceras jacksoni (Hyatt and Smith)
Keyserlingites stcphensoni n. sp.
Tirolites smithi n. sp.
Tirolites astakJjovi n. sp.
Tirolites sp. indet.
Ilcllenites idahoense (Smith)
Dalmatites kittli n. sp.
Ussurites mansfieldi n. sp.
Approximately 1000 feet above the mid-
dle shale member with the Columbites
fauna is a unit consisting of a couple of
hundred feet of gray-brown limestones and
shales that have yielded the following
fauna:
Pseiidosageceras drinense Artliaher
Isculitoides hanunondi n. sp.
Epiceltites gentii Artliabt-r
Svalbardiceras sp. indet.
C zekanow.skites cf. decipicns Nhjjsisox ics
Stachcites sp. indet.
Keyserlingites bearriverensis n. sp.
Keyserlingites bearlakensis n. sp.
Olenekites cf. spiniplicatus Mojsisovics
Prohungarites mckehei n. sp.
Prohungarites gutstadti n. sp.
Prohungarites sp. indet.
Above this fossiliferous unit with Pro-
hungarites are at least 600 feet more of
strata, but these are very poorly preserved
and have yielded no ammonites. The top
contact of the Thaynes Formation is not
exposed in the Bear River Range.
SYSTEMATIC PALEONTOLOGY^
Class CEPHALOPODA Cuvier, 1797
Subclass AMMONOIDEA Zittel, 1884
Order PROLECANITIDA Miller and Furnish,
1954
Superfamily MEDUCOTTIACEAE Karpinsky,
1889
Family SAGECERATIDAE Hyatt, 1900
Genus Pseuc/osogeceros Diener, 1895
Type species, Pseudosageceras multilobatum
Noetling, 1905
No other genus of Scythian ammonoid
is as long ranging or as widely and abun-
dantly distributed as Pseudosageceras. Ap-
proximately a dozen species have been
recognized, most of which, however, were
based on one or few specimens and are
quite restricted in distribution. I recognize
\\'ithin the upper Scythian faunas of the
world the following five species:
Pseudosageceras multilobatum Noethng
Pseudosageceras drinense Arthaber
Pseudosageceras albanicum (Arthaber)
Pseudosageceras pasiptcuii Renz and Renz
Pseudosageceras sin)))lex Kiparisova
Among these species pasquaiji and sim-
plex arc known onl\' from single specimens
at single localities. Tire species albanicum
^ Abbreviations in this paper: MCZ = Museum
of Comparative Zoolou>'; BMNH = Briti.sh Museum
( Natural History ) ; GSI = Geological Sur\'ey of
India; PIUV = Paleontological Institute, ITniver-
sity of Vienna; GPIBo = Geological Institute,
Bonn University; NHMB = Natural History Mu-
seum Basel; USNM = United States National
Museum; USGS — United States Geological Sur-
vey. No data on repository are given for species
not personally examined In the author.
Ammonoids of the Late Scythian (Lower Triassic) • Kximmel
361
is fairly well represented in the Siihco-
Jiimhites fauna of Albania and Chios, as
is drinense, but the latter species is also
known from a single specimen of late
Scythian age from southeast Idaho. The
above species are only known from upper-
most Scythian horizons. The species which
is universal in its distribution is midti-
lobatum. There are few fossiliferous marine
formations of Scythian age that have not
yielded specimens of this species.
Pseudosageceras multilobatum Noetling
Plate 34, figure 6; Text-figure 2
Pseudosageceras midtilohatum Noetling, 1905:
181, pis. 19-27; Freeh, 190,5: pi. 23, figs. 4,
.5, pi. 25, fig. 1, pi. 26, fig. 3; Krafft and Diener,
1909: 145, pi. 21, fig. 5; Wanner, 1911: 181,
pi. 7, fig. 4; Diener, 1915: 2.37; Diener, 1917:
173, pi. 1, fig. 13; Welter, 1922: 94, fig. 3;
Diener, 1925: 96, fig. 26; Smith, 1932: 87,
pi. 4, figs. 1-3, pi. 5, figs. 1-6, pi. 25, figs. 7-16,
pi. 60, fig. 32, pi. 63, figs. 1-6; Kiitassy, 1933:
630; Collignon, 1933: 24, pi. 11, fig. 2; Spath,
1934: 54, fig. 6a; Kiparisova, 1947: 127, pi.
25, figs. 3, 4; Kummel, 1954: 185-187; Chao,
1959: 183, pi. 1, figs. 9, 12; Silbeding in
Hose and Repenning, 1959: 2194; Jeannet,
1959: 30, pi. 6, fig. 1; Tozer, 1961a: 44, pi.
13, figs. 8, 9; Kummel, 1966: 388, pi. 1, figs.
11, 12; Kummel and Erben, 1968: 112, pi. 19,
fig. 9; Kummel, 1968b: 489.
Pseudosageceras intermontanutj} Hyatt and Smith,
1905: 99, pi. 4, figs. 1-3, pi. .5, figs. 1-6, pi.
63, figs. 1, 2; Mathews, 1929: 3, pi. 1, figs.
18-22; C. Renz, 1945: 301; C. Renz, 1947: 147;
Renz and Renz, 1947: 62; Renz and Renz,
1948: 90, pi. 16, figs. 4, 7.
Pseudosageceras multilobatum var. giganteum
Kiparisova, 1947: 127, pi. 26, figs. 2-5; Popov,
1961: 13, pi. 2, figs. 1, 2.
Pseudosageceras of. multilobatum, — Kiparisova,
1961: 30, fig. 3.
Pseudosageceras schamarense Kiparisova, 1961: 31,
pi. 7, figs. 3, 4.
Pseudosageceras of. clavisellatum, — Renz and Renz,
1948: 90, pi. 16. fig. 3.
Pseudosageceras longilobatum Kiparisova and
Kiishtofovich, 1954: 20, pi. 11, fig. 3; Kipari-
sova, 1961: 29, pi. 6, figs. 1, 2, text-fig. 2;
Popov, 1961: 12, pi. 10, fig. 1, text-fig. 2.
This is without doubt the most common
and longest ranging of all Scythian am-
monoids. The species multdohattim differs
from P. drinense in the retention of a
narrow tabulate venter; however, the su-
tures are not all that different. Including
Fseudosageceras longilobatum Kiparisova in
this species is done mainly on the claim
of Popov (1961, p. 13) that the venter
on the holotype of longdobatum is not
preserved, but on the specimen figured by
Popov (1961, pi. 10, fig. 1), which has a
suture identical to the holotype of longi-
lobatum, the venter is tabulate. The lobes
and saddles of this species are not much
different from those of multilobatum (Fig.
2A, I).
Occurrence. Worldwide in distril:)ution,
found in all Scythian zones. From the
uppermost Scythian, the species is known
from the Subcohimbites fauna of Chios;
the upper Scythian of the Mangyshlak
Peninsula; the Narmia Member of the
Mianwali FonTiation in the Surghar Range
and Salt Range of West Pakistan; Nifoe-
koko, Timor; the Subcohimbites fauna of
the Primorye Region; the Olenekites and
Dieneroceras zones of the Olenek River
region; the Columbites fauna of southeast
Idaho; the Upper Thaynes Formation, Con-
fusion Range, Nevada; Upper Tobin For-
mation, south end of Tobin Range, Nevada.
Repository. Specimens from Columbites
Zone at MontpeHer Canyon — suture speci-
men (Fig. 2C) MCZ9628, unfigured speci-
mens MCZ 9549; from Hot Springs MCZ
9550; specimens from upper Thaynes For-
mation, Confusion Range, plesiotype (PI.
34, fig. 6) USNM 153072; unfigured speci-
men from Tobin Formation, Nevada, MCZ
9650; from Narmia Member, Mianwali For-
mation, Salt Range and Surghar Range,
West Pakistan, MCZ 9576-9580; from Sub-
columbites fauna Kotal-e-Tera, Afghanis-
tan, MCZ 10166, 10173; from Subcohimbites
fauna of Chios, plesiotype P. intermontanum
(Renz and Renz, 1948: pi. 16, fig. 4)
NHMB J13813; (Renz and Renz, 1948: pi.
16, fig. 7) NHMB J13814, unfigured speci-
mens NHMB J13815; P. cf. clavisellatum
(Renz and Renz, 1948: pi. 16, fig. 3)
NHMB J13812; specimen from Olenekites
Zone, Olenek River region MCZ 8678.
362 BuUetin Museum of Comparative Zoology, Vol. 137, No. 3
A\
D
Figure 2. Diagrammatic representation of the suture of: A, Pseudosageceras wultilobatum, — Krafft and Diener (1909: pi.
21, fig. 5c), from Hedenstroemia beds, Muth, Himalayas, at a diameter of approximately 70 mm; B, fioiotype Pseudosogeceros
inlermontanum Hyatt and Smith (1905: pi. 4, fig. 3), from Mee/toceros limestone, Thaynes Formation, southeast Idaho,
at a diameter of 65 mm; C, Pseudosogeceros multilobatum Noetling (MCZ 9628), from Columbites fauna, Montpelier Can-
yon, southeast Idaho, at a whorl height of 10.5 mm; D, Pseudosogeceros schamarense Kiparisovo (1961: fig. 4), from
mid-Scythian strata in the Primorye Region, at a whorl height of 18 mm; E, Pseudosogeceros drineme Arthaber (1911:
pi. 17(1), fig. 7), from Subco/umbifes fauna of Albania; F, Pseudosogeceros drinense Arthaber, from Upper Thaynes
Formation, Hammond Creek, southeast Idaho (MCZ 9489), at a whorl height of 13 mm; G, Pseudosogeceros simplex
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel
363
Pseudosageceras drinense Arthaber
Plate 12, figures 4, 5; Text-figure 2
Pseudosageceras multilohatum, — Arthaber, 1908:
279, pi. 12(2), figs. 3a-c.
Pseudosageceras drinense Arthaber, 1911: 201, pi.
17(1), figs. 6, 7; Diener, 1915: 236; C. Renz,
1928: 155; Kutassy, 1933: 639; Spath, 1934:
55, fig. 6c; Renz and Renz, 1947: 62; Renz and
Renz, 1948: 92, pi. 16, figs. 6-6a.
Pseudosageceras drinense Arthaber var. incen-
trolata Renz and Renz, 1948: 92, pi. 16, figs.
11-lla.
Metahcdenstroemia n. sp. Kummel, 1954: 187.
Arthaber (1911: 201) stated he had 13
specimens of this species but only the
holotype specimen (PL 12, figs. 4, 5) is still
preserved. The principal distinguishing
feature of this species is the acute venter
developed in the later growth stages. The
suture (Fig. 2E) is quite similar to that of
Fseudosageceras multilohatum. A small
fragmentary specimen from the upper
Thaynes Formation can be assigned to this
species. Its suture (Fig. 2F), though taken
at a whorl height of 13 mm, is like that of
the Albanian specimens.
Occurrence. Subcolumhites fauna of
Albania and Chios; the upper Thaynes For-
mation, Hammond Creek, southeast Idaho.
Repository. The holotype is in the
Paleontological Institute, Vienna; the Brit-
ish Museum has 15 paratypes BMNH
C22984-98, C23002; the figured specimens
from Chios (Renz and Renz, 1948: pi. 16,
fig. 6) NHMB J13816, unfigured specimens
from Maradovuno on Chios NHMB J13817,
from Kephalovuno NHMB J13818; the
specimen from southeast Idaho is MCZ
9489.
Pseudosageceras albanicum (Arthaber)
Plate 21, figures 5, 6; Text-figure 2
Sageceras albanicum Arthaber, 1908: 281, pi.
13(3), figs, la-c; Arthaber, 1911: 203, pi.
17(1), figs. 4, 5; Diener, 1915: 249; C. Renz,
1928: 155; Kutassy, 1933: 651; Renz and
Renz, 1947: 62; Renz and Renz, 1948: 94,
pi. 16, figs. 5-5a, 10-lOa.
Pseudosageceras albanicum, — Spath, 1934: 56,
fig. 6b.
Arthaber (1911: 203) stated he had 15
specimens of this species. There is today
in the Paleontological Institute, Vienna,
only the holotype figured by Arthaber and
also figured here (PL 21, figs. 5, 6). This
specimen measures 64.3 mm in diameter,
12 mm for the width of the adoral whorl,
and approximately 40 mm for the height
of the adoral whorl. Arthaber's representa-
tion of the suture ( Fig. 2M ) is accurate.
The principal distinguishing feature of this
species is its suture.
Occurrence. Subcolumhites fauna of
Albania and Chios.
Repository. The holotype is in the
Paleontological Institute, Vienna; the Brit-
ish Museum has 13 paratypes BMNH
C22999-23001, 23003^12; the plesiotypes
from Chios (Renz and Renz, 1948: pi. 16,
fig. 5) NHMB J13821, (Renz and Renz,
1948: pi. 16, fig. 10) NHMB J13822;
unfigured specimens from Maradovuno
NHMB J 13823, from Kephalovuno NHMB
J13824.
Kiparisova (1961: fig. 1), from Subco/umbifes fauna, Primorye Region, Siberia, at a whorl fieigfit of 14.5 mm; H,
Pseudosageceras pasquay/' Renz and Renz (1948: pi. 16, fig. 2b), from Subco/umbifes fauna of Chios, at a diameter of 35
mm; I, Pseudosogeceras longi/obatum Kiparisova (1961: fig. 2), from Subco/umb/'tes fauna of Primorye Region, Siberia,
at a whorl height of 19 mm; J, holotype, Pseudosageceras fsofengense Chao (1959: fig. 5b), from Owen/tes zone,
Kwangsi, China, at a diameter of approximately 30 mm; K, Psevdoiagecerai longilobatum var. (cwongs/ense Chao (1959:
fig. 5c), from Owenites Zone, Kwangsi, China, at a diameter of approximately 35 mm; L, Pseudosagecerai compressus
(Mathews, 1929: pi. 1, fig. 17), from /4nasibib/(es fauna. Fort Douglas, Utah, at a diameter of approximately 20 mm;
M, Pseudosogeceras albanicum (Arthaber, 1911: pi. 17(1), fig. 5), from Subco/umb/fes fauna of Albania at a diameter
of 55 mm; N, holotype Pseudosogeceras curvafum Chao (1959: fig. 52), from Flemingites fauna, Kwangsi, China, at a
diameter of approximately 70 mm; O, Pseudosageceras cf. c/ovise/Zatum, Renz and Renz (1948: pi. 16, fig. 3a), from
Subco/umbi/es fauna of Chios, at a diameter of 24 mm; P, Pseudosageceras clavisellatunt Diener (1913: pi. 4, fig. 5c), from
Ophiceras layer, Pastannah, Kashmir, at a diameter of approximately 30 mm.
364 Bulletin Museiint of Comparative Zoology, Vol. 137, No. 3
Pseudosageceras pasquayi Renz and Renz
Text-figure 2
Pseudosageceras (Metasapeceras) pasquayi Renz
and Renz, 1947: 62, 79; Renz and Renz, 1948:
93, pi. 16, fig. 2.
Pseudosageceras pasquayi, — Kunimel, in Arkell et
al., 1957: L75.
A species based on a single specimen,
distinctive for its unusual suture ( Fig. 2H ) .
The holotype measures 37.7 mm in diam-
eter, 6.3 mm for the width of the adoral
whorl, 22.5 mm for the height.
Occurrence. Stibcolumbites fauna, Chios.
Repository. Holotype NHMB J13S20.
Pseudosageceras simplex Kiparisova
Text-figure 2
Pseudosageceras siinpJex Kiparisova, 1947: 128,
pi. 25, fig. 2, text-fig. 6; Kiparisova, 1961: 28,
pi. 6, fig. 3, text-fig. 1.
Another species established for a single
incomplete specimen. Its special features
are a narrow rounded venter and a quite
simple suture (Fig. 2G). The suture is very
much like that of P. alhanicum except for
its lack of curvature (Fig. 2G, M). It is
highly possible that this difference is not
of specific importance and that additional
samples of each of these species would
show the curvature and alignment of the
suture to be highly variable.
Occurrence. ?>uhcolumhitcs Zone, Pri-
morye Region, Siberia.
Genus Cordillerites Hyatt and Smith, 1905
Type species, Cordillerifes anguiatus Hyatt
and Smith, 1905
Cordillerites anguiatus Hyatt and Smith
Plate 20, figures 5, 6; Plate 51, figures
6, 7; Text-figure 3
Cordillerites anguiatus Ilvatt and Snn'tli, 1905:
110, pi. 2, figs. 1-8, pi. 68, figs. 1-10, pi. 71,
figs. 1-6, pi. 85, figs. 14-20; Freeh, 1908:
pi. 63, fig. 2; Diener, 1915: 112; Diener, 1917:
175, pi. 1, fig. 11; Smith, 1932: 96, pi. 2,
figs. 1-8, pi. 42, figs. 14-20, pi. 60, fig. 14,
pi. 68, figs. 1-10, pi. 71, figs. 1-6; Spath, 1934:
61; C. Renz, 1947: 176; Kuniniel, in Arkell,
et al., 1957: L75.
Iledenstroemia ski])etarcnsis Arthaber, 1911: 208,
pi. 17(1), fig. 13.
Taisle 23. Measurements of 15 Specimens
Assigned to Cordillerites angulatus Hyatt
AND Smith.
D
w
H
u
W/D
H/D
U/D
1.
97.0
28.0
59.4
0
28.9
61.2
0
2.
42.0
14.0
28.0
0
33.3
66.6
0
3.
35.0
10.0
23.0
0
28.6
65.9
0
4.
35.0
11.0
22.4?
0
31.4
64.0?
0
5.
33.0
6.3?
20.0
0
19.1
60.6
0
6.
29.5
5.8
17.4?
0
19.7
59.0?
0
7.
24.2
6.2
15.6
0
25.6
64.5
0
8.
22.6
5.3
13.7
23.5
60.6
0
9.
22.0
4.5?
13.4
20.5
60.9
0
10.
21.3
5.3
14.0
24.9
65.6
0
11.
19.4
4.7
12.0?
24.2
61.9?
0
12.
15.8
4.1
9.4
25.9
59.5
0
13.
15.3
4.0
9.4
0
26.1
61.5
0
14.
14.4
4.5
9.3
0
31.1
64.5
0
15.
14.2
3.1
8.0
0
21.8
56.4
0
1. Paralectotype, Hvatt and Smith (1905: pi. 68, figs.
1-3).
2. Holotype, Pseudosageceras hkarhuitum To/.er (1965:
16). '
.3. Paralcctot\pe, Hvatt and Smith (1905: pi. 2, figs.
4, 5).
4. Lt'ctotvpe, Hvatt and Smith (1905: pi. 2, figs. 1-3).
5. Plesiotype (PL 51, fig. 1), MCZ 9569.
6. 10, 15. Specimens from Cohimbitcs fauna. Bear
Lake region, southeast Idaho.
7. Paralectotvpe, Hyatt and Smith (1905: pi. 68, figs.
8-10).
8. Plesiotype, — Renz and Renz (1948: pi. 16, fig. 9).
9. 13. Unfigured specimens from Chios, NHMB.
11. Paralectotype, Hyatt and Smith (1905: pi. 2, fig. 6).
12. Holotvpe, Hcdenstrocniia skipctarcnsis Arthaber (1911:
pi. 17(1), fig. 13).
14. Paralectotvpe, Hvatt and Smith (1905: pi. 42, figs.
14-16).
Epihedenstroemia sM])ctarensis, — Spath, 19.34: 222,
fig. 71.
ef. Cordillerites anguiatus, — Renz and Renz, 1947:
67; Renz and Renz, 1948: 88, pi. 16, fig. 9.
Cordillerites kwangsianus Chao, 1959: 33, 188,
pi. 44, figs. 7, 8, text-fig. 6b; Knnimel and
Steele, 1962: 645.
Cordillerites orientalis Chao, 1959:
I, figs. 10, 11, text-fig. 6a.
Pseudosageceras bicarinatuin Tozer,
pi. 2, figs. 8a-d, text-fig. 1.
34, 188, pi.
196.5a: 16,
Cordillerifes up until now has been
known only through its type species, C.
aniitdatus, from the Mcckoccra.s limestone
of southeast Idaho. The description and
illustrations of this species by Fhatt and
Smith (1905) and Smith (1932) arc quite
adeciuate. The measurements of the speci-
mens assigned to this species are given
on Table 23.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel
365
/N
G
H
^
J{AAAr[
Figure 3. Diagrammatic representation of the sutures of two species of Cordillerites. A-K, Cordillerites angulatus Hyatt
and Smitfi; A, paralectotype, at a diameter of 90 mm (Hyatt and Smitfi, 1905: pi. 68, fig. 3, USNM 75300a); B,
poralectotype, at a diameter of 80 mm (Hyatt and Smith, 1905: pi. 68, fig. 6, USNM 75300b); C, lectotype, at a diameter
of 35 mm (Hyatt and Smith, 1905: pi. 2, fig. 3, USNM 75247a); D, paralectotype, at a diameter of 25 mm (Hyatt and
Smith, 1905: pi. 68, fig. 10, USNM 75300c); E, paralectotype, at a diameter of 17 mm (Hyatt and Smith, 1905: pi. 71,
fig. 5); F, paralectotype, at a diameter of 10 mm (Hyatt and Smith, 1905: pi. 71, fig. 2); G, holotype of Hedensfroemio
s/tipetorensis Arthaber (1911: pi. 17(1), fig. 13), new drawing at a diameter of 15.7 mm; H, specimen from Columbites
Zone, southeast Idaho, at a diameter of 29 mm (MCZ 9569); I, cf. Cordillerites angulatus, — Renz and Renz (1948: pi.
16, fig. 9b), at a diameter of approximately 20 mm; J, holotype C. orientalis Chao (1959: fig. 6a), at a diameter of 55
mm; K, holotype C. /twongsianus Choo (1959: fig. 6b), at a diameter of 19 mm; L, Cordillerites conclnnus Kiparisova (1961:
fig. 5), at a diameter of 42 mm.
Specimens of figures A-F, from Meekoceras fauna of southeastern Idaho; G, from Subco/umfa/fes fauna of Albania; H,
Columbites fauna of southeastern Idaho; I, from Subco/umbites fauna of Chios; J, from Subco/umb/tes fauna of Kwangsi,
China; K, from Owenites fauna of Kwangsi, China; L, from the Proptychites Zone of the Primorye Region.
366 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
The Columhites fauna of southeast Idaho
has yielded six specimens of this species.
One of them is ilhistrated here on Plate
51, figures 6, 7 and its suture on Figure
3H. These specimens agree remarkably
with all the essential features of the types
from the Meekoceros limestone.
Hedenstroemia skipetarensis Arthaber
(1911: 208) was established for a small,
involute, compressed specimen of 15.8 mm
in diameter (PI. 20, figs. 5, 6). Arthaber
(1911) considered this small specimen to
be unique and possibly deserving separate
generic status. Spath (1934: 222) estab-
lished the genus Epihedenstroemia with
Hedenstroemia skipetarensis Arthaber as
type. The uniqueness of the species centers
around the curvature of the suture as
depicted by Arthaber (1911: pi. 17(1),
fig. 13c). Re-examination of Arthaber's
type specimen clearly shows that this rep-
resentation of the suture was not one of
the most successful in his monograph. A
new drawing of the suture of this type
specimen is shown here on Figure 3G. The
suture is nearly identical to the suture of
the American specimens of Cordillerites
anii^ulatus from the Meekoceras limestone
at a diameter of 17 mm (Fig. 3E) and 25
mm (Fig. 3D). The shape of the conch,
nature of the venter, etc., also fit in per-
fectly with the American type specimens.
The Subcohnnbites fauna of Chios also
contains this species. Renz and Renz ( 1948:
89) had one specimen they assigned to
this species with the symbol "cf." There
are in addition four specimens from the
Subcohimbites fauna of Chios in the Nat-
ural History Museum, Basel. The suture
(Fig. 31), as noted by Renz and Renz
(1948: 89), is very much like that of the
lectotype (Fig. 3C). There is no question
but that these Subcohimbites fauna speci-
mens from Chios are conspccific with tlie
American type specimens.
Tozer (1965a: 16), in the discussion of
his new species Pseudosa<ieceras bicarina-
tmn, stressed the j^ossible allinities of
his species with Hedenstroemia skipetaren-
sis Arthaber. The suture of Tozer's speci-
men is nearly identical to that of C.
auii^idatus illustrated by Hyatt and Smith
( 1905: pi. 68, fig. 6; Fig. 3B of this report).
The general shape of the conch, etc., fits
in perfectly.
The descriptions and illustrations of
Cordillerites kicani:,sianus Chao (1959)
from the Oicenites Zone, Kwangsi, China,
and Cordillerites orientalis from the Sub-
eolumbites fauna of Kwangsi, China, leave
much to be desired. The first of these
species was based on a single specimen,
the second on only two. The sutures of
these species (Figs. 3J, K) are very com-
parable to the mature sutures of the Ameri-
can type specimens (Figs. 3A, B). I can
find no justification for not considering
these two species as synonyms of Cordil-
lerites angulatus.
The only other species of Cordillerites
recognized here is C. eoncinnus Kiparisova
from the early Scythian Froptijehites Zone
of Kiparisova (1961: 33) in the Primorye
Region, Siberia. This species differs mainly
in the nature of its suture.
Cordillerites eompressus Mathews (1929:
3) from the Anasibirites fauna of Fort
Douglas, Utah, had been accepted as a
valid species of Cordillerites by Smith
(1932) and Spath (1934). On the basis
primarily of the much greater elaboration
of the suture, I consider this to be a species
of Fseudosaii^eeeras.
Oceurrence. This species is now known
from the mid-Scythian Meekoceras Zone of
southeast Idaho and the equivalent horizon
in Kwangsi, China (Chao collection 542b);
from the Columbites fauna of southeast
Idaho; from the Subcohimbites fauna (or
its equivalent) in Albania, Chios, Kwangsi,
and British Columbia.
Repositorij. The new specimens from
the Columbites fauna of southeast Idaho
recorded Ikmc are MCZ 9569 (PI. 51, figs.
5, 6), unfigured specimens from Hot
Springs MCZ 9627.
Ammonoids of the Late Scythian (Lower Triassic) • KummcJ 367
Order CERATITIDA Hyatt, 1884
Superfamily OTOCERATACEAE Hyatt,
1900
Family DIENEROCERATIDAE Kummel, 1952
Tliis family was originally introduced for
the single genus Dicneroccras which is in-
terpreted as a persisiting stock of the
ophiceratids and a probable root of later
ornamented stocks. Dieneroceras is one of
the simplest of Scythian ammonoids and
quite common throughout the Scythian. In-
cluded in the family are Suhvishmiites and
Hemilecanites. Both of these genera are
much like Dieneroceras in the simplicity of
their sutures and conch form. They are
characterized by acute venters; Siibvish-
nuites has a more inflated whorl section;
Hemilecanites a highly compressed whorl
section. Hemilecanites is restricted to the
late Scythian Prohun<i,arites Zone; Dienero-
ceras and Siibvislmuites are quite common
in mid-Scythian faunas.
Genus Dieneroceras Spath, 1934
Type species, Ophiceros dieneri Hyatt and
Smith, 1905
This genus is not nearly as common nor
as widely distributed in the late Scythian
as it is in the Owenites Zone of mid-
Scythian age. There are two species (D.
mediterranean D. skutarensis) represented
by few specimens in the Subcohimbites
faunas of Albania and Chios. The correla-
tive fauna in the Primorye Region contains
a single specimen assigned to D. karazini
n. sp. In the underlying Columbites Zone
there are only two species: D. demokidovi
from northern Siberia, and D. apostolicus
from southeast Idaho and northern Siberia.
Dieneroceras mediterranea (Arthaber)
Plate 4, figures 7-10; Plate 19, figures
3, 4; Text-figure 4.
Xenaspis mediterranea Arthaber, 1908: 260, pi.
11(1), figs. 3a-c; Arthaber, 1911: 231; Diener,
1915: 311; Spath, 1934: 134, 136, 293.
Celtites kcirensis Arthaber, 1908: 273, pi. 11(1),
figs. 8a-c; Diener, 1915: 75; Renz and Renz,
1948: 42, pi. 3, fig. 6 ( non 3).
Xenodiscus kcirensis (Arthaber) 1911: 181.
Ophiceras sakiiutaJa Arthaber (non Diener), 1911:
239, pi. 21(5), fig. 4.
Ophiceras of. sakuniula Diener, 1915: 212.
The type specimen measures 53.7 mm
in diameter, 11.4 mm for the width of the
last whorl, 13.8 mm for the height, and
26.5 mm for the diameter of the umbilicus.
The whorls are subtrapezoidal in cross-
section, the flanks converging slightly
towards a low, arched venter. The ventral
and umbilical shoulders are rounded. The
specimen had been ground and polished
on part of the phragmocone to expose the
suture, but in this case I do not believe
there is any appreciable distortion to the
suture. A new drawing of this suture is
shown on Figure 4A.
The specimen Arthaber (1908: 273) as-
signed to Celtites kcirensis is a small in-
dividual with the following measurements:
Diameter 28.8, Width 8.8, Height 9.3,
Umbilicus 12.8 mm. The apparent sharpen-
ing of the venter in the adoral quarter
volution I believe is due to factors of preser-
vation and is not the true outline of the
whorl at that stage (Pi. 4, figs. 9, 10).
The suture is identical to that of the type
specimen. The specimen clearly is a juve-
nile form of D. mediterranea. A compa-
rable specimen from the Subcolumbites
fauna of Chios was assigned by Renz and
Renz (1948: 42, pi. 3, fig. 6)' to Celtites
kcirensis, and this I believe also to be a
juvenile form of D. mediterranea. One of
the specimens from Chios that Renz and
Renz (1948: 42, pi. 3, fig. 3) assigned to
Celtites kcirensis is a specimen of Hemile-
canites discus.
The specimen Arthaber assigned to
Ophiceras sakuntala Diener is incomplete
and sHghtly crushed (PI. 19, figs. 3, 4).
Direct comparison of this specimen with
the other specimens assigned to D. mediter-
ranea convinces me that they are con-
specific.
Occurrence. Subcolumbites fauna of
Albania and Chios.
Repository. The Paleontological Institute
of Vienna has the holotype of Xenaspis
368
Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
B
A
Figure 4. Diagrammatic representation of the suture of: A, fiolotype, Dieneroceras mediterranea (Arthaber, 1908: pi.
11(1), figs. 3a-c), from Subco/umbifes fauna of Albania, new drawing at a diameter of 36 mm; B, fiolotype D/eneroceros
s/(utarensis (Artfiaber, 1911: pi. 21(5), fig. 1), from Subco/umb;fes fauna of Albania, at a diameter of 18 mm; tfie
goniatitic aspect of tfie lobes is most probably due to excessive grinding in tfie preparation of tfie specimen; C,
D/eneroceros demokidovi Popov (1961: fig. 6f), from "D/eneroceros" Zone, Olenek River basin, Siberia; D, D/eneroceros
karazini nov. nom. (Kiparisova, 1961: fig. 14), from Subco/umb/fes fauna, Primorye Region, Siberia, at a wfiorl fieigtit
of 7 mm; E, fiolotype Hem/lecon/fes discus (Arthaber, 1908: pi. 11(1), fig. 5), from Subco/umb/tes fauna of Albania ot a
diameter of approximately 20 mm; F, poratype Hemilecanites pctradiscus n. sp. from Subco/umb/tes fauna of Tobin Forma-
mation, Nevada, at a diameter of 16 mm (MCZ 9631); G, poratype Hemilecanites paradiscus n. sp. from Subco/umbifes
fauna of Tobin Formation, Nevada, at a diameter of 14 mm (MCZ 9483); H, holotype Xenasp/s enver/s Arthaber (1911: pi.
20(4), fig. 3), from Subco/umbifes fauna of Albania.
mediterranea and Ccltite.s kciren.si.s, and the
plesiotype of Ophiceras sakuntaki Arthaber
(noil Diener). The ple.siotype of Celtitcs
kcirensis from Chios i.s NUMB J13654.
Dieneroceras skufarensis (Arthaber)
Plate 20, figures 3, 4; Text-figure 4
Lecanites skntarctisis Arlhabcr, 1911; 237, pi.
21(5), lis. 1; Spath, 1934: 135, 136: Rciiz
and Ren/., 1947: 61; Renz and Ren/,, 1948;
55, pi. 3, fijr. 5.
Prodvitcs .skuldrciisis, — Dieiier, 1915: 228.
This is a dieneroceratid very much on the
pattern of Dieneroceras knechti ( Hyatt and
Smith) from the Oioenites Zone; it could
possibly be eonspecifie with tliat species.
However, because of tlie smallness of the
sami:)le axailable, a tendency for D. sku-
Ammonoids of the Late Scythian (Lower Triassic) • Kummel
369
tarensis to have a slightly more inflated
whorl section, and the great age difference
between these forans I believe it best to
recognize the independent status of this
species. Another factor is the goniatitic
character of the lobes (Fig. 4B). The
specimen has been excessively ground and
polished to expose the suture, and this
could entirely account for the smooth lobes.
Occurrence. Subcolumbites fauna of
Albania and Chios.
Repository. The holotype, and only
specimen from Albania, is in the Paleonto-
logical Institute, Vienna. The figured speci-
men from Chios (Renz and Renz, 1948:
pi. 3, fig. 5) is NHMB J13700, unfig-
ured specimens from Maradovuno NHMB
J13701, from Kephalovuno NHMB J13702.
The MCZ has three specimens from Chios,
MCZ 10027, 10029.
Dieneroceras karazini n. sp.
Text-figure 4
Dieneroceras dieneri Kiparisova (non Hyatt and
Smith) 1961: 47, pi. 9, fig. 2.
Kiparisova had only a single specimen
of this species and this is indeed quite
similar to D. dieneri. However, because of
the great difference in age of the American
D. dieneri {Owenites fauna) and the Pri-
morye specimen (Sidjcohnnhitcs fauna) I
believe it best to consider them as separate
species. There are differences in involu-
tion, shape of the whorl section and suture
( Fig. 4D ) , but the smallness of the sample
prevents any evaluation of these differ-
ences.
Occurrence. Primorye Region, from Sub-
columbites fauna between Cape Mushketov
and Cape Karazin.
Dieneroceras demokidovi Popov
Text-figure 4
Dieneroceras demoMdovi Popov, 1961: .36, pi. 12,
figs. 1, 5.
Dieneroceras nikabitensis Popov, 1962a: 184, pi.
3, fig. 1.
The conch of this species is very much
on the pattern of that of D. knechti from
the Owenites Zone, except that the whorls
are slightly more inflated. The conch is
apparently completely smooth. It is really
only in the smoothness of the conch that
one can readily separate this species from
D. apostolicus. The suture is illustrated on
Figure 4C.
Occurrence. Dieneroceras Zone of Popov
( 1961 ) from a number of localities in the
Olenek and Kolyma river basins, Siberia.
Repository. Popov's specimens are in the
Tchernyshev Central Geological Museum
of Leningrad. The Museum of Compara-
tive Zoologv has three topotvpe specimens,
MCZ 6105,' 6106, 8679.
Dieneroceras aposfolicus (Smith)
Plate 53, figures 1-12; Text-figures 5, 6
Celtites apostolicus Smith, 1932: 104, pi. 48,
figs. 1-10.
"C elates" apostolicus, — Kummel, 1954: 187;
Kmiimel, 1961: 519.
Dieneroceras apostolicus, — Popov, 1961: 37, pi.
12, fig. 6.
Celtites planovolcis Smith, 1932: 104, pi. 48,
figs. 11-20.
''Celtites" planovolvis, — Kummel, 1954: 181;
Kummel, 1961: 519.
Celtites ursensis Smith, 1932: 104, pi. 47, figs.
11-23.
"Celtites" ursensis, — Kummel, 1954: 187; Kum-
mel, 1961: 519.
Dieneroceras khelaliensis Popov, 1961: 37, pi.
12, fig. 4.
Smith's ( 1932 ) analysis and description
of his three species of Celtites were not the
most successful of his efforts. The three
species were differentiated on slight dif-
ferences in height and width of the whorls.
The measurements of 50 specimens from
the Columbites fauna of southeastern Idaho
are listed on Table 24 and plotted on Figure
6. It can readily be seen that the umbilical
diameter and whorl height show only a
small range of variability, clearly intra-
specific. Smith (1932: 104) likewise stated
the suture to be goniatitic. The sutures of
six specimens plus five sutures reproduced
by Smith (1932) are shown on Figure 5.
370 Bulletin Museum of Comporative Zoology, Vol. 137, No. 3
A
B
H
A
D
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 371
ZZr
20
18
16
14
12
10
8
6
4
2
0
.6 .
u
I I I I I \ I I I L_
10 20
30
DIAMETER
Figure 6. Variation in umbilical diameter (U) and whorl height (H) of Dieneroceras opos/o/icus (Smith) from Columbifes
fauna of Bear Lake region, southeast Idaho. The data on this graph are from Table 24.
All the specimens I studied have denticu- One additional factor of the suture is the
lated lobes. It should be remarked, how- great variability in the shape and size of
ever, that the preservation in dense, fine the various elements.
grained, black limestone often makes fine Smith noted the weak, forward projecting
details of the suture very hard to observe, constrictions; however, these are absent
Figure 5. Diagrammatic representation of the sutures of Dieneroceras apostolicus (Smith). A, at a diameter of 14.2 mm
(MCZ 9524); B, at a diameter of 16.3 mm (MCZ 9525); C, at o diameter of 24.1 mm (MCZ 9526); D, at a diameter of 18 mm
(USNM 74989c), from Smith (1932: pi. 48, fig. 7); E, at a diameter of 7 mm (USNM 74989d), from Smith (1932: pi. 48, fig.
10); F, at a diameter of 20.9 mm (MCZ 9527); G, at a diameter of 15 mm (MCZ 9528); H, at a diameter of 5.5 mm
(USNM 74987e), from Smith (1932: pi. 47, fig. 23); I, at a diameter of approximately 22 mm (USNM 74987c), from
Smith (1932: pi. 47, fig. 17); J, at a diameter of 15 mm (USNM 74988c), from Smith (1932: pi. 48, fig. 16); K, at a
whorl height of 5.2 mm (MCZ 9529). All specimens from Co/umb/fes fauna, Thaynes Formation of southeast Idaho; speci-
mens A, C, F, K from Montpelier Canyon; B, G, from Hot Springs; D, E, H, I, J from Paris Canyon.
372 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Table 24. Measurements of Dieneroceras apostolicus (Smith) from
Around North End of Bear Lake, Southeast Idaho.
COLUMBITES pAUiNA
No.
D
w
H
u
W/D
H/D
U/D
No.
D
w
H
U
W/D
H/D
U/D
1.
37.0?
9.2
8.5
21.8
24.9?
22.9
58.9
26.
22.0
7.7
6.3
11.1
35.0
28.6
50.5
2.
33.8
8.1
9.3
18.5
23.9
27.5
54.7
27.
21.8
5.0
5.8
11.8
22.9
26.6
54.1
3.
33.7
7.9
7.9
19.4
25.4
25.4
57.5
28.
21.4
6.3
5.8
11.9
39.4
27.1
.56.2
4.
33.6
7.5
8.2
19.3
27.3
24.4
57.5
29.
21.2
7.1
5.1
11.7
33.5
24.0
55.2
5.
32.2
?
8.6
17.2
26.7
53.5
30.
21.1
5.8
5.8
10.4
27.5
27.5
49.3
6.
31.5
7.9
7.5
18.4
25.1
23.8
58.4
31.
21.0
5.2
5.8
11.2
24.8
27.6
53.4
7.
30.5
8.7
7.7
16.7
28.5
25.2
21.9
32.
20.8
5.7
5.4
11.3
27.4
25.9
54.3
8.
29.4
6.8
7.5
15.3
23.1
25.5
52.0
33.
20.7
6.1
5.2
12.1
.38.4
25.1
58.5
9.
29.2
6.8
6.8
16.0
23.3
23.3
54.8
34.
20.0
8.3
5.4
11.4
41.5
27.0
57.0
10.
29.0
6.6
7.8
14.2
22.7
26.8
49.0
35.
19.0
5.5
5.0
10.8
28.9
26.3
56.9
11.
29.0
8.4
8.0
15.2
28.9
27.6
52.4
36.
19.0
7.4
5.0
10.3
38.9
,36.3
54.2
12.
26.7
7.0
6.8
14.8
26.2
25.4
57.6
37.
18.8
7.0
4.3
10.8
37.2
22.8
57.5
13.
25.8
6.0
7.5
13.3
23.2
29.0
51.6
38.
18.7
5.7
5.4
9.2
30.5
28.9
49.1
14.
25.5
6.3
6.4
14.0
24.7
27.2
54.9
39.
18.5
6.4
5.4
10.0
34.6
29.2
54.0
15.
25.2?
7.7
6.6
14.4
30.5
25.2
57.2
40.
18.0
5.0
4.6
9.8
27.7
25.5
54.4
16.
25.0
?
6.2
14.0
24.8
56.0
41.
17.5
5.5
4.6
9.4
31.4
26.3
.53.7
17.
25.0
6.5
5.9
14.5
26.0
23.6
58.0
42.
17.1
5.3
4.7
9.0
31.0
27.5
52.5
18.
24.7
7.5
6.9
12.4
31.6
24.3
50.2
43.
16.5
4.6
4.0
8.7
27.8
24.2
.54.3
19.
24.4
6.1
6.6
12.6
25.0
27.1
51.7
44.
16.3
5.0
4.4
9.4
30.6
27.0
.57.7
20.
24.2
7.0
6.8
12.5
28.9
28.1
51.6
45.
15.2
5.5
4.4
8.2
36.2
28.9
53.9
21.
23.5
8.8
6.8
12.4
37.4
28.9
52.8
46.
14.9
5.8
3.2
8.2
38.9
21.4
55.0
22.
22.8
5.8
6.1
12.6
25.4
26.7
55.3
47.
14.1
6.5
4.1
7.8
46.1
29.0
55.3
23.
22.7
6.5
6.9
11.4
28.6
30.4
50.2
48.
13.2
5.6
3.2
7.0
42.4
24.2
53.0
24.
22.4
p
5.6
12.0
25.0
53.7
49.
8.4
4.6
3.7
6.9
54.8
44.1
82.2
25.
22.3
5.8
5.8
12.1
26.0
26.0
54.3
50.
6.9
3.8
2.0?
3.2
.55.1
28.9?
46.4
1. Paratype, Celtites ursensis Smith (1932: pi.
6. Holotype (Smith, 1932: pi. 48, figs. 1, 2)
7. Holotype, Celtites ttrsensis Smith (1932; pi.
11. Paratype, Celtites ursensis Smith (1932: pi.
21. Holotype, Celtites phn^ovohis Smith ( 1932:
23. Paratype (Smith, 1932: pi. 48, figs. 3, 4)
34. Paratype, Celtites phiiwvolvis Smith (1932:
41. Paratype (Smith, 1932: pi. 48, figs. 5-7)
45. Paratype, Celtites ursensis Smith (1932: pi. 47, figs. 18, 19),
50. Paratype (Smith, 1932: pi. 48, figs. 8-10), USNM 74987d.
47, figs. 13, 14), USNM 74987b.
, USNM 74989a.
47, figs. 11, 12), USNM 74987a.
47, figs. 15, 16), USNM 74987c.
pi. 48, figs. 11, 12), USNM 74988u.
USNM 74989h.
pi. 48, figs. 13, 14), USNM 74988b.
USNM 74989c.
USNM 74987cl.
completely on a large number of speci-
mens, and it is quite clear that this is a
highly variable feature. This feature would
generally suggest relationship to Xeno-
celtites; however, in this case the whole
aspect of the whorls is different from that
of "typical" xenoceltitids.
Even though the descriptions and illus-
trations of Popov's two Siberian species
leave much to be desired, I believe they
are conspecific with the Idaho forms.
This species is quite similar in its basic
conch shape to D. demokidovi Popov, but
that species apparently is smooth, lacking
anv fonn of constrictions.
Occurrence. Tlie middle shale member
of the Thaynes Formation, Cohimbitcs
fauna, Paris Canyon, Montpelier Canyon,
Hot Springs, and Draney Creek, south-
eastern Idaho. The two species recorded
by Popov are from the Dieneroceras Zone
of Popov in three different localities in the
Kolyma River basin, Siberia.
Repository. Holotype USNM 749S9a (PI.
53, figs. 10," 11); paratypes USNM 74989b
(PI. 53, fig. J2), USNM 74989c (Smith,
1932: pi. ^48, figs. 5-7), USNM 74989d
(Smith, 1932: pi. 48, figs. 8-10); suture
specimens MCZ 9524 (Fig- 5A), MCZ 9525
(Fig. 5B), MCZ 9526 (Fig. 5C), MCZ 9527
Ammonoids of the Late Scythian (Lower Triassic'
Kummel
373
(Fig. 5F), MCZ 9528 (Fig. 5G), MCZ
9529 (Fig. 5K); unfigured specimens from
Montpelier Canyon MCZ 9530, from Hot
Springs MCZ 953L Holotvpe Celtites
planovohis, USNM 74988a (PI. 53, figs. 7,
8); paratype USNM 74988b (PL 53, fig. 9);
suture specimens USNM 74988c (Smith,
1932: pi. 48, fig. 16), USNM 74988d (Smith,
1932: pi. 48, figs. 17-20). Holotype Celtites
iirsensls, USNM 74987a (PI. 53,' figs. 1, 2);
paratypes USNM 74987b (PI. 53, figs. 3, 4),
USNM 74987c (PI. 53, figs. 5, 6), USNM
74987d (Smith, 1932: pi. 47, figs. 18-20),
USNM 74987e (Smith, 1932: pi. 47, figs.
21-23).
Genus Subvishnuifes Spath, 1930
Type species, Subvishnuifes welteri Spath,
1930 (= Vishnuifes sp. Welter, 1922)
Records on this genus have increased
considerably in recent years. Tlie type
species of Subvishmiites was based on a
single specimen from the Ouenites fauna of
Timor. Conspecific forms have been de-
scribed by Kummel ( 1959 : 443 ) from an
Owenites fauna of South Island, New Zea-
land, by Popov (1962b: 43 — as Paiinyoites
mastykensis) from an Owenites fauna of
the Caucasus Mountains, and from an
Owenites fauna of Afghanistan (Kummel
and Erben, 1968). In addition, the Owenites
Zone of Kwangsi, China, contains Sf//;-
vishntiites tientiingensis Chdo (1959). The
specimen from the Dieneroceras Zone of
Siberia (Popov 1962a) described as Inyoites
eiekifensis is a species of this genus. The
Columhitcs Zone of southeast Idaho has
yielded one fragmentary specimen that is
described here. This specimen is quite like
the Siberian S. eiekitensis. Finally, the
Narmia Member of the Mianwali Forma-
tion in the Trans-Indus Surghar Range,
West Pakistan, has yielded fragmentary
representatives of this genus described as
S. sp. indet. (Kummel, 1966). Tliis horizon
contains Procornites kokeni, ProJiuniiarites
cf. crasseplicatus, etc., and is late Scythian
in age. Xenaspis enveris Arthaber (1911)
from the Suhcolumbites fauna of Albania
is considered here to be a species of Suh-
vishnuites. The Subcoltimbites fauna at
Kotal-e-Tera, Afghanistan, has also yielded
a couple of small specimens that have been
recorded as Subvislinuites sp. indet. (Kum-
mel, 1968a). There is also an interesting
allied species described as Subvishmiites cf.
enveris (Kummel, 1968b) from the Subco-
ltimbites fauna of Kotal-e-Tera, Afghanistan.
This genus thus ranges in age through
the whole of the upper half of the Scythian.
As yet few specimens have been recovered
and few species described.
Subvishnuifes enveris (Arthaber)
Text-figure 4
Xenaspis enveris Arthaber, 1911: 230, pi. 20(4),
fig. 3; Dicner, 1915: 311; Spath, 1934: 293.
The type and only specimen of this
species is not in the collection of the
Paleontological Institute, Vienna. The re-
lationships of this species and its assign-
ment to SitJ)vishniiifes are discussed fully
below in the description of Subvishnuites
cf. enveris from Afghanistan. Spath (1934:
293) considered this species to have a re-
lationship to EophyUites, but this I find
difficult to understand.
Occurrence. Subcolumbites fauna, Kcira,
Albania.
Repository. The specimen is apparently
lost, as it is not in the Paleontological In-
stitute, Vienna.
Subvishnuifes cf. enveris (Arthaber)
Subvishnuites cf. enveris (Arthaber), Kummel,
1968b: 491, pi. 1, figs. 8, 9.
This is a most interesting specimen from
a Subcohimbifes fauna at Kotal-e-Tera,
Afghanistan. The specimen measures 45
mm in diameter, approximately 20 mm for
the width of the adoral whorl, 21 mm for
the height, and 11.7 mm for the width of
the umbilicus. The whorl sides are broadh'
arched, converging on to a rounded venter.
The whorl sides bear widely spaced radial
ribs that commence and are most con-
spicuous on the umbilical shoulder and
decrease in intensity toward the venter
374 Bulletin Miisctini of Comparative Zoology, Vol. 137, No. 3
which is smooth. The adoral half volution
has four such ribs. The ribs are also present
on the inner whorls as far as they are pre-
served. The umbilical shoulder is abruptly
rounded and the umbilical wall, nearly
vertical. The suture is not preserved.
Arthaber's specimen of Xenaspis enveris
is slightly more evolute than my Afghan
specimen ( 34 percent versus 27 percent )
and has an acute venter on the adoral part
of the living chamber. The ribbing, ac-
cording to Arthaber, is developed only on
the living chamber. The absence of the
ribs on the phragmocone could well be a
matter of preparation or preservation. The
suture of the Allxmian specimen consists
of two denticulated lateral lobes (Arthaber,
1911: pi. 20(4), fig. 3c).
It appears quite probable that the Al-
banian Xenaspis enveris is not conspecific
with this Afghan specimen, though they
are most probably congeneric. However,
the assignment of these two specimens may
be open to question. The type specimen
of Su])vis]intiites is a smooth form with an
acute venter as are all the other specimens
assigned to this genus. Tlie Albanian and
Afghan specimens have prominent radial
ribs. A case could be made that the Al-
banian and Afghan species are generically
distinct from the more typical species of
Su1)vis]inuites. However, data are so in-
complete on both the Albanian and Afghan
species that it would be imprudent to
establish a new genus with either of these
specimens as type. Because of these factors,
and because there are no other late Scythian
genera to which these specimens have any
similarity, it seems best to assign them to
Suhvishntiites.
Occttrrenee. Suhcolunihites fauna, Kotal-
e-Tera, Afghanistan.
Repositorij. MCZ 1()14<S.
Subvishnuites eiekitensis (Popov)
Inyoites eickitcii.sifi Popov, 19fi2a: IcS4, \i\. 3, fiu.
5, text-fig. 6.
The description and illusliatioii ol this
species leave much to be desired; even so,
the whole appearance of the conch and the
plan of the suture show this species to be
a member of SuJ)vis]inuites. This species is
very similar in general appearance to Sub-
vishnuites sp. indet. from the Cohimbifes
Zone of southeast Idaho. However, that
species is known only from a single frag-
mentary specimen, and data are insufficient
to evaluate the relationships of the two
forms.
Occurrenee. Dieneroceros Zone of Popov
(1961, 1962a), Lena and Olenek river
basins, Siberia.
Subvishnuites sp. indet.
Plate 53, figure 15
A single fragmentary specimen of ap-
proximately 34 mm in diameter. The
whorls are compressed, convex, converging
on an acute venter. The lateral areas bear
weak, broad, radial folds which are more
apparent on the cast than they are on the
shell. Tlie umbilical shoulders are broadly
rounded. The suture is only vaguely visible
but it does show two lateral lobes.
This specimen is very similar in its form,
etc., to S. eiekitensis (Popov) from the
Dieneroeeras Zone of Siberia. It could well
be conspecific with that species but data
are not sufficient to allow a detailed analy-
sis.
Oeeurrenee. Coluinhites Zone, Thaynes
Formation, Montpelicr Canyon, southeast
Idaho.
Repository. MCZ 9512.
Genus Hemilecanites Spath, 1934
Type species, Lecanites discus Arthaber,
1908
Hemilecaniies discus (Arthaber)
Plate 25, figures 9, 10; Text-figure 4
Lccaiiitcs disctis ArtlialxT, 1908: 268, pi. 11(1),
figs. .5a-c; ArtlialuT, 1911: 181, 238; Renz and
Rcnz, 1947: (>1; Hni/. and Ren/., 1948: 55.
Proavitcs di.scu.s, — Dinner, 1915: 228.
ilctiiilccanitcs discus, — Spath, 1934: 135, p\. 13,
ligs. 7a-d; Kunnncl, in Arkell vt al., 1957: L136,
ligs. 109, 3: Chao, 1959: 41, 196, pi. 3, figs. 1,2.
Ammonoids of the Late Scythian (Lower Triassic) • Ktanmel 375
C elates kcirensis, — Renz and Renz ( non Aithaber ) ,
1948: 42, pi. 3, fig. 3 (non fig. 6).
This is another very distinctive species
first described from the Siibcohnnhites
fauna of Albania. Unfortunately, none of
Arthaber's types are preserved in the
Paleontological Institute, Vienna. The speci-
men recorded by Renz and Renz (1948:
55) is illustrated here on Plate 25, figures
9, 10. Tlie inner whorls of this specimen,
up to a diameter of 14 mm, are exposed,
and these show the whorl section to be
approximately as wide as it is high and
the venter to be broadly rounded. It is
in the following volutions that the whorls
become compressed and the venter acute.
The sutiu-e is nearly identical to that of
Arthaber's holotype (Fig. 4E). Another
representative of this species in the Chios
fauna is one of the specimens Renz and
Renz (1948: 42, pi. 3, fig. 3) assigned to
Ccltites kcirensis.
The specimen recorded by Chao ( 1959 )
yielded no suture, but the distinctive
morphology of this species suggests that
the specimen from Kwangsi, China, is con-
specific with the Albanian and Chios speci-
mens of H. discus.
The only other species of Hemilccanites
is H. paradiscus n. sp. from the Subcolum-
bites fauna of the Tobin Fonnation, Ne-
vada. This new species is quite similar to
H. discus, but it is a more compressed and
involute form with commensurate changes
in the localization of the sutural elements
(Fig. 4F).
Occurrence. Siibcolumbites fauna of Al-
bania, Chios, and Kwangsi, China (Chao
collection 610).
Repository. The holotype is supposed to
be in the Paleontological Institute, Vienna,
but has not been located and is presumed
lost; however, four topotype specimens are
in the British Museum (Natural History),
C22875-8. The plesiotype (PI. 25, figs.,
9, 10) from Chios is NHMB J13703; the
plesiotype of Celtites kcirensis, — Renz and
Renz (non Arthaber) 1948: pi. 3, fig. 3,
is NHMB J13653.
Hemilecanites paradiscus n. sp.
Plate 29, figures 11, 12; Plate 31,
figures 15, 16; Plate 35, figure 12;
Text-figure 4
There are nine, mostly incomplete, speci-
mens in the collection from the Tobin For-
mation, only two of which are complete
enough to yield the following measure-
ments :
D W H U W/DH/DU/D
Holotype
(MCZ 9465) 2L1 3.4 8.0 6.8 16.2 37.9 32.2
Paratype
(MCZ 9451) 15.8 3.2 6.7 4.8 20.2 42.4 30.3
The conch is much compressed, lenticular
in cross section. The venter is narrowly
rounded to oxynote. The flanks are only
slightly convex and the umbilical shoulder
and wall very low. The shell is perfectly
smooth except for extremely fine radial
growth lines. None of the specimens are
sufficiently complete to show the apertural
regions. The sutures are shown on Figure
4F, G.
There is no question bvit this species is
closely related to H. discus. It differs in
its greater compression and involution and
in the spacing of the sutural elements.
Occurrence. Tobin Formation, Pershing
County, Nevada; south tip of Tobin Range,
Cain Mountain 1:62,500 quad., center NW
% sec. 9, T. 26N, R. 39E, 5,500 ft. S., 27.5
ft. W from elevation point 5088 on range
crest.
Repositori/. Holotype MCZ 9465 (PI. 31,
figs. 15, 16); paratype MCZ 9451 (PL 29,
figs. 11, 12), MCZ 9473 (PL 35, fig. 12);
suture specimens MCZ 9631, 9483; unfig-
ured paratypes MCZ 9483.
Superfamily NORITACEAE Karpinsky, 1889
Family XENOCELTITIDAE Spath, 1930
Genus Xenocelfites Spath, 1930
Type species, Xenoceltifes subevolutus
Spath, 1930
The genus Xenoceltifes is very widely
distributed in the mid-Scythian Owenites
Zone where it is represented by a fairly
376 BiiUetin Museum of Comparative Zoology, Vol. 137, No. 3
large number of species. In the overlying
CoUimhites Zone the genus is represented
by a single species (X. spencei) known only
from southeast Idaho. In the late Scythian
ProhuHiiarites Zone there are only three
species, one each from the Salt Range,
China, and Primorye Region. In each case
these species are represented by very few
specimens. An indeterminant species of
this genus has been recorded from the
Subcolumbites fauna at Kotal-e-Tera, Af-
ghanistan (Kummel, 1968b).
Xenoce/f/tes sinuafus (Waagen)
Dinarites sinuatus Waagen, 1895: 33, pi. 10, fig.
4; Diener, 1915: 122.
Xcnoccltites .sinuatus Cliao, 1959: 194; Kummel,
1966: 389, pi. 1, figs. 1-8.
Lecanites laqueus Waagen, 1895: 285, pi. 38,
figs. 9, 10.
Xcnodiscus laqueus Diener. 1915: 313.
This species has been recently described
and illustrated (Kummel, 1966: 389) on
the basis of new specimens. Among the
few upper Scythian species of Xenoceltites
known, this species is most similar to X.
c reno ve nt wsii.s- Chdo (1959) from Kwangsi,
China. There is close agreement on the
suture and general conch shape, but the
pattern of ornamentation is different.
Occurrence. Sandstone beds of Narmia
Member of Mianwali Formation, just above
hard Bivalve Limestone, Chhidru Nala,
Salt Range, West Pakistan.
Repo.s-itory. Holotype GSI 7110; topo-
types MCZ 9581, 9582.
Xenoceltites crenoventrosus Chao
Xenoceltites crenoventrosus Chao, 1959: 38, 194,
pi. 3, figs. 14-15, pi. 42, figs. 2-6, text-fig.
8a-e.
The description and illustraticm of this
species are inadeciuate. The conch is of tlK>
general form of X. .sinuatus, but the ribs
cross the venter forming a disthictive
crenulated pattern. The suture is quite
like that of X. .sinuatus.
Occurrence. SuhcoUnnhites fauna,
Kwangsi, China (Chao collection 542b).
Xenoceltites spitsbergensis Spath
Xenoceltites spitsbergensis Spath, 1934: 128, pi.
9, figs. 1, 2, pi. 11, figs. 5, 7, 8; Kiparisova,
1961: 50, pi. 9, figs. 7, 8.
This species is recognized on the basis
of two very small specimens of 21 and 15
mm in diameter. Even so, the similarity to
the Spitsbergen xenoceltitids is quite strik-
ing, and I can do no more than agree with
Kiparisova's detemiination. The type speci-
mens of this species, from Spitsbergen, are
of mid-Scythian Oacnites Zone age and
this specimen is said to have come from
strata with Subcolumbites.
Occurrence. Subcolumbites fauna, Pri-
morye Region, Siberia.
Xenoceltites spencei (Hyatt and Smith)
Plate 48, figures 5-9; Plate 52, figures
1-7; Text-figures 7, 8
Ophiccras spencei Hvatt and Smith, 1905: 119,
pi. 62, figs. 1-10; Diener, 1915: 212; Smith,
19.32: .50, pi. 62, figs. 1-10.
The middle shale member of the Thaynes
Formation around Bear Lake, southeast
Idaho, that contains the Columbites fauna
has yielded an abundance of specimens of
this species. Measurements of 99 speci-
mens are given in Table 25 and the varia-
tions in whorl width and height are plotted
on Figure 8. The forward projecting con-
strictions that cross the venter are quite
variable in their intensity. As noted b>-
Smith (19.32: 50) these constrictions are
most noticeable on the cast; in many of
the specimens they arc not at all apparent
or expressed on the shell. The suture con-
sists of two denticulated lateral lobes and
is quite variable in details of the basic
pattern (Fig. 7).
This species is not really comparable or
very close to any of the upper Sc\thian
species assigned to Xenoceltites, as .sinuatus,
crenoventrosus, or tlu> species assigned by
Kiparisova (1961) h) X. s})it.sbergen.sis. It
is however, quite similar in its over-all
aspect to the Spitsbergen forms Spath
(19.31) assigned to X. subeiolutus, X. gre-
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 377
Figure 7. Diagrammatic representation of the sutures of Xenoce/f/tes spencer (Hyatt and Smitfi), from tfie Columbites fauna,
Thoynes Formation of southeastern Idaho. A, at a diameter of 17 mm (MCZ 9558); B, at a diameter of 21 mm (MCZ
9559); C, at a diameter of 18 mm (MCZ 9560); D, at a diameter of 22 mm (MCZ 9561); E, at a diameter of 13 mm
(MCZ 9562); F, at a diameter of 24 mm (MCZ 9563); G, at a diameter of 34 mm (MCZ 9551, PI. 52, fig. 1); H, para-
lectotype (Hyatt and Smith, 1905: pi. 62, figs. 5-7; PI. 48, figs. 5, 6 of this report), at a diameter of 28 mm (USNM 75291b).
Specimens of figures A, C, D, from Hot Springs, B, E, F, G from Montpelier Canyon, H, from Paris Canyon.
gonju or X. spitsbergensis, all of mid-
Scythian, Owenites Zone, age.
Occiincnce. Middle shale member of
Thaynes Formation, Columbites Zone, at
Paris Canyon, Montpelier Canyon, and Hot
Springs, Bear Lake region, and Draney
Creek, southeast Idaho.
Repository. Lectotvpe (PI. 48, figs. 7-9)
USNM 75291a; paralectotype (PI. 48, figs.
5, 6) USNM 75291b, paralectotype (Hyatt
and Smith, 1905: pi. 62, figs. 8-10) USNM
75291c; plesiotvpes (PI. 52, fig. 1) MCZ
9551, (PL 52, fig. 2) MCZ 9552, (PI. 52,
fig. 3) MCZ 9553, (PI. 52, fig. 4) MCZ
9554, (PI. 52, fig. 5) MCZ 9555, (PI. 52,
fig. 6) MCZ 9556, (PI. 52, fig. 7) MCZ
9557; suture specimens Figure 7, MCZ 9551,
9558-9563; specimens from Montpelier Can-
yon MCZ 9630, from Hot Springs MCZ
9629.
378 Bulletin Museum of Cotnjniidtive Zoology, Vol. 137, No. 3
Table 25. Measurements of Xenoceltites spencei (Hyatt and Smith) from the Columbites
Fauna, Bear Lake Region, Southeastern Idaho.
D
W
H
U
W/D
H/D
U/D
D
W
H
u
W/D
H/D
U/D
1.
51.5
?
16.2
19.8
?
31.5
38.4
47.
20.1
5.8
7.8
6.6
28.9
,38.8
32.8
2.
49.8
10.5
14.5
21.8
21.1
29.1
43.8
48.
20.0
5.2
7.6
6.4
26.0
,38.0
,32.0
3.
46.4
11.0
16.2
19.2
23.7
.34.9
41.4
49.
20.0
5.3
8.2
6.6
26.5
41.0
,33.0
4.
45.8
11.4
17.4
16.1
24.9
37.9
35.2
50.
20.0
6.3
8.0
6.7
31.5
40.0
33.5
5.
42.3
9.4
15.0
16.3
22.2
35.5
38.5
51.
19.8
5.6
8.2
6.3
28.3
41.4
31.8
6.
42.3
10.5?
13.8
18.6
24.8?
,32.6
43.9
52.
19.8
5.5
7.8
6.9
27.8
.39.4
.34.8
7.
40.8
10.9
15.7
13.4
26.7
.38.5
32.8
53.
19.7
5.8
7.3
7.0
29.4
,37.1
,35.5
8.
40.7
10.0
14.4
15.2
24.6
35.4
37.3
54.
19.7
6.0
8.5
5.7
30.5
43.1
28.9
9.
40.0
10.0
16.0
12.2
25.0
40.0
30.5
55.
19.7
6.6
7.2
6.1
33.5
,36.5
30.9
10.
39.4
9.4
12.7
17.3
23.9
32.2
43.9
56.
19.5
6.0
8.3
5.9
30.8
42.6
,30.3
11.
.33.4
9.0
15.0
8.7
26.9
44.9
26.0
57.
19.5
5.7
8.2
5.7
29.2
42.1
29.2
12.
.33.2
7.8
10.8
14.0
23.5
32.5
42.2
58.
19.2
6.2
8.0
5.9
32.2
41.7
,30.7
13.
31.8
7.8
12.2
11.3
24.5
38.4
35.5
59.
19.1
6.0
8.7
5.2
31.4
45.5
27.2
14.
31.7
8.2
12.3
10.3
25.9
38.8
32.5
60.
19.0
6.0
7.2
6.5
31.6
37.9
34.2
15.
30.6
8.1
10.6
12.6
26.5
34.6
41.2
61.
19.0
6.0
7.3
6.4
31.6
,38.42
33.7
16.
30.5
6.6
10.2
12.6
21.6
.33.4
41.3
62.
19.0
6.2
8.0
5.6
.32.6
42.1
29.5
17.
29.7
8.3
13.1
7.7
27.9
44.1
25.9
63.
18.8
5.6
7.7
6.0
29.8
40.9
31.9
IS.
29.2
7.5
10.7
10.8
25.7
.36.6
.36.9
64.
18.4
5.6
7.1
6.6
.30.4
.38.9
35.9
19.
29.0
7.4
11.4
9.7
25.5
.39.3
33.4
65.
18.2
5.7
7.0
6.3
31.3
38.5
.34.6
20.
28.8
7.6
10.2
11.0
26.4
35.4
38.2
66.
18.0
4.6
7.1
6.5
25.6
,39.4
36.1
21.
26.7
7.5
9.8
10.0
28.1
.36.7
37.5
67.
17.8
5.8
7.3
5.7
,32.6
41.0
32.0
22.
26.5
7.1
10.2
9.3
26.8
38.5
35.1
68.
17.8
5.5
7.1
6.1
30.9
39.9
34.3
23.
25.4
7.2
9.0
9.4
28.3
35.4
.37.0
69.
17.5
5.3
6.6
6.4
30.3
.37.7
36.6
24.
25.3
6.8
6.8
9.6
26.9
26.9
37.9
70.
17.5
5.8
6.3
6.8
33.1
,36.0
38.9
25.
25.3
6.9
10.6
8.3
27.3
41.9
32.8
71.
17.5
5.7
7.2
5.4
32.6
41.1
,30.9
26.
25.3
7.3
9.7
9.3
28.9
38.3
36.8
72.
17.5
5.2
6.1
6.7
29.7
,34.9
.38.3
27.
24.6
6.4
9.4
8.7
26.0
38.2
35.4
73.
17.4
4.8
7.2
6.1
27.6
41.4
,35.1
28.
24.3
6.8
9.1
9.8
27.9
37.4
40.3
74.
17.3
5.4
7.4
5.3
31.2
42.8
30.6
29.
24.0
6.8
10.2
7.0
28.3
42.5
29.2
75.
17.2
5.5
7.0
.5.6
31.9
40.7
32.6
30.
23.5
?
10.0
7.0
?
42.6
29.8
76.
17.1
5.0
6.5
5.3
29.2
38.0
30.9
31.
23.5
6.7
8.3
9.3
28.5
35.3
39.6
77.
17.0
6.0
6.8
5.1
35.3
40.0
30.0
32.
23.3
6.5
9.4
7.4
27.9
40.3
31.8
78.
16.9
5.3
6.7
6.1
31.4
.39.6
,36.1
33.
23.3
6.5
9.3
8.0
27.9
39.9
34.3
79.
16.8
5.5
6.5
6.0
32.7
38.7
35.7
34.
23.3
6.4
9.6
7.8
27.5
41.2
33.5
80.
16.6
5.2
5.6
7.1
31.3
33.7
42.8
35.
23.2
6.2
9.0
6.2
26.7
38.8
26.7
81.
16.5
5.1
6.2
5.8
30.9
.37.6
,35.2
36.
23.0
6.4
8.7
8.2
27.8
.37.8
35.7
82.
16.3
5.3
6.3
5.3
.32.5
,38.7
32.5
37.
23.0
6.3
9.1
7.2
27.4
.39.6
31.3
83.
1.5.9
5.2
6.7
5.0
32.7
42.1
31.4
.38.
21.7
?
9.0
6.9
?
41.5
31.8
84.
15.6
5.2
6.7
4.8
33.3
42.9
30.8
39.
21.7
6.2
8.3
8.2
28.6
38.2
.37.8
85.
15.2
5.1
6.2
4.6
.33.6
40.8
30.3
40.
21.5
7.5
9.4
6.6
86.
15.0
5.3
6.1
5.0
35.3
40.7
33.3
41.
21.3
6.6
9.0
7.1
30.9
42.3
33.3
87.
1.5.0
5.2
5.7
5.7
,34.7
.38.0
,38.0
42.
21.1
6.3
8.1
7.6
29.9
38.4
36.0
88.
14.8
5.1
5.9
.5.1
,34.5
.39.9
.34.5
43.
21.0
6.3
9.7
5.0
30.0
46.2
23.8
89.
14.7
4.7
5.7
.5.5
31.9
38.8
.37.4
44.
20.8
5.2
8.7
6.4
25.0
41.8
30.8
90.
14.6
4.5
6.1
5.0
30.8
41.8
34.2
45.
20.7
6.0
9.4
6.1
28.9
45.4
29.5
91.
14.6
5.2
5.6
4.8
35.6
38.4
32.9
46.
20.6
5.4
8.4
6.8
26.2
40.7
.33.0
92.
14.4
4.9
5.8
4.4
34.0
40.3
30.6
4.
7.
9.
21.
23.
24.
.36.
59.
7fi.
Plcsiotvpc, MCZ 9.5.51 (PI. .52, fig. 1).
Parah'ctotype, USNM 75291b (PI. 48, fi«s. 5,
LcctotMX', U.SNM 75291a (PI. 48, figs. 7-9).
MCZ 9550 (PI. 52, fig. «).
6).
Pk'siot) pf
Pksiotype
Pksiotype
Suture
Suturt-
MCZ
MCZ
specimen,
specimen.
.Suture specimen,
Plesiotype, MCZ
9554
9552
MCZ
MCZ
MCZ
9555
(PI. 52, fig.
(PI. ,52, tin.
9563 (Fiji.
9561 (Fin.
9560 (FiK.
(PI. 52, fin.
4).
2).
7F).
7D).
7C).
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 379
Table 25. Coi^tinued
D
w
H
u
W/D
H/D
U/D
D
w
H
U W/D
H/D
U/D
93.
14.2
4.8
4.8
5.3
33.8
33.8
37.3
97.
12.2
4.1
4.2
4.0? 33.6
.34.4
.32.8?
94.
14.0
4.9
5.1
5.4
35.0
36.4
38.6
98.
11.0
4.0
4.2
3.9 36.4
.38.2
28.0
95.
13.6
4.9
5.2
4.8
36.0
38.2
35.3
99.
8.3
3.1
2.8
3.2 37.3
33.7
38.6
96.
13.4
4.5
5.1
4.8
33.6
38.1
35.8
96. Suture specimen, MCZ 9.562 (Fig. 7E).
98. PlesioHpe, MCZ 9557 (PI. 52, fig. 7).
Genus Preflorianifes Spath, 1930
Type species, Danubifes strongi Hyatt and
Smith, 1905
Evokite forms with arched venters, often
tendmg to become acute; with radial
ribbing, most pronounced on inner lateral
areas near umbilical shoulders and not
crossing the venter; suture with two lateral
lobes.
Preflorionites is quite a generalized am-
monite. This, accompanied by the fact that
it is not particularly abundant in the upper
half of the Scythian, has made analysis
of the species described to date very diffi-
cult. Tlie late Scythian species of Pre-
florianifes recognized here are:
Prcflorianitcs sulioticus ( Arthaber)
Preflorianites ^arhinus ( Renz and Renz)
Prcflorianitcs tniiJtipIicattis ( Mojsisovics )
Prcflorianitcs intermedins Tozer
In addition, the CoJumhites fauna of
southeast Idaho has yielded one species,
P. montpelierensis.
I am not at all certain as to the merits of
all these species. Each is known from only
one or very few specimens; thus little or
no data are available on the range of varia-
tion in rib patterns, degree of involutions,
shape of whorl sections, suture, etc. Few
meaningful comparisons can be made be-
tween one species group and the others.
The late Scythian species are known from
the Snbcohimhifes fauna of Albania (siilio-
ficiis), the Sii])coJumJ)ites fauna of Chios
(sidioticiis, fi,arbiniis), the Olenekifes fauna
of the Olenek River region ( multiplicattis ) ,
from a late Scythian fauna of British Co-
lumbia (intermedins), and from the Co-
Jumbites fauna of southeast Idaho {montpe-
lierensis).
Preflorianifes sulioficus (Arthaber)
Plate 4, figures 5, 6; Plate 19, figures
1, 2, 5-8; Text-figure 9
Xcnodiscus snlioticus Arthaber, 1911: 229, pi.
19(3), fifis. 6a, b, pi. 20(4), figs. 2a, b;
Diener, 1915: 315; C. Renz, 1928: 155; Renz
and Renz, 1947: 61; Renz and Renz, 1948:
56, pi. .3, figs. 1, 2.
Preflorianites snlioticus, — Spatb, 1934: 133, pi.
12, figs. 2a-d.
Ophiccras cfr. Nangaensis Arthaber (1911: 239,
pi. 21(5), fig. 5) non Waagen.
Xcnodiscus sp. ind. aff. Nangaensis, — Diener,
1915: 313.
Arthaber (1911: 230) states he had four
specimens of this species; only the two
specimens he illustrated are still preserved.
It is of interest that whereas Arthaber had
only four specimens for study, the British
Museum ( Natural History ) has fifteen topo-
types obtained by purchase. The descrip-
tion of the species by Arthaber (1911) and
the comments by Spath ( 1934 : 133 ) are
adequate for the two syntypes even though
the specimens are poorly preserved. The
specimens of this species figured by Renz
and Renz (1948, pi. 3, figs. 1, 2) are very
typical fonns. They clearly show that the
adoral decrease in ribbing intensity is
highly variable, taking place at different
growth stages.
The two specimens which Arthaber
(1911) assigned to Ophiccras cfr. nangaensis
are both poorly preserxed and owe many
of their morphological features to excessive
grinding and polishing. One of the speci-
mens (PI. 19, figs. 1, 2) shows indications
380 BuUctin Museum of Comparative Zoolo'^ij, Vol. 137, No. 3
18
17
16
15
14
13
12
10
9
8
7
6
5
4
• - • • •
I • • •
•• • •
:.
X X
• • •
X „x
X
X X
X
• »xX X
-• .X X ^
• • ^ X XX V
• • . xx X 'y
r M
XKX
X^ X
X
XX
X'
X X
XX
X X
1
1
H
w
I I I I \ 1 I I
0 5 10 15 20 25 30 35 40 45 50 55 SO
DIAMETER
Figure 8. Variation in whorl height (H) and whorl width (W) of Xenoce/tites spencei (Hyatt end Smith) from Columbites
fauna, Bear Lake region, southeast Idaho. The data on thi; graph are from Table 25.
Ammonoids of the Late Scythian (Lower Triassic) • Ktimmel 381
A
A
Figure 9. Diagrammatic representation of the suture of: A, Preilorianiles su/iof/cus (Arthaber, 1911: pi. 20(4), fig. 2b),
from Subco/umbifes fauna of Albania, at a diameter of approximately 20 mm; B, Preilorianites garbinus (Renz and Renz),
from Subco/umb/'tes fauna of Chios (NHMB J13696), at a diameter of 25 mm; C, Preilorianites montpelierensis n. sp., from
Co/umbites fauna, southeast Idaho (MCZ 9498), at a whorl height of 5 mm; D, Pretlorianites strongi (Hyatt and Smith,
1905: pi. 9, fig. 6), from Meelcoceras fauna, Inyo Range, California, at a diameter of approximately 30 mm.
of ribs on the umbilical shoulder; the
ribbing has been completely ground away
on the flanks. Both these specimens I be-
lieve are representatives of P. sulioticus.
A suture of Prcflorianites sulioticus is il-
lustrated on Figure 9 A.
Occurrence. Siibcohnnbites fauna of Al-
bania and Chios.
Repository. The two syntypes from Al-
bania and the two specimens of Ophiceras
cfr. non^aensis are in the Paleontological
Institute, Vienna. The specimens from
Chios are NHMB J13704 (Renz and Renz,
1948, pi. 3, fig. 1), NHMB J13705 (Renz
and Renz, 1948, pi. 3, fig. 2); unfig-
ured specimens from Maradovuno NHMB
J 13706.
Preflorianites garbinus (Renz and Renz)
Plate 20, figures 10, 11; Text-figure 9
Imjoites n. sp. C. Renz, 1947: 176.
Intioites garbinus Renz and Renz, 1947: 60, 76;
Renz and Renz, 1948: 53, pi. 12, figs. 10-lOb.
The single specimen in the Chios fauna
studied by the Renzes for which they in-
troduced this new species name is indeed
superficially similar to Imjoites. However,
the venter is acute but does not bear a
keel as in the typical Imjoites from the
Meekoceras faima of western North Amer-
ica. Likewise, the typical Imjoites have a
vertical umbilical wall whereas this Chios
specimen has a rounded umbilical wall.
The specimen is slightly weathered, and
thus the fine denticulations of the suture
are lost and the suture as drawn (Fig. 9B)
is that of the weathered surface. A second
specimen in the Chios collection but not
mentioned by the Renzes is illustrated here
on Plate 20, figures 10, 11. It is a slightly
more inflated form than the holotype but
even more preflorianitid in aspect.
It is for the above reasons that this
species is believed to be a much compressed
species of Preflorianites. The Chios fauna
contains a very "typical" species of Pre-
florianites— P. sulioticus (Arthaber) — which
has a much more inflated whorl section.
Unfortunately, P. sulioticus is represented
by only two specimens in the Chios fauna,
and P. fS,arhinus by only two specimens;
thus the relationship between these two
382 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
species must remain uncertain until more
material becomes available.
Occurrence. Subcolumbites fauna, Mara-
dovuno, Chios.
Repositon/. Holotype NHMB J13696;
paratype (PI. 20, figs. 10, 11) NHMB
J 13697.
Preflorianifes multiplicatus (Mojsisovics)
Cerotites multiplicatus Mojsisovics, 1886: 25, pi.
9, figs. 15a, h.
Xenodiscus multiplicatus. — Diener, 1915: 313;
Popov, 1961: 7.
"Ceratites" multiplicatus, — Spatli, 1934: 128.
XenoccltUcs multiplicatus, — Kiin^mel, 1961: 521.
Ceratites fissi}>licatus Mojsisovics, 1886: 26, pi.
9, figs. 18a, b, 19c.
Xenodiscus fissiplicatus, — Diener, 1915: 312;
Popov, 1961: 7.
Ceratites discretus Mojsisovics, 1886: 27, pi. 9,
figs. 20a-c.
Xenodiscus discretus, — Diener, 1915: 312; Popov,
1961: 7.
Xenoceltites discretus, — Kiimmcl, 1961: 521.
Ceratites ht/pcrhoreus Mojsisovics, 1886: 26, pi.
9, figs. 16, 17.
Xenodiscus hyperhorcus, — Diener, 1915: 313.
Xenoceltites hijperhoreus, — Kummel, 1961: 521.
It is apparent from the five specimens
illustrated by Mojsisovics ( 1886, pi. 9, figs.
15-20) and brought together here as repre-
senting a single species that there is con-
siderable variability in the ribbing patterns.
The Olenek fauna has recently been mono-
graphed by Popov (1961), but in this ex-
tensive revision none of these species are
described or illustrated; they are merely
mentioned in a summary list of species from
the Olenekites Zone.
Occurrence. Olenekites Zone in region
of Ol(Mick River, northern Siberia.
Preflorianifes intermedius Tozer
Prcflorianites intcrmcdius Tmer, 1965a: 18, pi. 11,
figs. 92-6, text-fig. 2.
This species is based on a single frag-
mentary specimen and in outward appear-
ance is very much like nearly all other
upper Scythian specimens ot this genus.
The smallness of all the samples, however,
prevents any meaningful comparisons.
Occurrence. Toad Formation, Halfway
River area, British Columbia, associated
with Popovites occidentalis and Mona-
canthites monoceros.
Preflorianifes monfpelierensis n. sp.
Plate 43, figures 2, 3; Plate 44, figures
11-13; Text-figure 9
The Cohitnbites fauna of southeastern
Idaho has yielded thirteen specimens of
this interesting species. The conch is
evolute, compressed, and small. The inner
volutions have whorls which are approxi-
mately as wide as high with rounded ven-
ters, and slightly convex lateral areas and
rounded umbilical shoulders. The lateral
areas bear radial ribs that are projected
forward on the ventral shoulder. In general
the venter is smooth, but occasionally
there are constrictions crossing the venter,
constrictions which are the continuation
of rib interspaces on the lateral areas.
The number and prominence of these
constrictions are highly variable; the most
pronounced development is in the speci-
men shown on Plate 43, figiu-es 2, 3.
Tlie length of the body chamber is not
known for sure, but it is at least more than
half a volution. The body chamber is much
more compressed than the phragmocone
and the venter becomes narrowly rounded,
the lateral areas convergent. Likewise, the
radial rilxs become very subdued.
The suture is of a simple ceratite plan
with t\vo lateral lobes (Fig. 9C).
This species is not unduly different from
the other upper Scythian species of Pre-
florianites. In its somewhat compressed
whorls this species is especially like P.
garhinus from the Subcolumbites fauna of
Chios.
Occurrence. Middle shale member of
Thaynes Formation, Cohnnbites fauna at
Montpelier Can)oii and Hot Springs, south-
east Idaho.
Repositon/. Holotype, MCZ 9494 (PI.
44, fig. 13); figured paratvpes MCZ 9495
(PI. 43, figs. 2, 3), MCZ 9498 (PI. 44, fig.
12), MCZ 9635 (PI. 44, fig. 11); unfigured
paratvpes from Montpelier Can\'on MCZ
9497,' from Hot Springs MCZ 9496.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 383
A
Figure 10. Diagrammatic representation of the suture of: A, holotype of Pseudaspidites popovi n. sp., from Columbiles
fauna of southeast Idaho, at a diameter of 48 mm (MCZ 9575); B, paratype Pseudaspidites popovi n. sp., from Columbiles
fauna of southeast Idaho, at a diameter of 28 mm (MCZ 9636); C, holotype Pseudaspidites pos/erius (Popov, 1961: fig.
9), from Dienercceros Zone, northern Siberia; D, Pseudospidites yudishthira, — Krofft and Diener (1909: pi. 15, fig. 5), from
Hedemtroemia fauna of the Himalayas.
Family PARANORITIDAE Spath, 1930
Genus Pseudaspidifes Spath, 1934
Type species, Aspidites muthianus Krafft
and Diener, 1909
Pseudaspidifes popovi n. sp.
Plate 44, figures 14, 15; Plate 55,
figures 8, 9; Text-figure 10
This species is based on two specimens
that are all phragmocone. The larger speci-
men and holotype has an approximate
diameter of 63 mm, the width of the most
adoral part of the last volution is 14.8 mm,
the height is 34.7 mm, and the diameter
of the umbilicus is 4.9 mm. Tlie venter is
narrowly rounded, the lateral areas broadly
arched. Tlie umbilical shoulders are sharply
rounded. The lateral areas bear slightly
sinuous, narrow, low ribs and fine growth
lines. The suture is shown on Figure lOA.
The smaller specimen measures 32.3 mm
in diameter, 7.8 mm for the width of the
adoral whorl, 17.4 mm for the height, and
4.5 mm for the diameter of the umbilicus.
The suture is shown on Figure lOB.
The general configuration of the conch
and the suture clearly allies this species
with Pseiidaspiditcs. This genus is a fairly
common member of the mid-Scythian
Owenites Zone; species of it are known
from the Mcekoceras Zone of Idaho and
Nevada (P. wheeleri Kummel and Steele,
1962: 673), from the Hedenstrocmia fauna
of the Himalayas (P. muthianus and P.
yudishthira, — Krafft and Diener, 1909), and
from mid-Scythian horizons in northern
Siberia (Chjpeoceras ^antmani Popov, 1961:
49). The basic outline and elements of the
suture are quite similar in all these species,
but there is a particularly close similarity
of the suture of this species to the suture
of P. yudishthira Diener from the Heden-
stroemia fauna of Muth, Himalayas (Fig.
lOD).
The present species is from the Cohim-
bites Zone of southeastern Idaho and thus
is younger than the species mentioned
above. This, in fact, is the first recognition
of a species of Pseudaspidites from a zone
younger than that of Meekoceras (or
Owenites).
The fauna of the Dieneroceras Zone of
northern Siberia described by Popov ( 1961 )
I believe to be contemporaneous with that
384 BiiUetin Museum of Comparative Zoology, Vol. 137, No. 3
of the Columhites fauna of southeastern
Idaho. This Siberian fauna contains one
species — Koninckites po.sterhis Popov (1961:
51, pi. 4, fig. 2) — that is very similar to
Pscudaspiditcs popovi. This species clearly
])e]ongs in Fscudaspiditcs and not in
Koninckites. The principal differences be-
tween these species are in details of the
suture (Fig. IOC).
Occurrence. Middle shale member of
Tha\'nes Formation, Columhites Zone, Hot
Springs, southeastern Idaho.
Repositorii. Holotype, MCZ 9575; para-
type MCZ 9636.
Pseudaspidites posterius (Popov)
Text-figure 10
Koninckites posterius Popov, 1961: 51, pi. 4,
fig. 2.
This species is remarkably similar in
conch form to P. popovi described above.
The sutures are likewise quite similar (Fig.
10); the differences center mainly in the
length of the lobes and in the auxiliary
series. Popov compared his species with
Koninckites septentrioncdis Diener (1895:
pi. 1, figs, la-c) from the Primorye Region
and Meekoceras timorensis Wanner (1911:
185, pi. 6, fig. 2, pi. 7, figs. 5, 6) from
Timor. There is a superficial resemblance
between these forms but both these species
are approximately mid-Scythian in age,
whereas P. posterius comes from a younger
horizon contemporaneous with the Co-
lumhites fauna of Idaho.
Occurrence. Dieneroceras Zone of Popov
(1961), delta of the Lena River, Siberia.
Family PROPTYCHITIDAE Waagen, 1895
Genus Propfychifoides Spath, 1930
Type species, Proptychitoides decipiens
Spath, 1930
(= Propfychifes lafifimbriatus Arthaber,
191 1, non de Koninck)
Some ol the more conspicuous upper
Scythian ammonites belong to this genus;
they are likewise often some of the bigg{\st
fonns in a fauna. Arthalx'r (1911) first
recognized the group in the Suhcohimhites
fauna from Albania. At that time he allied
the group to the Proptychites described by
Waagen (1895) from the Salt Range,
noting, ho\\'e\'er, that his Albanian species
differed from the Salt Range species in
the more coarsely denticulated lobes and
the club-shaped, asymmetrical saddles of
the suture. Spath (1930: 30; 1934: 171)
has correctly summarized the major aspects
of these two species groups and introduced
the genus Proptychitoides for the Albanian
forms. Tliis change has been widely ac-
cepted.
Since the first recognition of the group
within the SuhcoJumI)ites fauna of Albania,
it has been recorded from the same horizon
in Kwangsi, China, and Timor. In addi-
tion, it was recorded from a horizon of un-
certain position in the upper Scythian in
northern Siberia. Tlie previous studies of
forms in these several faunas which I be-
lieve to belong in Proptychitoides have re-
sulted in the introduction of 22 species
names included within seven genera. Re-
examination of all the available specimens
from all but the fauna from Kwangsi,
China, leads me to conclude that there are
only five valid species of Pro))tycJiifoides.
The species of Proptychitoides recognized
as valid are:
Pio))tyc]iitnides decipiens Spath
Proplycliifoidcs trifionalis (Arthaber)
PropUjcMtoides arttuil^eri ( Welter )
Propftjcljitnicles tuniilancnsis Chao
Proplycliitoidcs l<nnini<'li ( PopoN' )
It is only in the Suhcohind)itcs launas of
Albania and Chios that two species {de-
cipiens and triiiomdis) occur together. In
all the other faunas mentioned above onh
a single species is present. The Albanian
and (>hi()s species differ in the degree of
iinolution, degrt>e of inilation of the whorls,
and in ornamentation, but their sutiu'es are
essentially identical. Pro])iychitoides de-
cipiens is the more involute, comi^ressed
species, and P. triii,on(dis is the more e\olute
form with a more inf]at(>d whorl. Pro-
))tychitoi(h's (Uth(d)cri from Timor and P.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel
385
tuuiijanensis from Kwangsi, China, have the
same general shell architecture as P.
(Iccipiens; unfortunately, the suture in both
of these species is not as well known as
\\ould be desired. The suture of P. artha-
heri, reproduced on Figure IIL, had, ac-
cording to Welter (1922), less distinctly
rounded saddles. Spath, however (1934:
177), on examination of one topotype speci-
men found this not to be the case. Due
to poor preservation and incomplete data,
it is not possible to make detailed com-
parisons with P. tunglanensis. Both these
species could well be conspecific with P.
dccipicns from Albania and Chios; they are
unfortunately known from so few speci-
mens that a complete analysis of their char-
acters is not yet possible.
There is one other Eurasian species of
Prupfychitoides, P. kummeli (Popov), from
the Olenek River region of northern Siberia.
This species has the general shell architec-
ture of P. trUgonalis, that is, somewhat evo-
lute with an inflated whorl, and a suture
very much like that given by Welter ( 1922 )
for P. arthaheri (Fig. UK, L), and thus, is of
the general jDlan of the Albanian and Chios
species of Propfychitoides.
Proptychifoides decipiens Spath
Plate 8, figures 1-4; Plate 12, figure 3;
Text-figure 1 1
Froptijchites lotifimbriatus, — Arthaber (non de
Koiiinck), 1911: 223, pi. 19(3), figs. 1, 2;
Renz, 1928: 155.
Proptychite.s sp. ind. aff. laiifiwhrkito, — Arthaber
(non de Koninck), — Diener, 1915: 231.
Propfychitoides decipiens Spath, 1930: 39; Spath,
1934: 171, figs. 51a, b; Kummel, in Arkell et
al, 1957: L138, fig. 171, 6.
Proptychites kraffti Arthaber, 1911: 224, pi. 19(3),
fig. 3; Diener, 1915: 231.
Pioptychitoides kraffti, — Spath, 1934: 174, fig.
51c.
Mcekoceras hakki Arthaber, 1911: 247, pi. 22(6),
figs. 1, 2; Diener, 1915: 192.
Proptychifoides hakki,—Spath, 1934: 176, fig.
51e.
Proptychites halcanicus Renz and Renz, 1947: 61,
77; Renz and Renz, 1948: 66, pi. 5, figs. 9-9a.
Proptychites laicrencianus (de Koninck). mut.
postindica Renz and Renz, 1947: 61, 77; Renz
and Renz, 1948: 67, pi. 5, figs. 8-8a.
Table 26. Measurements of Proptychitoides
DECIPIENS Spath from the Subcolumbites
Faunas of Albania and Chios.
D
w
H
U
W/D H/D U/D
1. 178.0 50.0 85.0 37.0 28.1 47.8 20.8
2. 153.0 45.00 75.00 35.0 29.4 49.0 22.9
3. 81.2 ? 42.0 11.1 ? 51.7 13.7
4. 79.0? 24.0 45.8 7.0 .30.4? 57.9? 8.9?
5. 61.4 18.6? .33.1 7.7 .30.3 53.9 12.5
6. .36.4 ? 19.3 5.8 ? .53.0 15.9
1. Svntvpe, Meekoceras hakki Arthaber (1911: pi. 22(6),
fig. la-c), PIUV.
2. Synt>'pe, Meekoceras hakki Arthaber (1911: pi. 22(6),
fig. 2), PIUV.
3. Lectotype, Proptychitoides decipiens Spath [= Propty-
chites latifimbriatus (de Koninck)] Arthaber (1911:
pi. 19 (3), figs. 2a-c), PIUV.
4. Holot\pe, Proptychites kraffti Arthaber (1911: pi. 19
(3), figs. 3a-c), PIUV.
5. Holotype, Proptychites lawrencianus (de Koninck) mut.
postindica Renz and Renz (1948: pi. 5, fig. 8,
NHMB J13725.
6. Proptychites halcanicus Renz and Renz (1948: pi. .3,
fig. 9), NHMB J1372I.
Much of the difficulty in the inteipreta-
tion of the species of the genus Propty-
chitoides from Albania has been due to the
generally poor preservation of the speci-
mens. Features of surface ornamentation,
the suture, and measurements of the stan-
dard conch dimensions are seldom clearly
preserved. In addition, the described
species are represented by one or very few
specimens.
The type specimen of P. decipiens is re-
figured here on Plate 8, figures 3, 4. Tlie
second specimen of this species illustrated
by Arthaber (1911: pi. 19(3), figs, la-c)
is apparently lost. The independent status
of this species and related forms from the
Salt Range Proptychites is most apparent
in the suture with its asymmetrical, phylloid
saddles (Fig. IIG-J).
Proptychites kraffti (Arthaber, 1911: pi.
19(3), figs. 3a-c; Plate 8, figures 1, 2 of
this report) is merely a slightly more in-
volute member of P. decipiens (see Table
26). Arthaber had only three specimens
of this species, of which only one is still
preserved; there is in addition a single
specimen in the British Museum of Natural
386 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Figure 11. Diagrammatic representation of the sutures of: A-F, Proplychitoides tngonatis (Arthaber). A, holotype, at a
diameter of 74 mm (Artfiaber, 1911: pi. 19(3), fig. 4c; PI. 9, figs. 3, 4 of tfiis report); B, fiolotype of Proptychites bertisci
Arthaber (1911: pi. 19(3), figs. 5a-c; PI. 9, figs. 1 , 2 of this report), at a diameter of 68 mm; C, holotype, Meekoceras
mohomed/s Arthaber (1911: pi. 22(6), figs. 3a-c; PI. 10, figs. 1, 2 of this report), at a diameter of 90 mm; D, holotype
of Proptychites buxtorii Renz and Renz (1948: pi. 7, fig. lb), at a diameter of approximately 50 mm; E, holotype of
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 387
History (Spath, 1934: 174). The small of the pattern of the type of P. decipiens.
number of available specimens does not The contemporaneous SuhcoJumhites
permit construction of a meaningful graph, fauna from Chios includes what I consider
On the basis of the available specimens the typical representatives of P. decipiens, which
differences in the width of the umbilicus was at least partially recognized by Renz
between P. decipiens and P. kraffti are 5 and Renz (1948). The papers by Spath
percent or less; from our experience with (1930, 19.34) strangely enough were not
many other species of these Scythian am- known to Renz and Renz (1948). From
monoids this is well within the range of study of their Chios fauna they came to
intraspecific variation seen in most species, the conclusion that the Albanian Proptij-
The slight difference in the suture, espe- chites latifimbriatus (non de Koninck)
cially in the auxiliary series (Fig. IIG-J), Arthaber was not conspecific with the Salt
I likewise do not consider of specific im- Range type of this species, and introduced
portance. the new name Proptij chites halcanicus
Arthaber's two illustrated specimens of Renz and Renz (1948: 66). A substitute
Proptychites hakki (1911: pi. 22(6), figs, name for the Albanian species, P. ffec/p/en5,
1, 2) are very poorly preserved. The holo- had already been introduced by Spath in
type (Arthaber, 1911: pi. 22(6). fig. 1) 1930. A second species in the Chios fauna —
is illustrated here on Plate 12, figure 3. Proptychites lawrencianus (de Koninck)
The specimen is preserved only on one mut. postindica Renz and Renz — was rec-
side and is slightly crushed. The paratype ognized, but this is no more than a poorly
(Arthaber, 1911, pi. 22(6), fig. 2) is so preserved P. decipiens lacking the radial
badly preserved and weathered that only ribs.
a portion of the phragmocone with the Occurrence. Stibcolumhites fauna of Al-
suture exposed offers data of value. The bania and Chios.
most apparent difference between P. hakki Repository. Tlie following specimens are
and P. decipiens is in the relative diameter -^^ ^^le Paleontological Institute, Vienna:
of the umbilicus. In this regard, hoxyever, i^^i^^y^^^ Proptychitoides decipiens Spath
the two specimens of P. hakki are larger Proptychites htifimbriatus,-Arth:xher,
by a factor of at least two than all the other ^^^^_ 2a-c [non de Ko-
specimens from the Subcolumbites fauna . , ,,^ -r, j ., 7 te^- a .1
£ All 1 r-i • 1.1 4. 1 ^ r.-r.^^A nmckl); syntype, Proptychites kraffti ArthsL-
or Albania and Chios that are here assigned ■'^' ' -"^ ' \ r o /di
to P. decipiens. From the observations that ^er, 1911: pi. 19(3), tigs. 3a-c (PI. 4,
can be made on these specimens, it appears f^gs- 1, 2 of this report); holotype Meeko-
that the relative size of the umbilicus in- ceras hakki Arthaber, 1911: pi. 22(6), fig.
creases as the conch gets larger. The other la-c (PI. 18, fig. 3 of this report); paratype,
features of the conch— whorl shape, degree Meekoceras hakki Arthaber, 1911: pi. 22(6),
of compression, and suture — are distinctly fig. 2. The following specimens are in the
Flemingites pseudorussei/i Renz and Renz (1948: pi. 7, fig. 3c), at a diameter of approximately 50 mm; F, type speci-
men of Propfych/tes mohamedis var. applanata Renz and Renz (1948: pi. 6, fig. 3b), at a diameter of approximately 40
mm; G-J, Propfych/fo/des dec/p/ens Spatti; G, syntype of Meekoceras hakki Artfiober (1911: pi. 22(6), fig. 2], at a
diameter of 99 mm; H, syntype of Meekoceras hakki Artfiaber (1911: pi. 22(6), fig. Ic), at a diameter of 60 mm; I,
holotype [^Proptychites latiiimbriatus non de Koninck, Arthaber (1911: pi. 19(3), fig. 2c), at a diameter of 73 mm,
J, syntype Proptychites kraffti Arthaber (1911: pi. 19(3), fig. 3c), at a diameter of 65 mm; K, holotype Procarnites kummeli
Popov (1962: 188, fig. 10), at a diameter of 100 mm; L, holotype of Proptychites orthoben Welter (1922: pi. 2, figs.
1-3), at a diameter of 45 mm.
Specimens of figures A-C, G-J from Subco/umbites fauna of Albania, specimens of figures D-F from same fauna on Chios,
specimen of figure K from Olenek River region, Siberia, specimen of figure L from Block E, Nifoekoko, Timor.
388
Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Natural History Museum, Basel: plesio-
tvpe, Propti/cliites halcanicus Renz and
Renz (1948: pi. 5, fig. 9) NHMB J13721;
unfigured paratypes NHMB 13722; holo-
type, Froptijchites Jaivrcncianus ( de Kon-
inck) mut. postindica Renz and Renz
(1948: pi. 5, fig. 8) NHMB J13725.
Proptychltoides trigonalis (Arthaber)
Plate 9, figures 1-4; Plate 10, figures
1-4; Plate 11, figure 5; Text-figure
11
Propiijchites trigonalis Arthaber, 1911: 225, pi.
19('3), fig. 4; Diener, 1915: 232.
ProptycJiitoidcs trigonalis, — Spath, 1934: 174.
Profiti/cJtites hertisci Arthaber, 1911: 225, pi. 19(3),
fig. 5; Diener, 1915: 231.
ProptycJiitoides hertisci, — Spath, 1934: 174.
Proptiichites ohU(iin'plicatiis, — Arthaber (non Waa-
geii), 1911: 226, pi. 20(4), fig. 1.
ProptycJiitoides (?) nopcsai Spath, 1934: 175
( = Proptychitcs obliqueplicatus, — Athaber, non
Waagen ) .
Meekoceras inohanicdis Arthal)er, 1911: 248, pi.
22(6), fig. 3.
Proptychitcs mahomedis, — Diener, 1915: 232.
Proptyciiitoides mahomedis, — Spath, 1934: 175,
fig."51d.
Proptychitcs mahomedis var. applanata Renz and
Renz, 1947: 61, 76; Ren/, and Renz, 194.S:
64, pi. 6, figs. 3-3b.
Proi)tycJiitcs ktcnasi Renz and Renz, 1947: 61,
76;" Renz and Renz, 1948: 65, pi. 6, figs. 1-lb.
Proptychitcs artJiahcri, — Renz and Renz (non
Welter) 1947: 61; Renz and Renz, 1948: 65,
pi. 7, figs. 4-4b; pi. 5, figs. 7-7b.
Proptychitcs mistardisi Renz and Renz, U)47: 61,
77; Renz and Renz, 1948: 67, pi. 5, figs.
lO-lOb.
Proptycliitcs httxtorfi Henz and Renz, 1947: 61,
77; Renz and Renz, 1948: 68, pi. 7, figs. 1-lb.
Koiiinckites J)ernoitUii Renz and Renz, 1947: 61,
76; Renz and Renz 1948: 58, pi. 5, figs. 5-5a,
pi. 6, figs. 2-2a, pi. 7, figs. 2-2a (var.).
Flemingites pscudorusselli Renz and Renz, 1947:
60, 76: Renz and Renz, 1948: 54, pi. 7, figs.
3-3e.
Moiiophyllitcs {LeiopJujIlilcs} dicneri Arthaber
var. irwohita, — Renz and Renz, 1947: 61; Renz
and Renz, 1948: 75, pi. 5, figs. 1-lb.
Mono))hylIites {?Sc}iizoi)1iyllitcs) psendohara Henz
and lienz, 1947: 61, 78; Henz and I^miz, 1948:
78, pi. 5, figs. 3-3b.
In contrast to the compressed, more or
less in\'olute forms of Propfijcltifoides (e.g.,
P. decipicns) in the Suhcolumhitcs fauna
of Albania and Chios, there is a more in-
flated form, with a trigonal whorl section,
and generally some form of radial ribs.
