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SMITHSONIAN INSTITUTION
UNITED STATES NATIONAL MUSEUM
PROCEEDINGS
OF THE
UNITED STATES NATIONAL MUSEUM
VOLUME 91
UNITED STATES
GOVERNMENT PRINTING OFFICE
WASHINGTON : 1943
ADVERTISEMENT
The scientific publications of the National Museum include two
series, known, respectively, as Proceedings and Bulletin.
The Proceedings, begun in 1878, are intended primarily as a medium
for the publication of original papers, based on the collections of the
National Museum, that set forth newly acquired facts in biology,
anthropology, and geology, with descriptions of new forms and re-
visions of limited groups. Copies of each paper, in pamphlet form,
are distributed as published to libraries and scientific organizations
and to specialists and others interested in the different subjects.
The dates at which these separate papers are published are recorded
in the table of contents of each of the volumes.
The present volume is the ninety-first of this series.
The Bulletin, the first of which was issued in 1875, consists of a
series of separate publications comprising monographs of large zoo-
logical groups and other general systematic treatises (occasionally in
several volumes), faunal works, reports of expeditions, catalogues of
type specimens, special collections, and other material of similar na-
ture. The majority of the volumes are octavo in size, but a quarto
size has been adopted in a few instances in which large plates were
regarded as indispensable. In the Bulletin series appear volumes un-
der the heading Contributions from the United States National Her-
barium, in octavo form, published by the National Museum since
1902, which contain papers relating to the botanical collections of the
Museum.
ALEXANDER WETMORE,
Assistant Secretary, Smithsonian Institution.
qi
536587—43
1
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CONTENTS
Pages
Basster, R. S. The Nevada Early Ordovician (Pogonip)
sponge fauna. No. 3126. November 1, 19413_---_--___-- 91-102
New species: Patellispongia magnipora.
CrargE, J. F. Gates. The North American moths of the
genus Arachnis, with one new species. No. 3123. Novem-
REPRE P 092 god 2 oes Vee eae a 59-70
New species: Arachnis apachea.
Fraser, C. McLuan. New species of hydroids, mostly from
the Atlantic Ocean, in the United States National Museum.
Novos. “November 14-1941 =i sse>ce~-=--ssseeesens04 77-89
New family: Symplectaneidae.
New genus: Symplectanea.
New species: Symplectanea bracteata, Hydractinia valens, Cory-
morpha adventitia, Lampra wuvularis, Tubularia crassa,
2? Campanularia fasciculata, ? Obelia racemosa, Egnundella
grandis, Halecium dubium, Halecium tensum, Lictorella
crassitheca, Aglaophenia inconstans, A. transitionis, Plumu-
laria polynema.
Gazin, C. Lewis. The mammalian faunas of the Paleocene of
central Utah, with notes on the geology. No. 3121. October
2. 105 lel een a ht, Gr IAS ap 2 IE 1-53
New genera: Draconiolestes, Oxytomodon, Desmaioclaenus.
New species: Ptilodus ferronensis, Dracontolestes aphantus,
Protogonodon biatieles, Oxyclaenus pearcei, Tricentes elassus,
Ozytomodon perissum, Desmatoclaenus hermaeus, D. para-
creodus, Ectoconus symbolus, Carsioptychus hamaaxitus, Ani-
sonchus oligistus, Haploconus ? elachistus.
GitmorE, CHartes W. Some littie-known fossil lizards from
the Oligocene of Wyoming. No. 3124. November 13,
Diet imeepeee 8 He or aed nou ANT oP Ain ay wie hel Doe tee 71-76
HrouiéKa, ALES. Catalog of human crania in the United
States National Museum collections: Eskimo in general.
Nosoldie, August 11940 2 ones 5 i net peti Ks 169-429
1 Date of publication.
VI PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
’ 4 , Pages
Loveripcr, ArrHur. Report on the Smithsonian-Firestone
Expedition’s collection of reptiles and amphibians from
Liberia. No. 3128. November 14, 1941 4.2 2 ee 113-140
New species: Typhlops manni, Hylambates cochranae, Leptopelis
bequaerti.