Arthaber (1911) recognized four species
within this "inflated" group, and Renz and
Renz ( 1948 ) recognized six species and one
new variety. The main differences between
all these "species" are in the degree of
inflation of the whorls and degree of lateral
ornamentation. There are thus 10 species
plus one variety recognized for this group.
Of these, seven species and one variety
were established on the basis of a single
specimen each, one species was recognized
from two specimens, one species on the
basis of three specimens, and one species
on the basis of five specimens; in summary
18 specimens of this inflated fonn of Pro-
ptijchitoides gave rise to 10 species and one
variety. Thirteen of these specimens are
still preserved and were re-examined for
this study. The measurements of these
specimens are tabulated in Table 27.
The holotype (Plate 9, figures 3, 4) is a
poorly preserved phragmocone showing the
trace of the umbilical seam for an additional
three-quarter volution. The umbilicus is
broad and open with steep umbilical walls.
The whorl section is trigonal in outline
with a narrowly roimded venter. The flanks
are slightly crushed but, in spite of this,
on the adoral half xolution the flanks are
slightly concave. This specimen appears
to be the inner whorls of what was originall\'
a very large specimen like tlie holotype
of Proptijchiics ktcnasi Renz and Renz
( 1948: pi. 6, fig. 1). Due to poor preserva-
tion no siuface ornamentation is preserved
on the type specimen of P. triiiomdis.
Proptijchitoides bciiisci Arthaber (1911:
pi. 19(3), fig. 5; PI. 9, figs. 1, 2 of this
report) represents one direction of varia-
tion in its \er\ broad, depressed whorl sec-
tion. Most of the other forms included
in Proptychltoides tri^otudis arc \'ariants
towards more compressed whorl sections
and (he deNcloiiincnt ol radial ribs. One
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel
389
Table 27. Measurements of Proptychitoides
TRIGONAUS (ArTHABEr) FROM THE SUBCOLVM-
BiTES Faunas of Albania and Chios.
No.
D
w
H
u
W/D
H/D
U/D
1.
123.0
45.7?
60.2
24.2
.37.2?
48.9
19.7
2.
90.5
30.0
38.8
28.4
33.8
42.9
31.4
3
77.8
?
37.3
21.4
?
47.9
27.5
4.
77.0
35.0
.38.5
16.2
45.5
50.0
21.0
5.
76.2
28.5?
35.0
19.1
37.4
45.9
25.1
6.
73.8
38.0?
31.7
21.7
51.5?
42.9
29.4
7.
68.6
?
32.1
16.0
?
46.8
23.3
8.
65.8
?
28.1
19.0
?
42.7
28.9
9.
65.6
17.8
26.7
19.8
27.1
40.7
.30.2
10.
61.4
21.7
28.7
14.6
.35.3
46.7
23.8
11.
60.6
21.0
26.3
16.0
.34.7
43.4
26.4
12.
31.8
10.4
14.8
9.3
.32.7
46.5
29.2
13.
30.3
12.6?
13.5
8.4
41.6?
44.6
27.7
1. Holotvpe, Proptychites ktenasi Renz and Renz (1948:
pi. 6,' fig. 1), NHMB J 13715.
2. Lectotvpe (herein designated), Meekoceras mahomc-
dis Arthaber (1911: pi. 22(6), fig. .3), PIUV.
3. HoloUpe, Koninckites bcrnotiUii Renz and Renz
(1948: pi. 6, fig. 2), NHMB J 13707.
4. HoloUpe, Proptychites trigonalis Arthaber (1911: pi.
19(3)^ fig. 4a-c), PIUV.
.5. Holot\pe, Proptychites buxtorfi Renz and Renz (1948:
pi. 7,' fig. 1), NHMB J13726.
6. Holotspe, Proptychites hertisci Arthaber (1911: pi.
19(3)', fig. 5), PIUV.
7. Paratype, Koninckites bernouUii Renz and Renz
(1948: pi. 7, fig. 2), NHMB J13709.
8. Paratype, Koninckites bernouUii Renz and Renz
(1948: pi. 5, fig. 5), NHMB J13708.
9. Holotvpe, Flemingites psetidonisscUi Renz and Renz
(1948: pi. 7, fig. 3), NHMB J 1.3698.
10. Plesiotvpe, Proptychites arthaberi, — Renz and Renz
(non Welter) (1948: pi. 7, fig. 4), NHMB J13717.
11. Holotvpe, Proptychites mohamedis Arthaber var. ap-
phmaia Renz and Renz (1948: pi. 6, fig. 3), NHMB
J13714.
12. Plesiotvpe, Proptychites arthaberi, — Renz and Renz
(non Welter) (1948: pi. 5, fig. 7), NHMB J13718.
13. Holotvpe, Proptychites mistardisi Renz and Renz
(1948: pi. .5, fig. 10), NHMB J13723.
of the more extreme forms is Meekoceras
mahomcdis (Plate 10, figs. 1, 2). Many of
the specimens from the Suhcolumhites
fauna of Chios described as species of
Proptijchites, Koninckites, and Flemingites
as hsted in the synonymy are intermediate
in degree of whorl inflation and ornamenta-
tion between P. trigonalis and P. mahomedis.
The number of available specimens does
not allow a statistical analysis of this varia-
tion in conch fonn. Unless one is willing
to take note of the large amount of varia-
tion potentially possible within many am-
monite species where there is an abundance
of data, one is left with a name per speci-
men, as has happened \\'ithin this group.
The pattern of the suture (Fig. IIA-F)
and degree of evolution in all these speci-
mens ties them together; on this back-
ground the variation in whorl width and
ornamentation is much more understand-
able.
The associated P. decipiens differs in its
greater involution and pattern of ornamen-
tation. The basic pattern of the suture in
these two species remains the same (Fig.
IIG-J). The suture and general conch
features in the other species of Propty-
chitoides easily distinguish them from P.
trigonalis.
Occurrence. Suhcohimbites fauna of Al-
bania and Chios.
Repository. The following specimens are
in the Paleontological Institute, University
of Vienna: holotype, P. trigonalis Arthaber,
1911: pL 19(3), fig. 4 (PI. 5, figs. 3, 4
of this report); holotype P. hertisci Artha-
ber, 1911: pi. 19(3), fig. 5 (Pk 5, figs. 1, 2
of this report); holotype, P. (?) nopcsai
Spath, 1934: 175 {^Proptychites oh-
UquepUcatus,—AYthAher, 1911: pi. 20(4),
fig. 1, non Waagen (Pi. 6, figs. 3, 4 of this
report); holotvpe, Meekoceras mahomcdis
Arthaber, 1911: pi. 22(6), fig. 3 (PI. 6,
figs. 1, 2 of this report). The following
specimens are in the Natural History
Museum, Basel: holotype, Proptychites
mohamedis var. applanata Renz and Renz
(1948: pi. 6, fig. 3) NHMB J13714; un-
figured paratypes NHMB J 13835; holotype,
Proptychites ktenasi Renz and Renz (1948,
pi. 6, fig. 1) NHMB J13715; unfigured
paratype NHMB J 13716; plesiotypes, Pro-
ptychites arthaberi^ — Renz and Renz (1948:
pi. 5, fig. 7) NHMB J13718, (pk 7, fig. 4)
NHMB J13717; unfigured paratypes from
Maradovuno NHMB J 137 19, from Kepha-
lovuno NHMB J13720; holotype, Propty-
chites mistardisi Renz and Renz (1948: pk
5, fig. 10) NHMB J13723; unfigured para-
types NHMB J 13724; holotype Proptychites
huxtoiii Renz and Renz (1948: pk 7, fig.
1) NHMB J13726; holotype, Koninckites
bernouUii Renz and Renz (1948: pk 6,
390
BuUetin Museum of Comparative Zoology, Vol. 137, No. 3
fig. 2) NHMB J13707; paratypes (pi. 5,
fig. 5) NHMB J13708, (pi. 7, fig. 2) NHMB
J 13709; holotype Flcmin^j^ites pseiidortisselli
Renz and Renz ( 1948: pi. 7, fig. 3) NHMB
J13698; unfigured paratype NHMB J13699;
figured specimen MonopliijIIites (Leiopliyl-
lites) cliencri var. invohita Renz and Renz
(1948: pi. 5, fig. 1) NHMB J13747; un-
figured paratype NHMB J13748; holotype,
MonophijUites (ScJiizopltt/Uite.s) pscudohara
Renz and Renz ( 1948: pi. 5, fig. 3) NHMB
J 13763.
Propfychitoides arfhaberi (Welter)
Plate 25, figures 1, 2; Text-figure 11
Proptychites arthaberi Welter, 1922: 102, pis.
156(27), figs. 1-4; Kutassy, 1933: 625.
Proptijcliitoidcs arthaheri, — Spath, 1934: 177;
Kummel, 1961: 525.
This species is of the general conch form
of Vroptijchitoides tri^onalis. Welter's type
and only specimen is well preserved but
incomplete; the illustration of the complete
specimen (Welter, 1922: pi. 156(2), fig.
4) is slightly inaccurate in that the um-
bilicus is shown too small. The umbilicus
is approximately 21 percent the diameter of
the conch rather than 18 percent as indi-
cated by Welter's figure. An additional
specimen that is of considerable interest,
illustrated here on Plate 25, figures 1, 2, is
available in the collections of the Geological
Institute, Amsterdam. First, it has a suture
nearly identical to that of the type speci-
men illustrated by Welter (Fig. IIL). The
whorl section, however, is more inflated
and more trigonal in cross-section. Like-
wise, the whorl sides bear more prominent
radial folds. However, the character of
the venter, umbilical shoulder and wall,
and degree of involution are the same in
the two specimens.
Occurrence. Welter's type specimen
came from Block E, Nifoekoko, Timor, with
the manganese coated fossils. The speci-
men in the Amsterdam collections has no
label but is a manganese coated specimen
like those from Block E at Nifoekoko.
Repo.siforij. Holotype in the Paleontologi-
cal Institute, Bonn; two topotypes are in
the British Museum (Natural History)
C33748-9; the specimen figured here is in
the Geological Institute, University of Am-
sterdam.
Propfychitoides tunglanensis Chao
Pioptijchitoides tunglanensis Chao, 1959: 80, 245,
pi. 20, figs. 11-12, text-fig. 25a.
Proptychitoidcs compressus Chao, 1959: 80, 246,
pi. 20, figs. 9-10, text-fig. 25b.
Pwptijchitokles ? simplex Chao, 1959: 81, 246, pi.
20, figs. 7-8, text-fig. 25c.
The three species brought together here
were each based on single, poorly preserved
specimens from three different localities
of the Subcohimbites fauna in Kwangsi,
China. They likewise represent three dif-
ferent growth stages. Chao (1959) made
no reference to the differences between his
three species. On the basis of the very
poor illustrations, the brief and incom-
plete descriptions, and taking into account
the poor preservation, there appears to be
little real basis for separating these three
forms.
This Kwangsi species is of the general
morphological type of P. dccipiens from
Albania and Chios. The available speci-
mens and their poor preservation do not
allow more detailed comparison. This
species could l)e conspecific with the Al-
banian and Chios P. decipiens or possibly
P. arthaheri from Timor. For the moment
it is thought best to retain an independent
status for these Kwangsi specimens.
Occurrence. The holotype of P. tung,-
lanensis came from the Subcohtnibites
fauna on the western side of Chashanao
between Tunglan and Hochich districts
(Chao collection 610); the holotype of P.
compres.s-us came from a limestone block
in the Lolou xillage in the Linglo district,
associated with Proearnites kukeni (Chao
collection 542b); the holotype of P. .simplex
came from the Su]}eohimhite.s fauna 1.5
km north of Yali in the Fengshan district
(Chao colU^ction 546), all in K\\'angsi,
China.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 391
Propfychitoides kummeli (Popov)
Text-figure 1 1
Procarnites kuiunicli Popov, 1962a: 187, pi. 2,
fifi. .5.
This species has the general conch
arcliitecture of P. artJmhcri and P. triiS,onalis.
Its suture is very much like that of the
Timor P. cirtJmberi (Fig. IIK) and not Hke
that of any known species of Procarnites.
Occurrence. From along the Nikabit
River in the Olenek River region, Siberia.
Popov (1961: 177, 188) Hsts the specimens,
with questions, as having come from his
Olcnekitcs Zone.
Genus Procarnites Arthaber, 1911
Type species, Porapopanoceras kokeni
Arthaber, 1908
Procarnites kokeni (Arthaber)
Plate 11, figures 1-4; Plate 12, figures
1, 2; Plate 13, figures 1-8; Text-
figures 12, 13
Faiapopauuccras kokeni Arthaber, 1908: 259, pi.
11(1), figs, la-c, 2a, b.
llcdcnstroemia sp. Arthaber, 1908: 284, pi. 3,
fig. 2.
Procarnites kokeni (Arthaber) 1911: 215, pi.
17(1), figs. 16, 17, pi. 18(2), figs. 1-5; Diener,
1915: 228; Diener, 1917: 167; C. Renz, 1928:
155; Renz and Renz, 1947: 61; Renz and Renz,
1948: 81, pi. 8, figs. 5, 6-6a, 7-7a, 8-8a, 9-9a,
pi. 9, figs. 2-2a; Kummel, 1966: 390, pi. 2,
figs. 10-13; Kummel, 1968b: 493, pi. 1, fig. 16.
Procarnites kokeni var. evoluta Renz and Renz,
1947: 61; Renz and Renz. 1948: 82, pi. 9,
figs. 1-la.
Procarnites kokeni var. panteleimonensis Renz and
Renz, 1947: 61, 78; Renz and Renz, 1948: 82,
pi. 8, figs. 3-3a, pi. 9, figs. 3-3a.
Procarnites acutiis Spatli, 1934: 183, pi. 5, figs.
4a, b {^ Hedenstroemia sp. Arthaber, 1908:
284, pi. 3, fig. 2): Chao, 1959: 89, 255, pi.
32, figs. 8, 9, pi. 33, figs. 1-8.
Procarnites skanderJ^egis Arthaber, 1911: 216, pi.
18(2), figs. 6, 7; Diener, 1915: 229; C. Renz,
1928: 155; Renz and Renz, 1947: 61; Renz and
Renz, 1948: 82, pi. 8, figs. 4-4a.
Procarnites andrusovi ( Bajarunas, 1936: nomen
nudum) Kiparisova, 1947: 1.32, pi. 28, figs. 2-4,
text-figs. 11-13; Astakhova, 1960b: 149.
Procarnites oxi/nosiiis Chao, 1959: 88, 254, pi.
32, figs. 1-7, 10-12, text-fig. 28a-d.
Species of Procarnites have been differ-
entiated on the basis of the suture, the
degree of inflation of the conch, the nature
of the venter, and to some extent on the
ornamentation. Much of the misconcep-
tion that has existed about this species has
been due directly to the relatively poor
preservation and to the manner of treat-
ment of this material.
The types of Procarnites kokeni (Artha-
ber) are two small, immature specimens,
refigured here on Plate 13, figures 1^.
Only one side of either of these specimens
is preserved. Noteworthy of the smaller
of these two specimens are the broadly
arched lateral areas and the round umbili-
cal shoulders. The larger specimen has
broader lateral areas and abruptly rounded
umbilical shoulders and nearly vertical
umbilical walls.
In his monograph on the Kcira, Albania,
fauna Arthaber (1911) stated he had 45
specimens for study, and he illustrated six
of these. Unfortunately, only three of the
illustrated specimens are still available,
and these are refigured here. Arthaber did
not give measurements of any of his speci-
mens.
In addition to Procarnites kokeni, Artha-
ber (1911) recognized one additional
species, P. skanderheg,is. For this species
he records eight specimens of which the
two illustrated types are available for study.
This species was differentiated on the basis
of suture, degree of conch inflation, and
ornamentation. In regards to the suture it
was the absence of minor adventitious ele-
ments in the ventral lobe to which Arthaber
pointed. The modification and develop-
ment of the suture in the ventral region
progressively change with growth, and com-
paring the suture of P. skanderhegis with
that of P. kokeni at a comparable growth
stage shows that they differ in only the
smallest details (Fig. 12A-C, E, F).
Assessment of the significance of conch
shape and ornamentation was not possible
until the discovery of the Siibcolumbites
fauna of Chios which yielded a large num-
ber of specimens of Procarnites. On Table
392 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Figure 12. Diagrammatic representation of the sutures of Procarnites kokeni (Arthaber), Procarn/tes immaturus (Kipari-
sova), and Procarnites /o/ouensis (Chao). A-G, Procorni/es kokeni, A, lectotype (Arthaber, 1908: pi. 11(1), fig. Ic), at
a diameter of 33 mm; B, plesiotype (Arthaber, 1911: pi. 18(2), fig. 3), at a diameter of 34 mm; C, plesiotype (Arthaber,
1911: pi. 18(2), fig. 2c), at a diameter of about 21 mm; D, holotype of Procarnites oxynosfus Chao (1959: fig. 28a), at
a diameter of approximately 80 mm; E, paralectotype of Procarnites skanderbegis Arthaber (1911: pi. 18(2), fig. 6c], a\
a diameter of 50 mm; F, lectotype of Procarnffes skanderbegis Arthaber (1911: pi. 18(2), fig. 7c), at a diameter of 55 mm;
G, syntype of Procarnites andrusovi Kiparisova (1947: 132, fig. 12), at a diameter of approximately 50 mm; H, holotype of
Megaphyilites immaturus Kiparisova (1947: 130, fig. 8), at a diameter of approximately 40 mm; I, J, Digitophyllites
/o/ouensis Chao (1959, figs. 29a, b), both sutures from whorl height of approximately 10 mm.
Specimens of Figures A-C, E, F from Subco/umbites fauna of Albania, of figure G, from upper Scythian of the Man-
gyshlak Peninsula, D, I, J, from Subco/umb/tes Zone of Kwangsi, China, and hi, from Subco/umbites fauna of the Pri-
morye Region.
Figure 13. Variation in the width (W) and height (H) of whorls, and umbilical diameter (U) of Procarn/tes kokeni (Arthaber).
Specimens from Albania and Chios are marked with a dot, those of Procarnites skanderbegis with a cross, specimen 69
from the /v\angyshlak Peninsula with a triangle, and specimens 64-68 and 70-74 from Kwangsi, China, with an X. The
data on this graph are from Table 28.
65
60
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 393
55
H
50
45
40
35
30
25
20
15
10
1
1
J L
0 10 20 30 40 50 60 70 80 90 100 110 120 130
DIAMETER
394 Biilleiin Museum of Comparative Zoology, Vol. 137, No. 3
Table 28. Measurements of Procarnites kokeni (Arthaber) from the Subcolumbites Faunas
OF Albania and Chios. Specimens 1-63: Those Identified by Arthaber and by Renz and Renz
AS P. KOKENI Marked with a Dot, Those as P. skanderbegis with a Cross. Specimens 64-74
Marked by an X; Specimen 69 is from Southern U.S.S.R., Specimens 64-68, 70-74 are from
Kwangsi, China.
D
W
H
u
W/D
H/D
U/D
D
w
H
u
W/D
H/D
U/D
1.
114.2
21.0?
61.5
11.6
18.4?
53.9
10.2
.38.
31.9
9.3
17.0
4.4
29.2
53.3
13.8
2.
90.5?
31.5?
49.5?
8.2
34.8?
54.7?
9.1?
39.
31.2
10.1
13.9
6.3
32.4
44.6
20.2
3.
80.0
19.7
41.6?
9.0?
24.6
.52.0?
11.3?
40.
30.7
8.6
1.5.6
4.4
28.0
50.8
14.3
4.
72.2
13.1
40.6
9.2
18.1
56.2
12.7
41.
30.3
12.6
16.7
3.8
41.6
55.1
12.5
5.
67.7
13.9
.37.2
6.8
20.5
54.9
10.0
42.
29.9
7.7
1.5.2
4.7
25.8
50.8
15.7
6.
64.0
16.2
.34.5
5.6
25.3
53.9
8.8
43.
29.1
9.3
15.0
3.3
31.9
51.5
11.3
7.
62.0
15.7?
32.8
6.8
25.3?
52.9
10.9
44.
28.3
9.3
14.0
5.0
.32.9
49.5
17.7
8.
61.1
16.5
31.7
7.1
27.0
51.9
11.6
45.
28.0
8.0
14.7
4.0
28.6
52.5
14.3
9.
54.5
15.2
28.8
6.7
27.9
52.8
12.3
46.
27.8
8.0
14.6
3.5
28.8
52.5
12.6
10.
54.5
14.1
28.9
8.0
25.9
53.0
14.7
47.
27.3
9.3
14.4
3.0
.34.1
52.7
10.9
11.
53.0?
17.0?
29.5
6.9
32.1?
55.7?
13.0?
48.
26.8
8.6
14.8
4.3
32.1
55.2
16.0
12.
51.8
10.7
29.0
4.7
20.7
56.0
9.1
49.
26.2
8.9
13.8
3.0
33.9
52.7
11.5
13.
50.8
12.1
25.6
7.3
23.8
50.4
14.4
50.
25.8
8.8
12.1
5.6
34.1
46.9
21.7
14.
48.7
15.0?
25.8
6.3
30.8?
52.9
12.9
51.
25.5
9.7
12.3
4.0?
.38.0
48.2
15.7?
15.
48.4
15.6
27.2
4.8
32.2
56.2
9.9
52.
25.3
8.5
11.7
5.9
33.6
46.2
23.3
16.
48.4
11.9
27.2
4.4
24.6
56.2
9.1
53.
24.5
6.4
12.7
2.8
26.1
51.8
11.4
17.
47.3
12.1
24.5
7.0?
25.6
51.8
14.8?
54.
23.6
8.0
11.0
4.7
33.9
46.6
19.9
18.
47.2
9.2
25.2
6.0
19.5
53.4
12.7
55.
21.6
6.6
10.5
4.1
.30.6
48.6
18.9
19.
46.5
13.7?
24.0
5.4
29.5?
51.6
11.6
56.
20.8?
p
10.4
4.2
p
50.0
20.2
20.
46.4
12.0
22.8
7.0
25.9
49.1
15.1
57.
19.3
5.8
9.5
3.5
.30.1
49.2
18.1
21.
46.2
?
21.7
9.0
?
46.9
19.5
58.
18.6
6.4
9.8
2.7
.34.4
52.7
14.5
22.
46.1
13.8
23.1
6.8
29.9
50.1
14.8
59.
18.2
6.0
13.8
3.7
.32.9
75.8
20.3
23.
45.4
9.9
26.0
4.7
21.8
57.3
10.4
60.
16.8
5.2?
8.5
3.6
30.9
50.6
21.4
24.
44.5
13.7
23.8
5.0
.30.8
53.5
11.2
61.
16.1
6.4
7.2
4.3
39.8
44.7
26.7
25.
37.7
?
20.1
3.8
?
53.3
10.1
62.
14.0
5.5
6.3
3.5
39.3
45.0
25.0
26.
37.2
11.6
18.8
5.7?
31.2
50.5
15.3?
63.
12.0
3.9
5.7
2.1
32.5
47.5
17.5
27.
.36.8
11.0
18.6
5.5
29.8
50.5
14.9
64.
105.0
16.7
59.2
9.4
16.0
56.0
8.9
28.
36.5
11.0
18.5
5.6
30.1
50.7
15.3
65.
100.0?
17.0
56.0?
8.2
17.0
56.0
8.2
29.
35.7
7.7
19.6
5.1
21.6
54.9
14.3
66.
80.0
16.8
41.5
7.3
20.0
52.0
9.0
30.
35.6
9.7
19.0?
5.7?
27.2
53.4?
16.0?
67.
69.2
16.2
33.6
8.2
16.6
48.0
12.0
31.
35.4
9.0
19.0
2.8
25.4
53.7
7.9
68.
68.0
15.1
,36.2
7.5
22.2
.52.0
10.0
32.
33.0
?
16.8
5.2
?
.50.9
15.8
69.
56.0
14.0
27.4
9.0
33.
32.8
12.5
17.0
4.1
38.1
51.8
12.5
70.
46.6
9.7
23.5
6.8
20.0
50.0
14.6
34.
32.8
9.2
15.6
5.3
28.0
47.6
16.2
71.
27.4
8.6
13.0
4.5
21.2
47.8
16.0
4.
5.
6.
7.
10,
11.
1,3
14
l.^.
19.
20,
21.
24,
2.5.
.32.
34.
.39.
Plesiotype (Arthaber, 1911: pi. 18(2), figs. .5a, h), PIUV.
Svnt\'pe, P. skanderbegis Arthaber (1911: pi. 18(2), figs.
8, 9, 12, 16, 18, 23, 27, 29-31, 35, 45, 46, 49, 53,
dovuno, Chios, NHMB J 13774- 13779.
Plesiotype, — Renz and Renz (1948:
Plesiotype, — Renz and Renz (1948:
Plesiotype, — Renz and Renz (1948:
Plesiotype, — Renz and Renz (1948:
17, 22, 37. Unfigured paratypes,
7a, b), PIUV.
58, 59, 62, 63.
Unfigured specimens, P. kok(ui Mara-
pl. S, fig.
pl. 8, fig.
pl. 8, fig.
pl. 8, fig.
P. kokeni
Chios, MIMH J13786.
6), NHMB II 3769.
7), NHMB 113772.
9), NHMB j 13773.
8), NHMB J 13770.
var. ixinteleinioiiensifi, .MaradoN uno,
Syntype, P. .skanderbegit Arthaber (1911: pl. 18(2), figs. 6a, b), PIUV.
Plesiotype, — Renz and Ren/, (1948: pl. 9, fig. 2), NHMB J13771.
Plesiotype (Arthaber, 1911: pl. 17(1), figs. 17a, b), PIUV.
Plesiotype, P. sktiuderliegis, Renz and Renz (1948; pl. 8, fig. 4),
Plesiotype,— Ren/, and Renz (1948: pi. 8, fig. 5), NHMB .113768.
28, 38, 40, 41, 42, 48, 50, 55, 57, 60. Unfigured paratvpes, /
NHMB Jl,3782.
Syntype, P. kokeni var. paiiteleinionensi.s Renz and Renz ( 1947: 61,
26, 33, 36, 43, 44, 47, 51, 52. 54, 61. Unfigured speeimens, P. skanderbegis Maradovuno, Chios, NHMB J13789.
Plesiotype (Arthaber, 1911: pi. 17(1), figs. 16a, b), PIUV.
Syntype (Arthaber, 1908: pl. 11(1), figs, la-c), PIUV.
Syntype, P. kokeni var. panteleimonensis Renz and Ben/ ( 1917: 61, 78; 1948: pl. 8, fig. 3), NHMB J1,37S4.
Holotype, P. kokeni var. evohilii Ren/ and Hen/ ( 1947; 61; 1948; pl. 9, fig. 1), NHMB JI,3781.
NHMB 1I37SS.
kokeni \ ar. eiohita,
8; 1948; pi. 9, fig. 3),
M;irado\'Mn(). Chios,
NHMB .|I;57.S5.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 395
Table 28. Continued
D W H U W/D H/D U/D D W H U W/D H/D U/D
35. 32.7 7.7 16.7 5.5 23.5 51.1 16.8 72. 22.6 5.5 12.5 3.2 23.8 53.4 13.2
36. 32.5 11.2 16.0 4.7 34.5 49.2 14.5 73. 21.0 7.5 10.7 2.7 34.0 51.0 13.0
37. 32.2 8.9 16.1 4.8 27.6 50.0 14.9 74. 17.2 5.0 10.4 2.4 27.8 60.0 14.0
56. Syntvpe ( Arthaber, 1908: pi. 11(1), figs. 2a, b), PIUV.
64, 65, 70, 72. Procamites actitus, — Chao (1959: 255-256).
66. Holotvpe, P. oxtjnostiis Chao (1959: 255).
67, 68, 71, 73, 74. Parat>pes, P. oxynostus Chao (1959: 255).
69. Procamites aiulnisovi Kiparisova (1947: 132).
28 are the measurements of 44 specimens tion. Thus the criteria used to distinguish
assigned by Renz and Renz (1948) to P. ska nderbegis horn P. kokeni do not stand
Procamites kokcni and 12 specimens these up on close examination,
authors assigned to P. .s/crtnf/c;-Z?(?gi5. These Spath (1934: 183) separated one of
data are plotted on Figure 13. It can readily Arthaber's varieties of Procamites kokeni
be seen that no distinction can be made as a new species — P. acutits. The distinc-
between these two species on whorl height tion was made on the basis of an acute to
or umbilical diameter. In respect to whorl oxynote venter. He (Spath) also mentions
width the specimens assigned to P. skan- a form which he considered transitional
derbegis tend to be thicker, but there is with P. kokeni from the same Albanian
complete gradation \\'ith more compressed Siibcolumhites fauna. Examination of the
forms which had been placed in P. kokeni. large number of specimens of P. kokeni from
The plot of Arthaber's two figured types Chios clearly shows complete gradation
on Figure 13 likewise shows that these are from forms with acute venters like Spath's
no more than slightly inflated forms which holotype of P. actitus (Spath, 1934: pi. 5,
can much better be considered as part of figs. 4a, b ) to forms with broadly rounded
P. kokeni. venters. Arthaber insisted on minor aspects
The ornamentation of the larger of of the suture as important distinguishing
Arthaber's two figured types (PI. 11, figs, features. Spath (1934: 183) was not able
1, 2) consists of very faint radial folds and to see these differences clearly and cor-
slightly accentuated growth lines every rectly brought attention to the frequent
millimeter or so. The specimen is not well loss of detail entailed in the grinding neces-
preserved so the complete pattern of this sary to bring out the sutures on these Al-
ornamentation is not known. None of the banian specimens.
specimens from Chios assigned by Renz Procamites andriisovi Bajavunas (Kipari-
and Renz to P. skanderbegis show any sova, 1947) from the Mangyshlak Peninsula
ornamentation. Two specimens of P. kokeni of the Caspian region is clearly conspecific
(Renz and Renz, 1948: pi. 8, figs. 3, 5 and with P. kokeni. Kiparisova distinguished
pi. 9, fig. 3 ) show faint radial ribs or falcoid her species on the basis of the greater width
ribs. The poor state of preservation of the of the umbilicus and the addition of an
Albanian and Chios Subcolumbites fauna adventive element in the ventral lobe. The
is an important factor which does not allow umbilical width of P. andriisovi falls well
full evaluation of the nature of and varia- within the variability in this feature in the
tion of the ornamentation patterns. On the population of P. kokeni from Chios (Fig.
basis of the data available, ornamentation 12G). It has already been pointed out
does not appear to be a criterion which can that variations in the ventral lobe are a
be used in this case for species discrimina- function of ontogeny.
396 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Unfortunately, the stratigraphic relations
of the Mangyshlak Scythian ammonoids de-
scribed to date are not well known. The
data given by Bajarunas ( 1936 ) and Kipari-
sova (1947) are ambiguous. Recent publi-
cations on the Triassic strata of Mangyshlak
did not discuss P. andrusovi but added some
additional stratigraphic data (Astakhova,
1960a, b).
Chao ( 1959 ) recognized two species of
Procarnitcs from Kwangsi, China. He had
four specimens he assigned to P. aciitiis
Spath and six specimens for which he
erected a new species, P. oxynosttis. The
measurements of these forms are plotted
on the graph of Figure 13. It can readily
be seen that in terms of whorl heights and
umbilical diameter these specimens are
quite like P. kokenl from Chios. In terms
of whorl width they are clearly of the com-
pressed variety but yet within the range
of variation in this feature in the material
of P. kukeni from Chios. The minor dif-
ferences in the suture pointed to by Chao
are more likely expressions of poor preserva-
tion than any true genetic significance.
These Kwangsi specimens are considered
to l)c valid representatives of Procarnitcs
kokcni.
This species was reported from Timor by
Spath (1934, p. 182) on the basis of two
specimens from Nifoekoko {Prohun^arites
fauna) in the British Museum (Natural
History). This species is also present in
Afghanistan (Kummel, 196Sb) and West
Pakistan (Kummel, 1966).
Occurrence. Siibcolumlntes faunas of Al-
bania, Chios, Afghanistan, Kwangsi, China
(Chao collection 542b), and Timor; from
the Pseiidusagcceras Zone of Astakhova
( 196()b ) in the Mangyshlak Peninsula,
Caspian region; from the Proliuuii^arilcs
fauna. Salt Range, West Pakistan.
Repository. The following specimens are
in the Paleontological Institute, Vienna:
lectotype (Arthaber, 19()(S, pi. 11(1), figs,
la-c); paralectotypc (Arthaber, 190S, pi.
11(1), figs. 2a, b); plesiotvpes (Arthaber,
1911, pi 17(1), figs. 16, 17; pi. 18(2), figs.
5, 6, 7). Topotvpes, BMNH C22700-05,
C22706-24, C22761-2, C22694-9, C34116-7,
C22725, C22882. The specimens from Chios
are in the Natural History Museum, Basel,
and are as follows: plesiotypes P. kokeni,
Renz and Renz (1948, pi. 8, fig. 5) NHMB
J13768, (pi. 8, figs. 6-6a) NHMB J13769,
(pi 8, figs. 7-7a) NHMB J13772, (pi 8,
figs. 8-8a) NHMB J13770, (pi 8, figs. 9-9a)
NHMB J13773, (pi 9, figs. 2-2a) NHMB
J 13771; unfigured specimens from Mara-
dovuno NHMB J13774-13779, from Kep-
halovuno NHMB J13780; var. pantcleimo-
nensis Renz and Renz ( 1948, pi 8, figs.
3-3a) NHMB J13784, (pi 9, figs. 3-3a)
NHMB J13785; unfigured specimens from
Maradovuno NHMB J13786, from Kep-
halovuno NHMB J13787; var. evoluta Renz
and Renz (1948, pi 9, figs. 1-la) NHMB
J13781; unfigured specimens from Mara-
dovuno NHMB J13782, from Kephalovuno
NHMB J13783; plesiotype P. skanderbegis,
Renz and Renz (1948, pi 8, figs. 4-4a)
NHMB J13788; unfigured specimens from
Maradovuno NHMB J13789, from Kepha-
lovuno NHMB J 13790; topotypes MCZ
10021, 10022; specimens from West Paki-
stan, MCZ 9593-9595; specimens from
Kotal-e-Tera, Afghanistan, MCZ 10154,
10155.
Procarnifes immoturus (Kiparisova)
Text-figure 12
McfiapJujUitCi iiuiiuituni.s Kiparisova, 1947: 130,
pi. 27, figs. 1, -2, text-fig. 8; Kiparisova, 1954:
22, pi. 12, fig. 4; Kipari.sova, 1961: 172, pi.
35, figs. 3-5, text-figs. 115-117; Tozer, 1965a:
39.
Procarnitcs inndcsfus Tozer, 1965a: 38, pi. 1, figs.
1-6, text-fig. 12.
This species can be distinguished on the
basis of faint radial constrictions; in all
other features it is quite similar to P. kokeni
(see illustration of suture, Figure 12H).
Tozer ( 1965a ) recognized the close resem-
blanc(> between his P. modestus and P.
imniaturus but relied primariK- on a slight
difference in the umbilical diameter to
separate the two species. Considering that
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 397
the specimens of P. modestus have crushed
l)ody chambers and the near identity in
the other morphologic features, I can see
no justification for separating these species.
Occurrence. Subcohmibites fauna, Cape
Zhitkov, Primorye Region, Siberia, and Toad
Formation, Halfway River area, British
Columbia.
Procarnites lolouensis (Chao)
Text-figure 12
Digitophi/Uites lolouensis Chao, 1950: 5, pi. 1,
tigs. 4^6; Chao, 19.59: 90, 256, pi. 32, tigs.
13-19, text-figs. 29a, b.
Chao ( 1959 ) recognized that this species
was a procarnitid but established a new
genus for it because it differed from Pro-
carnites in ( 1 ) having a wider umbilicus,
(2) having a broadly arched venter rather
than a narrow or acute one, encompassing
a subquadratic whorl section, and (3)
having a different suture. In the first place,
the umbilicus compares very favorably in
diameter to the specimen of Procarnites
kokeni of comparable size from Chios.
Chao gave measurements on four speci-
mens of diameters from 30 to 27 mm; the
umbilical diameters ranged from 6.0-4.5
mm. Among the specimens of Procarnites
kokeni from Chios, those of a diameter of
30.3 to 26.8 mm have a range of diameter
of 5-3 mm.
The more inflated, subquadrate whorl is
quite distinctive. Finally, a comparison of
the sutures of these Kwangsi specimens
and those at the same relative conch size
is shown on Figure 121, J. The adventitious
elements mentioned by Chao do not appear
until a much later stage of growth. All in
all I can see no justification for establishing
a new genus for this species. Chao's speci-
mens could well be juvenile forms, but
they are no more than a more inflated form
of Procarnites and in this respect differ
from kokeni and immaturus.
Occurrence. Subcolumbites fauna, Nali-
ling sections near Lolou village (Chao col-
lections 541a, b, 542b), Kwangsi, China.
Family PARANANNITIDAE Spath, 1930
Genus Arnautocelfites Diener, 1916
Type species, Celfiies arnauticus Arthaber,
1911
This genus is confined to the late Scythian
Prohungarites Zone where it is represented
by five species: A. jnediterraneus from the
Subcolumbites fauna of Albania and Chios,
A. bajarunasi from the late Scythian strata
of the Mangyshlak Peninsula, A. involutus
from the Subcolumbites fauna of Kwangsi,
China, A. gracilis from the Subcolumbites
fauna of the Primorye Region, and A.
teichei-ti from the Subcolumbites fauna of
Nevada. Of special interest in this group
of species is the very large degree of in-
traspecific variation in the Albanian and
Chios A. mediterraneus, in contrast to a
very limited degree of such variation in
A. teicherti from Nevada. The other species
are represented by very few specimens.
Arnautoceitites mediterraneus (Arthaber)
Plate 6, figures 7-13; Plate 7, figures
5, 6; Text-figure 14
Paranannites lucclitcnaneus Arthaber, 1911: 220,
pi. 18(2), fig. 8; Diener, 1915: 216; C. Renz,
1928: 155; Renz and Renz, 1947: 61, 66; Renz
and Renz, 1948: 69, pi. 1, figs. 12, 12b, 13,
13a, 17.
Arnautoceitites mediterraneus, — Spath, 1934: 193,
pi. 14, figs, la-c, text-fig. 59f.
Paranannites mediterraneus Arthaber var. media
Renz and Renz 1947: 77; Renz and Renz,
1948: 70, pi. 1, figs. 11-llb, 14-14b.
Paranannites cliionensis Renz and Renz, 1947: 66,
77; Renz and Renz, 1948: 70, pi. 1, figs. 10-lOc.
Paranannites aspenensis Hyatt and Smidi var.
europaca Renz and Renz, 1947: 61; Renz and
Renz, 1948: 71, pi. 1, figs. 16-16c.
Paranannites compressus Renz and Renz, 1947:
61, 77 (non Smith, 1932: 99, pi. 31, figs. 19-20);
Renz and Renz, 1948: 71, pi. 1, fig. 15-L5b.
Paranannites chiosensis Kiparisova, 1961: 130
( = P. compressus Renz and Renz, 1948, non
Smith, 1932).
Celtites arnauticus Arthaber, 1911: 267, pi. 24(8),
fig. 7; Diener, 1915: 73; Smith, 1932: 37.
Arnautoceitites arnauticus, — Spath, 1934: 192, pi.
13, figs. 6a-f.
Puragoceras dukagini Arthaber, 1911: 182, 188,
265, pi. 24(8), fig. 6; Diener, 1915: 366; C.
Renz, 1928: 1.55; Kutassy, 1933: 607; Spath,
398 Bulletin Museum of Comporative Zoology, Vol. 137, No. 3
Ammonoids of the Late Scythian (Lower Triassic) • Kummel
399
1934: 199, fig. 60; Kiimniel, in Arkell, et al.,
1957: L139, figs. 172, 3; Renz and Renz, 1948:
96.
Nannites herherti, — Aithaber ( non Diener), 1908:
274, pi. 11 ( 1), figs. 7a, b; Arthaber, 1911: 220.
A characteristic ammonite of the Sub-
coJiimbites fauna of Albania is a small form,
generally involute, with prosiradiate con-
strictions. Arthaber (1908, 1911) classified
ammonites of this general morphology in
four species in four different genera. These
species are:
Paraiuinnites mediterraneus Arthaber
Ccltitcs ariiatiticus Arthaber
Panigoccni.s dukagini Arthaber
Nannites herberti Diener
The first of the species was based on 5
specimens of which only 2 are preserved;
the second species was based on 17 speci-
mens of which only one exists; the last
two species were each based on a single
specimen and these are still preserved. The
basic difference between these species as
recognized by Arthaber lay in the suture.
Paranannites mediterraneus was stated to
have a single, serrated, lateral lobe, Cel-
fitcs arnauticiis a goniatitic lateral lobe,
Para{i,oceros dukag,ini a particularly unique
suture (of which more later), and finally,
no suture was available on the specimen
assigned to Nannites herberti.
The suture of the lectotype of Paranan-
nites mediterraneus reproduced by Arthaber
(1911: pi. 18(2), fig. 8c) is somewhat
poorly executed, but more significant, it is
incomplete. A new drawing of the suture
of this type specimen is reproduced here
on Figure 14J. The principal new datum
on the suture is the presence of a broad,
shallow, second lateral lobe on the umbilical
shoulder and wall.
The goniatitic suture of Celtites arnau-
ticiis needs verification. The only suture
known of this species is that reproduced
by Arthaber (1911: pi. 24(8), fig. 7d).
It is only by implication that one would
believe that this suture is from the figured
specimen of Arthaber's plate 24(8), figures
7a-c. However, this specimen is preserved
and shown here on Plate 6, figures 12, 13;
no sutures are visible on the specimen.
Likewise, Spath (1934: 192) did not un-
cover a suture among the topotypes in the
British Museum, and resorted to repro-
ducing Arthaber's data. The hard lime-
stone preservation of the Albanian Stib-
cohimbites fauna requires grinding and
polishing to expose the suture. It has been
demonstrated in this report that the sutures
of many of the species described by Artha-
ber are to a greater or lesser extent distorted
and inaccurate representations mainly due
to excessive preparation. There is every
reason to believe this is the case with the
suture of Celtites arnauticiis. Tliis sugges-
tion is strengthened on consideration of
Figure 14. Diagrammatic representation of the sutures of various species of Arnautoce/fites. A-l, Arnaufoce/t/tes teicherti
n. sp., from Tobin Formation of Nevada; A, at a diameter of 11 mm (MCZ 9617); B, at a diameter of approximately 11
mm (MCZ 9618); C, at a diameter of 10.5 mm (MCZ 9619); D, at a diameter of 9.5 mm (MCZ 9620); E, at a diameter
of 9 mm (MCZ 9621); F, at a diameter of 9 mm (MCZ 9622); G, at a diameter of 8.5 mm (MCZ 9623); H, at a diam-
eter of 8.5 mm (MCZ 9624); I, at a diometer of 7 mm (MCZ 9625); J, lectotype of Paranannites mediterraneus Artfiaber
(1911), from Subco/umbites fauna, Albania; K, plesiotype of Poronannites mediterraneus, — Renz and Renz (1948: pi. 1, fig.
17), at a whorl height of approximately 9 mm, from Subco/umbites fauna of Chios (NHMB-Jl 3729); L, syntype of Paranan-
nites mediterraneus var. media Renz and Renz (1948: pi. 1, fig. lib), at a diameter of approximately 20 mm, from Sub-
columbifes fauna of Chios (NHMB-Jl 3732); M, holotype of Paranannites chionensis Renz and Renz (1948: pi. 1, fig. 10c),
at a diameter of approximately 16 mm, from Subco/umbifes fauna of Chios (NHMB-J13737); N, holotype of Parononnites
compressus Renz and Renz (1948: pi. 1, fig. 15b), at a diameter of approximately 20 mm, from Subco/umb/fes fauna of
Chios (NHMB-J13736); O, holotype of Paranannites gracilis Kipariso^c (1961: fig. 93), at a diameter of approximately 10
mm; P, holotype of Poronannites involutus Chao (1959: fig. 37d), at a whorl height of 9 mm from beds with Subcolum-
bites, Kwangsi, China; Q, holotype of Porononnifes minutus Chao (1959; fig. 37b), at a whorl height of 10.5 mm, from
Subcolumbites fauna, Kwangsi, China; R, holotype of Parogoceros dul<agini Arthaber (1911), at a diameter of approximately
11 mm, from Subco/umbifes fauna, Albania.
400 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
the remaining morphological features of
the shell. In this regard, the type specimen
of Celtites ornauticus differs from the type
specimen of Paranannites mediterraneus
only in being slightly more evolute. It is
unfortunate that the sample from the SfJ;-
columbitcs fauna of Albania is so small,
but Spath (1934: 192), in commenting on
the few topotype specimens of Celtites
arnauticus and Paranannites mediterraneus
in the collections of the British Museum,
states "there are thin and thick varieties of
each." I cannot consider the differences
in degree of involution of these two species
(specimens) as a distinction warranting
separation of the two forms.
The single specimen Arthaber ( 1908 ) as-
signed to Nannites herherti is illustrated
here on Plate 6, figures 7, 8. It does not
preserve a suture but in all other morpho-
logical features it is nearly identical to the
specimen assigned to Paranannites mediter-
raneus.
Paragoeeras dukagini (Arthaber, 1911:
265) was based on a single, poorly pre-
served specimen (Pi. 7, figs. 5, 6). The
conch dimensions, degree of involution, and
pattern of constrictions show the specimen
to be identical with A. mediterraneus. It
was primarily on the basis of the unusual
suture reproduced by Arthaber (1911: pi.
24(8), fig. 6c) that Spath (1934) and
others have accepted the genus. Re-ex-
amination of the type specimen shows that
the suture has been damaged in prepara-
tion, but a new drawing of what is visible
is shown on Figure 14R. The suture is that
of a typical representation of A. mediter-
raneus and very imlike that reproduced by
Arthaber.
The SuJ)C()]u)nJ)ites fauna of Chios
studied by Renz and Renz ( 1948 ) contains
19 specimens which they assigned to four
species of Paranannites (= Arnautoceltites).
Of these, however, only six of the figured
specimens are sufficiently well preserved
and complete to yield significant measure-
ments, which are given on Table 29. Each
of th('S(» specimens which yielded a suture
Table 29. Measurements of Arnautoceltites
MEDITERRANEUS ( ArTHABER ) FROM THE SUBCO-
LUMBITES FAUNAS OF ALBANIA AND ChIOS.
D
w
H
u
W/D
H/D
U/D
1.
26.1
9.5
10.9
7.6
36.4
41.8
29.1
0_
23.1
p
9.2
7.2
?
39.8
31.2
3.
23.0
7.0?
10.0
3.5
.30.4?
43.5
15.2
4.
19.3
10.0
6.6
8.0
51.8
34.2
41.5
5.
18.5
11.0
6.6
6.4
59.5
.35.7
34.6
6.
18.0
11.2
6.8
6.5
62.2
37.8
36.1
7.
17.4
11.2
7.0
5.0
64.4
40.2
28.7
8.
17.1
9.7
6.8
6.0
56.7
39.8
.35.1
9.
16.5
?
6.4
4.9
?
38.8
29.7
10.
16.0
?
6.1
5.2
?
38.1
32.5
11.
15.6
?
7.3
3.7
?
46.8
23.7
1.
Holotvpe, Paranannhcs chionensis Renz and Renz
(1948, pi. 1, fig. 10), NHMB J13737.
2. Plesiotvpe, P. mediterraneus var. media Renz and
Renz (1948, pi. 1, fig. 11), NHMB J13732.
3. Holotvpe, Paranannites compresstis Renz and Renz
(1948, pi. 1, fig. 15), NHMB J13736.
4. Lectotype, Arnautoceltites arnauticus (Arthaber, 1911,
pi. 24(8), fig. 7).
5. Holotvpe, Paragoeeras dtikagini Arthaber (1911, pi.
24(8); fig. 6).
6. Plesiotype, Paranannites mediterraneus, — Renz and
Renz (1948, pi. 1, fig. 12), NHMB J13727.
7. Lectotvpe, Arnautoceltites mediterraneus (Arthaber,
1911, pi. 18(2), fig. 8).
8. Plesiotype, Na)tnites hcrberti, — Arthaber ( 1908. p. 274,
\A. 11(1), fig. 7) ( non Diener).
9. Plesiotvpe, P. mediterraneus var. media Renz and
Renz (1948, pi. 1, fig. 14), NHMB J13733.
10. Paralectotvpe, Arnautoceltites mediterraneus (Art-
haber, 1911, p. 220).
11. Plesiotype, P. aspetiensis var. eurojiaea Renz and Renz
(1948, pi. 1, fig. 16), NHMB J13735.
Specimens 1-3, 6, 9, 11 are from the Suhcolumbites
fauna of Chios; specimens 4, 5, 7, 8, 10 are from the
Suftenlumhites fauna of Albania.
has denticulated lobes. The specimens as-
signed to Paranannites mediterraneus (Renz
and Renz, 1948: pi. 1, figs. 12, 13, 17),
Paranannites mediterraneus var. media
(Renz and Renz, 1948: pi. 1, figs. 11, 14),
and ParatuDinites aspenensis Hyatt and
Smith var. europaea (Renz and Renz, 1948:
pi. 1, fig. 16) are clearly conspecific with
the Albanian type specimen of Arnautocel-
tites mediterraneus. One highly compressed
specimen was separated as Paranannites
eonipressus (Renz and Renz, 1948: pi. 1,
fig. 15) and another specimen intermediate
in degree of compression between com-
pressus and niediterranetis was set aside as
Paranannites chiosensis. The sutures of
these Chios specimens (Fig. 14M) are all
essentiallv the same, differing in minor
Ammonoids of the Late Scythian (Lower Triassic) • Ktimmel 401
details. The prosiradiate constrictions are
variable in both the number per volution
and in the degree of fonvard projection.
It is indeed unfortunate that the sample
available from the Siibcolumbites fauna of
Chios is so limited. However, on the basis
that these specimens came from one horizon
and locality and were most probably part
of a single population unit, and that other
species of this population (for example
Isciilituidcs originis) show a comparable
range in shell variation as between mediter-
r uncus and cornpressus, I believe it best to
consider all these Chios specimens as mem-
bers of a highly variable species — mcditer-
rancus.
Occurrence. Subcolumbites fauna of Al-
bania and Chios.
Repository. The following specimens
from Albania are in the Paleontological
Institute, Universitv of Vienna: lectotvpe
(Arthaber, 1908: pi. 18(2), fig. 8) and
one paralectotype; lectotype of Celtites
arnauticus Arthaber (1911: pi. 24(8), fig. 7);
plesiotype of Nannites herberti, — Artha-
ber (1908: pi. 11, fig. 7) (non Diener);
holotype of Farag,oceras dukaii'mi Arthaber
(1911: pi. 24, fig. 6). In addition, the
British Museum (Natural History) has
some topotype specimens. The following
specimens from Chios are in the Natural
History Museum of Basel: plesiotype (Renz
and Renz, 1948: pi. 1, fig. 12) NHMB
J13727, (Renz and Renz, 1948: pi. 1, fig.
13) NHMB J13728, (Renz and Renz, 1948:
pi. 1, fig. 17) NHMB J13729; Paranannites
mediterraneus var. media (Renz and Renz,
1948: pi. 1, fig. 11) NHMB J13732, (Renz
and Renz, 1948: pi. 1, fig. 14) NHMB
J13733; unfigured specimen NHMB J13734;
holotype Paranannites cJiionensis Renz and
Renz (1948: pi. 1, fig. 10) NHMB J13737;
unfigured paratype NHMB J13738; type of
Paranannites aspenensis Hyatt and Smith
var. europaea Renz and Renz (1948: pi. 1,
fig. 16) NHMB J13735; holotype Paranan-
nites cornpressus Renz and Renz (1948:
pi. 1, fig. 15) NHMB J13736; specimen of
Paragoceras dukagini Arthaber (Renz and
Renz, 1948: p. 96) NHMB J13838; topo-
types from Chios MCZ 10115.
Arnautoceltites bajarunasi (Astakhova)
Nannites bajarunasi Astakhova, 1960a: 145, pi. 34,
figs. 2, 3; Astakhova, 1960b: 150.
This species has been based on two small
specimens that morphologically are not too
different in appearance from the Albanian
A. mediterraneus. There appears to be a
tendency on the outer volution for the
whorls to be vaulted. The most significant
characteristic is the goniatitic suture with
a large, smooth lateral lobe followed by
a much smaller lobe on the umbilical
shoulder.
Occurrence. In Stacheites Zone of Astak-
hova (1960a, b) on the Mangyshlak Penin-
sula, said to be associated with Stacheites
prionoides.
Arnaufocelfifes involufus Chao
Text-figure 14
Paranannites invohitus Chao, 1959: 113, 285, pi.
24, figs. 13-15, 18, 20, 25, text-fig. 37d.
Paranannites niinutus Chao, 1959: 114, 286, pi.
24, figs. 16, 17, text-fig. 37b.
A very involute species most comparable
to (and possibly conspecific with) Paranan-
nites gracilis Kiparisova from the Primorye
Region. The suture (Fig. 14P, Q) has a
distinct second lateral lobe. Data on this
species are very incomplete.
Occurrence. Subcolumbites fauna,
Kwangsi, China (Chao collection 546).
Arnautoceltites gracilis (Kiparisova)
Text-figure 14
Paraminnites gracilis Kiparisova, 1947: 140, pi.
28, fig. 1, text-fig. 25; Kiparisova and Krishtofo-
vich, 1954: 21, pi. 12, fig. 1; Kiparisova, 1961:
125, pi. 28, figs. 3, 4, text-figs. 92, 93.
Paranannites minor Kiparisova, 1961: 129, pi. 28,
figs. 1, 2, text-fig. 98.
An involute species quite similar to A.
involutus Chao and possibly even con-
specific with it. The suture (Fig. 140)
is different in that the second lateral lobe
occupies all of the umbilical shoulder and
402 BiiUeiin Miiscmn of Comparative Zoology, Vol. 137, No. 3
Table 30. Measurements of Arnautoceltites
teicherti n. sp. from the tobin formation
OF Nevada.
D
w
H
u
W/D
H/D
U/D
1.
21.3
12.1
8.2
6.2
56.8
38.4
29.1
2.
21.3
11.3
?
?
53.5
?
?
.3.
20.2
12.1
8.7
4.8
59.4
43.0
23.2
4.
20.2
10.3
7.7
5.7
50.9
38.1
28.2
5
20.0
11.9
9.6
4.9
59.5
48.0
24.5
6.
19.7
12.4
7.7
5.9
62.9
39.0
29.9
7.
19.7
11.3
7.6
5.7
52.2
38.6
28.9
8.
19.6?
10.4
7.6
6.4
53.0?
38.7
32.6
9.
19.0
11.0
8.0
5.2
57.8
42.1
27.3
10.
18.8
11.2
8.0
5.0
59.5
42.5
26.5
11.
18.8
10.8
6.8
5.7
57.4
36.1
30.3
12.
18.1
9.6
7.3
5.3
53.0
40.3
29.2
13.
17.7
9.6
7.8
4.8
54.2
44.0
27.0
14.
17.5
11.7
7.4
3.8
66.8
42.2
21.7
15.
17.5
10.2
6.7
4.6
58.2
38.2
26.2
16.
16.5
9.2
6.6
4.3
55.7
40.0
26.6
17.
16.4
10.3
6.5
4.2
62.8
.33.5
25.6
18.
16.2
9.5
6.7
4.3
.58.6
41.3
26.5
19.
16.0
9.1?
7.1
4.0
56.8?
44.3
25.0
20.
15.6
9.4
6.7
4.1
60.2
42.9
26.2
21.
15.5
9.0
6.5
4.1
58.0
41.9
26.4
22.
15.3
9.0
6.5
4.1
58.8
42.4
26.7
23.
15.0
9.1
6.0
3.6
60.6
40.0
24.0
24.
14.8
9.3
6.0
3.6
63.5
40.5
24.3
25.
14.8
8.7
6.9
3.5?
56.0
46.6
23.6?
26.
14.6
9.0
5.4
3.7
61.6
36.9
25.3
27.
14.5
8.5
5.9
3.5
58.6
40.6
24.1
28.
14.4
8.2
6.1
3.1
56.9
42.3
21.5
29.
14.3
8.5
5.8
3.8
59.4
40.5
26.5
30.
14.1
9.4?
5.7
3.5
66.6?
40.4
24.8
31.
13.8
9.2
6.1
4.0
66.6
44.2
28.9
32.
13.3
8.8
5.7
3.3
66.1
42.8
24.8
33.
12.6
7.7
5.6
3.0
61.1
44.4
23.8
34.
11.1
7.6
4.6
3.0
68.4
41.4
27.0
35.
10.8
7.3
4.8
2.3
67.5
44.4
21.2
36.
9.6
6.5?
4.4
2.2
67.7
45.8?
22.9
37.
9.0
6.2
4.0
2.2
68.8
44.4
24.4
38.
8.4
6.4
4.2
1.2
76.1
50.0
14.2
3.
4.
Paratype,
Paratype,
Paratype,
Paratype,
Paratype,
Paratype,
All iitlicr
9490}.
8.
9.
12.
MCZ 9460
MCZ 9457
MCZ 9458
MCZ 9462
MCZ 9461
MCZ 9459
speciniens are
(PI. 31, figs. 6, 7).
(PI. 31, figs. 1, 2).
(PI. 31, figs. 3, 4).
(PI. 31, figs. 13, 14).
(PI. 31, figs. 9, 10).
(PI. 31, fig. 5).
iiiifigured iiarat> pes
(MCZ
wall. B(Uh lliis spctics and the Kwangsi
A. invohitiis diflei- from A. meditcrrancus
and A. teicherti in the .suture and degree
of involution of the eoneh.
Occurrence. Su])C(>ltun1)ile.s- fauna. Cape
Zhitkov, Primorye Region, Siberia.
Arnautoceltites teicherti n. sp.
Plate 31, figures 1-7, 9-14; Text-
figures 14, 15
The Siibcohimhite.s fauna from the Tobin
Formation of Nevada has yielded approxi-
mately 75 specimens of this new species,
of which 38 are sufficiently well preserved
and complete to obtain measurements. The
measurements are given on Table 30 and
the plot of these data shown on the graph
of Figure 15.
The basic morphology of the shell, that
is, degree of involution, whorl cross-section,
pattern of constrictions, etc., is very much
like that of the Albanian Arnautoceltites
meditcrrancus. It is only in the suture that
a small but subtle difference between these
species can be recognized. On Figure 14
are 9 sutures of A. teicherti. It is readily
seen that there is a high degree of vari-
ability. This is especially noted in the
shape and pattern of denticulation of the
first lateral lobe. The second lateral lobe,
lying on the umbilical shoulder and wall,
is likewise variable but in all cases more
pronounced than in the Albanian and
Chios representations of A. mediterraneus.
It is primarily on this feature that I con-
clude the two species are specifically dis-
tinct but closely related. These two species
are quite distinct from the other species of
Arnautoceltites so far recorded.
Occurrence. Lower part Tobin Forma-
tion, U.S.G.S. Mesozoic locaHty M2562,
Pershing County, Nevada; south tip of
Tobin Range, Cain Mountain 1:62,500
quad., centei- NW M sec. 9, T. 26N, R. 39E,
5,500 ft. S, 27.5 ft. W from elevation point
5088 on range crest.
Repository. Holotype, MCZ 9457 (PI. 31,
figs. 1, 2); paratypes, MCZ 9458 (PI. 31,
figs. 3, 4), MCZ 9459 (PI. 31, fig. 5),
MCZ 9460 (PI. 31. figs. 6, 7), MCZ 9461
(PI. 31, figs. 9, H)),\\CZ 9462 (PI. 31,
figs. 13, 14); suture specimens Figures
31A-I, MCZ 9617-9625; unfigured para-
tN'pcs MCZ 9490.
Ammonoids of the Late Scythian (Lower Triassic) • Kiiwmel 403
10
9
7
6
5
4
3
2
W
H
U
0
13 14 15 16 17
D I A METER
8
19 20
22 23
Figure 15. Variation in whorl height (H), whorl width (W) and umbilical diameter (U) in Arnoufoce/t/fes teicherti n.
sp. from the Tobin Formation of Nevada. The data on this graph are from Table 30.
Genus Prosphingites Mojsisovics, 1886
Type species, Prosphingites czekanowskii
Mojsisovics, 1886
Prosphingites has until the last few
years been considered mainly a late Scy-
thian genus {Suhcolwnbites — Prolnmgarites
Zone). The type species, P. czekanowskii
Mojsisovics (1886: 64, pi. 15, figs. 10-12)
came from the Olenekites fauna of northern
Siberia, of upper Scythian age. Hyatt and
Smith (1905: 72, pi. 7, figs. 1-4) later
described P. austini from the Meekoceras
fauna of southern California. Spath ( 1934 )
was uncertain as to the status of P. austini
and in his interpretation of the zonal range
of the genus tended to place more reliance
on the type species from the Olenek beds
and P. spathi (Frebold, 1930) from the so-
called Fish beds of Spitsbergen, which he
considered to be late Scythian in age, more
or less contemporaneous with the Olenek
fauna. Both Kummel ( 1961 ) and Tozer
( 1961a ) have presented convincing argu-
ments that this particular Spitsbergen
horizon is mid-Scythian {Owenites Zone)
in age. In recent years a number of new
404 Bulletin Museum of Comparative Zoolofiy, Vol. 137, No. 3
species of Prosphin^ites have been de-
scribed from Chios ( Renz and Renz, 1948 ) ,
Kwangsi, China (Chao, 1959), eastern
Siberia (Kiparisova, 1961), Arctic Canada
(Tozer, 1961a), and western United States
(Kmnmel and Steele, 1962).
From the mid-Scythian, Owenites Zone,
the following species of Prospliingites are
known: Prosphing,ites ousiini Hyatt and
Smith, P. ovalis Kiparisova, P. orientalis
Kiparisova, P. sinensis Chao, P. invohitus
Chao, P. kwan<i,sianus Chao, P. radians
Chao, P. spathi Frebold, P. slossi Kummel
and Steele. The documentation and illus-
tration of all these species from essentially
contemporaneous deposits in western United
States, Arctic Canada, Spitsbergen, eastern
Siberia, and southern China has added
much to our understanding of Prosp]}inp,ites
during the mid-Scythian. When I intro-
duced the species P. slossi (Kummel and
Steele, 1962) I was fully cognizant of its
close relationship and perhaps identity to
P. austini Hyatt and Smith; however, on
the argument that P. austini was known
only from a single, not very well preserved
specimen, I considered it best to ignore the
species. However, subsequently, on study
of all the new Scythian species that have
been introduced, and thorough restudy of
P. austini, I am convinced that P. ovalis,
P. orientalis, P. sinensis, P. radians, P.
spathi, and P. slossi are synonyms of P.
austini. Large populations of these species
are known only for P. spathi and P. slossi.
Large numbers of specimens of P. spathi
are in the British Museum (Natural His-
tory), but only a few measurements are
available. Nearly all the Spitsbergen speci-
mens are small phragmocones or juvenile
specimens.
Kummel and Steele (1962: 683) have
presented measurements on 49 specimens
of P. slossi from the Meekoceras beds at
Crittenden Spring, Nevada. A plot of the
available measurements of the other species
of Prosphinf^ites shows them to fall within
the limits of variations for P. slossi. The
Nevada fauna likewise shows that the pat-
tern of constrictions is highly variable.
Evaluation of the suture in all these species
is more difficult, as generally only one
pattern is given for any species by most
authors. However, on the basis of the
variability in suture within the Crittenden
Spring faima and within P. spathi from
the Canadian Arctic described by Tozer
(1961d), I believe that the pattern for all
the various species placed in the synonmy
of P. austini is essentially the same, and
what variation is present is partly due to
the small size of the sample (generally one
suture per species) and, more fundamen-
tally, is no more than one should expect.
The suture of these mid-Scythian species
of the Owenites Zone is in general simpler
than for species in higher Scythian zones.
This is especially marked in the nature of
the ventral lobe and in the auxiliary lobe
(Fig. 16).
No species of Prosphin0tes are known
from the Columhitcs Zone. The following
species have been described from the upper
Scythian Suhcolumhites-Olenekites fauna:
Prosphing,ites czekanowskii Mojsisovics, P.
pjohosus Kiparisova, P. insularis Kiparisova,
P. lolouensis Chao, P. ali Arthaber, P.
vondersehmitti Renz and Renz, and P.
eoomhsi Kummel.
Prosphingites czekanowskii is unique in
its compressed living chamber and acute
venter, but the suture is relatively simple
( Fig. 16G, H ) and not unlike that of species
in the Owenites Zone. Prosphingites
g,lohosus is based on a few, very small, im-
mature specimens and is too incompletely
known to make meaningful comparisons;
the same can be said for P. ali, but in each
case the suture is of an advanced type with
a more distinct auxiliary lobe and more
elaborate ventral lobe (Fig. 16D-F). Pro-
sphimgites lolouensis is based on a few
poorly preserved specimens. Isculitoides
<^lohosus Chao (1959: 292, pi. 26, figs.
9-13) appears to be a synonym of P.
lolouensis Chao. Pros))hin[!.ites vonderseh-
mitti from the Suhcolunibites fauna of
Chios is a species of Zenoites.
Ammonoids of the Late Scythian (Lower Triassic) • Kinnincl 405
Thus, there are now 16 described species of the venter. The suture is shown on
of Prosphingites. Analysis of the nine Figures 6G, H. I include within this species
species described from the mid-Scythian the specimen Tozer ( 1965a ) records from
Oicenites Zone leads me to accept only British Columbia. Even though this speci-
four of these as valid; the remaining five men is partially crushed, the visible conch
are synonyms. The valid species are: P. features and the suture indicate this identi-
atistini, known from western United States, fication.
Ellesmere Island, Spitsbergen, Primorye Re- Tliis species is unique in that in no other
gion, and Kwangsi; P. involutus Chao, an species assigned to Prosphingites does the
incompletely known species from Kwangsi, venter become acute. In the suture, also,
China; P. kwangsianus Chao, another in- this species differs from most other species
completely known species from Kwangsi; of Prosphingites from the upper Scythian
P. magmimhiUcattis Kiparisova, presum- in the reduced character of the auxiliary
ably from the Pro.?/;/i/ng;Ye5 Zone (=Oticn- lobe. In this aspect it is more comparable
ites Zone) of the Primorye Region. All of to the suture found in species of Pro-
the species from the upper Scythian Sub- sphingites from the Oicenites Zone. There
cohimbites Zone are believed to be valid is also no development of phylloid saddles
excejDt P. voncJcrschmitti Renz and Renz, such as characterize P. glohosiis, P. instdaris
which is a species of Zenoites. and P. coombsi, all of late Scythian age.
The principal difference between most Occurrence. Olenekifes fauna, northern
of these species of Prosphinigites of the Siberia, and Toad Fonnation, Halfway
upper Scythian and those of the mid- River area, British Columbia.
Scythian lies in the suture. The species Repository. The Museum of Compara-
from the upper Scythian tend to have su- tive Zoology has one topotype specimen
tures with phylloid saddles, more elaborate (8677).
dcnticulation of the lobes, and a distinct
auxiliary lobe. Prosphingites ali Arthaber
Plate 20, figures 12, 13; Text-figure 16
Prosphingites czekanowskii Mojsisovics Prosphingites ali Arthaber, 1911: 252, pi. 22(6),
Plate 26, figure 8; Text-figure 16 figs. 6, 7; Diener, 1915: 233; Spath, 1934:
Prosphingites czekanowskii Mojsisovics, 1886: 64,
pi. 15, figs. 10-12; Noetling, in Freeh, 1905: Arthaber ( 1911: 252) states he had three
^!lo'. P^^^'/j- ^'.P^'^t"''''' ^^^'^^ ?n?; ^^f!n' specimens of this species but only the
1934: 33, 195, 196; Kiparisova, 1937: 140, ^ . i -n . . j • j -ru
pi. 1, fig. 2; Kiparisova, 1947: 142, pi. 32, specimen he illustrated is preserved. The
figs. 4, 5, text-fig. 28; Kummel, 1961: 522; dimensions of this specimen are as follows:
Popov, 1961: 58, pi. 13, fig. 4. Diameter 15.3, Width 10.2, Height 6.0,
Prosphingites ci. P. czekanowskii,— Tozer, 1965a: Umbilicus 4.7 mm. The specimen pre-
. , p . -, igs. . a-c, ex - ig. . . serves much of the shell but this has been
This very characteristic and almost polished and is perfectly smooth. This
unique type species is known from rela- specimen likewise does not show any trace
tively few specimens. In tlie two main of a suture; it is presumed that the suture
discussions of the species by Mojsisovics Arthaber figured (1911, pi. 22(6), fig. 6c;
(1886) and Popov (1961) it appears there Fig. 16F of this report) came from one of
were only a total of five specimens. One the other two specimens he had in his
topotype specimen is now in the Museum original collection.
of Comparative Zoology. The major fea- This species was based on what are ob-
tures of the species are the globose, de- viously juvenile forms, making comparisons
pressed inner whorls grading adorally to with other species of the genus tenuous at
more compressed whorls, and a sharpening best. It is of special interest tliat the closely
406 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
1
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 407
related Subcohimbites fauna of Cliios does
not contain any true Frosphinii^ites. The
specimen described by Renz and Renz
(1948: 39, pi. 15, figs. 13, 15) as ?ro-
spJiinfiitcs vonderschmitti is a species of
Zenoites; the specimen they assigned to
Prosphingites ex aff. czekonowskii is a rep-
resentative of Zenoites helenae Renz and
Renz.
Occurrence. Subcolumbites fauna, Kcira,
Albania.
Repository. The holotype is in the
Paleontological Institute, Vienna.
Prosphingites lolouensis Chao
Prosphingites lolouensis Chao, 1959: 123, 298, pi.
27, figs. 18-24, text-fig. 39b.
Isctilitoides globosus Chao, 1959: 119, 292, pi. 26,
figs. 9-13.
This is a most unsatisfactory species
group. Prosphingites lolouensis was estab-
lished for a "great number" of deformed
specimens tliat never should have been
given a new specific name. The specimens
assigned by Chao (1959) to Isculitoides
globosus have a prosphingitid suture, as
noted by Chao (1959: 293), but he dis-
tinguished it on differences in the nature
of the umbilical shoulders. Even consider-
ing the state of preservation of the Kwangsi
material, I cannot separate these specimens
from those assigned to P. lolouensis.
Occurrence. Subcolumbites fauna of
Chashanao sections (Chao collection 610)
and northeast of Lolou village (Chao col-
lection 541a, b), Kwangsi, China.
Prosphingifes subglobosus (Chao)
Paranannites subglobosus Chao, 1959: 113, 285,
pi. 24, figs. 21-24, text-fig. 37a.
The most distinctive aspect of this species
is the eccentricity of the umbilicus on the
last volution. In this respect this species
is quite different from all others assigned
to Prosphingites. The suture is quite like
that of the type species P. czekanoivskii
with a weakly developed auxiliary series.
Occurrence. Subcolumbites fauna, NaH-
ling section (Chao collections 542a, b),
Kwangsi, China.
Prosphingites globosus Kiparisova
Text-figure 16
Pros^phingites globosus Kiparisova, 1947: 142, pi.
32, figs. 6, 7, text-fig. 29; Kiparisova and Krish-
tofovich, 1954: 21, pi. 12, figs. 2, 3; Kummel,
1961: 523; Kiparisova, 1961: 108, pi. 25, figs.
1, 2, text-figs. 69, 70.
Prosphingites aff. globosus Kiparisova, 1961: 109,
pi. 25, fig. 3, text-fig. 71.
A species based on two specimens, one
of which is a very small juvenile. Main
features are a very depressed whorl section
with an involute conch and showing the
beginnings of eccentricity of the umbilicus.
The suture (Fig. 16D, E) is characterized
Figure 16. Diagrammatic representation of the sutures of several species of Prosphingites. Sutures A-G are of species from
the upper Scythian Subco/umbifes Zone, sutures l-P are of species from the mid-Scythian Owenites Zone. A, P. coombsi
Kummel, paratype |OU3861), at a diameter of 21 mm, from near Kaka Point, New Zealand; B, P. coombsi Kummel, para-
type (OU3863), at a diameter of 20 mm; C, P. insularis Kiparisova (1961: fig. 74), at a whorl height of 9 mm from Sub-
columbites fauna, Primorye Region; D, P. globosus Kiparisova (1961: fig. 69), at a whorl height of 3 mm, from Sub-
columbites fauna, Primorye Region; E, P. globosus Kiparisova (1961: fig. 70), holotype, at a whorl height of 7 mm, from
Subco/umbites fauna, Primorye Region; F, P. ali Arthaber (1911: pi. 22, fig. 6c), from Subco/umbifes beds Kciro, Albania;
G, P. cze/<anows(cii Mojsisovics (1886: pi. 15, fig. lie), from Olenekites fauna, Olenek River, Siberia; H, P. cze/canows/t//
Mojsisovics, topotype (MCZ 8677), at a diameter of 31 mm; I, P. austini Hyatt and Smith (1905: pi. 7, fig. 4), from
A4ee/(oceras fauna. Union Wash, Inyo Range, California (USNM 75256), at a diameter of 20 mm; J, P. slossi Kummel and
Steele, paratype, at a diameter of 18 mm from Mee/toceras fauna, Crittenden Spring, Nevada; K, P. slossi Kummel and
Steele (1961: text-fig. 15a), at a diameter of 30 mm, from Mee/toceras fauna, Crittenden Spring, Nevada; L, P. spathi
Frebold, — Tozer (1961: pi. 13, fig. Ic), at a diameter of 16 mm, from Mee/coceros fauna, Blind Fiord Formation, Elles-
mere Island; M, P. ova/is Kiparisova (1961: fig. 79), at a whorl height of 5 mm, from Prosphingites Zone, Primorye Re-
gion; N, P. involutus Chao (1959: pi. 28, fig. 9), from Owenites lone, Kwangsi, China; O. P. kwangsianus Chao (1959:
text-fig. 39d), from Owenites fauna, Kwangsi, China.
408 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
by slightly phylloid saddles and a well de-
veloped auxiliary lobe.
It is inescapable that perhaps both of
the specimens of this species are juvenile
fonns. Until more material becomes avail-
able an assessment of this species is most
difficult. The whorl section is more de-
pressed than in any other species of Pro-
sphingites recorded to date, and it is this
feature which can be looked upon as the
most distinguishing for the species.
Occurrence. Subcohimbitcs fauna, Rus-
sian Island, Cape Zhitkov, Primorye Region,
eastern Siberia.
Prosphingifes insula ris Kiparisova
Text-figure 16
Prosphingifes imiilaris Kiparisova, 1961: 112, pi.
24, figs. 2-4, 6, text-figs. 74-76.
Prospliingites aff. insukiris Kiparisova, 1961: 114,
pi. 24, fig. 5. te.xt-fig. 77.
Prospliingites magnuml)ilicatus Kiparisova, 1961:
114, pi. 25, fig. 4, te.xt-fig. 78.
A more evolute prosphingitid with de-
pressed rounded whorls and a broad deep
umbilicus. Its most characteristic feature
is the suture with well denticulated lobes
and phylloid saddles (Fig. 16C). In its
external conch features this species is nearly
identical with P. coomhsi Kummel (1965);
it is only in the dorsal suture that differ-
ences can readily be seen. This species has
adjacent to the dorsal lobe two lateral lobes
whereas in P. coomhsi there are four. It
differs both in conch shape and suture
from P. czekanoicskii. It differs from P.
glohosus mainly in conch shape and in-
volution.
Occurrence. Suhcolumhitcs fauna, Pri-
morye Region, Siberia.
Prosphingifes coombs/ Kummel
Text-figure 16
Prosphingitc's cooinl)si Kuniinel, 1965: 538, figs.
1-5.
This species, in its general conch archi-
tecture, is nearly identical to P. insukiris
Kiparisova. It differs mainly in a more
elaborate dorsal suture (Fig. 16A, B).
Occurrence. This species was established
for specimens encountered in an isolated
pocket within disturbed beds between Kaka
Point and Nugget Point, south Otago, New
Zealand. On the basis of the general
morphology of the species, I have (Kum-
mel, 1965) interpreted this horizon as late
Scythian in age comparable to the horizon
of Subcolumbites of the Tethyan and
circum-Pacific region.
Repository. Department of Geology,
Otago University, New Zealand. Four
paratypes are in the Museum of Compara-
tive Zoology (MCZ 10113).
Genus Vickohlerifes Kummel, 1968
Type species, Prenkites sundaicus Welter,
1922
Vickohlerifes sundaicus (Welter)
Prenkites sundaicus Welter, 1922: 150, pi. 168(4),
figs. 18-21; Kutassy, 1933: 621; C. Renz, 1945:
301; Renz and Renz, 1947: 60; Renz and Renz,
1948: 29, pi. 12, fig. 1; Chao, 1959: 306.
''Prenkites" sundaicus, — Spath, 1930: 77; Spatli,
1934: 188, 209.
Vickohlerites sundaicus; — Kununel, 1968a: 9, pi.
1, figs. 10, 11.
Welter (1922) described this species on
the basis of a single specimen from Noel
Niti, Timor, and was quite positive as to
the close relationship of his specimen to
Prenkites malsorensis Arthaber from the
Subcoluuibites fauna of Albania. This close
relationship is difficult to see. Prenkites
is a more involute form, with depressed
whorls \\'hich contract on the adoral quarter
volution. The umbilical shoulders are sub-
angular and bear fine nodes. The Timor
specimen has a diameter of 40.7 mm, an
adoral whorl width of 21.5 mm, a height
of the adoral \\'horl of 13.2 mm and an
umbilical diameter of 20.3 mm. The conch
is evolute, the umbilicus comprising ap-
proximately 50 percent of the conch diam-
eter. The \\'h()rls are depressed with the
maximum width at the umbilical shoulder.
The venter is arched and grades with no
perceptible ventral shoulders to the um-
bilical shoulder which is acutely rounded.
The ninbilical wall is steep but not vertical.
Ammonoids of the Late Scythian (Lower Triassic) • Ktimmel
409
The last half volution of the specimens is
body chamber and shows traces of delicate
growth lines. The penultimate half volu-
tion bears a series of weak, fonvard pro-
jecting ridges which are most prominent
on the center part of the venter and disap-
pear completely midway between the venter
and the umbiHcal shoulder.
It is in Welter's (1922, pi. 14, fig. 21)
representation of the suture that one can
pinpoint the uncertainty in interpretation
of this form. Welter's drawing of the suture
covers only the portion from the venter
to the uml^ilical shoulder, but the drawing
implies that it was a complete suture. A
new drawing of the suture is given in
Kummel ( 1968, fig. 1 J ) . There are two
prominent lateral lobes but the umbilical
wall bears a good portion of a fairly large
auxiliary saddle and a small but very
distinct denticulated lateral lobe. The first
author after Welter to comment on Prenkites
sundaicus was Spath (1930, p. 77) who
wrote, "'Prenkites' sundaicus Welter, in
whorl shape resembles Cohimbites, but in
suture line it is closer to Subcohimbifes. . . ."
In terms of Welter's representation of the
suture this statement is correct. In Co-
lumbites the second lateral lobe is gen-
erally very small, consisting of a single
prong, whereas in Subcohimbites the sec-
ond lateral lobe, though much smaller
than the first, is more highly developed in
terms of its breadth and pattern of dentic-
ulation. On the basis of suture alone,
Prenkites sundaicus can not be attached to
either Cohimbites or Subcolumbites.
There is a general similarity in conch
shape of Prenkites sundaicus with some
groups of Subcolumbites. Among the sub-
columbitids three distinct groups can be
recognized. There is first of all the per-
rinismithi group with a tendency for carina-
tion of the venter; secondly the dusmani
group with a more marked development
of the reticulate ornamentation, a com-
pressed whorl section, but lacking the
tendency toward carination; finally there
is the robustus-multiformis group charac-
terized mainly by their depressed whorl
section. It is to this last group that Vickoh-
Jerites sundaicus has great resemblance
in conch fonn. Within the two subfamiHes
of the Paranannitidae those genera assigned
to the Columbitinae tend to have sutures
lacking an auxiliary lobe, whereas within
the Paranannitinae an auxiliary lobe is
commonplace, as in Prosphingites, Zenoites,
Chiotites, etc. It is within this subfamily
that Vickohlerites sundaicus belongs.
Renz and Renz (1948: 24, pi. 12, fig. 1)
have described and illustrated a single
specimen from the Subcolumbites fauna of
Chios as a representative of this species.
The general conch form of their specimen
is the same as that of the type specimens
from Timor. This Chios specimen mea-
sures 55.5 mm in diameter, 21.4 mm for
the width of the adoral whorl, 19 mm for
the height, and 23.5 mm for the width of
the umbilicus. The dimensions of the whorl
height and umbilical diameter in the two
specimens are reasonably similar. How-
ever, the Timor specimen has a broader
whorl than the Chios specimen by approxi-
mately 14 percent. This difference in whorl
width is difficult to evaluate, as each lo-
cality has yielded only a single specimen.
In addition, the Chios specimen, apparently,
does not show the suture. The overall simi-
larity of the Chios specimen to that from
Timor is such that, in spite of the differ-
ences and lack of data mentioned above,
the two specimens should be considered as
conspecific.
The new specimen recorded by Kummel
(1968) as Vickohlerites cf. sundaicus from
a Subcolumbites fauna at Kotal-e-Tera,
Afghanistan, is clearly congeneric with the
Timor and Chios specimens discussed
above but is not conspecific. Critical com-
parison of the Afghan and Timor forms
is difficult, as each is represented by a
single specimen. There is an overall simi-
larity between the two specimens, but at
the same time there are intriguing differ-
ences in whorl shape and suture.
Occurrence. The holotype is from Noel
410 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Niti, Timor. Welter (1922: 85, 86) con-
sidered this specimen to come from his
lowest Triassic horizon. The forms as-
sociated with tliis specimen he lists on
page 150 as Meekoceros sp. indet. The
available evidence does not allow any
precise determination of the age, bnt the
biological affinities of this species suggest
that it is late Scythian in age. The speci-
men recorded by Renz and Renz (1948)
is from the Suhcolumhitcs fauna of Chios.
Repository. The holotype is in the
Paleontological Institute of Bonn Univer-
sity; the specimen from Chios is in the Nat-
ural History Museum, Basel J13576.
Genus Zenoiies Renz and Renz, 1947
Type species, Prosphingites (Zenoites)
helenoe Renz and Renz, 1947
Generally small, somewhat evolute form
with rounded, depressed whorl sections
bearing irregularly spaced radial to pro-
siradiate constrictions that encircle the
wliorl section; suture consists of two dentic-
ulated lateral lobes, one auxiliary lobe on
the umljilical shoulder and the appearance
of another lobe on the umbilical seam.
The suture and general shape of the
conch indicate a relationship to Pros})]iing.-
ites. This genus was first recognized from
the Suhcolu?n1)ites fauna of Chios by Renz
and Renz (1948). Since then it has been
recognized from a late Scythian horizon on
Ellesmere Island (Tozer, 1965a).
Zeno(7es helenae Renz and Renz
Text-figure 17
Fros])hingHcs (Zcnoilcs) hclcnac Renz and Renz,
1947: 60, 75; Renz and Renz, 1948: 41, pi.
15, fig.s. 8-8a, pi. 16, figs. 1-lc.
Prospliiufiitcs (Zenoites) helenae var. maradocun-
ensis Renz and Renz, 1947: 60, 75; Renz and
Renz, 1948: 42. pi. 15, fig.s. 12-12a, 14-14a.
Zenoites helenae, — Kummel, in Arkcll et al., 1957:
L139, figs. 172, 4.
Prosphingites ex aff. czekanowskii,- — Renz and
Renz, 1948: .39, pi. 15, figs. 11 -lib.
The variety estabHshed for this species
{maradovAinensis) consists merely of two
slightly more compressed forms. The mea-
surements of the figured specimens are as
follows :
D
W H U W/D H/D U/D
1.
24.0
13.4
8.7
9.4
55.8
36.3
39.2
2.
17.4
?
5.3
6.5
p
30.5
37.4
3.
21.6
10.2
7.2
9.3
47.2
33.3
43.1
4.
17.7
9.0
6.4
6.1
50.8
36.2
34.5
1. Holotype, NHMB J13648, Renz and Renz,
1948: pi. 16, fig. 1.
2. Paratvpe, NHMB T13649, Renz and Renz,
1948: pi. 15, fig. 8.
3. var. maradovunensis, NHMB J 13652, Renz and
Renz, 1948: pi. 15, fig. 14.
4. var. maradovunensis, NHMB J13651, Renz and
Renz, 1948: pi. 15, fig. 12.
This species is not common in the Chios
fauna; Renz and Renz record only five
specimens, and there are 9 additional speci-
mens in the Natural History Museum,
Basel. The suture is shown on Figure 17 A.
Occurrence. Suhcolumhitcs fauna, Mara-
dovuno, Chios.
Repository. Holotype, NHMB J13648;
figured paratypes NHMB J13649, J13651,
J13652; unfigured paratypes NHMB J13650.
Zenoites vonderschmitii (Renz and Renz)
Text-figure 17
Prosphingites voiidersehniitti Renz and Renz, 1947:
60, 74; Renz and Renz, 1948: 39, pi. 15, figs.
13, 15.
This is a highly compressed form in com-
parison to the genotype helenae. The holo-
type (Renz and Renz, 1948: pi. 15, fig.
13) measures 24.6 mm in diameter, 9.5
mm for the width of the adoral whorl, 7.3
for the height, and 10.8 mm for the diam-
eter of the uml)ilicus. The paratype (Renz
and Renz, 1948: pi. 15, fig. 15) measures
15.7 mm in diameter, 5.2 mm for the height
of the adoral whorl and 7.1 mm for the
diameter of the mnbilicus ( faulty preser-
vation pre\'ents obtaining a whorl width
measurement). Aside from the very dif-
ferent whorl dimensions, this species has a
different pattern of constriction. The su-
ture, however, (Fig. 17B) is almost identical
in these two Chios species.
Occurrence. Suhcolumhitcs fauna, (Jliios.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 411
A
C F
Figure 17. Diagrammatic representation of the suture of: A, holotype Zenoites helenae Renz and Renz (1948: pi. 16,
fig. Ic), at a diameter of approximately 18 mm; B, fiolotype of Prosphingites vonderschrnitti Renz and Renz (1948: pi. 15,
fig. 13c), at a diameter of approximately 18 mm; C, paratype Columbites huangi Chao (1959: pi. 29, fig. 11), at a
diameter of approximately 20 mm; D, paratype Chiotites g/obu/oris Renz and Renz (1948: pi. 15, fig. 9c], at a diameter
of approximately 15 mm; E, paratype Popov/fes occidenta//s Tozer (1965: fig. 5b), at a diameter of approximately 30 mm;
F, paratype Monaconffi/fes monoceras Tozer (1965: fig. 8), at a diameter of approximately 15 mm.
Specimens of figures A, B, C from Subco/umbites fauna of Cfiios; D, E from Toad Formation, British Columbia; F, from
upper Scythian of Ellesmere Island.
Repository. Holotype NHMB J13639
(Renz and Renz, 1948: pi. 15, fig. 13);
paratype NHMB J13640 (Renz and Renz,
1948: pi. 15, fig. 15); unfigured paratypes
NHMB J13641.
Zenoites arcticus Tozer
Zenoites arcticus Tozer, 1965ii: 25, pi. 2, figs.
6, 7, te.xt-fig. 7.
Only two specimens of this species are
known. It differs from Z. helenae mainly
in the pattern of constriction.
Occurrence. Blaa Mountain Formation,
lower shale member, Ellesmere Island.
Genus isculitoides Spath, 1930
Type species, Isculiies originis Arthaber,
1911
This is another of the genera which are
confined to the late Scythian Prohungarites
Zone, where it is represented by six species.
In the Suhcohimhites fauna of Albania and
Chios, /. originis is one of the most common
species. This species in the Chios fauna
shows a very large degree of intraspe-
cific variation. Isculitoides eUipticus from
Kwangsi, China, I. minor from British Co-
lumbia, and /. suhoviformis from the Pri-
morye Region are represented by very few
specimens. Additional collections of these
three species are badly needed to clarify
their relationships. In the western United
States we have I. wasserbergi and I. ham-
mondi, both species known from a fair
number of specimens. Specimens assigned
to Isculitoides cf. originis have been re-
corded from the Suhcohimhites fauna at
Kotal-e-Tera, Afghanistan (Kummel, 1968b)
and specimens of indeterminant specific
identity from the Prohungarites fauna of
the Salt Range, West Pakistan.
412 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Ammonoids of the Late Scythian (Lower Triassic) • Kummcl 413
Isculitoides originis (Arthaber)
Plate 5, figures 1-10; Plate 6, figures
1-6; Text-figures 18-20
Isculites originis Artliaber, 1911: 259, pi. 23(7),
figs. 1-10; Diener, 1915: 157; C. Renz, 1928:
155; Kutassy, 1933: 540; Renz and Renz, 1947:
60; Renz and Renz, 1948: 33, pi. 13, figs.
7-7a, 9-9a, 11-1 lb, 12-12b, pi. 14, figs. 6-6a,
9-9a.
Isculitoides originis, — Spath, 1934: 198, pi. 14,
figs. 2a-d, te.\t-fig. 59b, c.
Iscuhites globulus Renz and Renz, 1947: 60, 74;
Renz and Renz, 1948: 34, pi. 14, figs. 10-lOc,
4-4a, 5-5a, 8-8b, 11-1 lb.
Iscuhites antiglohulus Renz and Renz, 1947: 60,
74; Renz and Renz, 1948: 35, pi. 13, figs. 1-la,
10-lOa, pi. 13, figs. 2-2a, 3-3a, 5-5a, 8-8a.
Iscuhites glohuhis-originis Renz and Renz, 1947:
60; Renz and Renz, 1948: 35, pi. 13, figs. 6-6a,
pi. 14, figs. 1-la, 2-2a, 3-3a.
Iscuhites glohuhis-antiglohuhis Renz and Renz,
1947: 60; Renz and Renz, 1948: 35, pi. 13, figs.
4-4a, pi. 14, figs. 7-7a.
Arthaber (1911: 259) stated he had 54
.specimens of this species, 10 of which he
illustrated. The type specimen and seven
paratypes, all among those illustrated by
Arthaber (1911), are still preserved. The
preservation of all these specimens leaves
much to be desired and most are too badly
preserved or incomplete to yield signifi-
cant measurements. Arthaber's (1911: pi.
23(7), figs. 1-10) illustrations are highly re-
touched and idealized; unre touched photo-
graphs of the existing original specimens
are reproduced here on Plates 5 and 6.
One of the most common elements in
the Stihcolinnbites fauna of Chios is this
species. Renz and Renz ( 1948 ) had avail-
able for study several hundred specimens.
Of this vast number of specimens 126 are
sufficiently complete and well preserved to
yield measurements of the basic conch
dimensions. These data are tabulated on
Table 31, and plotted on Figures 19 and
20. Renz and Renz ( 1948 ) recognized five
species of this genus within the Chios fauna.
These are: originis, globulus, antiglohulus,
glohulus-originis, and gl oh ul us- a nti globulus.
The latter two species were named for
intermediate forms between the first three
species. These "Formenreihen" are distin-
guished basically on conch thickness and
size of the umbilicus. Isculitoides glohulus,
considered by Renz and Renz as the point
of origin of their "FoiTnenreihen," is a
globular form with a relatively small um-
bilicus. They recognized one series of
transitional forms to Isculitoides anti-
glohulus, which has a more compressed
conch and a more open umbilicus. Another
series extended from Isculitoides glohulus
to I. originis, which is a compressed form
with no appreciable difference in the
diameter of the umbilicus. A plot of the
whorl width (Fig. 19) and the umbilical
diameter (Fig. 20) of these species recog-
nized bv Renz and Renz demonstrates the
Figure 18. Diagrammatic representation of the sutures of various species of /scu/ifo/des. A-l, Isculitoides originis (Artfia-
ber), A-E from Subco/umbites fauna, Albania, F-l from Subco/umbifes fauna, Chios. A, holotype (Arthaber, 1911: pi.
23(7), fig. Ic), no sutures ore visible on the type specimen; B, paratype (Arthaber, 1911: pi. 23(7), fig. 9), redrawn from
type specimen: C, paratype (Arthaber, 1911: pi. 23(7), fig. 6c), no sutures are visible on the type specimen; D, paratype
(Arthaber, 1911: pi. 23(7), fig. 8), specimen apparently lost; E, paratype (Arthaber, 1911: pi. 23(7), fig. 9), specimen ap-
parently lost; F, plesiotype (Renz and Renz, 1948: pi. 13, fig. 12b, NHMB J-13590), at a diameter of approximately 35 mm;
G, plesiotype (Renz and Renz, 1948: pi. 13, fig. lib, NHMB J-1 3591 ), at a diameter of approximately 35 mm; H, para-
type Iscuhites globulus Renz and Renz (1948: pi. 14, fig. 8b, NHMB J-13602), at a diameter of approximately 25 mm;
I, paratype Iscuhites globulus Renz and Renz (1948: pi. 14, fig. lib, NHMB J-13600), at a diameter of approximately 25
mm; J-L, /. ellipticus Chao (1959), from Subco(umb/tes fauna, Kwangsi, China; J, holotype (Chao, 1959: fig. 38a), at
a whorl height of 7 mm; K, paratype (Chao, 1959: fig. 38b), at a whorl height of 5 mm; L, paratype (Chao, 1959: pi.
30, fig. 3), at a diameter of approximately 25 mm; M, /. subov/formis (Kiparisova, 1961: fig. 97], from Subco/umb/fes
fauna, Primorye Region, at a diameter of approximately 15 mm; N-P, /. wasserbergi n. sp. from Subcolumbifes fauna,
Tobin Formation, Nevada, N, paratype, at a diameter of 8.5 mm (MCZ 9626); O, paratype, at a diameter of 8 mm (MCZ
9450, PI. 29, figs. 9, 10); P, paratype, at a diameter of 8 mm (MCZ 9657); Q, paratype /. hammondi n. sp., from Upper
Thaynes Formation, Hammond Creek, southeast Idaho, at o diameter of 13 mm (MCZ 9480).
414 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
20
18
16
14
12
10
8
6
w
.0 0
0 +,
0 0 0 +
I I I I 1 I I I I I L^
Ill
I I I I 1 \ \ I L
0
15
20
25 30
DIAMETER
35
40
45
50
Figure 19. Variation in whorl width (W) of Isculitoides originis (Arthaber), from the Subcolumbifes fauna of Chios. The
data on this graph are from Table 31.
completely tran.sitional nature of these two
characters.
■ Renz and Renz (1948: 33) recognized
that the basic plan of the suture was the
same in the Chios and Albanian popula-
tions (Fig. 18 A-I). What variations do
occur are minor changes in the shape of
the elements and in the denticulation of
the lobes.
Among late Scythian ammonite faunas,
species of Isculitoides are recognized from
Timor, Kwangsi, Primorye Region, British
Columbia, Nevada, and southeastern
Idaho. The Timor forms belong in /.
on<iinis (Spath, 1934: 198). None of the
remaining species are known by more than
a few specimens, which makes comparison
to /. oriii^inis difficult. Isculitoides suhovi-
formis Kiparisova from the Primorye Re-
gion and 7. icasscrhcr<ii n. sp. from Nevada
are extremely depressed forms; /. cllipticus
Chao from Kwangsi and 7. hammondi n.
sp. are similar in conch shape to 7. or0nis
but arc much more involute, not showing
as strong an exccntruinbilication ol the
umbilicus.
Occurrence. SiiheohiinhUes fauna of
Albania and C'hios, and from the man-
16r
14-
12
10
8
6
4
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 4L5
U
• o
0
A
+
+
0
0
0
I I I I I I
I I I I I I \ LJ 1 LJ I L
10
15
20
25 30 35
DIAMETER
I I I I I I I I I I i I L
40
45
50
Figure 20. Variation in umbilical diameter (U) of hculitoides originis (Arthaber), from Subco/umb/'/es fauna of Chiios.
Tfie data on this graph are from Table 31 .
ganese coated fauna with ProJiungaiites of
Nifoekoko and Toeboelopo, Timor.
Repository. The genotype and 7 para-
types are preserved in the Paleontological
Institute, University of Vienna. Tliese are
the specimens of Arthaber's plate 23(7),
figs. 1-7, 10.
The specimens of this species from
Chios described by Renz and Renz ( 1948 )
are in the Natural History Museum, Basel;
these are as follows: plesiotypes, I. orig-
inis (pi. 13, fig. 7) NHMB J13584, (pi. 13,
fig. 9) NHMB J13585, (pk 13, fig. 11a)
NHMB J13586, (pk 13, fig. lib) NHMB
J13591, (pk 13, fig. 12a) NHMB J13587, (pk
13, fig. 12b) NHMB J13590, (pk 14, fig. 6)
NHMB J13588, (pk 14, fig. 9) NHMB
J13589; unfigured specimens of /. originis
from Maradovuno NHMB J13592, J13593,
J13594, from Kephalovuno NHMB J13595;
holotype, /. globulus (pk 14, fig. 10a, b)
J13596, (pk 14, fig. 10c) J13601; paratypes.
416 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Table 31. Measurements of Iscuutoides originis (Arthaber) from the Subcolumbites fauna
OF Chios.
The species names on this hst are those recognized by Renz and Renz (1948). The measurements
are chstinguished on Figure 19 and Figure 20.
D
w
H
u
W/D
H/D
U/D
D
w
H
U
W/D
H/D
U/D
1.
47.2
15.5
20.0
10.2
32.8
42.4
21.6
33.
33.8
16.5±
14.8
7.5
48.8i
43.8
22.2
2_
45.7
16.1
17.8
11.1
35.2
38.9
24.3
34.
31.2
19.8
13.0
6.7
63.5
41.7
21.5
3.
45.0
14.6
19.2
10.7
32.4
42.7
23.8
35.
29.9
19.8
13.3
7.5
66.2
44.5
25.1
4.
44.4
?
19.6
9.3
?
44.1
20.9
36.
27.8
16.6
11.5
7.0
59.7
41.4
25.2
5.
41.7
13.8
16.3
9.6
33.1
39.1
23.0
37.
25.0
15.2
9.3
7.2
60.8
37.2
28.8
6.
39.8
14.6
18.1
7.0
36.7
45.5
17.6
38.
24.2
13.8?
11.6
4.5
57.0?
47.9
18.6
7.
39.8
14.3
15.2
11.0
35.9
38.2
27.6
39.
23.8
15.0
10.5
7.4
63.0
44.1
31.1
8.
37.0
?
14.0
10.2
p
37.8
27.6
40.
22.7
16.7
11.5
3.8
73.6
50.7
16.7
9.
35.6
12.7
13.9
10.4
35.7
39.0
29.2
41.
17.1
11.7
8.0
1.8
68.4
46.8
10.5
10.
35.0
11.7
13.5
9.0
33.4
38.6
25.7
42.
17.0
11.1
7.4
2.8
65.3
43.5
65.0
11.
34.2
11.5
14.8
7.5
33.6
43.3
21.9
43.
15.5
10.7
7.2
3.2
69.0
46.5
20.6
12.
33.8
12.5
14.2
7.0
37.0
42.0
20.7
44.
15.4
9.3
8.8
2.9
60.4
57.1
18.8
13.
33.6
12.1
14.2
6.8
36.0
42.3
20.2
45.
14.6
9.7
6.4
3.8
66.4
43.8
26.0
14.
31.5
11.2
12.8
7.8
35.6
40.6
24.8
46.
14.5
9.8
6.6
2.4?
67.6
45.5
16.6?
15.
31.4
12.1
13.2
8.2
38.5
42.0
26.1
47.
13.8
10.0
5.6
3.6
72.5
40.6
26.1
16.
31.3
11.8
12.3
7.4
37.7
39.3
23.6
48.
13.8
8.3
6.2
2.4
60.1
44.9
17.4
17.
30.7
11.7
12.1
7.0
38.1
39.4
22.8
49.
13.8
9.8
5.0
4.0
71.0
36.2
29.0
18.
29.3
11.2
12.0
6.7
38.2
41.0
22.9
Isculitoidcs
antiglobuJus
19.
29.3
10.2
12.0
6.0
34.8
41.0
20.5
20.
29.3
10.6
13.2
5.4
36.2
45.1
18.4
50.
46.3
13.7
16.4
15.0
29.6
35.4
32.4
21.
29.0
10.7
11.1
6.7
36.9
38.3
23.1
51.
45.5
12.6
16.8
12.8
27.7
36.9
28.1
22.
29.0
11.8
12.3
6.7
40.7
42.4
23.1
52.
43.3
11.9
16.6
10.8
27.5
38.3
24.9
23.
27.6
10.5
11.5
6.4
38.0
41.7
23.2
53.
43.2
14.3
16.1
12.5
33.1
37.3
28.9
24.
26.0
10.3
11.0
5.8
39.6
42.3
22.3
54.
39.5
?
14.0
13.6
p
35.4
34.4
25.
25.7
9.8
10.1
5.6
38.1
39.3
21.8
55.
36.0
12.2
11.9
12.1
33.9
33.1
33.6
26.
23.1
9.8
9.0
4.5
42.4
40.0
19.5
56.
35.3
11.4
13.7
9.4
32.3
38.8
26.6
27.
20.2
7.9
8.3
4.0
39.1
41.1
19.8
57.
35.2
12.1
14.7
9.4
34.4
41.8
26.7
28.
19.8
8.0
8.2
5.0
40.4
41.4
25.3
58.
35.0
13.2
12.5
12.5
37.7
35.7
35.7
29.
19.7
7.3
8.7
4.0
.37.1
44.2
20.3
59.
34.1
11.0
13.4
7.9
32.3
39.3
23.2
30.
16.0
6.6
7.0
2.0
41.3
43.8
12.5
60.
33.6
12.3
13.8
7.4
36.6
41.1
22.0
61.
32.7
10.3
13.3
7.9
31.5
40.7
24.2
Isci
ilitoide
s' p.Job
tiJu.s
62.
32.7
11.1?
11.4
11.4
33.9?
34.9
32.9
31.
37.8
15.0^
■ 17.3
6.7
39.7^
= 45.8
17.7
63.
31.1
10.0
12.2
8.0
32.2
39.2
25.7
32.
35.2
17.8
12.6
10.0
50.6
35.8
28.4
(>4.
31.0
10.7
12.2
6.5
34.5
39.4
31.0
1.
2.
4.
5.
6.
11.