New name: Rana albolabris parkeriana.
Moog, Cartes C. A new fossil crocodilian from Colombia.
No. 3128 ous anuany 1; 1949 Vn chev e¥l et 2. 55-58
New species: Dinosuchus neivensis.
Ruoapes, Renpevy. Notes on some crayfishes from Alabama
caves, with the description of a new species and a new sub-
species. No. 3129. November 6, 1941 3-----_._.-----.---- 141-148
New species: Cambarus (Cambarus) cahni.
New subspecies: Cambarus (Favonius) pellucidus australis.
Scumirr, Watpo L. The species of Aegla, endemic South
American fresh-water crustaceans. No. 3132. August 18,
ONE oc A Se Se A 431-520
New species: Aegla parana, A. sanlorenzo, A. platensis, A.
uruguayana, A. prado, A. castro, A. franca, A. jujuyana, A.
papudo, A. neuquensis, A. affinis, A. humahwaca, A. riolimayana.
New subspecies: Aegla odebrechtii paulensis, A. laevis
talcahuano.
Suiru, Hopart M. Notes on the snake genus 7'rimorphodon.
Wo."Sis0.” November 10; 9st 722522 ee ee 149-168
New species: Trimorphodon fasciolata, T. forbesi.
New subspecies: Trimorphodon biscutatus quadruplez.
The Mexican subspecies of the snake Conio-
phanes fissidens. No. 8127. November 18, 19411__---_-_- 103-111
New subspecies: Coniophanes fissidens dispersus.
ILLUSTRATIONS
PLATES
Following
1. View of principal fossiliferous exposures of Dragon Paleocene in
Dragon Canyon, Utah__.-____+2-_--1_>- -----~-+---~~__--=~-=..
2. General view of Wagon Road Ridge and Ferron Canyon localities___
3. View over newly discovered Paleocene locality, Dragon Canyon_-__-
4-9, Dinosuchus neivensis, a new fossil crocodile from Colombia__------
10-12. Species. of Arachnis.___._-~-+----=-~----------+---~+-+----+=+----
13-18. New hydroids, mostly from the Atlantic Ocean___-------___-------
19-24. Harly Ordovician sponges__._--_--____----~---------------+~----
2, ected Of ACGU LL 20. oe sate ee ee Se
16.
ane
18.
19.
TEXT FIGURES
. Geologic map of the Dragon Canyon area, Utah_---------------------
. Lower molar portion, occlusal view, of T'aeniolabis; occlusal views of
Catopsalis utahensis, type---=.----=-=-----_----+-----___--—_--==
. Jaw fragment of Ptilodus ferronensis, new species, type-------------
. Left ramus of mandible of Aphronorus simpsoii Gazin, type---------~-
. Maxillary portion of pantolestid (a), genus and species undetermined_-
. Left ramus of mandible of Dracontolestes aphantus, new genus and
SPECS SSE re eee eee ee eee
. Maxillary portion of mixodectid? (b), with one upper molar and part
Ca Feb CIO). EN Geer al te an eee
. Maxillary portion of Conoryciella dragonensis Gazin, type-------------
. Left ramus of mandible of Protogonodon? spiekeri Gazin, type-------
10.
Ade
12:
138.
14,
15.
Left ramus of mandible of Protogonodon biathcles, new species, type---
Left maxillary portion of Protogonodon? sp-------------------------
Right ramus of mandible of Oryclaenus pearcei, new species, type_----
Upper molars of Tricentes elassus, new species______------__--.---__-_
Dentition of Dracoclaenus griphus Gazin_..-=._.- -_--__-=__________
Fragment of left ramus of mandible of Oxytomodon perissum, new
Penns and ‘SPeCles,, LY PCs- == =~ =n eee a ee ee eee
Dentition of Hilipsodon shepherds Gazin__--_=____--=..=_-_-=-_-____.
Portion of right ramus of mandible of Hllipsodon sternbergi Gazin, type-
Right maxillary portion of Jepsenia mantiensis Gazin, type--—_---- =
Left upper dentition and right lower dentition of Desmatoclaenus
hermaeis-new genus. and. species, type-==——--=-—==-=-=-—_=_-___-
. Dentition of Desmatoclaenus paracreodus, new species__-_------------
2a,
22,
23.