13.
26.
32.
33.
35.
37.
3.S.
-10.
Plesiotype, Renz and Renz (1948: pi. 13, fig. 12), NHMB .T13587.
Plesiotype, Renz and Renz (1948: pi. 13, fig. 11), NHMB J135S6.
Plesiotype, Renz and Renz (1948: pi. 13, fig. lib), NHMB .113591.
Plesiotype, Renz and Renz (1948: pi. 13, fig. 9), NHMB jl35S5.
Plesiotype, Renz and Renz (1948: pi. 13, fig. 12b), NHMB .113587.
Plesiotype, Renz and Renz (1948: pi. 13, fig. 7), NHMB .113584.
Plesiotype, Renz and Renz (1948: pi. 14, fig. 9), NHMB J135S9.
Plesiotype, Renz and Renz (1948: pi. 14, fig. 6), NHMB J13588.
30, unfignred parat>pes from Maradovimo,
(1948: pi. 14, fig. 8b), NHMB 113602.
(1948: pi
(1948: pi.
(1948: pi.
(1948: pi.
(1948: pi.
(1948: 1
Remaining specimens from .3
•31. Parat>pe, Renz and Renz
Renz
Renz
Renz
Renz
Renz
Renz
Chios, NHMB .T13592, 13593, 13594.
Paratype,
Parat>pe,
lIolot\ pe,
Paratyjie,
Parat\pe,
Paratype,
and
and
and
and
and
and
Renz
Renz
Renz
Renz
Renz
Renz
14,
14,
14,
14,
II,
)1. It, f
fig. 11), NHMB 113600.
fig. 8), NHMB 113599.
fig. 10), NHMB T13596.
fig. 4), NHMB T 13597.
fig. 5), NHMB 113598.
ig. 10c), NHMB 113596.
Remaining specimens from 34—49, unfignred paratypes from Marado\imo, Chios, XI 1MB J13603.
.50. Paratype, Renz and Renz (1948: pi. 13, fig. 1), NHMB .113606.
53. Paratvpe, Renz and Renz (1948: pi. 13, fig. 2), NHMB 113607.
54. Paratype, Renz and Renz (1948: pi. 13, fig. 3). NHMB 113608.
56. Paratvpe, Renz and Renz (1948: pi. 13, fig. 8), NHMB 113610.
58. Paratvpe, Renz and Renz (1948: pi. 13, fig. 5), NHMB 113609.
62. llolotype, Renz and Renz (1948: pi. 13, fig. 10), NUMB .113605.
Maradovimo,
Remaininu specimens from 51-72, unfignred ])aratypes frc
Chios, NHMB 11361'
Ammonoids of the Late Scythian (Lower Triassic) • Kttmmel 417
Table 31. Contintied
D
w
H
u
W/D
H/D
U/D
D
W
H
u
W/D
H/D
U/D
65.
31.0
10.3
12.2
7.3
33.2
39.4
23.5
96.
12.9
6.8
6.8
2.0?
52.7
52.7
15.5?
66.
28.0
9.6
11.0
6.6
34.3
39.3
23.6
97.
10.8
6.7
5.5?
l.,3?
67.
68.
69.
70.
71.
72,
28.0
26.8
26.6
24.8
20.4
18.4
9.7
9.3
9.4
9.1
7.8
7.3
11.2
11.0
12.2
9.1
8.3
9.3
6.6
7.1
5.4
7.5
4.4
1.5?
34.6
34.7
35.3
36.7
38.2
39.7
40.0
41.0
45.9
36.7
40.7
50.5
23.6
26.5
20.3
30.2
21.6
8.2?
98.
99.
100.
101.
102.
Isc
41.5
.39.1
39.0
38.6
37.0
ulitoides glol
16.3 15.3
13.9 16.0
12.9 15.7
15.0 15.3
15.2 14.6
ndusantigloh
13.3 39.3
9.1 35.5
9.2 33.1
8.5 .38.9
10.3 41.1
ulus
36.9
40.9
40.3
39.6
39.5
32.0
23.3
23.6
22.0
27.8
J
Isculitoides glohulusoriginis
103.
35.5
13.1
14.5
9.0
36.9
40.8
25.4
73.
39.6
17.9
16.8
8.9
45.2
42.4
22.5
104.
.35.3
12.7
13.5
8.8
36.0
.38.2
24.9
74.
37.5
13.6
14.3
9.8
36.3
38.1
26.1
105.
.32.8
12.3
13.3
8.4
37.5
40.5
25.3
75.
37.2
13.7
13.1
10.8
.36.8
.35.2
29.0
106.
.32.7
13.2
14.0
7.0
40.4
42.8
21.4
76.
31.2
13.3?
13.2
8.1
42.6?
42.3
26.0
107.
31.7
11.5
12.6
8.5
36.3
39.7
26.8
77.
30.7
14.0
13.7
6.9
45.6
44.6
22.5
108.
31.3
12.3
12.7
8.0
39.3
40.6
25.6
78.
30.5
14.1
11.7
7.8
46.2
38.4
25.6
109.
31.2
11.8
13.7
6.2
.37.8
43.9
19.9
79.
30.4
13.8
12.5
6.1
45.4
41.1
20.1
110.
31.1
12.8
12.3
6.9
41.2
39.5
22.2
80.
29.7
12.1
11.4
7.8
40.7
38.4
26.3
111.
.30.3
12.3
12.3
7.1
40.6
40.6
23.4
81.
27.4
11.0?
12.4
5.5
40.1?
45.3
20.1
112.
30.3
14.2
11.7
10.0
46.9
38.6
33.0
82.
27.0
13.3
11.2
7.5
49.3
41.5
27.8
113.
29.3
14.3
11.5
7.8
48.8
.39.2
26.6
83.
27.0
12.0
11.1
6.2
44.4
41.1
23.0
114.
29.3
11.5
11.1
8.2
39.2
,37.9
28.0
84.
27.0
15.5
11.2
5.6
57.4
41.5
20.7
115.
29.2
10.8
13.0
5.2
37.0
44.5
17.8
85.
27.0
14.2
11.2
6.1
52.6
41.5
22.6
116.
29.0
13.0
12.4
5.7
44.8
42.8
19.7
86.
26.2
13.8
11.2
6.2
52.7
42.7
23.7
117.
28.8
12.7
9.8
8.4
44.1
34.0
29.2
87.
25.6
12.6
10.1
6.0
49.2
39.5
23.4
118.
28.4
10.2
10.9
6.7
35.9
38.4
23.6
88.
25.0
13.0?
11.6
4.0
52.0?
46.4
16.0
119.
26.4
11.7
11.6
5.2
44.3
43.9
19.7
89.
24.3
13.0
10.4
5.1
53.5
42.8
21.0
120.
25.4
11.9
11.0
6.1
46.9
43.3
24.0
90.
22.3
?
11.0
2.8
p
49.3
12.6
121.
25.0
12.2
11.4
5.5
48.8
45.6
22.0
91.
22.2
11.7
10.0
5.2
52.7
45.0
23.4
122.
24.5
10.4
10.3
5.8
42.4
42.0
23.7
92.
20.4
12.3
9.2
4.5
60.3
45.1
22.1
123.
24.2
10.7
10.2
4.3
44.2
42.1
17.8
93.
18.3
10.9
7.2
4.6
59.6
39.3
25.1
124.
23.0
8.5
10.3
4.5
36.9
44.8
19.6
94.
15.6
7.3
7.6
2.2
46.8
48.7
14.1
125.
21.8
11.1
9.3
3.9
50.9
42.7
17.9
95.
15.5
7.7
7.6
1.8
49.7
49.0
11.6
126.
21.7
11.5
11.1
2.8
52.9
51.2
12.9
74. Plesioti'pe, Renz and Renz (1948: pi. 14, fig. 2), NHMB T1361.5.
75. Plesiot^'pe, Renz and Renz (1948: pi. 14, fig. 1), NHMB J13614.
76. Plesiotype, Renz and Renz (1948: pi. 14, fig. 3), NHMB J13616.
Remaining specimens from 73—97: unfigured paratvpes from Maradovuno, Chios, NHMB J13617.
98. Plesiotvpe, Renz and Renz (1948: pi. 13, fig. 4), NHMB J13619.
112. Plesiotype, Renz and Renz (1948: pi. 14, fig. 7), NHMB J13620.
Remaining specimens from 99—126 unfigured paratypes from Maradovuno, Chios, NHMB J13621.
(pl. 14, fig. 4) J13597, (pi. 14, fig. 5)
J13598, (pl. 14, fig. 8a) NHMB J13599,
(pi. 14, fig. 8b) NHMB J13602, (pl. 14,
fig. 11) NHMB J13600; unfigured para-
types from Maradovuno NHMB J 13603,
from Kephalovuno NHMB J13604; holo-
type, 1. antiglohulus (pl. 13, fig. 10) NHMB
J13605; paratypes (pl. 13, fig. 1) NHMB
J13606, (pl. 13, fig. 2) NHMB J13607, (pl.
13, fig. 3) NHMB J13608, (pl. 13, fig. 5)
NHMB J 13609, (pl. 13, fig. 8) NHMB
J13610; unfigured paratypes from Mara-
dovuno NHMB J13612, from Kephalovuno
NHMB J 13611; svn types of 7. glohulus-
originis, (pl. 13, fig. 6) NHMB J13613,
(pl. 14, fig. 1) NHMB J13614, (pl. 14,
fig. 2) NHMB J13615, (pl. 14, fig. 3)
NHMB J13616; unfigured specimens from
Maradovuno NHMB J13617, from Kep-
halovuno NHMB J13618; syntypes of 7.
glohidus-antiglobulus, (pl. 13, fig. 4)
NHMB J 13619, (pl. 14, fig. 7) NHMB
J 13620; unfigured specimens from Mara-
dovuno NHMB J13621, from Kephalovuno
NHMB J13622;" topotypes from Chios
MCZ 10019, 10020. The specimens from
418 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Timor are in the British Museum (Natural
History ) .
Isculifoides ellipficus Chao
Text-figure 18
Isciilitoides ellipticus Chao, 1959: 119, 29.3, pi.
26, figs. 24-28, pi. 30, figs. 1-5, text-flu. 38a, h.
Isculifoides aff. ori^inis (Arthaber), Chao, 1959:
118, 292, pi. 26, figs. 22, 23.
Chao had one specimen of his /. aff.
originis and four specimens assigned to
I. ellipticus; for none of the specimens are
any measurements given. All the speci-
mens are compressed with rounded venters.
The specimen assigned to /. aff. originis
is slightly more evolute than I. ellipticus.
We have already seen from the abundant
material of Z. originis from Chios that
there can be a large amount of intraspecific
variation. These Kwangsi specimens
could well belong to /. originis or 7. ham-
moncli but the smallness of the sample does
not allow a critical comparison. It seems
best for the moment to recognize this
species with the reservation that it is very
close to 7. originis and could well be con-
specific. The sutures are illustrated on
Figure 18 J-L.
Occurrence. The four specimens of 7.
ellipticus came from a loose block of lime-
stone, stratigraphic position unknown, from
near the Lolou village, Linglo district,
Kwangsi, China (Chao collection 542b).
Associated forms include species of Fro-
earnites, Proptt/chifoides, etc., which indi-
cate a late Scythian Suhcolunibites age.
The single specimen of 7. aff. originis came
from the same place.
Isculitoides suboviformis (Kiparisova)
Text-figure 18
Paranannites suhovifonnis Kiparisova and Kiish-
tofovich, 1954: 21, pi. 11, figs. 4, 5; Kiparisova,
1961: 127, pi. 27, figs. 8, 9, text-figs. 94, 95.
PdrauannUcs aff. sulxniforniis Kiparisova, 1961:
128, pi. 27, figs. 10, 11, text-figs. 96, 97.
This species has an extremely depressed
whorl section as in /. wasserbergi and dif-
fers from that Nevada species mainly in
lacking any radial ornamentation. Kipari-
TA13LE 32. Measurements of Isculitoides
WASSERBERGI N. SP. FROM THE TOBIN FORMATION
OF Nevada.
D
w
H
u
W/D
H/D
U/D
1.
14.7
13.8
6.1
3.0
93.9
40.5
20.45
2.
14.1
13.8
6.5
1.8
97.9
46.1
13.8
3.
13.0
11.0
6.1
2.0
84.7
46.9
15.4
4.
13.0
11.4
6.2
2.0
87.6
47.7
15.4?
5.
8.5
8.9
3.5
1.0?
105.2
41.1
11.8?
6.
7.3
8.1
3.7?
■?
110.9
50.7?
?
7.
5.6
6.1
2.8
0.6?
108.3
50.0
10.7?
1. Holotvpe, MCZ 9447 (PI. 29, figs. 1-3).
2. Parat\'pe, MCZ 9448 (PI. 29, figs. 4-6).
.3. Paratype, MCZ 9449 (PI. 29, figs. 7, 8).
4, .5, 7. Unfigured paratvpes, MCZ 9485.
6. Paratype, MCZ 9450 (PI. 29, figs. 9, 10).
sova (1961: 127) had only six specimens
of this species; additional material may
establish that the Nevada species is con-
specific with this Siberian form. Tlie
suture is shown on Figure 18 M.
Occurrence. Suhcolunibites fauna, Cape
Zhitkov and Amur Bay, Primorye Region,
Siberia.
Isculitoides minor Tozer
Isculitoides minor Tozer, 1965a: 20, pi. 2, figs.
1-3.
This is an involute Isculitoides on the
pattern of 7. ellipticus Chao, 7. suboviformis
Kiparisova, and 7. hammondi n. sp. The
species was based on only three specimens
\\'hich makes comparisons difficult. It is
highly possible that 7. ellipticus, I. minor,
and 7. hammondi are conspecific, but for
the moment at least it appears best to
tentatively recognize all three species.
Occurrence. Toad Formation, Halfway
River area, British Colvmibia.
Isculifoides wasserbergi n. sp.
Plate 29, figures 1-10; Text-figure 18
Small, tightly involute, globular conchs
with highly depressed whorl sections, and
excentric umbilici on the ontcM' volutions.
There are 13 specimens in the collection,
of \\'hich 7 are sufficiently well preserved
to yield the measurements shown in
Table 32. The umbilical shoulder is well
rounded, sloping gradualK up to the very
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 419
l)ioadly arched venter and flanks. The
umbilical wall is steep. The conch of most
specimens is smooth, except for faint
growth lines. On some specimens, as that
of Plate 29, figures 1, 2, there are a few
narrow, low, rounded ridges extending
straight across the venter from one umbili-
cal shoulder to the other. The body
chamber is at least one volution in length.
The suture is simple, ceratitic with single
pronged ventral lobes, a denticulated first
lateral lobe lying midway between the
ventral lobe and the umbilical shoulder,
and a second lateral lobe on the umbilical
wall (Fig. 18 N-P).
Isculitoides orig,inis from the Subcolum-
bitcs faunas of Albania and Chios is ex-
tremely variable in conch dimensions, a
variability typical of many excentrumbili-
cate ammonites. IscuUtoides wasserhcrgi
tends to resemble the more involute and
globose variants, but still has a much more
depressed whorl section and is more in-
volute. Likewise, even though the Nevada
sample is small, there is a suggestion at
least that the amount of intraspecific varia-
tion is much less than that in 7. originis.
This new species is most similar to I.
suboviformis ( Kiparisova ) from the Siib-
cohimbites fauna of the Primorye Region.
In fact these two species have the most
depressed whorls of any known IscuUtoides.
Occurrence. Lower part of Tobin For-
mation, uses Locality M2562, Pershing
County, Nevada; south tip of Tobin
Range, Cain Mountain 1:62,500 quad.,
center NW % sec. 9, T. 26N, R. 39E, 5500
ft. S, 27.5 ft. W from elevation point 5088
on range crest.
Repositorij. Holotype, MCZ 9447 (PI.
29, figs. 1-3); paratypes, MCZ 9448 (PI.
29, figs. 4-6), MCZ 9449 (PI. 29, figs. 7, 8),
MCZ 9450 (PI. 29, figs. 9, 10); suture
specimens MCZ 9626, 9657; unfigured
paratypes, MCZ 9485.
IscuUtoides hammondi n. sp.
Plate 36, figures 8-13; Text-figure 18
proximately 40 specimens, most of which
are poorly preserved. The conch is sub-
globular to compressed and smooth. The
venter is rounded, merging onto convex
lateral areas. The umbilical shoulders are
broadly rounded. The umbilicus is very
small and markedly excentric on the outer
volutions. Because of the weathered
nature of most of the specimens, measure-
ments are not possible. The conch, how-
ever, varies significantly from depressed
forms (e.g. PI. 36, figs. 11, 12) to com-
pressed forms (e.g. PL 36, fig. 10). On
the basis of the available specimens there
appears to be a complete gradation be-
tween these extremes.
The suture is ceratitic with the first
lateral lobe on the mid-part of the lateral
areas, and a second lateral lobe on the
umbilical shoulder ( Fig. 18 ) .
This species is quite distinct from
IscuUtoides wasserbergi, which is more de-
pressed and constant in conch form, more
evolute, and has at least some ornamenta-
tion. IscuUtoides originis (Arthaber) from
Albania and Chios displays a wide range
in conch shape from depressed to com-
pressed similar to that of 7. luimmondi.
This Tethyan species tends to be more evo-
lute and more markedly excentric than the
Idaho species recorded here. The data on
IscuUtoides eUipticus Chao ( 1959 ) are
limited, but there is a strong similarity to
the Idaho species. It is not at all im-
possible that the Kwangsi and Idaho
specimens are conspecific.
Occurrence. Uppermost member of
Thaynes Formation, Hammond Creek,
Bear River Range, southeast Idaho.
Repository. Holotype, MCZ 9477 (PL
36, fig. 8); paratypes MCZ 9478 (PL 36,
figs. 9, 10), MCZ 9479 (PL 36, fig. 11),
MCZ 9480 (PL 36, figs. 12, 13); unfigured
paratypes MCZ 9486.
Genus Chiofifes Renz and Renz, 1947
Type species, Prosphingites (Chiotites)
globularis Renz and Renz, 1947
This new species is represented by ap- Small, very involute excentrumbilicate
420 BiiUetin Muscuni of Comparative Zoology, Vol. 137, No. 3
forms, with contracting living chamber; a
few prosiradiate constrictions near aper-
ture; whorls depressed with broadly arched
venter and rounded shoulders; livinu
chamber bears prominent strigations.
Suture with two denticulated lateral lobes,
a denticulated auxiliary lobe above the
umbilical shoulder and another one below
the umbilical shoulder.
In its involute, excentrumbilicate conch
this genus is quite similar to Isciilifoides
and Protropitcs, but the sutures of the
latter two genera are very different. The
suture of Chiotitcs is most similar to that
of Prosphingites except for the additional
auxiliary lobe.
The genus is known from a single species
in the Siibculiiiuhifcs fauna of Chios.
Chiotites globularis Renz and Renz
Text-figure 17
Prosphingites (Chiotitcs) globularis Renz and
Renz, 1947: 60, 74; Renz and Renz, 1948: 40,
pi. 15, figs. 9-9c.
Chiotites glohularvi, — Kummel, in Arkell et al.,
1957: L139, fig. 172, 11.
Prosphingites (Chiotitcs) siiperglobosus Renz and
Renz, 1947: 60, 75; Renz and Renz, 1948: 41,
pi. 15, figs. 7-7b, 10.
The species C. siiperglobosus was in-
troduced for the more inflated forms in the
Chios fauna from Maradovuno. However,
from the numerous paratypes in the Nat-
ural History Museum, Basel, it is quite
clear that there is a complete gradational
series from forms assigned to glohularis
to those placed in super glohosiis. In their
monograph the Renzes record only two
specimens for each of their two species.
There are in addition a number of speci-
mens in Basel. These additional specimens
are for the most part poorly preserved or
unprepared. A suture is illustrated on
Figme 17D. The measurements of the
figured specimens are as follows:
D
W
II
U W/D H/D U/D
1. 22.2 13.3 8.5 8.7 .59.9 .38.3 .39.2
2. 20.3 13.6 9.9 5.1 67.0 48.8 25.1
3. 21.8 16.8 7.3 5.7 77.1 .33.5 26.1
4. 22.6 ? 7.8 7.0 ? 34.5 31.0
1. Holotype, Renz and Renz (1948: pi. 15, fig.
9), NHMB J13642.
2. Paratype, Renz and Renz (1948: pi. 15, fig.
9c), NHMB J13643.
3. Holotvpe, C. superglohosus Renz and Renz
(1948: pi. 15, fig. 7), NHMB J13645.
4. Paratype, C. siiperglobosus Renz and Renz
(1948: pi. 15, fig. 10), NHMB J13646.
Occurrence. Stihcolumbites fauna, Mar-
adovuno, Chios.
Repository. Holotype, NHMB J13642;
paratypes NHMB J 1.3643 (Renz and Renz,
194(S: pi. 15, fig. 9c); unfigured paratypes
from Maradovuno NHMB J13644; holo-
type siiperglobosus Renz and Renz (1948:
pi. 15, fig. 7) NHMB JL3645; paratype
(Renz and Renz, 1948: pi. 15, fig. 10)
NHMB J 1.3646; unfigured paratypes from
Maradovuno NHMB J 13647.
Genus Cze/conows/c/fes Diener, 1915
Type species, Ceratites decipiens Mojsisovics,
1886
Czekonowskites decipiens (Mojsisovics)
Ceratites decipiens Mojsisovics, 1886: 27, pi. 6,
fig. 9.
Czekanoicskitcs decipiens, — Diener, 1915: 115;
Spath, 1934: 264; Kummel in Arkell et al.,
19.57: L143, fig. 175, 2; Kunimek 1961: 521.
Ceratites inostrauzeffi Mojsisovics, 1886: 28, pi.
6, fig. 10.
Czckanowskites inostranzeffi, — Diener, 1915: 115;
Kummel, 1961: .521.
It is unfortunate that in the extensive
revision of the Siberian Olenek fauna
Popov (1961, 1962a) apparently had no
additional specimens of this interesting
genus and species. There have been few
other records of the species since its intro-
duction by Mojsisovics. The specimen
from thc> Arctoceras horizon on Spitsbergen
that Frebold (1930) described as Czekan-
oicskitcs (?) sp. nov. does not belong in
this genus. Kummel (1954) recorded a
fragmentary specimen that probably be-
longs to this genus and is described below.
Occurrence. Olenckites Zone, Olenek
River region, Siberia.
Czekonowskites cf. decipiens (Mojsisovics)
Czckanouskilcs? sp. Kummel, 1954: 187.
A Iragiiu'iil consisting of a (juarter volu-
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 421
Hon of living chamber is believed to belong
to this genus, which is known from only
one species in the upper Scythian strata
at the mouth of the Olenek River, Siberia.
The adoral part of the fragment measures
19 mm high and 17.5 mm in width. The
whorl section is subquadrate in outline
with a broad, low arched venter, rounded
ventral shoulders, lateral areas that are
only slightly convex and convergent,
rounded umbilical shoulders, and a steep
umbilical wall. The lateral areas bear
radial ribs that enlarge slightly towards
the ventral shoulders. There is a faint
trace of the ribs across the venter, which
gives the venter an undulating appearance.
The posterior end of this fragment has
some traces of the last septum, enough to
indicate that there is a large first lateral
lobe, adjacent to large rounded saddles.
The second lateral lobe is high on the
lateral areas and is much smaller, and
there is probably a very small auxiliary
lobe on the umbilical wall.
This fragment is assigned to Czekanow-
skites on the basis of the great similarity
of the whorl outline and pattern of ribs
to the Siberian species of this genus. The
suture, insofar as it can be observed, is also
similar in basic pattern to that of the Si-
berian species. The major difference ap-
pears to be in the nature of the venter.
The Siberian species appears to have a
smooth venter, whereas the form recorded
here has traces of the ribs across the
venter.
Occurrence. Upper member of the
Thaynes Formation, Hammond Creek, Bear
River Range, southeast Idaho.
Repository. MCZ 9649.
Genus Popovites Tozer, 1965
Type species, Popovites occidentalis Tozer,
1965
"Inner whorls globose, outer whorl of
approximately equal height and width,
with a perpendicular umbilical wall, prom-
inent, rounded umbilical shoulder, flat
sides, and a broadly arched or slightly
flattened venter. Sculpture consists of
regular growth lines; radial wrinkles or
faint ribs may also occur on the venter.
Constrictions are absent. Body chamber
about one whorl in length. The suture line
comprises: a deep external lobe, with in-
cised branches; two ceratitic lateral lobes;
a suspensive lobe with one or more aux-
iliarv incisions; and one internal lateral
lobe" (Tozer, 1965a: 21).
This new genus shows marked affinities
to Frosphiniiites, on the one hand, and to
Czekanoiuskites, on the other. The suture
of Popovites is quite similar to that of
Prospliingites and at the same time not as
elaborate as the suture of Czekanowskites.
There is, however, in the fairly large size of
the first lateral lobe a similarity in the
suture of Popovites and Czekanotoskites
( Fig. 17E ) . The general conch form of
this new group of ammonites is like that
in various species of Prosphingitcs and
Czekano wskites.
The genus is known by two species from
an upper Scythian horizon in British Co-
lumbia and Ellesmere Island.
Popovites occidentalis Tozer
Text-figure 17
Popovites occidentalis Tozer, 1965a: 22, pi. 3,
figs. 2-12, text-figure 5.
The type species is well illustrated and
described by Tozer ( 1965a, Fig. 17E of
this report). It differs from P. boreolis
only in being slightly more involute; how-
ever, since that species is based on a single
specimen, there is no way of evaluating
the significance of this difference. Among
the measured specimens of P. occidentalis
there is only a variation of 5 percent in the
diameter of the umbilicus (Tozer, 1965a:
p. 23). The umbilical diameter of the holo-
type of P. boreolis is 31 percent at a diam-
eter of 31 mm and 35 percent at a diameter
of 43 mm. This is 5 and 10 percent, re-
spectively, greater than the maximum di-
ameter of the umbilicus for P. occidentalis.
Occurrence. Toad Formation, Halfway
River area, British Columbia, associated
422 Bulletin Museum of Comparative ZooJogij, Vol. 137, No. 3
with Prcflorianitcs intermedins Tozer and
Monacanthites monoceros Tozer.
Popovifes borealis Tozer
Popovites borealis Tozer, 1965a: 24, pi. 3, figs,
la, b, text-figure 6.
See above for discussion of this species.
Occurrence. Blaa Mountain Formation,
lower shale member, Ellesmere Island.
Genus Monacanthites Tozer, 1965
Type species, Monacanthites monoceros
Tozer, 1965
"Globose ammonoids with sculpture of
widely spaced, unbranched ribs that are
curved to form a sharp ventral sinus. On
the outer whorl each rib, at the ventral
mid-line, carries a single, solid spine. Body
chamber one whorl in length. Suture
ceratitic, with two lateral lobes, both in-
ternally and extemallv" (Tozer, 1965a:
27).
Monacanthites monoceros Tozer
Text-figure 17
Monacanthites monoceros Tozer, 1965a: 27, pi. 1,
figs. 8-10, pi. 2, fig. 4, text-fig. 8.
A unique species, quite unlike any other
ammonite of comparable age (Fig. 17F).
Occurrence. Toad Formation, Halfway
River area, British Columbia, associated
with Preflurianites interniedius Tozer and
Popovites occidentalis Tozer.
Genus Tunglanites Chao, 1959
Type species, Tunglanites lenticularis Chao,
1959
Involute, compressed conch with nar-
rowly rounded to acute venter; inner
volutions more inflated, bearing oblique
constrictions. Body chamber of one volu-
tion. Suture with single ceratitic lateral
lobe, two rounded lateral saddles.
The combination of conch shape and
suture of this genus is unique among late
Scythian ammonoids. The group was first
recognized from the Std)cohimhites fauna
of Albania by Arthabcr ( 1911 ) who placed
it in the genus Siyriles of much younger
age. This generic assignment for this late
Scythian form has never been accepted.
Diener (1915: 271) hsted it with question
in the genus Styrifes, and Spath (1934:
197) clearly recognized the independent
status of this form but refrained from pro-
posing a new generic name, most likely
because data on the Albanian forms were
very incomplete. Renz and Renz ( 1948 )
gave no indication that they were familiar
with any literature on this form after the
publication of Arthaber's paper in 1911.
The discovery of eight specimens in the
SubcoIumJntes fauna of Kwangsi, China,
has enabled a much clearer understanding
of the group, and a new generic name is
well justified.
There are at present two species of the
genus Tunglanites, T. lenticularis Chao,
the type species, and T. alexi n. sp. for the
Albanian and Chios forms assigned to
Sfyrites lilangensis Diener by Arthaber and
Renz and Renz, respectively. Direct com-
parison of these two species is handicapped
by the incompleteness in our kiiowledge of
the Albanian and Chios specimens. Un-
fortunately, one of the specimens studied
by Arthaber is lost and the Chios fauna
has yielded only two specimens. These
western Tethyan specimens were thought
by both Arthaber and the Renzes to have
a goniatitic lateral lobe. However, the
preservation, in hard red limestone, makes
development of the suture extremely dif-
ficult, and the smooth character of the lobe
may be due to over grinding or etching.
Regardless of this there can be no doubt
of the close relationship, and perhaps even
identity, of these two species.
In regard to the genetic relations of the
group, Spath (1934: 197) suggested they
represent a specialized offshoot of Isctdi-
toides. The recognition of faint constric-
tions on the early volutions of the Kwangsi
specimens, the compression of the conch,
and nature of the suture make this sug-
gestion perfectly reasonable.
In the three localities where Tunglanites
has been recorded, Albania, Clhios, and
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 423
Kwangsi, China, the associated fauna in-
cludes such typical upper Scythian genera
as Suhcohimhifes, Propiijchitoidcs, and
Ucnulccanites.
Tunglanifes lenficularis Chao
Text-figure 21
Tunglanitcs lenticularis Chao, 1959: 120, 294, pi.
27, figs. 25-32, pi. 28, figs. 23-25.
Involute, compressed, small conch with
rounded venter on early volutions be-
coming acute on last volution. Living
chamber one volution or more in length.
Surface may bear fine striae of growth
which curve backward along flanks; con-
strictions may be present on early volu-
tions. Suture with broad ventral lobe, two
rounded lateral saddles and single lateral
lobe; occasionallv a second lateral lobe is
present on the umbilical shoulder and wall.
Chao established his new genus and
species on the basis of eight specimens of
which three were illustrated. It appears
that the umbilicus is completely closed on
the earlier volution and gradually opens
slightly on the more adoral volutions. The
only other recorded occurrence of this genus
is that of specimens from Albania and
Chios which are xery similar in over-all
appearance to this Kwangsi species. The
western Tethyan specimens have a more
open umbilicus at a smaller diameter; how-
ever, the significance of this difference is
questionable. More material is needed to
evaluate the ontogenetic changes in growth.
Another distinctive feature is the suture
(Fig. 21A). The Chios and Albanian
specimens are reported to have smooth
lobes. However, the preservation of these
specimens in hard red limestone requires
grinding and acid to observe the suture in
most cases. It is not at all certain that the
lobes are really goniatitic. Tlie Albanian
specimen described by Arthaber (1911:
260, pi. 23(7), fig. 12) as Styrifes Ulangen-
sis Diener does not show a suture; just
where Arthaber obtained the suture of his
figure 12c is a puzzle ( Fig. 21B ) . The speci-
men from Chios described by Renz and
Renz (1948: 31, pi. 12, fig. 4) does not
show the suture clearly enough to establish
whether it is really goniatitic. I strongly
suspect that the lateral lobe is indeed
denticulated.
Occurrence. From a black, thin bedded
limestone 0.6 meters thick and 14 meters
above the lower Permian Maokou Lime-
stone (Chao collection 542b); the Chas-
hanao section of Chao (1959: 162) at the
border of Hochich and Tunglan districts
(Chao collection 610), western Kwangsi,
China. The Scythian strata at this locality
comprise only about 16 meters of shale and
limestone. The only fossils present are
from this 0.6 meter bed, which in addition
to Tunpjanitcs contains Subcohimbitcs,
PropfycJiitoides and Hemilecanites.
Tunglanifes alexi n. sp.
Plate 20, figures 1, 2; Text-figure 21
Styrites lilangensis, — Arthaber (non Diener), 1911:
260, pi. 23(7), figs. 11a, b, 12a, b, c; Renz and
Renz, 1947: 60; Renz and Renz, 1948: 31, pi.
12, figs. 4-4a.
Styrites (?) cf. lilangensis Diener, 1915: 271.
Gen. nov. "Stt/rites" lilangensis, — Spath, 1934:
189, 197.
This species is established for the speci-
mens Arthaber (1911) and Renz and
Renz (1948) described as Styrites lilangen-
sis Diener from the Siibcohimbites faunas
of Albania and Chios. Arthaber had two
specimens from Albania both of which he
illustrated; the smaller of his two speci-
mens (Arthaber, 1911: pi. 23(7), fig. 11)
is lost, but the larger specimen (Arthaber,
1911: pk 23(7), fig. 12) is available and
is selected as the type specimen. Tlie di-
ameter of this specimen is approximately
30 mm, the width of the adoral whorl 8.5
mm, the height 12.5 and the diameter of
the umbilicus 6.7 mm. A portion of the
phragmocone is broken off and missing.
The whorls are compressed and convergent,
forming an acute venter. The umbilical
shoulders are very low but rounded; the
umbilicus is excentrumbilicate. Arthaber
(1911: pi. 23(7), fig. 12c) shows a suture
(Fig. 21B of this report) which presum-
424 Bulletiii Museum of Comparotivc Zoology, Vol. 137, No. 3
B
Figure 21. Diagrammatic representation of the suture of: A, Tunglanites lenficu/or/s Choo from an upper Scytfiian hori-
zon in Kwangsi, China (1959: pi. 27, fig. 32), at a diameter of approximately 15 mm; B, Tunglanites alexi n. sp. (:=
Styrites lilangensii, — Arthaber, 1911: pi. 23(7), fig. 12c) from Subco/umbites fauna of Albania, suture presumably from
paratype of Arthaber (1911: pi. 23(7), fig. 11) which Is apparently lost; C, Paradinantes sum Chao (1950: fig. 4), holo-
type, at a diameter of approximately 30 mm; D, Pseudoceltites conztrictilis (Astakhova, I960; fig. 8) from Columbites
Zone of Astakhova (1960) on the Mangyshlak Peninsula, at a diameter of approximately 15 mm; E, Pseudoceltites nevadi
n. sp., from Upper Thaynes Formation, Confusion Range, Utah, un-numbered paratype at whorl height of approximately
10 mm; F, Pseudoce/fifes nevadi n. sp., from Upper Thaynes Formation, Confusion Range, Utah, un-numbered paratype at
whorl height of approximately 10 mm.
ably was taken off the other type specimen.
Careful examination of this specimen has
failed to show any trace of a suture.
The specimen from the SuhcohimJ)itcs
fauna of Chios illustrated by Renz and
Renz (1948: pi. 12, fig. 4) measures 17.1
mm in diameter, 5.2? mm for the width of
the adoral whorl, 7.0 mm for the height
of the adoral whorl and 3.5 mm for the
diameter of the umbilicus. Renz and Renz
state that their Chios specimen had a
goniatitic suture; however, my examination
of the specimen reveals that the suture is
not well preserved and it cannot be estab-
lished whether it is goniatitic or not. I
strongly suspect that the lateral lobe is
indeed denticulated.
The general shape of the conch is very
much like that of T. lenticiilaris except for
an apparently slightly larger umbilicus.
Occurrence. Suhcoliimhifes fauna of Al-
bania and Chios.
Repository. Holotypc (PI. 20, figs. 1, 2),
Paleontological Institute, University of
Vienna; specimens from Chios NHMB
J13582 (Renz and Renz, 1948: pi. 12, fig.
4), unfigured specimen NHMB J13583.
Genus Columbites Hyatt and Smith, 1905
Type species, Columbifes parisianus Hyatt
and Smith, 1905
Columbites parisianus Hyatt and Smith
Plate 39, figures 1-10; Plate 40,
figures 1-11; Plate 41, figures 1—7;
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 425
Plate 42, figures 1-9; Plate 43,
figures 4, 5; Text-figures 22, 23.
Coliimbites parisianus Hyatt and Smith, 1905: 51,
pi. 1, figs. 9-14, pi. 61, figs. 1-21, pi. 72, figs.
1-24; Freeh, 1908: pi. 42, fig. 2; Diener,
1915: 112; Diener, 1925: 69, pi. 24, fig. 2;
Smith, 1932: 107, pi. 1, figs. 9-14, pi. 61, figs.
1-21, pi. 72, figs. 1-24; Spath, 1934: 201, pi.
13, fig. 3, text-fig. 61; Kummel, in Arkell,
et al., 1957: L140, fig. 172, 2.
Coliimlntes spencei Smith, 1914: 36, pi. 70, figs.
1-16, pi. 71, figs. 1-16; Smith, 1932: 108, pi.
77, figs. 1-21, pi. 78, figs. 1-16; Kutassy,
1933: 490.
Cohnnlntes consanquinciis Smith, 1932: 106, pi.
46, figs. 1-13.
Cohimhites minimus Smith, 1932: 106, pi. 47,
figs. 9, 10.
Cohimhites hpatus Smith, 1932: 106, pi. 47, figs.
1-8.
Cohimhites ornatus Smith, 1932: 107, pi. 46,
figs. 14-21.
All the specimens assigned to the six
species of Columhiies by J. P. Smith came
from outcrops of the middle shale member
of the Thaynes Formation {Cohimhites
fauna) in Paris Canyon, southeast Idaho.
The original description of the type species
by Hyatt and Smith (1905), later slightly
enlarged by Smith (1932), is quite ade-
quate. The five additional "species" intro-
duced by Smith in 1932 were distinguished
on differences in whorl dimensions and
on ornamentation. On Table 33 are given
the measurements of 107 specimens of
CoJumhitcs from the same horizon, at three
localities around the north end of Bear
I^ake (including Paris Canyon), southeast
Idaho. These data are plotted on Figure
23. It can readily be seen from this large
sample that slight differences in conch
dimensions are meaningless. The dif-
ferences in ornamentation are more dif-
ficult to quantify. But here again, as with
most ornamented ammonites, there is com-
plete gradation from very weak ribs to
strong ribs; within this species there is also
an ontogenetic variable, that is, successive
patterns of ribs appear in differing orders.
All of these features, however, are com-
pletely gradational. Close study of the
specimens figured here will bring this fact
out.
Smith ( 1932 ) did not make particular
note of the suture of his six species but
this feature is also quite variable. On
Figure 22 are 13 sutures of Cohimhites.
As can be seen, the variation is expressed
in the shape and size of the first and
second lateral lobe and thus also in the
saddles.
There are no other well authenticated
species of Cohimhites recorded to date.
Coliimhifes sp. described by Kiparisova
(1961:119) from an uncertain horizon
(CoJiimhites Zone?) in the Scythian of the
Primorye Region is based on a single poorly
preserved specimen. It is possible this is
a species of Cohimhites, but it is much
more involute than C. i)arisionus. At the
same time it could well be a species of
Pseiidocchites. From northeiTi Siberia,
Popov (1961) has described two species
of Cohimhites, Cohimhites (?) aff. ornatus
Smith and Cohimhites morpheos Popov.
The first of these records is based on two
poorly preserved casts showing no suture;
I believe these to be unidentifiable. The
second, C. morpheos, is a species of Tiro-
hfes and is discussed under that genus in
this paper. Finally, there are Cohimhites
doJnapaensis Kiparisova (1947: 143) and
Cohimhites constrictilis Astakhova (1960:
pi. 140) from the Mangyshlak Peninsula
of southern Russia. These two species
belong in Pseiidocehites and are discussed
under that genus. Cohimhites parisiamis is
thus far only known from southeast Idaho.
Occurrence. Middle shale member,
Thaynes Formation, Cohimhites fauna at
Paris Canyon, Montpelier Canyon, and Hot
Springs, all around north end of Bear Lake,
southeast Idaho, and same horizon along
Draney Creek, Stewart Flat Quadrangle,
southeast Idaho (USGS locality M98).
Repository. Holotype (PI. 39, figs. 3,
4) USNM 75246a; paratypes (PI. 39, figs.
8, 9) USNM 75246b, (PI. 41, fig. 7) USNM
75286a, (PI. 39, figs. 1, 2) USNM 75286b,
(PI. 39, figs. 5-7) USNM 75286c, (Smith,
426 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 427
1932, pi. 61, figs. 8, 9) USNM 75286d, (PI.
39, fig. 10) USNM 75286e; all the remain-
ing small juvenile specimens of this species
studied by Hyatt and Smith (1905) and
Smith (1932) are in the U.S. National
Museum but are not formally numbered;
suture specimens of Figure 22, MCZ 9532-
9538; specimens from Montpelier Canyon
MCZ 9633, from Hot Springs 9634; from
Draney Creek, Stewart Flat Quadrangle,
southeast Idaho, USGS; holotype C. con-
sanquinciis Smith, (PI. 41, figs. 1, 2) USNM
74983a; paratypes (PI. 41, figs. 3, 4) USNM
74983b, (PI. 41, figs. 5, 6) USNM 74983c,
(Smith, 1932: pl. 46, figs. 7-9) USNM
74983d, (Smith, 1932: pl. 46, figs. 10-13)
USNM 74983e; holotype C. liiiatus Smith,
(Pl. 43, figs. 4, 5) USNM 74985a; paratypes
(Pl. 42, fig. 7) USNM 74985b, (Pl. 40,
figs. 7-9) USNM 74985c; holotype C.
minimus (Smith, 1932: pl. 47, figs. 9, 10)
USNM 74986; holotype C. omatus Smith,
(Pl. 40, figs. 1, 2) USNM 74984a; para-
types (Pl. 40, figs. 10, 11) USNM 74984b,
('smith, 1932: pl. 46, figs. 19-21) USNM
74984c; holotype C. spencei Smith (Pl. 42,
figs. 1, 2) USNM 75309; paratypes (Smith,
1932: pl. 78, fig. 3) USNM 75309b, (Pl.
42, figs. 3, 4) USNM 75309c, (Pl. 42, figs.
8, 9) USNM 75309d, (Pl. 42, figs. 5, 6)
USNM 75309e, (Pl. 40, figs. 5, 6) USNM
75309f, (Pl. 40, figs. 3, 4) USNM 75309g,
specimens of Smith (1932: pl. 77, figs.
1-21 ) USNM 75309h-i.
Genus Subcolumbites Spath, 1930
Type species, Co/umb/fes perrinismithi
Arthaber, 1908
The most common elements in many
late Scythian faunas are species of Sub-
columbites. The five species of this genus
recognized to date can be separated into
three distinct groups. The first group con-
tains only the type species, which is known
from Albania, Chios, China, and Japan,
and is characterized by carination of the
venter. The second group contains only
S. dus77Uini, and is characterized by a more
marked development of reticulate orna-
mentation. The third group contains S.
robustus from China, S. multiformis from
the Primorye Region, and S. americanus
from Nevada. This third group is char-
acterized by a more depressed whorl sec-
tion.
Subcolumbites perrinismifhi (Arthaber)
Plate 1, figures 1-9; Plate 2, figures
5-8; Plate 3, figures 1-9; Plate 4,
figures 1-4; Text-figure 24
Columhites perrini-smiihi Arthaber, 1908: 277,
pl. 12, fig. 1; Arthaber, 1911: 262, pl. 23(7),
figs. 19, 20; Diener, 1915: 112; C. Renz, 1928:
155; Renz and Renz, 1947: 59; Renz and
Renz, 1948: 20, pl. 11, figs. 7-7a.
Subcolumbites pcrriui-.srnithi, — Spath, 1930: 77;
Spath, 1934; 203, pl. 12, figs. 5a, b; Kummel,
in Arkell et al., 1957; L140, figs. 172, 15a, b;
Kummel, 1968b: 495, pl. 1, figs. 1-3.
Columbites europacus Arthaber, 1908; 278, pl.
12, fig. 2; Arthaber, 1911; 261, pl. 23(7),
Figure 22. Diagrammatic representation of the sutures of Columbites parisianus Hyatt and Smith. A, suture of holotype
at a diameter of 35 mm (USNM 75246a), from Smith 11932: pl. 1, fig. 11); B, at a diameter of 35 mm (USNM 75286b),
from Smith (1932: pl. 61, fig. 4); C, suture of holotype of C. ligatus Smith (1932: pl. 47, fig. 3), at a diameter of 40
mm (USNM 74985a); D, at a diameter of 46.7 mm (MCZ 9532); E, at o diameter of 22.3 mm (MCZ 9533); F, paratype
of C. spencei Smith (1932: pl. 78, fig. 4), at an approximate diameter of 35 mm (USNM 75309b); G, paratype of C.
consanguineus Smith (1932: pl. 46, fig. 4), at a diameter of approximately 40 mm (USNM 74983b); H, at a diameter of
42.0 mm (MCZ 9534); I, paratype of C. ornafus Smith (1937: pl. 46, fig. 18), at a diameter of 25 mm (USNM 74984b);
J, at a diameter of 28.6 mm (MCZ 9535); K, at a diameter of 33.7 mm (MCZ 9536); L, at a diameter of 23.0 mm (MCZ
9537); M, at a whorl height of 10 mm (MCZ 9538). All specimens from Co/umbites fauna, Thaynes Formation, southeast
Idaho; A, B, C, F, G, I, are from Paris Canyon, the remaining specimens from Hot Springs.
428 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Table 33. Measurements of Columbites parisianus Hyatt and Smith from Columbites
FAUNA FROM THREE LOCALITIES AROUND NORTH END OF BeAR LaKE, SOUTHEAST IdAHO.
D
w
H
u
W/D
H/D
U/D
D
w
H
u
W/D
H/D
U/D
1.
67.9
17.3
18.4
34.8
25.5
27.1
51.4
41.
40.3
0
10.8
20.4
?
27.0
50.5
2.
66.2
16.0
19.5
32.2
24.2
29.4
49.4
42.
.39.4
12.0
10.5
20.8
30.5
26.7
52.8
3.
65.0
16.5
18.8
31.8
24.4
39.0
49.0
43.
39.0
11.4
11.9
17.7
29.2
30.5
45.5
4.
63.0
16.0
19.4
27.4
25.4
30.8
43.5
44.
38.7
11.0
12.5
16.5
28.4
32.3
42.7
5.
62.5
16.9
16.7
30.4?
27.1
26.7
54.4?
45.
38.3
11.0
11.7
18.6
28.7
30.6
48.6
6.
61.7
15.3
20.6
25.9
24.8
33.4
42.0
46.
37.8
11.9
12.0
17.8
31.5
31.7
47.1
7.
59.7
15.0
19.1
25.0
25.1
32.0
41.9
47.
.37.6
11.9
11.3
17.8
31.6
30.1
47.4
8.
59.0
15.4
16.1
29.5
25.8
26.9
49.3
48.
37.3
11.9
10.0
18.5
31.9
26.8
49.6
9.
58.7
15.8
15.0
.32.7
27.1
25.5
55.6
49.
36.7
11.8
9.4
18.7
32.2
25.1
51.0
10.
58.5
16.6
16.4
30.2
28.4
28.1
51.6
50.
36.4
12.0
10.3
17.8
33.0
27.5
48.9
11.
58.3
16.7
18.4
28.3
28.6
31.6
48.6
51.
35.8
10.6
10.1
17.5
30.5
28.2
48.9
12.
56.5
17.3?
15.6
29.0
30.6?
27.6
51.3
52.
35.7
11.4
10.1
18.0
31.9
28.3
50.5
13.
54.5
14.2
17.1
24.2
26.1
31.4
44.4
53.
.35.6
11.2
10.3
18.4
31.3
28.9
50.6
14.
54.0
15.5?
17.2
23.7
28.7?
31.9
43.9
54.
35.3
13.3
9.3
18.5
.37.7
26.3
52.4
15.
54.0
14.4
17.4
25.3
26.7
32.3
46.9
55.
35.0
10.5
9.8
16.4
.30.0
28.0
46.9
16.
53.5
15.4
15.0
27.8
28.8
28.0
52.0
56.
33.8
12.7
8.8
17.8
37.6
26.0
52.7
17.
52.4
15.7
14.3
26.1
30.0
27.3
49.8
57.
33.4
10.5
9.2
17.3
31.5
27.5
51.8
18.
50.7
14.4
14.0
25.8
28.4
27.8
50.8
58.
33.2
9.7
11.1
14.5
29.2
33.4
43.7
19.
50.5
14.0
15.0
24.3
27.7
29.7
48.1
59.
.32.1
10.5
10.0
15.0
.32.7
31.2
46.7
20.
50.5
14.7
13.5
27.3
29.2
26.7
54.1
60.
31.4
11.7
8.9
16.1
37.3
28.3
51.3
21.
49.8
12.8
15.5
23.4
25.7
31.2
47.0
61.
31.4
11.0
11.2
12.5
35.0
.35.7
39.9
22.
49.7
?
14.8
24.7
?
29.7
49.5
62.
30.9
13.9
8.4
16.2
45.0
27.2
52.5
23.
49.4
15.8
13.8
25.2
32.0
28.0
50.6
63.
.30.8
11.3
10.5
13.0
36.9
.34.1
42.3
24.
49.0
13.7
15.0
20.8
27.9
30.6
42.4
64.
30.5
10.8
7.5
16.2
34.8
24.6
53.2
25.
48.7
15.0
12.5
27.8
30.7
25.6
56.9
65.
30.3
11.5
10.2
13.2
.38.0
33.6
43.6
26.
48.4
13.4
16.3
21.1
27.7
33.7
43.6
66.
29.0
10.0
8.7
13.7
34.5
30.0
47.3
27.
48.2
9.4
12.9
24.4
19.5
26.7
50.6
67.
29.0
10.0
7.7
15.0
34.5
26.5
51.7
28.
48.2
13.7
13.7
23.1
28.4
28.4
48.0
68.
28.8
11.1
8.7
9.1
.38.6
.30.2
.39.1
29.
48.0
14.3
12.5
24.8
29.8
26.1
51.6
69.
28.7
10.6
8.2
15.0
36.9
28.6
52.3
30.
47.8
14.7
13.7
24.7
.30.8
28.7
51.6
70.
28.5
10.8
8.0
14.4
37.8
28.1
,50.6
31.
47.7
14.0
12.7
25.4
29.4
26.6
53.3
71.
28.0
11.7
9.1
13.0
42.3
32.5
46.4
32.
46.6
15.0
14.4
23.4
.32.2
30.9
50.2
72.
27.0
9.6
7.3
12.5
35.5
27.0
46.3
33.
44.2
13.3
13.5
21.0
30.1
30.6
47.6
73.
26.4
10.0
7.0
14.0
37.9
26.5
53.0
34.
43.7
11.9
13.4
20.0
27.3
.30.7
45.8
74.
26.1
11.0
7.2
13.8
42.2
38.7
55.0
35.
43.0
12.3
12.0
22.8
28.6
27.9
53.0
75.
26.0
9.7
9.4
11.8
,37.2
36.2
45.4
36.
42.7
13.5
11.7
22.1
31.6
27.4
51.8
76.
25.3
9.4
7.3
13.3
.37.1
28.8
52.5
37.
42.3
11.1
13.5
18.2
26.2
31.9
43.0
77.
25.0
10.7
7.0
12.8
42.8
28.0
51.3
38.
41.5
11.5
11.4
19.6
27.7
27.5
47.2
78.
24.6
9.5
8.4
10.9
38.8
34.1
44.3
39.
41.5
12.3
12.4
21.3
29.6
29.9
51.4
79.
24.5
10.5
7.0
13.0
42.9
28.5
53.1
40.
41.0
10.9
12.8
18.7
26.6
31.2
45.6
80.
24.3
11.5
9.2
10.3
47.3
27.9
42.4
3.
6.
7.
11.
13.
1.5.
26.
40.
4.5.
46.
59.
61.
63.
65.
78.
SO.
Plesiotype, .Smith (1932: pi. 61, fig. 1), USNM 75286a.
Holotype, C. ligatus .Smith (1932: pi. 47, fins. 1-3), USNM 74985a.
Paratype, C. comanguincus Smith (1932: pi. 46, fig. 3), USNM 74983b.
Holotype, C. con.saufiuincus
Holotypo, C. spcncci Smith
Holotype, C. ornaliis Smith
Ple.siotype, Smith (1932: pi
Plesiotype, Smith ( 1932: pi.
Holotyiie, Smith ( 1932: pi
Smith (1932: pi. 46, fifis. 1, 2), USNM 74983a.
(1932: pi. 78, fijrs. 1, 2), USNM 75309a.
(1932: pi. 46, fius. 14, 15), USNM 74984a.
61, fius. 2-4), USNM 75286b.
61, fijjs. .5-7), USNM 55286f.
.1, figs. 9-11), USNM 75246a.
Paratype, C. ligaUis Smith (1932: pi. 47, figs. 4, 5), USNM 7498,5b.
Paratype, C. spencei Smith (1932: pi. 78, figs. 5, 6), USNM 75309c.
Paratype, C. ligaius Smith (1932: pi. 47, figs. 6-8), USNM 7498.5c.
Plesiotype, Smith (1932: pi. 61, figs. 8, 9), USNM 75286(1.
Paratype, Smith (1932: pi. 1, figs. 12-14), USNM 752 161).
Paratype, C. spencei Smith (1932: pi. 78, figs. 7, 8), USNM 75309(1.
Paratype, C. ornalns Smith (1932: pi. 46,
Paratype, C. spencei Smith (1932: pi. 78, f
fiRs. 16-18), USNM 74984b.
i«s. 9, 10), USNM 75309c.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 429
Table 33. Continued.
D
w
H
u
W/D
H/D
U/D
D
w
H
u
W/D
H/D
U/D
81.
23.7
10.0
7.2
10.3
42.2
30.1
43.5
95.
14.3
7.0
4.6
5.7
48.9
32.2
.39.8
82.
23.0
10.0
8.0
9.4
43.5
.34.8
40.7
96.
14.3
7.8
5.1
6.4
54.5
35.6
44.7
83.
22.8
9.4
6.5
11.1
41.2
28.5
48.7
97.
13.8
8.0
4.7
6.5
58.0
.34.0
47.1
84.
22.7
9.8
6.5
10.7
43.2
28.6
47.2
98.
13.5
7.0
5.0
5.2
56.0
.37.0
.38.5
85.
22.7
9.3
6.5
11.2
40.9
28.6
49.3
99.
13.0
4.0
4.0
6.7
30.8
30.8
51.5
86.
90 0
11.8
6.7
10.2
53.3
30.2
46.0
100.
11.4
4.9
3.8
4.5
43.0
.33.3
.39.4
87.
21.9
9.4
8.3
8.5
42.9
.37.8
.38.8
101.
10.6
6.6
3.7
4.8
62.2
.34.9
45.3
88.
21.8
10.4
6.2
10.0
47.7
28.4
45.9
102.
9.7
5.5
3.5
3.7
56.7
36.1
.37.8
89.
20.5
9.8
6.0
10.8
47.8
29.3
52.7
103.
9.5
5.5
3.5
4.1
58.0
36.9
43.2
90.
19.3
9.2
5.8
9.3
47.6
.30.1
48.2
104.
8.8
6.1
3.2
3.5
69.2
.36.4
39.8
91.
18.7
9.5
7.2
8.2
50.8
38.4
43.8
105.
8.4
5.1
3.0
2.8
60.7
.35.7
33.3
92.
17.0
8.8
4.7
8.4
51.7
27.6
49.4
106.
8.0
4.5
2.8
3.6
56.3
35.1
45.0
93.
15.8
7.3
4.3
7.3
46.2
27.2
46.2
107.
7.2
4.4
2.5
2.3
61.1
.34.7
32.0
94.
15.3
7.5
5.2
6.7
49.0
34.0
.33.8
81. Paratvpe, Smith (19.32: pi. 61, figs. 11-13), USNM 75286f.
82. ParaHpe, C. spencei Smith (1932: pi. 78, figs. 13-16), USNM 75309g.
87. Parat\pe, C. consanguineus Smith (1932: pi. 46, figs. .5, 6), USNM 74983c.
91. Paratope, C. spencei Smith (1932: pi. 78, figs. 11, 12), USNM 75309f.
94. Paratvpe, Smith (1932: pi. 61, figs. 14, 15), USNM 75286g.
96. Paratvpe, Smith (1932: pi. 72, figs. 22-24), USNM 7.5286q.
97. Paratvpe, C. spencei Smith (1932: pi. 77, figs. 1-4), USNM 7.5309h.
98. Paratvpe, C. consanguineus Smith (1932: pi. 46, figs. 7-9), USNM 74983d.
99. Holotype, C. minimus Smith (1932: pi. 47, figs. 9, 10), USNM 74986.
100. Paratvpe, C. ornatus Smith (1932: pi. 46, figs. 19, 20), USNM 74984c.
101. Paratype, C. spencei Smith (1932: pi. 77, figs. 5-8), USNM 7.5309i.
102. Paratype, Smith (1932: pi. 61, figs. 16-18), USNM 75286h.
103. Paratype, Smith (1932: pi. 72, figs. 19-21), USNM 75286p.
104. Paratyiie, C. spencei Smith (1932: pi. 77, figs. 9012), USNM 75309k.
105. Paratvpe, C. consanguineus Smith (1932: i^l. 46, figs. 10-13), USNM 74983e.
106. Paratvpe, Smith (1932: pi. 72, figs. 16-18), USNM 75309o.
107. Paratype, C. spencei Smith (1932: pi. 77, figs. 13-15), USNM 753091.
All other specimens are from the Columbites fauna of Paris Canyon, Montpelier Canyon, and Hot Springs, around
north end of Bear Lake, southeast Idaho.
figs. 13-18; Diener, 1915: 112; C. Renz, 1928:
1.55; Renz and Renz, 1947: 59; Renz and Renz,
1948: 19, pi. 11, figs. .3-3a, 4-4a, 5-5a, 6-6a.
SiiJjcoltnubitcs europaeiis, — Spath, 1934: 204, pi.
12, figs. 6a, b, text-fig. 62c.
Columbites europaeus-perrini-smithi Renz and
Renz, 1947: 59; Renz and Renz, 1948: 20,
pi. 11, figs. 1-lb, 2-2b.
Columbites mirditensis Arthaber, 1911: 263, pi.
24(8), fig.s. 2, 3, 4; Diener, 1915: 112; C. Renz,
1928: 1.55; Renz and Renz, 1947: 59; Renz
and Renz, 1948: 21.
Subeohnnbites mirditensis, — Spath, 1934: 205.
Sul)cohimbites kwangsianus Chao, 1959: 128, 304,
pi. 30, figs. 14-17, text-fig. 41c.
Columbites asijmmetricus Chao, 1959: 127, 303,
pi. 30, figs. 10-13.
Subcolumbites cf. peninismithi, — Bando, 1964a:
99, pi. 3, figs. 18, 19, pi. 4, fig. 3.
Arthaber (1908, 1911) had 70 specimens
from the Kcira, Albania, fauna that he
assigned to four species of Columbites.
Of this original collection there are avail-
able today only 16 specimens, most of
which were illustrated by Arthaber. The
unretouched photographs reproduced here
clearly demonstrate the poor preservation
of these Albanian specimens, and that
Arthaber's illustrations are highly re-
touched.
Both Arthaber (1911) and Spath (1934)
recognized the gradational nature of the
"species" established for these Albanian
forms. Three of these so-called species
(S. europaeus, S. peninismithi, and S.
mirditensis) vary mainly in the degree of
compression of the whorls. As the whorls
become more compressed the umbilical
shoulder is more rounded, as the whorls
become more depressed the umbilical
shoulder becomes more acutely rounded.
Subcolumbites perrinismithi is the more
compressed form, S. mirditensis the more
430 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
35r-
30
25
20
15
10
•
.65
X X
•
•
•63
X40
•
•61
X
"?? X
X
•75
45x
X
\
x6i ,
X
X
• . '78
X
81
X6i
X
.. ••80
65" X50
X
• ^2- X80^^^'
«" , '
X ^
87*
W X
•
•p, x87 X78
•81 x82
X X
X
" X
•
X9I BIX ^^ X "
.99 .94
97«.96
X
•
X "
58^¥«94
100. 97«x ^ X
°^* . »l(52
A%^4,S?'
XI07
•|07
•26
X26
•15
•13
•2
•3
•7
u
"2
13 V X,
•40 3
«ll
_I6
H
0 10 20 30 40 50 60 70
DIAMETER
Figure 23. Variation in whorl height (H) and umbilical diameter (U) in Columbites parisianus Hyatt and Smith from Co-
lumbites Zone, Thaynes Formation, southeast Idaho. The data on this graph are from Table 33.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 431
depressed form. Restudy of the available
specimens from Kcira, conspecific forms
from Chios, and topotypes of the Kcira
forms in the British Musemn (Natural
History) clearly show that there is com-
plete gradation in the degree of compres-
sion of the whorls. The measurements of
the types available from Kcira and Chios
are given on Table 34. The sutures of
S. minlitensis and S. europaeus are illus-
trated on Figures 24 A, B.
The pattern and intensity of ornamenta-
tion is also quite variable. In terms of the
forward projecting ribs, they range from
the fine, regular pattern, as illustrated by
S. peninismithi (PL 3, figs. 6, 7), to one
where in the adoral volutions the ribs are
bunched, especially on the venter, forming
chevrons (Pi. 1, figs. 7, 8).
Among the four species established by
Arthaber (1908, 1911), there is one (S.
diismani) which is quite distinct from the
other three. Arthaber had only two
specimens, both of small size and poor
preservation. This species differs from S.
perr'mismithi, as interpreted here, in lack
of a tendency toward carination of the
venter and in the more conspicuous re-
ticulate ornamentation. I cannot agree at
all that this form is "scarcely more than a
variety of S. europaeus" (Spath, 19.34: 205).
This conclusion is strengthened by the
discovery of better preserved conspecific
forms on Chios and other species of this
general ornamentational pattern from
Kwangsi, China, and Nevada.
Subcohimbitcs kwangsianus Chao ( 1959)
was established for two poorly preserved
specimens from Kwangsi, China. Though
very poorly preserved, the whorl shape,
carination of the venter, and ornamentation
are like those features in S. perrinismithi
of Albania and Chios. Chao (1959: 304)
recognized the affinities of his species to
the Albanian forms but concluded that the
elliptical coiling of his species served to
distinguish it. The so-called elliptical coil-
ing is no more than that apparent in some
Table 34. Measurements of Subcolumbites
PERRINISMITHI (ArTHABEr) FROM ALBANIA AND
Chios.
D
w
H
U
W/D
H/D
U/D
1.
60.4
17.5
19.0
28.7
28.9
31.5
47.5
2_
58.0
16.8
17.5
27.5
28.9
.30.2
47.4
3.
55.0
17.7
17.2
23.1
32.2
31.3
42.0
4.
54.4
18.1
20.0
24.6
33.3
.36.8
45.2
5.
53.7
13.0
17.4
23.8
24.2
32.4
44.3
6.
52.3
17.2?
16.6
24.2
32.9?
31.7
46.3
7.
49.3
?
14.4
22.8
?
29.2
46.2
8.
48.8
?
13.6
23.9
?
27.9
48.9
9.
47.5
?
16.0
22.0
?
33.7
46.3
10.
46.0
?
15.0
21.3
?
.32.6
46.3
11.
43.5
?
14.7
20.0
?
33.8
45.9
12.
40.7
15.4
14.1
18.3
37.8
.34.6
44.9
13.
37.0
?
13.0
16.3
?
35.1
44.1
14.
34.4
?
11.8
15.7
p
34.3
45.6
15.
.34.3
15. r?
11.2
14.8
44.0?
.32.7
43.1
16.
28.3
11.7
10.3
12.0
41.3
36.4
42.4
17.
27.3
14.1
10.1
9.7
51.6
36.9
35.5
1. Plesiotype, Cohunbitcs europaeus Arthaber (1911: p\.
23(7), figs. 18a, b), PIUV.
2. Plesiotype, Cohimbites europaeus-perrini-smithi Renz
and Renz (1948: pi. 11, figs. 1, la), NHMB J13538.
3. Plesiotype, Cohimbites europaeus-perrini-smithi Renz
and Renz (1948: pi. 11, figs. 2, 2a), NHMB J13539.
4. Plesiotype, Cohimbites europaeus Arthaber (1911: pi.
23(7),' figs. 15a, b), PIUV.
5. Plesiotype (Arthaber, 1911: pi. 23(7), figs. 20a, b),
PIUV.
6. Plesiotype, Cohimbites europaeus, — Renz and Renz
(1948? pi. 11. figs. 3-3a), NHMB J13533.
7. Plesiotype, Cohimbites perrinismithi, — Renz and Renz
(1948.' pi. 11, figs. 7, 7a), NHMB J13537.
8. Plesiotype, Columbites europaeus, — Renz and Renz
(1948: pi. 11, figs. 4, 4a), NHMB J13534.
9. Paratvpe, Cohimbites mirditensis Arthaber (1911: pi.
24(8'), figs. 3a, b), PIUV.
10. Hokmpe (Arthaber, 1908: pi. 12, figs, la-c) , PIUV.
11. Plesiotype (Arthaber, 1911: pi. 23(7), figs. 19a, b),
PIUV.
12. Plesiotype, Cohimbites europaeus Arthaber (1911: pi.
23(7)', figs. 16a, b), PIUV.
13. Holotvpe, Cohimbites mirditensis Arthaber (1911: pi.
24(8), figs. 2a, b), PIUV.
14. Paratvpe, Cohimbites mirditensis Arthaber (1911: pi.
24(8), figs. 4a, b), PIUV.
15. Plesiotype, Cohimbites europaeus, — Renz and Renz
(1948: pi. 11, figs. 5, 5a), NHMB J13535.
16. Plesiotype, Cohimbites europaeus Arthaber (1911: pi.
23(7)', figs. 13a-c), PIUV.
17. Plesiotype, Cohimbites europaeus, — Renz and Renz
(1948i pi. 11, figs. 6, 6a), NHMB J13536.
mens of S. perrinismithi (PL 1, figs. 3, 9).
Subcolumbites kwangsianus is considered
to be a synonym of S. perrinismithi. The
two specimens from Japan recorded by
"'as Subcohimbites cf.
Bando (1964a: 99) a.5 ^uiy<.Lyci4i,.iy.n.o ^x.
perrinismithi, though poorly preserved, are
of the poorly preserved Albanian speci- surely conspecific with this species. The
432 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
A
H
D
Ammonoids of the Late Scythian (Lower Triassic) • Kitmmel 433
sutures of these species are shown on
Figures 24A, B, E, J.
Occurrence. Subcolumbites faunas of
Albania, Chios, Afghanistan, China, and
Japan.
Repository. The following specimens are
in the Paleontological Institute, Vienna:
holotype Cohimbites perrini-smithi Art-
haber, 1908: pi. 12(2), figs, la-c (PI. 4,
figs. 1, 2 of this report); topotype, Cohim-
bites perrini-smithi Arthaber, 1911: pi.
23(7), figs. 19a, b (PL 3, figs. 1-3 of this
report); topotype CoJumbites perrini-smithi
Arthaber, 1911: pi. 23(7), figs. 20a, b (PI.
3, figs. 6, 7 of this report); holotype, Cohim-
bites europaeiis Arthaber, 1908: pi. 12 (2),
figs. 2a-d (PI. 4, fig. 3 of this report);
topotvpes, Cohimbites europaeus Arthaber,
1911:' pi. 23(7), figs. 13-18 (PI. 1, figs. 1-9,
PI. 2, figs. 5-6 of this report); syntype,
Cohimbites mirditensis Arthaber, 1911: pi.
24(8), fig. 2 (PI. 3, figs. 8, 9 of this report);
syntype, Cohimbites mirditensis Arthaber,
1911: pi. 24(8), fig. 3 (PI. 2, figs. 7, 8 of
this report); type specimen CoJumbites
mirditensis var. Arthaber, 1911: pi. 24(8),
fig. 4 (PI. 3, figs. 4, 5 of this report).
The following specimens are in the
Natural History Museum, Basel: plesiotype
Cohimbites perrini-smithi, — Renz and Renz
(1948: pi. 11, fig. 7) NHMB J13537;
plesiotypes CoJumbites europaeus, — Renz
and Renz (1948: pi. 11, fig. 3) NHMB
J13533, (pi. 11, fig. 4) NHMB J13534,
(pi. 11, fig. 5) NHMB J13535, (pi. 11, fig.
6) NHMB J13536; unfigured specimens
from Maradovuno NHMB J13543, from
Kephalovuno NHMB J13544; syntypes
CoJumbites europoeus-perrini-smitJi i Renz
and Renz (1948: pi. 11, fig. 1) NHMB
J13538, (pi. 11, fig. 2) NHMB J13539;
unfigured specimens from Maradovuno
NHMB J13547, from Kephalovuno J13548;
recorded specimen of CoJumbites mirditen-
sis—Renz and Renz (1948: 21) NHMB
J 13550.
The British Museum (Natural History)
contains the following topotype specimens
from the SubcoJumbites fauna of Albania:
S. perrini-smitJii, C911-15, C22916-23,
C22924-6; S. europaeus, C22890-900,
C22901-10; S. mirditensis, C22883-6,
C22887-9. Paratype from Albania MCZ
6723, from Chios MCZ 10026, specimens
from Afghanistan MCZ 10138, 10146.
Subcolumbites dusmani (Arthaber)
Plate 2, figures 1-4; Text-figure 24
Cohimbites du-smanl Arthaber, 1911: 263, pi.
24(8), figs, la-d; Diener, 1915: 112; Renz
and Renz, 1947: 73.
Suhcohimbites- chismani, — Spath, 1934: 204.
Columhites duinac Renz and Renz, 1947: 59, 73;
Renz and Renz, 1948: 21, pi. 10, figs. 6-6b,
7-7b.
Columhites diamie var. invohita Renz and Renz,
1947: 59.
CoJumljites diamie var. evohita Renz and Renz,
1948: 22.
Cohimbites graeco-americanus Renz and Renz,
1947: 59, 73; C. Renz, 1947: 176; Renz and
Renz, 1948: 27, pi. 10, figs. 4-4b.
Figure 24. Diagrammatic representation of the sutures of species of Subco/umb/fes. A, syntype of Co/umfaites mird/'fensis
Arthaber (1911: pi. 24(8), figs. 2a-c; PI. 3, figs. 8, 9 of this report), at a diameter of 25 mm; B, Co/umb/tes europaeus
Arthaber (1911: pi. 23(7), figs. 15o-c; PI. 1, figs. 1 , 2 of this report), at a diameter of 35 mm; C, paratype of Columbites
d/anoe Renz and Renz (1948: pi. 10, fig. 7b), at a diameter of 25 mm; D, paratype of feng%harii\es robustus Chao (1959:
129, fig. 42a), at a diameter of approximately 30 mm; E, holotype of Subco/umb/tes /twangs/anus Chao (1959:
128, fig. 41c), at a diameter of approximately 40 mm; F, Subco/umbifes multiformis Kiparisova (1947: 144, fig. 32),
at a diameter of 31 mm; G, Subco/umbites multiformis Kiparisova (1947: 144, fig. 31), at a diameter of approxi-
mately 15 mm; H, paratype of Subco/umbites americonus n. sp. (MCZ 9435, PI. 30, fig. 8], at a diameter of 31 mm; I,
paratype of Subco/umb/fes americanus n. sp. (MCZ 9438, PI. 30 figs. 13, 14), at a diameter of 14 mm; J, syntype of
Co/umb/tes europaeus-perrini-smittii Renz and Renz (1948: pi. 11, fig. 2b), at a diameter of approximately 40 mm.
Specimens of figures A, B from Subco/umb/fes fauna of Albania; C, J from same horizon on Chios; D, E from same horizon
in Kwangsi, China; F, G from same horizon in the Primorye Region, eastern Siberia; H, I from same horizon in Tobin For-
mation, Nevada.
434 BuUeiin Museum of Comparative Zoology, Vol 137, No. 3
Columhites aithuliae Renz and Renz, 1947: 59,
74; Renz and Renz, 1948: 28, pi. 10, figs.
3-3b, 5-5b.
Columhites parisianus, — C. Renz, 1945: 301; C.
Renz, 1947: 176; Renz and Renz, 1947: 59;
Renz and Renz, 1948: 22, pi. 11, figs. 8-8b.
Cohitnhites spencei Smith var. chiotica Renz and
Renz, 1947: 59, 73; Renz and Renz, 1948: 22,
pi. 3, figs. 7-7b.
The two syntypes of this species are
relatively small specimens preserved only
on one side and that only modestly well.
These two specimens differ from S. perrini-
smiihi in the strong reticulate ornamenta-
tion and the lack of any tendency toward
carination. The reticulate ornamentation
is more conspicuous on one syntype (PI. 2,
figs. 1, 2) than on the other (PI. 2, figs.
3, 4) where it is only faintly visible.
The Suhcolumhitcs fauna of Chios has
yielded a number of what are believed
to be conspecific forms. Renz and Renz
( 1948 ) had nine specimens of the morpho-
logical type of S. dusmani which they
placed in five different species. Three of
these species were based on one specimen
each, one species was based on two speci-
mens, and one species on four specimens.
The large number of species introduced by
Renz and Renz for this group reflects
pronounced morphological differences from
one specimen to the other. Whereas the
sample comprising all these species from
Chios is extremely small, a case can be
made that the patterns of morphological
differences are most likely gradational and
that we are dealing with a single variable
species. Likewise, study of large popula-
tions of other species of a similar morpho-
logical type, Columhites parisianus for
instance, offers an insight into the poten-
tial variability that is possible in some of
these Scythian ammonoids.
The holotype of Columhites dianac Renz
and Renz is a large well preserved speci-
men clearly conspecific with S. dusmani.
Renz and Renz (1918: 21) did not con-
sider Arthaber's two syntypes of S. dusnuDii
to be conspecific. On this basis they
designated one of the specimens (Arthalier,
1911: pi. 24(8), figs, la, b; PI. 2, figs. 3,
4 of this report) as the type (lectotype)
of S. du.smani and considered the other
specimen as conspecific with their C.
dianae from Chios. The separation of
Arthaber s two syntypes of S. dusmani can-
not be accepted. The principal difference
in these two specimens is in the expression
of the reticulate ornamentation which is
most pronounced on the ventral region.
The specimen which Renz and Renz (1948:
21) designated as the type of S. dusmani
is very badly weathered over most of the
the venter; however, small traces of the
shell are present and these clearly show a
nice reticulate pattern. There is no justifi-
cation for separating these two specimens
into different species.
In addition to the well developed re-
ticulate pattern on the holotype of Colum-
hites dianae, there is on the adoral quarter
volution a bundling of the prosiradiate
ribs and a decrease in the strigations. In
Columhites aithaliac there is a very pro-
nounced bundling of the ribs producing
strong prosiradiate folds over the ventral
regions and extending half way up the
flanks on the last quarter volutions. This
change in ornamentational patterns takes
place at an approximate diameter of 28.0
mm, whereas in Columhites dianae this
change takes place at approximately 70
mm in diameter.
Columhites graecoamericanus Renz and
Renz has the coarse bundled ribs developed
on the whole adoral \-olution. Columhites
speneei var. ehiotiea Renz and Renz is
nothing more than a sparsely ribbed graeeo-
amerieanus. Each species was established
on a single specimen. Finally, the Colum-
hites parisianus of Renz and Renz is based
on a small inner whorl of C. dianae. The
sutures of these species are all of the same
basic pattern and vary only in details
(Fig. 24C).
The few specimens that are available
suggest that the differenc(\s in ornament
pattern are most probabl) a reflection of
(liflerenees in ontogenetic- growth, that is.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 435
Table 35. Measurements of Subcolumbites
DUSMANI (ArTHABEr) FROM ALBANIA AND ChIOS.
D
w
H
u
W/D
H/D
U/D
L
74.2
25.5
31.5
21.7
34.4
42.5
29.2
■1
54.8
22.5
19.3
21.5
41.1
35.2
39.2
3.
51.3
17.8
17.7
20.7
34.7
34.5
40.4
4.
38.8
?
14.5
15.3
?
37.4
39.4
5.
37.7
?
14.5
14.9
p
38.5
39.5
6.
35.7?
p
15.3
10.4
?
59.5?
29.1?
7.
33.0
25.4
12.7
10.8
76.9
38.5
.32.7
8.
26.6
14.8
15.2
10.0
55.6
57.1
37.6
9.
23.7
11.6
7.6
11.7
48.9
32.1
49.4
1. Holotvpe, Columbites dianae Renz and Renz (1948:
pi. id, fig. 6), NHMB J13540.
2. Hc)l((t\pe, Columbites graeco-americanus Renz and
Renz (1948: pi. 10, figs. 4-4b), NHMB .T13564.
.3. Plesiotvpe, Columbites spencei var. chiotica Renz and
Renz (1948: pi. 3, figs. 7-7b), NHMB J135.53.
4. Svntvpe (Arthaber, 1911: pi. 24(8), figs, la, b), PIUV.
.5. Syntype (Arthaber, 1911: pi. 24(8), figs. Ic, d), PIUV.
6. Holotype, Columbites aithaliae Renz and Renz (1948:
pi. 10, figs. 5-5b), NHMB J13568.
7. Paratvpe, Columbites aithaliae Renz and Renz ( 1948:
pi. 10, figs. 3-3b), NHMB J13.570.
8. Paratype, Columbites dianae Renz and Renz (1948:
pi. 10, fig. 7), NHMB J 13541.
9. Plesiotvpe, Columbites parisianus, — Renz and Renz
(1948! pi. 11, figs. 8-8b), NHMB J13542.
different rates of appearance of the suc-
cessive stages. This type of variabihty in
gro\\'th patterns is well illustrated in
Columbites parisianus where we have a
large population to work with and can
document all of the transitional forms
convincingly. It is largely on the basis of
my studies of Columbites parisianus that
I have come to the conclusion that these
five species from Chios of Renz and Renz
are all conspecific. Measurements of these
species and of Arthaber's are given on
Table 35. This interpretation, of course,
needs confirmation, which can only be done
when additional collections of these species
are assembled.
SubcoJmnbites dusmani can be easily
distinguished from S. perrinismithi in the
lack of any tendency toward carination of
the venter and in the general greater de-
velopment of the reticulate ornamentation.
Subcolumbites robustus Chao has a much
more depressed whorl and low, broad,
radial folds on the flanks, as does S. ameri-
canus from Nevada. Subcolumbites multi-
formis is also a species with a depressed
whorl section but has no lateral nodes or
folds.
Occurrence. Subcolumbites fauna of Al-
bania and Chios.
Repository. Arthaber's two syntypes
are in the Paleontological Institute, Uni-
versity of Vienna. The Chios fauna studied
by Renz and Renz ( 1948 ) is in the Natural
History Museum, Basel — holotype, Colum-
bites dianae Renz and Renz (1948: pi. 10,
fig. 6) NHMB J13540; paratype (pi. 10,
fig. 7) NHMB J13541; unfigured paratypes
NHMB J13551; holotype, Columbites
graecoamericamis Renz and Renz (1948:
pi. 10, fig. 4) NHMB J13564; unfigured
paratypes J13565; holotype, Columbites
aithaliae Renz and Renz (1948: pi. 10,
fig. 5) NHMB J13568; paratype (pi. 10,
fig. 3) NHMB J13570; unfigured paratypes
from Maradovuno NHMB J 13571, from
Kephalovuno NHMB J13572; plesiotype,
Columbites parisianus, — Renz and Renz
(1948: pi. 11, fig. 8) NHMB J13542; un-
figured specimens NHMB J13552; type
specimen, Columbites spencei var. chiotica
Renz and Renz (1948: pi. 3, fig. 7) NHMB
J 13553.
Subcolumbites robusfus (Chao)
Text-figure 24
Fengshanitcs rohustiis Chao, 1950: 4, pi. 1, figs.
2, 3; Chao, 1959: 129, 305, pi. 8, figs. 1, 2,
pi. 29, figs. 21-22, text-fig. 42a.
This species is of the same general
morphological type as S. americanus from
Nevada. Although the species is based on
only two specimens, its distinctness is
readily apparent. The whorls are more
inflated and depressed than in S. dusmani.
In addition, there are low irregular folds
on the flanks. The suture is illustrated on
Figure 24D.
Occurrence. 1.5 km north of Yali, Feng-
shan district, associated with Dagnoceras
and Hellenites (Chao collection 546),
Kwangsi, China.
436 BuUetin Museum of Comparative Zoology, Vol. 137, No. 3
Subcolumbifes multiformis Kiparisova
Text-figure 24
Subcolumbites multiformis Kiparisova, 1947: 144,
pi. 32, figs. 8-11, text-figs. 31-34; Kiparisova
and Krishtofovich, 1954: 22, pi. 13, figs. 1-3;
Kiparisova, 1961: 121, pi. 27, figs. 1-7, text-
fig. 82-88.
SuhcoJumhitcs solittis Kiparisova, 1961: 123, pi.
26, figs. 4, 5, text-fig. 89, 90.
SubcoJtimbites anomalus Kiparisova, 1961: 123,
pi. 26, figs. 6, 7, text-fig. 91.
Kiparisova ( 1961 ) had 35 specimens
from what I judge to l)e the same horizon
and locahty at Cape Zhitkov, Primorye
Region, eastern Sil:)eria. Of these speci-
mens, she placed 25 in S. multiformis, first
described by her in 1947; seven specimens
were placed in a second species, S. solittis:
and three specimens were placed in an-
other new species, S. anomalus. The latter
two species were distinguished on the
basis of slightly greater whorl depression
and minor differences in the nature of the
ornamentations. The suture, however, in
all three species is essentially the same
(Fig. 24F, G).
Unfortunately, Kiparisova supplied mea-
surements for only 10 specimens of her
three species of Subcolumbites; these data
are tabulated on Table 36. On the basis
of these data, there does not appear to be
any real difference in relative whorl pro-
portions. Kiparisova recognized that her
S. multiformis was a highly variable form
and that nearly all her specimens of Sub-
columbites were juvenile specimens. This
does not appear to be any justification for
recognizing any more than one specie's of
Subcolumbites within this faunal group.
Subcolumbites multiformis does not
show any particularly close relationship
to the group of S. perrinismithi or S. dus-
mani, but is very similar to S. robustus
from Kwangsi, China, and S. americanus
from western United Stales. The similarity
to the latter two species is expressed in the
depressed whorl section, reticulate orna-
mentation, and in the suture. It differs
from these two species, however, in the
lack of any lateral folds or nodes.
Table 36. Measuhements of Subcolumbites
multiformis, s. solitus, and s. anomalus from
UPPER Scythian beds, Primorye Region, eastern
Siberia.
D
w
H
U
W/D
H/D
U/D
1.
45.0
24.8
15.3
18.9
55.0
34.0
42.0
2.
32.0
22.7
10.9
11.8
71.0
34.0
37.0
3.
29.0
18.9
9.9
11.3
65.0
34.0
39.0
4.
24.5
14.5
9.2
8.3
59.0
38.0
34.0
5.
17.0
11.9
5.4
6.5
70.0
32.0
38.0
6.
16.5
14.9
5.0
5.0
90.0
30.0
30.0
7.
16.0
13.4
3.5
5.9
84.0
22.0
37.0
8.
16.0
10.1
5.0
5.9
63.0
31.0
37.0
9.
11.5
10.4
3.0
4.5
90.0
26.0
39.0
10.
8.0
8.6
2.6
2.6
108.0
33.0
33.0
1-3, 5, 7, 9. Stibcolmnbites tntiltifonnis, data from Kipari-
sova, 1961: 121.
4, 8. Stibcohimhites soUtus, data from Kiparisova, 1961:
123.
6, 10. Stibcohiuibitcs anomalus. data from Kiparisova,
1961: 124.
Occurrence. Subcolumbites fauna at
Cape Zhitkov, Primorye Region, eastern
Siberia.
Subcolumbites americanus n. sp.
Plate 30, figures 1-14; Text-figure 24
This is one of the common ammonites
in the Tobin Formation fauna from Ne-
vada. The collections contain a large
number of well preserved but mainly frag-
mentary specimens. However, ten speci-
mens are sufficiently complete to yield
measurements (Table 37).
The conch is evolute, robust, and with
a characteristic pattern of ornamentation.
The venter is arched and grades imper-
ceptibly onto convex lateral areas. The
umbilical shoulders are sharply rounded
and the umbilical wall steep but not verti-
cal. The conch bears low nodes situated
just on the ventral side of the umbilical
shoulder. In addition, there are prominent
striae of gro\\th, often bundled, which are
rectiradiate on the umbilical wall and pro-
siradiate on the lateral areas, completely
crossing the venter. There are also periodic
constrictions and fine strigation.
The suturc> is shown on Figures 24H, I.
Clearly, this species is closely allied to
S. multiformis from the Primorye Region
and S. robusftis from Kwangsi, China.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 437
Table 37. Measubements of Subcolumbites
americanus n. sp. from the tobin formation
OF Nevada.
D
w
H
u
W/D
H/D
U/D
1.
47.3
25.0?
19.3
16.3
52.9?
40.8
34.5
2.
47.0
?
17.1
16.4
?
36.4
34.8
3.
42.2
20.8?
16.8
15.6
49.3?
39.8
36.9
4.
35.2
?
12.0
13.2
?
34.1
37.5
5.
31.8
16.1
12.6
9.7
50.6
39.6
30.5
6.
31.1
16.7
11.4
10.2
53.7
36.7
22.5?
7.
20.4
11.8?
7.3
7.0?
57.8
35.8
34.3?
8.
20.1
11.2
7.6
6.6
55.7
37.8
32.8
9.
20.1
10.3
7.8
5.7
51.2
38.8
28.4
10.
17.0
10.8
6.5
5.5
63.5
38.2
32.4
1. Holotype,
2. Paratype,
3. Parat>pe,
4. ParaHpe,
5. Paratvpe,
6. Paratype,
7. Paratype,
8. Parat\pe,
9. Paratype,
10. Paratype,
MCZ
MCZ
MCZ
MCZ
MCZ
MCZ
MCZ
MCZ
MCZ
MCZ
9430
9431
9492.
9433
9434
9435
9436
9432
9437
9438
(PI.
(PI.
(PI.
(PI.
(PI.
(PI.
(PI.
(PI.
(PI.
30,
30,
30,
30,
30,
30,
30,
30,
30,
figs.
fig. ;
fig.
figs.
fig.
figs.
fig.
figs.
figs.
1, 2).
•5).
6,
8).
9,
4).
11,
13,
7) .
10).
12;
14;
Occurrence. Basal part of Tol:)in FoiTna-
tion, USGS locality M2562, Pershing
County, Nevada; south tip of Tobin Range,
Cain Mountain 1:62,500 quad., center NW
Vi sec. 9, T. 26N, R. 39E, 5,500 ft. S, 27.5
ft. W of elevation point 5088 on range
crest.
Repository. Holotype (PL 30, figs. 1, 2)
MCZ 9430; figured paratypes (PI. 30, fig.
3) MCZ 9431, (PI. 30, fig. 4) MCZ 9432,
(PI. 30, fig. 5) MCZ 9433, (PI. 30, figs. 6,
7) MCZ 9434, (PL 30, fig. 8) MCZ 9435,
(PL 30, figs. 9, 10) MCZ 9436, (PL 30, figs.
11, 12) MCZ 9437, (PL 30, figs. 13, 14)
MCZ 9438; unfigured paratypes MCZ 9492.
Genus Paradinarifes Chao, 1950
Type species, Paradinarifes suni Chao, 1950
Paradinarifes suni Chao
Text-figure 21
Paradinarites suni Chao, 1950: 6, pL 1, figs, 7a,
b, text-fig. 4; Chao, 1959: 98, 330, pi. 41,
figs. 9-12.
Chao (1959: 331) recognized that this
new genus and species was quite similar
to ''Cohimbites' in conch form and gross
aspect of the suture but considered the
goniatitic character of the lobes to indicate
affinity with the dinaritids. The general
shape of the conch is that of a Siihcolum-
bifes but lacks strigations. The suture
(Fig. 21C), also, with its large first lateral
lobe is columbitid in plan and, in fact,
quite like the suture of Procohimbites
karataucihis Astakhova (1960a, b) from
the upper Scythian Formation of the
Mangyshlak Peninsula. The species is
known from only t\\'o not very well pre-
served specimens but on the basis of the
data available it clearly appears to be a
columbitid.
Occurrence. Subcolumbites iciuna (Chao
collection 610), Kwangsi, China.
Genus Pseudocelfifes Hyatt, 1900
Type species, Celfifes mulfiplicafus Waagen,
1895
Evolute, ribbed ammonites of rather
simple design are very common in mid-
Scythian formations. This has led to a
proliferation of specific and generic names,
the relationships of which are seldom
understood or appreciated. The type
specimen of Celtites multiplicatus is a
poorly preserved specimen from the Upper
Ceratite Limestone of the Salt Range (PL
27, figs. 5, 6). Celtites armatus Waagen
(1895: 75, pi. 7, figs. 1, 7), the type species
of Kashmirites Diener (1913) is clearly a
synonym of Celtites multiplicatus Waagen
(PL 27, figs. 7-10). Spath (1930: 35)
introduced the name Anakash mi rites (type
species Danubites nivalis Diener, 1897: 51,
pi. 15, figs. 17-19) for ammonoids not too
different in appearance from Celtites mul-
tiplicatus. Examination of the Salt Range,
Himalayan, and Timor specimens that
have been assigned to one or more of
these three genera by Waagen (1895),
Diener (1897, 1909, 1913), Welter (1922),
and Spath (1930, 1934) show that the
ornamentational pattern is highly variable.
It is in the suture pattern that one can
readily separate Pscudoceltites and Ana-
kashmirites. The former genus has a very
reduced second lateral lobe that rests on
or near the umbilical shoulder, whereas in
438 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Anakashmirites there is a "normal" second
lateral lobe and generally an auxiliary
lobe at or near the umbilical shoulder.
As mentioned above, the type species is
from the Upper Ceratite Limestone of the
Salt Range, West Pakistan. In the upper
part of the Scythian, three species are
recognized: one (cheneyi) from the Co-
Inmhites fauna of southeastern Idaho, the
second (ncvadl) from the Upper Thaynes
Formation, western Utah, and the third
{dohuipaensis) from the upper Scythian
formation of the Mangyshlak Peninsula.
This genus is grouped here with the
columbitids on the basis of the gross aspect
of the conch and especially the basic pat-
tern of the suture. In these aspects this
genus shows similarities to such columbi-
tids as CoJumhites and Procohunhites.
Pseudocelfifes cheneyi n. sp.
Plate 44, figures 4-10; Text-figure 25
A number of exposures of the Cohnn-
hites fauna of the Thaynes Formation in
southeastern Idaho have yielded approxi-
mately 50 generally well-preserved speci-
mens of this distinctive species. The conch
is evolute and compressed. Measurements
on 15 well-preserved specimens from one
horizon and locality are given on Table 38.
As can be readily seen, there is very little
variation in relative thickness and height
of the whorls or in umbilical diameter. The
lateral areas of the whorls are flattened
and slightly convergent; the venter is
broad and arched. Tlie ventral shoulder
is broadly rounded, and the umbilical
shoulder is more abruptly rounded with a
short, steep umbilical wall. The umbilicus
is broad and shallow.
Table 38. Measureaients of Pseudoceltites
cheneyi n. sp. from the columbites fauna of
Draney Creek, southeastern Idaho. The width
dimension inclt-tdes the lateral ribs.
D
W
H
u
W/D
H/D
U/D
1.
29.2
8.4
9.6
12.0
28.8
32.9
41.1
2,
29.0
?
10.0
12.1
?
47.8
57.9
3.
27.8
8.6
9.8
12.1
30.9
.35.3
43.5
4.
27.5
7.6?
10.0
10.0
27.6?
36.4
36.4
5.
26.5
7.8
7.6
11.6
29.4
28.7
43.8
6.
26.0
8.0
9.6
9.2
30.8
36.9
35.4
7.
25.7
7.8
9.4
9.1
30.4
36.6
.35.4
8.
22.7
7.4
7.8
9.0
32.6
.34.4
.39.6
9.
21.7
7.3
7.5
9.6
33.6
.34.6
44.2
10.
21.4
7.0
6.9
8.3
32.7
32.2
38.8
11.
20.1
7.4
6.9
8.5
36.8
34.3
42.3
12.
20.1
7.5
6.5
9.2
.37.3
32.3
45.8
13.
16.6
6.5
5.7
6.8
.39.2
34.3
40.9
14.
14.0
6.4
4.5
5.3
45.7
.32.1
37.9
15.
13.0
5.1
4.6
4.9
39.2
.35.4
,37.7
The lateral areas bear straight prosi-
radiate ribs that expand slightly toward
the ventral shoulder where they project
slightly forward. The ribs do not extend
on to the venter, which is smooth except
for growth lines and occasional constric-
tions which join the interrib areas from
the opposite flanks. Tliere are approxi-
mately 20 such ribs on the outer volution.
There is some variation in the spacing of
the ribs and in the relative strength or
prominence of the ribs, but this variation
is very slight and appears to be of no
significance.
The suture consists of two rounded
lateral saddles, a prominent first lateral
lobe, with some denticulations, and a
small, generally pointed second lateral lobe
on the umbilical shoulder. The variation
in the shape and proportions of the sutural
elements is quite spectacular (Fig. 25).
Figure 25. Diagrammatic representation of the sutures of Pseudoceltites cheneyi n. sp. A, at a diameter of 13 mm (MCZ
9507); B, at a diomefer of 17 mm (MCZ 9508); C, at a diameter of 20 mm (USNM 153074); D, at a diameter of 17 mm
(MCZ 9503, PI. 44, fig. 4); E, at a diameter of 20 mm (USNM 153075); F, at a diameter of 17 mm (USNM 153076); G, at
a diameter of 16 mm (WSU); H, at a diameter of 15 mm (MCZ 9509); I, at a diameter of 14 mm (MCZ 9506); J, at a
diameter of 17 mm (USNM 153077); K, at a diameter of 20 mm (USNM 153073, PI. 44, figs. 8, 9), fioiotype; L, at a diam-
eter of 17 mm (MCZ 9574).
All specimens from Columbites fauna, Thaynes Formation, southeastern Idaho; specimens A, B, D, H, I, J, and L are from
Hot Springs; C, E, F, G, and K from Draney Creek.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 439
B
440 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
The variation as illustrated by these 12
sutures should be sobering to those who
tend to erect species on minor differences
in the suture.
The type specimen of Celtites nudtipli-
catus Waagen is a weathered specimen, in
which only the outer volution is present;
the inner whorls are completely weathered
out (PL 27, figs. 5, 6). Waagen s illustra-
tion (1(S95: pi. 7, fig. 2) is highly restored
and actually misleading. Even though the
preservation of the Salt Range specimen
leaves much to be desired, it is morpho-
logically very similar to Pseudoccltites
cheneiji. The Idaho species differs in its
prosiradiate ribs rather than the radial ribs
of /-*. multiplicatus. Waagen mentions
slight nodes on the umbilical and ventral
shoulders associated with the ribs; these
nodes, however, are not present on the
type specimen. The basic pattern of the
suture of these two species is the same
( Mg. 25). Celtites multiplieatus came
from the Upper Ceratite Limestone of the
Salt Range, which is mid-Scythian in age.
The present species is from the Coliim1)ite.s
Zone.
Occurrence. Middle shale member of
Thaynes Formation, Columlntcs fauna,
along Draney Creek, Stewart Flat, Quad-
rangle ( uses locality M9<S ) ; and in Mont-
pelier Canyon, Hot Springs, and Paris
Canyon, all in the Bear Lake region of
southeastern Idaho.
Repository. Ilolotype USNM 153073
(PI. 44, figs. 8, 9); figured paratypes, MCZ
9503 (PI. 44, fig. 4), MCZ 9504 (PI. 44,
fig. 5), MCZ 9505 (PI. 44, figs. 6, 7), MCZ
9506 (PI. 44, fig. 10); suture specimens
MCZ 9507 (Fig. 25A), MCZ 9508 (Fig.
25B), USNM 153074 (Fig. 25C), MCZ
9503 (Fig. 25D), USNM 153075 (Fig.
25E), USNM 153076 (Fig. 25F), Depart-
ment of Geology, Washington State Uni-
versity (Fig. 25G), MCZ 9509 (Fig. 25H),
MCZ 9506 (Fig. 251), USNM 153077 (Fig.
25J), USNM 153073 (Fig. 25K), MCZ
9574 (Fig. 25L); unfigun^d specimens
from riot Springs, southeast Idaho M(-Z
9510; unfigured specimens from Mont-
pelier Canyon, southeast Idaho MCZ 9511.
Pseudocelfites dolnapaensis Kiparisova
Text-figure 21
Coliimhites dolnapaensis Kiparisova, 1947: 143,
pi. 30, fig. 3, text-fig. 30.
Coliuubites constrict (lis Astakhova, 1960a: 140,
pi. 33, fig. 6, text-fig. 8.
This species is remarkably similar to
Pseudoccltites cJieneiji n. sp. from the
Cohimbites fauna of southeastern Idaho in
its conch shape, ornament, and suture. The
differences are primarily centered on the
pattern of forward projecting constrictions
on the adoral part of the conch. This
species cannot be assigned to Cohimbites
as the patterns of ribs and constrictions are
very different. Astakhova ( 1960a ) dis-
tinguished her species constrictdis from
Cohn)d)ites dolnapaensis Kiparisova partly
on the basis that constrictdis had two den-
ticulations on the first lateral lobe, and
dolnapaensis had three. Examination of
Figure 25, with 12 sutures of F. cheneiji,
will give some indication of the variations
possible within the basic pattern of the
suture.
Occurrence. Mangyshlak Peninsula, Co-
himbites Zone of Astakhova (1960a), as-
sociated with Albanites, Epicehites and
OJenekites.
Pseudocelfites nevadi n. sp.
Plate 34, figures 1-5; Text-figure 21
Xcnoccltitcs et. A', spitshcr^cnsis, — Sill:)eding, in
Hose and Repenning, 1959: 2189, 2194.
The collections contain a large number
of fragmentary, poorly preserved speci-
mens. The basic form of the conch, whorl
cross-section, pattern of ribs, and suture is
very much like that of Pseudoccltites
cheneiji from the Cohimbites fauna of
southeastern Idaho. The suture (Figs. 21
and 25) likewise is very similar. It is
possible that these two species are con-
specific, but nevadi attains a much greater
si/(^ than cJienciji, and on this basis it is
thought best to keep the forms separate.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 441
One fragment must be from a specimen of
a diameter of approximately 65 mm.
Occurrence. USGS fauna Mill, from
upper part of Thaynes Formation, in sec-
tion 15 of Hose and Repenning (1959:
2187), Confusion Range, western Utah.
Repository. Holotype USNM 153078
(PI. 34, fig. 1); paratvpes USNM 153079
(PI. 34, figs. 2, 3), USNM 153080 (PL 34,
figs. 4, 5).
Genus Procolumb'iies Astakhova, 1960
Type species, Procolumbifes karataucikus
Astakhova, 1960
Procolumbifes karataucikus Astakhova
Text-figure 26
Procolumbites karataucikus Astakhova, 1960a:
142, pi. 34, figs, la-c, text-fig. 9 {nowcu
nudum Bajarunas, 1936: 547).
A columbitid with low \entral keel on
phragmocone, venter on body chamber
rounded. Ornamented with radial ribs and
constrictions that cross the venter. Suture
(Fig. 26G) with single, pointed, lateral
lobe.
Occurrence. This genus and species is
only known from the upper Sc\thian for-
mation of the Mangyshlak Peninsula in
beds associated with Pseudoceltites dol-
napacnsis, Olenekites tnangysJiIakensis, and
Albanites triadicus.
Genus Prenkifes Arthaber, 1911
Type species, Prenkites malsorensis
Arthaber, 1911
There are only three species of this
genus. One of these (timorensis) is known
from Chios, Timor, and China. The type
species is represented in the Suhcolumhitcs
fauna of Albania and Chios, whereas the
third species (helenoe) is only known
from the Chios fauna.
Prenkifes malsorensis Arthaber
Plate 7, figures 7-10; Text-figure 26
Prenkites malsorensis Arthaber, 1911: 258, pi.
22(6), figs. 17-19; Diener, 1915: 226; C.
Renz, 1931: 344; Spath, 1934: 208, pi. 12,
figs. 7a-c, text-figs, lllf-h; Renz and Renz,
1947: 59; Renz and Renz, 1948: 29, pL 12,
figs. 11-lla, 12.
Arthaber (1911: 258) stated he had 29
specimens of this species, but of these only
6 are still preserved in the Paleontological
Institute, Vienna. In contrast to this fair
representation, the Subcolumbites fauna of
Chios has yielded only two specimens. The
general character of the conch and the
suture ( Fig. 26A, B ) is quite similar to
that of P. timorensis which is distinguished
on the basis of its ornamentation. Pren-
kites helenae Renz and Renz ( 1948 ) is a
more broadly evolute form \\'ith rounded
umbilical shoulders and lacking the small
nodes on the umbilical shoulder.
Occurrence. Subcolumbites fauna of Al-
bania and Chios.
Repository. Si.x specimens, including the
two figured by Arthaber (1911: pi. 22(6),
figs. 17, 19; PI. 7, figs. 7-10 of this report)
are in the Paleontological Institute, Vienna.
The two plesiotypes from Chios, NHMB
J13574, J13575.
Prenkifes helenae Renz and Renz
Text-figure 26
Prenkites helenae Renz and Renz, 1947: 60, 74;
Renz and Renz, 1948: 30, pi. 12, figs. 2-2a,
5-5a.
This species is an evolute malsorensis
with rounded umbilical shoulders that lack
the small nodes. The species is based on
only two specimens; the measurements of
the holotype are Diameter 49.3 mm. Width
18.5 mm. Height 18.3, Umbilicus 17.5, the
same for the paratype are Diameter 35.8,
Width 17.8, Height 12.1, Umbilicus 14.6.
The suture is shown on Figure 26C.
Occurrence. Subcolumbites fauna, Chios.
Repository. Holotype NHMB J13577;
paratype NHMB J13578; unfigured para-
types from Maradovuno J13579, 13580.
Prenkifes timorensis Spath
Text-figures 17, 26
Columbites nov. sp. indet. Welter, 1922: 150,
pi. 168(14), figs. 12, 13.
442 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
M
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 443
Prenkites timorensis Spath, 1930: 77; Spath,
1934: 208, fig. 62d, e.
Cohimbites malaijanus C. Renz, 1945: 301; C.