Dentition of Hctoconus symbolus, new species____--------------------
Dentition of Carsioptychus hamazitus, new species_--------------------
Right upper dentition of Periptychus gilmorei Gazin, type----------~--
page
Some
89
102
520
Page
10
1
12
13
1h
16
18
19
20
21
25
27
30
40
41
43
Vill PROCEEDINGS OF THE NATIONAL MUSEUM VOL, o4
Page
24, Left ramus of mandible of Periptychus gilmorei Gazin-_-_--__________ 44
25. Left maxillary portion of Anisonchus dracus Gazin, type------_______ 46
26. Portion of left ramus of mandible of Anisonchus onostus Gazin, type____ 4%
27. Left maxillary portion and portion of left ramus of mandible of Anison-
chus oligistus, new. Species; tyWe@==- == ae a ee ee 4T
28. Left maxillary portion of Haploconus inopinatus Gazin, type__-_______ 49
29. Portion of left maxilla of Haploconus? elachistus, new species, type____ 51
30 wSkull of Acipnion: jormosum Cope: a ee eee 72
381. Skull and lower jaw of Aciprion formoswm Cope--_-___---________ 73
32. Anterior part of the skull of Hxostinus serratus Cope_______._. TD
33. Distribution of the Mexican forms of Coniophanes fissidens (map)-. 109
34. Map? showing river systems Of iberiasss2= === eee eee 126-127
35. Cambarus pellucidus australis, new subspecies____--________________ 143
36. CAaMmbaris VCO, Mew (SPECIES 22 Sa aaa es ae asa ee ee eee ee 147
37. Diagram of the possible phylogeny of Trimorphodon____--.-_--____ 151
58. Distribution of the species of Trimorphodon___.__~-~_-+---—_-~..=- 153
ao) DhesregionsoL chevwestern! Hskimom (may) See eee eee 173
40 SDistributionsot-Aicgia (nap) -22 52. a= = eee ee eee ee 441
41. Diagram of Aegia carapace, illustrating some of the terms used in
describing ispeciessa22 Aves Se lee oe ee Ee ee eee sos 447
42, 43. Aegla parana, new species, male holotype_-_---------_----_-_-~____ 459
44. Aegla sanlorenzo, new species, male holotype-------------------___ 462
45. Aegla platensis, new species, male holotype-—--—-----___-__-__-____ 465
46. Aegla platensis, new species, male paratype--------------------__-_ 466
47. Aegla uruguayana, new species, male holotype___---____-_-~-__----___~_ 468
48, 49. Aegla prado, new species, male holotype___—~-__-----_-----_----=- 471
50, Aegia castro; new Species; male holotypel 2 eee 4T4
51. Aegla franca, new species, male holotype——_---—-----=--=-22_L 22 477
52. Aegla jujuyana, new species, male holotype_-_--------_----__--_--__- 479
5s) Aegladenticulata Nicolet, male neotypes= 2-24 ae eee 481
54. Aegla papudo, new species, male holotype-------------------------_ 484
bo. Alegia odebnechis Miller, male neoty pease -s-s eee eee 487
56. Aegla odebrechtii paulensis, new subspecies, male holotype_-----~- 491
57. Aegla neuquensis, new species, male holotype_-—----_------_--_____- 493
58. Aegla ajfinis, new species, male holotype=—=—--_ 2-2 a a Se ee 496
59. Aegla humahuaca, new species, male holotype__------------------_~~ 499
60. Aegla concepcionensis Schmitt, male holotype_---_-_---_---------_---- 501
GE Aegia laevis (hatreille); malemmeoty pena === =— eae ae eee eee 505
62, Aegla laevis talcahuano, new subspecies, male holotype-__----------~- 509
Gay Aeglazaviao Schmitte mialesnoloty eee eee 511
64. Aegla riolimayana, new species, male holotype______------~--------- 514
PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM
THs XR y
TV Te
SHIRA Oa
eonses?!
SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
Vol. 91 Washington: 1941 No. 3121
THE MAMMALIAN FAUNAS OF THE PALEOCENE OF
CENTRAL UTAH, WITH NOTES ON THE GEOLOGY
By C. Lewis Gazin
Fourtuer investigation of the Paleocene deposits of central Utah by
the 1939 and 1940 Smithsonian Institution expeditions has added
considerably to the collections representative of the upper portion of
the North Horn deposits and has resulted in the discovery of a second
and distinct horizon for mammals within the Paleocene series. The
investigations of these years have led also to a better understanding
of the geologic relations pertaining to the fossil-bearing deposits in
and about Dragon Canyon and North Horn Mountain.
The area investigated lies within the region of the Manti National
Forest and along the eastern part of the Wasatch Plateau. Physio-
graphically, it belongs to the High Plateaus of Utah section of the
Colorado Plateaus province, as defined by Fenneman and Johnson.
North Horn Mountain (T. 18 S., R. 6 E.), due west of the towns of
Orangeville and Castledale, is an outlying remnant of the plateau to
the west, being separated from it by the troughlike depression known
as North or Upper Dragon. Dragon Canyon, or the Lower Dragon,
hes principally in the western half of T. 19 S., R. 6 E., and together
with North Dragon is primarily the result of a complex graben struc-
ture extending for a considerable distance both north and south.
The writer wishes to acknowledge the courtesy extended by Dr.
Walter Granger and Dr. G. G. Simpson in permitting him to make
further comparisons with Paleocene materials in the American Mu-
seum of Natural History. The drawings illustrating the specimens
were made by Sydney Prentice.
302662—41——1 1
2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
HISTORY OF THE INVESTIGATION
The occurrence of fossil vertebrates in this region was first recog-
nized in 1935 with the discovery, by Dr. J. B. Reeside, Jr., and Dr.
E. M. Spieker, of the U. S. Geological Survey, of fragmentary dino-
saur remains in exposures around North Horn Mountain and of incom-
plete mammalian remains at a locality high on Wagon Road Ridge
across the Dragon depression, to the west of North Horn Mountain.
These materials were all from beds that had been earlier regarded
as “Wasatch” in geological investigations pertaining to coal resources
of the region.
In 1937 a Smithsonian Institution expedition under the direction of
C. W. Gilmore, and with the aid of Dr. Spieker, made a collection of
dinosaurian remains from the Cretaceous of the region, and was also
successful, through the particular efforts of George B. Pearce, a
member of the party, in discovering a fruitful locality for Paleocene
mammals in lower Dragon Canyon. A popular account of this ex-
pedition by C. W. Gilmore and a description of the Paleocene fossils
by the writer were published in 1938.
During the summer season of 1938 a Smithsonian party under the
writer’s direction further investigated Paleocene and Cretaceous de-
posits and was successful in considerably enlarging the fauna known
from the previously described Dragon Canyon locality. A popular
description of the 1938 expedition and descriptions of the Paleocene
collections by the writer were published in 1939.
The success of the parties in the 1937 and 1938 expeditions, and
at the same time the fragmentary nature of many of the new finds
discovered during these seasons, made it imperative that further work
be done at these localities; hence, the 1939 and 1940 expeditions
undertook more thorough investigations of both the Cretaceous and
Paleocene. Accounts by the writer of the 1939 and 1940 expeditions
were published in 1940 and 1941, respectively.
FAUNAL RELATIONS
Contributory to the more outstanding results of further investiga-
tion of the Paleocene in 1989 was the finding of a new fossiliferous
locality in the upper portion of the North Horn series. The new
locality is in a patch of exposures in the western half of section 7,
T. 19 S., R. 6 E., about a mile nearly due west of the previously
described Dragon Canyon locality, which is in the northwest portion
of section 8. Fossils were found to occur at two levels in the new
locality, the upper of which, though relatively less productive, is
believed to represent the same stage as that at the old Dragon Canyon
locality, the Dragon horizon, as indicated by the occurrence there of
PALEOCENE MAMMALS OF CENTRAL UTAH—GAZIN a
Catopsalis utahensis, Oxyclaenus pearcei, Haploconus inopinatus, and
Ellipsodon cf. shepherdi. The lower level, stratigraphically about 165
feet lower, has produced a new fauna that is more nearly equivalent to
that of the Puerco but may be somewhat younger than the latter.
This lower horizon, which may be known as the Wagonroad stage,
is perhaps 10 or 15 feet above a level that may be arbitrarily defined
as the base of the Paleocene in this region.
Lists of the forms recognized in the two faunas are given below:
DRAGON FAUNA WAGONROAD FAUNA
MOULTITUBERCULATA :
Taeniolabididae :
Catopsalis utahensis Gazin Taeniolabis species
Ptilodontidae:
Ptilodus ferronensis, new species
INSECTIVORA:
Pantolestidae:
Aphronorus simpsoni Gazin
Pantolestid (a), genus and species
undetermined
Pantolestid (b), genus and species
undetermined
Mixodectidae:
Dracontolestes aphantus, new
genus and species
Mixodectid (a), genus and species | Mixodectid? (b), genus and species unde-
undetermined termined
TAENIODONTA :
Stylinodontidae:
Conoryctella dragonensis Gazin
Stylinodont, near Psittacotherium
CARNIVORA:
Arctocyonidae:
Protogonodon? spiekeri Gazin Protogonodon? species
Protogonodon biatheles, new species
Oxyclaenus pearcei, new species Oavyclaenus species
Oxyclaenid
Tricentes elassus, new species
Goniacodon? species
Miacidae:
Didymictis? species
CONDYLARTHRA:
Hyopsodontidae :
Dracoclaenus griphus Gazin
OCxytomodon perissum, new genus
and species
Ellipsodon shepherdi Gazin
Ellipsodon? sternbergi Gazin
Ellipsodon? species (a) Ellipsodon? species (b)
Jepsenia mantiensis Gazin
Phenacodontidae :
Desmatoclaenus hermaeus, new genus and
species
Desmatoclaenus cf. paracreodus Desmatoclaenus paracreodus, new species
Periptychidae :
Ectoconus symbolus, new species
Periptychus gilmorei Gazin Carsioptychus hamavitus, new species
Anisonchus dracus Gazin Anisonchus oligistus, new species
Anisonclus onostus Gazin
Haploconus inopinatus Gazin Haploconus? elachistus, new species
4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
Indicative of an earlier age than that of the Dragon level and
approaching more closely that of the Puerco is the presence in the
Wagonroad fauna of forms representative of the genera Zaeniolabis,
Ectoconus, and Carsioptychus. However, the separation in time of
the two levels in the Dragon Canyon area is not great, as a relation-
ship between the two stages is seen in the materials of Protogonodon?,
Haploconus, and of the new form Desmatoclaenus. 'The Wagonroad
is obviously more nearly comparable to the Puerco stage than it is to
that of the Torrejon.
Reviewing the list of forms now known from the Dragon it would
seem that the fauna was closely related to that of the Torrejon or
Crazy Mountain Fort Union; however, a closer study of the individual
forms in many cases shows them to be less distinctly removed from
related types in the Puerco. This is noticeable in the periptychids,
certain of the carnivores, and most markedly in the taeniodonts, the
latter group apparently having undergone considerable change in at
least two lines during lower Paleocene time. Many of the forms
present, such as the multituberculates and insectivores, can be com-
pared only with later types as ancestral stages of these are
not known in the Puerco. The conclusion is that the Dragon fauna
is intermediate between Puerco and Torrejon faunas in stage of
development, perhaps a trifle closer to the Torrejon, whereas the
Wagonroad fauna is definitely closer, if not equivalent, to that of the
Puerco.
GEOLOGIC RELATIONS
Work during the summer season of 1939 included an investigation
of the geologic relations existing in and around the Dragon in order
to show the distribution of certain formations and to account for
the otherwise anomalous position of many of the fossil localities.