Renz, 1947: 176; Renz and Renz, 1947: 59,
73; Renz and Renz, 1948: 24, pi. 9, figs. 5,
6-6c, 8-8b, 9-9d, 10.
Cohimbites malaijanus var. crassa Renz and Renz,
1947: 59, 73; Renz and Renz, 1948: 26, pi. 9,
fifis. 4-4 b.
Cohimbites bubidinac Renz and Renz, 1947: 59,
73; Renz and Renz, 1948: 26, pi. 9, figs. 7-7a,
pi. 10, figs. 1-lb.
Cohimbites levantiniis Renz and Renz, 1947: 59,
74; Renz and Renz, 1948: 27, pi. 10, figs. 2-2b.
Cohimbites heUeniciis Renz and Renz, 1947: 59,
74; Renz and Renz, 1948: 28, pi. 11, figs. 9-9c.
Cohimbites ex aff. phcatidi Smith, — Renz and
Renz, 1948: 23.
Prenkites kwan^siamis Chao, 1959: 130, 307, pi.
29, figs. 15-20, text-fig. 42b.
Cohimbites huangi Chao, 1959: 126, 301, pi. 29,
figs. 6-11, text-fig. 41a.
Cohimbites costatiis Chao, 1959: 126, 302, pi.
29, figs. 1-3, text-fig. 41b.
Cohimbites ijaUensis Chao, 1959: 126, 302, pi. 29,
figs. 12-14.
Thi.s i.s the "ornamented" species of
Prenkites. It has a conch essentially like
that of P. malsorensis except for constric-
tions and associated ribs which extend over
the flanks and project adorally as they
cross the venter. Welter ( 1922 ) had only
one specimen of this species and this is the
type; a second specimen is in the British
Museum (Natural History). The Sul)-
cohimbites fauna from Chios has yielded
eight specimens which I believe are con-
specific with the Timor form originally
Table 39. Measurements of Prenkites timor-
ensis Spath.
D
w
H
u
W/D
H/D
U/D
1.
45.2
22.8
23.0
15.5
50.4
50.9
.34.3
2.
45.2
?
17.4
16.0
?
38.5
.35.4
3.
42.0±
24.4
15.6
15.7
58. 1±
37. 1±
47.4±
4.
.38.5
22.7
15.3
13.5
59.0
.39.7
35.1
5.
36.5
21.7?
14.4
12.3
59.5?
.39.5
.33.7
6.
34.8?
24.7?
13.0
14.7
71.0?
37.4?
42.2?
7.
32.0
15.1
13.2
10.0
47.2
41.3
31.3
8.
29.1
20.5
12.8
8.2
70.4
44.0
28.2
6.
Paratv'pe,
(1948: pi
Holotv'pe,
(1948: pi
Paratope,
(1948: pi.
and
Holotype, Cohimbites httbulinae Renz and Renz (
pi. 10, fig. 1), NHMB J13.561.
Paratope, Cohimbites btibiilinae Renz and Renz (
pi. 9, fiK. 7), NHMB J13562.
malaijanus, — Renz
J13.55S.
mal-atjantis, — Renz
J135.5.5.
ntalayaniis, — Renz
.113557.
and
and
Cohimbites
9, fig. 9),
Cohimbites
9, fig. 6),
Cohimbites
9, fig. 8l,
Type specimen, Cohimbites malai/atms var. crassa
and Renz (1948: pi. 9, fig. 4), jl3549.
Holotvpe, Cohimbites heUeniciis Renz and Renz (
pi. li, fig. 9), J13.573.
ParaU'i^e, Cohimbites malaijanus, — Renz and
(1948: pi. 9, fig. 5), .113556.
1948:
1948:
Renz
Renz
Renz
Renz
1948:
Renz
described by Welter ( 1922 ) . Their mea-
surexnents are given on Table 39. Renz and
Renz (1948: 24) were not familiar with
Spath's publications ( 1930, 1934 ) in which
he introduced the name Prenkites timoren-
sis for Columhites sp. indet. Welter (1922:
150, pi. 168(14), figs. 12, 13). Columhites
malaijanus Renz and Renz was introduced
for a series of Chios forms that were be-
lieved to be conspecific with the Timor
Columhites sp. indet. of Welter. A second
species, Columhites huhulinae Renz and
Figure 26. Diagrammatic representation of the suture of: A, Prenkites mo/sorens/s Arthaber (1911: pi. 22(6), fig. 17c),
at G diameter of 20 mm; B, Prenkites malsorensis, — Renz and Renz (1948: pi. 12, fig. 12), at a diameter of approximately
15 mm; C, Prenkites helenae Renz and Renz (1948: pi. 12, fig. 5b), at a diameter of approximately 25 mm; D, Prenkites
timorensis, — Renz and Renz (1948: pi. 9, fig. 9d), at a diameter of approximately 27 mm; E, Prenkites timorensis, — Cfioo
(1959: fig. 42b), at a diameter of approximately 20 mm; F, Protropites liilmi Artfiaber (1911: pi. 22(6), fig. 16), at an un-
known diameter; G, Proco/umbites korataucikus Astakhova (1960a: fig. 9), at a diameter of approximately 20 mm; H,
Chioceras mitzopouloi Renz and Renz (1948: pi. 12, fig. 13), at a diameter of approximately 30 mm; I, Chioceras nodosum
Renz and Renz (1948: pi. 12, fig. 7c), at a diameter of approximately 20 mm; J, Arianites musacchi Arthaber, new su-
ture from holotype (PI. 2, figs. 9, 10), at a diameter of 20 mm; K, A4erope//a plejanae Renz and Renz (1948, pi. 3, fig.
4b); L, Ep/ce//ifes gentii Arthaber (1911: pi. 24(8), fig. 8d), at a diameter of 20 mm; M, Epiceltites subgracilis (Astakhova,
1960a: fig. 132), at a diameter of approximately 15 mm.
Specimens of figures A, F, J, L from Subco/umbifes fauna of Albania; specimens of figures B, C, D, H, I, K from Sub-
columbites fauna of Chios; specimen of figure E from Subco/umbifes fauna of Kwangsi, China; G, M from upper Scythian
horizon on the Mangyshlak Peninsula.
444 Bulletin Miiscmn of Comparative Zoology, Vol. 137, No. 3
Renz was introduced for slightly more
compressed forms. A third species, CoJiim-
bites helletiictis Renz and Renz is a slightly
aberrant form, compressed, with more
rounded umbilical shoulders and with a
more subdued pattern of ornamentation.
This species is based on a single specimen
and is considered here as falling within
the pattern of variation of P. fimorensis.
Cohimbites ex aff. pUcatuli, Renz and
Renz (1948: 23) is based on two fragmen-
tary specimens that have nothing in com-
mon ^^'ith Cohrmhites plicatuhis Smith
(1914: 37). They are actually more like
the forms Renz and Renz assigned to
Columhites hubidinae.
Prenkitcs kwan^,.siamts Chao (1959) was
established on three specimens from SiiJ)-
cohnnbifcs horizons in Kwangsi, China.
The descriptions and illustrations of this
species leave much to be desired. In spite
of this, I believe this species to be con-
specific with the forms from Timor and
Chios assigned to P. fimorensis. The in-
sight one can get on the variation within
this species from a study of the Chios
fauna lends support to the conclusion that
the differences in the ribbing used by Chao
to distinguish his species are not of specific
importance. The Kwangsi species of Co-
Jiimbitcs — Juianis,!, costafus, and yaliensis
— described by Chao ( 1959 ) occur together
at least in some outcrops and with Pren-
kites kwong,sianus at the Yali section.
These three species are clearly conspecific;
they differ only in the degree of ribbing.
These three forms are remarkably similar
to CoJumbiies Icvantiniis Renz and Renz
(1948: pi. 10, fig. 2) from the Siibcolum-
bites fauna of Chios. Analysis of the whole
Chios fauna suggests that Prenkites fi-
morensis is a highly variable species that
can and should include Cohimbites levati-
tiniis. These Kwangsi species (C. hiunii^i,
costafus, and yaliensis) are here considered
to be a variant similar to C. lecantinus
which belongs within the scope of /'.
timorensis.
The suture of ColiDnbites liuani:,i Chao
is shown on Figure 17C; Prenkites fimoren-
sis Spath is illustrated on Figures 26D, E.
Occurrence. Siibcolumbites fauna of
Chios; from block E, Nifoekoko, Timor;
from Suhcolumbifes fauna in the Linglo
and Fengshan districts of Kwangsi, China
(Chao ct)llections 542a, 546, 610); Pren-
kites aff. fimorensis is present in the Sub-
columbifcs fauna of the Primorye Region.
Repository. Holotype, Paleontological
Institute, Ronn; topotype RMNH C33714;
holotvpe, Cohimbites malayanus Renz and
Renz' (1948: pi 9, fig. 6)'nHMR J13555;
paratypes (pi. 9, fig. 9) NHMR J13558,
(pi. 9, fig. 8) NHMR J13557, (pi. 9, fig. 5)
NHMR J13556; var. crassa (pi. 9, fig. 4)
NHMR J13549, (pi. 9, fig. 10) NHMR
J13559; unfigured paratypes NHMR
J13546; holotype Cohimbites bubulinae
Renz and Renz (1948: pi. 10, fig. 1)
NHMR J13561; paratype (pi. 9, fig. 7)
NHMR J13562; unfigured paratypes from
Maradovuno NHMR J13563, from Kep-
halovuno NHMR J 13569; holotype Cohim-
bites helleniciis Renz and Renz (1948: pi.
11, fig. 9) NHMR J13573.
Genus Protropifes Arthaber, 1911
Type species, Proiropites hilmi Arthaber,
1911 (lectotype selected by Spath, 1934)
Involute, inflated forms \\'ith cadicone
inner whorls and carinate outer whorls.
With strongly prosiradiate growth lines,
occasionally enlarged on crossing the keel.
Sutme with single wide lateral lobe.
The type species is the only one known
for this genus and has been recognized
onlv in the Sitbcohimbitcs fauna of Kcira,
Albania. Spath (1934: 206) considered
Vrofropites to be an extreme development
of Subcohim])itcs mirdifensis. In this con-
clusion I concur. The rather poor preser\'a-
tion of the available material docs not
permit analysis of the ontogeny of the
only known species; however, in its gross
morphological features and the suture it
does appear to be a columbitid though its
exact relationships remain uncertain.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 445
Protropifes hilmi Arthaber
Plate 14, figures 1-8; Text-figure 26
Protropites hilmi Arthaber, 1911: 256, pi. 22(6),
figs. 9-16; Diener, 1915: 235; Renz, 1928:
155; Spath, 1934: 206, pi. 13, figs. 4a-c.
Arthaber's illustrations of the Subcolum-
hitcs fauna of Albania arc retouched photo-
graphs which on the whole are successful.
Arthaber illustrated six specimens plus a
suture (Fig. 26F) from an unspecified
specimen. Two of these specimens plus
the specimen which yielded the suture are
missing in the collection and presumed
lost. These are the specimens of figures
10, 11, and 16 of plate 22(6) in Arthaber
(1911). Tlie collections of the Paleonto-
logical Institute at the University of Vienna
contain approximately 20 specimens of the
species wdth no label as to origin, collector,
etc. It is presumed that these are part of
the original collection from Kcira, studied
by Arthaber. Unfortunately, the preserva-
tion of these specimens is uniformly bad,
none yielded any sutures, and none could
yield any useful measurements.
None of the four surviving primary
types studied by Arthaber are particularly
well preserved. The measurements of
these specimens are as follows:
D
W
H
U W/D H/D U/D
1. 33.2 ? 12.1 12.6 ? 36.4 38.0
2. 32.2 10.5? 11.9 12.8 32.6? 37.0 39.8
3. 29.2 10.2 11.4 10.8 34.9 39.0 37.0
4. 25.5 11.2 10.5 10.6 43.9 41.2 41.6
1. Lectotype, Arthaber (1911: pi. 22(6), fig.
15a, b).
2. Paralectotvpe, Arthaber (1911: pi. 22(6),
fig. 13a, b).
3. Paralectotype, Arthaber (1911: pi. 22(6),
fig. 12a, b).
4. Paralectotvpe, Arthaber (1911: pi. 22(6),
fig. 14a, b).
The lectotype (Pi. 14, figs. 3, 4) was
selected by Spath (1934: 206). Only the
venter and one side of the specimen are
preserved. The keel is very well developed
and present on the whole adoral volution.
Because of poor preservation, surface fea-
tures of the shell are obscure. Faint
patches of growth lines are present and
on the most adoral part of the last volution
there are two broad radial folds. The um-
bilical shoulder of the inner volutions
appears to bear small nodes, but these are
obscured by the poor preservation.
The largest of Arthaber's figured para-
lectotypes (Pi. 14, figs. 1, 2) is a more
compressed form with less inflated whorls.
Likewise only one side and the venter of
the specimen is preserved. The next para-
lectotype (PI. 14, figs. 5, 6) is of the same
preservation and is an even more com-
pressed form. The inner whorls are not
as depressed and cadicone as in the lecto-
type. The growth lines are in places well
preserved and show the strong forward
projection on the keel. Some of the growth
lines on crossing the keel are enlarged.
The specimen figured by Spath (1934: pi.
8, fig. 4a ) is comparable to this compressed
form. The fourth specimen (Pi. 14, figs.
7, 8) is the most inflated, with strongly
depressed cadicone inner whorls. It shows
the gradual contraction of the body cham-
ber very well. The conch is smooth except
for very faint and indistinct growth lines.
Occurrence. SuhcoJumhitcs fauna, Kcira,
Albania.
Rcpo.sitoiy. Lectotype, three figured
paralectotypes, and approximately 20 topo-
type specimens are in the Paleontological
Institute, University of Vienna. There are
eight topotype specimens in the British
Museum (Natural History) (C22838-45).
Genus Chioceras Renz and Renz, 1947
Type species, Chioceras mitzopouloi Renz
and Renz, 1947
Smooth, evolute conchs, inner \\'horls
rounded, slightly depressed, showing pro-
gressive expansion; outer whorls with con-
spicuous, smooth rounded keel; body
chamber slightly more than one volution
in length; suture with two denticulated
lateral lobes, third lobe on umbilical
shoulder, saddles rounded. One species
bears large lateral nodes.
This genus is known only from two
446 Biilk'tin Miiscuni of Comparative Zoology, Vol. 137, No. 3
species trom the Suhcohimhites fauna of
Chios. It is, however, fairly well repre-
sented in numbers of individuals in the
Chios fauna, as preserved in the Natural
History Museum in Basel.
The rounded keel immediately reminds
one of Protropiic's and at first these forms
were thought to belong to that genus.
However, the cadicone inner whorls of
Protropifes and the completely different
suture preclude any genetic relationship.
It is interesting to note that whereas Pro-
tropifes JiiJuii is fairly well represented in
the SuJ)cohimI)itcs fauna of Albania, it is
not present in the fauna from Chios.
The character of the suture appears to
be the best indication of genetic relations
and this suggests Prosphina^ites. The conch
can be looked upon as an evolute form of
Prosj)Jiin^ites with the marked addition of
the rounded keel.
Chioceras mitzopouloi Renz and Renz
Text-figure 26
Chioceras niitzoiiotiloi Renz and Renz, 1947: 60,
74; Renz and Renz 1948: 37, pi. 12, figs. 3-3b,
6-6a (hnlotvpe), 9-9a, 13; Kummel, m Arkell
et al., 1957:' L140, figs. 172, lOa-c.
Chioceras mitzopoaloi var. iiieridioiialis Renz and
Renz, 1947: 10; Renz and Renz, 1948: 38, pi.
12, figs. 8-8b.
This is the smooth species of Chioceras.
The measurements of the holotype and
figured paratypes are as follows:
D
W
H
U W/D H/D U/D
1.
49.0
?
16.6
19.7
?
33.9
40.2
2.
44.3
14.8
15.9
16.7
33.4
35.9
37.7
3.
43.5
13.7
13.1
18.4
31.5
.30.1
42.3
4.
33.9
13.8
10.5
15.5
40.7
31.0
45.7
5.
3().fi
10.8
10.7
13.0
35.3
35.0
42.5
1. Ilolotvpc, Renz aTid Renz (1948: pi. 12, fig.
6) NUMB .113626.
2. Paratype, Renz and Renz (1948: pi. 12, fig.
13) NHMB j 13627.
3. var. OTen'(:/K))u///.s Renz and Renz (1948: pi. 12,
fig. 8) NHMB .11.36.32.
4. Paratype, Reirz and Heiiz ( 1948: p\. 12, fig.
9) NUMB .113628.
5. Paratype, Renz and Henz (1948: pi. 12, fig.
3) NHMB .113629.
There are numerous unfigured j^aratypes
in the Chios collection in Basel that are
too poorly preserved or prepared to yield
useful measurements. The suture is shown
on Figure 26H.
Occurrence. SuhcolumI)ites fauna, Mara-
dovuno, Kcphalovuno, and Marmarotrap-
eza, Chios.
Repository. The figured specimens are
listed in the table of measurements; in
addition there are unfigured paratypes
from Maradovuno NHMB J13630, and
from Kcphalovuno NHMB J13631.
Chioceras nodosum Renz and Renz
Text-figure 26
Chioceras nodosum Renz and Renz, 1947: 60, 74;
Renz and Renz, 1948: 38, pi. 12, figs. 7-7c.
This species was established for a single
species that is like C. mitzopouhii except
for the presence of large, rounded nodes
on the lateral areas. The measurements
of the holotype are:
D W H U W/D H D U/D
.30.5 12.1 10.7 12.5 .39.7 35.1 41.0
The suture is illustrated on Figure 261.
Occurrence. Subcohimbites fauna, Mara-
dovuno, Chios.
Repository. Holotype NHMB J13633;
unfigured paratype NHMB J 13634.
Genus Arionites Arthaber, 1911
Type species, Arianites musacchi Arthaber,
1911
This genus and species are based on a
single poorly preserved specimen from the
Su])cohnnJ)ites fauna of Albania. I agree
with Spath (1934: 209) that it is mo.st
probabh' a columbitid.
Arianites musacchi Arthaber
Plate 2, figures 9, 10; Text-figure 26
Arianites musacchi Arthaber, 1911: 264, pi. 24(8),
fig. 5; Diener, 1915: 53; Spath, 19.34: 209;
Kummel, in Arkell el al., 1957: L14(), fig. 172,
14.
The type, and onl) specimen, of this
species is incomplete and generally poorly
preserxc'd. The last half xolution is body
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 447
chamber; the inner whorls are for the most
part l)uried in matrix. The outer volution
is completely smooth except for vague
traces of growth lines. A small portion of
the penultimate volution is visible, and
this shows sharp radial ribs on the flanks
and umbilical shoulder. A new drawing of
the suture is reproduced here on Figure
26J. Arthaber's drawing of the suture is
inaccurate, especially in his treatment of
the first lateral lol^e.
Occurrence. Suhcolumhites fauna, Kcira,
Albania.
Repository. Paleontological Institute,
Vienna.
Genus Meropella Renz and Renz, 1947
Type species, Arianites (Meropella) plejanae
Renz and Renz, 1947
Evolute conchs with rounded whorl sec-
tion of approximately equal width and
height dimensions. Ventral and umbilical
shoulders rounded. Early volutions bear
slight umbilical nodes.
The suture consists of two bifid lateral
lobes and a smooth, small auxiliary lobe
on the umbilical shoulder.
This genus and species is known only
from the Siibcolumbites fauna of Chios.
A fragmentary specimen from the Sul)-
columhites fauna of Kotal-e-Tera, Afgani-
stan, has been described by Kummel
(1968b) as Meropella cf. plejanae.
Meropella plejanae Renz and Renz
Plate 20, figures 14, 15; Text-figure 26
Arianites {Meropella) plejanae Renz and Renz,
1947: 67, 79; Renz and Renz, 1948: 95, pi. 3,
figs. 3-3b, 11-1 lb.
Meropella plejanae, — Knmmel, in Arkell et al.,
1957: L14(), fig. 172, 12.
There are only four specimens of this
species known. The measurements of
these are:
1.
2.
3.
4.
D
22.5
20.0
14.6
12.5
W
5.7
4.6
4.0
3.2
H
U W/D
5.8 11.8
5.7 9.8
4.0 7.3
25.3
23.0
27.5
H/D U/D
25.8 52.4
28.5 49.0
27.5 50.0
3.6
5.6 25.6 28.8 44.8
1. Figured paratype, Renz and Renz (1948: pi.
3, fig. 11) NHMB J 13826.
2. Holotype, Renz and Renz (1948: pi. 3, fig.
4) NHMB J 13825.
3. 4. Unfiguied paratypes, NHMB J13827.
One of the unfigured paratypes of the
Renz and Renz monograph is figured here
on Plate 20, figures 14, 15. Neither of
these small paratype specimens shows the
umbilical nodes as present in the holotype,
nor is the suture preserved. However,
aside from the lack of nodes, the remaining
features of the conch are identical to those
of the holotype. The drawing of the suture
(Renz and Renz, 194S: pi. 3, fig. 4b) is
accurate and is reproduced here as Figure
26K.
Occurrence. Suhcolumhites fauna, Mara-
dovuno, Chios.
Repository. Holotype NHMB J13825;
figured paratypes NHMB J13826 (Renz
and Renz, 1948: pi. 3, fig. 11), NHMB
J19550 (PI. 20, figs. 14, 15); unfigured
paratypes NHMB J13827.
Genus Epiceltites Arthaber, 1911
Type species, Epiceltites gentii Arthaber,
1911
Epiceltites genfii Arthaber
Plate 3, figures 10, 11; Plate 35, figures
6, 7; Text-figure 26
Epieeltites gentii Arthaber, 1911: 268, pi. 24(8),
fig. 8; Diener, 1915: 131; C. Renz, 1928: 155;
Kutassy, 1933: 510; Spath, 1934: 210, pi. 13,
figs. 5a-d; Renz and Renz, 1947: 60; Renz
and Renz, 1948: 43, pi. 1, figs. 9a-d.
Epiceltites n. sp. cf. E. gentii, — Kummel, 1954: 187.
Arthaber (1911) had five specimens of
this unique species of which only the holo-
type (PI. 3, figs. 10, 11) is still preserved
in the Paleontological Institute, Vienna.
The evolute, compressed conch with peri-
odic flares or constrictions that are strongly
projected forward on the venter makes
identification of this species easier than
with most species of Scythian ammonoids.
The holotype measures 36.3 mm in di-
ameter, 9.3 mm for the width of the last
whorl, 12.8 mm for the height of the last
448 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
whorl, and 14 mm for the diameter of the
umbilieus.
The Suhcolumbites fauna of Chios also
contains this species. The specimen figured
by Renz and Renz (1948: pi. 1, fig. 9)
measures 31 mm in diameter, 7.3 mm for
the width of the last whorl, 10.3 mm for
the height of the last whorl and 12.6 mm
for the diameter of the umbilicus.
The Thaynes Formation of southeastern
Idaho has yielded two incomplete and
poorly preserved specimens. One speci-
men (PI. 35, fig. 6) consists of only a
portion of one side of the conch, thoroughly
embedded in matrix; the other specimen
(PI. 35, fig. 7) shows only the venter and
part of the lateral area of one-half volution.
As no suture is preserved on either of the
specimens, there has been some hesitation
as to their specific identity. However, with
due consideration of the preservation and
incompleteness of the specimens, I feel
that the shape and involution of the conch
and the pattern of ornamentation are so
similar to the specimens of this species
from Albania and Chios that one must
assign these specimens to Arthaber's
species. Arthaber's suture is reproduced
on Figure 26L.
Occurrence. The holotype is from the
Suhcolumbites fauna of Albania. The
species is also known from that same fauna
from Chios, and from the uppennost mem-
ber of the Thaynes Formation, Hammond
Creek, Bear River Range, southeastern
Idaho.
Rcpositonj. Holotype, Paleontological
Institute, Vienna; topotypes BMNH C-
22867-74. Specimens from Chios, NHMB
J13657 (Renz and Renz, 1948: pi. 1, fig.
9); unfigured specimens NHMB J13658,
J 13659. Specimens from southeastern Idaho
MCZ 9470 (PI. 35, fig. 6), MCZ 9471
(Pi. 35, fig. 7).
Epicelfites subgracilis (Astakhova)
Text-figure 26
Anasihiritcs sttbgracilis Astakliova, 1960a: 147,
pi. 34, fig. 8, text-fig. 13.
The general shape etc. of the conch and
the suture ( Fig. 26M ) is very much like
that of the type species. The pattern of
ornamentations is sufficientlv different to
warrant separation.
Occurrence. From Columhites Zone of
Astakhova (1960a), Mangyshlak Penin-
sula.
Family USSURIIDAE Spath, 1930
Genus Parussuria Spath, 1934
Type species, Ussuria compressa Hyatt and
Smith, 1905
Parussuria latilobata Chao
Parmsuria latilobata Chao, 1959: 94, p. 260, pi. 31,
figs. 14, 15, text-fig. 31.
The few species that previously have
been assigned to this genus are of mid-
Scythian age; this is the first species from
a late Scythian horizon, and it is known
from only a single specimen. Chao (1959:
261 ) concluded that his species was most
closely related to Forussuria iwanovi
(Diener, 1895) from the Primorye Region.
In this I am in complete agreement; in
fact, these two specimens could well be
conspecific. Unfortunately, precise data on
the stratigraphic position of P. iuanovi are
lacking. This species was not in the faunas
described by Kiparisova ( 1961 ) from the
Primorye Region.
Occurrence. Limestone block in Lolou
village, Kwangsi, China (Chao collection
542b).
Family HEDENSTROEMIIDAE Waagen, 1 895
Genus Mefahedenstroemia Spath, 1934
Type species, Hedensfroemia kasfriotae
Arthaber, 1911
This is a very unsatisfactorily defined
genus because the type specimen is most
probably an immature form of onl\ modest
preservation. Spath (1934) considered the
suture to be the most unique aspect of
his new genus. The suture as illustrated
by Arthaber (1911) is highh idealized. As
with practicalK all the specimens from the
Suheoluni])ife.')- fauna of Albania and Chios,
Ammonoids of the Late Scythian (Lower Triassic) • Kummel
449
A
A
\.rv-
Figure 27. Diagrammatic representation of the suture of:
A, holotype of Metohedensfroemio (cosfriofoe (Arttiaber),
from tfie Subco/umbifes fauna of Albania, at a diameter
of 36 mm; B, fiolotype Hedenstroemia pityoussae Renz and
Renz (1948: pi. 16, fig. 8b), from tfie Subco/umb/fes fauna
of Cfiios, at a diameter of approximately 20 mm; C, fiolo-
type Beo//fes berthae Arthober (1911), from Subco/umbifes
fauna of Albania, new drawing at a diameter of 32 mm.
the suture can be exposed only by grind-
ing. In the case of the type specimen of
this genus the grinding has been excessive.
A new drawing of the suture, as exposed,
is shown on Figure 27A. The Chios species,
Hedenstroemia pityoussae Renz and Renz
( 1948 ) , differs from the type species of
Metahedenstroemia in details of the suture.
However, as I believe the differences are
due mainly to the results of specimen
preparation and are more apparent than
real, I consider the two species conspecific.
Tliis genus and species is known only from
the Subcolumbites fauna of Albania and
Chios.
Metahedenstroemia kastriotae (Arthaber)
Plate 14, figures 9, 10; Text-figure 27
Hedenstroemia kastriotae Arthaber, 1911: 208,
p. 17(1), figs. 14a-c; Diener, 1915: 148.
Metahedenstroemia kastriotae, — Spath, 1934: 223,
fig. 72; Kummel, in Arkell et al., 1957: LI 40,
fig. 173, 10.
Hedenstroemia pityoussae Renz and Renz, 1947:
61, 78; Renz and Renz, 1948: 83, pi. 16, figs.
8-8b.
Arthaber (1911: 208) had two speci-
mens of this species but only the specimen
Spath (19.34: 223) selected as the type
of his genus Metahedenstroemia is still
preserved in the Paleontological Institute,
University of Vienna. This specimen is all
phragmocone and measures 36.4 mm in
diameter, 22.0 mm for the height of the last
whorl, and 4.7 mm for the width of the
last whorl. It is illustrated here on Plate
14, figures 9, 10 and the suture on Figures
27 A, B.
The species Hedenstroemia pityoussae
described by Renz and Renz (1948: 83)
from the Suhcohimhites fauna of Chios is
based on three small fragmentary speci-
mens. These authors differentiated their
species on the basis of the suture; these
differences are, however, partly due to
preservation and preparation of the speci-
men.
Occurrence. Subcolumbites fauna of Al-
bania and Chios.
Repository. Holotype is in the Paleon-
tological Institute, Vienna; specimens from
Chios, holotype, NHMB J13791; unfigured
paratypes NHMB J 13792.
Genus Beatites Arthaber, 1911
Type species, Beatites berthae Arthaber,
1911
Highly compressed, involute form with
oxynote venter; sinuous growth lines.
Suture long with low goniatitic lobes and
saddles.
Beatites berthae Arthaber
Plate 21, figures 3, 4; Text-figure 27
Beatites berthae Arthaber, 1911: 210, pi. 17(1),
fig. 15; Diener, 1915: 66; Diener, 1917: 169;
Welter, 1922: 98; Kummel, in Arkell et al.,
1957: L142, fig. 173, 4.
The dimensions of the holotype (and
only specimen of this genus and species)
are: Diameter 32.3 mm, Width 3.7 mm,
450 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Height 16.0 mm, Umbilicus 5.8 mm. The
specimen is essentially all phragmocone.
The flanks of the penultimate half volution
have the wrinkle layer well presei-ved. The
suture has been exposed by grinding;
whether this has affected the details of
the lobes is not known. Arthaber's suture
drawing is not as accurate as it should be
and a new drawing is reproduced here on
Figure 27C.
Occurrence. Subcohimhitcs fauna, Kcira,
Albania.
Repository. Paleontological Institute,
University of Vienna.
Genus Lanceolites Hyatt and Smith, 1905
Type species, Lanceolites compactus Hyatt
and Smith, 1905
Until recently this genus has been known
only from the Meekoceras limestone of
Idaho, Nevada, and California. In addi-
tion to the type species. Smith (1932: 90)
has described one other (L. bicarinotus)
which is clearly closely related to the type
species and may be conspecific with it.
Neither of these species are very common
in the Meekoceras limestone of western
United States. Recently Ganev (1966) has
described L. discoidalis on four specimens
from the Campil Member of the Werfen
Formation of eastern Bulgaria. As these
forms are associated with a typical Werfen
tirolitid fauna, they are considered to be
of Frohun'^aritcs Zone age. This new record
considerably extends the range of the
genus.
Lanceo//7es discoidalis Ganev
Lanceolites discoidalis C.anv\\ 1966: 23, pi. 1,
figs. 1, 2, 4.
The conch shape of this species is nearly
identical to that of the species from the
Meekoceras limestone of western United
States. In this respect the Bulgarian
species is more similar to L. hicarinatus
than to L. compactus, which appears to have
a slightly more inflated conch. However,
so few specimens of an>' of these species
are known that there are no data available
on the amount of variability possible in
conch inflation. The suture of L. discoidalis
has the same basic pattern as that of the
American species but details are quite
different. Through the kindness of Dr.
Ganev I have plastotypes of his specimens.
I suspect, from studying these, that the
lack of fine denticulations in the suture is
the result of poor preservation and weath-
ering. At the same time, the suture is not
too unlike that of L. hicarinatus reproduced
by Smith (1932: pi. 55, figs. 3, 5, 7, 10),
or that of weathered specimens of L. com-
pactus reproduced bv Smith (1932: pi. 5,
fig. 9).
Occurrence. Campil Member, Werfen
Formation, perhaps as exotic blocks, Luda-
Kamcija region of eastern Bulgaria.
Repository. Primary types in Geological
Institute of the Bulgarian Academy of
Science; plaster casts are in the MCZ.
Family MEEKOCERATIDAE Waagen, 1895
Genus Svalbardiceras Frebold, 1930
Type species, Lecanites (?) spifzbergensis
Frebold, 1929
Svalbardiceras spifzbergensis (Frebold)
Plate 26, figures 1-4; Text-figure 28
Lecanites (?) spitzber^cnsis Frebold, 1929b: 299,
pi. 1, fig. 1; Kuta.ssy, 1933: 577.
Ammonites sp. iiidet. Frebold, 1929a: 14, pi. 1,
fiS. 12.
Annnonites sp. indet. Frebold, 1929ii: 15, pi. 1,
fig. 13.
Svalbardiceras s))Hz]}erfiensis (Freliold), 1930:
24, pi. 6, tigs. 1-3; Spath, 1934: 251, fig. 85;
Kummel, in Arkell et al., 1957: L142.
Interpretation of this species has becMi
in doubt amongst sexeral authors, mainh-
due to the uncertaint\' as to the nature
of the suture. The holotxpe is the speci-
men d(>scribed by Frebold in 1929 (1929b:
pi. 1, fig. J ) and not that figured by Fre-
bold in 1930 (pi. 6, fig. 1-la), as indicated
by Spath (1934: 251). The holotype
specimens came Irom Agardhberge on
Storfjord, Sjiitsbergen. The specimens de-
scribed 1)\ Frebold (1929a) as Aminonites
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 451
/N
Figure 28. Diagrammatic representation of the sutures of several species of Svalbardiceras. A, topotype of Sva/bard/'ceros
schmtdti (Mojsisovics), from Olenekites Zone, Olenek region, Siberia, at c whorl height of 13 mm (Popov, 1961: 39, fig.
6b); B, Svo/bord/ceros sp. indet., from Upper Thoynes Formation, Hammond Creek, southeastern Idaho, at a whorl height
of 15 mm (MCZ 9488); C, holotype of Svalbardiceras s/ieldon/ n. sp., from Co/umbifes fauna, Thaynes Formation, Draney
Creek, southeastern Idaho, at a diameter of 38 mm (MCZ 9493); D, Svalbardiceras spitzbergemis (Frebold, 1929a: pi. 1,
fig. 13; PI. 26, figs. 3, 4 of this report), from upper Scythian beds. Cape Thordsen, Spitsbergen, at a diameter of 37 mm.
sp. indet., which I believe to be conspecific
with the holotype, came from Cape Thord-
sen. The specimens described in 1930 by
Frebold came from Afj^ardh Bav, Milne
Edwardsberg, and Botneheia (Corrie
Down). All these specimens came from
what is interpreted as a late Scythian
horizon. The specimens from Cape Thord-
sen were associated with a KcyscrUn^itcs.
All of the known specimens of this
species show it to be an evolute form with
compressed whorls, higher than wide, and
with a flattened venter. The inner whorls
bear slight radial ribs which are strongest
near the mnbilical shoulder and decrease
down the flank; the weak ribs can be
observed throughout the phragmocone. On
the living chamber, the pattern of orna-
mentation is more irregular and is more like
irregular bundled growth lines. The full
details of the suture were not available to
Frebold, but on one of his specimens —
Ammonites sp. indet. (Frebold, 1929a: pi.
1, fig. 13; PI. 26, figs. 3, 4 of this report) —
it has been possible to develop the suture
(Fig. 28D). The suture bears two lateral
lobes that are clearly denticulated, and a
smooth auxiliary lobe on the umbilical
wall. The highlv crvstalline nature of the
phragmocone on many of the Spitsbergen
ammonites makes development of the
suture generally very difficult.
This Spitsbergen species of Svalbardi-
ceras is quite similar to the other species
of the genus but tends to be more involute
and with more development of an orna-
mentational pattern than the Siberian S.
scliniidti. The Idaho S. shcldoni has very
different whorl dimensions and a more
elaborate suture.
Occurrence. From presumably the up-
permost Scythian horizon at Agardh Bay,
Milne Edwardsberg, Botneheia (Corrie
Down ) , and Cape Thordsen, Spitsbergen.
Repository. The holotype was in the
Mineralogisch-Geologischen Staatsinstitut.
Hamburg, but was destroyed in the great
fire of 1943; the specimens described by
Frebold ( 1929a ) are in the Paleontologisk
Museum, Oslo; the specimens described by
Frebold (1930) are in the Geologisk In-
stitut, Uppsala.
S\/a\bard]ceras schmidfi (Mojsisovics)
Plate 26, figure 5; Text-figure 28
Xenodiscus schmidti Mojsisovics, 1886: 77, p\. 11,
figs. 8-11.
Gijronites mojsisovicsi Waagen, 1895: 297 (=:
452 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Xenodisciis schmidti Mojsisovics, 1886: pi. 11,
fijis. lla-c): Spath, 1934: 90, 251.
Piionolobus schmidti, — Noetling, in Freeh, 1905:
pi. 28, figs. 6a, b.
Mcekoceras (Ci/ronitcs) schmidti, — Diener, 1915:
197.
Meekoceras sp. indet. Frebold, 1929a: 13, pi. 1,
fig. 11.
Soalhardiccras spitzbcrgcnsis (Frebold) 1930: 24,
pi. 6, fig. 2.
Gijronitcs (?) schmidti, — Kiparisova, 1947: 134,
pi. 30, figs. 4, 5, text-fig. 16.
Svcdhaidiceras scJimidti, — Tozer, 1961a: 32.
Nordophiceras schmidti, — Popov, 1961: 39, pi. 12,
fig. 3.
Popov (1961: 39) recognized that his
Siberian species was very similar to Sval-
lyardiccras spitzheriien.sis (Frebold) but
on the argument that the suture of this
latter species was "unsatisfactorily defined"
and that the suture was unknown he felt
a comparison was not possible. It is true
that no drawing of a suture was presented
with the original description of Lccanitcs
(?) .spHzbcrgcnsis Frebold (1929b: 299,
pi. 1, fig. 1), but they are visible on the
photograph of the specimen. Frebold,
however, interpreted the lobes as being
goniatitic. The two specimens Frebold
(1929a: 14, 15, pi. 1, figs 12, 13) described
as Ammonites sp. indet. were considered
by their author as most probably closely
related to Leconites{?) spitzhergensis; I
believe them to be conspecific to that
species. One of these specimens does pre-
serve the suture ( Fig. 28D ) and this
clearly shows denticulated lobes; the gen-
eral pattern of the suture is identical to
that shown on the photograph of the holo-
type (Frebold, 1929b: pi. 1(36), fig. 1).
It does not, on this basis, seem justified to
disregard Svalhardiceras as a valid generic
name.
Acceptance of Sialbardiccra.s can and
does clarify the genus Nordopluceras
Popov ( 1961 ) from the Olenek region,
Siberia. Within this new genus, Popov
( 1961 ) included a variety of species de-
scribed by Mojsisovics ( 1S(S6) as XemHlis-
cus schtuidti, Xenodi.scus dentosus, Xen-
odisciis eiiom])hahis and the holotype.
Xenodisciis karpinskii; in addition, Popov
( 1961 ) described one new species, Nord-
ophiceras alexeevoe. Thus within his genus
Nordopluceras, Popov ( 1961 ) combined
what I consider to be a heterogeneous as-
semblage of species. The group includes
round ventered forms as illustrated by
Xcnodiscus karpinskii, the holotype of
Nordophiceras, and forms with truncate
venters as illustrated by Xenodisciis
schmidti and X. dentosus. It is this latter
group which is more properly allied to
the Spitsbergen species of Svalbardiceras.
Mojsisovics ( 1886 ) illustrated three
specimens of his Xenodisciis schmidti; the
principal variation observable is in the
sharpness of the ventral shoulders. Waagen
(1895: 297) renamed the form shown on
Mojsisovics' plate 11, figure 11a, b, as
Gyronites mojsisovicsi. This additional
name did not add clarity to the under-
standing of the group. Spath (1934: 251)
lists the specimen of Mojsisovics (1886: pi.
11, fig. Sa, b) as lectotype.
Of the species of Nordopluceras de-
scribed by Popov (1961), his specimen of
Nordophiceras scJimidti (Fig. 28A) is like
that of Mojsisovics (1886: pi. 11, fig 9) in
which the venter is slightly rounded but
the ventral shoulders still quite distinct. I
would, however, also include in this species
Meekoceras sp. indet. Frebold (1929a: pi.
I, fig. 11; PI. 26, fig. 5 of this report). This
specimen has the very flat venter with
angular \'entral shoulders shown by Moj-
sisovics' specimen of schmidti on his plate
II, figure 11. Likewise the abrupt umbili-
cal shoulder and the nature of the growth
lines are nearly identical. The suture is
unfortunately not preserxed. In addition,
the specimen of Svalbardiceras spitzbcr-
gcnsis (Frebold, 1930: pi. 6, fig. 2) should
be included. Both of these specimens dif-
fer from the other Spitsbergen spi>cimens
assigned to S. spitzhergensis in being more
involute and having a higher height-width
relationship in the whorl dimensions. This
is the primary difference between S. sch-
midti and S. spitzbcrgcnsis. The general
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 453
conch form of S. sheldoni is like that of S.
schmidti, but in this species there are con-
spicuous transverse ribs on the venter and
differences in the spacing of the suture ele-
ments.
Occurrence. Mojsisovics' species are only
listed as coming from the region of the
mouth of the Olenek River. Popov ( 1961 )
lists his specimens as from his Olenckites
Zone. Frebold's specimen of Meekoceras
sp. indet. is from Cape Thordsen, associ-
ated with Keyserlingites; his specimen of
Sva]I)(irdiccras spitzbergensis (Frebold,
1930: pi. 6, fig. 2) is from Milne Edward
Mountain.
Repository. The only specimen of this
species personally handled is Frebold's
Meekoceras sp. indet., and this is in the
Paleontological Museum, Oslo, Norway.
Svalbardiceras sibiricum (Mojsisovics)
Meekoceras sihirieinn Mojsisovics, 1886: 85, pi.
11, figs. 1-6; Spath, 1934: 224, 246, 254, 274,
341.
Aspidites sibirictis, — Freeh, 1905: pi. 28, fig. 11.
Meekoceras ( Koninckifes) sibiricum, — Diener,
1915: 198.
Meekoceras ? sibiricum. — Kiparisova, 1947: 150,
pi. 35, fig. 2.
Hemiprionites sibiricus, — Popov, 1962a: 176, 187,
pl. 2, fig. 3.
This species has been a puzzle to most
students of Triassic ammonoids. Tozer
(1965a: 37) suggested assignment of this
species to Svalbardiceras and I believe
there is much merit in this suggestion. The
umbihcal width of slightly less than 10 per-
cent the diameter of the conch makes this
the most involute species of the genus. As
already pointed out by Tozer ( 1965a : 37 ) ,
Popov's ( 1962a ) assignment of this species
to Hemiprionites cannot stand for many
reasons.
Occurrence. Olenek fauna, mouth of
Olenek River, northern Siberia.
Svalbardiceras dentosus (Mojsisovics)
Xenodiscus dentosus Mojsisovics, 1886: 78, pl. 11,
figs. 12a, b.
Gouiodiscus dentosus, — Diener, 1915: 135; Spath,
19.34: 330.
This species is like S. schmidti except for
long, low clavi along the sharp ventral
shoulders. It is based on a single specimen.
Occurrence. From near the mouth of the
Olenek River.
Svalbardiceras freboldi Tozer
Svalbardiceras freboldi Tozer, 1965a: 36, pl. 4,
figs. 12, 13, pl. 5, fig. 2, text-fig. 11.
With an umbilical width approximately
20 percent the diameter of the conch, this
is one of the more involute species of
Svalbardiceras. Aside from the slightly
greater involution, it is very similar to
schmidti and spitzbergensis.
Occurrence. Uppermost Scythian, Blaa
Moimtain Formation, lower shale member,
northern Ellesmere Island.
Svalbardiceras chov/adei Tozer
Svalbardiceras chowadei Tozer, 1965a: 37, pl. 4,
figs. 9-11.
This species could very well be con-
specific with S. spitzbergensis. The dif-
ferences are mainly in a slight difference
in the degree of involution. However, with
both species the number of specimens
known is so small that really no data are
available on the range of variability of any
of the morphological features.
Occurrence. Toad Formation, Halfway
River area, British Columbia.
Svalbardiceras sheldoni n. sp.
Plate 43, figure 1; Text-figure 28
The Cohtmbites fauna of southeastern
Idaho has yielded three specimens which
form the basis for this new species. The
two larger specimens are embedded in
matrix with only one side completely ex-
posed and a portion of the venter on the
phragmocone, and the body chamber ex-
posed. The body chamber is crushed. The
holotype specimen measures 70 mm in
diameter; the height of the adoral volution
is 30 mm and the umbilicus has a diam-
eter of 22.7 mm. The conch is evolute,
compressed, and essentially smooth. The
454 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
venter is flat, and aligned by angular
ventral shoulders. Tlie lateral areas are
broadly convex with the maximum breadth
at about mid-way between the ventral
and umbilical shoulders. The umbilical
shoulders are likewise sharply rounded and
the umbilical wall is vertical. The body
chamber occupies approximately two-thirds
of a volution. The shell bears fine growth
lines that are slightly prosiradiate. On
the phragmocone the venter has narrow,
shallow, transverse grooves that extend
from one ventral shoulder to the other.
The suture is shown on Figure 28C. It
consists of a wide ventral lobe, highly
denticulated, two lateral lobes, and a ser-
rated shallow lobe adjacent to the umbili-
cal shoulder. The large paratype has an
inner core of 59 mm in diameter of ex-
cellent preservation. Attached to it is a
portion of body chamber with a whorl
height of 39 mm. The specimen must
have had a diameter of at least 100 mm.
The small paratype has a diameter of 25.2
mm, and the whorl shape of the large
specimens. At a diameter of 4 mm the
venter is rounded; it appears that in the
following volutions the venter begins to
become flattened.
This species has a remarkable similarity
to Svalbardicems schmidti ( Mojsisovics,
1886: 77, pi. 11, figs. 11a, b), that is, in
the identity of the greatly compressed,
evoliite conch, and the flattened venter.
Svalhardiceras schmidti appears to have
a smooth venter, lacking transverse fur-
rows. The essential plan of the suture is the
same except that the ventral lobe lies
within the venter and does not spread out
on the lateral areas as in S. slicldoni. Moj-
sisovics noted a fair degree of variability
in the nature of the venter of his species,
some forms having rounded venters and
ventral shoulders. This observation has
been confirmed by Popov (1961).
The Spitsbergen species, S. spitzberii^en-
sis (Frebold, 1930: 24, pi. 6, figs. 1-3)
tends to be slightly more inflated than the
species described here. Likewise, the
lateral area bears weak radial ribs, widely
spaced on the inner whorls, but becoming
more bunched and sinuous on the body
chamber. The venter bears weak trans-
verse furrows.
Occurrence. From middle shale member
of Thaynes Formation {Columbites iauna) ,
on hillside north of Sage Creek, Stewart
Flat Quadrangle, and at Hot Springs,
southeast Idaho.
Repusiturij. Holotype MCZ 9493; para-
types MCZ'9643, 9644.
Svalhardiceras sp. indet. (S.E. Idaho)
Text-figure 28
Svalbardiceras sp. Kummel, 1954: 187.
This specimen is recognized on the basis
of a fragment of phragmocone consisting of
only five camerae. The whorl height is
14 mm, and the width is 7.3 mm. The
cross section of the whorl is compressed,
the lateral areas broadly convex. The
venter is slightly concave, measuring about
2 mm in breadth, and lined by angular
ventral shoulders. The umbilical shoulder
is broadly rounded. The suture is shown
on Figure 28B. The specimen recorded
here is much too fragmentary to compare
in a detailed fashion with the Spitsbergen
and Olenek species, but in its whorl shape
and suture it agrees well with the basic
conch pattern for the genus.
Occurrence. Upper member Thaynes
Formation, Hammond Creek, Bear River
Range, southeast Idaho.
Repositonj. MCZ 9488.
Svalbardiceras sp. indet. (Pakistan)
Svalburdiccia.s sp. indrt., KniuiiR'l, 196(i: 394, pi.
■1. litis. 6-9.
This identification was based on a com-
plete phragmocone and a fragment of an-
other. The specimens are complete enougli
to confidently assign them to the genus
Svalhardiceras but are not sufficiently com-
plete to make any meaningful comparisons
with ()th(>r species of the genus.
Occurrence. Narmia Member of Mian-
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 455
wall Formation, Nammal Gorge, Salt
Range, West Pakistan.
Repository. MCZ 9587, 9588.
Genus Stocheites KittI, 1903
Type species, Stacheites prionoides KittI,
1903
Compressed, involute forms with tabu-
late, often sulcate venter. Suture with
prominent first lateral lobe and shallow,
broad, second lateral lobe; first lateral
saddle narrow, second lateral saddle
broadly rounded.
The type specimens came from the
Werfen Formation in Dalmatia. The type
species, S. prionoides, is also stated to occur
in the Mangyshlak Peninsula in the Caspian
region, but unfortunately no specimens
from there have been described or illus-
trated (Astakhova, 1960b). A second
species, S. floweri, from the Subcohimhitcs
fauna of the Tobin Fonnation of Nevada
is described here. An indeterminate species
is known from the Nannia Member of the
Mianwali Formation in the Salt Range of
West Pakistan (Kummel, 1966); likewise,
two specifically indeterminate forms are
recorded from the Thaynes Formation of
southeastern Idaho.
Stacheites dionysi (Renz and Renz,
1948: 50) is here considered to be a
synonym of Metadagnoeeras tcrhunieum
(Arthaber). I agree with Spath (1934:
267) that Stacheites webhianus Diener
( 1907 ) from the Himalayan Anisian is a
completely unrelated stock to the Scythian
species discussed here.
Stacheites prionoides KittI
Plate 56, figures 9, 10; Text-figure 29
Stacheites prionoides KittI, 1903: 27, pi. 4, fig. 8;
Diener, 1915: 266; Spath, 1934: 267, fig. 92.
Kittl's original figure was a highly re-
constructed drawing with no view of the
venter. The type specimen is illustrated
here for the first time. It can be seen that
the specimen is a compressed form but
obviously crushed. The opposite side of
the conch is completely missing. The
Figure 29. Diagrammatic representation of the suture of:
A, Stac/ieites prionoides KittI (1903: pi. 4, fig. 8), at o
diameter of 33 mm, from Werfen Formation, Dalmatia;
B, S. Iloweri n. sp., composite suture from specimens MCZ
9439 and 9491, at a diameter of approximately 35 mm, from
Subco/umb/'tes fauna, Tobin Formation, Nevada; C, S. sp.
indet. II, at a diameter of 21 mm, from uppermost Thaynes
Formation, Sublette Ridge, western Wyoming (MCZ 9501).
venter is narrow, flattened and sulcate,
bordered by acute ventral shoulders. The
measurements of Kittl's type are as follows:
D
W
H
U
W/D
H,/D U/D
45.8
?
24.2
5.0
?
52.8 10.9
The suture is slightly weathered and the
ventral lobe could well be denticulated,
but is too weathered to preserve such
features. I consider Kittl's illustrations a
fairly good representation of this specimen.
Stacheites prionoides is morphologically
quite similar to the only other species of
the genus, S. floweri, from the Tobin
Fonnation of Nevada. The sutures espe-
cially are similar (Fig. 29). The shape
456 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Table 40. Measurements of Stacheites flow-
eri n. sp. from tobin formation, nevada.
D
w
H
u
W/D
H/D
U/D
1.
48.5
?
25.7
5.2?
?
52.9
10.6?
2.
40.5
?
20.8
5.5
?
50.2
13.2
3.
39.7
10.5
22.0
3.9
26.45
54.0
9.9
4.
31.7
8.2?
17.1
3.0
25.9?
53.9
9.48
5.
28.5
8.2?
16.7
2.6
28.75?
58.6
9.1
1. Paratvpe, MCZ 9440 (PI. 28, fig. 2).
2. Parat\pe, MCZ 9444 (PI. 28, fig. 8).
3. Holotvpe, MCZ 9441 (PI. 28, figs. 3, 4).
4. ParaUpe, MCZ 9445 (PI. 28, fig. 9).
.5. ParaUpe, MCZ 9443 (PI. 28, figs. 6, 7).
of the conch, degree of inflation and in-
volution are also strikingly similar. The
Nevada species, however, does bear faint
sigmoidal ribs on the flanks whereas S.
prionoidcs, as far as can be told, is smooth.
Astakhova ( 1960b ) records this species
from the Mangyshlak Peninsula; in fact
she used this as the name species of her
highest local zone. Unfortunately, no de-
scriptions or illustrations of these Mangy-
shlak specimens are available.
Occurrence. Werfen Fonnation, Muc,
Dalmatia.
Repository. The holotype and only
specimen of this species is in the Natural
History Museum, Vienna.
Siacheiies floweri n. sp.
Plate 28, figures 1-10; Text-figure 29
This is the second species of the genus
Stacheites to be recorded. The species is
represented by a fairly large number of
specimens, mainly fragmentary. The mea-
surements of five of the most complete
specimens are given on Table 40.
The conch is very involute and discoidal.
The venter is narrow, typically slightly
concave, and bordered by a fairly sharp
ridge. In some forms (e.g. PI. 28, figs. 6,
7) the venter loses its concavity, and the
adjoining ventral shoulders are rounded
and not angular; on the earlier volutions,
however, the fomner condition prevails.
The material available does not allow an
evaluation as to the nature and extent of
variations in the character of the venter.
The flanks are slightly arched, with the
maximum width in the dorsal third of the
flank. The umbilical shoulders are abruptly
rounded, and the umbilical wall vertical.
The flanks bear interesting low falci-
form ribs. These commence above the
umbilical shoulders as very low, nar-
row, radial ribs. At about the mid part of
the flanks, the ribs broaden greatly, and
inscribe a concave arc. Where the venter
is concave, it is perfectly smooth; on the
specimen ( PL 28, figs. 6, 7 ) where the
venter is not concave, there are faint
ridges crossing the venter. In some speci-
mens the falciform ribs are extremelv faint.
The suture consists of a fairly broad
first lateral lobe, a second lateral saddle
occupying about one-third of the width of
the flank, and a second lateral lobe that is
low, and with a characteristic pattern of
denticulation (Fig. 29B).
The basic features of the conch and
suture place this species morphologically
very close to S. prionoidcs. The falcifomn
ribs are the most obvious distinguishing
features.
Occurrence. Tobin Fonnation, Pershing
County, Nevada; south tip of Tobin Range,
Cain Mountain 1:62,500 quad., center NW
¥4 sec. 9, T. 26N, R. .39E, 5,500 ft. S, 27.5
ft. W from elevation point 5088 on range
crest.
Repositori/. Holotype, MCZ 9441 (PI.
28, figs. 3, "4); paratypes MCZ 9442 (PI.
28, fig. 5), MCZ 9443 (PI. 28, figs. 6, 7),
MCZ ^9444 (PI. 28, fig. 8). MCZ 9445 (PI.
28, fig. 9), MCZ 9446 (PI. 28, fig. 10),
MCZ 9439 (PI. 28, fig. 1), MCZ 9440 (PI.
28, fig. 2); suture specimen MCZ 9491;
unfigured paratypes MCZ 9500.
Stocheifes sp. indet. (S.E. Idaho)
Plate 37, figs. 7-10; Text-figure 29
Two spccimcMis from the Thaynes For-
mation of southeastern Idaho can be as-
signed to Stacheites. The first of these
specimens (PI. 37, figs. 7, 8) is a weathered
individual of oiiK' a (juarter volution. The
narrow concaNc \cnter, angular ventral
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 457
shoulder, slender compressed conch and
distinctive suture are characteristic of the
genus. The specimen is too incomplete to
allow a specific determination or com-
parison.
The second specimen (PI. 37, figs. 9,
10) is a small weathered individual of 33.6
mm in diameter that could well be con-
specific with Stacheitcs floweri of the
Tobin Fonnation of Nevada. The venter
is flat with angular ventral shoulders. The
lateral areas are broadly convex and
smooth. The adoral half of the specimen
is body chamber. The suture is shown on
Figure 29C. The suture is quite similar
to that of S. floweri except in the ventral
lobe, but the differences here could well
be the result of weathering.
Occurrence. The first specimen (PI. 37,
figs. 7, 8) came from the Upper Tliaynes
Formation, Hammond Creek, Bear River
Range, southeast Idaho. The second speci-
men (PI. 37, figs. 9, 10) came from a float
150-200 feet from top of upper calcareous
siltstone member of Thaynes Formation
(Kummel, 1954), Spring Canyon, Sublette
Ridge, western Wyoming.
Repositori/. MCZ 9487 (PI. 37, figs. 7,
8), MCZ 9501 (PI. 37, figs. 9, 10).
Sfacheites sp. indet. (West Pakistan)
Stacheites sp. indet., Kummel, 1966: 396, pi. 3,
fig. 13.
This record is based on a single, poorly
preserved specimen. The basic conch out-
line and pattern of suture generically
identify the specimen.
Occurrence. The single specimen came
from a five-foot limestone bed 38 feet
above the base of the Narmia Member of
the Mianwali Formation, Narmia Nala,
Surghar Range, West Pakistan.
Repository. MCZ 9609.
Genus Dagnoceras Arthaber, 1911
Type species, Dagnoceras nopcsanum
Arthaber, 1911
When Arthaber established this genus
he included within it a varied group of
species which are as follows:
Dagnoceras nopcsanum Arthaber
Dagnoceras zappanense Arthaber
Dagnoceras terhunicum Arthaber
Dagnoceras komanum Arthaber
Dagnoceras lejanum Arthaber
As Arthaber (1911) did not select a
type species, Diener (1915: 115) selected
D. nopcsanum. This selection was ap-
parently overlooked by Smith (1932: 65)
who selected D. komanum as type on the
principle that this was the first mentioned
species; this is, of course, an invalid selec-
tion.
Interpretation of the genus then rests
first on a consideration of D. nopcsanum.
The principal features are a moderately
involute conch, with rounded flanks, a
rounded umbilicus, and a suture with a
large first lateral lobe and small second
lateral lobe on or near the umbilical
shoulder. Arthaber's (1911) sutures of this
group were not accurately reproduced, and
new drawings of his type specimens are
offered here (Fig. 30). The sutures of all
these specimens are to a greater or lesser
extent altered by excessive grinding and
polishing. Dagnoceras nopcsanum and D.
zappanense are congeneric and actually
quite similar. These two species differ
mainly in the width of the first lateral lobe.
Dagnoceras lejanum is a synonym of D.
zappanense. Dagnoceras komanum is com-
pletely different in the aspect of the conch
morphology and suture and is considered
here to be a representative of AJhanites
triadicus. The specimen Arthaber (1911:
254, pi. 22(6), fig. 8) assigned to Fseudo-
sihirites cf. dichotomus Waagen is also
considered to be Albanifes triadicus. Fi-
nally, there is D. terhunicum, a species
with a narrow truncate venter and with a
more elaborate suture. This species is as-
signed to Metadagnoceras. This genus is
characterized by elaborate denticulation of
the first lateral saddle and first lateral lobe.
The conch is not too unlike that of a species
458 Bulletin Miiseutii of Comparative Zoology, Vol. 137, No. 3
4\
Figure 30. Diagrammatic representation of the sutures of species of Dognoceros. A, syntype of D. zopponense Arthaber
(1911: pi. 21(5|, fig. 8; PI. 15, figs. 3, 4 of this report), original drawing at a whorl height of 8 mm; B, syntype of
D. zappanense Arthaber (1911: pi. 21(5), fig. 9; PI. 15, figs. 5, 6 of this report), original drawing at a whorl height of
11.5 mm; C, syntype of D. iejanum Arthaber (1911: pi. 21(5), fig. 13; PI. 15, figs. 7, 8 of this report), original drawing at
a whorl height of 11 mm; D, holotype of D. nopcsanum Arthaber (1911: pi. 21(5), fig. 6; PI. 15, figs. 1 , 2 of this report),
original drawing at a whorl height of 9 mm; E, D. ellipticum Chao (1959: fig. 47a), at a whorl height of 10 mm; F,
D. latilobatum Chao (1959: fig. 47b), at a whorl height of 8 mm.
Specimens of figures A-D from Subco/umbites fauna of Albania; specimen of figure E from isolated block containing
typical upper Scythian fauna in Kwangsi, China; specimen of figure F from a Subco/umbi/es fauna in Fengshan District
of Kwangsi, China.
of Dciiinoccras, but some species do have
truncate venters, at least during a part of
their ontogeny. Metada^noccra.s appears
to be a vahd genus, distinct from DaiS,no-
ceras. Hov^ever, it needs to be emphasized
that the sutures of the two type specimens
of D. iw))csanum are affected liy grinding
and polisliing, and one cannot be sure as
to how much these factors have given the
suture its apparent simple pattern. None
of the specimens of D. nopcsanum in the
British Museum (Natural History) show
the suture.
In addition to Albania, Diiiinoccms is
known from a single specimen ( D. zap-
panense) from the Frohuniiarites faima of
Timor. The genus is known also from two
species from late Scythian faunas in
Kwangsi, C'hina. A fragmentar\' specimen
assigned to D. cf. zappanen.se has been
recorded from the Narmia Member of the
Mianwali Formation in the Surghar Range
of West Pakistan ( Kummel, 1966). It is
of interest to note that no species of this
genus has been recorded from the Suh-
cohnnhites fauna of Chios. Da'^noceras
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 459
(?) uniciim Kiparisova from the Primorye
Region is here assigned to Metadagnoceras.
The three species from the Mcekoceras
faima of sontheastern Idaho that Smith
(1932) assigned to Dciii^noceras (D. bonne-
villense, D. bridg,esi, and D. haijdeni) are
completely unlike the type species and
are excluded from this genus.
Dagnoceras nopcsanum Arthaber
Plate 15, figures 1, 2; Plate 16, figures
1, 2; Text-figure 30
Dagnoceras nopcsanum Arthaber, 1911: 241, pi.
21(5), figs. 6, 7; Diener, 1915: 115; Spath,
19.34: 268-271, pi. 8, fig. 1, text-fig. 93a;
Kummel, in Arkell, et al., 1957: L144, fig.
175, 1.
The two specimens of this species that
Arthaber figured are the only specimens
remaining in the original collection. They
have the following measurements:
D W H U W/D H/D U/D
Holotype 28.5 9.1 11.1 10.0 49.2 38.9 35.1
(Arthaber, 1911: pi. 21(5), fig. 6)
Paratype 36.3? ? 16.2 11.2 ? 44.6? 30.9?
(Arthaber, 1911: pi. 21(5), fig. 7)
The holotype is a specimen of only fair
preservation. The conch is evolute with a
broad and fairly deep umbilicus. The
whorls are oval in cross-section with a
narrowly rounded venter, broadly convex
lateral areas, an acutely rounded umbilical
shoulder, and a steep umbilical wall. The
greatest width of the whorls is at the
umbilical shoulder. The specimen is es-
sentially all phragmocone and devoid of
any ornamentation or growth lines. The
suture is shown on Figure 30D. As with
all the specimens from the Subcohimbites
fauna of Albania, the suture is revealed
only by grinding and polishing, and in
this case it is not possible to evaluate the
full extent of the damage. The paratype
has been so badly ground and polished that
the suture is essentially meaningless. The
holotype was prepared in slightly less
damaging fashion but there is grave un-
certainty as to whether the first lateral
saddle and the first lateral lobe are as
simple as shown on Figure SOD. The
character of this part of the suture is
critical for determining the relations of
this species to those species assigned to
Metada^noceras.
Of the five species established by Art-
haber from the Albanian Kcira fauna, only
the type, D. nopcsanum, and D. zappanense
are accepted as valid members of Dagno-
ceras. Dag,noceras nopcsanum differs
from D. zappanense mainly in the suture,
especially in the character of the first
lateral lobe (Fig. SOB, D).
Occurrence. Subcohimbites fauna, Kcira,
Albania.
Repository. Arthaber's two figured speci-
mens are in the Paleontological Institute,
University of Vienna.
Dagnoceras zappanense Arthaber
Plate 1 5, figures 3-1 1 ; Plate 24, figures
4, 5; Text-figure 30
Dagnoceras zappanense Arthaber, 1911: 241, pi.
21(5), figs. 8, 9; Diener, 1915: 115; Spath,
1934: 268-271, pi. 7, fig. 2, text-fig. 93d, e, f.
Dagnoceras cf. zappanense, — Kummel, 1966: 396,
pi. 3, figs. 9, 10.
Dagnoceras lejanum Arthaber, 1911: 242, pi.
21(5), figs. 12, 13; Diener, 1915: 115; Spath,
19.34: 269 (footnote), 271, text-fig. 93c.
The four specimens of this species
figured by Arthaber are available for
study. This species differs from D. nopc-
sanum mainly in the very slender first
lateral lobe (Figs. SOA, B). In addition,
there is some ornamentation. One speci-
men (PI. 15, fig. 11) has faint umbilical
nodes, and another faint sigmoidal ribs
(PL 15, fig. 6). The other two specimens
are smooth, which may be due to preserva-
tion or excessive preparation.
Arthaber distinguished between D. zap-
panense and D. lejanum on the basis of
degree of involution and ornamentation.
Both these characters are known to be
highly variable in most ammonite species,
and though the sample available is much
too small to see "gradations," it is felt
460 Bulletin Museum of Comparative Zoologij, Vol. 137, No. 3
that we are dealing here with a single
species complex.
The measurements of the four illustrated
types are as follows:
D
W
H
U
W/D H/D U/D
1. 42.7 12.1 17.5 13.3 28.3 40.9 31.1
2. 37.2 12.3 18.4 9.1 .33.1 49.5 24.5
3. 33.1 11.5 16.2 7.2 34.7 48.9 21.8
4. 30.1 10.7 13.7 8.5 35.5 45.5 28.2
1. Lectotype, D. lejaniiin Arthaber (1911: pi.
21(5), fig. 13).
2. Lectotype, D. zappanense Arthaber (1911: pi.
21(5), fig. 9).
3. Paralectotvpe, D. zappanense Arthaber (1911:
pi. 21(5), fig. 8).
4. Paralectotype, D. lejanuni Arthaber (1911:
pi. 21(5), fig. 12).
The sutures of these two species are like-
wise nearly identical (Figs. 30A, C).
Arthaber's drawings of the sutures for
these species are deceptive; his drawing
of the suture of D. zappanense ( Fig. 30B )
actually terminates on the umbilical
shoulder, while that for D. lejanum (Fig.
30C) terminates on the umbilical seam.
The specimen from Timor identified by
Spath (1934: 271, 272) as inseparable from
D. lejanum is figured here on Plate 24,
figures 4, 5. I completely agree with Spath
on this conclusion; the specimen, though
smaller, is nearly identical to the smaller
paralectotype of D. lejaninn illustrated here
on Plate 15, figures 9-11. A fragmentary
specimen of a form quite similar to this
species has been recorded by Kummel
(1966) from the Narmia Member of the
Mianwali Formation, Surghar Range, West
Pakistan, in association with Procarnites
kokeni.
Occunenee. Siibeohnnhites fauna, Kcira,
Albania, Alhanites fauna, Nifoekoko,
Timor; Narniia Member of Mianwali For-
mation, Surghar Range, West Pakistan.
Repository. The four types of Arthaber's
are in the Paleontological Institute, Uni-
versity of Vienna; the specimen from
Timor is BMNH C33713; the specimen from
West Pakistan is MCZ 9565.
Dagnoceras laiilobatum Chao
Text-figure 30
Dagnoceras latilobatiim Chao, 1959: 142, 322,
pi. 18, figs. 6-8.
This species was founded on two speci-
mens with a "narrow, subtruncate venter"
and "obtusely rounded" ventral shoulders.
The suture (Fig. 30F) is characterized by
a very large first lateral lobe. On the basis
of the data available, I believe it to be a
valid species of this genus.
Occurrence. From black, thick-bedded
limestone about 1.5 km north of Yali,
Kwangsi, China, associated with Siiheo-
lumbites, Hellenites, etc. (Chao collection
546).
Dagnoceras ellipticum Chao
Text-figure 30
Dagnoceras ellipticum Chao, 1959: 143, 323, pi.
18, figs. 3-5, text-fig. 47a.
This species is based on a single speci-
men which, however, is poorly illustrated.
From the data available, it is clearly a
species of Dagnoceras of the general type
of nopcsanum, the type species. It differs
from nopcsanum in its suture (Fig. 30E)
but not enough data are available to ana-
lyze the relationships.
Occurrence. Upper limestone bed, Nali-
ling section near village of Lolou, Linglo
District, Kwangsi, China, associated with
Hellcjiites, Prenkites, etc. (Chao collection
542a).
Genus Metadagnoceras Tozer, 1965
Type species, Metadagnoceras pulcher
Tozer, 1965
This genus is characterized by its large,
highK denticulated first lateral lobe and
narrow first lateral saddle, that may or
may not be denticulated. This basic plan
of the suture is like that of Daii.noeeras
except for the pattern ot denticulation.
T1k> suture of Dajinoceras nopcsanum is
known only from the type specimen, and
this sutur(> has been affected by grinding.
Takinii; this into consideration, the dif-
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 461
ferences between Dagnoceras and Meta-
dagnoceras in regards to the suture are not
all that great.
The genus Metadagnoccras includes the
following species:
xA/. pulcher Tozer
M. tohini n. sp.
M. iinicum (Kiparisova)
M. freemani Kummel
A/, terbiinictim (Arthaber)
Mefadagnoceras pulcher occurs in the
Toad Formation of British Columbia closely
associated with species of Isculitoides and
Keyserlingites; Metadagnoceras tohini is
from the Tobin Formation of Nevada,
associated with Suhcolumhites etc.; Meta-
dagnoceras unicum is from the Primorye
Region, presumably from a late Scythian
horizon (Cohimbites Zone); Metadagno-
ceras freemani is from the Prohungarites
fauna of Timor; and Metadagnoceras ter-
hunicum is from the Suhcolumhitcs fauna
of Albania and Chios.
This newly recognized genus is as yet
incompletely known; three of the species
{unicum, freemani, and pulcher) are known
from only one specimen, and the other
two species by a few specimens of only
modest preservation.
Metadagnoceras pulcher Tozer
Text-figure 31
Metadagnoceras pulcher Tozer, 1965a: 29, pi. 1,
figs, lla-e.
This unique species was based on a
single, well preserved specimen. It is
readily differentiated by its suture (Fig.
31A) and its delicate strigate sculpture.
Occurrence. Toad Formation, Halfway
River area, British Columbia.
Metadagnoceras tobini n. sp.
Plate 27, figures 1-4; Text-figure 31
This species is based on four fragmen-
tary specimens of fair preservation. The
conch is moderately involute with com-
pressed whorls. The lateral areas are
broadly arched, the venter low but rounded
and bordered by rounded but distinct
ventral shoulders. The umbilical shoulder
is more abruptly rounded and the umbilical
wall is nearly vertical. The flanks bear
sinuous growth lines and the living chamber
of one of the larger specimens has faint
indications of widely separated low radial
undulations. The venter on the two largest
specimens is unfortunately not preserved;
it is assumed that in these mature speci-
mens the venter is like that of the smaller
specimen illustrated on Plate 27, figures
1, 2.
The suture (Fig. 31B) consists of a
large first lateral lobe which has dentic-
ulations extending up the ventral side to
a narrow and irregular first lateral saddle.
There is a second lateral lobe on the
umbilical shoulder and wall. The second
lateral saddle is asymmetric and rounded.
In its conch shape and dimensions, Meta-
dagnoceras tobini is distinct from all the
other species of this genus. However, its
suture is nearly identical to the suture of
M. freemani (Fig. 31D).
Occurrence. Lower part of Tobin For-
mation, uses Locality M2562, Pershing
County, Nevada; south tip of Tobin Range,
Cain Mountain 1:62,500 quad., center NW
% sec. 9, T. 26N, R. 39E, 5,500 ft. S, 27.5
ft. W of elevation point 5088 on range
crest.
Repository. Holotype (PI. 27, fig. 3)
MCZ 9637; paratypes (PI. 27, figs. 1, 2)
MCZ 9638, (PI. 27, fig. 4) MCZ 9639;
unfigured paratype MCZ 9640.
Metadagnoceras unicum
Text-figure 31
Dagnoceras (?) unicum Kiparisova, 1961: 74, pi.
13, fig. 6, text-fig. 35.
This species was based on a single well
preserved specimen. In its general conch
morphology it differs from other species
of the genus in the great width of the whorl
in the area of the umbilical shoulder, in
the strongly converging whorl sides and
462 Bulletin Museum of Comparative Zoology, Vol. 137. No. 3
Figure 31. Diagrammatic representation of the sutures of species of A/lefodagnoceros. A, holotype of M. pulcher
Tozer (1965a: fig. 9), from British Columbia, at a whorl height of approximately 20 mm; B, paratype of M.
tobini n. sp., from Tobin Formation, Nevada, at a whorl height of 11 mm (MCZ 9638); C, M. sp., undescribed species
collected by N. J. Silberling from Star Peak Formation, Nevada, at a whorl height of 22 mm; D, holotype M. Ireewani
Kummel, from Nifoekoko, Timor, at a whorl height of 14 mm (BMNH C33701); E, holotype of M. lerbunicum (Arthaber),
from Subco/umbi/es fauna, Albania, at a whorl height of 18 mm; F, plesiotype M. terbunicum, — Renz and Renz (1948: pi.
1, fig. 7b), from Subco/umb//es fauna of Chios at a whorl height of 14 mm (Nf-IMB J13692); G, holotype of Sfacheifes
dionyii Renz and Renz (1948: pi. 1, fig. 6b), from Subco/umbi/es fauna of Chios, at a whorl height of 1/ mm (NHMB
J13689); H, M. unicum (Kiparisova, 1961: fig. 35), from late Scythian horizon, Primorye Region, Siberia, at a whorl height
of 1 9 mm.
the narrowly rounded venter. It.s suture tious in a dark siltstone together with
(Fig. 31II) i.s hkewise quite diflerent. Colmnhitcs )>ari.sianu.s. Tlii.s horizon is
Occurrence. Russki Ostrov, Cape i:)r()bably ec^uivalent to the Coliimhites
Sehmidt, Priniorve Region, from eonere- Zone of southeast Idalio.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel
463
Metadagnoceras freemani Kummel
Plate 24, figures 1, 2; Text-figure 31
Metadagnoceras freemani Kummel, 1968a: 11, pi.
1, iiiis. 12, 13.
This species is proposed for the speci-
men from Timor mentioned by Spath
(1934: 269, footnote) as "an isolated ex-
ample of a new species from Timor . . .
which is very close to Dagnoceras terbuni-
cum." The specimen measures 47.8 mm
in diameter, 14.7 mm for the width of
the last whorl, 21.7 mm for the height of
the last whorl, and 10.4 mm for the diam-
eter of the umbilicus. The conch is com-
pressed with a low, arched venter and
rounded ventral and umbilical shoulders.
The suture (Fig. 31D) has a very large
first lateral lobe with denticulations ex-
tending all along the ventral side to a
narrow irregular first lateral saddle. There
is a small denticulated second lateral lobe
on the umbilical shoulder and wall.
This species does not resemble Dagno-
ceras tcrbuniciim in conch shape; the
differences are centered mainly in the
character of the venter. In the suture the
first lateral saddle and the first lateral lobe
of the two species are quite similar (Figs.
31D-F) but the remainder of the suture is
quite different. In general conch morphol-
ogy Metadagnoceras freemani is quite
similar to the type species, M. ))ulcher
Tozer. The sutures are similar in basic
plan but differ in significant details. The
suture of M. tohini n. sp. is very nearly
identical to that of M. freemani, but in
the Nevada species the whorls are of quite
different proportions and the conch much
more evolute.
Occurrence. Nifoekoko, Timor, from bed
with manganese coated fossils, including
Albanites, Prohungarites, etc.
Repository. BMNH C33701, holotype.
Metadagnoceras terbunicum (Arthaber)
Plate 16, figures 7, 8; Plate 18, figures
9, 10; Text-figure 31
Dagnoceras terbimiciim Arthaber, 1911: 242, pi.
21(5), figs. lOa-c; Diener, 1915: 115; Spath,
1934: 269; Rcnz and Renz, 1947: 60; Renz
and Renz, 1948: 51, pi. 1, figs. 7-7b.
Meekoceras radiosum, — Arthaber (non Waagen)
1911: 246, pi. 21(5), fig. 14; Diener, 1915:
194.
Dagnoceras aff. ierhunico, — C. Renz, 1928: 155.
Dagnoceras nopcsanum Arthaber var. involuta
Renz and Renz, 1947: 60; Renz and Renz, 1948:
52, pi. 1, figs. 5-5a.
Stacficites dionysi Renz and Renz, 1947: 60, 75;
Renz and Renz, 1948: 50, pi. 1, figs. 6-6b,
8-8b.
When Arthaber proposed his genus
Dagnoceras, he included within it a hetero-
geneous group of species. One of these
was D. terbunicum, characterized by a flat
venter and angular ventral shoulders, which
is in striking contrast to the rounded venter
of the more typical species. Arthaber
(1911: 240) recognized the similarity of
his new generic group to Stacheites but
was uncertain of the character of the
venter of Kittl's S. prionoides. My own
examination of S. prionoides clearly estab-
lishes that Kittl's type specimen has a
slightly sulcate venter with angular ventral
shoulders. The basic morphological fea-
tures of the conch of Dagnoceras terbuni-
cum do suggest, at first, the possibility of
this species being a more inflated repre-
sentative of Stacheites. Tlie basic plan of
the suture, however, is more like that found
in Metadagnoceras than that found in
Stacheites. Arthaber's (1911: pi. 21(5),
fig. 10c) representation of the suture is
misleading. A new drawing of the suture
from this type specimen is shown here on
Figure 31E. There is a large first lateral
lobe, with prominent denticulations on the
base of the lobe and extending up on the
ventral side of the lobe to a narrow, ir-
regular first lateral saddle. There is a
small second lateral lobe above the umbili-
cal shoulder that apparently has a few
denticulations which are, however, indis-
tinct because of excessive grinding; this is
followed over the umbilical shoulder and
wall by an irregular series of large dentic-
ulations. The character of the first lateral
saddle and first lateral lobe is very much
Hke that of the suture of M. tobini, M.
464 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
freemani, and the specimen from Nevada
collected by N. J. Silberliiig from the base
of the Star Peak Group, Humboldt Range,
Nevada (Fig. 31C). It differs from these
species in the greater elaboration and
length of the remainder of the suture.
The measurements of the holotype of D.
terbunicum are: Diameter 43.0?, Width
12.0?, Height 21.8, Umbilicus 7.8 mm (the
second specimen of this species mentioned
by Arthaber is not in the collection of the
Paleontological Institute, Vienna, and is
presumed lost).
With some misgivings I include in M.
terbunicum, Meekoccras mdiosum, — Artha-
ber (non Waagen). This species has the
same general shell form as M. terbunicum
( compare PI. 16, figs. 7, 8 and PI. 18, figs.
9, 10) except for being more involute. The
dimensions of this specimen are: Diameter
45.0?, Width 13.0?, Height 24.0?, Umbilicus
3.6? mm. The suture had been exposed by
grinding and the pattern reproduced by
Arthaber is not at all a correct representa-
tion. One can never be absolutely sure in
cases like this but it appears that the suture
is actually like that of the holotype of M.
terbunicum. Thus on the basis of conch
form and probable nature of the suture,
this species is considered a synonym of
M. terbunicum.
In the fauna from Chios, the Renzes
(1948: 51) record one specimen as being
conspecific with D. terbunicum Arthaber.
The measurements of this specimen ( Renz
and Ren/, 1948: pi. 1, figs. 7-7b) are as
follows:
D W H U W/DH/DU/D
NHMR .113692 42.0 14.2 20..3 8.3 33.8 48.3 19.8
In addition, the Renzes assigned another
specimen to D. no})csantim Arthaber var.
involuta Renz and Renz ( 1948: 52, pi. 1,
figs. 5-5a). The measurements of this spe-
cies are as follows:
D W II U W/DH/13U/D
NHMB Ji3f)94 28.8 10.7 13.7P7.2 37.2 47.6?25.0
The Albanian s[)ecimens of D. nopcsa-
num have rounded venters, the Chios speci-
men has a truncate, flattened venter and is
merely a smaller specimen of terbunicum.
The Chios collection contained two
specimens which the Renzes assigned to a
new species of Stacheites — S. dionysi. The
measurements of these two specimens are
as follows:
D W H U W/DH/DU/D
NHMB J13689 34.8 11.4 16.4 8.1 32.8 47.1 23.3
(Renz and Renz, 1948: pi. 1, fig. 6)
NHMB J13690 45.8 ? 21.2 10.4 ? 46.3 22.7
(Renz and Renz, 1948: pi. 1, fig. 8)
The conch form of S. dionysi is identical
to that of Metadagnoceras terbunicum; the
only difference between these forms is in
the slightly greater individualization of
the second lateral lobe (Fig. 31G), a dif-
ference I can hardly consider of specific
significance and surely not of generic rank.
Preparation of the suture in these speci-
mens preserved in hard red limestone is
very difficult, and is generally done b\
acid etching or grinding. Also, with only
four specimens assigned to these three dif-
ferent species (and two genera), it is
hardly justifiable to put such importance
on minor differences in the suture. All
three of the species are considered to be
conspecific with Metadagnoceras terbuni-
cum (Arthaber).
The Albanian and Chios specimens
assigned to terbunicum have flattened
venters. Silberling ( personal communica-
tion) has a specimen from the Star Peak
Formation, Nevada, that also has a flat-
tened venter. This specimen is an in-
complete phragmocone of approximately
44 mm in diameter. The other species of
Metadagnoceras have low, arched venters
to more highly vaulted venters. Consider-
ing the state and nature of preservation
of the Albanian and Chios specimens as-
signed here to tcrl)unicu}n, I belie\e they
are all incomplete and immature specimens.
It is, of course, uncertain as to what the
character of the venter is like on a mature
indi\idual. As it is now interpreted, in
Ammonoids of the Late Scythiax (Lower Triassic) • Kiimmel
465
conch form this species is convergent to
Stacheites but differs significantly in its
suture.
Occurrence. Subcohimbites fauna of Al-
bania and Chios.
Repositori/. Arthaber's (1911) figured
type (pi. 21(5), fig. 10; PI. 16, figs. 7, 8 of
this report) is in the Paleontological In-
stitute, University of Vienna. A second
specimen indicated by Arthaber (1911:
242) is apparently lost. The Chios speci-
mens are in the Natural History Museum,
Basel, and are as follows: holotype Dag-
noceras nopcsanum var. invohita Renz and
Renz (1948: pi. 1, fig. 5) NHMB J13694;
unfigured specimen NHMB J 13833; holo-
tvpe Stacheites cUonijsi Renz and Renz
('l948: pi. 1, fig. 6) NHMB J13689; para-
type (pi. 1, fig. 8) NHMB J13690;
unfigured paratypes NHMB J13691; plesio-
type Dagnoceras terbiiniciim (pi. 1, fig.
7) NHMB J13692; unfigured specimens
NHMB J13693.
Genus Balkanifes Ganev, 1966
Type species, Balkanifes fabulafus Ganev,
1966
Conch compressed with flat lateral areas
and broad flat venter. Ventral shoulders
weakly rounded. Umbilicus small, about
20 percent the diameter of the conch.
Umbilical wall nearly vertical, umbilical
shoulders weakly rounded. Conch smooth
with no ornamentation. Suture goniatitic
consisting of a single large, smooth, lateral
lobe with indication of the beginning of a
second lobe at the umbilical shoulder.
Gane\' did not include a drawing of the
suture in his report but he kindly sent me
a photo of the specimen, plus a plaster
cast which clearly shows the suture.
This is indeed a unique Scythian am-
monoid and is an addition to the fairly
large number of endemic genera which
characterize the Werfen fauna. Ganev
allied his new genus to Dognoceras pri-
marily on the basis of the single large
lateral lobe. In this relationship I concur
as being the most logical on the basis of
the data available. In Dagnoceras the
venter is rounded and the lobe denticulated.
Balkanifes fabulafus Ganev
Balkanites tabulatus Ganev, 1966: 24, pi. 2, figs,
la-d.
Discussion of this species is given above.
Occurrence. Campil Member of Werfen
Formation, perhaps as exotic blocks, Luda-
Kamcija region of eastern Bulgaria.
Repository. Holotype (and only speci-
men ) in Geological Institute of the Bul-
garian Academy of Science; plaster cast
in MCZ.
Genus Nordophiceras Popov, 1961
Type species, Cerafifes euomphalus Key-
serling, 1845
Nordophiceras euomphalus (Keyserling)
Plate 47, figures 6—8; Text-figure 32
Ccratites euomphalus Keyserling, 1845: 171, pi. 3,
figs. 7-9; Middendorff, 1860: 248, pi. 3, figs.
7-9; Eichwald, 1868: 1039.
Meekuceras ciiomphalitm, — Mojsisovics, 1882:
214; Waagen, 1895: 239, 246; Diener, 1915:
191.
Xenodiscus ciiunipfialtis, — Mojsisovics, 1886: 76,
pi. 11, fig. 7.
Nordophicera.'i ciuniipluihis, — Popov, 1961: 39.
Xenodiscus kuipiiiskii Mojsisovics, 1886: 75, pi.
11, fig. 13.
Meekoccras {Gi/ronites} karpinskii, — Diener, 1915:
196.
Nordophiceras karpinskii, — Popov, 1961: 41, pi.
25, fig. 6; Vozin and Tikhomirova, 1964: 50,
pi. 27, fig. 2.
Nordophiceras contratius Popov, 1962a: 177.
Meekoceras contrariiis Popov, 1962a: 185, pi. 3,
fig. 7, text-fig. 8.
Meekoceras? contrarium, — Vozin and Tikhomirova,
1964: 55, pi. 30, fig. 3, text-fig. 9a.
Popov ( 1961 ) selected Xenodiscus karp-
inskii Mojsisovics (1886: 75, pi. 11, fig.
13) as the type species of his genus
Nordophiceras. At the same time he in-
cluded within that genus Ceratites euom-
phahis Keyserling (1845: 171, pi. 3, figs.
7-9). The latter species differs from X.
karpin.skii in having a slightly more inflated
whorl section. The author has available
four topotype specimens of Nordophiceras
466 BuUetin Museum of Coniparativc Zoology, Vol. 137, No. 3
Figure 32. Diagrammatic representation of the suture of: A-C, Nordophiceras euompho/us (Mojsisovics), from Dieneroceras
Zone, norffiern Siberia; A, topotype, — Popov |196f: 39, fig. 6e), at a wfiorl fieight of 8 mm; B, topotype (MCZ 8670a),
at a diameter of 50 mm; C, topotype (MCZ 8670b), at a diameter of 26 mm; D, liolotype Nordophiceras contrarius (Popov
1962a; fig. 8), Dieneroceras Zone, northern Siberia, at a whorl height of 20 mm; E, hoiolype Nordophiceras o/exeevoe
Popov (1961: 39, fig. 6c), from Dieneroceras Zone, eastern Taymyr, Siberia, at a whorl height of 15 mm; F, Nordophiceras
jacksoni (Hyatt and Smith), from Columbites fauna, Montpelier Canyon, southeast Idaho (MCZ 9572), at a diameter of
36 mm; G, paratype of Prionolobus jacksoni Hyatt and Smith (1905: pi. 62, fig. 16), from Columbites fauna, Paris Canyon,
southeast Idaho, at a diameter of 45 mm (USNM 75292c); H, paratype Mee/toceros (Submee/coceros) compressum Chao (1959:
fig. 46a), from Subcolumbifes fauna, Kwangsi, China, at a whorl height of 15 mm; I, holotype Meekoceras (Submee/toceras)
Ammonoius of the Lath Scythian (Lower Triassic) • Kummel
467
kaiphi.skii identified by Dr. Popov. Ex-
amination of these specimens and of the
ilhistrations in Mojsisovics (1886: pi. 11,
fig. 7) clearly shows that these two so-
called species cannot be distinguished. The
combining of these t\\'o species thus makes
cuomphalus the type species of Nordophi-
ceras.
Two of the Siberian topotypes in the
collections of the Museum of Comparative
Zoology are illustrated here on Plate 47,
figures 6-8. The conch is moderately in-
volute, compressed, with a rounded venter.
The flanks are broadly convex and the
umbilical shoulders rounded. The suture
consists of t\\'0 lateral lobes and a serrated
lobe on the umbilical shoulder and wall.
The sutures of t\\'o of the topot\'pes in the
collection and that reproduced by Popo\-
( 1961: fig. 6e) are shown on Figure 32A-C.
The conch is smooth except for sinuous
growth lines.
Associated with this species in the
Olenek fauna is a closely related species,
N. olexeevae Popov, which differs in the
presence of closely spaced fine, prosirad-
iate ribs on the inner whorls and more
widely spaced, thin ribs on the outer
whorls. In nearly all other aspects these
two species are identical. A much closer
relationship exists with Noiclo))Juccros
jacksoni (Hyatt and Smith) from the
Columbites fauna of southeastern Idaho.
The basic plan of the smooth conch and
the suture is remarkably similar. The Idaho
fauna has yielded a fairly large number of
specimens which gives some appreciation
of the variations in conch proportions ( Fig.
33); this kind of data is lacking for the
Siberian species described here. It is very
possible that study of a large collection
of the Siberian N. etiomphahis could estab-
lish that N. j(icks())ii is a synonym; for the
moment at least it seems best to keep the
forms separate. It is because of this very
close relationship with N. jacksoni that I
believe Meekoceras contrarius is a synonym
of N. cuomphulus. The main difference in
these species lies in the degree of involu-
tion. However, the difference in umbilical
diameters lies well within the variation of
this parameter in the populations of N.
jacksoni from southeast Idaho. The other
species of ISlordophiccras in the Columbites
fauna of Idaho are ornamented forms more
related to N. aJexeevae.
This species is also quite similar to Nord-
ophiccras planorhis (Waagen) (Kummel,
1966: 397) from the Salt Range of West
Pakistan from an upper Scythian horizon.
The basic architecture of the conch is
most similar; however, the Salt Range
species is known from very few and gen-
erally poorly preserved specimens.
Occurrence. Mojsisovics' specimens are
from the region of the mouth of the Olenek
River, those described by Popov ( 1961 )
are from his Dieneroceras Zone in eastern
Taymyr, Chernokhrebetnaya River.
Rcpositon/. Topotvpe specimens MCZ
9655 (PI. 47, fig. 6)', MCZ 8680 (PL 47,
figs. 7, 8), MCZ 6107, 9656.
Nordophiceras a/exeevae Popov
Text-figure 32
Nordophiceras cdexeevae Popov, 1961: 39, pi. 25,
fig. 7.
Nordophiceras olenekensis Popov, 1961: 40, pi.
12, fig. 8.
This species from northern Siberia is of
particular interest because it is another
fonn closely related to a species in the
Columbites fauna of southeastern Idaho.
In fact, because the Idaho fauna has vielded
/o/ouense Chao (1959: fig. 45d), from Subcolumbites fauna, Kwangsi, China, at a whorl height of 12 mm; J, holotype
iMee/<oceros (Submeekoceras) longiseptatum Chao (1959: fig. 46c), from Subcolumbites fauna of Kwangsi, at a whorl height
of 22 mm; K, holotype Nordophiceras olene(:ens/s Popov (1961:40, fig. 6d), from Dieneroceras Zone, northern Siberia, at a
whorl height of 12 mm.
468 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
a fairly large number of specimens, the
data on that species ( N. pilotiim ) have
aided greatly in the interpretation of this
Siberian species. It should be pointed out
that Popov had available only one speci-
men of A^ alcxeevac and six specimens
of N. oJenekcnsis.
This species is essentially of the same
conch morphology, that is in compression,
involution, etc., as the associated N. euom-
pJiahis, except that on the inner wliorls
there are fine, closely spaced prosiradiate
ribs which on the outer whorls are more
widely spaced. The two species of Popov
differ only in the degree of this ornamen-
tation. Nordophiceras pilatum from the
Cohimhites fauna of southeastern Idaho
has a similar pattern of ornamentation. The
large number of specimens in the Idaho
collection clearly show a broad range of
variation in the degree of intensity of
ornamentation. The suture is illustrated on
Figure 32E, K.
Occurrence. Dieneroceras Zone of Popov
(1961), eastern Taymyr and the delta of
the Lena River, northern Siberia.
Nordophiceras pseudosimplex n. sp.
Arctocera.s simplex, — Popox- ( iion Mojsisovics ),
1961: 67, pi. 18, fiu. 1; Vozin and ^rikhoiniiova,
1964: 55, pi. 29, fiii. 1.
Arctoceras simplex Mojsisovics ( 1<SS6 )
was established for specimens which are
the inner, juvenile whorls of Arctoceras
hlomstrandi (Lindstrom), and is of mid-
Scythian Oicenites Zone age ( Kummel,
1961). The specimen Popov (1961) as-
signed to Arctoceras simplex is not at all
comparable. Popov's specimen is more
evolute, has flattened, more or less par-
allel, lateral areas. Popov ( 1961 ) states
his specimen has smooth inner whorls but
that one specimen had widely spaced
radial ribs which disappeared on the body
chamber. The ribs arc^ not apparent on
Popov's illustration. The suture is said to
be c{"ratitic but this is not borne out b\
the published text-figure.
Needless to say that much more data
are needed, but we can be sure that this
specimen is not conspecific with Arctoceras
simplex but rather is allied to such forms
as Nordophiceras euomphalus from the
underlying Dieneroceras Zone of Popov. It
is, likewise, quite similar to Nordophiceras
planorhis (VVaagen) from the Nannia
Member of the Mianwali Formation in the
Salt Range.
Occurrence. Olenekites Zone in basin of
Olenek River, northern Siberia.
Nordophiceras planorbis (Waagen)
Lccanitcs planorbis Waagen, 1895: 278, pi. 39,
fig. 3.
Mcckoccras (Gyionitcs) planorhis. — Diener, 1915:
197.
Nordophiceras planorhis, — Kuniniel, 1966: 397,
pi. 4, figs. 1-3.
New illustrations and descriptions of the
type specimens and two topotype speci-
mens have been published by Kummel
( 1966 ) . This is the only record of Nor-
dophiceras {ox the Tethyan region.
Occurrence. Narmia Member, Mianwali
Formation, Chhidru Nala, Salt Range, West
Pakistan.
Repositon/. Holotvpe GSI 7226; topo-
types MCZ"9611, 9612.
Nordophiceras jacksoni (Hyatt and Smith)
Plate 47, figures 1-5; Plate 48, figures
1-4; Text-figures 32, 33
Priouolohiis jacksoni Hvatt and Smith, 1905: 151,
pi. 151, figs. 11-21:' Kraltt and Diener, 1909:
to, 41.
Mcckoccras iacksoni. — Diener, 1915: 192.
Oj)hiccras jacksoni, — Snn'tli, 1932: 49, pi. 62,
figs. 11-21.
This species is abundanth' represented
in the (U)lund)ites beds exposed around
the north end of Bear Lake, southeastern
Idaho. The conch is smooth, CNolute,
compressed, with a rounded x'cnter. The
measurements of 4(S specimens are given
on Table 11 and plotted on Figure 33.
ihese data show that there is only a
moderate amount ol xariation in exolution
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel
469
Table 41. Measurements of Nordophiceras jacksoni (Hyatt and Smith) from Columbites
FAUNA, Bear Lake region, southeastern Idaho.
D
w
H
u
W/D
H/D
U/D
D
w
H
U
W/D
H/D
U/D
I.
62.0
14.2
20.7
25.1
22.9
33.4
40.5
25.
25.6
7.1
11.5
7.0
27.7
44.9
27.3
2.
61.0?
14.4
21.8
23.3
23.6?
35.7?
.38.2?
26.
25.3
7.2
9.6
9.0
28.5
37.9
.35.6
3.
55.7
12.2
21.0
20.0
21.9
.37.7
35.9
27.
25.2
7.2
11.1
6.8
28.5
44.0
26.9
4.
50.4
10.8
20.4
15.3
21.4
40.5
.30.4
28.
24.6
6.5
10.7
6.2
26.4
43.5
25.2
5.
47.0
11.0
18.4
15.0
23.4
,39.1
31.9
29.
24.6
6.3
11.5
5.4
25.6
46.7
21.9
6.
45.3
11.4
18.3
13.5
25.1
40.4
29.8
.30.
24.0
7.2
10.5
6.4
30.0
43.8
26.7
7.
44.4
11.5
16.2
16.7
25.9
36.5
.37.6
31.
21.1
6.5
8.7
6.6
.30.8
41.2
31.3
8.
44.2
10.6
17.5
12.5
23.9
.39.6
28.3
32.
20.1
6.0
8.4
6.5
29.9
41.8
32.3
9.
43.0
9.7
18.2
11.0
22.6
42.3
25.6
33.
20.0
5.4
8.2
5.3
27.0
41.0
26.5
10.
39.1
9.6
15.4
12.5
24.6
.39.4
31.9
34.
19.3
5.7
8.2
5.7
29.5
42.5
29.5
11.
36.2
8.5
14.4
11.9
23.5
.39.8
32.9
.35.
19.2
5.7
8.1
5.7
.39.7
42.2
29.7
12.
33.8
8.2
14.7
9.0
24.3
43.5
26.6
.36.
18.9
6.5
7.4
5.8
34.4
.39.2
.30.7
13.
32.6
?
13.0
9.9
?
.39.9
30.4
.37.
18.2
5.1
7.0
5.9
28.0
.38.5
32.4
14.
32.5
8.4
13.7
9.3
25.8
42.2
28.6
.38.
18.1
5.8
7.3
6.1
32.0
40.3
.33.7
15.
31.0
8.3
13.3
8.7
26.8
42.9
28.1
39.
18.0
6.0
7.7
6.1
33.3
42.8
33.9
16.
31.0
8.5
12.4
10.2
27.4
40.0
32.9
40.
17.5
6.0
7.4
5.7
.34.3
42.2
32.6
17.
29.0
7.2
10.5
10.7
24.8
36.2
.36.9
41.
16.7
5.3
6.4
5.7
31.7
.38.3
.34.1
18.
28.4
7.6
11.2
9.1
26.7
.39.4
.32.0
42.
16.3
5.7
6.4
5.1
.34.9
39.3
31.3
19.
28.0
7.0?
12.3
8.1
25.0?
43.9
28.9
43.
15.8
5.0
6.3
5.3
31.6
.39.9
33.5
20.
27.5
7.3
11.8
7.6
26.5
42.9
27.6
44.
13.7
4.8
5.4
4.7
.35.0
.39.4
34.3
21.
27.2
7.0
11.2
9.3
25.7
41.2
.34.2
45.
11.5
4.3
5.0
4.0
37.4
43.5
34.8
22.
26.2
7.2
11.4
7.0
27.5
43.5
26.7
46.
11.3
4.2
4.4
3.7
37.2
.38.9
32.7
23.
26.0
7.5
10.8
7.8
28.8
41.5
30.0
47.
10.8
4.3
4.3
3.6
39.8
39.8
33.3
24.
26.0
6.8
9.8
9.2
26.1
37.7
35.4
48.
10.1
4.4
3.9
3.2
43.6
.38.6
31.7
1. Plesiotype, MCZ 956.5 (PI. 47, fig. 2).
3. Plesiotype, MCZ 9.564 (PI. 47, fiR. 1).
4. Holotype, USNM 75292a (PI. 48, figs. 3, 4).
8. ParaUpe, USNM 75292c (PI. 48, figs. 1, 2).
15. Plesiotyi^e, MCZ 9567 (PI. 47, fig. 4).
24. Plesiotype, MCZ 9568 (PI. 47, fig. 5).
29. Paratvpe, USNM 75292d (Smith, 1932: pi. 62, figs. 17, 18).
45. Plesiotype, MCZ 9566 (PI. 47, fig. 3).
All other specimens are from the Columbites fauna, Thaynes Formation at Montpelier Canyon and Hot Springs, .south-
eastern Idaho.
or in the dimensions of the whorls. There
is a tendeney toward greater opening of
the iimbiHcus with growth. Two of the
largest specimens of this species are illus-
trated on Plate 47, figures 1, 2; both of
these specimens show larger relative diam-
eters of the umbilicus than more juvenile
forms.
The suture is shown in Figure 32F, G,
and consists of two denticulated lateral
lobes and an auxiliary lobe on the umbili-
cal shoulder and wall that is generally
irregularly straight and denticulated. The
smooth conch bears only fine growth lines.
Nordophiceras jacksoni is very similar to
Nordophiceras etiomphahis (Keyserling)
from northern Siberia. The similaritv is so
close that I consider it possible these two
species may be conspecific. The Siberian
species is known from very few specimens,
and no data are available on the variation
within those populations. For the moment
it seems best to keep the two species sep-
arate.
Nordophiceras jacksoni and N. eiiom-
j)hahis are the smooth species of this
genus; in contrast to these two species
there are two other described species — N.
alexeevae and N. pilafum — which are very
similar in general conch morphology but
bear some ornamentation in the form of
ribs.
Occurrence. Columbites fauna, Thaynes
Formation, at Paris Canyon, Montpelier
470 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
20-
15
10
H
W
? ^ X '
X *
X XkX * X
w X X
... ^ '<
XX X
X * X
1
1
1
0
10
15 20 25
30 35 40 45
DIAMETER
50 55 60
Figure 33. Variation In whorl height (H) and whorl width (W) of Nordophiceras jacksoni (Hyatt and Smith), from Co-
lumbites fauna, Bear Lake region, southeast Idaho. The data on this graph are from Table 41.
Canyon, and Hot Spring.s, .southeastern
Idaho.
Repository. Holotype, USNM 75292a
(PI. 48, figs. 3, 4); paratypes, USNM
75292b (Smith, 1932: pi. 62, fig. 14),
USNM 75292f (PI. 4<S, figs. 1, 2), USNM
75282d (Smith, 1932: pi. 62, figs. 17, 18),
USNM 75292e (Smith, 1932: pi. 62, figs.
19-21); plesiotypes, MCZ 9564 (PI. 47,
fig. 1), MCZ 9565 (PL 47, fig. 2), MCZ
9566 (PI. 47, fig. 3), MCZ 9567 (PI. 47,
fig. 4), MCZ 9568 (PI. 47, fig. 5); suture
specimen, MCZ 9572 (fig. 32F); un-
figured specimens from Montpelier C^anyoii
MCZ 9570, from Hot Springs MCZ 9571.
Nordophiceras piiafum (Hyatt and Smith)
Plate 46, figures 2, 3; Plate 49, figures
1-8; Plate 50, figures 1-11; Plate
51, figures 1-5; Text-figures 34, 35
Mcckoccrus pilaiuui ll\att and Smith, 1905: 144,
pi. 63, fi,u.s. 3-9; biencr, 1915: 193; Smith,
1932: 59, pi. 63, fi^s. 7-13.
Mcekoccras curticosfdtinti Smitli, 1932: 56, pi.
48, figs. 21-30.
Mcekoccras micri>iiii)h(ilu.s Smitli, 1932: 58, pi.
49, lig.s. 5-11.
Mcckoccr(is sunctonu}! Smith, 1932: pi. 49, figs.
1-4.
This is one ol the more abundant species
in the Columhiies fauna oi southeastern
Idaho. Basically this species has an in-
\()lute, eompresscxl eoneh, with a rounded
Ammonoids of the Late Scythian (Lower Tiuassic) • Kiimmel 471
venter, and bears prosiradiate ribs of vary-
ing strength. There are considerable varia-
tions in the degree of involntion. inflation
of the whorls, and ornamentation. The
measurements of 42 specimens, including
the primary types, are given on Table 42
and plotted on Figure 35.
All of Smith's (1932) collection from
the Columhites fauna of southeastern Idaho
came from one locality, namely Paris Can-
yon. In the description of the four species
now included in N. pilatiun. Smith (1932)
made a direct comparison only between
his Aleckoccras curticostatiim and Meeko-
ccras pilatiim; he states, ^^Meckoceras ciirti-
costatum has some resemblance to M.
pihitinu, with which it is associated, dif-
fering in the wider umbilicus, more com-
pressed whorls, weaker and more numerous
ribs." Smith considered his Mcekoceras
mkromphaJus to be closely related to
Mcekoceras kcyserlingi Mojsisovics (now
the type species of Boreomeekoceras
Popov, 1961 ) from the Olenek region of
Siberia. He considered his Mcekoceras
sanctorum to be closely related to Mceko-
ceras affinc Mojsisovics also from the
Olenek region of Siberia. Even though
Smith (1932) did not discuss the specific
criteria upon which he separated his
species of this group, it is clear that the
intensity of ribbing, degree of whorl in-
flation, and degree of involution were im-
portant considerations. On examination of
a fairly large suite of specimens it is
readily apparent that the primary types of
Mcekoceras sanctorum (PI. 49, figs. 1-3)
and Mcekoceras micrornphalus (PI. 50,
figs. 7-11) are juvenile specimens. Both
these specimens are characterized by rather
small umbilici; examination of large mature
specimens, such as the holotype of Mceko-
ceras curticostatum (PL 50, figs. 1, 2) and
the new plesiotypes (PL 51, figs. 1-5),
clearly shows how the umbilicus increases
in relative diameter with growth. This is
well shown on the graph of Figure 34.
The umbilical region also reflects another
facet of variation and this is in the nature
of the umbilical shoulder and wall. Gen-
erally, the umbilical shoulder is fairly
acutely roimded with a sloping umbilical
wall; at the same time, there are forms with
slightly more rounded umbilical shoulders
and lower, sloping umbilical walls. Among
the large number of specimens before me
one can recognize a continuous gradation
from one type to the other.
The most conspicuous variation within
this species is in the intensity of the ribbing.
Typically the ribbing pattern consists of
fine, prosiradiate ribs on the inner volu-
tions that become more widely spread and
more subdued with growth of the shell.
In some forms the ribbing on the inner
volutions is extremely faint and on the
outer volutions there is nothing more than
a vague bundling of the growth lines (see
PL 49, fig. 1). On the other hand the
ribbing on the inner volutions may be
coarse and more widely spaced (as in the
paralectotype of N. pilatum, PL 49, figs.
4-6) but on the mature volutions these
become inconspicuous broad folds (as in
the lectotype of N. pilatum, PL 49, figs. 7,
8). Finally, there are forms (as the speci-
men of PL 50, fig. 2) in which the ribs
continue to increase in intensity through-
out growth. The photographs here of the
primary types and of a few additional
plesiotypes illustrate the range of variation
in ornamentation to be observed in the
collection. Needless to say, there is com-
plete gradation from one type to the other.
The venter is another area in which one
can observe extensive variation. This may
be from forms with acutely rounded ven-
ters to forms with more broadly rounded
venters. Generally, the acutely rounded
venters are found in the more juvenile
specimens.
This species is very similar to N. alexee-
vae Popov from northern Sibera. Both
these species are the ornamented nordo-
phicerids associated in their respective
faunas with smooth species, ettompliaJus
in Siberia, and jacksoni in southeast Idaho.
The sutures are illustrated on Figure 34.
472 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Ammonoids of the Late Scythian (Lower Triassk:) • Kummel 473
Table 42. Measurements of Nordophiceras pilatum (Hyatt and Smith)
BITES FAUNA AROUND BeAR LakE, SOUTHEASTERN IdAHO.
FROM THE COLUM-
D
w
H
u
W/D
H/D
U/D
D
w
H
u
W/D
H/D
U/D
1.
87.0
17.8
37.2
21.4
20.5
42.8
24.6
22.
42.7
9.4
18.8
12.0
22.0
44.0
28.1
2.
78.0
15.6
33.4
18.0
20.0
42.8
23.1
23.
42.5
8.8
17.7
11.7
20.7
40.0
27.5
3.
73.7
16.0
30.5
20.6
21.7
41.4
27.9
24.
41.8
9.2
19.0
9.6
22.0
45.5
22.9
4.
72.5
13.7
29.0
21.0
18.9
40.0
28.9
25.
41.5
11.0
21.0
6.4
26.5
50.6
15.4
5.
70.2
16.0
30.2
20.0
22.8
43.0
28.5
26.
40.4
8.3
17.7
9.2
20.5
43.8
22.8
6.
68.5
13.8
28.8
19.0
20.1
42.0
27.7
27.
39.2
9.4
16.7
11.6
23.9
42.6
29.6
7.
60.0
15.8
29.0
12.3
26.3
48.3
20.5
28.
38.4
8.1
16.8
10.0
21.1
43.8
26.0
8.
57.8
12.0
22.6
16.7
20.8
39.1
28.9
29.
37.9
8.5
19.5
5.7
22.4
51.5
15.0
9.
57.4
13.2
24.8
14.2
22.9
43.2
24.7
30.
35.8
7.2
15.7
8.8
20.1
43.9
24.6
10.
57.3
11.7
26.9
11.2
20.4
46.9
19.5
31.
35.7
7.8
19.0
5.0
21.8
53.2
14.0
IL
55.0
?
22.3
16.2
?
40.5
29.5
32.
34.6
8.0
15.7
7.8
23.1
45.4
66.8
12.
54.4
9.5
23.0
15.0
17.5
42.3
27.6
33.
33.7
7.5
17.0
5.3
22.3
50.4
15.7
13.
54.4
10.8
23.7
13.0
19.9
43.6
23.9
34.
.33.4
7.2
15.4
6.5
21.6
46.1
19.5
14.
53.4
10.2
21.4
16.8
19.1
40.1
31.5
35.
33.2
?
15.2
8.0
?
45.8
24.1
15.
53.1
10.8
23.5
13.7
20.3
44.2
25.8
.36.
.32.3
6.5
13.9
8.9
20.1
43.0
27.6
16.
52.0
11.0
22.7
13.2
21.2
22.5
25.4
37.
31.4
6.8
14.2
7.3
21.7
45.2
23.2
17.
48.7
9.2
19.8
9.0
18.9
40.7
18.5
38.
30.3
7.2
16.0
5.2
23.8
52.8
17.2
18.
48.7
10.6
21.2
13.0
21.8
43.5
26.7
.39.
27.3
6.3
12.6
6.1
23.1
46.2
22.3
19.
48.6
10.2
21.4
13.7
20.9
44.0
28.2
40.
21.0
4.6
10.4
4.1
21.9
49.5
19.5
20.
46.7
9.6
21.0
10.0
20.6
44.9
21.4
41.
16.6
5.6
7.6
4.4
33.7
45.8
26.5
21.
43.3
12.1
20.7
9.4
27.9
47.8
21.7
42.
12.5
3.4
6.3
2.5
27.2
50.4
20.0
3. Holotvpe, Meekoceras ciirticostatum Smith (1932: pi. 48, figs. 21, 22), USNM 74990a.
5. Plesiotype, MCZ 9542 (PI. .51, figs. 2, 3).
6. Plesiotype, MCZ 9541 (PI. 51, fig. 5).
7. Syntvpe, Meekoceras pilatum Hvatt and Smith (1905: pi. 63, figs. 7, 8), USNM 75294a.
14. Plesi(it%pe, MCZ 9539 (PI. 51, fig. 1).
15. Plesiotype, MCZ 9540 (PI. 51, fig. 4).
20. Paratvpe, Meekoceras sanctorum Smith (1932: pi. 49, figs. 3, 4), USNM 74991b.
21. Svntvpe, Meekoceras pilatum Hyatt and Smith (1932: pi. 63, figs. 10-13), USNM 75294b.
25. Holotype, Meekoceras micromphalus Smith (1932: pi. 49, figs. 5-8), USNM 74992a.
29. Holotype, Meekoceras sauctorum Smith (1932: pi. 49, figs. 1, 2), USNM 74991a.
31. Para^i-pe, Meekoceras curticostatum Smith (1932: pi. 48, figs. 23, 24), USNM 74990b.
33. Plesiotype, MCZ 9544 (PI. 46, fig. 3).
34. Plesiot>pe, MCZ 9543 (PI. 46, fig. 2).
38. Paratype, Meekoceras curticostatum Smith (1932: pi. 48, figs. 25, 26), USNM 74990c.
40. Paratvpe, Meekoceras curticostatum Smith (1932: pi. 48, figs. 27, 28), USNM 74990d.
41. Paratvpe, Meekoceras micromphalus Smith (1932: pi. 48, figs. 9-11), USNM 74992b.
42. Paratype, Meekoceras curticostatum Smith (1932: pi. 48, figs. 29, 30), USNM 74990e.
All other specimens are from the Columbites fauna, ThavTies Fomiation at Hot Springs (MCZ 9546)
Canyon (MCZ 9545), southeastern Idaho.
and Montpelier
Figure 34. Diagrammatic representations of the sutures of Nordophiceras pilatum (Hyatt and Smitfi). A, from paratype
of Meekoceras micromphalus Smitfi (1932; pi. 49, fig. 8), at a diameter of 17 mm (USNM 74992b); B, from paratype of
Meekoceras curt/cosfofum Smitfi (1932: pi. 48, fig. 24), at a diameter of 33 mm (USNM 74990b); C, from paratype of
Mee/coceras sonctorum Smitli (1932: pi. 49, fig. 4), at a diameter of 29 mm (USNM 74991b); D, from lectotype of Mee-
koceras pilatum Hyatt and Smitfi (1905: pi. 63, fig. 9), at a diameter of 60 mm (USNM 75294a); E, at a diameter of 45
mm (MCZ 9573a); F, at a diameier of 31 mm (MCZ 9573b); G, at a diameter of 40 mm (MCZ 9573c); H, at a diameter
of 42 mm (MCZ 9573d); I, at a diameter of 35 mm (MCZ 9573e); J, at a diameter of 45 mm (MCZ 9573f); K, at a diam-
eter of 30 mm (MCZ 9573g); L, at a diameter of 52 mm (MCZ 9573fi); M, at a diameter of 34 mm (MCZ 9573i); N, at
a diameter of 40 mm (MCZ 9573i); O, at a diameter of 53 mm (MCZ 9573k); P, at a diameter of 41 mm (MCZ 95731);
Q, at a diameter of 26 mm (MCZ 9573m); R, at a diameter of 45 mm (MCZ 9573n).
All specimens from Columbites fauna, Thaynes Formation, southeastern Idaho; specimens A-D are from Paris Canyon,
H and L from Hot Springs, the remaining specimens from Montpelier Canyon. All diameters given are approximate.
474 BiiJJcfin Museum of Coivparativc Zoology, Vol. 137, No. 3
35
30
25
20
15
10-
•40
•41
•42
'42
•40
•25.^1 -20
■29
•20
•21
•25
■29
38 ■a
•7
. '3
H
U
15
10
- 41
42
4 0.
21
25
20
2 9.
38
I I
w
J L
20 30
40 50 60
DIAMETER
70 80 90
Ammonoids of the Late Scythian (Lower Triassic) • Kmnmcl 475
Occurrence. Cohimbites fauna, Thaynes
Formation at Paris Canyon, Hot Springs,
and Montpelier Canyon, southeastern
Idaho.
Repo.sitonj. Lectotype, Meckuccrus pilci-
tiim Hyatt and Smith ( 1905: pi. 63, figs.
7-9) USNM 75294a; paralectotvpe (Hyatt
and Smith, 1905: pi. 63, figs. 10-13) USNM
75294b; holotvpe, M. curticostaium Smith
(1932: pi. 48,' figs. 21, 22) USNM 74990a;
paratvpes (Smith, 1932: pi. 48, figs. 23,
24) USNM 74990b, (Smith, 1932: pi. 48,
figs. 25, 26) USNM 74990c, (Smith, 1932:
pi. 48, figs. 27, 28) USNM 74990d, (Smith,
19.32: pL 48, figs. 29, 30) USNM 74990e;
holotvpe M. sanctorum Smith (19.32: pi.
49, f^gs. 1, 2) USNM 74991a; paratvpe
(Smith, 19.32: pi. 49, figs. 3, 4) USNM
74991b; holotype M. micrompJuihis Smith
(1932: pi. 49, figs. 5-8) USNM 74992a;
paratvpe (Smith, 1932: pi. 49, figs. 9-11)
USNM 74992b; plesiotvpes MCZ 9539 ( PI.
51, fig. 1), MCZ 9542 (PI. 51, figs. 2, 3),
MCZ 9540 (PI. 51, fig. 4), MCZ 9541 (PI.
51, fig. 5), MCZ 9543 (PI. 46, fig. 2), MCZ
9544 (PL 46, fig. 3); suture specimens MCZ
9573 a-n; unfigured specimens from Mont-
pelier Canyon MCZ 9545; unfigured speci-
mens from Plot Springs MCZ 9546.
Nordophiceras compressum (Chao)
Text-figure 32
Mcckoceras (Siibineekoceras) comprcsstiin Chao,
1959: 140, 320, pi. 44, figs. 1-6, text-fig. 46a, b.
Meekoceras (Siibmeekoceras) lolouense Chao,
1959: 141, p. 320, pi. 10, figs. 7, 8, text-fig. 45d.
Meekoceras (Suhmeekoceras) longiseptatum Chao,
1959: 141, pi. 10, figs. 5, 6, text-fig. 46c.
The three species brought together here
are based on specimens that come from
the same horizon and locality; two of the
species were based on one specimen each,
the other on five specimens. Chao (1959)
made no mention of the specific criteria he
used to differentiate these species. I can
see no significant differences between any
of these species. The sutures ( Figs. 32H-J )
are remarkably similar.
This is a smooth form of Nordophiceras
of the general pattern of euomphalus.
pseudoshnplex, and planorhis. There are
slight differences in conch form among
these species but more than anything the
suture is quite distinctive and the most
useful in differentiating this species.
Occurrence. Limestone block containing
a Subcolumbites fauna in Lolou village
(Chao collection 542b) Kwangsi, China.
Genus Pseudokymatifes Spath, 1934
Type species, Kymatites svilajanus KittI,
1903
Pseudokymafites svilajanus (KittI)
Plate 62, figure 5, Text-figure 40
Kymatites .milajamis KittI, 1903: 69, pi. 4, fig. 3;
Diener, 1915: 181.
Pseudokifmatites svilajaiitis, — Spath, 1934: 265, fig.
91; Kummel, /(( Arkell et al., 1957: L143, fig.
175, 6.
Kittl's type and only specimen of this
species is incomplete, crushed, and at best
of only fair preservation. It measures 63.5
mm in diameter, 13.0? mm for the width of
the adoral whorl, 29.4 mm for the height,
and the umbilicus is 12.3 mm in diameter.
Spath (1934: 265) created a new genus for
this species in his belief that it was prob-
ably a "smooth meekoceratid" with smooth
lateral lobes. The goniatitic nature of the
lobes, however, is not at all certain. The
specimen is weathered and the smooth
nature of the lobes could well be the result
of this. It is not possible to settle this ques-
tion on the basis of the single specimen
available. Speculations of the genetic re-
lations of this fonn are pointless until the
Figure 35. Variation in whorl height (H), whorl width (W), and umbilical diameter (U) of Nordophiceros pilatum (Hyatt
and Smith), from Columbites fauna, Bear Lake region, southeast Idaho. The data on this graph are from Table 42.
476 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
A
Figure 36. Diagrammatic representation of the suture of: A, /Arctomee/toceras rotundatum, — Popov (1962a: fig. 9), from
Olenekites Zone, northern Siberia, at a whorl height of 20 mm; B, Arctomeefcoceros sp. indet. Kummel (1966: fig. 22E),
from Narmia Member of Mianwoli Formation, Surghar Range, West Pakistan, at a whorl height of 9 mm; C, D, two
sutures of topotype specimen of Boreomeefcoceros keyserlingi (Mojsisovics) from Olenekites Zone, northern Siberia, C, at a
whorl height of 26.5 mm, D, at a whorl height of 15 mm (MCZ 8684).
basic morphological elements are better
understood.
Occurrence. Werfen Formation, Muc,
Dalmatia.
Repository. Natural History Museum,
Vienna.
Genus Arcfomeekoceras Popov, 1961
Type species, Mee/coceras rofundotum
Mojsisovics, 1886
Compressed, involute conclis, with nar-
rowly rounded venters on earlier volutions
tending to become more broadly rounded
with age. Suture ceratitic with two lateral
lobes and auxiliary series. The type and
only species of th(> genus is known from
the Olcnekilcs fauna of Siberia. Indetermi-
nate species have also been recorded from
the upper part of tlie Lower Triassic suc-
cession in the Salt Range of West Pakistan
(Kummel, 1966).
Arcfomee/coceras rofundotum (Mojsisovics)
Text-figure 36
Meekoceras rotundaliiin Moj.sisovits, IcSSfi: (S3, pi.
10, fijr. 16; Dicncr, 1915: 194.
Borcomcckoccras roltiiidatinu, — Popov, I9()l: 42.
Arcfomeekoceras ri>lmi(l(ilu)it, — Popov, 1962a: 187,
pi. 1, fiirs. 1, ,5, pi. 3, ii.u. 4
This is as yet an incompletely known
species; Mojsisovics apparently had only
one specimen and Popov four specimens
but the illustrations and descriptions are
incomplete. The key feature of the genus
and species is the broadening of the venter
on the adoral whorls. The suture (Fig.
36A) has two serrated lateral lobes and a
small auxiliary on th(> umbilical wall.
Occurrence. Olenekites Zone, Lena River
delta and Olenek River, Siberia.
Arcfomee/coceros sp. indet. (West Pakistan)
Text-figure 36
Arctotncckoceras .sp. iiulct. Kuiiiincl, 1966: 398,
pi. 2, fi.us. 1-5.
Generally incomplete and poorh' pre-
ser\'c>d specimens that appear to be much
like the type species of this genus, Init more
material is needed from both faunas before
a significant comparison can be made. The
suture is shown on Figure 36B.
Occurrence. Narmia Member of Mian-
wali Information in Salt Range and Surghar
liange of West Pakistan.
Repository. MCZ 95S4-9586, 9589, 9590-
Genus Boreomee/coceras Popov, 1961
Type species, Mee/coceros keyserlingi
Mojsisovics, 1886
lii\()lutc, compressed conch, venter nar
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 477
rowly rounded, flanks convergent. Suture
ceratitic with two lateral lobes and ex-
tended auxiliary series. The type and only
species of this genus is from the OJcnckitcs
fauna of the Olenek region, northern Si-
beria.
Boreomeekoceras keyserlingi (Mojsisovics)
Text-figure 36
Meekoceras keyserlingi Mojsisovics, 1886: 81, pi.
10, figs. 13-15; Diener, 1915: 192; Spath, 1934:
158, 254.
Boreomeekoeeras keyserlingi, — Popov, 1961: 42,
pi. 10, fig. 4.
This is a unique species in the known
late Scythian faunas because of its tightly
inxolute lenticular conch; the illustrations
by Mojsisovics (1886: pi. 10, figs. 13-15)
and Popov (1961: pi. 10, fig. 4) are quite
satisfactory. The suture, however, in nei-
ther of these publications is well repre-
sented. A topotype specimen is available
and two sutures from this specimen are
shown on Figures 36 C, D.
Occurrence. OJenekites Zone near mouth
of Olenek River, northern Siberia.
Repository. The MCZ has one topotype
specimen, 8684.
Genus Arctotirolites Popov, 1963
Type species, Pseudotirolites menensis
Popov, 1962
Arctotirolites menensis (Popov)
Pseudotirolites menensis Popov, 1962a: 178, pi. 2,
fig. 4.
Arctotirolites menensis (Popov) 1963: 137.
This new genus and species was estab-
lished on the basis of a single specimen.
The illustration and description of the spe-
cies leave much to be desired. The conch
is moderately involute, compressed, and
with an arched venter. The lateral areas
bear sigmoidal folds which apparently end
at the ventral shoulders in small nodes.
The suture has two serrated lateral lobes
and a serrated auxiliarv lobe on the um-
bilical shoulder and wall.
On the basis of the nodes one is at first
tempted to think of this form as a tirolitid
of some sort. However, the greater involu-
tion of the conch and the suture does not
support such a conclusion. On the basis of
the very incomplete data, it seems more
probable that this form is related to Nor-
dophiceras. The suture of menensis is very
much like that of a typical nordophicerid.
In addition, some species of N ordophiceras
are ornamented with ribs.
Occurrence. Olenekites Zone, Olenek
River basin, Mene River, northern Siberia.
Family NORITIDAE Karpinsky, 1889
Genus Albanites Arthaber, 1909
Type species, Pronorites triadicus Arthaber,
1908
Albanites triadicus (Arthaber)
Plate 16, figures 3-6; Plate 17, figures
1-10; Plate 18, figures 7, 8; Plate
20, figures 7-9; Text-figure 37
Pronorites triadicus Arthaber, 1908: 264, pi. 11,
figs. 4a-c; Arthaber, 1911: 204, pi. 17(1), figs
8, 9; Diener, 1915: 231; C. Renz, 1928: 155
Kutassv, 1933: 624; Renz and Renz, 1947: 61
Renz and Renz, 1948: 84, pi. 14, figs. 14-14b
Albanites triadicus,— Spath, 1934: 275, fig. 95;
Kummel, 1968b: 498, pi. 2, figs. 1-9.
Pronorites osmanicus Arthaber, 1911: 205, pi. 17
(1), fig. 10; Diener, 1915: 231; C. Renz, 1928:
155.
Albanites osmanicus, — Spath, 1934: 276.
Pronorites cf. osmanicus, — Renz and Renz, 1947:
62; Renz and Renz, 1948: 86, pi. 15, figs. 6-6c.
Pronorites arhanus Arthaber, 1911: 205, pi. 17(1),
figs. 11, 12; Diener, 1915: 230: Welter, 1922:
94, pi. 155, figs. 10-14; C. Renz, 1928: 155;
Kutassv, 1933: 624; C. Renz 1945: .301; Renz
and Renz, 1947: 61; Renz and Renz, 1948: 85,
pi. 14, figs. 13-13)1, 15-15b, pi. 15, figs. 5-5c.
Albanites arbanus, — Spath, 1934: 277.
Pronorites arbanus var. mcditcrranea Renz and
Renz, 1947: 62; Renz and Renz, 1948: 85, pi.
14, figs. 12-12b.
Pronorites spec. ind. ex aff. arbani, — Welter, 1922:
95, pi. 155(1), fig. 9.
Anasibirites gracilis Kiparisova, 1947: 164, pi. 39,
figs. 3, 4, text-figs. 60, 61.
Pronorites arbanus var. sundaica Renz and Renz,
1948: 85.
Albanites welteri Spath, 1934: 278.
Pronorites orientalis Renz and Renz. 1947: 62;
Renz and Renz, 1948: 86, pi. 15, figs. 2-2b.
478 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Pronoritcs schaubi Renz and Renz, 1947: 62, 78;
Renz and Renz, 1948: 87, pi. 15, figs. 4-4a.
Pronoritcs schcmhi var. timorcnsis Renz and Renz,
1948: 87.
Pronoritcs schauhi \'ar. kephalovunensis Renz and
Renz, 1947: 62, 78; Renz and Renz. 1948: 87,
pi. 15, figs. 3— 3a.
Pronoritcs reicheli Renz and Renz, 1947: 62, 79;
Renz and Renz, 1948: 88, pi. 15, figs. 1-lc.
Albanitcs danispancnsis (Astakhova) 1960a: 143,
pi. 34, figs. 4, 5; Astakhova, 1960b: 150.
Aspiditcs hasscrti Arthaber, 1911: 249, pi. 21(5).
fig. 16; Spatli, 1934: 275.
Mcekoccras (Koninckitcsj hasscrti, — Diener, 1915:
198.
Dagnoccras konianuin Arthal)er, 1911: 242, pi.
21(5), fig. 11; Diener, 1915: 115; Smith, 19.32:
65; Spath, 19.34: 269, 275.
Pscudosihiritcs cfr. dicliotonuis Waagen, — Arthaber,
1911: 254, pi. 22(6), fig. 8.
Anasihiritcs cfr. dichotomus, — Arthaber, 1911: 273.
Sihiritcs cf. dichotomus, — Diener, 1915: 255.
The genii.s Albanitcs i,s a con.spiciious
member of the Suhcohim])ites fauna from
Albania and Chios, from the Mangyshlak
Peninsula, and from the ProJiun^arites
fauna of Timor, l)ut in none of these local-
ities is the form particularly abundant. The
genus is much better represented in Chios
than in Albania, but the preservation of
most of the specimens in these faunas often
leaves much to be desired. It is factors of
preservation and preparation of the Al-
banian specimens that have led to some of
the misunderstanding about the genus. Ex-
amination of all the s})ecimens in the Al-
bania, Chios, atul Timor collections leads
me to conclude that they all represent a
single species; there have been seven spe-
cies and lour variety names introduced for
this grouji.
The holotype is a small spc^cimen of onl\
fair preservation; the lateral area and
venter ol the adoral quarter volution has
been ground and polished to expose the
suture. The smoothness ol the Ncntcr com-
mented on by Arthaber (1911: 205) and
Spath ( 1934: 277) is the result of this
grinding and polishing. The dimensions of
the holotype are given on Table 43, and the
sutvue is shown on 1^'igure 37. At the time
he introduced the species triadicus in 190S,
Arthaber had only one specimen. In 1911
Table 43. Measurements of Albanites tri-
adicus FROM THE SUBCOLUMBITES FAUNAS OF
Albani.a and Timor, and from Block "E,"
NiFOEKOKO, TlMOR.
D
w
H
u
W/D
H/D
U/D
1.
55.8
26.1
22.4
15.8
46.8
40.1
28.3
2
53.3
18.7
23.4
12.8
35.1
43.9
24.0
3.
49.2
14.2
22.3
10.1
28.9
45.3
20.5
4.
49.0
15.5
22.7
11.8
31.6
46.3
24.1
5.
40.0?
16.5
16.2?
10.1
41.3?
40.5?
25.3?
6.
38.4
13.5
16.1
11.7
35.2
41.9
30.5
7.
34.8
14.1
16.4
7.2
40.5
47.1
20.7
8.
34.7
12.8
14.4
11.4
37.1
41.5
32.9
9.
31.0
10.1
11.7
10.7
32.6
37.7
34.5
10.
28.7
8.7
11.8
9.1
30.3
41.1
31.7
11.
28.4
10.8
12.4
7.3
38.0
43.7
25.7
12.
28.2
8.4
14.4
4.4
29.8
51.1
15.6
13.
27.7
8.7
11.2
8.4
31.4
40.4
30.3
14.
25.8
9.0
12.0
6.1
34.9
46.5
23.6
15.
25.7
10.8
12.4
6.4
42.0
48.2
24.9
16.
25.0
10.5
11.1
6.4
42.0
44.4
25.6
17.
24.6
7.7
12.4
4.0
31.3
50.4
16.3
18.
?
?
P
12.8
?
?
?
1. Holotvpe, Pronoritcs reicheli Renz and Renz (1948:
pi. 15, fig. 1), NHMB J13809.
2. Plesiotvpe, Pronoritcs sp. ind. ex aff. arbani, — Welter
(1922: pi. 15.5(1), fig. 9). GPIBo W2()fi.
3. Holotvpe, Pronoritcs schaubi Renz and Renz ( 1948:
pi. 1.5, fig. 4), NHMB J138()5.
4. Type specimen, Pronoritcs schdtibi var. kephahyvu-
ncnsis Renz and Renz (1948: i^l. 15, fig. 3), NHMB
113808.
5. T\pe specimen, Pronoritcs osni<iniciis .\rthaher (1911:
pi. 17(1), fig. 10), PIUV.
6. Plesi(it\pe, Pronoritcs arbnnus, — Renz and Renz
(1948:' pi. 14, fig. 15), NHMB J13795.
7. S\nt\pe, Alb/iniics iceltcri Spath (^^ Pronoritcs iir-
/«/m/.s.-— Welter, 1922: pi. 155, figs. 10-12t, GPIBo
W205b.
8. Type specimen, Protioritcs iirbanus \ar. ineditcrnineti
Renz and Renz (1948: pi. 14. fig. 12), NHMB
J 13800.
9. Plesiotvpe, Protioritcs nrbanus, — Renz and Renz
(1948:' pi. 14, fig. 13), NHMB .|1379fi.
10. Holotype, Pronoritcs orirnfalis Renz and Renz (1948:
pi. 1.5, fig. 2), NHMB j 13801.
11. Plesiotvpe, Pronoritcs trindiciis, — Renz and Renz
(1948:' pi. 14, fig. 14), NHMB J13793.
12. Svnt\pe, AUwnitcs iceltcri Spath (^^ Pronoritcs ar-
hmuis.- \\elter, 1922: pi. 155, figs. 13, 14), GPIBo
W 205a.
13. Plesiot\ pe, Pronoritcs (irbanus. — Renz and Renz
(1948: pi. 15, fig. 5), NHMB j 13797.
14. Holotvpe, Pronoritcs triiiiliiiis Arthaber (1908; pi.
11. fig. 4), PIUV.
15. l'lesiot\pe, Pronoritcs cf. osnuiniciis. -Ren/, and lUnz
(1948;' pi. 15, fig. 6c), NHMB .113804.
l(i. Plesiot\ jie, Protioritcs cf. osnxiniciis, — Renz and Ren/
( 1948; pi. 15, figs, fi-fib), NHMB j 13803.
17. I,ectot\pe, Pronoritcs tirlxituis Arth.dicr (U)ll: p]. 17
< 1 ), figs. 11 a-d), P1U\'.
IS. ParalectotNpe, Pronoritcs arbaniis Arlli;dii'r ( U)l 1 : pi.
17( 1 ), figs. 12 a, b). PIUV.
Specimens ol numbers .5, 14, 17, 18 are Irom the
Sii)Holtiitil>itcs fauna of Albania; those of 2, 7, 12 are
Iroui the .\lhiinitcs fauna of Timor; and the remaining
specimens ;ire from the Siil>iiihn)iliitcs r;iuna of (-hios.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 479
he had one additional specimen to which,
however, no specific reference was made.
This specimen is illustrated here on Plate
17, figures 3, 4, and the dimensions are
given on Table 43. As can be seen, it is a
small juvenile form with a fairly inflated
whorl section. The venter is broadly arched
with rounded ventral shoulders; there are
nimierous weak ridges across the venter
\\'hich disappear just above the ventral
shoulders. This specimen differs from the
holotype mainly in its slightly more inflated
conch.
In 1911, Arthaber introduced two addi-
tional species for this group from the Al-
banian SuhcoJumhiics fauna. For Prono-
rites osmanicus Arthaber (1911: 205, pi.
17(1), fig. 10; PI. 17, figs. 5, 6 of this re-
port) the author states he had six speci-
mens. The illustrated specimen is the only
one preserved in the collections of the Pale-
ontological Institute, University of Vienna.
There are two topotypes in the British Mu-
seum (Natural History). Arthaber's illus-
trated specimen is selected as the lecto-
type; this illustration is one of the least
successful in Arthaber's monograph as it is
highly modified. The specimen is all phrag-
mocone, distorted, and generally of poor
preservation. Much of the lateral area of
the conch has been ground and polished to
expose the suture. The adoral quarter volu-
tion bears prominent cross ridges, but the
remainder of the venter is too poorly pre-
served to show this feature. The radial
ribs on the flanks of the penultimate volu-
tion are not present on the specimen. The
dimensions of the specimen are given on
Table 43. This species was said to differ
from A. triadicus in the presence of cross
ridges on the venter and in slight differ-
ences in the suture ( Fig. 37c ) . The smooth
venter of the holotype of A. triadicus is
due to grinding and polishing. The ridges
are clearly present on the small second
specimen assigned by Arthaber to A. triadi-
cus (PI. 17, figs. 3, 4).
Pronorites arbanus (Arthaber, 1911: 205,
pi. 17(1), figs. 11, 12) was based on three
specimens of which two were illustrated
and are still preserved. This species was
said to differ from A. osmanicus in its more
compressed conch, greater involution, and
slightly different proportions of the suture.
The measurements of these two specimens
are given on Table 43; the smaller of the
two specimens (Pi. 17, figs. 9, 10) was se-
lected by Spath (1934: 278) as lectotype.
The paralectotype (Pi. 17, figs. 1, 2) has
weak falcoid folds on the adoral half volu-
tion. This species is clearly only a com-
pressed variant and is conspecific with the
remaining forms of Albanites from Albania.
The collections from the Suhcolumhitcs
fauna of Chi(« studied by Renz and Renz
( 1948 ) contain approximately 50 specimens
of Albanites which they placed in six spe-
cies and two varieties. One species (ar-
banus) is represented by perhaps 20 speci-
mens, two species (triadicus and schaubi)
were recognized on eight specimens each,
one species (oricntaUs) on four specimens,
one species (cf. osmanicus) on two speci-
mens, and one species and two varieties on
the basis of a single specimen each. The
measurements of the prepared and more
complete specimens of Albanites in the
Renz and Renz collections from Chios are
given on Table 43. The basic difference
between all these species lies in the degree
of compression of the conch, in degree of
ornamentation, and in degree of involution
of the conch. Examination of the Chios
collection indicates to me that these char-
acters are highly gradational and the num-
erous species names are merely labels
applied to possessors of a particular mor^Dho-
logical character within a completely gra-
dational complex. Much of the discussion
of these species by Renz and Renz ( 1948 )
is confusing and out of date due to their
oversight of some of the earlier literature.
There has been some question and doubt
as to whether the two specimens from
Timor, described by Welter ( 1922 ) as Pro-
norites arbanus, are the inner whorls of
the same species that Welter described as
Pronorites spec. ind. ex aff. arbani. Exami-
480 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Figure 37. Diagrammatic representation of the sutures of Albanitei triadicus (Artficber). A, liolotype (Artliaber, 1911:
pi. 17(1), fig. 8), redrawn suture at diameter of 30 mm; B, paratype (Arthaber's second and smaller specimen of friadicus),
at a diameter of 18 mm; C, figured type of A. osmanicus (Artfiaber, 1911: pi. 17(1), fig. 10), redrawn suture at a diam-
eter of 38 mm; D, lectotype of A. arbanus (Artfiaber, 1911: pi. 17(1), fig. 11), redrawn suture at a diameter of 23 mm;
E, holotype of Dagnoceros komanum Arthaber (1911: pi. 21(5), fig. 11), redrawn suture at a diameter of 18 mm; tfiis
specimen has been ground and polished so that the fine details of the sutural elements have been destroyed; F, type
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 481
nation of Welter's original specimens leaves
no doubt on this question. Spath (1934:
278) came to the same conclusion on ex-
amination of a fine series of specimens in
the British Museum ( Natural History ) .
Spath ( 1934 ) recognized the "external re-
semblance" to the Albanian A. arhanus, but
felt because there were differences in the
suture that a new name was needed, and
he introduced the name A. welteri. The
differences in the suture, however, are no
more than what one can observe within the
Albanian and Chios faunas.
Albonifcs danispancnsis (Astakhova,
1960a) was stated to resemble A. osmani-
cus Arthaber and A. arhanus Arthaber, dif-
fering, however, in size of the umbilicus,
ornamentation, and details of the suture.
On the basis of the illustrations and de-
scriptions of this species, none of these dif-
ferences appear to be significant against
the range of variation known to exist in the
Albanian and Chios forms. This species
from the Mangyshlak Peninsula can be
considered as a representative of A. tria-
dicus.
Finally, there is a group of species from
the Subcohimbites fauna, based on very
poorly preserved specimens; an inspec-
tion of the types suggests inclusion in
this species. First, there is the specimen
for which Arthaber (1911: 242, pi. 21(5),
fig. 11; PI. 18, figs. 7, 8 of this report) pro-
posed the name Dagnoceras komanum.
This specimen measures 27 mm in diam-
eter, 7.2 mm for the width of the adoral
whorl, 12.7 mm for the height, and 6.3 mm
for the diameter of the umbilicus. The
whorls are compressed, converging slightly
toward a flattened venter. Both the ven-
tral and umbilical shoulders are sharply
rounded. On the adoral one-third volution
the ventral shoulders bear diagonal ribs;
the venter is not well enough preserved to
tell just how these ribs cross the venter. In
the better preserved specimen of A. tria-
dicus, cross ridges on the venter are con-
nected with weak radial ribs on the flanks.
I am inclined to believe that the diagonal,
projecting ribs on the ventral shoulder of
the holotype of Dagnoceras komanum are
at least partially due to the poor state of
preservation of the specimen and possibly
to slight crushing. Thus the difference in
the rib pattern from that of the type speci-
men of A. triadicus is more apparent than
real.
The specimen Arthaber (1911: pi. 22(4),
figs. 8a-c; Pi. 20, figs. 7-9 of this report)
assigned to Pseudosibirifes cfr. dichotomus
Waagen I believe to be essentially identical
to the specimen he assigned to Dagnoceras
komanuj}}. This conclusion is made taking
into account the poor and slightly different
preservations of these two specimens. The
lateral ribs are more conspicuous, as is the
chevron aspect of the ribs across the venter.
The specimen is, however, slightly crushed
laterally.
The sutures of these species are shown
on Figure 37E, F. First of all, it can be
seen that they are essentially identical; any
differences can readily be explained by ex-
cessive grinding of the specimen. Secondly,
it can be seen that these two sutures are
essentially identical to the sutures of the
specimens of AII)anifes so far known from
the Albanian Subcohimbites fauna.
specimen of Pseudosib/rifes cfr. d/chofomus, — Arthaber (1911: pi. 22(6), fig. 8), redrawn suture at a diameter of 18 mm;
G, plesiotype of A. orfaonus,— Renz and Renz (1948: pi. 14, fig. 13b), at a diameter of 20 mm (NHMB J13796); H, plesio-
type of A. ofbanui, — Renz and Renz (1948: pi. 14, fig. 15b), at a diameter of 25 mm (NHMB J13795); I, plesiotype of
A. arbonus var. med'iierranea (Renz and Renz, 1948: pi. 14, fig. 12b), at a diameter of 24 mm (NHMB J13800); J, plesio-
type (Renz and Renz, 1948: pi. 14, fig. 14b), at a diameter of 17 mm (NHMB J13793); K, holotype of Pronorites reicheli
Renz and Renz (1948: pi. 15, fig. Ic), at a diameter of approximately 40 mm (NHMB J13809); L, paratype of A/ban/7es
danispanensis (Astakhova, 1960a: fig. 10), at a diameter of 30 mm.
Specimens of figures A-F from Subco/umbi/es fauna of Albania, those of G-K from Subco/umb/fes fauna of Chios; speci-
men of figure L from the Mangyshlak Peninsula.
482 Bulletin Mii.wuni of Comparative Zoolo'^ij, Vol. 137, No. 3
Considering all the factors available for
these two "species," I believe it best to
consider them as conspecific with Allianitcs
triadieus of the same fauna. The differ-
ences in the rib pattern on the venter are
difficult to evaluate on the basis of the
sample available. No purpose is served in
assigning these two forms to different
genera.
Further, I suggest that the two speci-
mens Arthaber (19fl: 249, pi. 21(5), fig.
16) assigned to Aspiditcs hasscrti Arthaber
are poorly-preserved representations of A.
triadieus. Arthaber's figured type (PI. 16,
figs. 3, 4) measures 44.6 mm in diameter,
14.3 mm for the width of the adoral whorl,
20.8 mm for the height, and 10.5 mm for
the diameter of the umbilicus. In addition
to being poorly preserved, the specimen
has been ground and polished in places.
The basic plan of the suture is that of A.
triadieus. The smaller unfigured specimen
referred to by Arthaber (1911: 249) is
illustrated here on Plate 16, figures 5, 6.
This specimen measures 22.4 mm in diam-
eter, 6.6 mm for the width of the adoral
whorl, 11.7 mm for the height and 2.5 mm
for the diameter of the umbilicus. In both
these specimens the basic conch form and
suture show them to be close to A. triadieus;
what differences are apparent can be read-
ily explained as due to the state of pres-
ervation or due to grinding and polishing.
Occurrence. The species is known from
the Suheoluinhites fauna of Albania, Chios,
and Afghanistan, from the Frohuuiiarites
fauna of Timor, and from the CUihnnhites
Zone of Astakhova { 196()a, b ) on the
Mangyshlak Peninsula.
Repository. The Paleontological Insti-
tu(e, University of Vienna contains the
holotype, and the second specimen referred
to as Pronorites triadieus by Arthaber (19 11:
204; figured here Pi. 17,' figs. 3, 4), the
lectotype of Pronorites osmanieus Arthaber,
the lectotype and paralectotype of Pro-
norites arJ)anus Arthaber. This institution
also contains the holotype ol Dat^iux-eras
komanum Arthaber, the figured s):)ecimen
of Pseudosibirites cfr. dieliotonuis Waa-
gen, and the lectotype and paralectotype
of As])idites hasserti Arthaber. The Natural
History Museum, Basel, contains the fol-
lowing specimens: plesiotype Pronorites
triadieus, — Renz and Renz (1948: pi. 14,
fig. 14) NUMB J13793; unfigured speci-
mens NHMB J 13794; plesiotypes Pronorites
arhanus, — Renz and Renz (1948: pi. 14,
fig. 15) NUMB J13795, (pi. 14, fig. 13)
NHMB J13796, (pi. 15, fig. 5) NHMB
J13797; unfigured specimens from Mara-
dovuno NHMB J 13798, from Kephalovuno
NHMB J13799; figured specimens Pro-
norites arhanus var. rnediterranea Renz
and Renz (1948: pi. 14, fig. 12) NHMB
J 13800; holotype Pronorites oricntalis Renz
and Renz (1948: pi. 15, fig. 2) NHMB
J 13801; unfigured paratypes NHMB J13802;
figured specimens of Pronorites cf. osmani-
eus Renz and Renz ( 1948: pi. 15, fig. 6-6b)
NHMB J13803, (pi. 15, fig. 6c) NHMB
J 13804; holotvpe Pronorites sehauhi Renz
and Renz (1948: pi. 15, fig. 4-4a) NHMB
J 13805; unfigured paratypes from Mara-
dovuno NHMB J13806, from Kephalo\uno
NHMB J 13807; figured specimen of Pro-
norites sehauhi var. kephalominensis Renz
and Renz (1948: pi. 15, fig. ;3-3a) NHMB
J138()8; holotvpe Pronorites reieheJi Renz
and Renz (1948: pi. 15, fig. 1-lc) NHMB
J 13809. The three specimens studied by
Welter ( 1922 ) are in the Paleontological
Institute, Bonn University; additional topo-
type specimens are in the British Museum
(Natural Historv); specimens from Af-
ghanistan MCZ 10136, 10145. 10152. 10153,
l0156, 10168.
Family PRIONITIDAE Hyatt, 1900
Genus Hemiprionifes Spath, 1929
Type species, Goniodiscus typus Waagen,
1895
Hemiprionifes costafus Popov
This species is based on a single spc>ci-
men which unfortunateK is incompletely
described and illustrated. On the basis of
the data axaikible, I can onlx concur in
Aaimonoids of the Late Scythian (Lower Triassic) • Kummel
483
Popov's generic assignment of this species.
The genus Herniprionites had previously
been known only from the Anasihiritcs
Suhzon(> of the Owenites Zone. In this
mid-Scythian horizon it is a very common
and quite distinctive form. Popov (1961)
states this species is associated with
Inyuitc'.s cickifcnsis Popov (here assigned
to Siil)cishnuites) and Dicnewccras nik-
(ihifcnsis Popov (here considered to be a
synonym of D. demokidovi Popov). Both
these associated species are generalized
forms, and it is possible all of these species
belong to the Owenites Zone and are not
equivalent to the CoJumhites Zone as are
most of the faunas assigned by Popov
( 1961 ) to his Dieneroceras Zone.
Occurrence. Olenek River basin, north-
ern Siberia.
Family SIBIRITIDAE Mojsisovics, 1896
Genus Sibirites Mojsisovics, 1886
Type species, Ceratifes eichwaldi Keyser-
ling, 1845
Sibirites eichwaldi (Keyserling)
Ceratites cicJnvahli Ke\seiling, 1845: pi. 3, fig.
14; Eichvvald, 1868:' 1040; Mojsisovics, 1882:
41.
Sibirites eichwaldi, — Mojsisovics, 1886: 59, pi. 10,
figs. 1-9; Freeh, 1905: pi. 28, fig. 10; Diener,
1915: 255; Spath, 1934: 342, 344, figs. 116d,
e; Kiparisova, 1947: 164; Popov, 1961: 31, pi.
14, fig. 2; Vozin and Tikhomirova, 1964: 63,
pi. 38, figs. 1, 2.
Sibirites cf. eichwaldi, — Popov, 1961: 31, pi. 14,
fig. 6.
Sibirites pretiosus Mojsisovics, 1886: 61, pi. 10,
fig. 10; Diener, 1915: 256; Spath, 1934: 343,
figs. 116a-c; Popov, 1961: 32, pi. 13, fig. la;
Vozin and Tikhomirova, 1964: 64, pk 38, fig. 3.
Sibirites ind. aff. pretioso Mojsisovics, 1886: 61,
pk 10, figs. 11, 12; Diener, 1915: 256.
Sibirites grambcrgi Popov, 1961: 31, pk 14, fig. 1.
Farasibirites grambergi (Popov), 1962a: 181, pk
l.fig. 3.
Sibirites grambergi var. rariaculeatus Popov, 1961:
31, pk 14, fig. 5.
Farasibirites rariaculeatus Popov, 1962a: 182, pk
1, fig. 2.
Sibirites grambergi var. mixta Popov, 1961: 31,
pk 14, fig. 7.
Farasibirites mixtus Popov, 1962a: 183, pk 1,
fig. 4.
Sibirites subprctiosus Popov, 1961 : 33, pk 14,
fig. 8.
Farasibirites subprctiosus (Popov), 1962a: 181.
The large number of species brought
together here reflect the usual typological
treatment of trachyostracan ammonites.
The "species" differ in degree of ornamenta-
tion, that is, in the relative prominence of
ribs, nodes, etc. These species have now
been recorded from a number of localities
in northern Siberia, especially at and
around the Olenek region. Neither Moj-
sisovics (1886) nor Popov (1961, 1962a)
had particularly large collections to study
nor did they present much data on the
variabilit)' within their samples. I shall
make the prediction that when large col-
lections of Sd)iritcs from the Olenek region
become available they will show a com-
plete gradational series from lesser to
stronger omamented forms, and that the
ontogenetic development will be found to
be also highly variable. I look upon
Sibirites as having the same range of
genetic variability as is seen, for instance,
in Anasihiritcs kingianus or Columhites
parisianus, both species known by large
samples.
Occurrence. Olenekites Zone, northern
Siberia, Olenek River, Verkhoyan region,
Kolyma River basin, and eastern Taymyr.
Sibirites renzi n. sp.
Plate 24, figures 6-9
Anasibirites aff. anguloso (Waagen), — Renz and
Renz, 1947: 60; Renz and Renz, 1948: .35, pk
11, figs. 10, 11.
The Renzes ( 1948 ) listed only two speci-
mens of this species, but there are in the
collections four additional paratypes. The
preservation of all the specimens leaves
much to be desired. The shape of the
conch, degree of involution, and pattern
of ornamentation place this species in
Sibirites. The ribs are slightly interrupted
in their passage over the venter; in Ana-
sibirites the ribs are continuous over the
venter.
Occurrence. Subcolumbites fauna, Chios.
484 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
D
Figure 38. Diagrammatic representation of the suture of: A, Keyserlinglles subrobustus (Mojsisovics, 1886: pi. 4, fig. 2c),
at a diameter of approximately 85 mm; B, Keyser/ingifes middendorfti (Keyserling) — Mojsisovics (1886: pi. 3, fig. Ic), at
a diameter of approximately 100 mm; C, Ceroti/es nikifini Mojsisovics (1888: pi. 1, fig. 13c), at a diameter of approximately
25 mm; D, Cerafifes bungei Moisisovics (1888: pi. 1, fig. 14c), at n diameter of ap|3roximately 50 mm; E, fiolotype Key-
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 485
Repository. Holotype, Renz and Renz
(1948: pi. 11, fig. 10) NHMB J13623;
paratype, Renz and Renz (1948: pi. 11,
fig. li) NHMB J13624; PI. 24, figs. 6, 7 of
this report, NHMB J 19551; PI. 24, figs. 8, 9
of this report, NHMB J19552; unfigured
paratypes NHMB J 13625.
Genus Keyserlingites Hyatt, 1900
Type species, Ceratifes subrobustus
Mojsisovics, 1885
There are now five late Scythian species
of this genus confined to the circum-Arctic
region and in western North America. Four
of these species (sub robust us, middendorffi,
bcarJakensis, and bearriverensis) are con-
fined to the Frohuniiaritcs Zone, and one
species (stephensoni) is from the Colum-
bites Zone. On the basis of studies on sev-
eral well preserved specimens of Keyser-
lingites subrobustus from British Columbia
and Ellesmere Island, Tozer (1965a) has
been able to clarify the relations between
KeyserUngitcs and Durgaites. Tozer's sug-
gestion, however, that the Himalayan
"Durgaites" dieneri and the Timor "D."
angustecostatus may be upper Scythian in
age rather than Anisian, as concluded by
Diener (1907, 1912), Spath (1934) and
Welter (1915), is here rejected. This
question has been fully discussed in the
introductory chapter (p. 348).
KeyserVingWes subrobusfus (Mojsisovics)
Plate 26, figures 6, 7; Text-figure 38
Ceratites middendorffi Kevserling, 1845: pi. 2,
fig. 4.
Ceratites subrobustus Mojsisovics, 1885: 155, pi.
6, fig. 3; Mojsisovics, 1886: 44, pi. 4, fig. 2,
pi. 5, figs, la, b, pi. 6, fig. 1; Noetling, in
Freeh, 1905: 194, 200, pi. 28, fig. 5a, b.
Keyserlingites subrobustus, — Hyatt, 1900: 559;
Diener, 1915: 172; Spath, 1934: 355, fig.
119a-c; Kummel, 1961: 521; Popov, 1961: 55,
pi. 15, fig. 1; Tozer, 1965a: 31, pi. 5, fig. 1,
pi. 6, figs. 1, 2, pi. 7, figs. 1-3, pi. 8, figs. 1, 2;
Tozer, 1965b: 5.
Robustites subrobustus, — Philippi, 1901 : 556.
Keyserlingites cf. subrobustus, — Frebold, 1929a:
12, pi. 2, figs. 8, 9; Tozer, 1962, pi. 4, figs. 6a-c.
Ceratites bungei Mojsisovics, 1888: 8, pi. 1, fig.
14; Spath, 1934: 356.
Keyserlingites bungei, — Diener, 1915: 178; Popov,
1961: 54.
This species has recently been exten-
sively described and illustrated by Tozer
(1965a) on the basis of specimens from
Ellesmere Island and British Columbia
which have contributed much to our
knowledge of this most interesting form.
Ceratites bungei (Fig. 38D) is believed
to be based upon a juvenile specimen of
K. subrobusius. The differences in the two
Arctic species of Keyserlingites are largely
in the nature of the whorl section. This
species has a subquadrate whorl section,
whereas middendoiifi is a much more com-
pressed form. Keyserlingites bearlakensis
n. sp. has a suture quite similar to sub-
robustus (Fig. 38A) but a simpler pattern
of ornamentation with only a single set of
lateral bullae. The other species of Key-
serlingites in the Prohungarites Zone of
southeast Idaho, bearriverensis, is a com-
pressed form like the Siberian midden-
dorffi.
Occurrence. The type specimens came
from the Olenekites Zone at the mouth of
the Olenek River, Siberia. The species is
also known from Spitsbergen at Cape
serlingites bearriverensis n. sp., at a diameter of 50 mm; F, paratype Keyserlingites bearriverensis n. sp., at a diameter of
9.5 mm (MCZ 9521); G, holotype Keyser/ingifes bearlakensis n. sp., at a whorl height of 24 mm (MCZ 9516); H, paratype
Keyserlmg/fes beor/alcensis n. sp., at a whorl height of 25 mm (MCZ 9523); I, paratype Keyser/ingifes beor/o/tensis n. sp.,
at a diameter of 7.3 mm (MCZ 9518); J, holotype Keyser/mgites Stephenson/ n. sp. at a diameter of 190 mm; K, holotype
Ceratites subrobustus Diener (1897: pi. 19, fig. 2) (=KeYserlingites dieneri], at a diameter of approximately 110 mm;
L, Keyserlingites angustecostatus Welter (1915: 108, fig. 12).
Specimens of figures A-D from Olenekites lone, Olenek River region, Siberia; of E-l from Upper Thaynes Formation, Ham-
mond Creek, southeast Idaho; J, from Thaynes Formation, Fort Hall Indian Reservation presumably from Co/umb/(es Zone;
K, lower Anisian, Himalayas and Tibet; L, lower Anisian, Nifoekoko, Timor.
486 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Thordson (Frebold, 1929a), and from the
Grippia beds, Botneheia, south of Sassan-
fiord (Tozer, 1962). On Ellesmere Island
the species is present in the Blaa Mountain
Formation and the BHnd Fiord Formation,
and in British Columbia in the Toad For-
mation in the Halfway River area ( Tozer,
1965a).
Repositonj. The Siberian specimens are
in the Central Geological Museum, Lenin-
grad. The Spitsbergen specimen reported
by Frebold ( 1929a ) is in the Paleontologi-
cal Museum, Oslo; another Spitsbergen
specimen, collected by Frebold and illus-
trated by Tozer ( 1965a ) is in the Geologi-
cal Survey of Canada and a plastotype in
the Museum of Comparative Zoology. The
Ellesmere Island and British Columbia
species are in the Geological Survey of
Canada.
Keyserlingites middendorffi (Keyserling)
Ccratites middendorffi Keyserling, 1845: 170, pi. 1,
fig. 1; pi. 2, tigs. 1, 3 (noil 2, 4); Mojsisovies,
1882: 11; Mojsisovies, 1885: 153, pi. 6, fig. 2;
Mojsisovies, 1886: 38, pi. 2, figs. 12, 13, pi.
3, figs, la-e, pi. 20, fig. 10; Freeh, 1905: 200;
Spath, 19.34: .359.
Ainniojiitc.s middendorffi, — v. Bueh, 1848: 15.
Ceratites ( ?Stci)Juinites) middendorffi, — Freeh,
1905: pi. 28, fig. 7.
Kei/serlingites middendorffi, — Diener, 1907: 44;
biener, 1915: 178; Spath, 1934: 33, 353, 355,
356, .363, 4.32, fig. 119cl; Kumniel, 1961: 521;
Popov, 1961: 54, pi. 15, figs. 2, 3.
Ceratites nikitini Mojsisovies, 1888: 6, pi. 1, figs.
12, 13.
Keyserlinf^ites nikitini, — Diener, 1915: 179; Popov,
1961: 56, pi. 15, fig. 4.
Ceratites selireid<i MojsisoN ies, 1886; 47, pi. 4,
fig^ 1.
Kei/.s-erlinf^ites sehrenki, — Diener, 1915: 179.
C)l the two species of Siberian keyser-
lingitids this is the compressed form. It
differs from suhrohustus also in ornamenta-
tion, with its i)rominent nodes just below
the umbilical shoulder and only weak ridges
crossing the venter. Ke[iscr]in0tcs nikitini
(Mojsisovies) is much like middendorffi
except for being more cNolute. fhat
species is known from onl\ three speci-
mens and the difference in involution of
the conch from middendorffi is a little less
than 10 percent. As these two "species"
are associated in the same beds, I feel it
more likely that nikititii is nothing more
than an evolute variant of middendorffi.
In the same vein one can look upon Key-
serlini!,itcs sehrenki (Mojsisovies) as an
involute variant of middendorffi. In its
compressed whorls K. middendorffi is simi-
lar to K. hearriverensis n. sp. from south-
east Idaho, but the two species differ
significantly in their ornament pattern.
The sutures of these species are illus-
trated on Figure 38.
Oceurrence. This species is only known
from northern Siberia where it occurs in
the Olenekitcs Zone at the mouths of the
Lena and Olenek rivers, in the Kolvma
River basin of the Verkhovan region, and
in eastern Taymyr.
Repositonj. The Siberian specimens are
in the Central Geological Museum, Lenin-
grad. The Museum of Comparative Zool-
ogv has a specimen from the delta region
of 'the Lena River (MCZ 6108).
Keyserlingifes bearlakensis n. sp.
Plate 37, figures 5, 6; Plate 38, figures
1-3; Text-figure 38
Keyserlingites n. sp. et. K. suhrohusltis (Moj-
sisovies),— Kuninicl, 1954: 187.
The largest ammonoids in the upper
member of the Thaynes Formation at Ham-
mond Creek, Bear River Range, southeast
Idaho, are species of Kcyserhniiites. This
species is represented by ten, mainly frag-
mentary, specimens. The conch is evolute
with an lunbilieus measuring appro.ximately
30 percent of the diameter. The w horls are
depressed, being wider than high, with a
broadly arched xenter. The lateral areas
are likewise convex and merge onto a
broadK rounded umbilical slope. The
conch is ornamented with largc>, j)rominent
bullae on the lateral areas. On the holo-
t\pe (PI. 38, ligs. 1, 2), there are nine
bullae on the hall \olution. The bullae
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 487
occupy the entire lateral area. Between
the bullae there may or may not be low,
weak ribs extending across the venter from
one ventral shoulder to the other. The
suture is shown on Figures 38G-L
The collections contain one small speci-
men of 7.8 mm in diameter, which is be-
lieved to be the juvenile whorls of this
species (PI. 37, figs. 5, 6). What is of
special interest in this specimen is that at
this diameter it already has all of the main
morphological features of the large adult
specimens. This includes the depressed
whorl section, bullae, and suture ( Fig.
381).
The Tobin Formation of the Tobin
Range, Nevada, has yielded a few poorly
preserved, fragmentary specimens that are
possibly conspecific with this species. In
its depressed whorls this species is much
like the Arctic K. suhrohustus but differs
in ornament pattern.
Occurrence. Upper member of Thaynes
Formation, Hammond Creek, Bear River
Range, southeast Idaho.
Repository. Holotype MCZ 9561 (PI. 38,
figs. 1, 2); figured paratypes MCZ 9517
(PI. 38, fig. 3), MCZ 9518 (PI. 37, figs. 5,
6); unfigured paratypes MCZ 9519.
Keyserlingifes bearriverensis n. sp.
Plate 37, figures 1-4; Text-figure 38
It was pointed out above that of the two
Siberian species of Keyserlingites, one was
a form with a depressed whorl section {K.
suhrohusius) and the other was a form
with a compressed whorl section (K. mid-
dcndorjfi). A similar relationship exists
with two species of Keyserlingites that
occur in the upper member of the Thaynes
Formation at Hammond Creek, Bear River
Range, southeast Idaho. Of the compressed
species the collections contain eight poorly
preserved specimens.
The venter is highly arched, grading onto
convex lateral areas; the umbilical shoulder
and wall are broadly convex. Prominent
bullae on the lateral areas extend from the
umbilical shoulder to the ventral shoulder;
they are essentially radial on the phragmo-
cone but on the living chamber become
slightly prosiradiate. The suture is shown
on Figure 38E, F.
The collection contains a small specimen
of 9.8 mm in diameter which is believed to
be a juvenile of this species. Like the
juvenile specimen of K. bearlakensis de-
scribed above, this specimen already has
the essential features of the adult. That is,
the compressed whorl section, high, arched
venter, and bullae are clearly developed.
The suture has the basic pattern of lobes
and saddles, but the lobes are only slightly
denticulated (Fig. 38F).
In the compressed nature of the conch
this species is similar to K. middcndorffi.
It differs from that species in the nature
of the bullae and of the venter.
The fact that each of the Siberian and
Idaho late Scythian faunas have two
species, one compressed in whorl section,
the other depressed, tempts one to con-
sider the possibility that these are dimorphs
of one species in each case. However, the
material available from each fauna is far
too limited to explore this problem further.
Occurrence. Upper member of the
Thaynes Formation, Hammond Creek, Bear
River Range, southeast Idaho.
Repository. Holotype MCZ 9520 (PI. 37,
figs. 1, 2); figured paratype MCZ 9521 (PI.
37, figs. 3, 4); unfigured paratypes MCZ
9522.
Keyserlingites sfephensoni n. sp.
Plate 46, figure 1; Text-figure 38
This species is based on the largest speci-
men of an ammonite discovered in the
Lower Triassic formations of western United
States. It was discovered by Gordon R.
Stephenson of the U.S. Agricultural Re-
search Service of Boise, Idaho, in whose
honor the species is named. The specimen
measures 268.0 mm in diameter, 106 mm
for the height of the last whorl, and 80 mm
for the diameter of the umbilicus. The
488 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
specimen was broken out of a large black
limestone concretion, and one side is em-
bedded in matrix. Even though a width
measurement is not possible, it is clear
that the whorls are higher than wide. The
lateral areas are broadly convex, and
slightly convergent; the venter is broadly
arched. Both the ventral and umbilical
shoulders are rounded.
The most striking feature of the speci-
men is its ornamentation. On the umbilical
shoulder are large nodes. On the last volu-
tion there are seven such nodes that
adorally increase in size. On the ventral
shoulder there are slightly elongated nodes
( clavi ) which also increase in size adorally.
There are approximately twice as many
nodes on the ventral shoulder as on the
umbilical shoulder.
Tlie shell is preserved over much of one
side of the specimen. The shell in the
region of the nodes measures as much as
5 mm in thickness. On the mid-part of the
venter the shell is only 2 mm thick.
The suture is only partially known, that
is, the first lateral lobe, second lateral sad-
dle, and the second lateral lobe are visible
(Fig. 38J). This species is remarkably
similar in its gross feature to the specimen
of Kciiserlingites middcndoiffi figured by
Mojsisovics (1886: pi. 3) from the Olenek
fauna of northern Siberia. The Idaho speci-
men lacks the transverse ventral ribs, and
has more prominent nodes on the ventral
shoulder. It differs from Kcys-crlingites
.mbwJnistu.s, also from the Olenek fauna,
and from Kcyscrliniiitcs hcarlakensis in its
compressed rather than depressed \N'horl
section.
Occurrence. From the lower black lime-
stone member of the Thaynes Formation,
Fort Hall Indian Reservation, southeast
Idaho. The specimen is presumably from a
horizon e((uivalent to the Columhitcs
fauna of the Bear Lake region.
Repository. This specimen has been pre-
sented by Mr. G. R. Stephenson to the
Department of Geology, Washington State
University, Pullman, Washington.
Keyserlingifes sp. indet. (Afghanistan)
Keijserlingites sp. indet. Kunimel, 1968b: 500,
pi. 1, figs. .^7.
A small specimen, 22 mm in diameter, is
the first record of the genus Keyscrlingites
in late Scythian strata of Tethys. The speci-
men appears to be all phragmocone and
has a whorl height of 9 mm and an umbili-
cal diameter of 7.5 mm. The whorl sides
are slightly convex and converge toward
a broadly rounded venter. The umbihcal
shoulder is abruptly rounded and the um-
bilical wall nearly vertical. There are large
nodes, one approximately every quarter
volution, that are anchored on the umbilical
shoulder and extend upward on the flanks.
The most adoral node, at a diameter of 21
mm, extends half way across the lateral
areas. The suture consists of a simple
pronged ventral lobe, a large first lateral
saddle and first lateral lobe, and much
smaller second lateral saddle and lobe; a
low denticulated auxiliary lobe occupies the
umbilical wall. The general shape of the
conch, the nodes, and the suture identifies
this specimen as Keyscrlingites. The speci-
men is most probably a juvenile torm.
Occurrence. Snbcolumbites fauna, Kotal-
e-Tera, Afghanistan.
Repository. MCZ 10139.
Genus Olenekites Hyatt, 1900
Type species, Dinarifes spiniplicafus
Mojsisovics, 1886
Olenekites spiniplicafus (Mojsisovics)
DiiKiritc.s .spiniplicafus Mojsisovics, 1886: 10, pi.
I, figs. 1-5, 8-16, 18-26, pi. 2, figs. 1-5, 7;
Mojsisovics, 1888: 2, pi. 1, figs. 1-3; Freeh,
1905: pi. 28, Fig. 9.
Olenekites spiuipliratus, — Hyatt. 1900: 559; Spath
1934: 361, pi. 7, fig. 3; Kiparisova, 1947: 166;
k'nminel, in Arkell et al., 1957: L146, figs. 178,
5; I'opoN, 1961: 34, pi. 14, figs. 3, 4; Vozin and
Tikhomirova, 1964: 67, figs. 4-7.
Dinarite.s (Olenekites) spiniiilieatiis, — Diener, 1915:
123.
Diniirites roiulus Mojsisovics, 1886: II, pi. 1.
fig. 6. pi. 2, fig. 6; Mojsisovics, 1888: 1, pi. 1,
figs. 5. 6.
Dinuritcs (Olenekites) lohitiis, — Diener, 1915:
124.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel
489
Dinaritcs densiplicatus Mojsisovics, 1886: 15, pi.
1, fig. 7; Mojsisovics, 1888: 4, pi. 1, fig. 4.
Ditmrites (Olcnekites) densiplicatus, — Diener,
1915: 123.
Dinaritcs altiis Mojsisovics, 1886: 16, pi. 2, fig. 8.
Dinaritcs {Olcnekites) altus, — Diener, 1915: 123.
Olcnekites fl/ft/.s,— Popov, 1961: 35, pi. 13, fig.
lb; Vozin and Tikhomirova, 1964: 68, pi. 33,
fig. 3.
Dinaritcs intermedius Mojsisovics, 1886: 17, pi.
2, fig. 9.
Dinaritcs (Olcnekites) intermedins, — Diener, 1915.-
123.
Dinaritcs glacialis Mojsisovics, 1886: 18, pi. 2,
fig. 11.
Dinaritcs (Olcnekites) glacialis, — Diener, 1915:
123.
Olcnekites glacialis, — Popov, 1961: 34, pi. 13,
fig. 16.
Both Mojsisovics ( 1886 ) and Spath
( 1934 ) have made particular note of the
large degree of variation in nearly all
morphological features in this species.
Mojsisovics had 64 specimens of spini-
plicafus, but for the remaining five species
he assigned to his Dmarites he had only
one, two, or three specimens per species.
Restudy of the type specimens, topotype
material, and the literature of Mojsisovics
(1886, 1888) and Popov (1961) demon-
strate clearly that there is but one species
of Olcnekites in this north Siberian region.
Occurrence. Northern Siberia, mainly at
and around mouth of Olenek River.
Repository. The Museum of Compara-
tive Zoology contains a number of topotype
specimens (MCZ 8682, 8683). The Central
Geological Museum, Leningrad, contains
the types of Mojsisovics and Popov.
Olenekites mangyshlakensis Astakhova
Olcnekites mangyshlakensis Astakhova, 1960a:
148, pi. 34, figs. 6, 7, text-fig. 14.
This species is obviously quite close to
O. spiniplicatus and could possibly be con-
specific. However, since data available on
this species are so incomplete, it seems best
to accept it as a separate and distinct
species.
Occurrence. Mangyshlak Peninsula, Kar-
atauchik Range, from Columbitcs Zone of
Astakhova, 600-650 m above base of Tyur-
Upa suite.
Olenekites canadensis Tozer
Olenekites canadensis Tozer, 1961a: 73, pi. 18,
figs. 1-3; Tozer, 1965a: 32, pi. 4, figs. 1-8,
text-fig. 10.
This is likewise a highly variable species,
differing from O. spiniplicatus mainly in its
truncate venter.
Occurrence. Blaa Mountain FoiTnation,
upper Scythian, Ellesmere Island.
Olenekites cf. spiniplicatus (Mojsisovics)
Plate 36, figures 4-6
Previously, the two available specimens
of this species were assigned to Olenekites,
with question, as no suture is preserved
( Kummel, 1954 ) . In recent years, I have
had the opportunity of examining numerous
specimens of Olenekites from the type
locality at the mouth of the Olenek River,
northern USSR, and no longer have any
doubt but that these specimens represent
a species of Olenekites. Each specimen
consists of only one-half volution. The
whorls are approximately as high as wide;
the venter is broadly rounded, as are the
ventral shoulders. The umbilical shoulders
are more abruptly rounded, and the umbili-
cal wall nearly vertical. The lateral area
bears bullae that begin on the umbilical
shoulder and decrease gradually toward
the ventral shoulder. On one specimen ( Pi.
36, figs. 4, 5) the bullae are rather robust
and widely spaced, there being approxi-
mately four per half volution. On the other
specimen the bullae are narrower, not as
robust, and there are approximately six
bullae per half volution. Both Mojsisovics
( 1886 ) and Spath ( 1934 ) have emphasized
the exceedingly variable character of this
species. The two specimens recorded here
can be compared favorably with one or
more of the specimens illustrated by Moj-
sisovics (1886: pi. 1). The species O.
canadensis, described by Tozer (1961a:
73) from Ellesmere Island, is also closely
490 BuUefin Museum of Comparative Zoology, Vol. 137, No. 3
related to Olcnckitcs spinipJicatus, differ-
ing mainly in the subtabulate character of
the venter on the later whorls.
Occurrence. Uppermost member of
Thaynes Formation, Hammond Creek, Bear
River Range, sontheast Idaho.
Repusitori/. MCZ 9482 (PL 36, figs. 3, 4),
MCZ 9476 (PI. 36, figs. 5, 6).
Genus Eukashmirifes n. gen.
Type species, Kashmirifes acutanguiatus
Welter, 1922: 125, pi. 9, figs. 9-12
The genus Kashmirites was introduced
by Diener (1913: 33) for the "group of
Celtites siibrcctang,uJari.s Waagen or Ccltitcs
(innoftis Waagen from the Ceratite Forma-
tion of the Salt Range." Diener at that
time was dealing with a Scythian fauna
from Kashmir. He had within his Kashmir
fauna fragmentary specimens which he felt
belonged to the Celtites of Waagen de-
scribed from the Salt Range. This new
generic name was introduced for these Salt
Range and Himalayan species as they no
longer could be accommodated in the genus
Celtites as redefined by Mojsisovics ( 1893).
In this revision it is quite apparent that
Diener overlooked the genus Pseudoccltites
Hyatt (1900) — type species Celtites miil-
tipJicatus Waagen. In the discussion of the
genus Pseudoceltites it has already been
pointed out that Celtites armatus (the type
of Kashmirites, so designated by Diener,
1915: 137) is a synonym of Celtites mul-
fiplicatus. There is still, however, a need
for separation of the group of ammonoids
with subquadrate whorl sections, lateral
ribs, occasional nodes which cross the
venter, and a simple two lobed suture. Both
Diener (1913) and Spath (1934) looked
upon Kashmirites htaschkei Dic>ner as a
typical species. As noted by Diener ( 1913),
the Kashmir specimens of hlaschkei and
related species are poorly preserved and
fragmentary. Because of this 1 si-lecl
Welter's Timor species acuta)i^iilattts as
the type species of this new genus.
Etikashmirites is fairly well represented
in the mid-Scythian zones of the Himalayas
and Timor. In the late Scythian Prohun-
garites Zone it is represented by only two
species, both from the Mangyshlak Penin-
sula in southern U.S.S.R.
Eukashmirifes subdimorphus (Kiparisova)
Kashmirites suhditijorpjuis Kiparisova, 1947: 148,
pi. 33, figs. 3-5, text-figs. 40, 41; Astakhova,
1960a: 140.
The overall shell morphology of this
species is very much like that of the type
species — E. acutanguiatus — from Timor ex-
cept the whorls tend to be broader on the
adoral volutions. The other species of this
genus, E. contortus Astakhova, also known
only from the Mangyshlak Peninsula, is a
much more compressed fonn with slightly
sinuous ribs on the adoral whorls.
Occurrence. — Scythian formation of Man-
gyshlak Peninsula but Kiparisova ( 1947 )
gives no precise horizon. Astakhova (1960b:
150) lists the species from her Tirolites
Zone.
Eukashmirifes conforfus (Astakhova)
Kashmirites contortus Astakhova, 196()a: 139, pi.
33, fig. 5, text-fig. 7.
This species is also quite similar to the
type species — acutanguiatus — but much
more compressed in its whorl section; the
lateral ribs also are sinuous on the adoral
whorls. The sample of this species available
to Astakhova and to Kiparisova when she
described E. subdimorphus was very small.
It is conceivable that a large sampk> would
show these two species to be synonyms.
Occurrence. Tirolites Zone of Astakhoxa,
Mangyshlak Peninsula.
Genus Anakashmirifes Spath, 1930
Type species, Danubifes nivalis Diener, 1897
Species of this genus are known mainly
from the mid-Scythian Ouciiifes Zone. The
only records of this genus are fragmentary
and specificalK' indeterminate forms ironi
the Narmia Member of (he Mianwali For-
mation ill the Sursihar Range of West
Ammonoids of the Late Scythian (Lower Triassic) • Kinnmcl
491
Pakistan (Kummel, 1966). These speci-
mens in conch form and ornamentation are
very much Hke the species from the
Owenites Zone but the suture is distinctly
more advanced.
Superfamily CERATITACEAE Mojsisovics,
1879
Family TIROLITIDAE Moisisovics, 1882
Genus Tirolifes Mojsisovics, 1879
Type species, Tirolifes idrianus Hauer, 1865
As with most genera of ornamented am-
monites, the genus Tirolifes includes a more
than generous number of species. The
great majority of these species were estab-
lished for forms from the Werfen Forma-
tion of the Alps and Dalmatia. Mojsisovics
in his classic monograph on "Die Cephalo-
poden der mediterranen Triasprovinz"
(1882) recognized 14 species of Tirolifes.
The ammonites of the Werfen Formation
were monographed by Kittl (1903), who
recognized 40 species of Tirolifes and three
subgenera! Most subsequent authors (e.g.
Smith, 1932; Spath, 1934) recognized that
many (or most) of Kittl's species were
very closely allied or identical, yet con-
tinued to recognize all the species — I pre-
sume as a matter of convenience.
I have had the opportunity of studying
the Werfen Formation collection described
by Kittl (1903), in the Natural History
Museum, Vienna. All the specimens fig-
ured by Kittl (1903) were photographed,
and measurements were made of all speci-
mens sufficiently well preserved. It should
be remarked here that Kittl's monograph
is profusely illustrated by line drawings,
and that in effect Kittl illustrated nearly
every specimen of fair to good preserva-
tion. Most of the unfigured paratypes and
topotypes are very poorly preserved speci-
mens. No satisfactory photographic illustra-
tions of the Werfen Formation Tirolifes
exist; for this reason the more important
types of Kittl's study are reproduced here.
Study of these illustrations first of all clearly
shows the relatively poor preservation of
most of the specimens, particularly since
these figured forms are the very best speci-
mens in the collection.
Tubercles on the ventro-lateral shoulder
form the basic ornamentation pattern for
Tirolifes. Very often there are radial or
prosiradiate ribs that extend from the
tubercles dorsally along the flanks for vari-
able distances. Study of the hundreds of
specimens in the Kittl collection soon gives
one the impression that they represent one
continuous gradational series. Mojsisovics
( 1882 ) recognized two primary groups
within the genus Tirolifes, the Spinosi with
ornamented inner whorls and the Seminudi
with smooth inner whorls. Of the 14 species
of Tirolifes that Mojsisovics (1882) recog-
nized from the Werfen Formation, 7 were
assigned to the Spinosi, and 7 to the
Seminudi. Kittl (1903) adopted the same
two groups and recognized three additional
groups as subgenera (Hololobus, Svilajifes,
Bitfnerifes).
Preservation of the Werfen Formation
ammonites is such that retention and ex-
posure of the inner whorls is not the usual
thing. There are likewise numerous transi-
tions between the two groups, Spinosi and
Seminudi, and between these and other
genera. One is tempted to consider all of
the tirolitids of the Werfen Formation as a
complex, variable, single species group.
This could very well be the case. At the
same time, in a general way, the Seminudi
seldom have ribs associated with the tuber-
cles as do the Spinosi. Clearly as a matter
of convenience and not because of any real
understanding of the relationships, I recog-
nize two major species of tirolitids from the
Werfen Formation — T. idrianus (Seminudi)
and T. cassianus (Spinosi).
The type specimen of Tirolifes {Svilajifes)
cingulafus is poorly preserved. Spath
(1934) raised this group to generic rank
and this was accepted by Kummel (in
Arkell, et al., 1957). Examination of the
type specimen leads me to believe that at
best this can be recognized only as a
distinct species of Tirolifes.
492 Bidletin Museum of Comparative Zoology, Vol. 137, No. 3
Tirolitoides prior ( Kittl, 1903 ) was said
to have umbilical nodes and a more ceratitic
suture. The so-called umbilical nodes are
no more than a reflection of poor preserva-
tion and the suture is no different from
that of many other specimens figured by
Kitd (1903).
The comprehensive interpretation of the
Werfen Formation tirolitids creates special
problems in assessing the relationships to
species from other horizons and localities.
In no other locality or formation are the
tirolitids as abundantly represented as they
are in the Werfen Fonnation. The other
species of Tirolites recognized here are
discussed below, but in almost every case
the species is known from very few speci-
mens.
The Siihcolumhites fauna of Albania has
yielded three specimens of Tirolites ( Artha-
ber, 1908, 1911) which are here considered
to be T. idrianus. The upper Scythian for-
mation of the Mangyshlak Peninsula has
yielded T. rossicus Kiparisova ( 1947 ) and
T. impolitus Astakhova ( 1960a ). Tliese two
species were said to be associated with T.
cassiamis and T. spinosus. The Narmia
Member of the Mianwali Fonnation in the
Salt Range of West Pakistan has yielded a
single indeterminate species of Tirolites
(Kummel, 1966). The Hedenstroemia fauna
of the Himalayas has yielded a single
species — T. injucundus Krafft and Diener
( 1909 ) . This is the oldest species of the
genus recorded to date. A fragmentary
specimen presumably from a Subeolutn-
I)ites horizon in Kwangsi was assigned b\
C^hao (1959) to Tirolites cf. dancini. Tiro-
litids are present in strata of Olenek age
in the basin of the Ivolyma River, Siberia
(Popov, 1961).
Finally, in western North America there
are several records of Tirolites. In fact the
stratigraphic position of the Tirolites fauna
as generally interpreted was established on
the basis of the sequence of faunal zones
in southeastern Idaho. In Paris C'anyon,
Smith (1932) discovered a small, poorly
preserved faima including three species of
Tirolites, above his Meekoceras Zone and
below his Columhites Zone. I have had
several occasions to verify this sequence.
These Paris Canyon species are very simi-
lar to the Spinosi of the Werfen Formation
and are treated here as a single but distinct
species — T. liarti Smith. From the over-
lying Columhites Zone, Smith ( 1932 ) de-
scribed a specimen as T. illyricus. Addi-
tional specimens of this species suggest its
separation as a distinct form. The Colum-
hites fauna contains two other species of
Tirolites very different from the form
Smith (19.32) assigned to T. illyrieus. Sil-
berling (in Hose and Repenning, 1959:
2194) has recorded indeterminate species
from the upper part of the Thaynes Fonna-
tion in the Confusion Range of western
Utah. These are recorded here as T. cf.
eassianus.
It can be seen from this brief summary
that species of Tirolites are now recognized
throughout the upper half of the Scythian.
Tirolifes idrianus (Hauer)
Plate 18, figures 1-6; Plate 66, figures
1-13; Plate 67, figures 1-9; Plate
68, figures 1-9; Plate 69, figures
1-10; Text-figure 39
Ccraiitcs idrianus Hauer, 1865: 610, pi. 1, figs.
4, 5.
Tirolites idrianus, — Mojsisovics, 1879: 138; Moj-
sisovics, 1882: 67, pi. 1, fig. 1: Kittl, 1903: 36,
pi. 5, figs. 8, 9; Arthaber. in Freeh, 1906: pi.
34, fig. 14.
Tirolites seniinudus Mojsisovics, 1882: 66, pi. 2,
fig. 11: Kittl, 1903: 40, pi. 6, figs. 3-10, 17,
18; Arthaber, 1908: 275, pi. 11(1), fig. 9;
Diener, 1915: 279; Spath, 1934: .375.
Tirolites seniinudus var. nttdior Kittl. 1903: 41,
pi. 6, fig. 3; Spath, 1934: 375.
Tirolites .wniinudus var. plieosus Kittl, 1903: 41.
pi. 6, figs. 5, 7; Diener, 1915: 279; .Spath, 19.34:
375.
Tirolites irn'reiirii Mojsisox ics, 1882: 68, pi. 1,
fig. 9; Kittl, 1903: .38, pi. 5, figs. 10, 11, pi. 6,
figs. 1, 2; Diener, 1915: 278; Spath, 19.34: 377.
Tirolites paueispinatus Kittl, 1903: .39, pi. 6, fig.
11, pi. 7, figs. 4-6; Diener, 1915: 279.
Tirolites distans Kittl, 1903: 42, pi. 6, figs. 12-16,
pi. 7, figs. 7, 8; Diener, 1915: 278; Spath, 19.34:
375.
Tirolites ijuetistedti Mojsisoxics, 1882: 66, pi. 2,
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel
493
fig. 12; Kittl 1903: 42, pi. 6, fiRs. 19, 20; Die-
ner, 1915: 279.
Tirolitcs wbiistus Kittl, 1903: 43, pi. 7, figs. 9-11,
pi. 8, fig. 1; Diener, 1915: 279; Spath, 1934:
377.
TiroUtes dimidiattis Kittl, 1903: 44, pi. 8, fig. 15;
Diener, 1915: 278; Spath, 19.34: 377.
TiroUtes stcichei Kittl, 1903: 45, pi. 7, fig. 14;
Diener, 1915: 280; Spath, 19.34: 377.
TiroUtes dinarus Mojsisovics, 1882: 74, pi. 2, fig.
9; Kittl, 1903: 45.
TiroUtes hyhridus Kittl, 1903: 46, pi. 8, fig. 2;
Diener, 1915: 278.
TiroUtes auf^ustus Kittl, 1903: 47, pi. 7, fig. 12;
Diener, 1915: 277; Spath, 1934: .377.
TiroUtes siibiUijrictts Kittl, 1903: 47, pi. 7, figs.
15, 16; Diener, 1915: 280.
TiroUtes iUt/ricus Mojsisovies, 1882: 68, pi. 2,
fig 10; Kittl, 1903: 48, pi. 8, figs. 3, 4, 6-9;
Arthaber, 1911: 250, pi. 22(6), fig. 4; Diener,
1915: 278; Spath, 1934: .373.
TiroUtes repuhus Kittl, 1903: 49, pi. 8, figs. 5,
10, 11, 14; Diener, 1915: 279.
TiroUtes rotiformis Kittl, 1903: 50, pi. 8, figs. 12,
13; Diener, 1915: 279.
TiroUtes reetanguhiris Mojsisovics, 1882: 69, pi. 3,
fig. 5; Kittl, 1903: 50, pi. 8, figs. 16, 17; Artha-
ber, 1911: 251, pi. 22(6), fig. 5; Diener, 1915:
279; Spath, 1934: .374.
TiroUtes undulatus Kittl, 1903: 52, pi. 7, fig. 13;
Diener, 1915: 280.
TiroUtes heterophanus Kittl, 1903: 38, pi. 5, fig. 7.
The species idrianiis includes all the
"species" that had been included by
Mojsisovics and Kittl in the Seminudi.
Measurements of 122 specimens from the
Werfen Formation of Muc, studied by
Kittl, are given on Table 44 and plotted
on Figure 39. There is clearly considerable
variation in these conch parameters. The
same is true of the pattern of ornamenta-
tion, the principal criterion used to dif-
ferentiate species within this group. The
illustrations given here should amply show
that the number, spacing, etc., of the tu-
bercles are highly variable. The sutures in
the Seminudi were claimed to be entire,
that is, goniatitic. This is not at all certain,
as details of fine denticulations are most
often not retained in the kind of preserva-
tion which characterizes the Werfen Forma-
tion fauna. The position of the lateral lobe
is likewise highly variable.
Occurrence. Werfen Formation of Alps,
Dalmatia, and associated regions. The
three specimens of TiroUtes recorded by
Arthaber ( IQO.S, 1911) from the Suhcolmn-
hites fauna of Albania are included in this
species.
Repository. All of the Kittl collection is
in the Natural History Museum, Vienna.
The specimens from the Subcolumbites
fauna of Albania are in the Paleontological
Institute, Vienna.
Tirolifes cassianus (Quenstedt)
Plate 63, figures 1-9; Plate 64, figures
1-4; Plate 65, figures 1-9; Plate 70,
figures 3-6, 11, 12; Text-figures
40, 41
Ceratites eassianus Quen^iedi, 1849: 231, pi. 18,
fig. 11; Hauer, 1865: 606, pi. 2, figs. 1, 2;
Laube, 1869: 61, pi. .37, fig. 1.
Ammonites {Ceratites) cassianus, — Hauer, 1851:
6, pi. 2, fig. 5.
TiroUtes cassianus, — Mojsisovics, 1882: 70, pi.
2, fig. 48, pi. 81, fig. 3; Tomniasi, 1895: 69,
pi. 4, fig. 15; Kittl, 1903: 54, pi. 9, figs. 4-6;
Arthaber, 1906: pi. 34, fig. 15; Wittenbnrg,
1908: 285, pi. 40(5), fig. 19; Diener, 1915:
278; Diener, 1925: 80, pi. 12, fig. 1; Ogilvie-
Gordon, 1927: 31, pi. 3, fig. .39; Kutassy, 1933:
674; Spath, 19.34: 369, fig. 126; Leonardi, 1935:
90, pi. 6, figs. 5, 6; Kollarova-Andrusovova,
1961: 56, pi. 1, figs. 1, 3, 4.
TiroUtes cassianus var. tenuis Mojsisovics, 1882:
71, pi. 2, figs. 4-6; Kittl, 1903: 55.
TiroUtes cassianus var. alpha Kittl, 1903: 55, pi.
9, figs. 4, 5.
TiroUtes angustilohatus Kittl, 1903: 54, pi. 9,
figs. 1-3; Diener, 1915: 277; Spath, 1934: 370.
TiroUtes angustilohatus var. alpha Kittl, 1903:
54, pi. 8, fig. 19.
TiroUtes spinosus Mojsisovics, 1882: 70, pi. 1, fig.
10, pi. 2, figs. 1-3; Tomniasi, 1895: 70, pi.
4, fig. 16; Kittl, 1903: 56, pi. 9, fig. 7; Diener,
1915: 279; Spath, 1934: 370; Leonardi, 1935:
91, pi. 6, figs. 7-9; Kollarova-Andrusovova,
1961: 57, pi. l,figs. 2a, b.
TiroUtes haueri Mojsisovics, 1882: 71, pi. 3, figs.
2-4; Kittl, 1903: 56, pi. 9, figs. 8-13; Diener,
1915: 278; Spath, 1934: 371.
TiroUtes haueri var. minor Kittl, 1903: 58, pi. 10,
figs. 1-3; Diener, 1915: 278.
TiroUtes multispinatus Kittl, 1903: 58, pi. 11,
fig. 9; Diener, 1915: 279.
TiroUtes percostatus Kittl, 1903: 58, pi. 10, fig.
6; Diener, 1915: 279.
TiroUtes turgidus Mojsisovics, 1882: 72, pi. 3,
figs. 6, 7; Kittl, 1903: 59, pi. 10, figs. 7, 8;
Diener, 1915: 280; Spath, 1934: 371.
TiroUtes darwini Mojsisovics, 1882: 73, pi. 2,
494 Bulletin Museum of Comparative Zoology, Vol. 137. No. 3
Table 44. Measurements of specimens of Tirolites wrianus (Hauer) from Muc in Dalmatia
STUDIED BY Ernst Kittl, 1903. All specimens are deposited in the Natural History Museu:m,
Vienna.
D
w
H
u
W/D
H/D
U/D
D
w
H
u
W/D
H/D
U/D
1.
66.0?
18.3
19.0
22.1
27.7?
28.8?
33.5?
25
44.9
12.8
15.5
17.7
28.5
34.5
39.4
2.
57.3
?
22.7
19.0
?
39.6
33.2
26
44.7
12.4
18.0
16.0
27.7
40.3
35.8
3.
55.5
18.5
19.9
22.7
33.3
35.9
40.9
27
44.5
14.4
16.0
16.3
32.4
36.0
36.6
4.
53.7
17.8
18.8?
197?
33.1
35.0?
36.7?
28
44.5
13.2
16.4
16.9
29.7
36.9
38.0
5.
52.6
?
20.7
19.3
?
39.4
36.7
29
44.5
?
18.4
15.4
?
41.3
34.6
fi.
52.5?
16.1
20.0
20.7
307?
38.1?
39.4
30
44.4
?
15.3
17.8
?
34.5
40.1
7.
52.4
13.1
19.9
20.0
25.0
38.0
38.2
31
44.3
12.1
16.7
19.4
27.3
37.7
43.8
8.
51.8
16.7
18.5
21.1
32.2
35.7
40.7
32
44.2
13.0
17.3
16.0
29.4
39.1
36.2
9.
51.7
15.5
17.0
21.0?
30.0
32.9
40.6?
33
44.1
12.3
16.2
17.5?
27.9
36.7
39.7?
10.
51.3
17.1
19.2
18.6
33.3
37.4
36.2
34
44.1
10.4
9.2
19.1
23.6
20.9
43.3
11.
51.2
14.7
17.4
22.0
28.7
34.0
43.0
35
43.7
12.6
15.2
17.3
28.8
34.8
39.6
12.
51.1
16.2
20.3
18.7?
31.7
39.7
36.6?
36
43.4
10.1
15.7
17.0
23.3
36.2
39.2
13.
50.2
10.0=±:
18.5
19.0
19.9±
36.9
37.8
37
43.3
14.3
16.8
16.6
33.0
38.8
38.3
14.
50.0
12.0
16.5
19.6
24.0
33.0
39.2
38
43.3
10.0
14.5
18.3
23.1
33.5
42.3
15.
49.7
9.9
17.3
21.1
19.9
34.8
42.5
39
43.2
14.0
17.3
15.4
32.4
40.0
35.6
16.
49.6
15.8
17.8
19.8
38.9
35.9
39.9
40
43.2
11.2
12.8
21.8
25.9
29.6
50.5
17.
48.4
13.2
18.0
18.1
27.3
37.2
37.4
41
43.2
8.5?
16.7
15.7
19.7?
38.7
36.3
18.
47.5
14.3
16.0
17.5?
30.1
33.7
36.8
42
43.0
10.4?
16.8
15.7
24.2
39.1
36.5
19.
47.2
10.8
17.2
18.4
22.9
36.4
39.0
43
43.0
?
17.1
15.2
?
39.8
35.3
20.
47.2
?
19.2
17.1
?
40.7
36.2
44
42.7
14.1
15.8
16.2
33.0
37.0
37.9
21.
46.3
15.5?
17.3
17.7
33.5?
37.4
38.2
45
42.7
13.3
15.7
16.0
31.1
36.8
37.5
22.
46.2
15.0
17.2
17.5
32.5
37.2
37.9
46
42.6
13.7
15.3
17.0
32.2
35.9
39.9
23.
45.3
16.6
16.7
16.8?
36.6
36.9
37.1?
47
42.6
13.1
14.7
16.0
30.8
34.5
.37.6
24.
45.0
13.9
16.8
18.4
30.9
37.3
40.9
48
42.4
13.4
14.6
17.7
31.6
34.4
41.7
1.
2.
3.
4.
5.
6,
7,
S,
9.
11,
13.
14.
1.5.
17,
19.
20.
21.
22.
2.3.
24.
2.5.
26,
27.
28.
29,
31,
32.
33,
34.
3.5.
38.
40.
41.
42.
43.
44.
47,
imicuni.
Lectotype, T. unduiattis Kittl (1903: pi. 7, Hg. 13).
Lectotype, T. stachei Kittl (1903: pi. 7, fig. 14).
Plesiotype, T. mercurii, — Kittl (1903: pi. 6, fig. 2).
Unfigured paratvpe, T. luihriduf. Kittl (1903: 46).
Lectotviie, T. hijbridus Kittl (1903: pi. 8, fig. 2).
16, 37, 45, 46, 48, 50, 64, 92. Unfigured para types,
10, 12, 39, 63, 74, 86, 88. Unfigured paratvpes, T.
18, 75, 77, 85, 99, 102. Unfigured paratypes, 7".
Plesiotype, T. mercurii, — Kittl (1903: pi. 6, fig. 1).
55, 61, 67, 70, 71, 76, 89, 90, 95, 103, 104,
mens, T. semintidus, — Kittl (1903: 40).
Plesiotype, T. quenstedti, — Kittl (1903: pi. 6, fig. 19).
Plesiotype, T. illyricus, — Kittl (1903: pi. 8, fig. 4).
Plesiotype, T. seminudiis, — Kittl (1903: pi. 6, fig. 3).
87, 98. Unfigured specimens, T. quenstedti, — Kittl ( 1903:
Plesiotype, T. quenstedti, — Kittl (1903: pi. 6, fig. 20).
Paralectotype, T. distans Kittl (1903: pi. 7, fig. 8).
Plesiotype, 7'. mercurii, — Kittl (1903: pi. 5, fig. 11).
Plesiotype, T. illyricus, — Kittl
Plesiotype, T. mercurii, — Kittl
Paralectotype, 7'. distans Kittl
Syntype, T. subilli/ricus Kittl (1903: pi. 7
30. Unfigured paratypes, 7'. subiUijricus
robust us Kittl (1903: 43).
Justus Kittl (1903: 47).
-Kittl ( 1903: 38).
105, 106, 110, 111. 115, 118, 121, 122. Unfigured speci-
42).
(1903: pi. 5, fig.
(1903: pi. 8, fig. 3).
(1903: pi. 5, fig. 10).
(1903: pi. 6, fig. 12).
fig. 15).
Kittl (1903: p
( 1903:
M).
Jig. 4).
Paralectotype, T. distans Kittl (1903: pi. 6, lig.
Plesiotype, 7'. semintidus, — Kittl ( 1903: pi. 6,
83, 113. Unfigured parat\ pes, 7'. i)aucisi)inatus Kittl (1903:
36, 60, 65, 66, 81. Unfigured specimens, T. illyricus, — Kittl
Plesiotype, 7'. illyricus, — Kittl (1903: pi. 8, fig. 8).
53, 69, 73, 94, 101. Unfigured paratypes, T. distans Kittl
Plesiotype, T. rectanaularis, — Kittl (1903: pi. 8, fig. 16).
Plesiotype, T. semintidus, — Kittl (1903: pi. 6, fig. 7).
Figured type, T. repulstis Kittl (1903: pi. 8, fig. 11).
Figured type, 7'. rotijormis Kittl (1903: pi. 8, fig. 12).
Plesiotype, T. illyricus, — Kittl (1903: pi. 8, fig. 7).
Plesiotype, 7'. seininudus, — Kittl (1903: pi. 6, fig. 8).
Plesiotype, T. seminudtis, — Kittl (1903: pi. 6, fig. 18).
Figured type, T. reptdsus Kittl (1903: pi. 8, fig. 5).
108. Unfigured paratypes, T. rotiforniis Kittl (1903: 50).
39).
( 1903:
48).
( 1903: 42).
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 495
Table 44. Continued.
D
w
H
u
W/D
H/D
U/D
D
w
H
u
W/D
H/D
U/D
49.
42.4
12.6
15.1
17.9
29.7
35.6
42.2
86.
38.0
11.7
14.4
14.6
30.8
37.9
38.4
50.
42.3
12.8
17.0
15.3
30.3
40.2
36.2
87.
38.0
8.2?
14.7
14.5?
21.6?
38.7
38.2?
51.
42.3
11.2
15.1
14.8
26.5
35.7
35.0
88.
37.8
8.4?
15.3
13.0
22.2?
40.5
34.4
52.
42.2
14.6
16.2
16.3
34.6
38.4
38.6
89.
37.5
12.0
12.8
15.4
32.0
34.1
41.1
53.
42.1
13.2
14.5
17.1
31.4
34.4
40.6
90.
37.3
10.0
13.3
16.0
26.8
35.7
42.9
54.
42.1
12.0
14.2
17.1
28.5
33.7
40.6
91.
37.0?
11.6
13.8
14.0?
31.4?
37.3?
37.8?
55.
42.0
10.2
15.0
16.1
24.3
35.7
38.3
92.
37.0
11.5
14.5
13.6
31.1
39.2
36.8
56.
41.7
13.8
16.5
14.5
33.1
39.6
34.8
93.
37.0?
11.3
12.6
12.2?
30.5?
34.1?
33.0?
57.
41.5
11.7?
16.1
15.4
28.2?
38.8
37.1
94.
37.0
10.2
13.6
13.3
27.6
36.8
35.9
58.
41.4
15.6
16.8
14.6
37.7
40.6
35.3
95.
36.3
11.5
12.7
14.7
31.7
35.0
40.5
59.
41.4
13.2
15.2
17.0
31.9
36.7
41.1
96.
36.0
8.9
14.0
14.4
24.7
38.9
40.0
60.
41.4
10.5
14.6
17.6
25.4
35.3
42.5
97.
35.7
10.7
12.3
14.8
30.0
34.5
41.5
61.
41.2
12.1?
14.1
16.1
29.4?
34.2
39.1
98.
35.7
9.8
12.0
14.4
27.5
33.6
40.3
62.
41.0
10.8
15.1
17.1
26.3
36.8
41.7
99.
35.6
12.0
14.1
11.7
33.7
39.6
32.9
63.
40.8
13.8
14.7?
15.3?
33.8
36.0?
37.5?
100.
35.6?
11.8
13.0
14.8
33.1?
36.5?
41.6
64.
40.7
13.0
14.0
15.8
31.9
34.4
38.8
101.
35.5
9.4
13.1
14.4
26.5
36.9
40.6
65.
40.5
11.6
9.2
16.1
28.6
22.7
39.8
102.
35.4
10.8?
13.6
14.0
30.5?
38.4
39.5
66.
40.5
11.3
13.8
18.2
27.9
34.1
44.9
103.
35.4
10.5
12.7
12.8
29.7
35.9
36.2
67.
40.5
8.8?
13.5
17.0
21.7
33.3
42.0
104.
35.2
11.3
13.0
13.3
32.1
36.9
37.8
68.
40.1
12.8
14.7
14.6
31.9
36.7
36.4
105.
35.2
9.3
12.6
13.8
26.4
35.8
39.2
69.
40.1
11.9
14.2
16.2
29.7
29.7
40.4
106.
35.0
8.7?
13.1
13.3
24.9?
37.4
38.0
70.
40.1
11.6
13.5
15.5
28.9
33.7
38.7
107.
34.9
8.7?
12.5
13.2
24.9?
35.8
37.8
71.
40.0
12.6
14.2
16.5
31.5
35.5
41.3
108.
34.8
9.5
11.1
15.2
27.3
31.9
43.7
72.
40.0
12.1
15.0
14.3
30.3
37.5
36.8
109.
34.6
10.7
10.5
15.5?
30.9
30.3
44.8?
73.
39.8
11.7
14.1
15.2
29.4
35.4
38.2
110.
34.1
9.6
12.5
13.4
28.2
36.7
39.3
74.
39.7
12.8
15.0
14.4
32.2
37.8
36.3
111.
34.0
10.1
12.6
13.2
29.7
37.1
38.8
75.
39.6
18.8
15.0?
14.5
47.5
37.9
36.6
112.
33.9
9.8
11.8
13.0
28.9
34.8
38.3
76.
39.4
12.3
14.5
15.7
31.2
36.8
39.8
113.
32.3
8.5
11.2
12.7
26.3
34.7
39.3
77.
39.2
14.0
15.0?
14.0?
35.7
38.3?
35.7?
114.
31.8
8.6
11.4
12.8
27.0
35.8
40.3
78.
39.0
13.8
14.1
14.7
35.4
36.2
37.7
115.
31.2
8.8
11.5
11.8
28.2
36.9
37.8
79.
38.8
?
16.1
13.6
?
41.5
35.1
116.
30.8
7.8?
9.6
8.7
25.3?
31.2
28.2
80.
38.6
10.2
13.2
16.2
26.4
34.2
42.0
117.
30.5?
7.5?
11.6
11.4
24.6?
38.0?
37.4?
81.
38.5
9.1
12.5?
17.4
23.6
32.5?
45.2
118.
30.0
7.9
11.2
11.3
26.3
37.3
37.7
82.
38.5
?
16.5
13.7
?
42.9
35.6
119.
28.7
8.9
9.3
12.3
31.0
32.4
42.9
83.
38.2
11.5
13.3
14.6
30.1
34.8
.38.2
120.
28.6
7.5
9.0
12.7
26.2
31.5
44.4
84.
38.2
11.4
13.8
14.7
29.8
36.1
38.5
121.
28.4
8.5
11.1
9.9
29.9
39.1
34.9
85.
38.0
13.1
15.3
13.4
34.5
40.3
35.3
122.
21.7
6.3
7.7
7.7
29.0
35.5
35.5
49.
51.
52,
54.
56.
57.
58.
59.
62.
68.
72.
79.
80.
82.
84.
91.
93.
96.
97,
100.
107.
112.
114.
116.
117.
120.
Untigured paratype, T. rectangiilaris, — Kittl (1903: 50).
Figured type, T. paucispinatus Kittl (1903: pi. 6, fig. 11).
78. Unfigured parat>'pes, T. undulatus Kittl (1903: 52).
Figured type, T. paucispinatus Kittl (1903: pi. 7, fig. 6).
Holot>pe, T. angustus Kittl (1903: pi. 7, fig. 12).
Figured type, T. paucispinatus Kittl (1903: pi. 7, fig. 4).
Lectot>pei T. robustus Kittl (1903: pi. 7, fig. 9).
Plesiotype, T. rectangiilaris, — Kittl (1903: pi. 8, fig. 17).
Plesiotype, T. quenstedti, — Kittl (1903: pi. 6, fig. 20).
Paralectotype, T. distans Kittl (1903: pi. 6, fig. 13).
Plesiotype, T. seminiidiis, — Kittl (1903: pi. 6, fig. 5).
Paralectotype, T. distans Kittl (1903: pi. 7, fig. 7).
Figured type, T. repulsus Kittl (1903: pi. 8, fig. 10).
Figured type, T. paucispinatus Kittl (1903: pi. 7, fig. 5).
Plesiotype, T. iZ/yncus,— Kittl (1903: pi. 8, fig. 6).
Lectotype, T. distans Kittl (1903: pi. 6, fig. 15).
Paralectotype, T. distans Kittl (1903: pi. 6, fig. 16).
Plesiotype, T. iUyricus, — Kittl (1903: pi. 8, fig. 9).
109, il9. Unfigured paratypes, T. repulsus Kittl (1903: 49).
Paralectotype, T. robustus Kittl (1903: pi. 7, fig. 11).
Syntype, T. subillyricus Kittl (1903: pi. 7, fig. 16).
Plesiotype, T. seminudus, — Kittl (1903: pi. 6, fig. 9).
Plesiotype, T. seminudus, — Kittl (1903: pi. 6, fig. 17).
Figured type, T. rotiformis Kittl (1903: pi. 8, fig. 13).
Plesiotype, T. seminudus, — Kittl (1903: pi. 6, fig. 10).
Figured type, T. repulsus Kittl (1903: pi. 8, fig. 14).
496 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
zz
•
•
u
21
•
•
20
—
•
•
19
—
•
•
X
•
•
•
•
•
•
X
•
x
18
—
•
•
•
• •
•
X
17
—
•
• •
• • • •
•
•
• X
•
X
X
w
16
—
• •
• • •
• • •
• •• • •
•
• ••
X
• •
• • • •
X
" X
X
X
15
—
• • •
•• •
• • • X
X
X
14
—
-
" « X
• •
. r • X ' X
X
X
13
—
•
•
«
• • XX X X
X X X
X
12
t 1
—
•
•
• •
X X '
X *x « « X
• X « XX *
X X X
« » ^ ^^ XX
11
10
—
•
" X X
X X
X
« X
X
X
x"
9
"
X
X
%
-" X
X
X
XX
X
8
•
X
X
7
K
■l
1 1 1
1 1
1
I 1
25 30 35 40 45 50 55 60 65 70
DIAMETER
Figure 39. Variation in whorl width (W) and umbihcal diameter (U) in Tnolilei idnanus, — Mojsisovics, from Werfen For-
mation of Muc, Dalmatic. The data on this graph are from Table 44.
fig. 13, pi. 3, fig. 1; Kittl, 1903: 60, pi. 10, Tirolitcs dancini var. co.stalu.s Kittl, 1903: 62,
fig.s. 4, 5, 11, pi. 11, figs. 1-3, 7; Diener, 1915: pi. 11, fig. 2; Spath, 1934: 372.
278; Patte, 1922: 54, pi. 3, fig. 16; Spath, Tirolitcs darwini var. ahhrcvians Kiltl, 1903: 62;
1934: 372. Spath, 1934: 372.
Tirolites darwini var. cinclti.s Kittl, 1903: 61, pi. Tirolitcs spinosior Kittl, 1903: 62, pi. 11, fig. 5;
10, fig. 4, pi. 11, fig. 3; Spath, 19,34: 372. Dii-ncr, 1915: 279.
Tirolitcs danvini var. rcrninisccns Kittl, 1903: 61, Ccralitcs siiiiri(iiJ.ini Aiu'rhath, 1S71: 50, pi. 4,
pi. 10, fig. 5; Spath, 1934: 372. fi.gs. 9-11.
Tirolitcs dancini var. modcstus Kittl, 1903: 61, Tirolitcs siiiiria^ini, — Mojsisovics, 1882: 73, pi.
pi. 11, fig. 7; Spath, 1931: 372. 81, figs. I, 2; Kittl, 1903: 63, pi. 11, fig. 6;
Ammonoids of the Late Scythian (Lower Triassic) • Kitmmel 497
Diener, 1915: 279; Spoth, 1934: 372.
Tirolitcs kemcri Kittl, 1903: 64, pi. 11, fig. 8;
Diener, 1915: 278; Spath, 1934: 373.
Tiiolites ioulai Kittl, 1903: 64, pi. 11, figs. 11, 12;
Diener, 1915: 280; Spath, 1934: 358, 379.
Tirolites (Scilajitcs) tictzci Kittl, 1903: 66, pi. 10,
fig. 9; Diener, 1915: 281.
Ccratites {Paraceratites) prior Kittl, 1903: 29, pi.
11, figs. 4, 13.
Xenodiscus prior, — Diener, 1915: 314.
Tirolitoidcs p;/or,— Spath, 1934: 378, fig. 127;
Kummel, in Arkell, et al., 1957: L147, fig.
180, 5.
Tirolites hispinatus Ganev, 1966: 25, pi. 1, fig. 5.
The species cassiamis includes all the
species that had been included by Moj-
sisovics and Kittl in the Spinosi. In general
the representatives of this species have
ribs associated with the ventrolateral tu-
bercles. There are, however, gradational
forms to idrianus. Measurements of 31
specimens from the Werfen Formation of
Muc, Dalmatia, are given on Table 45 and
plotted on Figure 4L Very few specimens
of this species in the Kittl collection that
were not figured are sufficiently well pre-
served to yield useful measurements; this
accounts for the fewer measurements of
this species than of idrianus. The many
species of this group were differentiated
on differences in ornamentation and suture.
There appears, however, to be complete
transition among the various so-called
species in these and other characters.
The types of Tirolitoidcs prior owe their
umbilical tubercles to preservation. The
suture likewise is no different from that of
several other tirolitids (Fig. 40B).
The type specimen of Tirolites (Svilajites)
tietzei Kittl, with its ventral cross-ribs, is no
more than a typical tirolitid. Many other
specimens have similar cross-ribs on the
venter though generally not as well de-
veloped as in the type specimen of tietzei.
The binodal aspect of Tirolitcs hispinatus
Ganev (1966) is apparently due to the
crushed nature of the specimen. There is a
plaster cast of the holotype, and only speci-
men, in the Museum of Comparative Zool-
ogy.
Occurrence. Werfen Formation of the
Alps, Dalmatia and the associated regions.
Also reported from eastern Bulgaria, from
southern U.S.S.R. at Bogdo, Mangyshlak
Peninsula, and Tuarkyr.
Repository. The specimens studied here
are from the Kittl ( 1903 ) collection which
is in the Natural History Museum, Vienna.
Tirolites cingulafus Kittl
Plate 70, figures 8-9
Tirolites (Svilajites) cingulatus Kittl, 1903: 65,
pi. 8, fig. 18; Diener, 1915: 280.
Svilajites cingidutiis,—Spath, 1934: 380, fig. 129;
Kummel, in Arkell, et al., 1957: L147, fig.
180, 1.
The type species of Svilajites is cingulatus.
The type specimen measures 37.3 mm in
diameter, 10.3 mm for the width of the
adoral whorl, 14.6 mm for the height, and
the umbilicus is 13.2 mm in diameter. It
is really only on the venter and one side
of the last half volution that the shape and
pattern of ornament are of at least fair
preservation. The prior volutions and the
opposite side from that shown on Plate 70,
figure 9, are completely destroyed by
weathering. The adoral half volution has
two radial ribs extending from the umbili-
cal shoulder to the ventral shoulder. These
ribs presumably cross the venter, but on
the type specimen this region of the venter
is broken. An unfigured paratype in the
collection of the Natural History Museum,
Vienna, is weathered and poorly preserved
but does show the ribs crossing the venter.
The second species that Kittl assigned
to his subgenus Svilajites is tietzei, and
it is illustrated here on Plate 70, figures 11,
12. The type specimen measures 41.7 mm
in diameter, 12.8 mm for the width of the
adoral whorl, 16.6 mm for the height, and
the umbilicus is 16.9 mm in diameter. This
is a typical tirolitid of much better preser-
vation than the type specimen of cingulatus.
There are broad folds extending across
the venter from the ventro-lateral nodes.
There is an indication that these cross folds
decrease in prominence adorally. The gen-
eral features of this species and especially
the folds across the venter are to be seen
498 Bulletin Museum of Comparative Zoologij, Vol. 137, No. 3
H
Figure 40. Diagrammatic representation of the suture of: A, Tirolites toulai KittI (1903: pi. 11, fig. 11), at a diameter of
45 mm; B, paratype Tirolitoides prior (KittI, 1903: pi. 11, fig. 4b), at a diameter of 35 mm; C, Tirolites inipolitus Astakfiovo
|1960a: fig. 16); D, Tirolites elegans Astakhova (1960a: fig. 15); E, Tirolites cf. cass/onus (PI. 34, figs. 9, 10), at a whorl
Ammonoids of the Late Scythian (Lower Triassic) • Kummel
499
Table 45. Measurements of specimens of Tirolites cassianus (Quenstedt) from Mug in Dal-
matia studied by Ernst Kittl, 1903. All specimens are deposited in the Natural History
Museum, Vienna.
D
W
H
U
W/D
H/D
U/D
D
W
H
u
W/D
H/D
U/D
1.
85.0
21.5
28.5
.37.2
26.7
.33.5
43.8
17.
49.8
13.1
17.7
21.0
26.3
,35.5
42.2
0
76.0
16.6
26.5
.34.0
21.8
34.9
44.7
18.
49.0
15.3
1,5.6
21.2
31.2
31.8
43.3
3.
66.0
16.8
22.9
28.2
25.5
.34.7
42.7
19.
49.0
17.0
17.3
20.7
34.7
35.3
42.2
4.
65.0
17.2?
22.5
27.2
26.5?
34.6
41.8
20.
47.7?
?
15.2
22.1
?
31.9?
46.3
5.
59.5
14.8
21.2
26.0
24.9
35.6
43.7
21.
47.1
?
16.3
20.6
?
34.6
43.7
6.
59.3
19.3
20.7
24.2
32.5
.34.9
40.8
22.
46.7
157
17.8
17.7
33.6
38.1
,37.9
7.
58.7
20.0
20.8
25.0
34.1
,35.4
42.6
23.
46.6
?
9.1?
22.8
?
19.5
48.9
8.
57.0
15.5?
20.1
22.4
27.2?
35.3
39.3
24.
46.2?
13.0
15.6
20.7
28.1?
,33.8?
44.8?
9.
56.6
15.3
15.4
21.7
27.0
27.2
,38.3
25.
45.0
11.3
14.2
20.9
25.1
31.6
46.4
10.
56.3
17.7
19.1
24.0
31.4
,33.9
42.6
26.
43.6
11.0?
15.7
19.5
25.2?
36.0
44.7
11.
55.7
14.7
19.3
23.7
26.4
.34.6
42.5
27.
43.5
15.2
15.6
17.2
,34.9
.35.9
,39.5
12.
55.0
17.0
18.6
23.2
.30.9
.33.8
42.2
28.
42.2
?
9.6
18.0
?
22.7
42.7
13.
51.0
17.2
17.3
22.8
33.7
33.9
44^7
29.
40.5
10.3?
13.7
18.4
25.4?
33.8
45.4
14.
50.4
16.3
17.0
20.7
.32.3
33.7
41.1
,30.
.38.4
12.4
12.8
17.4
.32.3
33.3
45.3
15.
50.4
13.5
17.2
20.7
26.8
.34.1
41.1
,31.
36.6
10.1
12.9
14.1
27.6
,35.2
38.5
16.
50.2
10.2
17.6
21.3
20.3
,35.1
42.4
1.
Plesiotype
, T. da
rwiru,-
—Kittl (
1903: pi. 10,
fig. 11).
2.
Lectotyi^e
T. sp
nosior
Kittl (1903: p
1. 11,
fig. 5).
3.
Plesiotype
, T. darwini,—
-Kittl (1903: pi
. 11, fig. 1).
4.
Plesiotype
, T. darwini
I'ar. cos
latus Kittl (1903: pi. 11
, fig.
2).
5.
Syntype,
T. toula
i Kittl
( 1903
pi. 11,
fig. 11).
6.
Pleisotjpe
, T. ha
ueri, —
-Kittl (1903: p
. 9, fig. 10).
7.
Plesiotjpe
, T. turgidus.
—Kittl
(1903:
pi. 10,
fig. 8).
8.
Plesiotype
, T. haueri, —
-Kittl (1903: pi
• 9, fig
. 9).
9.
Lectotype
T. percostatus Kittl
(1903:
pi. 10
, fig. 6).
10,
12, 14,
15. Ur
figured specimens, T.
haueri
,— Kittl (1903
: 56).
11.
Plesiot>pe
, T. spitwsus.
—Kittl
(1903:
pl. 9,
fig. 7).
13.
Plesiot>pe
, T. ha
ueri, —
-Kittl (1903: pi
9, fig
. 13).
16.
Plesiot>i5e
, r. cassianus
,— Kittl
(1903:
pl. 9,
fig. 4).
17.
Figured specimen, T.
darwini
var. modestus
Kittl (1903:
)1. 11, fig
7).
18.
Plesiotype
, T. ha
ueri, —
-Kittl (1903: pi
• 9, fig
. 8).
19.
Plesiotype
, T. ha
ueri, —
-Kittl (1903: p
. 9, fig. 12).
20.
Plesiot>'pe
, T. smiriagin
i,— Kittl
(1903:
pl. 11
, fig. 6).
21.
Lectotype
T. angustilobatus Kittl (1903
: pl. 9,
fig. 3).
22.
Figured specinier
1, T. haueri var. minor Kittl
(1903: p
. 10
, fig. 1).
23."
Figured specimer
, T. angustilobatus var. alpha Kittl (1903:
pl. 9, fig
1).
24.
Figured specimer
, T. darwini
var. rcminiscens Kittl (1903:
pl. 10, fig. 5).
25.
Figured specimer
, T. darwini
var. cinctus Kittl (1903
: pl.
10, fig. 4).
26.
Plesiot>'pe
, T. cassianus
, — Kittl
(1903:
pl. 9,
fig. 6).
27.
Figured specimen
, T. haueri var. minor
Kittl
1903: pl.
10,
fig. 3).
28.
Plesiotype
, T. cassianus
,— Kittl
(1903:
pl. 9,
fig. 5).
29.
Figured specimer
, T. angustilobatus var. alph
a Kittl (1903:
pl. 8, fig
19).
30.
Figured specimer
1, T. haueri var. minor Kittl
(1903: p
. 10
, fig. 2).
31.
Figured specimer
I, T. darwini
var. cinctus Kittl (1903:
pl.
11, fig. 3)
height of 19 mm; F, Tirolites sp. indet. II (PI. 55, figs. 4, 5), at a whorl height of 14 mm (MCZ 9502); G, holotype Tirolites
astakhovi n. sp. (PI. 55, figs. 1, 2), at o diameter of 45 mm (USNM 153081); H, Dinarites do/mofinus,— Kittl (1903: pl. 2,
fig. 4), at a diameter of approximately 40 mm; I, Dinarites dalmatinus, — Kittl (1903: pl. 3, fig. 3), at a diameter of ap-
proximately 30 mm; J, DJnarifes dalmatinus, — Kittl (1903: pl. 2, fig. 2), at a diameter of approximately 26 mm; K, holotype
Ho/o/obus monoptychus (Kittl, 1903: pl. 4, fig. 9), at a diameter of approximately 40 mm; L, holotype Pseudokymatites
svilajanus Kittl (1903: pl. 4, fig. 3), at a diameter of approximately 40 mm; M, Dinarites undatus Astakhova (1960a: fig. 17),
at a diameter of approximately 25 mm; N, Dinarites lialsika:i Renz and Renz (1948: pl. 1, fig. 3c); O, Dor/cron/tes
bogdoanus (Mojsisovics, 1882: pl. 80, fig. 4), at a diameter of approximately 80 mm.
Specimens of figures A, B, H, I, J, K, L from Werfen Formation, Daimatio; of C, D, M from upper Scythian strata Man-
gyshlak Peninsula; of N, from Subco/umbi/es fauna of Chios; of O, from upper Scythian strata Mount Bogdo, southern
Russia; E, Upper Thaynes Formation, Confusion Range, Utah; F, G, Columbiles fauna southeastern Idaho.
500 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
,:)0
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1 1
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40 45 50 55 60 66
DIAMETER
70
75
80
85
Figure 41 .
Formation
Variation in whorl width (W) and umbilical diameter (U) in Tirolites cassianus (Quenstedt), from the Werfen
of Muc, Dalmctia. The data on this graph are from Table 45.
in several of the so-called species of
Tirolites of the Spinosi. The ventral ridges
are related in some degree to the general
prominence of the ventro-lateral nodes.
The species tietzei belongs in the synonymy
of Tirolites cassianus.
The species cin^ulattis is thus known
from only two poorly preserved specimens.
These appear to be distinct from the highly
variable Tirolites cassian\is at a specific
level but not at a gcMieric le\'el as has been
advocated.
Occurrence. Werfen Formation, Muc,
Dalmatia.
Repository. Natural History Museum,
Vienna.
Tirolites rossicus Kiparisova
Text-figure 40
Tirolites ro.ssicus Kiparisova, 1947: 168, pi. 43,
litis. 2, 3, pi. 44, fijjr. 2, te.\t-fi,u. 66: Shcvyrev
and Slilezinj^er, 1960: 1418.
Tirolites clcfimi.s Astakhoxa, 196()a: 150, pi. 35,
by. 1; le.\t-fi.u. 15.
A robust species that in its slightly
Ammonoids of the Late Scythian (Lower Triassic) • Ktimmel 501
prosiradiate ribs on the inner whorls re-
minds one of the specimens assigned to
T. cf. cassiamis from the Tliaynes Forma-
tion of the Confusion Range (PI. 34, figs.
7-10). This species is clearly a member of
the Spinosi and is said by Astakhova ( 1960a )
to be associated with T. cassianus and T.
spinosus. The suture of T. elegans Astak-
hova is illustrated on Figure 40D.
Occurrence. Tirolitcs Zone of Astakhova
(1960a), Mangyshlak Peninsula.
Tirolites impolitus Astakhova
Text-figure 40
Tirolites impolitus Astakhova, 1960a: 151, pi. 35,
fig. 5, text-fig. 16.
This species comes from a horizon ap-
proximately 150 m above that which
yielded T. rossicus. It differs from that
species primarily in having more prominent
ribs and nodes. Too few data are avail-
able as yet on the Mangyshlak fauna. At
the moment it seems best to accept this
species; a great deal more collecting and
study are needed of these forms. The
suture is shown on Figure 40C.
Occurrence. Tirolites Zone of Astakhova
(1960a), Mangyshlak Peninsula.
Tirolifes morpheas (Popov)
Columbites morpheos Popov, 1961, p. 28, pi. 13,
fig. 8.
Popov assigned this species (based on
two specimens) to the genus Columbites
on the strength of an apparent similarity
of the suture to that of Columbites porisi-
anus. Popov, however, was not aware of
the tremendous variation that is present in
the suture of Columbites parisianus (Fig.
22). This suture is quite similar to that of
Tirolites in basic plan. A more reliable
indicator of genetic affinity is the oma-
mentational pattern. In this respect the
species morpheos is clearly a tirolitid.
Popov ( 1961 ) described two other tiro-
litids from Siberia on the basis of single
fragmentary and poorly preserved speci-
mens. Both specimens were collected from
alluvium in the basin of the Kolyma River,
Siberia. These two species — T. ex gr. cas-
sianus, Popov (1961: 29, pi. 13, fig. 7) and
T. gerbaensis Popov (1961: 29, pi. 13, fig.
5) — because of poor preservation and lack
of stratigraphic data are included in the
list of unrecognizable species.
Occurrence. Olenek stage, basin of
Kolvma River, Siberia.
Tirolifes harti Smith
Plate 71, figures 1-7
Tirolites harti Smith, 1932: 83, pi. 57, figs. 9, 10.
Tirolites kniphti Smith, 1932: 84, pi. 57, figs. 1-4.
Tirolites pealei Smith, 1932: 84, pi. 57, figs. 5-8.
Smith ( 1932 ) clearly recognized that the
few fragmentary specimens of Tirolifes he
had from the Tirolites Zone in Paris Canyon
were closely related to, if not conspecific
with, various species of the Spinosi of the
Werfen Fonnation. In this conclusion I am
in complete agreement. However, along with
Smith, I believe it best to keep these Idaho
fonns under a separate species name be-
cause the fragmentary specimens known
to date do not allow any kind of critical
analysis.
Occurrence. Tirolites Zone, Thaynes For-
mation, Paris Canyon, southeast Idaho.
Repository. Holotype, USNM 75022; T.
knighti, holotvpe, USNM 75020a; paratype,
USNM 75020b; T. pealei, holotype USNM
75021a; paratype, USNM 75021; topotypes
MCZ 9641.
Tirolifes smifhi n. sp.
Plate 54, figures 1-5
Tirolites ilh/riciis Mojsisovics, Smith, 1932: 84,
pi. 49, figs. 12-16.
Smith ( 1932 ) had a single specimen from
the Columbites fauna of Paris Canyon,
southeast Idaho, that he assigned to
Mojsisovics' species — illyricus. There is a
general resemblance to T. illyricus as there
is to many other closely similar Werfen
Formation forms. The Idaho species, how-
ever, has a very different aspect. First of
all. Smith's specimen has fine radial ribs
502 Bulletin Museum of Comparative Zoology, Vol 137, No. 3
on the inner whorls, that decrease adorally;
on the outer vokition the ribs are gone and
only sinuous growth lines are present with
widely spaced tubercles at the ventral
shoulders; the venter is highly vaulted.
Two additional specimens have since been
uncovered which are of special interest.
There is first of all a small juvenile form
(PI. 54, figs. 4, 5) measuring 31.4 mm in
diameter, 9.3 mm for the width of the
adoral whorl, 11.2 mm for the height, and
the umbilicus is 12.8 mm in diameter.
The radial ribs on the inner \\'horls and
the gradual adoral decrease in rib promi-
nence are well displayed on this specimen.
The ventral nodes are already well de-
veloped on the outer whorl of this speci-
men. The second specimen is a large one
measuring 102 mm in diameter, 32.7 mm
for the width of the adoral whorl, 39.5 mm
for the height, and the umbilicus is 35.3
mm in diameter. The adoral half \'olution
appears to be body chamber. This portion
of the conch is still characterized by the
prominent ventral tubercles and sinuous
growth lines.
The sum total of the characters of this
form set it apart from any of the Werfen
Formation species of Tirolitcs. It likewise
is totally different from Tirolitcs asfakhovi
which also occurs in the Colunibitcs fauna
of southeastern Idaho but at a different
locality from where this species has been
found.
Occurrence. The holotype (PI. 54, figs.
2, 3) comes from the CoUnuhitcs fauna,
Thaynes Formation, Paris (>anyon, south-
east Idaho. The two plesiotypes recorded
here (PL 54, figs. 1, 4, 5) come from the
same horizons in Montpelier Canyon, south-
east Idaho.
RcjK>sitoni. Holotype (PL 54, figs. 2, 3)
USNM 74993; plesiotypes (Pi. 54, fig. 1)
MCZ 9547, (PI. 54, figs. 4, 5) MCZ 9548.
Tirolifes ostakhovi n. sp.
Plate 55, figures 1-3; Text-figure 40
Three excellently preserved specimens
from the Cohnnhitcs fauna of southeast
Idaho are the basis for this new species.
The measurements of the specimens are as
follows.
D
H
W U
51.5 14.3 ? 26.0
82.4 22.2 ? 42.2
38.8 10.6 9.5 21.8
Holotype USNM 153081
Paratype Unfigured
Paratype USNM 153082
The conch is widely umbilicate with sub-
rectangular whorl sections. The lateral
areas are flattened, and the venter broadly
arched. Both the ventral and umbilical
shoulders are rounded. The lateral areas
bear slightly prosiradiate ribs that begin
on the umbilical shoulder and terminate at
the ventral shoulder in a prominent node.
On the smallest of the available specimens
the nodes are much more prominent than
on the other two specimens. Likewise on
this small specimen the lateral areas are
slightly divergent.
The suture of the holotype is illustrated
on Figure 40G. The first lateral lobe lies
just above the node on the ventral shoulder
and the second lateral lobe lies on the
umbilical shoulder and wall.
The morphological variations possible
within species of the genus Tirolitcs were
impressively illustrated by Kittl in his well-
kno\\'n monograph on the upper W^nfen
fauna from Muc. There is a strong tempta-
tion to assign these forms to one of the
species described by Kittl. In fact, there
is a marked morphological similarity with
practically all of the thirteen species Kittl
(1903: 52, 53) included in the Spinosi.
The separation of this form from the Wvy-
fen species is done more as a matter ol
convenience than of conviction. More
Idaho Columhites Zone material is needed,
and the pattern of intraspeeific variation in
the Werfen species needs to be better
understood before the relationships can be
approached objectively.
Occurrence. From middle shale member
of Thaynes Formation {Columhites fauna),
on hillside north of Sage Check, Stewart
Flat Quadrangle, southeast Idaho.
Repository. Holotype, USNM 153081
Ammonoids of the Late Scythian (Lower Triassic) • Ktimmel
503
(PI. 55, figs. 1-2); figured paratypc, USNM
153082 ( PI. 55, fig. 3 ) ; iinfigured paratype
USGS collections.
Tirolites cf. cassianus (Quenstedt)
Plate 34, figures 7-10; Text-figure 40
Tirolites cf. spinosus Mojsisovics, — Silberling, in
Hose and Repenning, 1959: 2194.
Tirolites aff. haueri Mojsisovics, — Silberling, in
Hose and Repenning, 1959: 2194.
The two specimens recorded here are
clearly representatives of the Spinosi and
could well be conspecific with the Werfen
Formation species — cassianus. These speci-
mens can be matched readily with several
of the specimens illustrated by Kittl ( 1903 ) .
However, because the sample consists of
only two fragmentary specimens it appears
best to do no more than indicate a close
affinity to the Werfen Formation species.
The suture is shown on Figure 40E.
Occurrence. Thaynes Formation, Con-
fusion Range, sample Mill in Hose and
Repenning (1959, p. 2194).
Repository. USNM 153083 (PI. 34, figs.
7, 8), USNM 153084 (PL 34, figs. 9, 10).
Tirolites sp. indet. II
Plate 55, figures 4, 5; Text-figure 40
A single specimen, consisting of approxi-
mately one half volution and much of the
inner whorls, is clearly a tirolitid, but of
uncertain specific affinities. The conch is
evolute, with whorls that are about as wide
as high on the earlier volutions but gradu-
ally become more compressed and higher
than wide. The outer half volution of the
specimen is half phragmocone and half
body chamber. The lateral areas are flat-
tened and the venter is broadly rounded.
Both the ventral and umbilical shoulders
are rounded. The umbilical wall slopes to
the seam at an angle of approximately 45
degrees. The lateral areas bear widely
spaced nodes which lie just dorsal of the
ventral shoulders. There are approximately
seven such nodes on the adoral half volu-
tion, and these show marked increase in
size adorally. The shell bears sinuous
growth lines that are slightly prosiradiate
on the lateral areas and strongly projected
forward in a broad arc over the venter.
The inner whorls, up to a diameter of
10 mm, are approximately as high as wide
and the lateral areas bear radial ribs, some
of \\'hich terminate in weak nodes at the
ventral shoulder. On the next volution, that
is, to a diameter of 20 mm, the whorls
gradually increase in height, and the oma-
ment consists only of widely spaced (four
per half volution) conspicuous nodes at
the umbilical shoulder.
The suture is shown on Figure 40F. The
large first lateral lobe enclosed the ventro-
lateral node, and the second lateral lobe
is on the umbilical wall. It is not difficult
to find, among the numerous Werfen speci-
mens illustrated by Kittl ( 1903 ) , forms
which are quite similar to the one men-
tioned here. However, because I am
dealing with a single specimen, it seems
best to merely document the occurrence
of this form.
Occurrence. Middle shale member of
Thaynes Foimation (Columhites fauna),
Montpelier Canvon, southeast Idaho.
Repository. MCZ 9502.
Genus Diaplococeras Hyatt
Type species, Ceratifes liccanus Hauer, 1865
Diaplococeras liccanum (Hauer)
Ceratites liccanus Hauer, 1865: 616, pi. 3, figs.
1-3.
Dinarites liccanus, — Mojsisovics, 1882: 10, pi. 4,
fig. 1; Tommasi, 1895: 69, pi. 4, fig. 14.
Diaplococeras liccanum, — Hyatt, 1900: 556; Spath,
1934: 379, fig. 128; Kmiimel, in Arkell et al.,
1957: L147, tig. 180,6.
Dinarites (Liccaites) liccanus, — Kittl, 1903: 26;
Diener, 1915: 123.
Hauer ( 1865 ) based his species on one
specimen. Mojsisovics apparently had three
specimens for study; however, none of these
specimens were available for study to the
writer.
Occurrence. Werfen Formation, Muc,
Dalmatia.
504 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Diaplococeras connecfens (Mojsisovics)
Plate 62, figures 1-4
Dinuriics (Ccratites) connectcns Mojsisovics, 1882:
9, pi. 3, fig. 10.
Diiwritcs (Liccaites) connecfens, — Kittl, 1903:
25; Diener, 1915: 123.
Diaplococeras connectcns, — Spath, 1934: 380.
Dinarites circumplicatus Mojsisovics, 1882: 8, pi.
3, figs. 8, 9.
Dinarites [Liccaites) circurn))licatus, — Kittl, 1903:
24; Diener, 1915: 123.
Diaplococeras circumplicatus, — Spath, 1934: 380.
Dinarites hianpulatus Kittl, 1903: 16, pi. 4, fig.
1; Diener, 1915: 120.
Dinarites (Hercegovites) diocletiani Kittl, 1903:
23, pi. 3, fig. 4; Diener, 1915: 122; Spath,
1934: 388.
Dinarites (Liccaites) progressus Kittl, 1903: 26,
pi. 4, fig. 2: Diener, 1915: 123; Spath, 1934:
123.
Dinarites progressus, — Ganev, 1966: 27, pi. 2,
fig. 2.
Poor preservation and misleading illus-
trations account for at least some of the
confusion that has surrounded the species
brought together here. The Werfen For-
mation contains a compressed form of am-
monite that is modestly involute with low
arched venter and conspicuous umbilical
shoulder and umbilical wall. The suture is
ceratitic with two lateral lobes. The flanks
bear slightly prosiradiate ribs. Five species
have been established for ammonites of the
above general design. The major difference
between these "species" is in their state of
preservation. The specimens of the two
species described by Mojsisovics — con-
tiectcns and circuni))lic<ifiis- — were not per-
sonally studied. The three species estab-
lislied by Kittl (1903) were studied in
detail.
The holotyjx' of Dinarites diocletiani is
shown here on Plate 62, figure 1. The
specimen is crushed and the opjiosite side
from that shown in the photograph has
been destroyed by weathering. There art-
low, narrow, prosiradiate ribs on the flanks
that arc most conspicuous at and near the
umbilical shoulder and decrease toward
the ventral shoulder. The ribs also de-
crease in intensitv adorallv.
The holotype of Dinarites hiangulatus
is shown here on Plate 62, figures 3, 4.
This specimen is highly weathered and
only the body chamber approximates the
original size and shape; the phragmocone is
completely distorted by weathering. The
basic outline of the whorls, degree of in-
volution, and pattern of the suture (taking
into account the weathering) is like that
of D. diocletiani. The weathered body
chamber has faint indications of ribs.
The holotype of Dinarites progressus
Kittl, is shown here on Plate 62, figure 2.
As can be seen, it also is a highly weathered
specimen in which all surface features are
obliterated or altered. In shape of whorl
section, degree of involution and suture,
it appears surely to be conspecific with the
other forms included in this species.
The forms brought together here as D.
connectcns differ from D. liccanum in lack-
ing peripheral clavi and umbilical tubercles.
Occurrence. Werfen Formation, Muc,
Dalmatia, and eastern Bulgaria.
Repository. The specimens described by
Kittl are in the Natural History Museum,
Vienna. The specimen described by Ganev
( 1966 ) is in the Geological Institute of the
Bulgarian Academy of Science; a plaster
cast is in the Museum of Comparative Zool-
ogy-
Genus Biffnerites Kittl
Type species, Tirolites (Biffnerites) biffneri
Kittl, 1903
Biffnerites biffneri (Kittl)
Plate 57, figures 1-6
Tirolites (Bittnerites) l)ittneri Kittl, 1903: 67,
pi. 11, fig. 10; Diener, 1915: 280.
Bittnerites iuttncri. — Sp;ith, 1934: 381, fig. 130:
Knnnnel, in Arkell, et al., 1957: 1.147, fig.
180, 3.
Tirolites {Bittnerites) malici Kittl, 1903: 67, pi.
3, fig. 8; Diener, 1915: 280.
Bittnerites malici, — Spath, 1934: .vSl, fig. 130.
Tirolites ( Bitlnerites?) telleri Kittl, 1903: 68, pi.
10, fig. 10; Diener, 1915: 280.
Of the three species Kittl (1903) as-
signed to his new genus Bittnerites, only
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel
505
the illustrated specimen of each species is
still preserved. The genotype specimen —
B. hittncri — is weathered, measuring 50.5
mm in diameter. 12.3 mm for the width of
the adoral whorl, 17.7 mm for the height,
and 23.1 mm for the diameter of the
umbilicus. None of the inner whorls are
preserved. Kittl (1903: pi. 11, fig. 10)
shows the venter to be shaipened; this,
however, is entirely due to weathering.
The normal condition of the venter is
rounded. Tlie large specimen which repre-
sents the type of Bittnerites malici is of
slightly better preservation in that at least
the inner whorls are preserved. This speci-
men measures 71.1 mm in diameter, 17.2
mm for the width of the adoral whorl, 23.1
mm for the height, and the umbilicus is
30.6 mm in diameter. Kittl (1903: pi. 2,
fig. 8) shows prominent prosiradiate shal-
low constrictions at least on the body
chamber of the specimen. A photograph
of the specimen reproduced here on Plate
57, figure 3, shows no such furrows. It
should again be emphasized that preserva-
tion of the Werfen Formation ammonites
leaves much to be desired and fine surface
details are seldom preserved.
The type specimen of Bittnerites telleri
is a very poorly preserved specimen con-
sisting only of the outer volution. It mea-
sures 59.0 mm in diameter, 17.3 mm for
the width of the adoral whorl, 18.7 mm
for the height, and the umbilicus is 25.4
mm in diameter. The prosiradiate shallow
furrows are visible on part of the outer
volution. This specimen differs from the
type specimen of B. malici only in having
slightly more inflated whorls; there is no
justification for keeping these specimens in
separate species.
The genus Bittnerites is known only
from its type species.
Occurrence. Werfen Formation, Muc,
Dalmatia.
Repository. Natural History Museum,
Vienna.
Genus Doricranifes Hyatt, 1889
Type species. Ammonites bogdoanus v.
Buch, 1831
Doricranifes bogdoanus (v. Buch)
Text-figure 40
Annnonites bogdoanus v. Buch, 1831: pi. 2, fig.
1; V. Buch, 1848: 16, pi. 5, figs. 6, 7.
Goniatites bogdoanus, — de Verneuil, in Muichison,
Verneuil and Keyserhng, 1845: 366, pi. 26,
fig. 1.
Ceratites bogdoanus, — Auerbach, 1871: 49, pi.
4, figs. 1-8.
Balatonites bogdoanus, — Mojsisovics, 1882: 87, pi.
80, figs. 1-4.
Dorikianites bogdoanus, — Hyatt, in Whiteaves,
1889: 145; Spath, 1934, p. 382, fig. 131;
Kummel, in Arkell et al., 1957: L147, fig.
180, 2.
Doricranifes l)Ogdoanus, — Diener, 1915: 129;
Kiparisova, 1947: 169, pi. 43, fig. 1; Astakhova,
1960a: 155; Astakhova, 1960b: 149; Shevyrev
and Shlezinger, 1960: 1418; Astakhova, 1962:
70, 75.
Balatonites rossicus Mojsisovics, 1882: 89, pi. 80,
fig. 5.
Dorikranites rossicus, — Hyatt, in Whiteaves, 1889:
145.
Doricranites rossicus, — Diener, 1915: 129; Kipari-
sova, 1947: 170, pi. 43, fig. 4; Astakhova,
1960a: 154, 157; Astakhova, 1960b: 149;
Shevyrev and Shlezinger, 1960: 1418; Astak-
hova, 1962: 75.
Doricranites tumulosus Astakhova, 1960a: 154,
pi. 35, fig. 2, text-figs. 18, 19; Astakhova, 1960b:
149; Astakhova, 1962: 75.
Doricranites lanceolatus Astakhova, 1960a: 155,
pi. 36, fig. 1, text-figs. 20, 21; Astakhova, 1960b:
149.
Doricranites schairicus Astakhova, 1960a: 156,
pi. 36, fig. 2, text-figs. 22, 23; Astakhova,
1960b: 149; Astakhova, 1962: 75.
Doricranites ovatus Astakhova, 1960b: 149.
Doricranites discus Astakhova, 1960b: 149.
Doricranites rarecostatus Astakhova, 1960b: 149.
The criteria used to differentiate these
several species of Doricranites are mainly
ornamentation and shape of the whorl sec-
tion. Examination of the few illustrations
of these species suggests that the degree of
ornamentation and compression of the
conch are highly variable features. One
gets the impression from Astakhovas dis-
cussion of the stratigraphy of the Scythian
formations of the Mangyshlak Peninsula
that Doricranites is a common fossil and
506
Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
that most if not all of the species listed
above occur together. On the basis of
experiences with other ornamented Scythian
ammonoids in which large numbers of
specimens are available, it appears much
more plausible that the species listed in the
synonymy are part of a single species com-
plex characteristic of a particular horizon
in southern U.S.S.R. The only other species
assigned to this genus is D. aciitus which
is considerably more involute. The suture
of D. bo^duanus is illustrated on Figure
40O.
Occurrence. Scythian strata, Mount Bo-
gdo, Mangyshlak Peninsula, and region
of Tuarkyr in Turkmenia, southern U.S.S.R.
Doricranites acufus (Mojsisovics)
Balatouitc.s acufus Mojsisovics, 1882: 89, pi. 80,
fitr. 6.
Dorikranltes acufus, — Ilvatt, in Whiteaves, 1889:
145.
Doricranites acufus. — Diener, 1915: 129; Astak-
hova, 196()a: 159; Schevyrev and Shlezinger,
19W): 1418.
SuJ)cioricranitcs discoidcs Astakliova, 1960a: 158,
pi. 35, figs. 3, 4, text-fig. 24 {nonien nudum
of Bajarunas, 1936).
Sulxloricranifcs orbiculatus Astakhova, 1964: 380,
pl. 1, fig. 2.
This species differs from hogdoanus in
being more compressed, more involute, and
generally with a more subdued pattern of
ornamentation.
Occurrence. This species along with D.
hoiidoanus is apparently ({uite common in
the Doricranites Zone of Astakhova ( 1960a,
b, 1962) at Mount Bogdo, Mangyshlak
Peninsula, and the region of Tuarkyr in
Turkmenia.
Family DINARITIDAE Mojsisovics, 1882
Genus Dinarifes Mojsisovics, 1882
Type species, Ceratites dalmatinus Hauer,
1865
I recognize here four species of Dimiritcs,
all confined to the western region of Tethys.
The genus is an important member of the
Werfen fauna where it is rejirescMited by
two species. One of these (carniolicus)
is the type species of CarnioUtes; this genus
is considered to be a synonym of Dinorites.
The presence of Dinorites dalmatinus in the
Sidjcohimhites fauna of Chios is an im-
portant link in dating the Werfen fauna.
The other two species of Dinarites {liat-
sikasi and undatus) are quite distinct from
dalmatinus and carniolicus but closely re-
lated to each other. Dinarites liatsikasi is
from the Subcolumhites fauna of Chios
and D. undatus from the upper Scythian
formations of the Mangyshlak Peninsula.
Dinarites dalmatinus (Hauer)
Plate 58, figures 1-10; Plate 59,
figures 1-11; Plate 60, figures 1-8;
Text-figure 40
Cerafitcs dalmafinus Hauer, 1865: 615, pl. 2,
figs. 3, 4.
Dinarifes dalmatinus, — Mojsisovics, 1882: 8, pl.
1, figs. 7, 8; Kittl, 1903: 18, pl. 2, figs. 1-11,
pl. 3, figs. 1, 2; Hyatt and Smith, 1905: 162;
Arthaber, in Freeh, 1906: pl. 34, fig. 17; Diener,
1915: 120; Kumniel, in Arkell, et al., 1957:
L148, fig. 181, 1; C.anev, 1961: 182, pl. 2,
figs. 4, 5, 8, pl. 4, fig. 6, pl. 6, fig. 2.
Dinarites dahnatinus var. extcnsus Kittl, 1903:
20, pl. 2, figs. 8, 9.
Dinarifes dahnatinus var. i)Iurimcostafus Kittl,
1903: 20, pl. 2, figs. 10, 11.
Dinarifes dalmatinus var. externeplanafus Kittl,
1903: 20, pl. 2, figs. 1, 2.
Plueoccras dalmaiinum, — Hvatt, 1900: 556: Spath,
1934: 388, fig. 134.
Dinarites nudus Mojsiso\ics, 1882: 6, pl. 1, figs.
5, 6; Kittl, 1903: 17, pl. 1, figs. 11-13; Diener,
1915: 122; Renz and Renz, 1948: 48, pl. 1,
figs. 1, 2.
Dinarites laevis Toniniasi, 1902: 347, pl. 13, figs.
4, 5; Kittl, 1903: 13, pl. 1, figs. 1-3, pl. 3, figs.
10, 11; Diener, 1915: 121; Spath, 1934: 386.
Ceratites inucliianus Hauer, 1865: 613, pl. 2,
figs. 5, 6.
Dinarites inucliianus, — Mojsis()\ ics, 1882: 6, pl. 1,
fig. 4; Kittl, 1903: 15, pl. 1, figs. 4-8; Artha-
ber, in Freeh, 1906: pl. 34, fig. 16; Witten-
burg, 1908: 285, pl. 40, fig. 20; Diener, 1915:
121; Spath, 19.34: .386, fig. 1,32; Canev, 1966:
26, pl. 1, figs. 3a-e.
Dinarifes evolutior Kittl, 1903: 16, pl. 1, figs. 9,
10; Diener, 1915: 121; Spath, 1934: ^384;
Renz and Renz, 1948: 49, pl. 1. fig. 4;
K()ll;irova-Andrusovov;i, 1961: 29, pl. 3, fig. 2.
Dinarites (?) an^ulatus Kittl, 1903: 22, pl. 3,
fig. 9.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel
507
Dinarites miilticostattis Kittl, 1903: 21, pi. 3, fig.
3; Diener, 1915: 122.
Dinarites tiwlitoides Kittl, 1903: 21, pi. 7, figs.
1-3; Diener, 1915: 122.
Dinarites bulgaricns Berndt, 1934: 8, pi. 2, fig. 8.
Aside from Tirolites the most common
element in the Werfen Formation fauna of
the Alps and associated regions is Dinarites.
A number of species names have been in-
troduced for the dinaritids of this region
but basically there are two main species,
the first, encompassing predominantly
smooth forms (miickiamis), and the second,
ornamented forms with ribs (dalmatinus).
This second species had been set aside
early (Hyatt, 1900) as the type of PIoco-
ceras. The latter genus had been accepted
by Diener (1915) and Spath (1934). How-
ever, the selection of dalmatinus as the
type of Dinarites Mojsisovics (1882), by
Hyatt and Smith (1905, p. 162), makes
Flococeras a synonym of Dinarites. Whereas
the earlier authors wished to separate the
smooth and ornamented dinaritids into
distinct genera, my own studies bring me
to the conclusion that we are dealing here
with a single species complex rather than
several species in two different genera.
The largest single collection of Werfen
Formation dinaritids is that studied by
Kittl ( 1903 ) and on deposit in the Natural
History Museum, Vienna. The measure-
ments of 38 specimens are listed on Table
46 and plotted on Figure 42. These 38
specimens represent the most complete
and best preserved specimens available to
Kittl, who illustrated most of them. Practi-
cally all of the remaining specimens that
Kittl assigned to this or that species of
dinaritid are very poorly preserved and of
somewhat doubtful value. Kittl's illustra-
tions are line drawings of better than
average quality but yet lacking in not
conveying the nature of preservation, im-
perfections in the conch, and at times with
errors in artistry. All of the earlier illustra-
tions for the Werfen Formation dinaritids
are line drawings (e.g. Mojsisovics, 1882),
and most of the more modem illustrations
are impossible to decipher due to faulty
printing and poor paper. Plates 58, 59,
and 60 have photographic prints of the
principal dinaritids studied by Kittl ( 1903 ) .
The smoothness of some of the specimens
is clearly due to weathering and preserva-
tion. The number and intensity of the ribs
is highly variable. Finally, there is a fair
degree of variation in the diameter of the
umbilicus (Fig. 42). Study of all of Kittl's
specimens assigned to these species of
dinaritids (smooth-muchianus; rihhed-dal-
matinus) leads me to believe there is com-
plete gradation from smooth to strongly
ribbed forms. This observation plus the
fact that all combinations of the morpho-
logical grades are found in nearly every
fossiliferous localitv in the Werfen Forma-
tion brings me to the conclusion we are
dealing with a single, variable species.
The suture is illustrated on Figure 40H-J.
The dinaritids are a very small element
in the SidjcoJumhites fauna of Chios. Renz
and Renz (1948) record two specimens of
Dinarites niidtis Mojsisovics and two speci-
mens of Dinarites evolutior Kittl. These
specimens fall well within the range of
variability recognized for Dinarites dal-
matinus and are thus considered synonyms.
The two specimens assigned to Dinarites
liatsikasi (Renz and Renz, 1948: 49, pi. 1,
fig. 3) are of special interest for the close
similarity they show to Dinarites undatus
Astakhova (1960a) from the Mangyshlak
Peninsula. Each of these species is distinct
from D. dalmatinus in ornamentation and
in suture.
Occurrence. Primarily in Werfen Forma-
tion of the Alps and related regions. A few
specimens are known from the Subcolum-
hites fauna of Chios.
Repository. The Werfen Formation speci-
mens are in the Natural History Museum,
Vienna; those from the Subcolumbites
fauna of Chios are in the Natural Histoiy
Museum, Basel, NHMB J13683-6; the
specimens from eastern Bulgaria recorded
by Ganev (1961, 1966) are in the Geologi-
cal Institute of the Bulgarian Academy of
508
Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Table 46. Measurements of Dinarites dalmatinus (Hauer) from Werfen Formation, Muc,
Dalmatia. All specimens are in Natural History Museu':m, Vienna; data plotted on gfl\ph o"
Figure 42.
D
w
H
u
\V/D
H/D
U/D
D
w
H
u
W/D
H/D
U/D
1.
63.7?
16.7
29.5
16.4
26.2
46.3
25.7
20.
45.4
14.4?
18.8
11.7
31.8
41.5
25.8
2.
59.7
?
27.1
14.4
p
45.5
24.1
21.
45.0
14.0
18.7
13.1
31.1
41.5
29.1
3.
58.7
16.8
26.0
16.4
28.6
44.3
27.9
22.
44.5
10.1
17.3
15.3
22.7
38.9
34.4
4.
56.4
?
23.9
15.7
?
42.4
27.8
23.
44.3
?
20.4
10.1
p
46.0
22.8
5.
56.2
?
25.3
13.0
?
45.0
23.1
24.
44.2
?
20.5
10.8
p
46.4
24.4
6.
55.6
16.6?
22.6
17.5
29.9
40.6
31.5
25.
44.0
8.5
20.7
8.4
19.3
47.0
19.1
7.
54.8?
?
24.8
14.7
?
45.1
26.8
26.
43.5
11.8?
17.4
11.0
27.1
40.0
25.3
S.
53.8
12.3
22.5
13.7
22.9
41.8
25.5
27.
42.8
13.3
16.8
12.8
31.1
39.3
29.9
9.
53.0
12.7
22.7
?
24.0
42.8
?
28.
41.8
p
20.1
6.4
?
48.1
15.3
10.
52.8
14.5
24.4
11.3
27.5
46.2
21.4
29.
41.7
13.0
17.2
11.9
31.2
41.2
28.4
11.
51.4
16.6
19.8
17.7
32.4
38.6
34.5
30.
41.4
9.5?
17.8
11.3
23.0
43.0
27.3
12.
51.4
11.8
23.0
11.2
22.9
44.7
21.8
31.
41.3
13.2
17.7
11.7
32.0
42.9
28.4
13.
50.6?
10.4?
20.8
9.1
20.6
41.1
18.0
32.
41.1
11.2
18.3
9.5
27.3
44.5
23.1
14.
49.0
14.1
21.1
15.0?
28.8
43.1
30.6
33.
39.5
12.3?
16.5
10.8
31.1
41.8
27.3
15.
47.8
?
21.2
12.8
?
44.4
26.8
34.
39.4
12.3
16.0
11.5
31.2
40.6
29.2
16.
47.4
?
24.6?
5.9?
?
52.0
32.3
35.
38.9
9.0
19.4
6.7
23.1
49.9
17.2
17.
47.0?
13.0
23.7
10.2
27.6
50.5
21.7
36.
37.5
6.8
16.3
10.0
18.1
43.5
26.6
18.
46.6
?
20.5
10.2
?
44.0
21.9
37.
34.7
p
15.5
9.2
p
44.7
26.5
19.
45.5
11.4
19.2
10.9
25.1
42.2
24.0
38.
26.2
6.3
13.5
3.8
24.4
51.5
14.5
1.
2
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
27.
28.
29.
30.
31.
32.
33.
34.
35.
36.
37.
38.
Plesiotvpe, — Kittl ( 1 903 :
Plesintvpe, — Kittl (1903:
Plesiotype, — Kittl (1903,
Plesiotvpe, — Kittl ( 1903,
pl. 2, fig.
pl. 2, fig.
pl. 2, fig.
pl. 2, fig.
5).
6).
4).
7).
Unfigured parat\pe, Dinarites
Unfigured parat\pe, Dinarites
Plesiotype, — Kittl (1903: pl.
Unfigured paratype. Dinarites
cvohitior Kittl.
mtdticoslatiis Kittl.
2, fig. 1).
evohitior Kittl.
fig.
pl. 1,
pl. 7
pl. 1,
10).
fi«.
, fig.
fiR.
1, fig. 1).
Syntyi^e, Dinarites evohitior Kittl (1903: pl. 1
Plesiotyjje, Dinarites mttchianus, — Kittl ( 1903:
Figured type, Dinarites tirolitoidcs Kittl (1903
Plesiotyiie, Dinarites nntchianus, — Kittl (1903:
Syntype, Dinarites cvolutior Kittl (1903: pl. 1, fig. 9).
Unfigured paratype, Dinarites tirolitoides Kittl.
Unfigured i^aratype, Dinarites cvohitior Kittl.
Plesiotvpe, Dinarites lacvis, — Kittl (1903: pl
Plesiotype, — Kittl (1903: pl. 2, fig. 8).
Plesiotype, Dinarites rnucliianns, — Kittl (1903: pl. 1, fig.
Plesiotyi^e, Dinarites nnchis, — Kittl (1903: pl. 1, fig. 13).
Figured type, Dinarites tiroUtoides Kittl (1903: pl. 7, fig.
Plesiot\i>e, — Kittl (1903: pl. 2, fig. 9).
Figured type, Dinarites niulticostatus Kittl (1903: i)
Unfigured paratvpe, Dinarites evohitior Kittl.
Plesiotype, — Kittl (1903: pl. 2, fig. 11).
Plesiotype, Dinarites laevis, — Kittl (1903: pl. 3,
Plesiotype, — Kittl (1903: pl. 2, fig. 2).
Unfigured paratype, Dinarites firohtoides Kittl.
Plesiotype, Dinarites laevis, — Kittl (1903: pl. 1,
Plesiotype, — Kittl (1903: pl. 3, fig. 1).
Plesiotype, — Kittl (1903: jil. 2, fig. 10).
Plesiotyiie, — Kittl (1903: pl. 3, fig. 2).
Plesiotype, Dinarites inurliianiis, — Kittl (1903:
Plesiotype, Dinarites nndns, — Kittl (1903: pl. 1
Figured type, Dinarites tirolitoides Kittl ( 1903:
Plesiotype, Dinarites laevis,- -Kiltl (1903: i)l.
Plesiotype, Dinarites laevis, — Kittl ( 1903: pl.
Unfigured paratype, Dinarites evohilior Kittl.
Plesiotype, Dinarites iniichianus, — Kittl (1903: pl. 1, fig.
6).
2).
8).
7).
3).
fig. 3).
ti
11).
lig. 2).
pl. 1,
fig.
5).
, fi.U.
11).
pl. 7,
tig.
1).
1, fig.
3).
3, fig
10)
4).
Ammonoids ok the Late Scythian (Lower Triassic) • Kummel 509
30
25
20
0
H
U
• •
30 35
40 45 50
DIAMETER
55
60
_l
65
Figure 42. Variation in whorl height (H) and umbilical diameter (U) of Dinarites dalmatinus (Hauer), from Werfen Forma-
tion, Muc, Dalmatia. The data on this graph are from Table 46.
510 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Table 47. Measubements of specimens of Di-
narites carniolicus ( mojsisovics ) from the
Werfen Formation, Dalmatia, figured by
KiTTL (1903). All specimens are in the Natu-
ral History Museum, Vienna.
D
w
H
u
W/D
H/D
U/D
1.
59.3
11.7
26.8
11.2
19.7
45.2
19.2
2.
58.5
?
24.6
15.5
?
42.1
26.5
3.
57.2
?
23.0
16.2
?
40.0
28.3
4.
55.2
10.0±
23.2
10.4
18.1±
42.0
18.8
5.
53.3
12.0?
24.2
12.2
22.5?
45.4
22.9
6.
52.7
10.5?
26.2
10.1
19.9
49.7
19.2
7.
52.6
12.0?
23.7?
10.3?
22.8
45.1?
19.6?
1. PlesioUpe, — Kittl (1903: pi. 5, fig. 1).
2. Plesiot\pe, — Kittl (1903: pi. 5, fig. 2).
3. Holotvpe, Tirolites heterophanua Kittl (1903: pi. 5,
fig. 7).
4. Svntv'pe, Tirolites scrratelobatus Kittl (1903: pi. 5,
fig. 6).
5. Ple.siot\pe, — Kittl (1903: pi. 5, fig. 3).
6. Svntyi3e, Tirolites scrratelobatus Kittl (1903: pi. 5,
fig. 5).
7. Plesintype,— Kittl (1903: pi. 5, fig. 4).
Science; plaster casts are in the Museum
of Comparative Zoology.
Dinarites carniolicus (Mojsisovics)
Plate 61, figures 1-8
Tirolites cariiioliciis Mojsisovics, 1882: 65, pi. 1,
figs. 2, 3; Kittl, 1903: 35, pi. 5, figs. 1-4;
Diener, 1915: 277.
Carniolitcs carniolicus, — Arthaber, 1911: 241, 250;
Spath, 1934: 392, fig. 136; Kummel, in Aikell,
et al., 1957: L148, fig. 181,3; Kollarova-Andm-
sovova, 1962: 31, pi. 2, figs. 6a-c, pi. 3, fig. 4.
Tirolites serrcitelohahis Kittl, 1903: 36, pi. 5, figs.
5, 6; Arthaber, 1911: 250; Diener, 1915: 279.
Tirolites heterophuniis Kittl, 1903: 38, pi. 5, fig.
7; Arthaber, 1911: 250.
This species differs from the "smooth"
class of dinaritids merely in the presence
of fairly prominent tubercles near the ven-
tral shoulder on the adoral part of the
phragmocone and body chamber. I can
see no justification for a separation at the
generic level. Kittls types of the species
brought together here as D. carniolicus are
illustrated here on Plate 61. The measure-
ments of these specimens are shown on
Table 47.
Occurrence. The specimens studied here
are from the Werfen Fonnation, Muc, Dal-
matia. The species is also recorded from
the same formation in Czechoslovakia
( Kollarova-Andrusovova, 1961, 1962).
Repository. The specimens from Muc,
studied here, are in the Natural History
Museum, Vienna.
Dinarifes liatsikasi Renz and Renz
Text-figure 40
Dinarites liatsikasi Renz and Renz, 1947: 60, 75;
Renz and Renz, 1948: 49, pi. 1, fig. 3.
Ornamentation and suture (Fig. 40N)
differentiate this species from Dinarites
dahuatintis. The ribbing consists of slightly
prosiradiate folds that cross a subtruncate
venter. The suture is goniatitic but with a
very broad first lateral lobe. The holotype
measures 35.7 mm in diameter, 11.2 mm
for the width of the adoral whorl, 16.2 mm
for the height, and 9.8 mm for the diameter
of the umbilicus. This species is very simi-
lar to Dinarites undatus Astakhova from
the Mangyshlak Peninsula. In that species
the radial ribs are slightly sinuous and are
somewhat enlarged at the umbilical shoul-
der. The suture, likewise, has a much
narrower first lateral lobe. There is a su-
perficial similarity in conch form to Cucoco-
ceras as indicated by Renz and Renz ( 1948 )
but the suture is completely different.
Occurrence. Subcolumhites fauna, Chios.
Repository. Holotype NHMB J 13687,
paratypes NHMB J 13688.
Dinarites undafus Astakhova
Text-figure 40
Dinarites undatus Astakho\a, 196()a: 152, pi. 34,
fig. 9, text-fig. 17.
A species of the general design and form
of D. liatsikasi from the Subcohnnbites
fauna of Chios. The general features are
discussed under the discussion of that spe-
cies. The suture is illustrated on Figure
4()M.
Occurrence. Tirolites Zone of Astakhova
(196()a), Mangyshlak Peninsula.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 511
Genus Hololobus (KittI)
Type species, Tirolites (Hololobus) mono-
pfychus KittI, 1903
Hololobus monopfychus (KittI)
Plate 70, figures 1, 2; Text-figure 40
Tirolites (Hololobus) monoptychus KittI, 1903: 33,
pi. 4, fiff. 9; Diener, 1915: 280.
Hololobus monoptychus, — Spath, 1934: 390, fig.
135; Kummel, in Arkell, et al., 1957: L148, fig.
181,5.
The type, and only specimen, of this
species and genus is of only fair preserva-
tion. It measures 61.8 mm in diameter, 15.7
mm for the width of the adoral whorl,
26.7 mm for the height, and the umbilicus
is 16.8 mm in diameter. The critical point
of interest with this species is the nature
of the ventral lobe. Examination of the
type specimen clearly shows that Kittl's
drawing of the suture (Fig. 40K) is ac-
curate. Like KittI, I could not observe any
trace of a ventral siphuncle, but this I
attribute more to faulty preservation than
to anything else.
Occurrence. Werfen Formation, Muc,
Dalmatia.
Repository. Natural History Museum,
Vienna.
Genus Pseudodinarifes Hyatt
Type species, Dinarifes mohamedanus
Mojsisovics, 1882
Pseudodinarifes mohamedanus (Mojsisovics)
Plate 57, figures 7-9; Plate 62, figure 6
Dinarites mohamedanus Mojsisovics, 1882: 7, pi.
40, fig. 12.
Pseudodinaiites mohamedanus, — Hyatt, 1900: 559:
Spath, 1934: 387, fig. 133; Kummel, in Arkell
etal., 1957: L148.
Dinarites (Hercegovites) mohamedanus, — KittI,
1903: 22, pi. 3, figs. 5-7; Diener, 1915: 122.
This species is supposedly characterized
by an evolute, smooth conch with rounded
but sHghtly compressed whorls and suture
with ceratitic lobes. Mojsisovics' type speci-
men was not studied; the specimens as-
signed to this species by KittI are illustrated
here. These specimens are so poorly pre-
served they add little to our understanding
of this genus or species. The few speci-
mens of this species have all come from the
Werfen Formation of the Alps and adjacent
regions. Simionescu (1908: 161) recorded
a specimen as Dinarites cfr. mohamedanus
from eastern Rumania. This is the only
species that has been assigned to this
genus. It is quite clear much more data
are needed to evaluate the generic relations
of the species.
Occurrence. Kittl's specimens illustrated
here are from the Werfen Formation, Muc,
Dalmatia.
Repository. Kittl's specimens are in the
Natural History Museum, Vienna.
Family HELLENITIDAE Kummel, 1952
Genus Hellenites Renz and Renz, 1947
Type species, Tropicelfiies praematurus
Arthaber, 1911
Evolute forms with subquadrate whorl
section. Lateral sides with rectiradiate to
radial ribs which curve adorally on ventral
shoulder forming an acute junction with a
median ventral keel; ribs may or may not
cross keel. Suture with simple pronged
ventral lobe, large, denticulated lateral
lobe and very simple second lateral lobe,
saddles rounded.
This extremely interesting and somewhat
anomalous group of late Scythian am-
monoids has been the source of an evolving
confusion. The group was first recognized
by Arthaber (1911) on the basis of two
specimens from the Subcolumbites fauna
of Albania. Neither of these two specimens
had the suture preserved, and on the basis
of the conch form and ornamentation, he
placed them in the genus Tropiceltites, a
common late Triassic (Carnian) group. A
more complete understanding of Arthaber's
species was not possible until Renz and
Renz ( 1948 ) described and illustrated a
number of conspecific forms from the Sub-
columbites fauna of Chios. The suture is
512 BtiUetin Museum of Comparative Zoology, Vol. 137, No. 3
well preserved on these Chios specimens
and shows that it is very different from the
suture of the late Triassic Tropiceltites. On
this basis, Renz and Renz erected the genus
HcUenites.
Neither of these authors, however, had
taken note of Psctidlmrpocems .s/>/n/gc'r
Waagen (1895: 130, pi. 21, fig. 1). This
species was based on a single specimen
from a horizon Waagen considered to be
near the top of his Ceratite formation in
the Sheik-Budin Hills of the Trans-Indus
Region. Spath (1951: 8) correctly pointed
out that the specimens may belong to an
Upper Triassic hildoceratid or some other
related form.
J. P. Smith (1932) assigned a single
specimen from the CoJumhiics fauna of
southeastern Idaho to the genus Fscudhar-
poccras. Spath (1951: 9) made the claim,
and correctly so, that since Vscudharpo-
ceras was based on a poorly preserved
specimen of uncertain stratigraphic posi-
tion this genus should be rejected. He then
introduced the genus Fscudarniotitcs with
Psendliarpoccras idaliocnse Smith (1932:
81, pi. 49, figs. 17-19) as type. Spath was
at this time unaware of the genus Hel-
lenites Renz and Renz, 1948. Fseudharpo-
ccras idahoensc is clearly a species of Hel-
Icnifcs.
The specimen Waagen described and
illustrated as Fseudharpoccras spiniiicr is
not in the collections of the Geological Sur-
vey of India. There is, however, a partial
mold of one side plus a plaster cast. These
have been illustrated and discussed by
Kummel (1966) and his conclusion was
similar to that of Spath: that the genus and
species should be rejected.
The Subcohn}d)il('s fauna of Chios has
two species of this genus. One or lioth of
these two species are recognized from the
same horizon in Kwangsi, (]hina, Irom the
Primorye Region, eastern Siberia, and the
Tobin Formation of Nevada. HcUenites
idahoense occurs in the Culumhites fauna
of southeastern Idaho.
Taele 48. Measurements of Hellenites prae-
MATURUS (ArthABEr) FROM SuBCOLUMBITES
FAUNAS, Albania and Chios.
D
w
H
u
W/D
H/D
U/D
1.
55.2
13.2
17.7
25.5
23.9
32.1
46.2
2.
53.4
14.3
15.0
29.0
26.6
27.2
,54.5
3.
52.3
15.9
16.8
26.2
30.4
.32.2
50.1
4.
48.2
12.2
14.0
23.0
25.3
29.1
47.8
5.
47.0?
13.2
14.5?
22.4
28.1?
.30.9?
47.6?
6.
46.2
11.7?
13.8
18.6
25.4?
29.9
41.0
7.
44.2
11.6?
14.4
20.5
27.2?
.32.6
46.4
8.
43.3
13.0
13.2
21.1
30.0
30.6
48.7
9.
42.1
12.1?
12.8
20.2
28.7?
.30.4
47.9?
10.
36.8
10.3
10.5
17.1
28.0
28.5
46.5
11.
36.7
10.7
10.7
17.4
29.2
29.2
47.4
12.
36.3
9.6?
11.9
16.6
26.4?
.32.8
45.7
13.
36.2
9.2
10.1
19.2
25.4
27.9
53.0
14.
35.8
10.8
11.0
16.0
30.2
30.7
44.7
15.
34.1
10.7
10.4
14.6?
31.4
.30.5
42.8
16.
33.4
10.1
11.1?
14.5
.30.2
33.2?
43.4
17.
33.0
9.8
10.9
14.7
29.7
33.0
44.5
18.
25.7
8.0
6.8
13.0
31.1
26.5
50.6
19.
25.7
8.2?
9.1
10.2
31.9?
.35.4
.39.0
20.
25.5
7.5
7.5
10.8?
29.4
29.4
42.4?
21.
23.3
6.8
7.7
9.6
29.2
.33.0
41.2
22.
19.1
7.8?
6.2
8.8
40.8?
32.4
46.0
23.
16.6
6.1
6.0
7.0
36.7
.36.1
42.2
24.
13.8
6.3
4.8
5.6?
45.7
34.8
40.6?
1.
•1
3,
4,
5.
6,
S.
11.
12.
13.
19.
S\aitvpe, H. praemiiturus var. acgaeica Renz and Renz
(1947: 60; 1948, pi. 2, fig. 6), NHMB J13664.
Holotvpe, H. trikkalinoi Renz and Renz (1947: 60,
75; 1948, pi. 2, fig. 2), NHMB J 13668.
9, 1.5, 18, 24. Unfigured parat\i5es, H. trikkalinoi,
Maradovuno, Chios, NHMB J 13672.
23. I'nfigiired paratypes, Kephal<ivuno, Chios, NHMB
J13673.
Svntvpe. H. praetnaturus var. acgaeica Renz and Renz
(1947: 60; 1948, pi. 2, fig. 8), NHMB J13665.
10, 14, 16, 17, 20, 21. Unfigured parat\pes,
Maradovuno, Chios, NHMB .113662.
T\pe specimen, H. trikkalinoi var. graeca Renz and
Renz (1947: 60; 1948, pi. 2, fig. 5), NHMB J13674.
Plesiotype, — Renz and Renz (1948: pi. 2, fig. 7),
NHMB J1366().
Plesiotype, — Renz and Renz (1948: pi. 2, fig. 3),
NHMB J13661.
Sviitspe, H. praentaturus var. acgaeica Renz and Renz
(1947: 60; 1948, pi. 2, fig. 9), NHMB J13667.
Paratvpe, H. trikkalinoi Renz and Renz (1948: pi. 2,
fig. 4), NHMB J 13671.
Holotvpe, H. pracmatiirti.s
(8), 'figs. 9a, b), PIUV.
HoloUpe, H. pracnuiiiirtis
(Arthaber, 1911: pi. 24
var. (Arthaber, 1911: pi.
24(8), figs. 10a, b), PIUV.
Hellenites praematurus (Arthaber)
Plate 7, figures 1-4; Text-figure 43
Tropiceltites praciuatunis .Arthaber, 1911: 268, pi.
24(8), liys. 9a, 1): C. Hen/, 1928: 1.55.
Tr()))iccltilc.s ? ]ircieiu(itiirns var. Artliaber, 1911:
269, pi. 24(8), fifrs. 10a, 1).
Tr()i>iccllit('s (?) )>r(i('i)ialunis. — Diener, 1915: 300.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 5L3
A
C
Figure 43. Diagrammatic representation of the suture of: A, Hellenites tnkkalmol Renz and Renz (1948: pi. 2, fig. 2b),
at a diameter of 35 mm; B, H. praemofurus (Arthaber),— Renz and Renz (1948: pi. 2, fig. 8b), at a diameter of 30 mm; C,
H. radlatus Renz and Renz (1948: pi. 2, fig. 13b), at a diameter of 20 mm; D, H. idahoense (Smith) (USNM 74994), at- a
diameter of 1 3 mm.
Specimens of figures A-C from Subco/umbifes fauna, Chios; specimen of figure D from Co/umb/tes fauna of southeast
Idaho.
Hellenites praematunis, — Renz and Renz, 1947:
60, 75; Renz and Renz, 1948: 44, pi. 2, figs.
3-3a, 7-7a; Kummel, in Arkell et al., 1957:
L149, figs. 181, 6a-c.
Hellenites praematuriis var. aegaeica Renz and
Renz, 1947: 60; Renz and Renz, 1948: 45, pi.
2, figs. 6-6b, 8-8b, 9-9a.
Hellenites trikkalinoi Renz and Renz, 1947: 60,
75; Renz and Renz, 1948: 46, pi. 2, figs. 1-la,
2-2b, 4-4b.
Hellenites trikkalinoi var. graeca Renz and Renz,
1947: 60; Renz and Renz, 1948: 46, pi. 2, figs.
5-5b.
Hellenites cf. praemattinis. — Chao, 1959: 145, pi.
41, figs. 1, 2 (not 3, 4).
This species was established by Arthaber
on the basis of two specimens, one of which
he considered to be a variety. The preser-
vation of both of these specimens leaves
much to be desired. The measurements
are as follows:
D W H U W/D H/D U/D
25.7 8.2? 9.1 10.2 31.9? 35.4 39.7
Holotype— Arthaber, 1911: pi. 24(8),
fig. 9a, b.
19.1 7.8? 6.2 8.8 40.8? 32.5 46.1
Paratype— Arthaber, 1911: pi. 24(8), fig.
10 a, b.
The suture is not preserved on either of
these specimens.
This anomalous late Scythian species
remained somewhat of a problem until the
discovery of the contemporaneous Sith-
coJumbites fauna from Chios. A number of
specimens of H. praematunis have been
well illustrated by Renz and Renz ( 1948 ) .
In addition to H. praematunis, these au-
thors recognized a variety and another
species based mainly on the relative promi-
nence of the ribs. It is apparent on exam-
ination of the large number of specimens
in the fauna studied by Renz and Renz
that there is wide variation in rib promi-
nence, and that there are gradational forms
connecting the species and varieties rec-
ognized. In the other morphological fea-
tures, as width, height and vimbilical di-
ameter, there is relativelv little variation
( see Table 48 and Figure 44 ) . Tlie sutures
are illustrated on Figures 43 A, B.
Chao (1959) obtained two fragmentaiy
514 Bulletin Museum of Comparative Zoology, Vol 137, No. 3
specimens from separate localities of the
SubcoUimhites horizon, which he described
as H. cf. pracmaturus. One of these speci-
mens (Chao, 1959: pi. 41, figs. 1, 2) is
identical in rib pattern, conch shape, etc.,
to H. praeniaturus. Though a suture is not
preserved on this specimen, I believe it to
be a valid representative of H. praema-
turus. The second specimen is more frag-
mentary and of poorer preservation. The
rib pattern, however, compares favorably
with H. radiatus.
Occurrence. Suhcohimhites faunas of Al-
bania, Chios, and Kwangsi, China (Chao
collections 542a, 546).
Repository. Holotype and paratype, in
the Paleontological Institute, Vienna; spec-
imens from Chios, plesiotypes, Renz and
Renz (1948: pi. 2, figs. 3-3a) NHMB
J13661, (1948: pi. 2, figs. 7-7a) NHMB
J 13660; unfigiued specimens from Mara-
dovuno NHMB J13662; unfigured speci-
mens from Kephalovuno NHMB J13663;
type specimens, //. praematxirus var. aegae-
ica Renz and Renz (1948: pi. 2, fig. 6)
NHMB J13664, (pi. 2, figs. 8-8a) NHMB
J13665, (pi. 2, fig. 8b) NHMB J13666, (pi.
2, fig. 9) NHMB J 13667; holotype H. trik-
kalinoi Renz and Renz (1948: pi. 2, fig.
2-2a) NHMB J 13668; paratypes, Renz and
Renz (1948: pj. 2, fig. 1) NHMB J13670,
(pi. 2, fig. 4) NHMB J13671, (pi. 2, fig.
2b) NHMB J13669; unfigured paratypes
from Maradoxuno NHMB J13672, from
Kephalovuno NHMB J13673; type speci-
men //. trikkalinoi var. groeco Renz and
Renz (1948: pi. 2, fig. 5) NHMB J13674.
Hellenites radiatus Renz and Renz
Text-figure 43
Hellenite.s (Pallasitc.s) radiatus Renz and Renz,
1947: 60, 75: Renz and Renz, 1948: 47, pi. 2,
fijis. 12-121), 13-131).
Hellenites (Pallasites) striates Renz and Renz,
1947: 60, 75; R(>nz and Renz, 1948: 47, pi. 2,
fifrs. 11-1 la.
Hellenites (Pallasites) striatus var. densicostata
Renz and Renz, 1947: 60, 75; Renz and Renz,
1948: 48, pi. 2, fi^s. 10-lOb.
Hellenites cf. praematurus Chao, 1959: 145, pi.
41, figs. 3, 4 (not 1 and 2).
Table 49. Measurements of Hellenites radi-
atus Renz and Renz from the Subcolumbites
FAUNA OF Chios.
D
w
H
u
W/D
H/D
U/D
1.
43.7
10.9
13.5
20.5
24.9
30.9
46.9
2.
37.1
10.5
12.7
15.7
28.3
34.2
42.3
3.
33.4?
8.5
11.7
8.2
25.4?
35.0?
24.6?
4.
31.2
8.5
10.1
14.1
27.2
32.4
45.2
5.
30.2
7.2
8.7
14.7
23.8
28.8
48.7
6.
29.5
8.5
9.8
12.6
28.8
33.2
42.7
7.
28.7
8.2
10.0
12.5
28.6
34.8
43.6
8.
27.8?
7.0
8.3
13.2
25.2?
29.9?
47.5?
9.
26.7
6.1
7.8
12.7
22.8
29.2
47.6
10.
22.8
5.4
7.4
10.3
23.7
32.5
45.2
11.
22.3
6.8
6.4
10.1
30.5
28.7
45.3
12.
17.8
5.1?
5.5
8.5
28.7?
30.9
47.8
1, 2. Unfigured paratypes, Maradovuno, Chios, NHMB
J13678.
3. HoIot>'pe, H. { Pallasites) striatus var. densicostata
Renz and Renz (1947: 60, 75; 1948: pi. 2, fig. 10),
NHMB J13681.
4. Paratype, Renz and Renz (1947: 60, 75; 1948, pi.
2, fig. 13), NHMB J13677.
5. 6, 11, 12. Unfigured parat>pes, H. (Pallasites)
striatus, Maradovuno, Chios, NHMB J 13680.
7. Holotvpe, Renz and Renz (1947: 60, 75; 1948: pi.
2, fig. 12), NHMB J13675.
8. Holotvi^e, H. (Pallasites) striatus Renz and Renz
(1947: 60, 75; 1948: pi. 2, fig. 11), NHMB J13679.
9. 10. Unfigured paratypes, H. (Pallasites) striatus var.
densicostata, Maradovuno, Chios, NHMB J13682.
This species is exactly like //. praema-
turus except that the ribs are radial instead
of rursiradiate. Renz and Renz ( 1948 )
distinguished within this group two species
under a separate subgenus of Hellenites.
The two species were separated on the
basis of rib prominence, but again the col-
lections have a number of transitional forms
in this character. The suture (Fig. 43H)
is essentially the same as in //. praema-
turus.
The only distinguishing feature between
//. praematurus and //. radiatus is the
character of the lateral ribs. This is pre-
sumably a valid species criterion but not
one to warrant erection of a separate sub-
genus.
This species is not as abundant in the
Chios fauna as H. praematurus. As in the
latter species, //. radiatus does not exhibit
any marked variability in the basic conch
dimensions. On Table 49 are measurements
of 12 specimens including the primary
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 515
30
25
20
15
10
5
0
15
20
30 35
DIAMETER
Figure 44. Variation in whorl width (W) and umbilical diameter (U) of Hellenltes praemoturus (Arthaber), from the Sub-
co/umbites fauna of Albania and Chios. The data on this graph are from Table 48.
types. The measurements are plotted on
Figure 44.
One of the specimens from a SuhcoJum-
bites horizon in Kwangsi, China, that Chao
(1959: pi. 41, figs. 3, 4) assigned to H. cf.
pracmaiurus has radial ribs and should be
assigned to H. radiatus. The specimen is
poorly preserved and lacks a suture, but
the pattern of ornamentation, etc., is so dis-
tinctive that this conclusion seems justified.
The Tobin Formation of Nevada has
yielded one specimen, 18 mm in diameter,
that appears to be identical to the other
representatives of this species. No suture
is preserved, but the degree of involution,
ornamentation, etc., are identical.
The species of Hellenites from Chios and
Kwangsi are from Subcoliimhites faunas.
There are two older species which need to
be attached to this genus. These are H.
idahoense (Smith, 1932: 81, pi. 49, figs.
13-19 ) from the Coliimbitcs fauna of south-
eastern Idaho, and H. inopinatus Kipari-
sova ( 1958a ) from the Primorye Region at
a horizon equivalent to the Cohimbites
Zone of Idaho.
516 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Occurrence. The holotype and paratype
from the SuhcoJumhitcs fauna of Chios;
from SuhcoJumhitcs fauna, Nahhng, one
kilometer northeast of Lolou, Lingolo dis-
trict, Kwangsi, China; upper part of Tol^in
Formation, south end of Tobin Range, Ne-
vada.
Repositon/. Holotvpe, Renz and Renz
(1948: pi. 2, figs. 12ll2a) NHMB J13675;
paratvpes, Renz and Renz (1948: pi. 2, fig.
12b)'NHMB J13676, (pi. 2, figs. 13-13b)
NHMB J13677; unfigured paratypes from
Maradovuno NHMB J13678; holotype, H.
(Palkisitcs) striatus Renz and Renz (1948:
pi. 2, figs. 11-lla) NHMB J13679; unfig-
ured paratypes NHMB J 13680; type speci-
mens //. (Pallasitcs) striatus var. dcnsicos-
tata Renz and Renz (1948: pi. 2, figs. 10-
10b) NHMB J13681; unfigured paratypes
NHMB J13682; specimen from Tobin For-
mation MCZ 9654.
Hellenifes idahoense (Smith)
Plate 53, figures 13, 14; Text-figure 43
Pseudliarpuccias idahoense Smith, 1932: 81, pi.
49, figs. 17-19.
Pseudarniotites idatioense, — Spath, 1951: 9.
Smith based his species on a small frag-
mentary specimen 19 mm in diameter, and
extensive collecting from the same horizon
in southeastern Idaho has not yielded any
additional specimens. This species is very
similar in its basic morphological features
to H. radiatus. The differences are mainly
seen in the character of the ribs on the
venter, the character of the keel, and the
suture (Fig. 43D).
Most other occurrences of Hellenifes are
from Suhcolumhites faunas of latest Scy-
thian age. This species is from the Colum-
hites Zone,
Occurrence. Middle shale member,
Thaynes Formation, CoJunihitcs Zone, Paris
Canyon, southeastern Idaho.
Repository. Holotype, USNM 74994.
Heilenifes inopinafus Kiparisova
Hellenifes (?) ino))inaius Kiparisova, 1958a: 13,
fig. 9; Kiparisova, 1961: 169, pi. 33, fig. 4.
Kiparisova had six specimens upon which
to base her species, but unfortunately none
of these yielded a suture, nor was the pres-
ervation of the specimens particularly
good. On the basis of the illustrations and
figures of the holotype, this species is quite
similar to H. radiatus Renz and Renz from
the Suhcolumhites fauna of Chios. Kipari-
sova ( 1961 ) did not give precise strati-
graphic data on her species, but Zakharov
(personal communication ) tells me that the
species is in his NeocoUimhites insignis Sub-
zone which is correlative with the Colum-
bites fauna of southeast Idaho. Kiparisova's
species is not very similar to Hellenifes
idahoense (Smith) from the Columhites
faima of southeast Idaho.
Occurrence. Neocolumbites insignis Sub-
zone, Primorye Region.
Family BEYRICHITIDAE Spath, 1934
Genus 6eyr/ch/fes Waagen, 1895
Type species. Ammonites reuftensis Beyrich,
1867
Beyrichifes laurae Renz and Renz
Text-figure 45
Bei/richites laurae Renz and Renz, 1948: 62, pi. 8,
fig. 1.
Bei/riehite.s prdenuifinus Renz and Renz, 1948: 61,
pi. 7, fig. 5.
The authors of this species were fully
cognizant that the presence of this typical
Anisian genus in their Chios fauna raised
serious questions. They briefly discussed
the possibility of mixing and reworking
versus the true existence of this genus in
the late Scythian. On the basis of the data
available, I have elected to accept this
species as a member of the late Scythian
Chios fauna. Each of the two species of
this genus recognized by Renz and Renz
was bas(xl on a single specimen; I can see
no signilicant difference between these two
specimens.
Occurrence. Suhcolund)ites launa of
Chios.
Repository. Holotype of Reyrichitcs
laurae, NHMB 113712, liolotvpe of B. prae-
77uiturus, NHMB J 13711.
Ammonoids of the Late Scythian (Lower Triassic) • Ktimmel 517
Figure 45. Diagrammatic representation of the suture of:
A, Eogymnites arihaberi (Diener), from fiolotype, at a diam-
eter of 75 mm, from Subcolumbites fauna of Albania; B,
Beyrichifes laurae Renz and Renz (1948; pi. 8, fig. lb), at
a diameter of 42 mm, from Subco/umb/fes fauna of Cfiios
(NHMB J13712).
Superfamily PINACOCERATACEAE Moj-
sisovics, 1879
Family GYMNITIDAE Waagen, 1895
Genus Eogymnifes Spath, 1951
Type species, Japonites arthaberi Diener,
1915
Eogymnifes arfhaberi (Diener)
Plate 21, figures 1, 2; Text-figure 45
Japonites sugriva Diener var. Arthaber, 1911: 231,
pi. 20(4), fig 4.
Japonites arthaberi Diener, 1915: 158 ( = /. sug-
riva,— Arthaber non Diener); Kunimel, in Arkell
et al., 1957: L185, fig. 214,4.
Japonites decipiens Spath, 1951: 172 (= /. sug-
riva,— Arthaber non Diener ) .
Arthaber's type and only specimen of
this unusual Scythian species is only mod-
estly well preserved, consisting mainly of
phragmocone; the adoral 30 mm is body
chamber. The specimen measures approxi-
mately 89 mm in diameter, 23.4 mm for the
width of the adoral whorl, 29 mm for the
height, and 36.4 mm for the diameter of
the umbilicus. The whorls are compressed
with convex whorl sides converging to a
narrowly rounded venter. The maximum
width is at the umbilical shoulder. The
umbilical wall is low, rounded, and slopes
to the umbilical seam at a steep angle. The
shell is preserved only in part, and very
poorly; it is apparently smooth except for
slightly sinuous growth lines.
Spath (1951: 172) expressed concern
over Arthaber's (1911: pi. 20(4), fig. 4c)
illustration of the suture. A new drawing
made from the type specimen is illustrated
here on Figure 45A. As with nearly all the
specimens from the Subcoliimbites fauna
from Albania and Chios, the suture can be
developed only by grinding. The suture
illustrated here I believe to be a fairly ac-
curate representation and does not differ
from that reproduced by Arthaber to any
significant degree.
There is no other Scythian ammonoid
comparable to this species.
Occurrence. Subcolumbites fauna, Kcira,
Albania.
Repository. Paleontological Institute,
University of Vienna.
Family HUNGARITIDAE Waagen, 1895
Genus Prohungarifes Spath, 1934
Type species, Prohungarifes similis Spath
(= Hungarifes cf. middlemissii Diener,—
Welter, 1922)
Prohungarifes crasseplicafus (Welter)
Text-figure 46
Hungarites cra.sseplicatus Welter, 1922: 147, pi.
168(14), figs. 1-6.
Prohungarites crasseplicatus, — Spath, 1934: 244;
Spath, 1951: 20; Kunimel, 1961: 525.
Hungarites cf. middlemissii Diener, — Welter, 1922:
146, pi. 13, figs. 6-9, 18.
Prohungarites similis Spath, 1934: 327; Spath,
1951: 19; Kunimel, /;i Arkell et al., 1957: L155,
fig. 186,7.
It was for this moiphologic group from
Timor that Spath (1934) introduced the
genus Prohungarites. Welter (1922) had
included in the genus Hungarites three
518 Bulletin Museum of Comparative Zoologij, Vol. 137, No. 3
species from the blocks with manganese
coated fossils from Nifoekoko, Timor; these
were cf. middlcmissii Diener, cmsseplicatus
and tiibcrculattis. Spath (1934: 327) in-
troduced the species name .similis for cf.
middlemissii, — Welter (non Diener). The
first two of these species are combined
here. They were originally separated
merely on differences in the intensity of
the ribbing. Even Welter (1922: 147)
discussed the gradational aspects of the
species and illustrated one specimen (Wel-
ter, 1922: pi. 167(13), figs. 10, 11) as a
transitional form between P. middlcmissii
and P. crasscplicotus. Restudy of Welter's
types show that his descriptions and illus-
trations are quite adequate. The measure-
ments of Welter's types are as follows:
D
W
H
U
W/D
H/D
U/D
1.
58.5
17.5
23.4
15.1
29.9
40.0
25.8
2.
38.7
14.2
16.5
10.6
36.7
42.6
27.4
3.
36.1
11.5
15.0
10.5
31.9
41.6
29.1
4.
35.4
11.4
16.0?
8.2?
32.2
45.2?
23.2?
5.
33.6
11.9
13.2
11.2
35.4
39.3
33.3
6.
30.0
10.1
11.9
9.5
33.7
39.7
31.7
1. Welter (1922: pi. 13, fifis. 6-9) GPIBo 226a.
2. Welter (1922: pi. 14, fi^s. 1-3) GPIBo 228a.
3. Welter (1922: pi. 13, fig.s. 10-11) GPIBo 227.
4. Welter (1922: pi. 13, ii^. 18) GPIBo 226b.
5. Welter (1922: pi. 14, fijis. 4, 5) GPIBo 228b.
6. Welter (1922: pi. 14, fi.g. 6) GPIBo 228c.
The upper part of the Thaynes Fomia-
tion along Hammond Creek in southeastern
Idaho has yielded about 30 poorly pre-
served specimens which are very similar
to the Timor P. crasseplicattis. The whorls
of the Idaho species are fastigate only on
the early volutions. The mature volutions
have rounded venters. The slightly sinuous
ribs on most of the specimens are compar-
able to those on Welter's transitional speci-
men between P. similis and P. crasseplicatus
but are prosiradiate, not radial. Tlie largest
specimen, approximately 65 mm in diam-
eter, consisting of a half volution of body
chamber, has the blunt lateral ribs like
Welter's P. cmsseplicatus (Welter, 1922:
pi. 14, figs. 1-3). The sutures in the Timor
and Idaho specimens are likewise similar
(Fig. 46).
Occurrence. The specimens from Timor
came from the limestone with manganese
coated fossils, Nifoekoko.
Repositonj. Holotype P. similis Spath ( =
Ilungorites cf. 7uiddlcmissii, — Welter, 1922:
pi. 13, figs. 6-9) GPIBo 226a; paratype,
Hun^arites cf. middlcmissii, — Welter (1922:
pi. 13, fig. 18) GPIBo 226b; syntype P.
crasseplicatus (Welter, 1922: pi. 14, figs.
1-3) GPIBo 228a, (pi. 14, figs. 4, 5) GPIBo
228b, ( pi. 14, fig. 6 ) GPIBo 228c; Welter's
transitional specimen (1922: pi. 13, figs.
10,11) GPIBo 227.
Prohungarifes cf. crasseplicatus (Welter)
Prolitiugarites cf. crasseplicatus, — Kumniel, 1966:
400, pi. 3, figs. 11, 12.
Four small phragmocones from West
Pakistan differ from the Timor P. cra.s-
seplicatus principally in the absence of any
indication of a keel along the central part
of the venter on the mature whorls. In all
other conch features, such as shape of the
whorl section, shape of the ventral and
umbilical shoulders, and nature of the um-
bilical wall, it is very similar. The suture
is likewise essentially the same. The other
Timor species, Prohung,arites tubcrculatus,
is more robust with a more highly de-
veloped ornamental pattern. There is a
strong moiphological similarity to Prohiin-
i!,arites gutstadti n. sp. from the Upper
Thaynes Formation of southeast Idaho.
Both species have acute venters only on
the earliest volutions and rounded venters
on the later volutions.
Occurrence. Narmia Member of the
Mianwali Formation, Narmia Nala, Surghar
Range, West Pakistan.
Repository. MCZ 9606, 9607.
Prohungarifes fuberculatus (Welter)
Text-figure 46
Ilungariles (ul)crciil(ilus Welter, 1922: 148, pi.
167( 13), figs. 12-17.
Prohtiufiarites tubcrculatus, — Spath, 19.34: 244;
Kiimiuel, 1961: 525.
The basic form of the conch in this
species is ver\ much like P. crasseplicatus
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 519
/N
A
Figure 46. Diagrammatic representation of the suture of: A, Hungor/fes cf. middlemissii Diener, — Welter (1922: pi. 13,
fig. 18), from Block E, Nifoekoko, Timor; B, Hungarites crasseplicatus Welter (1922: pi. 14, fig. 6), from Block E,
Nifoekoko, Timor; C, Hungarites tuberculaius Welter (1922: pi. 13, fig. 12), from Block E, Nifoekoko, Timor; D, Kipari-
sovitei carinatus Astakhova (1964: 379, fig. Id), from Dor/cronites Zone of Astakhova, Mangysfilak Peninsula; E, Prohun-
garifes (?) popovi Kiparisova (1961: fig. 113), from Anisian strata of the Primorye Region, at a whorl height of 13 mm;
F, Prohungarites mckelvei n. sp. from Upper Thaynes Formatioi, Hammond Creek, southeast Idaho, at a whorl height of
17 mm (MCZ 9646); G, Prohungar/fes gutstadti n. sp., from Upper Thaynes Formation, Hammond Creek, southeast Idaho,
at a whorl height of 13 mm (holotype MCZ 9475); H, Prohungori/es sp. indet., from Upper Thaynes Formation, Hammond
Creek, southeast Idaho, at a whorl height of 15 mm (MCZ 9648); I, Prohungarites sp. indet., from Upper Thaynes Formation,
Hammond Creek, southeast Idaho, at a whorl height of 26 mm; J, Dolmatites morlaccus KittI (1903: pi. 4, fig. 6), from
Werfen Formation, Dalmatia; K, Dolmatites kittii n. sp., from Co/umb;tes Zone, Paris Canyon, southeast Idaho, at a whorl
height of 10 mm (MCZ 9499).
520 Bulletin Museum of Comporative Zoology, Vol. 137, No. 3
except that it is more e volute and the
whorls more inflated. The big difference
in these species is in the ornamentation.
This species has coarse, conspicuous ribs
with a node just above the umbilical
shoulder. This pattern of ornamentation
commences at an early growth stage, as
can be seen in the umbilical region of
Welter's holotype and in the small speci-
men illustrated by Welter (1922: pi. 13,
figs. 16, 17). The measurements of Welter's
two specimens are as follows:
D
47.5
7.3
W
H
U W/D H/D U/D
16.6 18.0 17.5 34.9 37.9 36.8
? 2.5 2.8 ? 34.2 38.4
The first of these measurements is for the
holotype and the second is for the paratype.
The suture is illustrated on Figure 46C.
Occurrence. In the limestone with black
manganese coated fossils, Nifoekoko, Block
E, Timor.
Repositon/. Holotype GPIBo 229a; para-
type GPIBo'229b.
Prohungarites middlemissii (Diener)
Plate 25, figures 3-8
Hini^uritc.s middlemissii Diener, 191.3: 23, pi. 3,
figs. 5-7; Diener, 1915: 1.53; Spath, 1934: 33.
Prohungarites middlemissii, — Kimimel, 1961: 525.
Diener (1913) stated he had approxi-
mately 40 specimens of this species col-
lected from a loose block at Pastannah,
Kashmir. Of these only the three figured
syntypes (Diener, 1913: pi. 3, figs. 5-7) are
preserved in the collection of the Geologi-
cal Survey of India. The disposition of the
other specimens is not known.
Even though the suture is unknown, the
general conch morphology clearly indicates
that this is a valid species of Pro]mng,arites.
In the degree of involutions and ornamen-
tations, it is quite similar to P. juckehei but
is unique in the widening of the ventral
part of the whorl in the mature stages and
in the acquisition of nodes on tlu' ribs.
Prohimfi^arites crassepJicatus has a much
more robust whorl section, is more evolute,
and has more pronounced ornamentation. "^
This species strongly suggests the pres-
ence of a horizon younger than the Heden-
strocmia beds in Kashmir. Final evaluation
on the biologic affinities of this species and
its stratigraphic position will have to await
new field investigations and collections.
Occurrence. Loose block, Pastannah,
Kashmir.
Repositon/. Svntype (Diener, 1913: pi.
3, fig. 5) GSI il276, (pi. 3, fig. 6) GSI
11277, (pi. 3, fig. 7), GSI 11278.
Prohungarites carinatus (Astakhova)
Text-figure 46
Ki))arisovites carinatus Astakhova, 1964: .379, pi.
l,fig. 1.
This is clearly a species of Prohungarites
of the general form of P. mckelvei and P.
crasseplicatus; just why its author intro-
duced a new genus for her two fragmentary
specimens is hard to tell. The suture is
shown on Figure 46D.
Occurrence. Doricranites Zone (of Astak-
hova, 1960a, b), Mangyshlak Peninsula,
Karatauchik Range.
Prohungarites mckelvei n. sp.
Plate 35, figures 1-5, 8, 9; Text-
figure 46
Prohungarites n. sp. cf. P. similis, — Kunimel, 1954:
187.
This is the most common species in the
upper member of the Thaynes Formation
at Hammond Creek, Bear River Range,
southeast Idaho. The collection contains a
couple of hundred specimens, most of
which, however, are not well preserved.
In addition, the Tobin Formation, Tobin
Range, Nevada, has yielded 10 fragmentary
and poorly preserved specimens.
The conch is involute, compressed, with
a distinct knile-edged \'enter. A remarkable
feature of this species is the high degree
of constancy in basic conch proportions.
The venter has a distinct fastigate appear-
ance except that the ventral shoulders are
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 521
rounded and not angular. The lateral areas
are broadly convex. The umbilical shoulder
is acutely rounded and the umbilical wall
is vertical.
The conch is essentially smooth, except
for extremely faint, moderately spaced,
falcoid ribs. Within the small umbilicus
only the umbilical shoulders and wall of
the preceding volutions are visible. No
ornamentation of any kind is visible in the
umbilical area.
The suture is shown on Figure 46F.
This new Idaho species displays some
morphological similarity to Frohung^arites
crasscplicatus from the manganese coated
beds of Nifoekoko, Timor, but a more
marked similarity to Proliuniiiuitcs mid-
dJemissii from an unknown horizon in
Kashmir. The Timor species (P. cras-
seplicafus) is more inflated, more evolute,
with sharper ventral shoulders, and with a
tendency towards more robust ornamen-
tation. In spite of these differences, the
moiphological relationships of P. mckelvei
and P. crasseplicatiis are very close. The
Kashmir species (P. middlemissii) differs
mainly in that on the outer volutions the
greatest width of the whorls shifts to the
region of the ventral shoulders. The degree
of involution, the subdued falcoid ribs, and
nature of the venter are strikingly similar
to P. mckelvei. My specimens from the
Tobin Formation, though poorly preserved
and fragmentary, cannot be separated from
the Idaho forms. N. J. Silberling of the
U.S. Geological Survey has kindly shown
the author some specimens of ProJningorites
he collected from the lower part of the
Tobin Formation, Tobin Range, Nevada
(U.S.G.S. locality 2565). These specimens
are very much like the species described
here but are slightly more inflated in whorl
section; they possibly represent a new
species.
Occurrence. Upper member of Thaynes
Formation, Hammond Creek, Bear River
Range, southeast Idaho; upper part Tobin
Formation, Tobin Range, Nevada.
Repository. Specimens from Hammond
Creek: Holotype MCZ 9466 (Pi. 35, figs.
1, 2); figured paratypes MCZ 9467 (PI. 35,
fig. 3, 4), MCZ 9468 (PI. 35, fig. 5), MCZ
9469 (PI. 35, figs. 8, 9); unfigured para-
types MCZ 9646; unfigured specimens from
Tobin Formation, MCZ 9651.
Prohungarites gutstadti n. sp.
Plate 36, figures 3, 14, 15; Text-
figure 46
Prohungarites cf. crasseplicatiis Kummel, 1954:
187; Kummel, 1966: 400.
This species is represented by 30 or more
specimens most of which, unfortunately,
are poorly preserved. This species has the
general conch form of P. crasseplicatiis. It
differs in that the venter on the mature
volutions is rounded and not fastigate.
A blunt sharpened venter is present up to
a diameter of approximately 15 mm, as seen
in the specimen of Plate 36, figures 14, 15,
but at some stage after that the venter
becomes rounded. The second major dif-
ference is that the ribs are prosiradiate and
not radial. The range of ribbing pattern
is much the same in the two species. The
suture is shown on Figure 46G.
This species is remarkably similar to the
specimens assigned to Prohungarites cf.
crasseplicatiis from the Narmia Member
of the Mianwali Formation in the Surghar
Range of West Pakistan (Kummel, 1966).
In that species the sharpened venter like-
wise is confined to the earlier volutions, the
more mature volutions having a rounded
venter. The ribs in the Pakistan species
are radial rather than prosiradiate. The
restriction of the sharpened venter to the
earliest volutions differentiates these two
species from all others assigned to Pro-
hungarites.
Occurrence. Upper Tliaynes Formation,
Hammond Creek, Bear River Range, south-
east Idaho.
Repository. Holotype MCZ 9475 (PI. 36,
fig. 3); figured paratype MCZ 9481 (PI.
36, figs. 14, 15); unfigured paratypes MCZ
9645.
522 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Prohungarites sp. indet.
Plate 38, figures 4, 5; Text-figure 46
Tlie collections from the Hammond
Creek locality of the Thaynes Formation,
and from the upper part of the Tobin For-
mation in the Tobin Range, Nevada, con-
tain, in association with Frohuniia rites
mckelvei, several poorly preserved frag-
ments that are clearly a new and distinctive
species of ammonite. However, because
of the fragmentary nature of the specimens
no new name is introduced. The whorls
are very rapidly expanding with radial to
slightly prosiradiate ribs that are greatly
enlarged in the general region of the um-
bilical shoulder. The venter is fairly broad
and fastigate. The suture is shown on
Figure 46H, I.
This form is completely different from
the associated Prolnin^a rites mckclvei and
Frohunii^aritcs iiutstadti and is quite differ-
ent from the other species of Frohun^aritcs
reported to date. The pattern of ribbing
and the rapidly expanding whorls at first
suggested a relationship to Arctoprionites.
In that genus, however, in so far as we
know, the venter is truncate and never
fastigate. The suture of the form recorded
here is more prohungaritid in aspect.
Occurrence. Upper Thaynes Formation,
Hammond Creek, Bear River Range, south-
east Idaho; upper Tobin Formation, south
end Tobin Range, Nevada.
Repositorij. Figured specimens from
southeast Idaho MCZ 9474 (PL 36, figs.
1, 2), MCZ 9647 (PI. 38, figs. 4, 5); un-
figured specimens from southeast Idaho
MCZ 9648; specimens from Tobin Range
MCZ 9652.
Genus Dalmatites KittI, 1903
Type species, Dalmafiies morlaccus KittI,
1903
Dolmafifes morlaccus KittI
Plate 56, figures 1-8; Text-figure 46
Dalmatites morlaccus KittI, 1903: 73, pi. 4, tij^s.
3-7; Diener, 1915: 115; Spath, 1951: 20;
Kuinmel, in Arkell, et al., 1957: LI 56, f'i<,'.
187, 7.
1. 69.0 15.6 39.0
2. 53.0 10.0? 27.1
3. 49.1 8.4 27.1
4. 43.0 9.3 23.0?
5.1 22.6 56.5 7.4
2.7 18.9? 51.1 5.1
3.0 17.1 55.2 6.1
? 21.6 53.5? ?
1. Paialectotype, KittI (1903: pi. 4, fig. 7).
2. Paialectotype, KittI (1903: pi. 4, fig. 6).
3. Lectotype, KittI (1903: pi. 4, fig. 4).
4. Paialectotype, KittI (1903: pi. 4, fig. 5).
The suture is reproduced here in Figure
46J.
This species is known only from the
Werfen Formation of Europe. The only
other species of this genus, D. kittli, is
known only from a single specimen from
the Coliimbifcs fauna of southeastern
Idaho. The two species are very similar.
Occurrence. Werfen Fonnation, Muc,
Dalmatia.
Repository. Natural History Museum,
Vienna.
Dalmatites kittli n. sp.
Plate 55, figures 7, 8; Text-figure 46
This species is established on a single
specimen from the Colum1)itcs fauna of
southeastern Idaho. The conch is smooth,
compressed, involute, and is entirely
phragmocone. It measures 33.5 mm for the
width of the last whorl, and the umbilicus
is 3.8 mm in diameter. The venter is acute,
the lateral areas broadly convex. The
broadest part of the whorl is in the mid-area
of the whorl. The umbilical shoulder is
rounded with a fairly steep umbilical wall.
The conch is smooth except for growth
lines. The pattern of the growth lines,
however, is not \isible due to fault\' pres-
ervation of the shell.
The suture is illustrated on Figure 46K.
It is of a fairly simple pattern with two
lateral lobes, and an auxiliarv lobe on the
The four specimens of this species illus-
trated by KittI (1903: pi. 4, figs. 4-7) are ''^
still preserved. The photographs of these
type specimens reproduced here on Plate
56 show that the preservation is no more
than fair to poor. The measurements of
these four specimens are as follows:
D W H U W/D H/D U/D
Ammonoids of the Late Scythian (Lower Triassic) • Ktimmel 523
Figure 47. Diagrammatic representation of the sutures of three species of Eophyllitei. A-F, Eophyllites dieneri (Arthober);
A, paralectotype (Arthober, 1908; fig. 4), new drawing at a diameter of 42 mm; B, plesiotype (Arthaber, 1911: pi. 20(4),
fig. 6), new drawing at a diameter of 47 mm; C, type specimen of A4onophy//ites kingi, — Arthaber (non Diener) z= Ussurltes
(?) dec/'piens Spath (1934), at a diameter of approximately 30 mm; this suture is highly distorted due to excessive grinding of
the specimen; D, holotype of A4onophy///fes nopcsoi Arthaber (1908: pi. 12, fig. 5), nev/ drawing at a diameter of 28 mm;
E, holotype of Monop/iy//ifes (Schizophy/Zites) befi7/on/ Renz and Renz (1948: pi. 4, fig. 8b), at a diameter of 47 mm; F,
plesiotype (Renz and Renz, 1948: pi. 5, fig. lb), at an unknown diameter. G, Eophyllites orientalis Spath, — Welter (1922:
pi. 161(7), fig. 7), at an unknown diameter. H, Eophyllites amurensis Kiparisova (1961: text-fig. 104), at o whorl height
of 10 mm.
Specimens of A-D from Subco/umb/tes fauna of Albania; E, F, from same fauna on Chios; G, from Nifoekoko, Block E,
Timor; H, from Subco/umb/tes fauna, Primorye Region, Siberia.
dorsal areas of the flanks. It is difficult to
be sure whether the lobes are really
goniatitic as shown on Figure 46K. There
are faint indications of denticulations on
some of the first and second lateral lobes,
and I suspect the smooth aspect is due to
poor preservation.
Dalmatites morlaccus Kittl is morphologi-
cally very similar to D. kittli. The venter
on the type species becomes rounded on
the body chamber and the suture shows
minor differences in proportion and shape
of the elements.
Occurrence. Middle shale member of
524 Biilleiin Museum of Comparative Zoology, Vol. 137, No. 3
Thaynes Formation (Cohimbites fauna),
Paris Canyon, Bear River Range, South-
east Idaho.
Repository. Holotype, MCZ 9499.
Dalmatites attenuatus Smith
Plate 71, figures 8, 9;
Dalmatites attenuatus Smith, 1932: 81, pi. 57,
figs. n-13.
A highly compressed dalmatitid with a
sharp venter. It is more compressed than
either D. morlaccus or D. kittli and in
addition more e volute. The suture is on the
same basic pattern as the other species.
Occurrence. Smith's holotype came from
the Tirolites Zone in Paris Canyon, south-
east Idaho. He reports (Smith, 1932: 81)
that he had only one specimen from the
Tirolites Zone of Idaho but in addition he
states the species was also found in the
Meckoccras Zone at Phelan ranch, mouth
of Cottonwood Canyon, east of the Ruby
Range, Nevada; however, this latter speci-
men (or specimens?) is apparently no
longer preserved. Smith (1932: 81) did
describe a species — D. richardsi — from the
Meckoccras Zone.
Repository. Holotype USNM 75023.
Order PHYLLOCERIDA Arkell, 1950
Superfamily PHYLLOCERATACEAE Zittel,
1884
Family USSURITIDAE Hyatt, 1900
Genus Eophyllifes Spath, 1930
Type species, Monophyllites dieneri
Arthaber, 1908
Eophyllites dieneri (Arthaber)
Plate 22, figures 1-4; Plate 23, figures
1-7; Text-figure 47
Monu])hyllitcs dieneri Arthaber, 1908: 288, pi.
13(3), figs. 3a-c, 4a-c; Arthaber, 1911: 234,
pi. 20(4), figs. 5-8; Diciier, 1915: 203.
Eophyllite.s f/foicrj,— Spath, 1930: 89; Spatli,
1934: 293-295; Kvimmel, in Arkell et al., 1957:
LI 86.
Monop]}t)Uilcs Juira, — Arthaber (nun Dieiier),
1908:" 286, pi. 12(2), figs. 4a-c; Arthaber,
1911: 235.
Monophyllites (Ussurites) /lara Diener, 1915: 206.
Eophyllites refractus Spath, 1934: 295, pi. 3, fig. 4.
Monophyllites kingi, — Arthaber (non Diener),
1911: 235, pi. 20(4), figs. 12a-c.
Monophyllites {Usstirites) kingi Diener, 1915:
207.
Ussurites (?) decipiens Spath, 1934: 302 ( =
Monophyllites kingi, — Arthaber non Diener).
Monophi/lUtes nopcsai Arthaber, 1908: 287, pi.
12(2)", figs. 5a-c; Arthaber, 1911: 235; Diener,
1915: 203.
Eophyllites nopcsai, — Spath, 1934: 302.
Monophyllites (Leiophyllites) rosae Renz and
RcMiz,"l947: 61, 77; Renz and Renz, 1948: 74,
pi. 3, figs. 8-8a.
Mono})hyllitcs (Schizoj^luillitcs) hetilloni Renz and
Renz, 1947: 61, 78; Renz and Renz, 1948: 76,
pi. 4, figs. 8-8b.
Monophyllites (Schizoplujllites) hetilloni var.
cioluta Renz and Renz, 1948: 76, pi. 4, figs.
6-6a, pi. 5, figs. 2-2a, 4-4a, 6-6a.
The lectotype of this species is not in
the collections of the Paleontological In-
stitute, University of Vienna, and is pre-
sumed lost. Two figured paralectotypes are
available, but none of the unfigured speci-
mens noted by Arthaber. The one figured
paralectotype (Arthaber, 1908: pi. 13(3),
figs. 4a-c; 1911: pi. 20(4), figs. 5a-c; PI.
23, figs. 2, 3 of this report) is mainly
phragmocone. The lateral area of the il-
lustrated side has been polished to expose
the sutures, but at the same time this has
destroyed all surface markings. The op-
posite side of the conch is all matrix or
highly weathered. This specimen measures
49.5 mm in diameter, 20.6 mm for the
height of the last whorl and 15.7 mm for
the diameter of the umbilicus. The un-
retouched photograph of Plate 23, figures
2, 3 shows the main features of the conch
and the general state of preservation of
the specimen better than the retouched il-
lustration of Arthaber.
The second paralectotype is the speci-
men which \ ieklcd the suture of Arthaber's
plate 20(4), figure 6 and is illustrated here
on Plate 23, figures 4, 5. It is a fragmentary
specimen that is nearly all phragmocone.
Much of the lateral area has been ground
to expose the suture (Fig. 47B).
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 525
MonopJu/Uifes hara, — Arthaber, non Die-
ner (1908: 286, pi. 12(2), fig. 4), re-
named EophijUites refractus by Spath
(1934: 295) is unfortunately not preserved
in the collections of the Paleontological In-
stitute, University of Vienna. There are,
however, several topotypes in the British
Museum (Natural History). The species
was characterized mainly by its broader
venter. Spath (1934: 295) also mentions
differences with EopJiyUitcs dieneri in the
pattern of the growth lines. In regard to
the width of the venter, Spath himself
stated that among the topotypes he had,
they could not be distinguished satisfac-
torily from E. dieneri. As yet too few speci-
mens of EophylJitcs from Albania have
been studied, but it does not seem likely
that differences in the character of the
venter as used in this case are anything
more than the normal type of variation one
can expect and can demonstrate in many
cases within ammonite species. EophijUites
refractus is believed to be conspecific with
E. dieneri.
Ussurites (?) decipiens Spath (1934)
{= MonophyUitcs kingi, — Arthaber, 1911:
pi. 20(4), fig. 12) is likewise beheved to
be a specimen of E. dieneri. Proper under-
standing of this species (and specimens)
has not been possible on the basis of
Arthaber's illustrations, especially of the
suture. In the first place the type speci-
men (Plate 23, figures 6, 7) has the same
general shape, etc., as the types of E.
dieneri. The measurements are: diameter
50.8 mm, width of last whorl 12.0? mm,
height of last whorl 18.8 mm, and width
of umbilicus 18.3 mm. One of the most
conspicuous differences is that decipiens
is more evolute than dieneri; this differ-
ence, however, amounts to only 5 percent
of the conch diameter and this is hardly a
criterion of specific significance. The
whorls are slightly more inflated than in
decipiens and the venter is more broadly
rounded than indicated by Arthaber's
(1911: pi. 20(4), fig. 12b) drawing. The
whorl cross-section is intermediate between
that of E. decipiens and E. refractus. The
adoral quarter volution is marked by fine
radial lines some of which at intervals are
more conspicuous than others. The general
absence of these lines on the two available
specimens of decipiens is due to preserva-
tion plus grinding of the surface of the
conch.
It is in the suture, however, that most
authors recognized significant differences.
As with practically all of the Albanian ma-
terial, the suture can be made visible only
by grinding and polishing of the surface.
Arthaber's type specimen had been ground
for this purpose and in this case the grind-
ing was far too much, destroying many of
the details of the lobe denticulations. Faint
outlines of denticulations are visible on the
lower flanks of the lobes. Arthaber's suture
(1911: pi. 20(4), fig. 12c) actually ends
on the umbilical shoulder and does not
include anything for the umbilical wall.
The suture on the umbilical wall is, how-
ever, not preserved. The relative shortness
of the suture line reflects the difference
in degree of involution and whorl height.
The suture of decipiens illustrated by Ar-
thaber came from a whorl height of 10.5
mm; the suture of dieneri (Arthaber, 1911:
pi. 20(4), fig. 5) came from a whorl height
of 17 mm. All of the above data clearly
point to Ussurites decipiens as being part
of the dieneri complex.
EophijUites nopcsai (Arthaber), which is
associated with E. dieneri in the Albanian
fauna, is based on a single specimen; this
is illustrated here on Plate 22, figures 1, 2.
In suture ( Fig. 47D ) and ornamentation it
is very much like E. dieneri, but its conch
is very involute; its umbilicus is only 25
percent the diameter of the conch whereas
in dieneri the umbilicus measures 30-36
percent the diameter of the conch. The
association of this form in the same beds
with E. dieneri and the very close simi-
larity in all other conch features leads me
to conclude that nopcsai represents nothing
more than a variant toward involution of
the conch, as decipiens represents a variant
526 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
in the opposite direction — that is, a more
e volute conch.
The Subcolumbitcs fauna of Chios con-
tains an interesting assemblage of speci-
mens assigned to other species which I
believe are conspecific with the Albanian
E. dieneri. The specimens which Renz and
Renz (1948: 74) named MonophyUites
(Leiopliyllites) rosae I cannot separate
from E. dieneri. The specimens assigned
by Renz and Renz (1948: 76) to Mono-
pJiyllites {SchizopliyUitcs) bctilloni were
set aside into a new subgenus on the basis
of a slight saddle in the ventral lobe. The
suture of the subgenotype ( Fig. 47E ) is
nearly identical to that of the suture of
the paralectotype of E. dieneri (Fig. 47A).
Spath (1934: 294) had earlier called at-
tention to the variability in the suture of
E. die7}eri and especially to the ventral
lobe. He also rightly cautioned that some
of this variability is caused by preparation
of the suture with acids and grinding. In
all other features betilloni resembles E.
dieneri, and there appears little doubt but
that these two species are conspecific.
The form from Timor described by
Welter (1922: 118, pi. 161(7), figs. 5-7)
as MonophyUites nov. sp. ex aff. dieneri was
renamed EophyUites oriental is by Spath
(1934: 295). This species is based on a
single incomplete specimen that represents
the inner whorls of what was a much
larger form. It is unquestionably very close
to E. dieneri and possibly even conspecific,
but here again, since it is based on a single
specimen, it is considered best for the
moment to recognize it as a distinct species.
Occurrence. Subco]um])ites fauna of
Kcira, Albania, and Chios.
Repository. The Paleontological Insti-
tute, University of Vienna, contains two
paralectotypes of E. dieneri; the lectotype
is apparently lost. This collection also con-
tains the holotype of Ussnrites (?) decipicns
Spath ( = Monopliyllites kin^i, — Arthaber
non Diencn). A number of topotypes are
in the British Museum (Natural History),
C:22939-47, C22979. The Natural Tlistorv
Museum, Basel, contains the following
specimens from the Siibcohim])ites fauna
of Chios: holotype MonophyUites (Leio-
phyUites) rosae Renz and Renz (1948: pi.
3, fig. 8) NHMB J13746; holotype Mono-
phyUites (ScluzophyUites) betiUoni Renz
and Renz (1948: pi. 4, fig. 8) NHMB
J13756; var. evohita Renz and Renz ( 1948:
pi. 4, fig. 6) NHMB J13757, (pi. 5, fig. 2)
NHMB J13758, (pi. 5, fig. 4) NHMB
J13759, (pi. 5, fig. 6) NHMB J13760;
unfigured paratypes from Maradavuno,
NHMB J13761, from Kephalovuno NHMB
J13762.
EophyUites orientalis Spath
Text-figure 47
MonophyUites nov. spec, ex aff. dieneri, — Welter,
1922:" 118, pi. 161(7), figs. 5-7; Kutassy, 1933:
595.
EoplujUites orientalis Spath, 1934: 295.
This Timor species is based on a single
specimen that represents the inner whorls
of what was a much larger form. The speci-
men measures 56.7 mm in diameter, 16.0
mm for the width of the last whorl, 23.0
mm for the height of the last whorl, and
19.1 mm for the diameter of the umbilicus.
It shows a great similarity to E. dieneri
of Albania and Chios and perhaps is con-
specific with that form. However, because
only one specimen is known and because
there are minor differences in the suture
(Fig. 47C) and conch features, it is con-
sidered best to maintain the separate
identity of this species until more material
becomes available.
Occurrence. Manganese coated blocks
from Nifoekoko, Timor.
Repository. GPIBo-W215.
EophyUites amurensis Kiparisova
Text-figure 47
E()i>hi/llites anitircnsis Kipariso\a, 1961: 137, pi.
28," tigs. 7, 8, text-fig. 104.
luiphi/lliles cf. refractus, — Kiparisova, 1961: 136,
pi. 28, tig. 9, text-fig. 103.
Thc> two forms of E.ophyUites recognized
by Kiparisova from the Primorye Region
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 527
represent an inflated form (cf. refractiis)
and a more compressed form (amurensis).
The latter species was compared closely
with cUeneri from the Subcohimbites fauna
of Albania. One of these species (cf.
refractus) is based on a single specimen
and the other {amurensis) was based on
three poorly preserved specimens. They
are in the first place considered to repre-
sent a single species complex. These speci-
mens differ from E. dieneri in being slightly
more involute and in the slightly simpler
structure. They are clearly closely related
to the dieneri group of Albania and could
well be conspecific but much more material
is needed before this relationship can be
established with any degree of certainty.
Occurrence. The three specimens of
amurensis came from the Subcohimbites
fauna on the west coast of Amur Bay be-
tween Cape Atlasov and Cape Ugolny,
Ussuri Bay, Primorye Region. The single
specimen assigned by Kiparisova to cf.
refractus came from the east coast of Us-
suri Bay between Cape Kom-Pikho-Sakho
and Cape Chigan from an uncertain hori-
zon.
Genus Palaeophyllites Welter, 1922
Type species, Palaeophyllites sfeinmanni
Vv^elter, 1922
Palaeophyllites steinmanni Welter
Palaeophyllites steinmanni Welter, 1922: 119, pi.
162(8), figs. 5, 6, 7, pi. 163(9), figs. 3-6;
Kutassy, 1933: 606; Spath, 1934: 297, fig. 103.
Monophijllites (Palaeophyllites) thalmanni Renz
and Renz, 1947: 61, 78; Renz and Renz, 1948:
79, pi. 3, figs. 10-lOb ( = Palaeophyllites
steinmanni Welter, 1922: pi. 163(9), figs. .3-6
(nonpl. 162(8), figs. 5-7).
Monophyllites {Palaeophyllites} praekieperti Renz
and Renz, 1947: 61, 78; Renz and Renz, 1948:
80, pi. 4, figs. 5-5b.
Renz and Renz (1948) established the
species thahnanni for the specimen illus-
trated by Welter on his plate 163(9),
figures 3-6, at the same time designating
the specimen of Welter's plate 162(8),
figures 5, 6, as "holotype." They were un-
Table 50. Measurements of SPEcrNiENS of
Palaeophyllites steinmanni from Timor and
Chios.
D
w
H
u
W/D
H/D
U/D
1.
58.8
14.9
20.4
24.5
25.3
34.7
41.7
2.
48.2
12.1
16.4
20.5
25.1
34.0
42.5
3.
40.7
12.0
13.8
16.6
29.5
.33.9
40.8
4.
31.5
12.2
10.7
14.3
38.7
34.0
45.4
5.
26.0
8.7
8.7
10.8
.33.5
33.5
41.5
6.
23.4
7.9
10.1
7.9
33.8
43.2
33.8
1. Lectotype, Welter (1922: pl.l62(8), figs. .5-7),
GPIBo W2I6a.
2. Plesiotype, Palaeophyllites thalmanni Renz and Renz
(1948: pi. 3, fig. 10), NHMB J13764.
3. Paralectot>pe, Welter (1922: pi. 163(9), figs. 3-4),
GPIBo W216b.
4. Holotype, Monophyllites (Palaeophyllites) praekieperti
Renz and Renz (1948: pi. 4, fig. 5), NHMB J13766.
5. Paralectotvpe, Welter (1922: pl.l63(9), figs. .5-6),
GPIBo W216c.
6. Unfigured specimen from Chios, Monophyllites {Pa-
laeophyllites) thalman/U, NHMB J13765.
aware that Spath (1934: 298) had pre-
viously designated this specimen as lecto-
type. The smaller specimens of Welter's
plate 163(9), figures 3-6, are characterized
by a more subdued ribbing pattern. These
specimens are clearly juvenile forms; the
ornamentation increases on the mature
body chamber. The Chios and Timor speci-
mens of ^'thalmanni' are immature forms
but clearly conspecific. The species
praekieperti established by Renz and Renz
( 1948: 80) is merely a small, juvenile speci-
men of steinmanni. Measurements for 6
specimens from Timor and Chios are given
on Table 50.
Occurrence. Subcohimbites fauna of
Chios and the Frohungarites fauna with
manganese coated fossils of Timor.
Repository. Lectotype, GPIBo-W216a;
paralectotypes GPIBo- W216b, c; specimens
from Chios, plesiotype MonopJiyUites (Pal-
aeopJiyllites) tludmanni Renz and Renz
(1948: pi. 3, fig. 10) NHMB J13764; un-
figured specimen from Maradovuno NHMB
J13765, from Kephalovuno NHMB J13836;
holotype MonophyUites (PaleoplujUites)
praekieperti Renz and Renz ( 1948, pi. 4,
fig. 5) NHMB J13766; unfigured paratype
NHMB J13767.
528 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Table 51. Measurements of Ussurites sieveri
N. SP. FROM TOBIN FORMATION, NeVADA.
D
w
H
u
W/D
H/D
U/D
1.
62.5
20.0
25.8
17.3
32.0
41.2
27.6
2.
48.0
16.4
21.8
13.2
34.1
45.4
27.5
3.
48.0
15.3
21.4
13.7
31.8
44.5
28.5
4.
43.0
14.5?
19.5?
10.0
34.4?
45.3?
23.0
5.
38.7
?
16.5
11.5
?
42.6
29.7
6.
38.4
?
15.5
10.0
?
40.3
26.0
7.
33.5
11.4
14.4
9.3
34.0
42.9
27.7
8.
26.7
10.4
12.3
6.8
38.9
46.0
25.4
9.
25.0
•p
11.1
6.4
?
44.4
25.6
10.
21.0
7.8
9.1
6.1
37.2
43.3
29.0
11.
20.6
8.4
9.0
5.7
40.7
43.6
27.6
12.
20.0
7.7
9.0
4.8
38.5
45.0
24.0
1. Holotvpe, MCZ 9452 (PI. 32, figs. 1, 2).
3. Paratvpe, MCZ 9456 (PI. 32, figs. 6, 7).
4. Paratype, MCZ 9455 (PI. 32, fig. 5).
5. Paratype, MCZ 9454 (PI. 32, fig. 4).
6. Paratype, MCZ 9472 (PI. 35, figs. 10, 11).
10. Paratype, MCZ 9464 (Fig. 48 CD).
2, 6, 7, 9, 11, 12. Unfigured paratypes, MCZ 9484.
Genus Ussurites Hyatt, 1900
Type species, Monophyllites sichoticus
Diener, 1895
Ussurites sieveri n. sp.
Plate 31, figure 8; Plate 32, figures
1-7; Plate 35, figures 10, 11; Text-
figure 48
This .species is well represented in the
Tobin Fonnation fauna. There are twelve
specimens sufficiently well preserved and
complete to allow measurements, which
are given on Table 51.
The conch is moderately involute with
a rounded venter, broadly rounded lateral
areas, a well rounded umbilical shoulder,
and a vertical umbilical wall. The flanks
bear low, narrow, slightly prosiradiate folds
that cross the venter.
The sutures from a whorl height of 2.0
mm to a whorl height of 14.8 mm are shown
in Figures 48A-D. The suture is typical
for the genus, with the asymmetric, club-
shaped saddles, the large denticulated first
lateral lobe, etc.
This species differs from Ussurites mans-
ficldi n. sp. in conch form and suture. The
latter species is quite distinctive in its large
asymmetrical second lateral saddle; like-
wise the ontogenetic changes in the whorl
shape are quite different. Ussurites hosei
n. sp. is a much more robust species, with
a more inflated whorl section and a suture
with coarser denticulations on the lobes.
Occurrence. Tobin Formation, Pershing
County, Nevada; south tip of Tobin Range,
Cain Mountain 1:62,500 quad., center NW
% sec. 9, T. 26N, R. 39E, 5,500 ft. S, 27.5 ft.
W from elevation point 5088 on range
crest.
Repository. Holotype MCZ 9452 ( PI. 32,
figs. 1, 2); paratypes MCZ 9464 (PI. 31, fig.
8), MCZ 9453 (PI. 32, fig. 3), MCZ 9454
(PI. 32, fig. 4), MCZ 9455 (PL 32, fig. 5),
MCZ 9456 (PI. 32, fig. 6, 7), MCZ 9472
(PI. 35, figs. 10, 11); unfigured paratypes
MCZ 9484; suture specimen (Figures 48C,
D), MCZ 9464.
Ussurites hosei n. sp.
Plate 33, figures 1-6; Text-figure 48
This new species is based on a fairly
large assortment of fragmentary and par-
tially crushed specimens that are, however,
so distinctive that a description and new
name is warranted. The conch is robust,
and moderately involute. None of the
specimens are in a state of preservation or
completeness to yield any significant mea-
surements. The whorl section is broadly
oval and quite variable in relative width-
height dimensions. Most of the material
shows the whorl width to be approximately
75 per cent of the whorl height, thus
slightly compressed. On the other hand,
fragments of whorls are present in which
the whorl height and width are approxi-
mately the same, and in some specimens
the whorls are depressed, that is, the whorl
width is greater than the whorl height.
The venter is broadly rounded grading
onto broadly arched flanks. The umbilical
shoulder is rounded, and merges with a
broad, nearly vertical umbilical wall. The
umbilicus appears to measure approxi-
mately 20-25 per cent the diameter of the
conch.
None of the specimens are really suf-
ficiently well preserved to show surface
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 529
H
Figure 48. Diagrammatic representations of the suture of: A-D, Ussurites sieveri n. sp. A, holotype at a diameter of 35
mm (MCZ 9452); B, paratype at a diameter of 26 mm (MCZ 9472); C, paratype at a diameter of 21 mm; D, at o diameter
of 13 mm (MCZ 9464); E, paratype of Ussurites hoes/' n. sp. at a diameter of 53 mm jUSNM 153089); F-l, Ussurites mansHeldi
n. sp.; F, paratype (PI. 45, figs. 2, 3), at a diameter of 110 mm; G, at a diameter of 75 mm (MCZ 9515, PI. 44, fig. 1);
H, at a diameter of 43 mm (MCZ 9513, PI. 44, figs. 2, 3); I, at a diameter of 24 mm (USNM 153090).
Specimens of figures A-D are from Tobin Formation, Nevada, tfiat of figure E from Tliaynes Formation, Confusion Range,
Nevada, and tfiose of figures F-l, from Columbites fauna, Thaynes Formation, soutfieasfern Idaho.
530 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
Table 52. Measurements of Ussurites mans-
fikldi n. sp. from the colvmbites zone,
SOUTHEASTERN IdAHO.
D
W
H
U
W/D H/D U/D
1. 203.0 ? 93.0 53.0 ? 45.8 26.1
2. 126.0 39.5? 53.6 .38.0 31.3? 42.5 30.2
3. 48.0 ? 24.2 12.8 ? 50.4 26.7
4. 43.0 ? 20.6 13.6 ? 47.9 31.6
5. 37.7 18.0 18.7 11.4 47.7 49.6 30.2
1. Holotype, State Historical Museum, Boise.
2. Parat\pe, State Historical Museum, Boise.
3. Paratype, MCZ 9513 (PI. 44, figs. 2, 3).
4. 5. Paratypes, USGS.
marking of any kind. One of the larger
phragmocones (PL 33, fig. 1) does appear
to have broad, low, radial folds on the
flanks. The sntnre is typical for the genus,
and is illustrated on Figure 48E. It is some-
what like the suture of U. sieveri but has
much coarser denticulation of the lobes;
it differs from the suture of U. mansficldi
to a very marked degree.
Occurrence. Collection Mill, 1,420 to
1,530 feet above the base, Thaynes Forma-
tion, Confusion Range, Utah, from section
15 of Hose and Repenning (1959).
Repositon/. Holotype, USNM 153085
(PI. 33, fig. 1); paratypes USNM 153086
(PI. 33, fig. 2), USNM 153087 (PI. 33,
figs. 3, 4), USNM 153088 (PL 33, figs. 5,
6); suture specimen USNM 153089 (Fig.
48E).
Ussurites monsfieidi n. sp.
Plate 44, figures 1-3; Plate 45, figures
1—3; Text-figure 48
The Columhites fauna of southeastern
Idaho has yielded 10 specimens of this
most interesting species. The available
measurements of five of these specimens
are listed in Table 52. There is one ex-
ceptionally large specimen, one of inter-
mediate size, and the remaining forms of
relatively small diameter. The largest speci-
men, and holotype, is preserved only on
one side and the adoral one-half volution,
which is body chamber, is crushed. The
inn(M- volutions are not crushed. These
inner volutions have rounded lateral areas,
rounded umbilical shoulders and a steep,
rounded umbilical wall. The height of the
whorl in relation to the width increases
greatly during shell growth. On approxi-
mately the first two volutions, the whorl
width and height are much the same; on
later volutions the whorls increase rapidly
in height whereas the width increases quite
modestly. At all stages of growth the
venter is broadly rounded.
The shell of the body chamber of the
large holotype bears fine, slightly sinuous
growth lines which are periodically bun-
dled to give rise to faint broad ribs. On
the inner volutions the shell bears faint
strigations in addition to extremely fine
growth lines.
The large paratype (PL 45, figs. 2, 3)
is all phragmocone with much of the shell
preserved. The widely spaced, low, broad
radial ribs are more conspicuous on this
specimen, as are the fine, sinuous growth
lines.
The small specimens show the whorl
width to approximate the whorl height.
Likewise, the shell is smooth, except for
extremely fine growth lines and strigations.
Probably the most distinguishing feature
of this species is the suture ( Figure 48F-I ) .
There is a large denticulated first lateral
lobe, a much smaller second lateral lobe
and an auxiliary series on the umbilical
shoulder and wall. The saddles, however,
are unusual, especially the long asym-
metrical second lateral saddle. Figure 48
compares four sutures taken from a whorl
height of 12 mm ( diameter of approxi-
mately 24 mm) to one taken from the
paratype at a whorl height of 47 mm ( diam-
eter of approximateh' 110 mm). The dis-
tinctive character of ihv suture is already
well established at a small diameter. It
is interesting to note that on the late mature
suture (Fig. 48F) a "degeneration" occurs,
expressed in the wavy outline of the saddles
and to some extent in the lobes.
The large, asymmetrical second lateral
saddle sets this species apart from all other
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 531
species of Ussurites. In addition to the
suture, the general shape and configura-
tion of the conch is distinctive. The two
other American species of Ussurites, U.
sievcri and U. hosci, are quite different;
however, here the age factor may be sig-
nificant. Ussurites mans-fieldi is from the
Cohimhites Zone. Nearly all other Scythian
species of Ussurites are from the next
higher SuhcoJumhites Zone.
Occurrence. The holotype and paratype
were collected by Mr. Gordon R. Stephen-
son in Webster Canyon, Freedom Quad-
rangle, southeast Idaho ( Sec. 1, T. 8S, R.
45E and Sec. 6, T. 8S, R. 46E). The
paratype was found in a black limestone
concretion within a 42 foot dark shale and
thin limestone bed 120 feet above the
Meekoceras limestone. The holotype was
not found in place, but in a concretion at
the base of a cliff with the above mentioned
concretions which yielded the paratype.
In addition, the species is known from the
Columbites fauna at Hot Springs, southeast
Idaho, and along Draney Creek, Stewart
Flat Quadrangle, southeast Idaho ( USGS
Locality M98).
Repository. Holotype and large paratype
(Pi. 45, figs. 1-3) are in the Department
of Geology, Washington State University,
Pullman, Washington; figured paratypes
MCZ 9513 (PI. 44, figs. 2, 3) and MCZ
9515 (PI. 44, fig. 1); unfigured paratypes
from Hot Springs MCZ 9514; suture speci-
men (Fig. 481) USNM 153090.
Genus Leiophyllifes Diener, 1915
Type species, Monophyllites suessi Moj-
sisovics, 1882
Leiophyllifes variabilis (Spath)
Plate 22, figures 5-10; Text-figures
49, 50
MonoplujUitcs pitainalia, — Arthaber (non Diener)
1911: 234, pi. 20(4), figs. 9-11; C. Renz,
1928: 155.
Monopht/Uites (Leiophyllites) pitamaha Diener,
1915:" 205.
Monophi/llites {Leiophyllites) aff. pitamaha, — Renz
and Renz, 1947: 61- Renz and Renz, 1948: 76,
pi. 3, figs. 9-9a, pi. 4, figs. 7-7b.
Eophvllites variabilis Spath, 1934: 296, pi. 2,
fig. 3, pi. 6, fig. 1, pi. 7, fig. 1.
Eophyllites variabilis var. involuta Spadi, 1934:
296 (=Ardiaber, 1911: pl. 20(4), fig. 9).
Eophyllites variabilis var. cvoluta Spath, 1934:
296, pl. 4, fig. 1.
Monophyllites (Leiophyllites) praeconfiicii Renz
and Renz, 1947: 61, 77; Renz and Renz, 1948:
73, pl. 4, figs. 1-lb, 2-2a.
Monophyllites (Leiophyllites) georgalasi Renz and
Renz," 1947: 61, 77; Renz and Renz, 1948:
74, pl. 4, figs. 3-3a.
Monophyllites (Leiophyllites) palaeotriadicus Renz
and Renz, 1947: 61, 78; Renz and Renz, 1948:
75, pl. 4, figs. 4-4a.
Leiophyllites praematiinis Kiparisova, 1958b: pl.
7, fig. 13, text-fig. 17b; Kiparisova, 1961: 134,
pl. 28, figs. 5, 6, text-figs. 101, 102.
The lectotype of this species (Arthaber,
1911: pl. 20(4), fig. 11) is not preserved
in the collection of the Paleontological In-
stitute, Vienna; the two figured paralecto-
types (Arthaber, 1911: pl. 20(4), figs. 9,
10; Pl. 22, figs. 5-10 of this report) are
fortunately still preserved and available
for study. These two paralectotypes plus
one additional unfigured specimen (Pl. 22,
figs. 7, 8) and the many topotypes in the
British Museum (Natural History) are not
well preserved and the larger of the para-
lectotypes (Pl. 22, figs. 5, 6) has been
ground and polished.
The Chios fauna contains a fair number
of specimens which belong to this species
but which Renz and Renz ( 1948 ) placed
in four distinct species. The differences
^jetween these species are mainly in whorl
dimensions and expressions of fine orna-
mentation. The measurements of the
Chios and Albania specimens considered to
belong to this species are listed on Table
53 and plotted on the graph of Figure 50.
These data do not suggest that the differ-
ences in whorl dimensions are anything
more than what should be expected. It is,
however, possible that some of the un-
figured specimens in the Chios collection
placed here (e.g. the more involute fonns)
are really species of EophijUites. In regard
532 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
A
Figure 49. Diagrammatic representation of the suture of: A, paralectotype Eophyllitei variabilis Spatfi [=i Monophyllites
pitamaba Arthaber [non Diener] 1911: pi, 20(4), fig. lOc; PI. 22, figs. 5, 6 of this report), from Subco/umb/tes fauna of
Albania at a diameter of 28 mm; B, paratype Leiophyllites praematurus Kiparisovo (1961: 135, fig. 101), from Subco/um-
bites fauna, Primorye Region, Siberia; C, holotype Leiophylliles radians Astakhova (1960a: 146, fig. 12), from Stacheites
Zone of Astakhova (1960a) Mangyshlak Peninsula; D, holotype Danubites [Prellorianites] maritimus Kiparisovo (1961: 146,
fig. Ill), from Subco/umb/fes fauna of Primorye Region, Siberia, at a whorl height of 8 mm; E, holotype Danubites [Pre-
Horianites) inliatus Kiparisovo (1961: 145, fig. 110), from the Subco/umb/tes fauna of Primorye Region, Siberia, at a
whorl height of 7 mm; F, holotype Leiophyllites serpenfinus Chao (1959: 149, fig. 48a), from Subco/umbifes fauna of
Kwangsi, China, at a whorl height of 16 mm; G, holotype Leiophyllites serpent/nus Chao (1959: 149, fig. 45c), from Sub-
columbites fauna of Kwangsi, China, at a diameter of 23 mm; H, holotype Leiophyllites oxynotus Chao (1959: 150, fig.
48b), from Subco/umbites fauna of Kwangsi, China, at a whorl height of 10 mm; I, holotype Danubites [Danubites] incertus
Kiparisovo (1961: 143, fig. 108), from Subco/umbites fauna of the Primorye Region, Siberia, at a whorl height of 10 mm; J,
holotype Danubites (Donubifes) admaris Kiparisovo (1961: 142, fig. 106), from Subco/umbites fauna of the Primorye Region,
Siberia, at a whorl height of 7 mm.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 533
to surface ornamentation, the differences
noted by various authors are more a reflec-
tion of preservation than anything else.
The sutures, likewise, in all these species
have the same basic pattern with minor
differences which I consider to be intra-
specific. The suture of the paralectotype
as reproduced by Arthaber (1911: pi. 20(4),
fig. 10c) is idealized. The suture was ex-
posed by grinding and a new drawing is
reproduced here on Figure 49 A.
Comparison with the other upper Scy-
thian species of LeiophijUitcs, namely from
the Siibcohimbites fauna of Kwangsi,
China, and L. praematunis from the Siih-
coliimbifes fauna of the Primorye Region,
is difficult because of lack of information
on these species. Chao (1959) described
four species of Leiophijllifes on the basis
of seven specimens. These species were
separated by minor morphological features
that appear more understandable as reflec-
tions of the rather poor preservation. On
the features of conch evolution and whorl
shape and dimensions, this species cannot
be separated from L. variobilis from Al-
bania and Chios. However, the sutures of
these Kwangsi specimens are simpler than
the Albanian and Chios species (Fig. 49),
and on this basis I believe they should be
kept in a distinct species group.
A species which I believe to be con-
specific with L. variabilis is L. praematunis
Kiparisova ( 1958b ) . That author compared
her species mainly with Middle Triassic
species of LeiophijUitcs, merely noting the
similarity of the conch to that of L. vari-
abilis. In this regard she pointed to the lesser
denticulation of the lobes as an important
distinction. The suture pattern of L. prae-
matunis as illustrated by Kiparisova falls
well within the variations found within the
Chios specimens of L. variabilis (Fig. 49).
Occurrence. Siibcohimbites fauna of Al-
bania, Chios, and Primorye Region.
Rc])ository. The primary types are in
the Paleontological Institute, University of
Vienna; the lectotype (Arthaber, 1911: pi.
20(4), fig. 11) is apparently lost, but two
Table 53. Measurements of Leiophyllites
VARIABILIS (SpATh) FROM THE SUBCOLUMBITES
FAUNAS OF Albania and Chios.
D
w
H
u
W/D
H/D
U/D
1.
96.2
20.6
33.8
37.2
21.4
35.1
38.7
2.
68.8
16.7
20.0
34.3
24.3
29.1
49.9
3.
60.7
17.4
20.2
26.4
28.7
33.3
43.5
4.
52.2
13.5
15.0
27.0
25.9
28.7
51.7
5.
41.0
9.8
11.2
27.8
23.9
27.3
67.8
6.
39.5
10.5?
12.0
19.2
26.6?
30.4
48.6
7.
38.4
12.3
11.3
19.0
32.0
29.4
49.5
8.
37.4
12.5?
12.0
17.6
33.4?
.32.1
47.1
9.
35.2
9.1
13.1
13.8
25.9
37.2
39.2
10.
34.5
8.2
12.0
14.5
23.8
34.8
42.0
11.
33.7
6.7
10.0
15.5
19.9
29.7
46.0
12.
30.5
8.0
9.3
13.3
26.2
30.5
43.6
13.
28.0
7.4
10.4
11.0
26.4
37.1
39.3
14.
26.5
7.7
9.1
10.6
29.1
34 .3
40.0
15.
24.6
6.0
9.3
9.5
24.4
37.8
38.6
16.
21.5
6.3?
6.8
9.8
29.3?
31.6
45.6
17.
21.0
6.3
7.5
7.3
30.0
35.7
34.8
18.
19.1
5.8
7.4
7.0
30.4
38.7
36.6
19.
19.0
4.4
5.8
8.5
23.2
30.5
44.7
20.
18.0
5.3
7.0
5.6
29.4
38.9
31.1
21.
17.3
5.2
6.6
5.8
30.1
38.2
33.5
1.
3.
6.
Plesiotype, Monophtjllifes (Leiophyllites) aff. pita-
malw, — Renz and Renz (1948: pi. 3, figs. 9-9a),
NHMB J13752.
Holotvpe, MonophijUites (Leiophyllites) praecoiifticii
Renz and Renz (1948: pi. 4, figs. 1-lb), NHMB
.113740.
Plesiotype, MonophijUites (Leiophyllites) aff. pita-
maha, — Renz and Renz (1948: pi. 4, figs. 7-7b),
NHMB J 137.53.
5. Unfigured paratypes, MonophyUites (Leiophyl-
lites) praeconfucii Renz and Renz from Maradovuno,
NHMB J13742.
( Leiophyllites ) praeconfucii
pi. 4, fig. 2-2a), NHMB
MonophyUites
Renz (1948:
( Leiophyllites ) georgalasi
pi. 4, figs. 3-3a), NHMB
MonophyUites ( Leiophyllites)
Renz (1948: pi. 4, figs. 4-
palaeotriadi-
4a), NHMB
Paratype,
Renz; and
.T13741.
7. Holotvpe, MonophyUites
Renz and Renz (1948:
J13744.
8. Holotype,
CHS Renz and
J13749.
9-11, 13-15, 17-21. Unfigured specimens of Monophyl-
lites (Leiophyllites) aff. pitamaha from Maradovuno
NHMB J13754.
12. Paralectotype, Eophyllites variabilis Spath (^Mono-
phyUites pitamaha Arthaber (non Diener), 1911: pi
20(4), figs. 10 .i-c), PIUV.
16. Paralectotype, Eophyllites variabilis Spath (=Moiio-
phylUtes pitamaha Arthaber (non Diener), 1911: p,
20(4), figs. 9a, b), PIUV.
figured paralectotypes (Arthaber, 1911: pi.
20(4), figs. 9, 10) and one unfigured para-
lectotype of Arthaber are preserved in that
institution. A large collection of topotypes
is in the British Museum of Natural His-
534 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
^U
X
35
—
X
30
—
X
X
25
~~
X
20
—
xx
X
•
•
•
15
—
X
X
X
X
••
•
10
—
X
X •
X X
•
5
T
X
•
•
X •
•1 1 1
1
1
1 1
1 1
u
w
10 20 30 40 50 60 70
DIAMETER
80
90
100
Figure 50. Variation in umbilical diameter (U) and whorl width (W) of Leiophyllites variabilis from Subcolumbites faunas of
Albania and Chios. The data on this graph are from Table 53.
tory. Tlie Natural History Museum, Basel,
contains the following specimens from
C]hios studied hy Renz and Renz (1948):
plesiotype, MonopJujUitcs ( LciophijUitcs)
aff. pitamaha Renz and Renz (1948: pi. 3,
fig. 9) NHMB J13752, (pi. 4, fig. 7)
NHMB J13753; unfigured specimens from
Maradovuno NHMB J 13754, from Kep-
halovuno NHMB J13755; holotype, Mono-
phijUites (LeioplniUitcs) praeconfucii Renz
and Renz (1948- pi. 4, fig. 1) NHMB
J 13740; paratype (pi. 4, fig. 2) NHMB
J 13741; unfigured paratypes from Mara-
dox'uno NHMB J13742, from Kephalovuno
NHNHi J13743; holotype, Monopliyllitcs
(LeiopJit/IIilcs) ii,corfi,aIasi Renz and Renz
(1948: pi. 4, fig. 3) NHMB J13744; un-
figured paratypes NHMB J13745; holotype,
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 535
MonophijUitcs {LciophijUiics) paheotri-
adicus Renz and Renz (1948: pi. 4, fig. 4)
NHMB JL3749; iinfigiired paratypes from
Maradovuno NHMB J 13750, from Kep-
halovuno NHMB J13751.
Leiophyllites radians Astakhova
Text-figure 49
LciophijUitcs radians Astakhova, 1960a: 146, pi.
34, fig. 10, text-fig. 12.
This species has the general leiophyllitid
conch form but is characterized by broadly
spaced, low radial ribs that cross the venter.
Adorally the ribs tend to decrease in size
and eventually disappear. The suture is
shown on Figure 49C. Assuming that it is
accurately drawn, it is a much simpler
suture than that of most other species of
this genus. The only other species of
LeiophijUites that have ornamentation con-
sisting of ribs are the two species from
the Primorye Region (L. mcuitimus and L.
admcu'is). However, these two species
have quite different ribbing patterns and
very different sutures ( Fig. 49 ) .
Occurrence. Stacheites Zone of Astak-
hova (1960a) Mangyshlak Peninsula.
Leiophyllifes serpentinus Chao
Text-figure 49
Leiophi/llitcs serpentinus Chao, 1959: 149, 331,
pi. 42, figs. 7, 13-15, text-fig. 48a.
LeiophijUites oxijnotus Chao, 1959: 150, 332,
pi. 42, figs. 11, 12, text-fig. 48b.
LeiophijUites lolouensis Chao, 1959: 150, 332, pi.
42, figs. 8-10, text-fig. 48c.
LeiophijUites aff. pitamaha Chao, 1959: 150, 332,
pi. 42, fig. 1.
LeiopliijUites kwangsiensis Chao, 1959: 7, 160,
{nomen nudum).
LeiopliijUites verniifornns Chao, 1959: 7, 160
( nomen nudum ).
All the species listed above in the syn-
onymy are based on one or very few speci-
mens of only fair preservation, and all came
from the same horizon and locality. Chao
describes the venter on some of his species
as fastigate but this is not apparent on the
illustration of the species. On the basis of
the data available, all these species have
the general conch architecture of L. vari-
abilis and are considered to be conspecific.
They differ from /.. variabilis in the suture
(Fig. 49).
Occurrence. Limestone block (Chao col-
lection 542b) Kwangsi, China.
Leiophyllifes admaris (Kiparisova)
Text-figure 49
Danubites (Danubites) admaris Kiparisova, 1961:
142, pi. 28, fig. 11, text-fig. 106.
Danubites (Danubites) aff. floriani Mojsisovics, —
Kiparisova, 1961: 141, pi. 28, fig. 10, text-fig.
105.
Danubites (Danubites?) nice;t»s Kiparisova, 1961:
143, pi. 29, figs. 1, 2, text-figs. 107, 108.
The assignment of these species and
maritimus to LeiophijUites is done entirely
on the basis of the suture pattern ( Fig. 49 ) .
The ornamentation of these species and
especially that of maritimus would ally
these species to Prcflorianites. In the inter-
pretation followed here the suture is con-
sidered the more critical sign post of genetic
affinity and the ornamentation a case
of homeomorphy. The different species
brought together here differ mainly in the
degree of ribbing.
Occurrence. Subcohtmbites fauna, Pri-
moiye Region, Siberia.
Leiophyllifes marifimus (Kiparisova)
Text-figure 49
Preflorianites maritimus Kiparisova, 1958b: pi. 8,
fig. 3, text-fig. 22b.
Danubites (Prcflorianites) maritimus Kiparisova,
1961: 146, pi. 29, figs. 8, 9, text-fig. 111.
Danubites (Preflorianites) inflatus Kiparisova,
1961: 145, pi. 29, figs. 3, 4, text-figs. 109-110.
Danubites (Preflorianites) aff. maritimus Kipari-
sova, 1961: 147, pi. 29, fig. 10, text-fig. 112.
This species has a suture (Fig. 49D)
much like that of L. admaris but the rib
pattern is completely preflorianitid in as-
pect, that is, concentrated near the vunbili-
cal shoulder. The specimen from an upper
Scythian horizon in the Toad Formation
of British Columbia that Tozer (1965a:
40) assigned to LeiophijUites sp. indet. is
quite similar to L. maritimus.
Occurrence. Subcolumbites fauna, Pri-
morye Region, Siberia.
536 BiiUetin Museum of Comparative Zoology, Vol. 137, No. 3
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542 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
INDEX
345, 346, 347, 506
396
Acrochordiceras (Paracrochordiceras) anodosum,
351
acutangukitus (Etikashmiritcs), 490
(Kashmiritcs), 490
actitus (BaJatonites), 506
(Doricranitcs), 328,
(Dorikmnitrs), 506
— ■ (Procarnites), 352, 391, 395
adai (Columhites), 345
admaris {Dcmubites [Danuhites]) , 354, 532, 535
{LeiophylUtcs), 328, 355, 535
aithaUac (Columhites), 340, 434, 435
alhanicum {Pscudosageceras) , 325, 334, 340, 360,
363, 364, 582
( Sageccros ) , 338, 363
var. (Sageceras), 341
Albanites, 323, 327, 344, 440, 463, 477, 478, 479,
481, 572, 576, 580
arhanus, 480, 481
var. mediterranea, 481
danispanensis, 346, 478, 481
osmanicus. All, 479, 480, 481
■ triadicus, 327, 334, 340, 347, 350, 441, 457,
477, 478, 479, 480, 481, 482, 572, 574, 576,
580
welteri. All, 478, 481
alexeevae (Nordophiceias), 337, 357, 452, 466,
467, 468, 469, 471
(dcxi {Tiiugkmites), 326, 340, 422, 423, 424, 580
aJi (Prosphingites), 325, 338, 340, 405, 407, 580
(dtcrnccostatits (Dantdntcs), 350
altm (Dinarites), 356, 489
(Dinarite.s- [Olcnekites]) , 489
i Olcnekites-), 356, 489
and)ika { DatuiJ)ites), 350
ainericatuis (Std)cohim])ites), 326, 335, 358, 433,
435, 436, 437, 600, 602
Anunonites ])(>gd(>(iiius, 313, 505
niiddeiidorffi, 486
reuttensis, 516
sp. indet., 450, 451, 452, 592
(Ceratites) cassianiis, 493
aniuremi.s {Eopliullites), 328, 354, 523, 526, 527
Anakaslimiiites, 327, 329, 437, 490
nivalis (aff.), 352
sp. indet., 348
Ana.sihiriles, 483
cmgidoso (aff.), 340, 483
dicholDnms (cfr. ), 478
gmritis, 345, 346, 448, 477
kingianus, 483
nirieostatiis, 356
s-nbgracili.s, 346
(indnisovi ( Prncarnifes), 345, 346, 391, 392, 395,
396
angulatus {Cordille rites), 325, 337, 340, 353, 358,
360, 364, 365, 366, 580, 642
(cf. Curd ill elites), 341, 364, 365
? (Dinarites), 342, 506, 680
angtdoso aff. (Anusihirites), 340, 483
341
Pronorites), 341, 477,
angtistecostatus (Durgaites), 485
— (Keyserlingitcs), 350, 351, 485
angustilohatus (Tirolites), 342, 493, 499, 670
var. alpha (Tirolites), 493, 499, 670
angustus (Tirolites), 342, 493, 494, 495, 678
anodosum (Acrochordiceras [Paracrochordiceras] ),
351
anomalus (Subcolumbites), 354, 436
antiglobtdus (Isculitoides), 413, 416
(Iscuhites), 340, 413, 417
var. (Iscidtites), 340
apostolicus (Celtites), 360, 369, 646
("Celtites"), 369
(Dieneroceras), 337, 357, 360, 367, 369,
371, 372, 646
arhani sp. ind. ex aff. (Pronorites), 349, 477, 478,
479
arhanus (Albanites), 480, 481
var. meditrterranea (Albanites), 481
(Pronorites), 338, 341, 349, 477, 478 479,
482, 574
var. (Pronorites) .
var. mediterranea
478, 482
var. sundaica (Pronorites), All
arctieus (Zenoites), .326, 357, 411
Arctoceras blom.strandi, 468
simplex, 356, 468
Arctohungarites primoriensis, 355
Arctomeekoceras, 327, 476
rotundatum, 327, 356, 476
sp. indet., 348, 476
Aretoprionites, 522
ArctotiroJites, 327, 329, 332, 477
menensis, 327, 356, 477
Arianites, 326, 329, 446, 544
musaeehi, 326, 338, 340, 443. 446, 544
(MeropcUa) plcjanae, 341, 447
armatus (Celtites), 437, 490, 594
arnautieus ( Arnautoceltites) , 397, 400
(Celtites), 338, 397, 399, 400, 401, 552
Arnautoceltites, 325, 332, 335, 397, 400, 402, 550,
554, 602
arnautieus, 397, 400
baiarunasi, 325, 347, 397, 401
^'rac77z.v, 325, 355, 397, 401
invohitus, 325, 353, 397, 401, 402
mediterraneus, 325, 340, 397, 400, 401,
402, 552, 554
teicherti, 325, 358, 397, 399, 402, 403, 602
arthaberi (Eogijmnites) , 328, 340, 341, 517, 582
(Japonites), 517
(Prnptijehites), 341, 349, 387
390
( Propli/chitoides),
390, 391, 590
325, 350,
388, 389,
384, 385,
asiatieus (Columhites), 345
a.spencnsi.s (Paranannites), 346
\ar. europaea (Paranannites), 341, 397,
400, 401
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 543
Aspidites, 572
has-scrti, 338, 478, 482, 572
imimiualis, 338, 339, 572
muthianus, 383
sihiiicus, 453
(issi/mctiicus (Cohiinhites), 352, 429
astakhovi (Tirolites), 337, 360, 499, 502, 650
atkisovicmi.s (McpaphyUites) , 355
attcnmiius (Dalmatites), 342, 343, 359, 524, 684
au.stini (Prosphingites), 403, 404, 405, 407
hajarunasi (Arnaiitoceltites), 325, 347, 397, 401
(Nannitcs), 346, 401
Balatonitc's actittis, 506
bogdoanus, 505
wssictis, 505
halcanicus (Proptychites), 341, 385, 387, 388
Balkanites, 327, 343, 465
tahulatus, 327. 343, 465
bearlakeii.si.s (Keyserlingites), 327, 360, 485, 486,
488, 614, 616
hearriverensis (Keijserlingites) , 327, 360, 485, 486,
487, 614
Beatites, 327, 329, 449, 582
hcrtJwc, 327, 338, 340, 449, 582
hcnwullii (Koninckites), 341, 388, 389
var. (Koninckites), 341
herthac (Beatites), 327, 338, 340, 449, 582
bertisci (Pwptychites) , 338, 386, 388, 389, 558
(Proptychitoides), 388, 389
betilloni (Monophi/Ilites [SchizophyUites]), 341,
523, 524, 526 "
var. evoliita (MonoplniUites [Schizophyl-
lites]), 341, 524, 526
Beyhchites, 328, 329, 342, 516
khanikoffi, 349
laurae, 328, 340, 341, 516, 517
praematurus, 341, 516
Beyriehitidae, 328, 516
biangulatus (Dinarites), 342, 504, 664
bicariuatiim (Pseudosageceras) , 358, 364, 366
bicaiinatus (LaneeoUtes) , 450
binodosiis (Ceratites), 317
bispinatiis (Tirolites), 343, 497
bittneri (Bittnerites), 328, 343, 504, 505, 654
(Tirolites [Bittnerites]), 343, 504
Bittnerites, 328, 343, 491, 504, 505, 654
bittneri, 328, 343, 504, 505, 654
malici, 504, 505, 654
teUeri, 505. 654
blaschkei ( Kashmir ites), 490
blomstrandi ( Arctoceras) , 468
bogdoanus (Ammonites), 313, 505
(Balatonites), 505
(Ceratites), 505
(Doricranites), 328, 345, 346, 347, 499,
505, 506
(Dorikranites), 505
(Goniatites), 505
Boreomeekoccras, 327, 329, 332, 471, 476
keyserlingi, 327, 356, 476, 477
rottindatum, 476
bridgesi (Dagnoceras), 459
bubulinae (Cohnnbitcs) , 340, 443, 444
bulgariciis (Dinarites), 507
bungci (Ceratites), 356, 484, 485
(Keyserlingites) , 485
buxtorfi (Pwptychites), 341, 386, 388, 389
canadensis (Olenekites), 327, 357, 489
caprilense (Meekoceras), 343
carinatus (Kiparisovites), 346, 519, 520
(Prohungarites), 328, 344, 347, 520
carniolicus (Carniolites), 510, 660
(Dinarites), 328, 343, 506, 510
(Tirolites), 342
Carniolites, 506
carniolicus, 510
cassianus (Ammonites [Ceratites]), 493
(Ceratites), 493
(Tirolites), 317, 328, 342, 343, 345, 346,
491, 492, 493, 500, 501, 503, 666, 668, 670,
680
cf. (Tirolites), 359, 492, 498, 501, 503,
608
ex ,m-. (Tirolites), 356, 501
var. al))ha (Tirolites), 493
var. tenuis (Tirolites), 493
Celtites, 369, 490
aposiolicus, 360, 369, 646
armatus, 437, 490, 594
arnauticus, 338, 397, 399, 400, 401, 552
kcirensis, 338, 340, 367, 375, 548
. mulliplicatus, 437, 440, 490, 594
■ planovohis, 360, 369, 372, 373, 646
sp., 352
subrectangularis, 490
ursensis, 360, 369, 372, 373, 646
^'Celtites" apostolicus, 369
ursensis, 369
bonnevillense (Dagnoceras), 459
borealis (Popovites), 326, 357, 421, 422
Ceratitacae, 491
Ceratites binodosus, 317
bogdoanus, 505
bungei, 356, 484, 485
cassianus, 493
dalmatinus, 506
decipiens, 356, 420
discrctus, 356, 382
eichwaldi, 483
euon\phalus, 465
fissiplicatus, 356, 382
hyperboreiis, 356, 382
idrianus, 492
inostranzeffi, 356, 420
liccanus, 503
middendorffi, 356, 485, 486
muchianus, 506
multiplicatus, 356, 382
)i(/c(Y//i/, 356, 484, 486
normalis, 317
544 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
ravana, 349
schrenki, 356, 486
sigmatoideus, 356
smiriagini, 496
subwhu.stus, 356, 485
trinodosiis, 317
( Paraceratites ) prior, 342, 497, 680
"Ceratite.s" muJtiplicatiis, 382
Ccratitida, 367
cheneiii {Pseudoceltites) , 337, 359, 360, 438, 440,
628
Chioccras, 326, 329, 445, 446
mitzopouloi, 326, 334, 335, 340, 443, 445,
446
var. meridonalis, 340, 446
nodosum, 326, 340, 443, 446
chionen.sis (Paraminnitcs), [see also chiosensis],
341, 397, 399, 400, 401
chiosensis {Paraminnitcs), [see also chionensis],
397
Chiotites, 326, 329. 409, 419, 420
globidaris, 326, 340, 411, 420
superglohosus, 420
chowadei (Svalhardiccras) , 327, 358, 453
ciugidatas (Svilajitcs), 497
( Tirolites), 328, 343, 497, 680
{Tirolites [Svilaiites]), 343, 497, 500, 680
circum})Iicatiis {Diaplococcras), 504
{Dinarites), 504
( Dinarites [Liccaites] ), 342, 504
clavlsellntum { Pscndosageccras) , 363
cf. {Psemlosagcceras), 341, 361, 363
Columbites, 337, 338, 409, 424, 425, 429, 437,
440, 501, 626
adai, 345
aithaliac, 340, 434, 435
asiatictis, 345
a^symmetricus, 352, 429
hubulinac, 340, 443, 444
consanguincus, 360, 425, 427, 428, 429,
622
minimus, 360, 425, 427, 429
mirditensis, 338, 340, 429, 431, 433, 544,
546
— comtrictilis, 346, 425, 440
— costatus, 321, 352, 443, 444
— dianae, 340, 433, 434, 435
var., 340
var. evohita, 433
var. involuta, 433
— dolnapacnsis, 345, 346, 425, 440
dusmani, 338, 433
— curopacus, 338, 340, 427, 431, 433, 542,
544, 548
cur()))arus})('rrinismithi, 340, 429, 431, 433
gracilis, 345
— graccoamericanus, 340, 433, 434, 435
— hellcnicus, 340, 443, 444
— huangi, 352, 411, 443, 444
Icrantinus, 340, 443, 444
ligatiformis, 345
ligalus, 360, 425, 427, 428, 620, 624, 626
malayaitus, 340, 443, 444
var. crassa, 340, 443, 444
var., 433, 546
— morpheos, 356, 425, 501
nov. sp. indet., 441
— ornatus, 360, 425, 427, 428, 429, 620
( ? aff. ), 356, 425
— parisianus, 322, 337, 340, 346, 354, 355,
360, 424, 425, 427, 428, 430, 434, 435, 462,
483, 501, 618, 620, 622, 624, 626
(cf.),345
— pcrrinismithi, 338, 340, 427, 431, 433, 546
— plicatuli (ex aff.), 340, 443, 444
— plicatuhis, 444
— sp., 358, 425
— sp. (?),352
— sp. indet., 349, 354, 355, 443
— spencei, 360, 425, 427 428, 429, 434, 620,
624
var. chiotica, 340, 434, 435
tururpcnsis, 345
ijalienms, 352, 443, 444
compactiis {LanccoUtcs), 450
compressa ( Vssuria), 448
compressum {Meekoceras [Submeekoccras]), 466,
475
{Nordophiccras), 327, 353, 475
{Submeekoccras), 352
eomj)ressus {Cordillcrites), 366
{Danubites), 350
{Paranannitcs), 341, 397, 399, 400, 401
{Proptychitoides), 352, 390
{Pseudosageceras) , 363
{Xenoceltitcs), 352
concinnus {Cordillerites), 365, 366
confucii {LeiophyUites), 350
couneclcns {Diaplococcras), 328, 343, 504, 664
{Dinarites [Ceratites]), 504
{Dinarites [Liccaites]), 342, 504
cunsanguineus {Columbites), 360, 425, 427, 428,
429, 622
consirictilis {Columbites), 346, 425, 440
{Pseudoceitites), 424
contorttts {Eukashmirites), 327, 490
( Kashmirites ) , 346, 490
contrarium ? {Meekoceras), 465
contrarius {Meekoceras), 465, 467
{Niyrdophicera.s), 357, 466
eontratitis {Ncrrdophiceras), 465
coombsi {Prosphingites), 325, 351, 404, 405, 407,
408
Cordillcrites, 325, 332, 335, 337, 364, 366, 580,
642
angulatus, 325, 337, 340, 353, 358, 360,
364, 365, 366, 580, 642
( ef . ) , 34 1 , 364, 365
con}i)res.s-us, 366
concinnus, .365, 366
kuangsia)uis, 364, 365, 366
oricntalis, 352, 364, 365, 366
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 545
costatus (Columbites), 352, 443, 444
(Hemiprionites), 337, 357, 482
crassepUcatus (Hungarites), 349, 517, 518, 519
{Prohungarites), 328, 350, 517, 518, 520,
521
cf. (Prohungarites), 348, 373, 518, 521
crenoventrosus (Xenoceltites), 325, 352, 353, 376
Cucococeras, 510
curticostatum (Meekoceras), 359, 470, 471, 473,
475, 640
curvatuin (Psetidosageceros) , 363
czckanowskii (Prosphingites), 325, 356, 403, 404,
405, 407, 408, 592
cf. (Prosphingites), 358, 405
ex aff. (Prosphingites), 340, 407, 410
Czekanawskites, 326, 420, 421
decipiens, 326, 356, 420
( cf. ), 360, 420
— inostranzeffi, 420
sp., 420
sp. nov., 420
Dagnoceras, 327, 435, 457, 458, 459, 460, 461,
463, 465, 570, 572, 588
honnevillense, 459
hridgesi, 459
elUpticum, 327, .352, 353, 458, 460
hat/deni, 459
komonum, 338, 457, 478, 480, 481, 482,
576
hililobatum, 327, 352, 353, 458. 460
lejamim, 338, 457, 458, 459, 460, 570
nopcsanum, 327, 338, 340, 457, 458, 459,
460, 463, 464, 570, 572
var. invoJtita, .341, 464, 465
Danubites alternecostatus, 350
amhika, 350
comprcssus, 350
kansa, 350
nivalis, 437, 490
strongi, 379
• (Danubites) admaris, 354, 532, 5.35
incertus, 354, 532
floriani ( aff. ) , 535
? incertus, 535
(Prcfhnianitcs) infhitus; .354, 5.32, 535
inaritimus, 354, 532, 5.35
nuiritiinus (aff.), 535
darwini (Tirolites), 343, 493, 499, 666, 668
cf. ( Tirolites), 492
var. abbrevians (Tirolites), 496
var. cinctus ( Tirolites), 496, 499
var. costatus ( Tirolites), 496, 499
var. modestus ( Tirolites), 496, 499
var. reminisccns (Tirolites), 496, 499
decipiens ( Ceratites ) , 356, 420
(Czekanowskites), 326, 356, 420
cf. (Czekanowskites), 360, 420
(Japonites), 517
(Proptt/chitoides), .325, 340, 384, 385,
,387, 388, 389, .390, 556, 564
? (Ussurites), 423, 524, 525, 526, 586
sp. indet., .348
terbunico (aff.), 463
terbunicum, 338, 341, 457, 463, 464, 465,
572
— unicum, 355
— ? unicum, .354, 458, 461
— zappanense, 327, 338, 340, 350, 457, 458,
459, 460, 570, 588
(cf.),.344, 4.58, 4.59
dalnuitinum (Plococeras) , 506
dalmatinus (Ceratites) , 506
(Dinarites), 328, 340, 342, 343, .344, 499,
506, 507, 508, 509, 510, 656, 658, 660, 680
var. extensus (Dinarites) , 506, 660
var. externeplanatus (Dinarites), 506, 660
var. plurimcostatus (Dinarites), 506, 660
demissuni cf. (Ophiceras), 341, .345
demokidovl (Dieneroceras), 327, .357, 367, 368,
369, .372, 483
densiplicatus (Dinarites), 356, 489
( Dinarites [Olenekites]), 489
dentosus (Goniodiscus) , 453
(Svalbardiceras), .327, .356, 453
( Xenodiscus ) , 356, 452, 453
depauperatus (Gtjmnites), .350
dianae (Cohnnhitcs), .340, 4.33, 4.34, 435
var. (Columbites), .340
var. evoluta (Columbites), 433
var. involuta (Columbites), 433
Diaplococeras, 328, 329, 343, 503, 664
circumplicatus, 504
connectens, 328, 343, 504, 664
liccanum, 328, .343, 503, 504
dichotomus cfr. ( Anasibirites) , 478
cfr. (Pseudosibirites), 338, 457, 478, 481,
482, 580
cf. ( Sibirites), 478
Dalmatites, 321, 328, 329, 337, .343, 344, 522, 650,
652, 682
attenuatus, .342, .343, .359, 524, 682
kittli, 337, 343, 360, 519, 522, 523, 524,
650
morlaccus, 328, 343, 519, 522, 523, 524,
652
richardsi, 343, 524
ropini, .344, .349, .350
dnnispanensLs (Albanites), 346, 478, 481
dieneri (Dieneroceras), 354, 369
C^Durgaites"), 485
(Eophyllites), .328, 334, .335, 340, 523,
524, 525, 526, 527, 584, 586
(Keijserlingites), 348, 349, 350, .351, 485
(Monophyllites), 338, 524, 586
nov. sp. e.v. aff. (Monophyllites), 526
sp. ind. ex. aff. (Monophyllites), 349
var. involuta (Monophyllites [Leiophyl-
lites]), 341, 388, .390
— (Ophiceras), 367
546 BiiUetin Museum of Comparative Zoology, Vol. 137, No. 3
Dieneroceras, 325, 337, 355, 357, 367, 548, 578,
580, 646
apostolicus; 337, 357, 360, 367, 369, 371,
372, 646
dcmokidovi, 337, 357, 367, 368, 369, 372,
483
diencii, 354, 369
karazinU 325, 355, 367, 368, 369
khcUiVwmis, 357, 369
kncchti, 368, 369
mediterronea, 325, 340, 367, 368, 548, 578
uihdutnisi',; 357, 369, 483
skutari'u.si.s, 325, 340, 367, 368, 580
Dieneroceratidae, 325, 337, 367
Di^it(>))hiiUitcs loloticnsis; 352, 392, 397
dimidiatus ( Tiwlitcs), 342, 493, 674
Dmarites, 328, 343, 344, 345, 489, 506, 507, 680
(dtits, 356, 489
— ? anuidattis, 342, 506, 680
hiaiif^idatus, 342, 504, 664
bul^ciricus, 507
camiolicus, 328, 343, 506, 510, 662
circiimfylicattis, 504
dalmcitimi.% 328, 340, 342, 343, 344, 499,
506, 507, 508, 509, 510, 656, 658, 660, 680
var. extohsus, 506, 660
var, cxterniplanatus, 506, 660
var. )diirimcostattis\ 506, 660
densiplicatus, 356, 489
diocletiaui, 504
evohitior, 341, 342, 344, 506, 507, 508,
507,
658
^lacialis, 356, 489
inicniu'diiis, 356, 489
lacvis, 342, 356, 506, 508, 656, 658
- liutsika.si, 328, 340, 341, 344, 499, 506,
510
liccanii.s, 503
moliainedantts, 511
• (cfr.),511
muchimuis, 342, 343, 506, 507, 508, 658
midticostalus, 342, 507, 508, 660
nudii.s, 341, 342, 344, 506, 507, 508, 656
I^rof^re.ssiis, 343, 504
■,s/;in«/(/.v, 348, 376
spiuiplicatiis, 356, 488, 489
timlUnidcs, 342, 507, 508, 660
tolli, 356
undalus, 328, 344, 346, 347, 499, 506, 507,
510
vohilus, 356, 488
( Crralitcs) coiuicrtciis, 504
(Jh'rcc^ovitcs) dioclcluiiii, 342, 504, 664
moJuiDicdauus, 342, 511
( Liccaitc.s) ciiciiDiplicafus, 342, 504
coniu'clciis, 342, 504
liccanus, 342, 503
pw^re.'isus, 342, 504, 664
(Olnu'kitcs) alius, 4H9
dcnsiplicatus, 489
glacialis, 489
intcnncdius, 489
s))inii)Ucatus, 488
vohitus, 488
Dinaritidae, 328, 506
dinarus (TiwJites), 342, 493
dioclctiaui ( Dinarites), 504
i Dinarites [Hercegovites]), 342, 504,
diomjsi (Stacheites), 341, 455, 462, 463, 464,
var. ( Stacheites), 341
discoidalis (Laneeolites), 327, 343, 450
discoides (Suhdorieranites), 345, 346, 506
discretus (Ceratites), 356, 382
(Xenorcltites), 382
(Xenodiscus), 382
discus (Doricrauites) , 346
(Hemilecanities), 325, 340, 352, 353,
368, 374, 375, 590
(Lecauites), 338, 341, 374
(Proavitcs), 374
distaus (Tirolites), 342, 492, 494, 495, 672,
doJnapaensis (Cohimhites) , 345, 346, 440
(Pseudoceltites), 326, 347, 359, 425,
440, 441
Doricranites, 328, 329, 345, 346, 505
aeutus, 328, 345, 346, 347, 506
hogdoanus, 328, 345, 346, 347, 499,
506
discus. 346
Iduceolatus, 346, 505
ovatus, 346, 505
rarccostatus, 346, 505
rossicus, 345, 346, 505
schdiricus, 346, 505
tumulosus. 340, 505
Dorikrauites aeutus, 506
Ixigdoanus, 505
rossicus. 505
driiicu.se (Pscudosagecera.s), 325, 338, 340,
360, 361, 362, 363, 564
\ar. incentrolata (Pseudosageeeras) ,
664
465
367,
676
438,
505,
dukagini iParagoccras) 338, 397, 399, 400,
554
(cf. Paragoceras) , 341
Durgaites, 323, 485
"Durgaites" aiigustecuslaius, 485
diencri, 485
du.siiiaiii (CU>[uiid>ites), 338,433
{Suh,-clu,id)ilc.s). 326, 340, 431. 433,
341,
341
401,
434,
435, 436, 544
eiclucaldi (Ccralites), 483
(Sihirites), 327, 356, 483
v{. ( Sil)irilcs), 483
eiekiteii.si.s (Inyditcs), 357, 373, 374, 483
(Suhrislnuiites), 337, 357, 374
cicgaiis (TiroUh's), 346, 498, 500, 501
cllipticum (Daguoeeras), 327, 352, 353, 458, 460
cUipticus (Isculitoidcs), 326, 352, 353, 411, 414,
418, 419
eincris ( Suhri.sliiniilcs) , 325, 340, 373
Ammonoids of the Late Scythian (Lower Thiassic) • Kiimmel 547
cf. (Subvishmiites), 347, 373
(Xeuaspis), 368, 373, 374
Eo^i/mnitcs, 328, 329, 517, 582
— arthabeii, 328, 340, 341, 517, 582
Eophyllitcs, 328, 373, 523, 524, 525, 526, 531,
584, 586
amurensis, 328, 354, 523, 526, 527
dicncri, 328, 334, 335, 340, 523, 524, 525,
526, 527, 584, 586
nopcsai, 525
oricutalis, 328, 350, 523, 526
rcfmctus, 524, 525, 584
( cf . ) , 526, 527
vanabilis, 531, 532, 533
var. evoluta, 531
var. involuta, 531
Epiccltites, 326, 344, 440, 447, 546, 610
gentii, 326, 338, 340, 360, 443, 447, 546,
610
ficntii (n. sp. cf. ), 447
suhgmcilis, 326, 347, 443, 448
Epihcdenstrocm ia, 366
skipetarensis, 364
Eukoshmiritcs, 327, 329, 490
• acutdngulatus, 490
— contoitiis, 327, 490
subdimorphus, 327, 347, 490
ciiomphahim (Meekoceras) , 465
eiiomphahis (Ceratites), 465
(Nordophicera.s), 337, 357, 465, 466, 467,
468, 469, 471, 475, 634
iXcnodisciis), 356, 452, 465
ctiropaeiis (Coliimbites), 338, 340, 427, 431, 433,
542, 544, 548
(Subcolumbites), 429, 431, 433
curopaeuspeninismithi (Cohimbites) , 340, 429,
431, 433
emlutior (Dinaiites), 341, 342, 344, 506, 507,
508, 658
cvulutus {MegaphijlUtcs), 350
Fcngshanites robustus, 352, 433, 435
fislitac (Lccanites), 338, 339
fissiplicatus (Ceratites), 356, 382
flcmingianus (Flcmingites) , 317
Flemingites; 389
flemingianus, 317
psetidonmeUi, 341, 387, 388, 389, 390
radiatii.s, 317
floriani aff. (Daniibites [Danid)itcs] ), 535
floweri (Stacheites), 327, 344, 358, 455, 456, 457,
596
frebokli (Svalbardiccras), 327, 357, 453
freemani (Mctadagnoceras), 327, 350, 461, 462,
463, 464, 588
frequens (Gijronites), 317
garbiniis (Imjoites), 341, 381, 580
{Preflorianites) , 325, 340, 379, 381, 580
gentii (Epiceltites), 326, 338, 340, 360, 443, 447,
546, 610
n. sp. cf. (Epiceltites), 447
georgalasi (Moiwphi/lIHes [Lcio])hiiUites]) , 341,
531, 533, 534
gerbaensis (Tirolitcs), 356, 501
glacialis ( Di'naiites ) , 356, 489
(Dinaiites [Olcnekites] ), 489
(Olcnekites), 356, 489
globosiis (IscuIU aides), 352, 404, 407
(Prosphingites), 325, 354, 355, 404, 405,
407, 408
aff. (Prosphingites), 407
gluhuhiris (Chiotitcs), 326, 340, 411, 420
(Prosphingites [Chiotitcs]), 340, 419, 420
globtdtis (Isciditoidcs), 413, 416
(Iscultitcs), 340, 413, 415
var. (Iscidtitcs), 340
gjobtdtisantiglobulus (Iscultitcs), 340, 413, 417
gjobidusoriginis (Isciditoidcs), 417
( Iscultitcs), 340, 413, 417
Goniatites bogdoaniis, 505
Goniodiscus dentosus, 453
ttjpiis, 482
gracilis (Anasibiritcs), 345, 346, 477
(Arnautoccltitcs), 355, .397, 401
(Cohimbites), .345
(Paranannites), .354, .399, 401
gracilitatiis cf. (Meekoceras) , 341
graecoamericanus (Columbitcs) , .340, 433, 434,
435
grambcrgi (Parasihiritcs), 356, 483
(Sibirites) , AH3
\ar. mixta (Sibirites), 483
var. rariaculateus (Sibirites) , 483
gregoriji (Xenoccltitcs), 376
gutstadi (Prohimgarites), 328, 360, 518, 519, 521,
,522, 612
Gijmnites, 351
depauperatus, .350
sp., .349, .351
vasatasena, 349
volzi, 350
Gijmnitidae, 328, 517
Gijronites frequens, .317
mojsisovicsi, 451, 452
? sch m id ti, 452
hakki (Meekoceras), 338, 385, 387, 564
(Proptyehites), 387
(Proptijchitoides), 385, 387
hammoudi (Isciditoidcs), 326, .360, 411, 413, 414,
418,419, 612
hara (MonoplujUites), 338, 349, 524, 525, 584
(Monophidlitcs [Ussurites]), 524
( Ussurites), 350, 351
harti (Tirolitcs), 337, 342, 359, 492, 501, 682
ha.<!serti (Aspidites), 338, 478, 482, 572
(Meekoceras [Koninckites] ), 478
haiieri (Tirolitcs), 343, 49.3-, 499, 670
aff. (Tirolitcs), .359, 503
var. minor ( Tirolitcs), 493, 499
haydeni (Dagnoccras) , 459
548 Bulletin Museum of Comparative Zoologij, Vol 137, No. 3
Hedenstrocmia kastriotae, 338, 449
pitt/oussae, 341, 449
skipetarensis, 338, 364, 365, 366, 580
sp., 391
Hedcnstrocmiidae , 326, 448
hdenac {Picnkites), 326, 340, 441, 443
(Piosphingites [Zcnoites]), 340, 410
var. marodovunensis (Prosphingites [Zcrw-
/to]),340, 410
(Zenoites), 326, 340, 407, 410, 411
hellcnictis (Cohimbites), 340, 443, 444
Hellcnitcs, 321, 328, 332, 337, 355, 435, 460, 511,
512, 515, 516, 554, 646
idahocnse, 337, 360, 512, 513, 515, 516,
646
inopinatus, 337, 355, 515, 516
? inopinatus, 354, 516
pracmaturus, 328, 334, 335, 340, 352, 353,
512,513,514,515, 554
(tl.),513, 514, 515
var. acgacica, 340, 512, 513, 514
radiatm, 358, 513, 514, 515, 516
tchernqschewiensis, 355
trikkaiinoi, 340, 512, 513, 514
var., 341
var. graeca, 341, 512, 513, 514
iPaUusitcs) radiatus, 328, 340, 341, 514
striatus, 341, 514, 516
var., 341
var. den.sicostaia. 341, 514,
516
Ht'//t';u7/(/rt(', 328, 511
Hemilccanites, 325, 332, 335, 3.37, 344, 367, 374,
.375, 423, 590, 598, 602, 610
discus, 325, 340, 352, 353, 367, 368, 374,
.375, 590
paradiscus, 325, 358, 368, 375, 598, 602,
610
Hcmiprionites, 337, 453, 482, 483
cDstatus, 337, 357, 482
sibiricus, .356, 453
herberti (Naunites), 399, 400, 401, 552
hctcrophanus (Tiwlitcs), .342, 493, 510, 674
hilmi (Protropitcs), 326, 334, 335, 338, .340, 443,
444, 445, 446, 568
hindostanus (cf. Nannites), .341
hocsi (Ussurites), .328, 359, 528, 529, 531, 606,
608
Holland it cs, 349
vt/dsa, 350
Ilotolobus, .328, 343, 491, 511, 680
nwno))lychus, 328, .343, 499, 511, 680
huangi (Cohimbites), .352, 411, 443, 444
Ilungaiitcs, 517
cmsscplicatus, .349, 517, 518, 519
ndddlemissii, 520
(cf.),349, 517, 518, 519
tubcrculatus, .349, 518, 519
sp., .358
Hungaritidac, .328, 3.37, 517
lujpcrborcus (Ccrutitcs), 356, 382
(Xenoceltites), 382
(Xcnodiscus), 382
hybridus { Tirolitcs), .342, 493, 494, 676
idahocnse (HcUenites), 337, 360, 512, 513, 515,
516, 646
(Pseudarniotitcs), 516
(Pseudharpoccras), 319, 360, 512, 516
idrianus (Ceratites), 492
(Tiwlites), .328, 340, 342, .343, 491, 492,
493, 494, 496, 497, 576, 672, 674, 676, 678
iUyricus (Tiwlites), 338, 342, 492, 493, 494, 495,
501, 576, 648, 676
cf . ( Tiwlites ) , 360
immaturus (Meguphyllitcs) , 354, 392, .396
(Procarnites), 325, .355, 358, 392, 396,
.397
impolitus (Tiwlites), 328, .346, 347, 492, 498,
501
inceitus (Danubifes [Dantdntes]), 354, 532
(Danubites [Danubites ?]), 5.35
indoaustralica (Leiophyllites), 350, 351
infldtus (Danubites [Preflorianitcs]), 354, 532,
535
htjucundus ( Tiwlites), 344, 492
inopinatus (HcUenites), .3.37, .3.55, 515, 516
? (HcUenites), 354, 516
inostranzeffi (Ceratites), 356, 420
(Czekanowskitcs), 420
insignis (Neocolund)ites) , .355, 516
insuhiris (Prosphingites), 325, 351, 354, .355, 404,
405, 407, 408
aff. (Prosphingites), 408
intermedins (Dinarites), 356, 489
( Dinarites [Oletiekites] ), 489
(Pwfloiianitcs), 325, .358, 379, 382, 422
intermontanum (Pscudosageceras) , 341, 361 ,362
involutus (Arnautoceltitcs), 325, 353, 397, 401,
402
(Prosphingites), 352, 404, 405, 407
(Paranannites), 352, 399
Inyoitcs, 381
eiekitensis, 357, 373, 374, 483
garbinus, ,341, 381, 580
Isculites originis. [see also Iscullitcs], 338, 411,
413
Isculitoides, .326, 332, 3.35, 344, .353, 411, 414,
419, 420, 422, 461, 5.52, .598, 612
antiglobulus, 413, 416
cllipticus, .326, 352, 353, 411, 414, 418,
419
globosus, .352, 404, 407
globulus, 4\:], 416
globulusorigiuis, 411
hammondi, .326, 360, 411, 413, 414, 418,
419. 012
minor, 326, 358, 411,418
originis, 326, 334, .3.35, 340, .350, 401, 411,
413, 414, 415, 416, 418, 419, 550
(aff.), 352, 418
(cf.), 347, 411
Ammonoids of the Late Scythian (Lower Triassic) • Ktimmel
549
sp., 352
sp. indet., 348
suboviformis, 326, 355, 411 413, 414, 418,
419
wasserbcrgi, 326, 358, 411, 413, 414, 418,
419, 598
Iscultitcs atitiglobulits, [see also Isculites], 340, 413,
417
var., 340
globulus, 340, 413, 415
var., .340
globuhisantiglobiihis, 340, 413, 417
globitlusoiiginis, 340, 413, 417
originis, 340, 415
iwanovi [Parussuria) , 448
jacksoni ( Meekoccras ) , 468
(Nordophiceras), 337, 360, 466, 467, 468,
469, 470, 471, 634, 636
(Ophiceras), 359, 468
? r Ophiceras"), 358
(Frionohbus), 466
Japonites arthabcri, 517
decipiens, 517
meridiamis, 350
raphaclis zojae, 350
surgrim, 338, 517
ugra, .350, 351
kansa {Danubitcs), 350
karatauciki {Procohtmbites) , .345
karutauciktis (Prucohnnbites), 326, 346, 347, 437,
441, 443
karazini (Dieneroceras), 325, 355, 367, 368, 369
karpiii.skii (Meekoceras), 356
(Meekoceras [Gyronites] ), 465
(Nordophiceras), 357, 465, 466
(Xcnodiscus), 452, 465
Kashmirites, 437, 490
aciitangidatus, 490
blaschkei, 490
contortus, 346, 490
subdimorphus, 345, 346, 490
kastriotae (Hedcnstroemia), 338, 448, 449
(Metahedenstroemia), 326, 340, 449, 568
kcirensis (Celtites), 338, 340, 367, 368, 375, 548
(Xcnodiscus), 367
kerneri (Tirolites), 343, 497
ketjserlingi ( Boreomeekoceras ) ,
(Meekoceras), All, All
Keijserlingites, 321, 323, 327, 332, 337, 349, 355,
451, 453, 461, 485, 486, 487, 488, 592, 614,
616, 632
angustecosiatus, 350, 351, 485
bearlakensis, 327, 360, 485, 486, 488,
614, 616
bearriverensis, 327, 360, 485, 486, 487,
327, 356, 476,
614
meridiamis, 355
— middcndorffi, 327, 356, 484, 485, 486
487, 488
nikitini, 356, 486
pagoda, 350
pahari, 350
sclucnki, 486
— sp., 349
— sp. indet., 347, 358, 488
— stephcnsoni, 337, 360, 485, 487, 632
— subrobw^us, 327, 356, .357, 358, 484, 485,
486, 487, 488, 592
(cf.),485
(n. sp. cf.),486
khanikoffi (Bcyrichites), 349
khelaliensis (Dieneroceras), 357, 369
kingi (Monophyllites), 338, 349, 523, 524, 525,
526, 586
(Monophyllites [Usstirites]), 524
(Ussu rites), .350
kingianus (Anasibirites), 483
Kiparisovites carinatus, 346, 519, 520
kittli (Dalmatites), 337, .343, 360, 519, 522, 523,
524, 650
knechti (Dieneroceras), 368, 360
knight i ( Tirolites), .342, 501, 682
koeneni (Owenites), 354
kokeni (Parapopanoceras) , ,391
(Procarnites), 325, 3.34, 3.35, .340, .341,
344, 347, ,348, 350, 353, .373, 390, 391, 392,
393, 394, 395, 396, 397, 460, 562, 564, 566
var. (Procarnites) , .341
var. evoluta (Procarnites), .341, 391, .394,
.396
var. panteleimonensis (Procarnites), .341,
391, .394, 396
komanum (Dagnoceras) , 338, 457, 478, 480, 481,
482, 576
Koninckites, 384, 389
bernoullii, .341, 388, 389
var., .341
postcrius, .357, 384
septentrionalis; 384
timorensis, 341
kraffti (Proptychites), 338, 385, 387, 556
(Proptijchitoides), 385, 387
ktenasi (Proptychites), 341, .388, .389
kummeli (Procarnites), .356, 387, .391
(Proptychitoides), 325, .356, 384, .385,
391
kwangsianus (Cordillerites) , 364, 365, 366
(Prenkites), [see also kioangsiensis], 352,
443, 444
(Prosphingites), 404, 405, 407
(Subcolumbites), 352, 431, 4.33
bungei, 485
dieneri, .348, 349, 350, .351, 485
kwangsiensis (Leiophyllites), 352, 535
(Prenkites) [see also kwangsianus], .352
Kymatites svilajamis, 343, 475
laevis (Dinarites), .342, 356, 506, 508, 656, 658
(Leiophyllites), 350
550 Bulletin Museum of Comporative Zoology, Vol. 137, No. 3
kiiiccolatus (Doricranitcs), 346, 505
LaiKcalite.s, 327, 343, 344, 450
bicaiinaiiis, 450
compactus, 450
discoidalis, 327, 343, 450
huniciis (Lccanitcs), 348, 376
{Xeuodisciis), 376
latifimhrkita sp. ind. aff. {ProptychUcs), 385
latifimhriatiis (Proptychitcs), 338, 384, 385, 387,
556
laiihhata (Panissuria), 326, 352, 353, 448
lafilohatum (Damwccrm), 327, 352, 353, 458,
460
laurac {Beyiichites), 328, 340, 341, 516, 517
lawicuciaini.s {Pioiitycliites) , 317
nuit. pustindica {PiuptycJiitcs}, 341, 385,
387, 388
Lecaniics, 578
discus, 338, 341, 374
fishtae, 338, 339
liupieus, 348, 376
niuzi, 338, 339, 578
— planorbis, 348, 468
■ skutarcusis, 338, 341, 368, 580
sp., 352
■ — • ? .sj)itzbei<j.c'usis, 450, 452
LeiophylUtes, 328, 332, 335, 351, 531, 533, 535,
584
adinaris, 328, 355, 535
• confucii, 350
iiidoaustralica, 350, 351
kuan^sicnsis, 352, 535
laevis, 350
loloiicihsis, 352, 535
DKiiitinuis, 328, 355, 535
itiiddlcinissi, 350
oxyuDtus, 532, 535
pildinalui, 350
(aff.), 5.35
])i(idyuinna, 350
pracnuittini.s, 354, 355, 531, 532, 533
radians, 328, 346, .347, 532, 535
.scpniliiius, 328, 352, 353, 532, 535
sp. iiulct., .347, .358, 5.35
variabilis, 328, 3.34, 335, 340, 355, 531,
533, 5.34, 535, 584
vcriforiuis, 352, 535
Iciaiimii il^a^itoccras), .3.38, 4.57, 4.58, 4.59, 460,
.570
hiiliriilaris ( Tmmlaiiitcs) , .326, .3.52, .353, 422,
423, 424
levaiiliniis {Coliinibilcs), .340, 44.3, 444
liatsikasi (Dinaritcs), 328, 340, 341, 344, 499,
506, 507, 510
Viccanum {Diaidorocrras) , 328, 343, 503, 504
liccaiiiis {Ccralitcs}, 503
{Dinaritcs), 503
( Dinaritcs [Liccaitc.s] ), 342, 503
li^aliforniis ( C!()Iund)itcs) , 345
ligatus (Columbitcs), 360, 425, 427, 428, 620,
624, 626
lihngen.tis {Sty rites), 338, 422, 423, 424, 580
{"Styrites"), 423
(cf. Sty rites) , 340
? cf. {Styrites), 423
lohiohse {Meekoceras [Submcckoceras]), 466, 475
{Submcckoceras) , .352
lohuensis {Digitophyllitcs) , .352, 392, .397
{LciophyUitcs), 352, 535
{Procarnites), 325. .353, 392, 397
{Prosphingites), 325, 352, .353, 404, 407
longilobatum {Pscudosageceras) , .354, 356, 357,
361, 363
\ar. kwangsiensc {Pscudosageceras) , 363
longiseptatuiu {Meekoceras [Subnicckoccra.'i]), 475,
467
{Su1?niCckoccras), .352
n}agiunn])ilicatus {Prospliingites), 405, 408
inahojuedis {Meekoceras) [see also mohamcdis],
338, 386, 388, 389, 560
{Proptychitcs), .388
{Proptychitoides), 359, 388
malayanus {Columbitcs) , .340, 443, 444
var. {Columbitcs), .340
var. crassa {Columbitcs), 340, 443, 444
malici {Bittnerites), 504, 505, 654
{Tirolitcs [Biltneritcs]), .343, 504
mahoren.sis {Prenkites), 326, 334, 335, 338, 340,
408,441,443,554
maugyshlakensis {Olcuekitcs) , 327, 346, 347, 441,
489
mansficldi {Ussurites), 337, 360, 528, 529, 530,
531, 628, 630
marginalis {As))iditcs), 338. 3.39, 572
nuiritimus { Danubitcs [Prcflorianites]), .354, 531,
535
aff. {Danubitcs [Prcflorianites]), ,5.35
{LeiophylUtes), .328, .3.55, 535
{Prefhnianiles), 535
inastykensis {Parinyoites) , 373
mekelvci {Prohungarites), 328, .348, 358, .360,
519, 520,521,522, 610
mcdilcrranea {Dieneroceras), 325, 340, 367, 368,
.548, ,578
{Xcnaspi.s), .3.38, 367, ,548
mcditcrrancus {Arncnitoceltitcs), 325, .340. 397,
400, 401,402, .552, 5,54
{ Parammnitc.s) , 338, ,341, 397, .399, 400
\ ai\ niedia { Parananniles) , 341, .397, .399,
400, 401
Medlicottiaeeae, .360
mcdius ( cf . Nannites), .341
Meekoceras, 359
caprilcnse, .34.3
? eontrarium, 46.5
conlrarius, 465, 467
curticoslatum, 3.59, 470, 471, 473, 475, 640
cuom]>h(dum, 165
gracililatis ( ci. ), 341
Ammonoids of the Late Scythian (Lower Triassic) • Ktimmel 551
Iiakki, 338, 385, 387, 564
jacksoni, 468
kurpinskii, 356
kei/serlingi, 471, 477
mahomedis, 338, 386, 388, 389, 560
micromphalus, 360, 470, 471, 473, 475,
640
pilatum, 360, 470, 471, 473, 475, 638
rodiosuin, 338, 463, 464, 576
roiundatum, 356, 476
sanctorum, 360, 470, 471, 473, 475, 638
sibiricum, 356, 453
? sihirictim, 453
skodrensc, 338. 339
sp. indet., 352, 410, 452, 453, 492
timarensis, 384
( Gyronites ) karpiiukii, 465
piano rh is, 468
schmidti, 452
(Koninckites) hasserti, 478
sibiricum, 453
(Submeekoceras) compressum, 466, 475
lolouense, 466, 475
longiseptaium, 467, 475
Meekoccratidac, 327, 337, 450
Megaphyllites atlasoviensis, 355
cvohitus, 350
immaturus, 354, 392, 396
menensis {Arctotirolites), 327, 356, 477
(Pseudotirolites), 356, 477
mercurii (Tirolitcs), 342, 492, 494, 674
meridianus (Japonites), 350
( Kei/serlingites) , 355 ^
Meropella, 326, 329, 447, 580
plejanae, 326, 340, 443, 447, 580
( cf. ), 347, 447
Metadagnoceras, 321, 327, 332, 335, 337, 457,
458, 459, 460, 461, 463, 464, 572, 576, 588,
594
freemani, 327, 350, 461, 462, 463, 464,
588
pulcher, 327, 358, 460, 461, 462, 463
sp., 462
tcrbunicum, 327, 340, 455, 461, 462, 463,
464, 572, 576
tobini, 327, 358, 461, 462, 463, 594
unicum, 337, 354, 355, 461, 462
Metahedenstroemia, 326, 363, 448, 449, 568
kastriotae, 326, 340, 448, 449, 568
micromphalus (Meekoceras), 360, 470, 471, 473,
475
middendorffi (Ammonites), 486
iCeratites), 356, 485, 486
(Keyserlingites), 327, 356, 484, 485, 486,
487, 488
middlcmissii (Hungarites), 520
cf. (Hungarites), 349, 517, 518, 519
( Leiophyllites) , 350
(Prohungarites), 320, 328, 348, 518, 520,
521, 590
minimus (Columbites), 360, 425, 427, 429
jyiinor (IsculHoides), 326, 358, 411, 418
(Paranannites), 354, 401
minutus (Paranannites), 352, 399
mirditensis (Columbites), 338, 340, 429, 431, 433,
444, 544, 546
var. (Columbites), 433, 546
(Subcolumbites), 429, 431, 433
mistardisi (Proptychites), 341, 388, 389
mitzopauloi (Chioceras), 326, 334, 335, 340, 443,
445, 446
var. meridionalis (Chioceras), 340, 446
mioctus (Parasibirites), 356, 483
modestus (Procarnites), 358, 396, 397
mohamedanus (Dinarites), 511
cfr. (Dinarites), 511
(Dinarites [Hercegovites] ), 432, 511
(Pseudodinarites), 328, 343, 511, 654,
664
mohamedis var. applanata (Proptychites) [see also
mahomedis], 341, 387, 388, 389
mojsisovicsi (Gyronites), 451, 452
mongolica (Psilosiura), 350, .351
Monocanthites, 326, 329, 422
monoceras, 326, 358, 382, 411, 422
monoceras (Monocanthites), 326, 358, 382, 411,
422
Monophyllites dieneri, 338, 524, 586
( .sp. ind. ex aff. ) , 349
( nov. sp. ex aff. ) , 526
hara, 338, 349, 524, 525, 584
kingi, 338, 349, 523, 524, 525, 526, 586
nopcsai, 338, 523, 524, 584
pitamaha, 338, 531, 532, 533, 584
sichoticus, 528
suessi, 531
(Leiophyllites) dieneri var. involuta, 341,
388, 390
georgalasi, 341, 531, 533, 534
palaeotriadicus, 341, 531, 533, 535
pitamaha, 531
(aff.), 341, 531, 533, 534
praeconfucii, 341, 531, 533, 534
rosae, 341, 524, 526
Palaeophyllites) praekieperti, 341 527
thalmanni, 341, 527
526
(Schizophyllites) betilhni, 341, 523, 524,
var. evoluta, 341, 524, 526
-^ pseudohara, 341, 388, 390
( Ussiirites) hara, 524
kingi, 524
monopytchus (Hololobus), 328, 343, 499, 511,
680
(Tirolitcs [Hololobus]), 342, 511, 680
montpelierensis (Prefloriantes) , 337, 360, 379,
381, 382, 626, 628
morlaccus (Dalmatites), 328, 343, 519, 522, 523,
524, 652
morpheas (Columbites), 356, 425, 501
552 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
( TiwUtes), 328, 356, 501
muchianus (Ceratites), 506
(Dimirites), 342, 343, 506, 507, 508, 658
multicostutus {Dinaritcs), 342, 507, 508, 660
multiformis (Siihcolumbites), 326, 335, 354, 355,
433, 435, 436
multilobatum (Pseudosa^eccra-s), 323, 325, 337,
340, 345, 346, 347, 348, 355, 356, 357, 358,
359, 360, 361, 362, 363, 608
var. ^ifiantcum {Pscudo.saiJ^cccras}, 361
ct. ( P.scudosap.eceras) , 361
multiplicatu.s (CcltHcs), 437, 440, 490, 594
(Ceratites), 356, 382
rCeratite.i'), 382
(Prcflorionitcs), 325, 356, 379, 382
(Psetidoccltites), 440, 594
(Xenoceltites), 382
(Xenodiscus), 382
multispinatus (Tirolites), 343, 493, 666
musacchi (Arianites), 326, 338, 340, 443, 446,
544
muthianus { As])idites), 383
{Psvud(isi>idUes), 383
nangaensis cfr. (Ophiceras), 338, 379, 381, 548,
578
sp. iiid. aff. (Xenodiscus), 379
Nantiites haiarunusi, 346, 401
herherti, 399, 400, 401, 552
cf. Nanuitcs Iiindustauus, 341
medius, 341
Neocolumbites, 337
iiisignis, 355, 516
ncvadi (Pseudoceltitcs) , 326, 359, 424, 438, 440,
608
niazi (Lccanites), 338, 339, 578
nik(d)itensis (Dieneroccras) , 357, 369, 483
nikitini (Ceratites), 356, 484, 486
(Keijserliiigifes), 356, 486
nivalui aii. (Auakashuiirites), 352
(DanuhUes), 437, 490
m)dosum (Chiorera.s), 326, 340, 443, 446
nopcsai (Eofiht/llites), 525
(Mnuopluillites), 338, 522, 524, 584
? (Proplyehitoides), 388, 389, 560
iio))r.sai]um ( Dagiioceras) , 327, 338, 340, 457,
458, 459, 460, 463, 464, 570, 572
\ar. inrolula (Danmneras), 341, 464, 465
Nordophiceras, 321, 327, 337, 353, 355, 357, 452,
465, 467, 468, 475, 477, 632, 634, 636, 642
alcxeevae, 337, 357, 452, 466, 467, 468,
469, 471
compressum, 327, 353, 475
conlrarius, 357, 466
coutriilius, 465
euomphalus, 337, 357, 465, 466, 467,
468, 469, 471, 475, 634
jacks(mi, 337, 360, 466, 467, 468, 469,
— pilatum, 337, 357, 360, 468, 469, 470, 471,
473, 475, 632, 638, 640, 642
— plamvhis, 322, 327, 348, 467, 468, 475
ct. phinorhis, 348
— pseudosimplex, 327, 356, 468, 475
Schmidt i, 356, 452
Noritaceac, 375
Noritidae, 327, 477
normalis (Ceratites), 317
midus (Dimirites), 341, 342, 344, 506, 507, 508,
656
ohliqueplicatus (Proptychites), 338, 388, 389, 560
occidentalis (Popovites), 326, 358, 382, 411, 421,
422
oJenckensis (Nordophiceras), 357, 467, 468
Olem'kites, 323, 327, 332, 335, 337, 344, 358,
440, 488, 489, 612
altus, 356, 489
camidensis, 327, 357, 489
glacialis, 356. 489
maugysldakensis, 327, 346, 347, 441, 489
.sonticus, 355
spiniplicatu.s; 327, 356, 488, 489, 490
spiuiplicatus (cf.), 360, 489, 612
tururpcnsis, 346
Ophiceras, 345
demi.ssum ( l-L), 341, 345
dieneri, 367
jacksoni. 359, 468
mmgaensis (dr.), 338, 379, 381, 548, 578
sakuntala, 338, 367, 578
sakuntala ( cf . ) , 367
spencei, 359, 376
"Ophiceras"? jacksoni, 358
? spencei, 358
orhicidatus (Suhdoricranites), 346, 506
orientalis (Cordillcrites), 352, 364, 365, 366
(Eophyllites), 328, 350, 523, 526
(Pronorites), 341, 477, 478, 482
(Pro.s))hingites), 404
originis ( Isculiles), 338, 411, 413
(Isculitoides), 326, 334, 335, 340, 350,
401, 411, 413, 414, 415, 416, 418, 419, 550,
552
aff. (Isculitoides), 352, 418
cf. (Isculitoides), .347, 411
(/.sr(//f/7r.s),340, 415
470,471,6.34,636
— karpin.skii, 357, 465, 466
— olenekensis, 357, 467, 468
ornatus (Cohimhitcs), .360, 425, 427, 428, 429,
620
? aff. ( Cotui)d)ites), 356, 425
osmanicus ( Alhanites) , 477, 479, 480, 481
(Pronorites), 338, 477, 478, 479, 482, 574
cf. { Pronorites), 341, 477, 478, 482
Otoceras uoodirardi, .317
Otoceralacae, 367
ovalis (Prosphingites) , 404, 407
ovatus (Doricraniles), 346, 505
Owcnites, .338
koeneni, 354
r).v|/(i(..s7;/.v ( Procarnites) , 352, 391, 392, 395, 396
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 553
oxynotus (Leiophyllites), 532, 535
pagoda (Keyserlingitcs), 350
pahari {KeyscrUngitcs) , 350
PaJacophyUitcs, 328, 527
sicinmanni, 328, 340, 349, 350, 527
pdlucotiiadiciis (MonophyUitcs [Lcivphyllites]),
341,531,533,535
pandya (Sibiritcs), 350
Paradinarites, 326, 329, 437
suni, 326, 352, 353, 424, 437
paradiscus (Hcmilccanitcs), 325, 358, 368, 375,
598, 602, 610
Paragocems dukagiui, 338, 397, 399, 400, 401, 554
cf. Paragoccras dukagini, 341
Paranannites, 400
aspenensLs, 345
var. euiopaea, 341, 397, 400, 401
chionemis [see also chiosensis], 341, 397,
399, 400, 401
cliioscihsis [see also chionensis], 397
compressus, 341, 397, 399, 400, 401
gracilis, 354, 399, 401
involutus, 352, 399
mediterraneus, 338, 341, 397, 399, 400
var. media, 341, 397, 399, 400,
353, 427, 429, 431, 433, 434, 435, 436, 542,
544, 546, 548
cf. {Siibcohimhites), 429, 431
401
minor, 354, 401
minutiis, .352, 399
subglobosiis, 352, 407
suboviformis, 354, 418
suboviformis ( aff. ), 418
Paranannitidac, 325, 337, 397
Paranoritidae, 337, 383
Parapopanoceras kokeni, 391
Parasageceras, 35 1
Parasibirites grambergi, 356, 483
mixtus, 356, 483
rariaculcattis, 356
suprciiosiis, 483
Parimjoites mastykcims, 373
pamianus (Cohimbifes), 322, 337, 340, 346, 354,
355, 360, 424, 425, 427, 428, 430, 4.34, 435,
462, 483, 501, 618, 620, 622, 624, 626
cf. (Cohnnbites), .345
pariscnse (Svalbardiccras) , 354, 355
Parussuria, 326, 329, 448
iicanovi, 448
latilobata, 326, 352, 353, 448
pasquayi (Pseudosageceras), 325, 340, 360, 363,
364
(Pseudosageceras [Metasageceras]), 341,
364
paucispinatus (Tirolites), 342, 492, 494, 495, 674,
676
peali (Tirolites), 342, 501, 682
pcrcosiattis ( Tirolites), 343, 493, 499, 666
perrinismithi (Cohnnbites), 338, 340, 427, 431,
433, 546
(Subcolumbites) , 326, 334, 335, 340, 347,
Pliylloceratacae, 524
Phyllocerida, 524
pilutum (Meekoceras), 360, 470, 471, 473, 475,
638
(Nordophiceras), 3.37, 357, 360, 468, 469,
470, 471, 473, 475, 632, 6.38, 640, 642
Pinacoceratacae, 517
pitamaha (Leiophyllites), .350
aff. (Leiophyllites), 535
(MonophyUites), 338, 531, 532, 533, 584
(Mom^phyllites [Leiophyllites]), 531
aff. (MonophyUites [Leiopliyllites]), 341,
531, 533, 534
pityoussae (Hedenstroemia) , .341, 449
lilanorbis ( Lecanites) , 348, 468
( Meekoceras [Gyronites]) , 468
(Nordophiceras), 322, 327, 348, 467, 468,
475
cf . ( Nordophiceras ) , 348
phmovohis (CeJtites), 360, 369, 372, 373, 646
plejanae (Arianites [MeropeUa]), 341, 447
(MeropeUa), 326, 340, 443, 447, 580
cf. ( MeropeUa), 347, 447
plieatuhis (Cohnnbites), 444
pUeatuU ex aff. (Cohnnbites), 340, 443, 444
Plococeras, 507
dahnatinmn, 506
poi)ovi ? (Prohtingarites), 354, .355, 518
(Pseudaspidites), 337, 360, 383, 384, 628,
650
Popovites, .326, 421
borealis, 326, 357, 421, 422
oceidentahs, 326, 358, 382, 411, 421, 422
posterius (Koniiiekites), 357, 384
(Pseudaspidites), 3.37, 357, 383, 384
pradyumna (LeiophyUites) , .350
praeconfucii (MonophyUites [LeiophyUites]), 341,
531, 533, 534
praekieperti (MonophyUites [PalaeophyUites]), 341,
527
praematurus (Beyrichites) , 341, 516
(HeUenites), 328, 3.34, 3.35, 340, 352, 353,
512, 513, 514, 515, 554
cf. ( HeUenites), 513, 514, 515
var. aegaeica (HeUenites), 340, 512, 513,
514
(LeiophyUites), 354, .355, 531, 532, 533
( Tropieehites), 511, 512
? ( Tropieehites), 338, 512
? var. (Tropieehites), 512
prahJada (Sibiritcs), 350
Prefhmanites, 325, 3.37, 379, 381, 382, 535, 548,
578, 580, 626, 628
garbinus, .325, 340, 379, 381, 580
intermedius, 325, 358, .379, 382, 422
maritimus, 535
554 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
— monipelierensis, 337, 360, 379, 381, 382,
626, 628
— muUiphcutus, 325, 356, 379, 382
— stron^i, 381
sulioticu.s, 325, 340, 379, 391, 548, 578
Prcnkites; 326, 408, 441, 460, 554
hdenae, 326, 340, 441, 443
kwangsianus, 352, 443, 444
mahorensis, 326, 334, 335, 338, 340, 408,
441, 443,554
simdaicu.s; 340, 350, 408, 409
timoremis, 321. 326, 340, 350, 353, 441,
443, 444
( aff . ) , 354, 355, 444
^'Preukitcs" sinulaicus, 408, 409
prctioso ind. aff. (Sibiiitcs) , 483
pretiosus (Sihiritcs), 356, 483
primoriciisis { Arctohiingaritcs) , 355
Prionitidae, 337, 482
prioiwidcs (Stacheitcs), 327, 342, 343, 344, 345,
401, 455, 456, 463, 652
Prionolohiis jacksoni, 466
.schiuidti, 452
prior {Ceratites [Paraccratitcs]), 342, 497, 680
(Tirolites), 497
( Tirolitoidcs), 492, 497, 498, 680
( Xenodisciis ) , 497
Proavites disctis, 374
skutarensis, 368
Procarnites, 325, 332, 335, 391, 396, .397, 418, 562,
564
uciitus, 352, 391, 395, 396
andrusovi, 345, 346, .391, 392, 395, 396
immaturus, 325, .355, .358, 392, 396, 397
kukcni, 325, 334, 335, 338, .340, 341, .344,
347, 348, 350, .353, 373, 390, 391, 392, 393
394, 395, 396, 397, 460, 562, 564, 566
var., 341
var. evoluta, .341, 391, 394, 396
var. panteleimonensis, 341, 391,
394, .396
ktimmeli, 356, 387, .391
lolouensis, 325, 353, 392, 397
modest us, .358, 396, 397
oxijnostus, .352, 391, 392, .395, 396
skandcrhegis, 338, 341, .391, .392, .393, .394,
395, 396, 562
Procladiscitcs, 349
ijasoda, .350, 351
ProcoJumhitcs, 321, 326, 329, 337, 441
karutauciki, 345
karataucikus, 326, ,346, .347, 437, 441, 443
sp., 355
progresses ( Dinaritcs), 343, 504
( Dinarites [Liccaitcs] ), ,342, 504, 664
Prohungaritcs, 323, .328, 3.32, 3.35, 338, ,348, 360,
415, 463, 517, .520, 521, ,522, ,590, 610, 612,
616
carinatus, 328, 344, 347, 520
crasseplicatus, 328, .350, 517, 518, 520, ,521
612
( cf. ), ,348, ,373, 518, ,521
gutstadti, 328, .360, 518, 519, 521, 522,
— mckelvei, 328, ,348, ,358, 360, 519, 520,
521, 522, 610
— middlcmissii, 328, 348, 517, 518, 520, 521,
.590
? popovi, 354, ,355, 519
.v/;»//;s, 517, 518
( n. sp. cf. ) , 520
sp., 344
sp. indet., .3,58, 360, 519, 522, 612, 616
tuherculatus, 328, .350, .357, 518
Prolecanitida, 360
Pronorites arhani (.spec. ind. c\ aff.), 349, 477,
478, 479
arhamts, .338, ,341, ,349, 477, 478, 479,
482, .574
482
— var., ,341
— var. mcditcrranca, 341, 477, 478,
var. sundaica, All
oricntalis, 341, 477, 478, 482
osmanicus, 338, 477, 478, 479, 482, 574
(cf.), ,341, 477, 478, 482
rcicheli, ,341, 478, 481, 482
schauhi, 341, 478, 479, 482
var. kephaJovunensis, ,341, 478,
482
var. timorensis, 478
triadiciis, 338, .341, 477, 478, 479, 482,
574
Propti/chites, 384, 385, 389
(irdmhcri, 341, 349, 387, 388, 389, .390
hcdciinicus, 341, 385, 387, 388
hcrtisci, 338, 386, 388, 389, 558
hiixtorfi, ,341, 386, 388, 389
hakki, 387
kraffti, 338, 385, 387, 556
ktcnasi, 341, 388, ,389
latifimbriata (sp. ind. aff.), 385
latifimhriatus, 338, 384, 385, 387, 556
lawrencianus, 317
nuit. post ind ica, ,341, 385, 387, 388
niahomcdis, 388
var. upplanata, 341, 387, 388, 389
mistardisi, 341, 388, 389
oidUpn'phcatits, 3,38. 388, 389, 560
trigonalis, 338, 388, 389, 558
trilohatus, 317
Proptychitidac, 325, 384
Proptycluioidcs, 325, 332, 384, 385, 388, 389, 418,
423, 562, 564, 590
arthahcri, 325, 3,50, 384, 385, 390, 391,
590
hcrtisci, 388, 389
comprcsstts, 352, 390
decipiens, 325, ,340, 384, 385, 387, 388,
389, .390, 556, 564
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 555
— hakki, 385, 387
— kraffti, 385, 387
— kiimmcli, 325, 356, 384, 385, 391
inahomedis, 359
— ? nopcsai, 388, 389, 560
— ? simplex, 352, 390
— trigonalis, 325, 340, 384, 385, 386, 388,
389, 390, 391, 558, 560, 562
tunglanensis, 325, 352, 353, 384, 385, 390
ProsphiuRites, 325, 332, 335, 403, 404, 405, 407,
408, 409, 410, 420, 421, 446, 580, 590
all, 325, 338, 340, 405, 407, 580
austini, 403, 404, 405, 407
coomhsi, 325, 351, 404, 405, 407, 408
czekanowskii, 325, 356, 403, 404, 405,
407, 408, 592
( cf . ) , 358, 405
(ex aff. ), 340, 407, 410
globosus, 325, 354, 355, 404, 405, 407,
— (aff.), 407
408
— insularis, 325, 351, 354, 355, 404, 405,
407, 408
(aff.), 408
— involuttts, 352, 404, 405, 407
— kwaugsianus, 404, 405, 407
— hloticnsis, 325, 352, 353, 404, 407
magnumhilicatus, 405, 408
orientalis, 404
ovtilis, 404, 407
radians, 404
— sii\ensis, 404
— slossi, 404, 407
spathi, 403, 404, 407
— suhglohosus, 325, 353, 407
— vonderschmitti, 404, 405, 407, 410, 411
— (Chiotitcs) globidaris, 340, 419, 420
superglohosus, 340, 420
{Zcnoiies) helenac, 340, 410
var. viaradoviinensis, 340,
410
Pwptwpites, 326, 329, 334, 420, 444, 445, 568
hilmi, 326, 335, 338, 340, 443, 444, 445,
446, 568
Pscudaniiolites, 512
idahoense, 516
Pseudaspidites, 337, 355, 357, 383, 384, 628, 650
imitliianus, 383
popovi, 337, 360, 383, 384, 628, 650
posierius, 337, 357, 383, 384
whceleri, 383
yudishthira, 383
Pscudluiipoceras, 512
idahoeme, 319, 360, 512, 516
spiniger, 319, 348, 512
Pseudoceltites, 321, 326, 337, 359, 425, 437, 490,
594, 608, 628
cheneyi, 337, 359, 360, 438, 440, 628
constriciilis, 424
dolnapaensis, 326, 347, 359, 438, 440, 441
midtiplicattts, 440, 594
nevadi, 326, 359, 424, 438, 440, 608
Pseudodinarites, 328, 343, 511, 654, 664
mohamcdanus, 328, 343, 511, 654, 664
pseudohara (MonophyUites [? Schizophyllites]),
341, 388, 390
Pseudokymatites, 327, 343, 475, 664
svilajamts, 327, 343, 475, 499, 664
pseudortisseUi (Flcmingiies), 341, 387, 388, 389,
390
Pseudosageceras, 325, 332, 337, 345, 360, 366,
564, 582, 608
albanicum, 325, 334, 340, 360, 363, 364,
532
bicarinatum, 358, 364, 366
clavisellatum, 363
( cf. ), 341, 361, 363
compressus, 363
curvatum, 363
drincnsc, 325, 338, 340, 341, 360, 361,
362, 363, 564
var. incentrolata, 341
intcnuontaiutiii, 341, 361, 362
— longilobutum, 354, 356, 357, 361, 363
var. kwangsiense, 363
— midtihbatum, 323, 325, 337, 340, 345,
346, 347, 348, 355, 356, 357, 358, 359, 360,
361,362, 363,608
(cf.),361
var. gigantciim, 361
pasquayi, 325, 340, 360, 363, 364
schamarense, 361, 362
simplex, 325, 354, 355, 360, 362, 364
sp., 359
tsotengense, 363
(Metosageceras) pasquayi, 341, 364
Pseudosibirites dichotomous (cfr. ), 338, 457, 478,
481, 482, 580
pseudosimplex (Noidophiceras), 327, 356, 468,
475
Pseiidotriolites menensis, 356, 477
Psilosturia, 349
mongolica, 350, 351
Ptychites, 349
riigifer, 317
pulcher (Metadagnoceras), 327, 358, 460, 461,
462, 463
qiiensiedti (Tiwlites), .342, 492, 494, 495, 676
radians (Leiophyllites), .328, 346, 347, 532, 535
(Prusphingites), 404
radiatus (Flemingites), 317
(Hellenites), 328, 340, .358, 513, 514, 515,
516
{Hellenites [Pallasites]), 341, 514
radiosum (Meekoceras), 338, 463, 464, 576
raphaelis zojae (Japonites), 350
rarecostatus (Dorieranites), 346, 505
rariaculeatus (Parasibirites), 356
556
Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
ravana (Ceratites), 349
raricostatus (Anasihirites), 356
rectangularis (TiroUtes), 338, 342, 493, 494, 495,
576, 674
refractiis (EophyUites), 524, 525, 584
cf. {EophijUitcs), 526, 527
reicheli (Pronorites), 341, 478, 481, 482
renzi (Sibirites), 327, 340, 483, 588
repuhus {TiroUtes), 342, 493, 494, 495, 674
reuttensis (Ammonites) , 516
richardsi ( Dalmatites ) , 343, 524
Rohustites suhrohustus, 485
robustus [Fengshanites), 352, 433, 435
(Subcohmibites), 326, 353, 435, 436
( TiroUtes), 342, 493, 494, 495, 678
Rommanites simionescui, 350
ropini ( Dabuatites), 344, 349, 350
rosae (MonophtjlUtes [LeiophyUite.^]), 341, 524,
526
rossictis ( Balatonites ) , 505
( Doricranites), 346, 505
(Dorikranites), 505
(TiroUtes), 328, 345, 346, 347, 492, 500,
501
rotiformis (TiroUtes), 342, 493, 494, 495, 676
rotundatum { Arciomeckoeeras ) , 327, 356, 476
( Boreomeekoceras ) , 476
(Meekoceras), 356, 476
riigifer (Pfychites), 317
Sageceras aUxinicum, 338, 363
var., 341
Sageceratidae, 325, 337, 360
.mkuntala (Ophiceras), 338, 367, 368, 578
cf. (Ophieeras), 367
sanctorum (Meekoceras), 360, 470, 471, 473, 475,
638
schamarense (Psendosageceras) , 361, 362
scharicus (Doricranites), 346, 505
schaubi (Pronorites), 341, 478, 479, 482
var. kep]ialovunensis {Pronorites), 341,
478, 482
var. tin}orensis (Pronorites), 478
schmidti ? (C^yronites), 452
{Meekoceras [Gyronites] ), 452
(Nordoi)}iieeras), 356, 452
{ Prionoldbtis), 452
(SvaUMirdieeras), 327, 356, 357, 451, 452,
453, 454, 592
(Xenodisrus), 356, 451, 452
.schrcnki (Ceratites), 356, 486
(KeyserUngites), 486
.seminudus (TiroUtes), 342, 492, 494, 495, 576,
672
var. nudior ( TiroUtes), 492, 672
var. pUcosus ( TiroUtes), 492, 672
.seplenlrionaUs (Koninekites), 384
seriienlintts (LeiopltylUtes) , 328, 352, 353, 532,
535
serralelobatus (TiroUtes), 342, 510, 662
sheldoni (Svalbardiceras), 337, 360, 451, 453, 454,
626
sibiricum (Meekoceras), 356, 453
? (Meekoceras), 453
(Meekoceras [Koninekites] ), 453
(SvaUmrdiceras), 327, 356, 453
sibiricus (Asj)idites), 453
(Hemiprionites), 356, 453
Sibirites, 327, 332, 483, 588
dichotomus ( cf. ), 478
eichwaldi, 327, 356, 483
(cf.),483
grambergi, 483
var. mixta, 483
var. rariacideatus, 483
pandya, 350
prahlada, 350
pretioso ( iiid. aff. ), 483
pretiosus, 356, 483
renzi, 327, 340, 483, 588
spiniger, 348
subpretiosus, 356, 483
superbus, 317
Sibiritidae, 327, 337, 483
sichoticus (MonophyUites), 528
sieveri (Ihsuritcs),' 328, 358, 528, 529, 530, 531,
602, 604, 610
sigmotoidcus (Ceratites), 356
simiUs (Prohungarites), 517, 518
II. -sp. cf. (Prohungarites), 520
simionescui (Ro)nmanites), 350
simplex ( Arctoccras), 356, 468
? (Proptychitoides), 352, 390
(Pseudosageceras), 325, 354, 355, 360,
362, 364
sinensis (Prosphingites), 404
sinuatus (Dinarites), 348, 376
(XenoeeUitcs), 322, 325, 348, 376
skandcrbegis (Procarnites), 338, 341, 391, 392,
393, 394, 395, 396, 562
.^kipetaren.sis ( EpOiedenstroemia ) , 364
(Hedcnstrocmia), 338, 364, 365, 366, 580
.skodrense (Meekoceras), 338, 339
.skutaren.sis (Dieneroccras) , 325, 340, 367, 368,
580
( Lecunites), 338, 341, 368, 580
( Proavites ) , 368
slo.ssi (Prosi)hingites), 404 ,407
smiriagini ( Ceratites), 496
( TiroUtes), 343, 346, 496, 499, 666
smilhi (TiroUtes), 337, 360, 501, 648
soUtus (SulxoUimbitcs), 354, 436
sonticus (Olenekites), 355
spathi (Pro.sphingites), 403, 404, 407
spencei (Columbites), 360, 425, 427, 428, 429,
434, 620, 624
var. chiotiea (CoUind)ites), 340, 434, 435
(Ophiceras), 359, 376
? ("Ophiceras"), 358
Ammonoids of the Late Scythian (Lower Triassic) • Ktimmel 557
(Xenoceltites), 337, 360, 376, 377, 378,
380, 636, 644
spini^er (Psctidhaipoceras) , 319, 348, 512
— (Sibirites), 348
spiniplicaius (Dinarites), 356, 488, 489
(Dinarites [Olenekitcs] ), 488
(Olcnekites), 327, 356, 488, 489, 490
cf. (Olcnekites), 360, 489, 612
spiiwsior ( Tiwlites), 343, 496, 499, 668
spinosiis (Tiwlites), 342, 346, 492, 493, 499, 501,
670
cf. ( Tiwlites), 359, 503
spitsbergetisLs (Xenoceltites), 325, 354, 355, 376,
377
cf. (Xenoceltites), 359, 440
? (Lecanites), 450, 452
(Svalbardiceras), 327, 357, 450, 451, 452,
454, 592
stachei ( Tiwlites), 342, 493, 494, 678
Stacheites, 320, 327, 344, 455, 456, 463, 464, 465,
614, 652
cUonijsi, 341, 455, 462, 463, 464, 465
var., 341
floweri, 327, 344, 358, 455, 456, 457, 596
prionoides, 327, 342, 343, 344, 346, 401,
455, 456, 463, 652
sp. indet., 344, 348, 360, 455, 456, 457,
614
webbianiis, 350, 455
stcinmanni (Pcdaeophiillites) , 328, 340, 349, 350,
527
stephensoni (Keyserlingites), 337, 360, 485, 487,
632
striatus (Hellenites [Pallasites]), 341, 514, 516
var. densicostata (Hellenites [Pallasites]),
341, 514, 516
strongi (Danubites), 379
(Preflorianites), 381
Sti/rites, 422
mangemis, 338, 422, 423, 424, 580
? lilangensis ( cf . ) , 423
cf. Styiites lilangensis, 340
"Stijrites" hlangensis, 423
Subcolumbites, 323, 326, 3.32, 3.35, .344, .354, .376,
409, 423, 427, 436, 437, 460, 461, 544, 546,
548
americanus, 326, 335, 358, 433, 435, 436,
437, 600, 602
anomalus, .354, 436
diismani, .326, .340, 431, 433, 4.34, 435,
436, 544
europaeus, 429, 431, 433
kwangsianus, 352, 431, 433
mirditensis, 429, 431, 433, 444
multiformis, 326, 335, 354, 355, 433, 435,
436
perrinismithi, 326, 334, 335, 340, 347, 353,
427, 429, 431, 433, 4.34, 4.35, 436, 542, 544,
546, 548
( cf . ) , 429, 431
robiistiis, .326, .353, 435, 436
solittts, 354, 436
sp., .358
subdimorphus ( Eiikashmirites), .327, .347, 490
(Kashmirites), .346, 490
Siibdoricranites discoides, 345, 346, 506
orbiculatus, .346, 506
subevolutus (Xenoceltites), 375, 376
subglobosits (Paranannites), 352, 407
("Prosphingites), .325, .353, 407
siibgracilis ( Anasibirites ) , 346, 448
(Epiceltites), 326, .347, 443, 448
subillyriciis (Tirolites), 342, 493, 494, 495, 678
Siibmeekoceras compresstim, 352
loloitense, 352
longiseptatum, 352
suboviformis (Isciditoides), .326, .355, 411, 413,
414, 418, 419
aff. (Paranannites), 418
(Paranannites), .354, 418
subpretiosus (Parasibirites), 483
(Sibirites), 356, 483
subrectangularis (Celtifes), 490
subrobustus (Ceratites), 356, 485
(Keyserlingites), .327, .356, .357, 358, 484,
485, 486, 487, 488, 592
cf. (Keyserlingites), 485
n. sp. cf . ( Keyserlingites ) , 486
(Robustites), 485
Subvishnuites, .325, 3.37, 357, 367, 373, .374, 483,
646
eiekitensis, 337, .357, 374
enveris, 325, .340, 373
( cf. ) , 347, 373
sp. indet., 347, 348, 360, 373, 374, 646
tientungensis, 373
ivelteri, 373
suessi (Monophyllites) , 531
sugriva (Japonites), 338, 517
sidioticus (Preflorianites), 325, 340, 379, 381,
548, 578
(Xenodiscus), 338, .341, 379
stindaicus (Prenkites), .340, .350, 408, 409
("Prenkites"), 408, 409
(Vickohlerites), .326, .340, 408, 409
cf. (Vickohlerites), 347, 409
siini (Paradinarites), .326, 352, 353, 424, 437
superbus (Stephanites), 317, 318, 319
superglobosus (Chiotites), 420
(Prosphingites [Chiotites]), 340, 420
Svalbardiceras, .321, .327. 332, 3.37, 355, 450, 451,
452, 453, 454, 592, 626
chowadei, 327, 358, 453
dentosus, 327, 356, 453
freboldi, 327, .357, 453
pariscnse, 354, 355
schmidti, 327, .356, .357, 451, 452, 453,
454, 592
sheldoni, 3.37, 360, 451, 453, 454, 626
sibiricum, 327, 356, 453
558 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
sp., 454
sp. indet., 348, 360, 451, 454
spitzher^emis, 327, 357, 450, 451, 452,
453, 454, 592
svilaiamis (Kymatites), 343, ■i'^^
{Pseudokymatites), 327, 343, 475, 499,
664
Svilaiites, 491, 499
cingulatus, 497
tahuhtus (Balkdnite.s), 327, 343, 465
tchermjschewicnsis {Hellcnites), 355
teicherti (Arnautoceltites), 325, 358, 397, 399,
402, 403, 602
telleri {Bittncrites), 505, 654
? ( Tiiolites [Bittnehtes]), 343, 504
terhunico aff. ( Dagnoceras ) , 463
terbunicum (Dagnoceras), 338, 341, 457, 463,
464, 465, 572
(Metadagnoceras), 327, 340, 455, 461,
462, 463, 464, 572, 576
thaliiumni {Monuphyllites [PalacophyUites]), 341,
527
Tlwrmalites, 345
tientungcmis {Subvishnitites), 373
tietzei (Tirolites [SviJajites]), 343, 497, 500, 680
timoremis (Koninckitcs), 341
(Mcekocera-s), 384
(Prenkites), 321, 326, 340, 350, 353,
441, 443, 444
aff. (Prenkites), 354, 355, 444
Tirolites, 328, 337, 342, .343, 344, 345, 346, .359,
425, 491, 492, 499, 501, 502, .507, .576, 608,
65()', 680, 682
cmgustilobaius, .342, 493, 499, 670
var. alpha, 493, 499, 670
angusttis, .342, 493, 494, 495, 678
astakluwi, 3.37, 360, 499, 502, 650
hispinatus, .343, 497
carniolicus, .342
cassianus, 317, 328, .342, .343, 345, 346,
491, 492, 493, .500, 501, 503, 666, 668, 670,
] (ef.), .359, 492, 498, 501, 503, 608
( ex gr. ), .356, .501
var. alpha, 493
var. tenuis, 493
. cingulatus, 328, 343, 497, 680
. dancini, .343, 493, 499, 666, 668
(cf.),492
var. abbrevians, 496
var. ductus, 496, 499
var. costatus, 496, 499
— var. modeslus, 496, 499
var. reminiscens, 496, 499
dimidiatu.s, .342, 493, 674
diuarus, .342, 493
. distans, 342, 492, 494, 495, 672, 676
. clegans, .346, 498, .500, 501
- gerbacnsis, .356, 501
- harti. 337, .342, .359, 492, 501, 682
haueri, .343, 493, 499, 670
(aff.), 359, 503
var. minor, 493, 499
heterophanus, 342, 493, 510, 674
hnbridus, .342, 493, 494, 676
_ idrianus, 328, 340, .342, 343, 491, 492,
493 494, 496, 497, 576, 672, 674, 676, 678
_ \lhiricus, 338, .342, 492, 493, 494, 495,
501,576,648,676
(cf.),360
impolitus, .328, 346, 347, 492, 498, 501
injucundus, 344, 492
kerneri, .343, 497
knight i, 342, 501, 682
mcrcurii, .342, 492, 494, 674
morpheos, 328, 356, 501
multispinatus, 343, 493, 666
paucispinatus, .342, 492, 494, 495, 674, 676
pealei, .342, 501, 682
percostatus, 343, 493, 499, 666
})rior, 497
quenstedti, 342, 492, 494, 495, 676
rectangularis, 338, 342, 493, 494, 495, 576,
674
repulsus, 342, 493, 494, 495, 674
robustus, 342, 493, 494, 495, 678
nmicus, 328, 345, 346, 347, 492, 500, 501
wtiformis, .342, 493, 494, 495, 676
. seminudus, 342, 492, 495, 576, 672
var. nudior, 492, 494, 672
var. plicosus, 492, 672
. serratelobatus, 342, 510, 662
- smiriagini, .343. .346, 496, 499, 666
-,s-m/7/i/, .3.37, .360, 501,648
- sp. indet., 348, 360, 499, 503, 650
- spinosior, 343, 496, 499, 668
- spinosus, .342, .346, 492, 493, 499, 501, 6/0
(cf.), 359, .503
- stachei, .342, 493, 494, 678
- subiUiiricus, 342, 493, 494, 495, 678
- toulai, .343, 497, 498, 499, 668
- turgidus, 343, 493, 666
- undulatus, .342, 493, 494, 495, 678
- (Bittnerites) bittneri, 34.3, 504
mali£i, 343, 504
? telleri, 343, 504
680
(Hololohus) monoptyehus, 342, 511, 680
(Svilaiites) cingulatus, .343, 497, 499, 500,
— tietzei, 343, 497, 500, 680
Tirolitidae, 3.37, 491
Tirolitoides prior, 492, 497. 498, 6S0
iirolitoides (Dinarites), .342, 507, 508, 660
tobini (Metadagnoceras), 327, 358, 461, 462,
463, 594
lolli (Dinarites), .356
toulai ( Tirolites), .343, 497, 498, 499, 668
triadicus (Albanites), 327, 3.34, 340, 347, .350,
441, 457, 477, 478, 479, 480, 481, 482, 572,
,574, 576, 580
Ammonoids of the Late Scythian (Lower Triassic) • Kummel
559
(Pronorites), 338, 341, 477, 478, 479,
482, 574
trigonalis (Proptychites), 338, 388, 389, 558
(Proptychitoidcs), 325, 340, 384, 385,
386, 388, 389, 390, 391, 558, 560, 562
trikkalinoi (Hellenites), 340, 512, 513, 514
var. {Hellenites), 341
var. graeca {Hellenites), 341, 512, 513,
514
trilobatiis {Proptychites), 317
trinodusus {Ceratites), 317
Tropiceltites, 511,512
praematurus, 511, 512
? praematurus, 338, 512
praematurus var., 512
tsotengense {Pseudosageceras) , 363
tubcrculatus {Hungarites), 349, 518, 519
{Prolmngarites), 328, 350, 357, 518
twnidosus {Doricranites), 346, 505
tunglanensis {Proptychitoides) , 325, 352, 353,
384, 385, 390
Tunglanites, 326, 422, 423, 580
alexi, 326, 340, 422, 423, 424, 580
Icnticularis, 326, 352, 353, 422, 423, 424
turgidis { Tirolites), 343, 493, 666
tururpensis {Columbites), 345
{Olenekites), 346
typus {Goniodiscus), 482
ugra (]aponites), 350, 351
undulatus {Tirolites), 342, 493, 494, 495, 678
undatus {Dinarites), 328, 344, 346, 347, 499, 506,
507, 510
unictim {Dagnoceras), 355
? {Dagnoceras), 354, 458, 461
{Metadagnoceras) , 337, 354, 355, 461,
462
ursensis {Celtites), 360, 369, 372, 373, 646
{"Celtites'), 369
Ussuria compressa, 448
Ussuriidae, 326, 337, 448
Ussurites, 321, 328, 329, 337, 359, 528, 531, 602,
610, 628
? decipiens, 523, 524, 525, 526, 586
hara, 350, 351
hosei, 328, 359, 528, 529, 531, 606, 608
kingi, 350
man.^ieldi, 337, 360, 528, 529, 530, 531,
628, 630
sieveri, 328, 358, 528, 529, 530, 531, 602,
604, 610
Ussuritidae, 328, 524
variabilis {Eophyllites), 531, 532, 533
var. evoluta {Eophyllites), 531
var. involuta {Eophyllites), 531
• {Leiophyllites), 328, 334, 335, 340, 355,
531, 533, 534, 535, 584
vasantasena {Gymnites), 349
vermiformis {Leiophyllites), 352 ,535
Vickohlerites, 326, 350, 408
sundaicus, 326, 340, 408, 409
( cf . ) , 347, 409
volutus {Dinarites), 356, 488
{Dinarites [Olenekites]), 488
volzi {Gymnites), 350
vonderschmitti {Prosphingites) , 404, 405, 407,
410, 411
{Zenoites), 326, 340, 410
vyasa {Hollandites) , 350
wasserbergi {Iseulitoides), 326, 358, 411, 413,
414, 418,419,598
ivebbianus {Stacheites), 350, 455
welteri {Alhanites), 477, 478, 481
{Subvishnuites), 373
wheeleri {Pseudaspidites), 383
woodwardi {Otoceras), 316, 317
Xenaspis enveris, 368, 373, 374
mediterranea, 338, 367, 548
Xenoceltidae, 325, 337, 375
Xenoceltites, 325, 337, 353, 372, 375, 636
eompresstts, 352
crenoventrosus, 325, 352, 353, 376
discretus, 382
gregoryi, 376
hyperboreus, 382
multiplicatus, 382
sinuatus, 322, 325, 348, 376
spencei, 337, 360, 376, 377, 378, 380, 636,
644
sp. indct., 347, 348
spitsbergensis, 325, 354, 355, 376, 377
( cf . ) , 359, 440
subevolutus, 375, 376
Xenodiscus dentosus, 356, 452, 453
diseretus, 382
euomphulus, .356, 452, 465
hyperboreus, 382
karpinskii, 452, 465
keirensis, 367
lac/ueus, 376
multiplicatus, 382
nangaensis (sp. ind. aff. ), 379
prior, 497
schmidti, 356, 451, 452
sp., 345
sulioticus, 338, 341, 379
yaliensis {Columbites), 352, 443, 444
yasoda {Procladiscites) , 350, 351
yudishthira {Pseudaspidites), 383
zappanense {Dagnoceras), 327, 338, 340, 350,
457,458, 459, 460, 570, 588
cf. {Dagnoceras), 344, 458, 459
Zenoites, 326, 332, 404, 405, 407, 409, 410
arcticus, 326, 357, 411
helenae, 326, 340, 407, 410, 411
vonderschmitti, 326, 340, 410
560 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 1. SUBCOLUMBITES PERRINISMITHI
Figures Page
1-9 Subco/umbifes perrinhmilhi (Arthaber). 427
Figs. 1, 2, front and side view of Columbites europaeus Arthaber (1911: pi. 23(7), figs. 15 a-c), X 1- Figs.
3, 4, front and side viev/ of Columbites europaeus Arthaber (1911: pi. 23(7), figs. 13 a-c), X 1-5. Figs. 5, 6,
front and side view of Columbites europaeus Arthaber (1911: pi. 23(7), figs. 16 a, b), X 1- Figs. 7, 8, ventral
and side view of Columbites europaeus Arthaber (1911: pi. 23(7), figs. 17 a, b), X 1. Fig. 9, side view of
Columbites europaeus specimen which yielded the suture of Arthaber (1911: pi. 23(7), fig. 14), X 1-
All specimens are from the Subco/umb//es fauna of Kcira, Albania, and are deposited in the Poleontological
Institute, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 561
8
Plate
562 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 2. SUBCOLUMBITES and ARIANITES
Figures Page
1-4 Subcolumbites dusmani (Arthaber) 433
Figs. 1, 2, syntype (Arthaber, 1911: pi. 24(8) figs. Ic, d), X 1- Figs. 3, 4, syntype (Arthaber, 1911: pi.
24(8), figs, la, b; designated lectotype by Renz and Renz, 1948: 21), X 1.
5-8 Subcolumbi/es perrinismiihi (Arthaber) 427
Figs. 5, 6, front and side view of Columbites europaeus Arthaber (1911: pi. 23(7), figs. 18a, b), X 1- Figs. 7,
8, front and side view of syntype of Columbites mirdltensis Arthaber (1911: pi. 24(8), figs. 3a, b), X 1-
9, 10Ar/an;fes musacchi Arthaber 446
Holotype, Arthaber (1911: pi. 24(8), figs. 5a-c), X 1-5.
All specimens are from the Subco/umbiles fauna of Kcira, Albania, and are deposited in the Paleontological
Institute, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 563
\ .'^
Plate 2
564 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 3. SUBCOLUMBITES and EPICELTITES
Figures Page
1-9 Subcolumbites perrinismithi (Arthaber) 427
Figs. 1-3, front, ventral, and side view of Columbites perrini smithi Artfiaber (1911: pi. 23(7), figs. 19a, b),
X 1. Figs. 4, 5, front and side view of Columbites mirdltensis var. Arthaber (1911: pi. 24(8), figs. 4a, b),
X 1- Figs. 6, 7, ventral and side view of Columbites perrini smtthi Arthaber (1911: pi. 23(7), figs. 20a, b),
X 1- Figs. 8, 9, front and side view of syntype of Columbites mirditensis Arthaber (1911: pi. 24(8), figs.
2a-c), X 1.
10,11 Epiceltites genf/'i Arthaber 447
Side and ventral view of holotype, Arthaber (1911: pi. 24(8), fig. 8), X 1-
All specimens are from the Subcolumbi/es fauna of Kcira, Albania, and are deposited in the Paleontological
Institute, Vienna.
Ammonoids of the Late Scythian- (Lower Triassic) • Kummel 565
/ ...
Plate 3
566 Bulletin Museum of Comparative Zoology, Vol. 137, ISJo. 3
PLATE 4. SUBCOLUMBITES, PREFLORIANITES and DIENEROCERAS
Figures P°9e
1-4 Subcolumbites perr/n/smiffii (Arthaber) ^127
Figs. 1, 2, side and front view of holotype (Arthaber, 1908: pi. 12(2), fig. la-c), X 1- Figs. 3, 4, side and
ventral view of fiolotype of Co/umb/fes europoeus Arthaber (1908: pi. 12(2) figs. 2a, b], X 1.
5, 6 Prellorianites sulioticus (Arthaber) 379
Side and ventral view of unfigured specimen of Ophiceras cfr. nangoensis, — Arthaber (1911: 239), X 1-5
7-10 Dieneroceras wediterranea (Arthaber) 367
Figs. 7, 8, side and ventral view of holotype of Xenospis med/ferranea Arthaber (1908: 260, pi. 11(1), figs.
3a-c), X 1- Figs- 9, 10, side and ventral view of holotype of Ce/fites fccirens/s Arthaber (1908: 273, pi. 11(1),
figs. 8a-c), X 1.5.
All specimens are from the Subco/umbites fauna of Kcira, Albania, and are deposited in the Paleontological
Institute, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 567
\*"-
r •->«
Plate 4
568 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 5. ISCULITOIDES ORIGINIS
Figures Page
1-10 Isculitoides originis (Arthaber) 413
Figs. 1, 2, ventral and side view of genotype (Arthaber, 1911: pi. 23(7), fig. 1), X 1-5. Figs. 3, 4, ventral
and side viev/ of parotype (Arthaber, 1911: pi. 23(7), fig. 2), X 1- Figs. 5, 6, front and side view of paratype
(Arthaber, 1911: pi. 23(7), fig. 3), X 2. Figs. 7, 8, side and ventral view of paratype (Arthaber, 1911: pi.
23(7), fig. 5), X 1-5. Figs. 9, 10, ventral and side view of paratype (Arthaber, 1911: pi. 23(7), fig. 4), X 1-5.
All specimens are from the Subco/umb/tes fauna of Kcira, Albania, and are deposited in the Paleontological
Institute, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 569
/■
^""V
* .H
V.
L^
i
*::>
>
5 <i|k
^
/
; i •
■^ ^'i
•M
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Plate 5
570 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 6. ISCULITOIDES and ARNAUTOCELTITES
Figures Page
1-6 Isculitoides originis (Arthaber) 413
Figs. 1, 2, side and ventral view of paratype (Arthaber, 1911: pi. 23(7), fig. 6), X 2. Figs. 3, 4, side and
ventral view of paratype (Artfiaber, 1911: pi. 23(7), fig. 7), X 1-5. Figs. 5, 6, side and ventral view of
paratype (Arthaber, 1911: pi. 23(7], fig. 10), X 1-5.
7-13 Arnautoceltites mediterraneus (Arthaber) 397
Figs. 7, 8, plesiotype of Nannites herberti Diener, -Arthaber (1908: pi. 11(1), fig. 7), X 2. Figs. 9, 10, lecto-
type (Arthaber, 1911: pi. 18(2), fig. 8), X 2. Fig. 11, paralectotype, X 2. Figs. 12, 13, lectotype Ce///fes
arnauticus Arthaber (1911: pi. 24(8), fig. 7), X 2.
All specimens are from the Subco/umbifes fauna of Kcira, Albania, and are deposited in the Paleontological
Institute, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 571
^ ..
«."■
..»%
s^m^
^^. %^
«y ,■
11
•:->^'
>
tCiS^i^ '
Plate 6
572 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 7. HELLENITES, ARNAUTOCELTITES, and PRENKITES
Figures Page
1-4 Hellenites praematurus (Arthaber) 512
Figs. 1, 2, ventral side view of holotype (Arthaber, 1911: pi. 24(8), figs. 9a, b), X 1-5. Figs. 3, 4, ventral
and side view of paratype (variety) (Arthaber, 1911: pi. 24(8), figs. 10a, b), X 2.
5, 6 Arnaufoce/t/fes mediterraneus (Arthaber) 397
Ventral and side view of holotype of Paragoceras dukagini Arthaber (1911: pi. 24(8), figs. 6a-c), X 2.
7-10 Prenkites malsorensis Arthaber 441
Figs. 7, 8, ventral and side view of lectotype, Arthaber (1911: pi. 22(6), figs. 17a, b), X 1-5. Figs. 9, 10,
ventral and side view of paralectotype, Arthaber (1911: pi. 22(6), figs. 19a, b), X 1-5.
All specimens are from the Subco/umb/'fes fauna of Kcira, Albania, and are deposited in the Paleontological
Institute, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 573
f>^v
i ¥'
10
Plate 7
574 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 8. PROPTYCHITOIDES DECIPIENS
Figures Page
1-4 Proptychltoides decipiens Spath 385
Figs. 1, 2, ventral and side view of synfype of Proptychites kraftti Arthaber (1911: pi. 19(3), figs. 3a, b),
X 1- Figs. 3, 4, ventral and side vievv' of tiolotype of Proptychitoides decipiens [^zProptychites latilimbriatus de
Koninck,-Arttiaber, 1911: pi. 19(3), figs. 2a, b), X 1-
Both specimens are from the Subcolumbites fauna of Kcira, Albania, end are deposited in the Paleontological
Institute, Vienna.
Ammonoids of the Late Scythiax (Lower Triassic) • Kiimmel 575
\
\
A
Plate 8
576 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 9. PROPTYCHITOIDES TRIGONALIS
Figures Page
1-4 Proptychitoides trigonalis (Arthaber) 388
Figs. 1, 2, ventral and side view of holotype of Proptychites bertisci Arthaber (1911: pi. 19(3), figs. 5a-c),
X 1- Figs. 3, 4, ventral and side view of tiolotype of Proptychites trigonalis Artfiaber (1911: pi. 19(3), figs.
4a-c), X 1.
Both specimens are from the Subco/umb/fes fauna of Kcira, Albania, and are deposited in the Paleontological
Institute, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 577
\
'f'y
I
/.
Plate 9
578 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 10. PROPTYCHITOIDES TRIGONALIS
Figures Page
1-4 Proptychitoides trigonalis (Arthaber) 388
Figs. 1, 2, ventral and side view of holotype of Mee(coceros mahomedis Arthaber (1911: pi. 22(6), figs. 3a-c),
X 1. Figs. 3, 4, rigfit and left side views of Proptychitoides (?) nopcsoi Spath (r=Proptych/fes ob/iquep//ca-
fus,-Artfiaber [non Waagen], 1911: pi. 20(4), figs, la-c), X 1.
Both) specimens are from tfie Subco/umbites fauna of Kcira, Albania, and are deposited in the Paleontological
Institute, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 579
(
./
Plate 10
580 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 11. PROCARNITES and PROPTYCHITOIDES
Figures Page
1-4 Procarnites kokeni (Arthaber) 391
Figs. 1, 2, side and ventral view of lectotype of Procarnites skanderbegis Arthaber (1911: pi. 18(2), figs. 7a-c),
X 1. Figs. 3, 4, side and ventral vievv of paralectotype of Procarnites skanderbegis Arthaber (1911: pi. 18(2),
figs. 6a-c], X 1-
5 Proptychitoides trigonalis (Arthaber) 388
Ventral view of specimen shown on PI. 10, figs. 3, 4, XI.
All specimens are from the Subco/L/mfa/7es fauna of Kcira, Albania, and are deposited in the Paleontological
Institute, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 581
Plate 1'
582 BiiUetin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 12. PROCARNITES, PROPTYCHITOIDES, and PSEUDOSAGECERAS
Figures Page
1, 2 Procarnites kokeni (Arfhaber) 391
Side and ventral view of topotype (Arthaber, 1911: pi. 18(2), figs. 5a, b), X 0.7.
3 Proptychitoides decipiens Spath 385
Side view of tiolotype of //leefcoceros hakki Arthaber (1911: pi. 22(6), figs, la-c), X 0.5.
4, 5 Pseudosogeceras dr/nense Artfiober 363
Side and ventral view of holotype, Artfiober (1911: pi. 17(1), figs. 6a, b), X 1-
All specimens ore from the Subco/umbifes fauna of Kcira, Albania, and are deposited in the Paleontological
Institute, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 583
Plate 12
584 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 13. PROCARNITES KOKENI
Figures Page
1-8 Procornifes kokeni (Arthaber) 391
Figs. 1, 2, side and ventral view of lectotype (Arthaber, 1908: pi. 11(1), figs, la-c), X 2. Figs. 3, 4, side
and ventral view of parolectotype (Arthaber, 1908: pi. 11(1), figs. 2a, b), X 2. Figs. 5, 6, side and ventral view
of plesiotype (Arthaber, 1911: pi. 17(1), figs. 17a-c), X 1. Figs. 7, 8, side and ventral view of plesiotype
(Arthaber, 1911: pi. 17(1), figs. 16a, b), X 1.5.
All specimens are from the Subco/umbites fauna of Kcira, Albania, and are deposited in the Paleontologicol
Institute, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kinnmel 585
Plate 13
586 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 14. PROTROPITES and METAHEDENSTROEMIA
Figures Page
1-8 Profrop/fes hilmi Arthaber 445
Figs. 1, 2, ventral and side view of paralectotype, Arthaber (1911: pi. 22(6), figs. 13a, b), X 1-5. Figs. 3,
4, ventral and side view of lectotype, Arthaber (1911: pi. 22(6), figs. 15a, b), X 1-5. Figs. 5, 6, ventral
and side view of paralectotype, Arthaber (1911: pi. 22(6), figs. 12a, b), X 1-5. Figs. 7, 8, ventral and side
view of paralectotype, Arthaber (1911: pi. 22(6), figs, 14a, b), X 1-5.
9, 10 Melahedenstroemia kastriotae (Arthaber) 449
Side and front view of holotype (Arthaber, 1911: pi. 17(1), figs. 14a-c), X 1-
All specimens are from the Subco/umbifes fauna of Kcira, Albania, and are deposited in the Paleontological
Institute, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 587
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Plate 14
588 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 15. DAGNOCERAS
Figures Page
1,2 Dagnoceras nopcsanum Arthaber 459
Ventral and side view of holotype, Arthaber (1911: pi. 21(5), figs. 6a-c), X 1-5.
3-1 1 Dognoceras zappanense Arthaber 459
Figs. 3, 4, ventral and side viev/ of poralectotype, Arthaber (1911: pi. 21(5), figs. 8a, b), X 1- Figs. 5, 6,
ventral and side viev/ of lectofype, Arthaber (1911: pi. 21(5), figs. 9a, b), X 1- Figs. 7, 8, ventral and side
view of lectotype of Dognoceras lejanum Arthaber (1911: pi. 21(5), figs. 13a-c), X 1- Figs. 9-11, ventral and
side views of paralectofype of Dognoceras lejanum Arthaber (1911: pi. 21(5), figs. 12a, b), X 1-5.
All specimens are from the Suibco/umbites fauna of Kcira, Albania, and are deposited in the Paleontological
institute, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 589
8
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10
Plate 15
590 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 16. DAGNOCERAS, METADAGNOCERAS, and "ASPIDITES"
Figures Page
1 , 2 Dagnoceras nopcsanum Arthaber 459
Side and ventral view of variety of Arthaber (1911: pi. 21(5), figs. 7a-c), X 1-5.
3-6 Albanites triadicus (Arthaber) 477
Figs. 3, 4, side and ventral viev/ of lectotype of Aspidites hasserti Arthaber (1911: pi. 21(5), figs. 16 a-c),
X 1- Figs. 5, 6, side end ventral view of paralectotype of Aspidites hasserti Arthaber (1911: 249), X 2.
7, 8 Metadognoceros terbunicum (Arthaber) 463
Ventral and side view of lectotype of Dagnoceras terbunicum Arthaber (1911: pi. 21(5), figs. lOa-c),
X 1.
9, 10 "Aspidites" marginalis Arthaber 339
Ventral and side view of holotype, Arthaber (1908: pi. 11(1), figs. 6a-c), X 2.
All specimens are from the Subcolumbites fauna of Kcira, Albania, and are deposited in the Paleontological
Institute, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 591
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Plate 16
592 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 17. ALBANITES TRIADICUS
Figures ^ Page
1-10 Albanites triadicus (Arthaber) 477
Figs. 1, 2, side and ventral view of paralectofype of Pronorites orbonus Arttiaber (1911: pi. 17(1), figs. 12a,
b), X 1- Figs. 3, 4, unfigured specimen of Arthaber (1911), X 2. Figs. 5, 6, side and ventral view of lecto-
fype of Pronorifes osmanicus Artfiaber (1911: pi. 17(1), figs. lOa-c), X 1- Figs. 7, 8, side and ventral view of
fiolotype of Pronor/'fes triadicus Artfiaber (1908: pi. 11(1), figs. 4a, c; 1911: pi. 17(1), figs. 8, 9), X 1-5.
Figs. 9, 10, side and ventral view of lectotype of Pronorites arbanus Artfiaber (1911: pi. 17(1), figs. 11 a-c),
X 1.5.
All specimens are from tfie Subco/umb/fes fauna of Kcira, Albania, and are deposited in tfie Paleontological
Institute, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 593
■^V;,^^- -. ■
Plafe 17
594 Bullet in Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 18. Tl ROUTES, ALBANITES, and METADAGNOCERAS
Figures Page
1-6 Tirolites idrianus (Hauer) 492
Figs. 1, 2, side and ventral view of Tirolites recfongu/aris,-Arthaber (1911: pi. 22(6), figs. 5a, b), X 1.
Figs. 3, 4, side and ventral viev/ of Tirolites /7/yr/cus,-Arthaber (1911: pi. 22(6), figs. 4a, b), X 1.5. Figs. 5,
6, side and ventral view of Tirolites seminudus, -Arthaber (1908: pi. 11(1), figs. 9a-c), X 1-5.
7, 8 Albanites friadicus (Arthaber) 477
Side and ventral view of holotype of Dagnoceras komanum Artfiaber (1911: pi. 21(5), figs, lla-c), X 2.
9, 10 Mefodagnoceras terbunicum (Arthaber) 463
Side and ventral view of Mee/coceros radiosum, -Arthaber (1911: pi. 21(5), figs. 14a-c), X 1-
All specimens are from the Sufacolumb/tes fauna of Kcira, Albania, and are deposited in the Paleontological
Institute, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 595
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Plate 18
596 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 19. PREFLORIANITES, DIENEROCERAS, "LECANITES"
Figures Page
1 , 2 Preflorianites sulioticus (Arthaber) 379
Side and ventral view of Ophiceras cf. nangoensis,-Arthaber (1911: pi. 21(5), figs. 5a, b), X 1-5.
3, 4 D/eneroceros mediterranea (Arthaber) 367
Side and ventral view of Ophiceros sa/cunto/o, -Arthaber (non Diener) (1911: pi. 21(5), figs. 4a, b), X 1-
5-8 Prellorianites sulioticus (Arthaber) 379
Figs. 5, 6, syntype (Arthaber, 1911: pi. 20(4), figs. 2a, b), X 1-5. Figs. 7, 8, syntype (Arthaber, 1911: pi.
19(3), figs. 6a, b), X 1.
9, 10 "Lecon/fes" niozi Arthaber 338
Side and ventral view of holotype, Lecon//es niazi Arthaber (1911: pi. 21(5), figs. 3a-c), X 2.
All specimens are from the Subco/umb/tes fauna of Kcira, Albania, and are deposited in the Paleontological
Institute, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 597
8
Plate 19
598 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 20. TUNGLANITES, DIENEROCERAS, CORDILLERITES, ALBANITES, PREFLORIANITES, PROSPHINGITES,
and MEROPELLA
Figures Page
1,2 Tunglanites o/exi n. sp. 423
Side and ventral view of holotype {=:Styrites /i7angensis,-Arfhaber (non Diener) (1911: pi. 23(7), fig. 12),
X 1.5.
3, 4 Dieneroceras skutarensis (Arthaber) 368
Side and ventral view of fiolotype of Lecanites skutarensis Arthaber (1911: pi. 21(5), fig. 1), X 1-5.
5, 6 Cord/l/er/'tes angulatus Hyatt and Smith 364
Side and ventral view of holotype of Hedenstroemia skipetarensh Arthaber (1911: pi. 17(1), fig. 13); Fig.
5, X 2, Fig. 6, X 2.5.
7, 8, 9 Albanites friadicus (Arthaber) 477
Side and ventral views of Pseudosiblrltes cfr. dichotomus, -Arthaber (1911: pi. 22(6), fig. 8), X 1-5.
10, 11 Preflorianites garbinus (Renz and Renz) 381
Side and ventral view of Inyoites garbinus Renz and Renz (1948). NHMB 13697, X 1.5.
12, 13 Prosph'mgitez ali Arthaber 405
Side and ventral view of holotype, Arthaber (1911: pi. 22(6), fig. 6), X 2.
14, 15 Meropella p/e/onoe Renz and Renz 477
Side and ventral view of paratype, NHMB J19550, X 2.
Specimens of Figures 1-9, 12, 13 are from the Subco/umb/fes fauna of Albania and are deposited in the
Paleontological Institute, Vienna; specimens of Figures 10, 11, 14, 15 are from the Subco/umb/'/es fauna of
Chios and are deposited in the Natural History Museum, Basel.
Ammonoids of the Late Scythiax (Lower Triassic) • Kummel 599
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Plate 20
600 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 21. EOGYMNITES, BEATITES, and PSEUDOSAGECERAS
Figures Page
1,2 Eogymnites arthaberi (Diener) 517
Side and ventral view of hoiotype.-Arthaber (1911: pi. 20(4), fig. 4), X 1-
3, 4 Beatites berthae Arthaber 449
Side and ventral viev*' of holotype, Artfiaber (1911: pi. 17(1), fig. 15), X 1-5.
5, 6 Pseudosageceras albanicum (Arthaber) 363
Side and front view of holotype (Arthaber, 1908: pi. 13(3), figs, la, b; 1911: pi. 17(1), figs. 4, 5), X 1-
All specimens are from the Subcolumbifes fauna of Kclra, Albania, and are deposited in the Paleontological
Ini'titute, Vienna.
/
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 601
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Plate 21
602 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 22. EOPHYLLITES and LEIOPHYLLITES
Figures Page
1-4 Eophyllites dieneri (Arthaber) 524
Figs. 1, 2, side and ventral view of holotype of Monophyllites nopcsai Arthaber (1908: pi. 12(2), figs.
5a-c), X 1-5. Figs. 3, 4, side and ventral view of fiolotype of Eophyllites refroctus Spath (^ Monophy/Zifes
hora, -Arthaber [non Diener], 1908: pi. 12(2), figs. 4a-c), X 2.
5-10 Leiophy/Zifes variabilis (Spath) (= Monophy/lifes pifomoho, -Arthaber [non Diener]) 531
Figs. 5, 6, paralectotype, -Arthaber (1911: pi. 20(4), figs. lOa-c), X 1.5. Figs. 7, 8, unfigured specimen, -Artha-
ber (1911: 234), X 1-5. Figs. 9, 10, paralectotype, -Arthaber (1911: pi. 20(4), figs. 9a, b), X 2.
All specimens are from the Subco/umbites fauna of Kcira, Albania, and are deposited in the Paleontological
Institute, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 603
8
10
Plate 22
604 Bulletin Museum of Comparotivc Zoology, Vol. 137, No. 3
PLATE 23. EOPHYLLITES
Figures Page
1-7 Eophyllites dieneri (Arfhaber) 524
Fig. 1, suture specimen of Monophyllites dienen Arthaber (1911: pi. 20(4), fig. 8), X 1- Figs. 2, 3, syntype,
Arthaber (1911: pi. 20(4), figs. 5a-c), X 1- Figs. 4, 5, unfigured specimen of Arthaber, X 1- Figs. 6, 7,
side and front view of Ussurifes (?) decipiens Spath (^ Monophyllites >;ingi, -Arthaber [non Diener], 1911: pi.
20(4), figs. 12a-c), X 1.
All specimens are from the Subco/umb/fes fauna of Kcira, Albania, and are deposited in the Paleontological
Institute, Vienna.
Amjsionoids of the Late Scythian (Lower Triassic) • Kummel 605
^ ^ 5
Plate 23
606 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 24. METADAGNOCERAS, DAGNOCERAS, and SIBIRITES
Figures Page
1-3 Mefadagnoceras freemani n. sp. 463
Right and left side and ventral view of holotype, BMNH C33701 . From Nifoekoko, Timor, X 1-
4, 5 Dagnoceros zoppanense Arthaber 459
Side and ventral view of specimen from Nifoekoko, Timor. BMNH C33713, X 2.
6-9 Sibirifes renzi n. sp. 483
Side end ventral view of two paratypes. Figs. 6, 7, NHMB J19551, X 5; Figs. 8, 9 NHMB J19552, X 5 from
Maradovuno, Chios.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 607
\
8
Plate 24
608 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 25. PROPTYCHITOIDES, PROHUNGARITES, and HEMILECANITES
Figures Page
1,2 Proptychitoidei arthaberi (Welter) 390
Ventral and side view of manganese coated specimen, presumably same age as fauna from block E at
Nifoekoko, Timor, X 1-
3-8 Prohungantei middlemism (Diener) 520
Figs. 3, 4, side and ventral view of syntype (Diener, 1913: fig. 6) GSI 11277, X 1- Figs. 5, 6, side and
ventral view of syntype (Diener, 1913: fig. 5) GSI 11277, X 1- Figs. 7, 8, side and ventral view of syntype
(Diener, 1913: fig. 7) GSI 11278, X 2. Specimens from loose block, Pastannah, Kashmir.
9, 10 Hem/7ecanites discus (Arttiober) 374
Side and ventral view of specimen from Subco/umbifes fauna of Cfiios, NHMB J13703, X 1-5.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 609
Plate 25
610 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 26. SVALBARDICERAS, KEYSERLINGITES, and PROSPHINGITES
Figures Poge
1-4 Svalhardiceras spitzbergensis (Frebold) 450
Figs. 1, 2, plesiotype {^Ammonites sp. indet. Frebold, 1929b: pi. 1, fig. 12), X '• Figs. 3, 4, plesiotype
[ = Ammonites sp. indet. Frebold, 1929b; pi. 1, fig. 13), X 1.
5 Svalbardiceras ichmidti (Mojsisovics) 451
Side view of Mee/coceras sp. indet. Frebold (1929b: pi. 1, fig. 11), X 1-
6, 7 Keyser/ingifes subrobuitui (Mojsisovics) 485
Fig. 6, side viev^ of specimen figured by Frebold (1929b: pi. 2, fig. 9), X 1- Fig. 7, side view of specimen
figured by Frebold (1929b: pi. 2, fig. 8), X 1.
8 Proiphingitei czekanowskii Mojsisovics 405
Topotype specimen, MCZ 8677, X 1 5.
Specimens of Figures 1-7 are from upper Scytfiian liorizon at Cape Thorson, Isfjord, Spitsbergen; specimen
of Figure 8 is from the mouth of the Olenek River, Siberia.
Ammonoids of the Late Scythian (Lo^^•ER Triassic) • Kunwiel 611
Plate 26
612 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 27. METADAGNOCERAS and PSEUDOCELTITES
Figures Page
1-4 Metadagnoceras tobini n. sp. 461
Figs. 1, 2, side and ventral view of paralype, MCZ 9638, X 1- Fig- 3, holotype, MCZ 9637, X 1- Fig. 4,
paratype, MCZ 9639, X 1.
5-10 Pzeudocehites multiplicatus (Waagen) 440
Figs. 5, 6, side and ventral view of holotype of Celtites multiplicatus Waagen (1895: pi. 7, figs. 2a-c), GSI
7062, X 1. Figs. 7, 8, side and ventral view of syntype of Celtites armatus Waagen (1895: pi. 7, figs, la-c),
GSI 7061, X 1- Figs. 9, 10, side and ventral view of syntype of Celtites armatus Waagen (1895: pi. 7, figs.
7a-c), GSI 7067, X 1.5.
Specimens of Figures 1-4 are from the Tobin Formation, Tobin Range, Nevada; specimens of Figures 5-10 are
from the Mianwali Formation, Salt Range, West Pakistan.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 613
Plate 27
614 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 28. STACHEITES FLOWERI
Figures Page
1-10 Stacheites floweri n. sp. 456
Fig. 1, paratype, MCZ 9439, X 1- Fig. 2, paratype, MCZ 9440, X 1- Figs. 3, 4, holotype, MCZ 9441, X 1-
Fig. 5, paratype, MCZ 9442, X 1- Figs. 6, 7, paratype, MCZ 9443, X 1-5. Fig. 8, paratype, MCZ 9444,
X 1- Fig. 9, paratype, MCZ 9445, X 1- Fig. 10, paratype, MCZ 9446, X 1-
All specimens ore from the Tobin Formation, Tobin Range, Nevada.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 6L5
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Plate 28
616 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 29. ISCULITOIDES and HEMILECANITES
Figures Page
1-10 hculitoides wosserbergi n. sp. 418
Figs. 1-3, holotype, MCZ 9447, X 1-5. Figs. 4-6, poratype, MCZ 9448, X 2. Figs. 7, 8, paratype, MCZ
9449, X 2. Figs. 9, 10, paratype, MCZ 9450, X 3.
11, 12 Hemilecaniles paradiscus n. sp. 375
Paratype, MCZ 9451, X 2.
All specimens ore from the Tobin Formation, Tobin Range, hJevada.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 617
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Plate 29
618 Bulletin Museum of Comparative Zoology. Vol. 137, No. 3
PLATE 30. SUBCOLUMBITES AMERICANUS
Figures Page
1-14 Subcolumbites americanus n. sp. 436
Figs. 1, 2, holotype, MCZ 9430, X 1- Fig. 3, paratype, MCZ 9431, X 1. Fig. 4, paratype, MCZ 9432, X 1-5.
Fig. 5, paratype, MCZ 9433, X 1- Figs. 6, 7, paratype, MCZ 9434, X 1- Fig. 8, paratype, MCZ 9435, X 1.
Figs. 9, 10, paratype, MCZ 9436, X 1.5. Figs. 11, 12, paratype, MCZ 9437, X 1-5. Figs. 13, 14, paratype,
MCZ 9438, X 1.5.
All specimens are from the Tobin Formation, Tobin Range, Nevada.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 619
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620 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 31. ARNAUTOCELTITES, USSURITES, and HEMILECANITES
Figures Page
1-7, Arnautoceltites techerti n. sp. 402
9, 10,
13, 14 Figs. 1, 2, holotype, MCZ 9457, X 1-5. Figs. 3, 4, paratype, MCZ 9458, X 1.5. Fig. 5, paratype, MCZ 9459,
X 1-5. Figs. 6, 7, paratype, MCZ 9460, X 1.5. Figs. 9, 10, paratype, MCZ 9461, X 1-5. Figs. 13, 14,
paratype, MCZ 9462, X 1.5.
8 Ussurites sieveri n. sp. 528
Paratype, MCZ 9464, X 1-5.
11, 12 Subcolumbites americanus n. sp. 436
Paratype, MCZ 9463, X 1.
15, 16 Hemilecanites paradiscus n. sp. 375
Holotype, MCZ 9465, X 1.5.
All specimens are from the Tobin Formation, Tobin Range, Nevada.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 621
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622 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 32. USSURITES SIEVERI
Figures P°9e
1-7 Ussurites siever/' n. sp. ^■^°
Figs. 1, 2, holotype, MCZ 9452, X 1- Fig- 3, paratype, MCZ 9453, X 1- Fig. 4, paratype, MCZ 9454, X 1-
Fig. 5, paratype, MCZ 9455, X 1- Figs. 6, 7, paratype, MCZ 9456, X 1.
All specimens ore from the Tobin Formation, Tobin Range, Nevada.
Ammonoids of the Late Scythian* (Lower Triassic) • Kiimmel 623
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Plate 32
624 Bidk'tin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 33. USSURITES HOSEI
Figures Page
1-6 Ussuntes hosei n, sp. 528
Fig. 1, holotype, USNM 153085, X 1- Fig. 2, parafype, USNM 153086, X 2. Figs. 3, 4, paratype, USNM
153087, X 1- Figs. 5, 6, paratype, USNM 153088, X 1-
All specimens are from the USGS collection Mill, Confusion Range, Utah. From section 15 of Hose and
Repenning (1959).
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 625
Plate 33
626 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 34. PSEUDOCELTITES, PSEUDOSAGECERAS, and TIROLITES
Figures Page
1-5 Pieudoceltites nevadi n. sp. 440
Fig. 1, side view of holotype, USNM 153078, X 1- Figs. 2, 3, side and ventral view of fragment of paratype,
USNM 153079, X 1- Figs. 4, 5, side and ventral view of fragment of paratype, USNM 153080, X 1-
6 Pseudosageceras multilobatuw Noetling 361
Side view, USNM 153072, X 1.
7-10 Tirolites cf. coss;anus (Quenstedt) 503
Figs. 7, 8, side and ventral view of portion of body chamber, USNM 153083, X 1- Figs. 9, 10, side and
ventral view of fragmentary specimen, USNM 153084, X 1-
All specimens are from the USGS Collection Mill, Thaynes Formation, Confusion Range, Utah, associated
with Ussurites hose/ n. sp.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 627
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Plate 34
628 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 35. PROHUNGARITES, EPICELTITES, USSURITES, and HEMILECANITES
Figures Page
1-5, 8, 9 Prohungariles mckelvei n. sp. 520
Figs. 1, 2, holotype, MCZ 9466, X 1- Figs. 3, 4, porotype, MCZ 9467, X 1-5. Fig. 5, parotype, MCZ
9468, X 1.5. Figs. 8, 9, parofype, MCZ 9469, X 1.
6, 7 Epiceltites gentii (Arthober) 447
Fig. 6, plesiotype, MCZ 9470, X 1-5. Fig. 7, plesiotype, MCZ 9471, X 1-5.
10,11 Ussurites sieveri n. sp. 528
Paratype, MCZ 9472, X 1-5.
12 Hemilecanites porad/'scus n. sp. 375
Paratype, MCZ 9473, X 1-5.
Figures 1-9 ore from fhe upper member of Thaynes Formation, Hammond Creek, Bear River Range, south-
east Idaho. Figures 10-12 are from Tobin Formation, Tobin Range, Nevada.
Ammonoids of the Late Scythiax (Lower Triassic) • Kummel 629
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Plate 35
630 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 36. PROHUNGARITES, OLENEKITES, and ISCULITOIDES
Figures Page
1, 2 Prohungarites sp. indet. 522
MCZ 9474, X 1.
3 Prohungarites gutitadfi n. sp. 521
Side view of holotype, MCZ 9475, X 1-
4-7 Olenekites cf. spiniplicatus (Mojsisovics) 489
Figs. 4, 5, plesiotype, MCZ 9482, X 1.5. Figs. 6, 7, plesiotype, MCZ 9476, X 1.5.
8-13 Isculitoides hammondi n. sp. 419
Fig. 8, holotype, MCZ 9477, X 1-5. Figs. 9, 10, paratype, MCZ 9478, X 1-5. Fig. 11, paratype, MCZ
9479, X 1-5. Figs. 12, 13, paratype, MCZ 9480, X 1-5.
14, 15 Prohungarites gufstadt; n. sp. 521
Side and ventral view of paratype, MCZ 9481, X 2.
All specimens are from the upper member of Thaynes Formation, Hammond Creek, Bear River Range, south-
east Idaho.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 631
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Plate 36
632 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 37. KEYSERLINGITES and STACHEITES
Figures Page
1-4 Keyserlingites beorr/Verensfs n. sp. 487
Figs. 1, 2, side and ventral view of holotype, MCZ 9520, X 1- Figs. 3, 4, side and ventral view of para-
type, MCZ 9521, X 1.
5, 6 Keyserlingites bearlakemis n. sp. 486
Side and ventral view of paratype, MCZ 9518, X 3.
7, 8 Stacfieites sp. indet. I 456
Side and ventral view, MCZ 9487, X 1-
9, 10 Stacheites sp. indet. II 457
Side and ventral view, MCZ 9501, X 1-
Specimens of Figures 1-8 came from tfie upper member of Tfiaynes Formation, Hammond Creek, Bear River
Range, soutfieastern Idaho; specimens of Figures 9, 10 came from tfie upper part of Thaynes Formotion,
Sublette Ridge, Wyoming.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 633
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Plate 37
634 Bulletin Museum of Comparative Zoology, Vol 137, No. 3
PLATE 38. KEYSERLINGITES and PROHUNGARITES
Figures Page
1-3 Keyserlingifes bearlakensh n. sp. 486
Figs. 1, 2, side and ventral view of holotype, MCZ 9516, X '■ Fig- 3, side view of paratype, MCZ 9517,
X 1.
4, 5 Prohungorifes sp. indef. 522
Side and ventral view, MCZ 9647, X 1 •
All specimens are from the upper Thoynes Formation, Hammond Creek, Bear River Range, southeast Idaho.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 635
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Plate 38
636 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 39. COLUMBITES PARISIANUS
Figures Page
1-10 Co/umbifes por/s/anus Hyatt and Smith 424
Figs. 1, 2, front and side view of paratype, Hyatt and Smith (1905: pi. 61, figs. 2, 3), USNM 75286b, X 1.
Figs. 3, 4, front and side view of holotype, Hyatt and Smith (1905: pi. 1, figs. 9, 10), USNM 75246a, X 1-
Figs. 5-7, front, ventral, and side views of paratype, Hyatt and Smith (1905: pi. 61, figs. 5-7), USNM
75286c, X 1- Figs. 8, 9, front and side view of paratype, Hyatt and Smith (1905: pi. 1, figs. 12-14), USNM
75246b, X 1. Fig. 10, front view of paratype, Hyatt and Smith (1905: pi. 61, fig. 10), USNM 75286e, X 1-
All specimens are from the middle shale member of Thaynes Formation [Columbites fauna), Paris Canyon, Bear
River Range, southeast Idaho.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 637
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Plate 39
638 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 40. COLUMBITES PARISIANUS
Figures Page
1-11 Co/umb/fes porisfanus Hyatt and Smith 424
Figs. 1, 2, front and side view of holotype of Co/umbifes ornafus Smitfi (1932: pi. 46, figs. 14, 15), USNM
74984a, X 1- Figs. 3, 4, front and side view of paratype of Co/umbi/es spencei Smith (1932: pi. 78, figs.
13-15), USNM 75309g, X 2. Figs. 5, 6, front and side view of paratype of Co/umbi/es spence/ Smith (1932:
pi. 78, figs. 11, 12), USNM 753091, X 2. Figs. 7-9, side, ventral, and front view of paratype of Co/umb/fes
ligatus Smith (1932: pi. 47, figs. 6-8), USNM 74985c, X 1. Figs. 10, 11, side and ventral view of paratype
of Co/umbifes ornafus Smith (1932: pi. 46, figs. 16, 17), USNM 74984b, X 1.
All specimens are from the middle shale member of Thaynes Formation (Co/umb/fes fauna). Pans Canyon, Bear
River Range, southeast Idaho.
Ammonoids of the Late Scythian (Lower Triassic) • Knmmel 639
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Plate 40
640 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 41. COLUMBITES PARISIANUS
Figures Page
1-7 Columbites parisianus Hyatt and Smith 424
Figs. 1, 2, front and side view of fiolotype of Columbites consongu/neus Smitfi (1932: pi. 46, figs. 1, 2), USNM
74983a, X 1- Figs. 3, 4, ventral and side vievv* of paratype of Columbites consanguineus Smith (1932: pi. 46,
figs. 3, 4), USNM 74983b, X 1- Figs. 5, 6, front and side view of paratype of Columbites consanguineus
Smith (1932: pi. 46, figs. 5, 6), USNM 74983c, X 1- Fig. 7, side view of paratype of Columbites parisianus
Hyatt and Smith (1904: pi. 61, fig. 1), USNM 75286a, X 1.
All specimens ore from the middle shale member of Thaynes Formation [Columbites fauna) Paris Canyon, Bear
River Range, southeast Idaho.
Ammonoids of the Late Scythian (Lower Triassic) • Ktimmcl 641
Plate 41
642 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
^4
PLATE 42. COLUMBITES PARISIANUS
Figures Page
1-9 Columbites pan's/onus Hyatt and Smith 424
Figs. 1, 2, front and side view of holotype of Columbites spencei Smith (1932: pi. 78, figs. 1, 2), USNM
75309a, X 1- Figs. 3, 4, front and side view of paratype of Columbites spencei Smith (1932: pi. 78, figs. 5,
6), USNM 75309c, X 1- Figs. 5, 6, front and side view of paratype of Columbites spencei Smith (1932: pi.
78, figs. 9, 10), USNM 75309e, X 2. Fig. 7, side view of paratype of Columbites ligatus Smith (1932: pi. 47, fig.
4), USNM 74985b, X '■ Figs. 8, 9, front and side view of paratype of Columbites spencei Smith (1932: pi. 78,
figs. 7, 8), USNM 75309d, X 1-
All specimens are from the middle shale member of Thaynes Formation [Columbites fauna), Paris Canyon, Bear
River Range, southeast Idaho.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 643
8
Plate 42
644 Bullet in Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 43. SVALBARDICERAS, PREFLORIANITES, and COLUMBITES
Figures Page
1 Svalbardiceras sheldoni n. sp. 453
Side view of holotype, MCZ 9493, X 1-
2, 3 Pretlonanites monfpelierensis n. sp. 382
Side and ventral v\ew of paratype, MCZ 9495, X 1-5.
4, 5 Columbites par/sionus Hyatt and Smith 425
Side and front view of fiolotype of Columbites ligatus Smith (1932: pi. 47, figs. 1-3), USNM 74985a, X 1-
All specimens are from the middle shale member of Thaynes Formation (Co/umb//es fauna), southeast Idaho.
Specimen of Figure 1 from Sage Creek, of Figures 2, 3 from Montpelier Canyon, and of Figures 4, 5 from
Paris Canyon.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 645
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Plate 43
646 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 44. USSURITES, PSEUDOCELTITES, PREFLORIANITES, and PSEUDASPIDITES
Figures Page
1-3 Ussurites mansiield'i n. sp. 530
Fig. 1, side view of paratype, MCZ 9515, X 1- Figs- 2, 3, ventral and side view of parctype, MCZ 9513, X 1-
4-10 Pseudocelf/fes cheneyi n. sp. 438
Fig. 4, parctype, MCZ 9503, X 1. Fig. 5, paratype, MCZ 9504, X 1- Figs. 6, 7, paratype, MCZ 9505,
X 1. Figs. 8, 9, fiolotype, USNM 153073, X '• Fig. 10, paratype, MCZ 9506, X 1-
11-13 Preflorianites montpelierensis n. sp. 382
Fig. 11, paratype, MCZ 9635, X 1- Fig. 12, paratype, MCZ 9498, X 1- Fig. 13, holotype, MCZ 9494, X 1-
14-15 Pseudaspidites popovi n. sp. 383
Side and ventral view of juvenile specimen, MCZ 9636, X 1-
All specinnens came from tfie middle shale member of Thaynes Formation (Co/umbifes fauna), southeast Idaho.
Specimens of Figures 8, 9, from Draney Creek, of Figure 13 from Montpelier Canyon, all others from Hot
Springs.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 647
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648 Bulletin Museum of Comparative Zoology. Vol. 137, No. 3
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Ammoxoids of the Late Scythian (Lower Triassic) • Kitmmel 649
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650 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 46. KEYSERLINGITES and NORDOPHICERAS
Figures Page
1 Keyserlingitei stephensoni n. sp. 487
Side view of the holotype deposited in Department of Geology, Wasfiington State University. Specimen is
presumably from tfie Columbites fauna, Fort HaW Indian Reservation, soutfieast Idafio, X 0.3.
2, 3 Nordophiceras pilatum (Hyatt and Smith) 470
Fig. 2, MCZ 9543, X 1.5. Fig. 3, MCZ 9544, X 1.5.
Both specimens ore from the middle shale member of Thaynes Formation {Columbltes fauna) southeast Idaho,
Figure 2, from Montpeiier Canyon, Figure 3, from Hot Springs.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 651
Plate 46
652 BtiUetin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 47. NORDOPHICERAS
Figures Page
1-5 Nordophiceras jacksoni (Hyatt and Smith) 468
Fig. 1, MCZ 9564, X 1. Fig. 2, MCZ 9565, X 1. Fig. 3, MCZ 9566, X 3. Fig. 4, MCZ 9567, X 1-5.
Fig 5, MCZ 9568, X 1.5.
All specimens are from the middle shale member of Thaynes Formation (Co/umbites fauna), southeast Idaho.
Specimen of Figure 1 is from Montpelier Canyon, Figure 2 is from Pans Canyon, and the others are from Hot
Springs.
6-8 Nordophiceras euomphalus (Keyserling) 465
Fig. 6, MCZ 9655, X 1. Figs. 7, 8, MCZ 8680, X 1-5.
Topotype specimens are from the Olenekian fauna, Olenek River, Siberia.
Ammoxoids of the Late Scythian (Lower Triassic) • Kumme] 653
84
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Plate 47
654 Bulletin Miiscmn of Comparative Zoology, Vol. 137, No. 3
f ■
PLATE 48. NORDOPHICERAS and XENOCELTITES
Figures Page
1-4 Nordophiceras jacksoni (Hyatt and Smith) 468
Figs. 1, 2, side and front view of paratype (Hyatt and Smith, 1905: pi. 62, figs. 15, 16), USNM 75292c, X 1-
Figs. 3, 4, side and front view of holotype (Hyatt and Smith, 1905: pi. 62, figs. 11-13), USNM 75292a, X 1-
5-9 Xenoce/t/tes spencei (Hyatt and Smith) 376
Figs. 5, 6, side and front view of paralectotype (Hyatt and Smith, 1905: pi. 62, figs, 5-7), USNM 75291b, X 1.
Figs. 7-9, side, front, and ventral view ot lectotype (Hyatt and Smith, 1905: pi. 62, figs. 1-3), USNM 75291a,
X 1.
All specimens are from the middle shale mernber of Thaynes Formation (Co/umb/'tes fauna), Paris Canyon, Bear
River Range, southeast Idaho.
Ammonoids of the Late Scythiax (Lower Triassic) • Kiimmcl 655
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Plate 48
656 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 49. NORDOPHICERAS PILATUM
Figures Page
1-8 Nordophiceras pilafum (Hyatt and Smith) 470
Fig. 1, side view of porotype of A/leef;oceras soncforum Smith (1932: pi. 49, fig. 3), USNM 74991b, X 1-
Figs. 2, 3, side and front view of holotype of Meekoceras sanctorum Smith (1932: pi. 49, figs. 1, 2), USNM
74991a, X 1. Figs. 4-6, side, front, and ventral view of paralectotype of A4ee((oceros pilatum Hyatt and
Smith (1905: pi. 63, figs. 10-12), USNM 75294b, X 1. Figs. 7, 8, side and ventral view of lectotype of
Meekoceras pilatum Hyatt and Smith (1905: pi. 63, figs. 7, 8), USNM 75294a, X 1-
All specimens are from the middle shale member of Thaynes Formation [Columbites fauna), Paris Canyon, Bear
River Range, southeast Idaho.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 657
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658 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 50. NORDOPHICERAS PILATUM
Figures Page
1-11 Nordophiceras pilatum (Hyatt and Smith) 470
Figs. 1, 2, side and front view of fiolotype of Mee/<oceras curf/cosfotum Smith (1932: pi. 48, figs. 21-22),
USNM 74990a, X 1- Fig. 3, side view of paratype of A4ee(;oceras curf/cosfatum Smith (1932: pi. 48, figs. 27,
28), USNM 74990d, X 1- Fig. 4, side view of paratype of A4ee/(oceras curficostatum Smith (1932: pi. 48,
figs. 29, 30), USNM 74990e, X 2. Fig. 5, side view of paratype of Meefroceros curticostofum Smith (1932:
pi. 48, figs. 25, 26), USNM 74990c, X 1- Fig, 6, side view of paratype of Meekoceras curticostatum Smith
1932: pi. 48, figs. 23, 24), USNM 74990b, X 1- Figs. 7-9, side, front, and ventral view of holotype of
/Vlee/<oceras micrompho/us Smith (1932: pi. 49, figs. 5-7), USNM 74992a, X 1- Figs. 10, 11, side and front
view of paratype of Meekoceras micromphalui Smith (1932: pi. 49, figs. 9-11), USNM 74992b, X 2.
All specimens are from the middle shale member of Thaynes Formation (Co/umbifes fauna), Paris Canyon, Bear
River Range, southeast Idaho.
Ammonoids of the Late Scythian (Lower Triassic) • Kinnmel 659
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Plate 50
660 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 51. NORDOPHICERAS and CORDILLERITES
Figures Page
1-5 Nordoph/ceros pilalum (Hyatt and Smith) 470
Fig. 1, MCZ 9539, X 1. Figs. 2, 3, MCZ 9542, X 1. Fig. 4, MCZ 9540, X 1- Fig. 5, MCZ 9541, X 1.
6, 7 Cord/7/erifes angulatui Hyatt and Smith 364
Side and ventral view, MCZ 9569, X 1.
All specimens are from the middle shale member of Thaynes Formation (Co/umb/fes fauna), Montpelier Canyon,
southeast Idaho.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 661
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Plate 51
662 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 52. XENOCELTITES SPENCEI
Figures Page
1-7 Xenoceltltes spencei (Hyatt and Smith) 376
Fig. 1, MCZ 9551, X 1. Fig. 2, MCZ 9552, X 2. Fig. 3, MCZ 9553, X 1.5. Fig. 4, MCZ 9554, X 1.5.
Fig. 5, MCZ 9555, X 2. Fig. 6, MCZ 9556, X 1.5. Fig. 7, MCZ 9557, X 3.
All specimens ore from the middle shale member of Thaynes Formation (Co/umbites fauna), southeast Idaho.
Specimens of Figures 1, 4, 6, and 7 are from Montpelier Canyon; specimens of Figures 2, 3, and 5 are from
Hot Springs.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 663
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Plate 52
664 Bulletin Museum of Comparative Zoology, Vol 137, No. 3
Figures
1-12
13, 14
15
PLATE 53. DIENEROCERAS, HELLENITES, and SUBVISHNUITES
D/eneroceras aposfo/icus (Smith)
Figs. 1, 2, side and front view of holotype of Celtites ursensis Smith (1932: pi. 47, figs. 11, 12), USNM 74987a,
X 1. Figs. 3, 4, side and ventral view of paratype of Celtites ursensis Smith (1932: pi. 47, figs. 13, 14), USNM
74987b, X 1- Figs. 5, 6, side and front view of paratype of Celtites ursensis Smith (1932: pi. 47, figs. 15,
16), USNM 74987c, X 1- Figs. 7, 8, side and front view of holotype of Ce/(/fes p/onovolvis Smith (1932:
pi. 48, figs. 11, 12), USNM 74988a, X 1. Fig. 9, side view of paratype of Cehites planovolvis Smith (1932:
pi. 48, figs. 13, 14), USNM 74988b, X '. Figs. 10, 11, side and front view of holotype of Celtites apostoli-
cus Smith (1932: pi. 48, figs. 1, 2), USNM 74989a, X 1. Fig. 12, side view of paratype of Celtites apostoli-
cus Smith (1932: pi. 48, figs. 3, 4), USNM 74989b, X 1.
Helleniles idohoense (Smith)
Side and ventral view of holotype (Smith, 1932: pi. 49, figs. 17, 18), USNM 74994, X 2.
Sulbvishnuites sp. indet.
Side view, MCZ 9512, X 1-
All specimens are from the middle shale member of Thaynes Formation [Columbites fauna), Idaho. The speci-
mens of Figures 1-14 are from Paris Canyon, and the specimen of Figure 15 is from Montpelier Canyon.
Page
369
516
374
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 665
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Plate 53
666 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 54. TIROLITES SMITHI
Figures Page
]-5 Tirolites smithi n. sp. 501
Fig. 1, specimen from Montpelier Canyon, MCZ 9547, X 1. Figs. 2, 3, ventral and side view of Tirolites
illyricus Mo|sisovics from Paris Canyon, figured by Smith (1932: pi. 49, figs. 12, 13), USNM 74993, X 1-
Figs. 4, 5, side and ventral view of juvenile form from Montpelier Canyon, MCZ 9548, X 1-
All specimens ore from the middle shale member of Thaynes Formation (Co/umbifes fauna), southeast Idaho.
Ammonoids of the Late Scythian (Lower Triassic) • Kinnmel 667
Plate 54
668 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
'
PLATE 55. TIROLITES, DALMATITES, and P5EUDASPIDITES
Figures Page
1-3 Tirol/fes asfofihovi n. sp. 502
Figs. 1, 2, side and ventral view of holotype, USNM 153081, X 1- Fig. 3, side view of paratype, USNM
153082, X 1.
4, 5 Tirolites sp. indet. II 503
Side and ventral view, MCZ 9502, X 1-
6, 7 Do/mofifes kittli n. sp. 522
Side and ventral view of holotype, MCZ 9499, X 1-5.
8, 9 Pseudaspidites popovi n. sp. 383
Side and ventral view of tiolotype, MCZ 9575, X 1-
All specimens are from tine middle sfiale member of Tfiaynes Formation (Co/umbiJes fauna), Idalio. Speci-
mens of Figures 1-3, from Sage Creek, of Figures 4, 5, from Montpelier Canyon, of Figures 6, 7, from Paris
Canyon, of Figures 8, 9, from Hot Springs.
Ammonoids of the Late Scythian (Lower Triassic) • Kiunmel 669
Plate 55
670 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 56. DALMATITES and STACHEITES
Figures Page
1-8 Da/mof/fes morlaccus KittI 522
Figs. 1, 2, lectotype, Kitrl (1903: pi. 4, fig. 4), X 1- Figs. 3, 4, paralectotype, KittI (1903: fig. 5), X 1-
Figs. 5, 6, paralectotype, KittI (1903: fig. 6), X 1- Figs. 7, 8, paralectotype, KittI (1903: fig. 7), X 1-
9, 10 Sfocfieites prionoides KittI 455
Holotype, KittI (1903: pi. 4, fig. 8), X 1.
All specimens are from the Werfen Formation at Muc, Dalmatia, and are deposited in tfie Natural History
Museum, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 671
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Plate 56
672 Bulletin Muscinn of Comparative Zoology, Vol. 137, No. 3
i
PLATE 57. BITTNERITES and PSEUDODINARITES
Figures Page
1-6 Bittnerites bittneri KIttI 504
Figs. 1, 2, lectotype, KittI (1903: pi. 11, fig. 10), X 1. Figs. 3, 4, holotype, Bittnerites malici KittI (1903: pi. 3,
fig. 8), X 1- Figs. 5, 6, holotype, Bittnerites telleri KittI (1903: pi. 10, fig. 10), X 1-
7-9 Pseudodinantes mohamedanus (Mojsisovics) 511
Figs. 7, 8, plesiotype,-Kittl (1903: pi. 3, fig. 7), X 1; Fig. 9, plesiotype,-Kittl (1903: pi. 3, fig. 6), X 1-
All specimens are from the Werfen Formation at Muc, Dalmatia, and are deposited in the Natural History
Institute, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 673
Plate 57
674 Bulletin Museum of Comparative Zoologt/, Vol. 137, No. 3
PLATE 58. DINARITES DALMATINUS
Figures Page
1-10 DInarites dalmalinus (Hauer) 506
Fig. 1, plesiotype, Dinarites nudus Tommasi.-KittI (1903: pi. 1, fig. 13), X 1- Fig- 2, plesiotype, Dinarites
laevis Tommasi,-Kittl (1903: pi. 3, fig. 11), X 1- Figs. 3, 4, plesiotype, Dinarites laevis Tommasi,-Kittl (1903:
pi. 3, fig. 10), XI- Figs. 5, 6, plesiotype, D;nor/tes dalmalinui (Hauer), -KittI (1903: pi. 2, fig. 1), X 1-
Fig. 7, plesiotype, Dinarites dalmatinus (Hauer),-Klttl (1903: pi. 2, fig. 5), X 1. Fig. 8, plesiotype, Dinarites
dalmatinus (Hauer),-Kittl (1903: pi. 2, fig. 6), X 1- Figs. 9, 10, plesiotype, Dinorifes dalmatinus (Houer),-
Kittl (1903: pi. 2, fig. 3), X 1-
All specimens are from ttie Werfen Formation at Muc, Dalmatia, end are deposited in tHie Natural History
Museum, Vienna.
Ammoxoids of the Late Scythian (Lower Triassic) • Kuminel 675
Plate 58
676 BiiUciin Mmciun of Comparative Zoology, Vol. 137, No. 3
PLATE 59. DINARITES DALMATINUS
Figures
1-11 Dinarites dalmatinus (Hauer)
Figs. 1, 2, plesiotype, Dinarites laevis Tommasi,-Kittl (1903: pi. 1, fig. 1), X 1- Figs. 3, 4, plesiotype,
Dinarites laevis Tommasi-KittI (1903: pi. 1, figs. 2, 3), X 1. Figs. 5, 6, plesiotype, Dinarites much/onus (Hauer), -
KittI (1903: pi. 1, fig. 7), X 1- Figs. 7, 8, plesiotype, Dinarites muchianus (Hauer),-Kittl (1903: pi. 1, fig. 5),
X 1- Figs. 9, 10, syntype, Dinarites evolutior KittI (1903: pi. 1, fig. 10), X 1- Fig. 11, syntype, Dinarites
evolutior KittI (1903: pi. 1, fig. 9), X 1-
All specimens are from the Werfen Formation at Muc, Dolmotia, and ore deposited in tfie Natural History
Museum, Vienna.
i
Page
506
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 677
%e^
8
■fS
10
Plate 59
I
678 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 60. DINARITES DALMATINUS
Figures Page
1-8 Dinarifes dalmatinus (Hauer] 506
Fig. 1, plesiotype, war. plunmcoitatus KittI (1903: pi. 2, fig. 10), X 1- Fig- 2, plesiotype, var. externep!anatus
KittI (1903: pi. 3, fig. 1), X 1- Fig. 3, plesiotype, var. externeplanatui KittI (1903: pi. 3, fig. 2), X 1-
Fig. 4, plesiotype, war. extensus (KittI: 1903, pi. 2, fig. 8), X 1- Fig. 5, fiolotype Dinarttes multicostaius KittI
(1903: pi. 3, fig. 3), X 1- Fig. 6, syntype, Dinorifes tirolitoides KittI (1903: pi. 7, fig. 3), X 1- Fig. 7, syntype,
Dinarites tirolitoides KittI (1903: pi. 7, fig. 2), X 1- Fig. 8, plesiotype, -KittI (1903: pi. 2, fig. 7), X 1-
All specimens are from tfie Werfen Formation at Muc, Dalmatia, and are deposited in the Natural History
Museum, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 679
ff-'/"*-?
wT~^ -r****
Plate 60
680 Biillcliii MusiuDi of Coinpciidlivc Zoology, Vol. 137, No. 3
PLATE 61. DINARITES CARNIOLICUS
Figures ^°^^
1-8 Dmar/fes carniolicus (Mojsisovics) ^'0
Figs. 1, 2, plesiotype-Klttl (1903: pi. 5, fig. 1], X 1. Fig. 3, plesiotype-KittI (1903: pi. 5, fig. 2), X 1.
Fig. 4, plesiotype-KittI (1903: pi. 5, fig. 3), X 1- Fig. 5, plesiotype,-Kittl (1903: pi. 5, fig. 4), X 1. Figs.
6, 7, syntype, Tnolites serratelobatui Kitfl (1903: pi. 5, fig. 4), X 1- Fig- 8, syntype, Tirolltes serratelobatus
Kitfl (1903: pi. 5, fig. 6), X 1.
All specimens are from the Werfen Formation at Muc, Dalmatia, and are deposited in the Natural History
Museum, Vienna.
Ammonoids of tuk Late Scvthiax (Lower Tiuassic) • Kiimmel 681
Plate 61
682 BuUciin Museum of Comparative ZooJo<i.y, Vol. 137, No. 3
I
PLATE 62. DIAPLOCOCERAS, PSEUDOKYMATITES, and PSEUDODINARITES
Figures Page
1-4 D/op/ococeras connecfens (Mojsisovics) 504
Fig. 1, holotype, Dinarites [Hercegovites] dioclefiani KittI (1903: pi. 3, fig. 4), X 1- Fig. 2, lectotype, Dinarites
[Liccaites] progressus KittI (1903: pi. 4, fig. 2), X 1- Figs. 3, 4, lectotype, Dinarites biangulatus KittI (1903: pi. 4,
fig. 1), X 1.
5 Pseudol<Ymatites svilajanus (KittI) 475
Holotype (KittI, 1903: pi. 4, fig. 3), X 1.
6 Pseudodinarites mohamedanui (Mojsisovics) 511
Plesiotype,-Kittl (1903: pi. 3, fig. 5), X 0.6.
All specimens are from the Werfen Formation at Muc, Dalmatia, and are deposited in the Natural History
Museum, Vienna.
Ammoxoids of the Late Scythiax (Lower Triassic) • Kiimmel 683
M
f
Plate 62
684 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
I
PLATE 63. TIROLITES CASSIANUS
Figures Page
1-9 Tirolites cassianus (Quenstedt) 493
Fig. 1, plesiotype, Tirolites dorwm/,-Kittl (1903: pi. 11, fig. 1), X 1- Figs. 2, 3, plesiotype, Tirolites darwini,-
Kitfl (1903: pi. 10, fig. 5), X 1- Fig. 4, lectotype, Tirolites multispinatus KittI (1903: pi. 11, fig. 9), X 1.
Figs. 5, 6, lectotype, Tirolites percostatus KittI (1903: pi. 10, fig. 6), X 1- Fig. 7, plesiotype, Tirolites
turgidus, -KittI (1903: pi. 10, fig. 7), X 1. Fig. 8, plesiotype, 7;>o//fes dorw/ni,-Kittl (1903: pi. 11, fig. 3), X 1-
Fig. 9, plesiotype, Tirolites sm;riagin/, -KittI (1903: pi. 11, fig. 6), X 1-
All specimens are from the Werfen Formation at Muc, Dalmatia, and are deposited in tfie Natural History
Museum, Vienna. -
Ammonoids of the Late Scythian- (Lower Triassic) • Kummcl 685
i
Plate 63
686
Bulletin Museum of Comparative Zoology, Vol 137, No. 3
PLATE 64. TIROLITES CASSIANUS
c Paqe
Figures =• =
1-4 Tirolites cassionus (Quenstedt) '*°3
Figs. 1, 2, lectotype, Tirolites sp/nosior KittI (1903: pi. 11, fig. 5), X 1- Fig- 3, syntype, Tirolites loulai KittI
(1903: pi. 11, fig. 11), X 1- Fig- 4, plesiotype, Tirolites dor^v in/, -KittI (1903: pi. 10, fig. 11), X 1-
All specimens are from the Werfen Formation at Muc, Dalmatia, and are deposited in the Natural History
Museum, Vienna.
Ammonoids of the Late Scythian: (Lower Triassic) • Kiimmel 687
Plate 64
688 Bulletin Museum of Comparaiive Zoology, Vol 137, No. 3
PLATE 65. TIROLITES CASSIANUS
Figures Page
1-9 Tirolites cassianus (Quenstedt) 493
Fig. 1, figured specimen, Tirolites angustilobatus vor. alpha KittI (1903: pi. 8, fig. 19), X 1- Fig. 2,
lectotype, Tirolites ar)gustilobatus KittI (1903: pi. 9, fig. 3), X 1- Fig. 3, plesiotype, -KittI (1903: pi. 9, fig.
5), X 1- Fig. 4, figured specimen, Tirolites angustilobatus var. alpha KittI (1903: pi. 9, fig. 1), X 1- Fig- 5,
plesiotype,-Kittl (1903: pi. 9, fig. 4), X 1- Fig. 6, plesiotype, -KittI (1903: pi. 9, fig. 6), X 1. Figs. 7, 8,
plesiotype, Tirolites spinosus, -KittI (1903: pi. 9, fig. 7), X 1- Fig. 9, plesiotype Tirolites houer;, -KittI (1903:
pi. 9, fig. 10), X 1.
All specimens are from ttie Werfen Formation at Muc, Dalmatia, and are deposited in tlie Natural History
Museum, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 689
Plate 65
690 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 66. TIROLITES IDRIANUS
Figures Page
1-13 Jirolites idrianus (Hauer) 492
Fig. 1, plesiotype, Tirolites seminudus Mojsisovics var. nud/or KittI (1903: pi. 6, fig. 3), X 1- Fig. 2, plesio-
type, T. seminudus var. pl/cosus KittI (1903: pi. 6, fig. 7), X 1- Fig. 3, plesiotype, 7. seminudus var. plicosus
Kittl (1903: pi. 6, fig. 5), X 1- Fig. 4, plesiotype, T. seminudus,-Kittl (1903: pi. 6, tig. 6), XL Fig. 5, plesio-
type, T. seminudus,-Kittl (1903: pi. 6, fig. 8), X 1- Fig. 6, paralectotype, T. distans KittI (1903: pi. 6, fig.
12), X 1- Fig. 7, plesiotype, T. seminudus, -KittI (1903: pi. 6, fig. 4), X 1- Fig. 8, plesiotype, T. seminudus,-
Kittl (1903: pi. 6, fig. 18), X 1- Fig. 9, lectotype, T. distans KittI (1903: pi. 6, fig. 15), X 1- Figs. 10, 11,
plesiotype, T. semmudus,-Kittl (1903: pi. 6, fig. 10), X 1- Fig. 12, plesiotype, T. seminudus,-Kittl (1903:
pi. 6, fig. 9), X 1- Fig. 13, plesiotype, T. seminudus, -Kittl (1903: pi. 6, fig. 17), X 1-
All specimens are from the Werfen Formation at Muc, Dalmatia, and are deposited in the Natural History
Museum, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 691
Plate 66
692 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 67. TIROLITES IDRIANUS
Figures Page
1-9 Tirolites idrianus (Hauer) 492
Figs. 1, 2, pleslotype, Jirolitei mercurii-K'tWl (1903: pi. 6, fig. 1), X 1- Fig. 3, holotype, Tirolites hetero-
phanus Kittl (1903: pi. 5, fig. 7), X 1- Fig. 4, figured type, Tirolites paucispinatus KittI (1903: pi. 6, fig. 11),
X 1- Fig. 5, plesiotype, Tirolites mercuri/,-Kitfl (1903: pi. 6, fig. 2), X 1- Fig. 6, figured type, Tirolites
repulsus KittI (1903: pi. 8, fig. 10), X 1- Fig. 7, figured type, Tirolites dimidiatus KittI (1903: pi. 8, fig. 15),
X 1- Fig. 8, plesiotype, Tirolites rectongu/or/s, -KittI (1903: pi. 8, fig. 16), X 1- Fig. 9, figured type, Tiro-
lites repulsus KittI (1903: pi. 8, fig. 9), X !■
All specimens are from the Werfen Formotion at Muc, Dalmatic, and are deposited in the Natural History
Museum, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kiimmel 693
Plate 67
694 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 68. TIROLITES IDRIANUS
Figures Page
1-9 Tiro/ifes idrianus (Hauer) 492
Fig. 1, paralectotype, Tiro/ifes d'ntans Kitfl (1903: pi. 7, fig. 8), ^«' 1. Fig. 2, paralectotype, TiVo/ifes distans
KittI (1903: pi. 7, fig. 7), X 1- Fig. 3, figured type, Tiro/ifes paucisp/nofus KittI (1903: pi. 7, fig. 5), X 1-
Fig. 4, plesiotype, Tirolites i7/yricus, -KittI (1903: pi. 8, fig. 3), X 1- Fig. 5, plesiotype, T/ro/ifes quensfedf/,-
Kittl (1903: pi. 6, fig. 20), X 1. Fig. 6, plesiotype, Tirolites quenstedfi,-Kittl (1903: pi. 6, fig. 19), X 1-
Fig. 7, lectotype, Tirolites hybridus KittI (1903: pi. 8, fig. 2), X 1. Fig. 8, figured type, Tirolites rotiformis
KittI (1903: pi. 8, fig. 12), X 1- Fig. 9, figured type, Tirolites rotilormis KittI (1903: pi. 8, fig. 13), X 2.
All specimens are from the Werfen Formation at Muc, Dalmatia, and are deposited in the Natural History
Museum, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 695
Plate 68
696 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 69. TIROLITES IDRIANUS
Figures Page
1-10 Tirolites idrianus (Hauer) 492
Fig. 1, lectotype, Jirolites robustus KittI (1903: pi. 7, fig. 9), X 1- Fig. 2, paralectotype, Tirolites rofausfus
KittI (1903: pi. 7, fig, 10), X 1. Fig. 3, paralectotype, Tirolites robustus KittI (1903: pi. 7, fig. 11), X 1- Fig.
4, paralectotype, Tirolites robustus KittI (1903: pi. 8, fig. 1), X 1- Fig. 5, syntype, Tirolites subillyricus KittI
(1903: pi. 7, fig. 15), X 1- Fig. 6, fioiotype, Tirolites ongusfus KittI (1903: pi. 7, fig. 12), X 1- Fig. 7,
lectotype, Tirolites stachei KittI (1903: pi. 7, fig. 14), X 1- Figs. 8, 9, syntype, Tirolites subillyricus KittI
(1903: pi. 7, fig. 16), X 1- Fig. 10, lectotype, T;>o//tes undulatus KittI (1903: pi. 7, fig. 13), X 1.
All specimens are from tfie Werfen Formation at Muc, Dalmatic, and are deposited in tlie Natural History
Museum, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kummel 697
Plate 69
698 Bulletin Museum of Comparative Zoology^ Vol. 137, No. 3
PLATE 70. HOLOLOBUS, TIROLITES, and DINARITES
Figures Page
1,2 Hololobus monopfychus KIttI 511
Genotype, Jirolites [Hololobus] monoptychus KittI (1903: pi. 4, fig. 9), X 1-
3-6 T/ro/ifes cass/onus (Quenstedt) 493
Figs. 3, 4, lectotype, Ceratites [Paraceratiles] prior KittI (1903: pi. 11, fig. 13) (^ Tirolitoides prior -
Spath, 1934), X 1- Figs. 5, 6, paralectotype, Ceratites [Paraceratites] prior KiftI (1903: pi. 11, fig. 4), X 1-
7, 8 Dinarites dalmatinus (Hauer) 497
Holotype, Dinantes (?) angulatus KittI (1903: pi. 3, fig. 9), X 1.
9, 10 Tnolites cingulatus KittI 497
Holotype, Tirolites [Svilajifei] cingulatus KittI (1903: pi. 8, fig. 18), X 1.
11, 12 Tirolites cass/onus (Quenstedt) 493
fHolotype, Tirolites [Svilaiites] tietzei KittI (1903: pi. 10, fig. 9), X 1-
All specimens are from the Werfen Formation at Muc, Dalmatia, and are deposited in tfie Natural History
Museum, Vienna.
Ammonoids of the Late Scythian (Lower Triassic) • Kunimel 699
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Plate 70
700 Bulletin Museum of Comparative Zoology, Vol. 137, No. 3
PLATE 71. TIROLITES and DALMATITES
Figures Page
1-7 Jirolites harti Smith 501
Figs. 1, 2, side and ventral view of holotype, Tiroliles harti Smith (1932: pi. 57, figs. 9-10), USNM 75022,
X 1- Figs. 3, 4, side and ventral view of holotype, Tirolites knighti Smith (1932: pi. 57, figs. 1, 2), USNM
75020, X 1. Figs. 5, 6, side and ventral view of holotype, Tirolites pealei Smith (1932: pi. 57, figs. 5, 6),
USNM 75021a, X 1- Fig. 7, side view of paratype, Tirolites pealei Smith (1932: pi. 57, figs. 7, 8), USNM
75021b, X 1.
8, 9 Dalmatifes attenuatus Smith 524
Side and ventral view of holotype, Smith (1932: pi. 57, figs. 11, 12), USNM 75023, X 1.
All specimens are from the Tirolites Zone, Thaynes Formation, Paris Canyon, Bear River Range, southeast Idaho.
Ammonoids of the Late Scythian (Lower Triassic) • Kummcl 701
Plate 71
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