For this purpose a small map has been prepared (fig. 1), using an
enlargement of a portion of the topographic and geologic map of
EK. M. Spieker as a base. The later Cretaceous and Paleocene beds
previously undifferentiated are here distinguished and the distribution
of these together with that of the Flagstaff limestone and later
deposits is more accurately shown. Moreover, a greater refinement
of the fault pattern is indicated.
Stratigraphy.—The older rocks, including the Blackhawk and Price
River formation, and a limited exposure of Star Point sandstone in
Ferron Canyon are all of Cretaceous age and have not been dis-
tinguished on the map. They consist principally of massive buff
sandstones with interbedded clay shale, sandy clay, and coal (in the
lower part), and with a certain amount of conglomeratic material
in the Price River formation.
U. S. NATIONAL MUSEUM FROGEEDINGS,-VOL.391 (PEATE 1
is
we
4, View northwestward of principal fossiliterous exposures of Dragon Paleocene in Dragon Canyon
(loc. 2 in fig. 1 and pl. 2, B), NW% sec. 8, T. 19S., R. 6 E.
B, View northward in northerly pocket of exposure seen in upper photograph. Figure in middk
foreground is approximately at fossiliferous horizon. A large portion of the remains of the
Dragon fauna was found in the small area shown in this view. Caprock of Flagstaff limestone
is seen in right background.
U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 91 PLATE 2
A, General view northward of Wagon Road Ridge locality, near Sanpete-—Emery County line and
probably in sec. 36, T. 18 S., R. 5 E. The first Paleocene materials from this region, though
fragmentary and undeterminable, were discovered at this locality by Drs. Reeside and Spieker
in 1935. Subsequent small collections are indicative of the Dragon horizon.
B, General view northward across Ferron Canyon and up Dragon Canyon, showing the principal
localities for fossil vertebrates, numbered as on the geologic map (fig. 1): (1) Cretaceous expo-
sures at southwest portion of North Horn Mountain, which produced sauropod and ceratopsian
dinosaur remains; (2) principal Dragon Canyon Paleocene locality, Dragon horizon (pl. 1);
(3) Cretaceous exposures in lower part of Dragon Canyon, which produced the fossil lizard col-
lection; (+) new Paleocene locality, with both Dragon and Wagonroad horizons (pl. 3). Original
discovery locality, shown above, is indicated by arrow in left background on Wagon Road Ridge.
PALEOCENE MAMMALS OF CENTRAL UTAH—GAZIN
AW ff Wf
WU
/ YI
/
ys VEN YM
Y YY
ML ae
A] Terrace
| deposits
post-
Flagstaff
Flagstaff
limestone
7) North Horn
Liisi, Paleocene
J North Horn
Cretaceous
Price River,
} Blackhawk
Cretaceous
IN) yn) // f
Y ) if
AV) DHE pe Ry
) We L(x ; mile i
HA ATX wy eM / contour interval = 250 ft.
A » fH ‘
Figure 1.—Geologic map of the Dragon Canyon area, showing principal fossil localities
WY Ni
Li
Or
6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
Overlying the Price River formation, apparently in conformable
relation, is the fossiliferous series of clays, sandy clays, and sand-
stones that have been designated by Spieker as the North Horn forma-
tion. The use of this name should in the opinion of the writer have
been restricted so as to include only the Cretaceous or Paleocene
beds and not both. However, since the U. S. Geological Survey has
adopted the more inclusive definition for North Horn, the name Joes
Valley is proposed as a member to include the Paleocene portion of
the North Horn formation. The clays and sandy clays in the Creta-
ceous portion of the North Horn are varied in color with thick beds
of gray, green, and brown shades of clay with occasional thinner
zones of more reddish clay. Near the top the buff sandstones become
more conspicuous, forming cliffs below the Paleocene deposits.
The Joes Valley member exposed high on the mountain slopes
adjacent to Joes Valley has been more critically observed farther
south on North Horn Mountain, and particularly in Dragon Canyon,
where the Paleocene fossils occur. The member is defined as begin-
ning with the highly colored clay and sandy clay, locally black
carbonaceous shales, resting abruptly but without apparent discon-
formity on the massive sandstones capping the dinosaur-bearing
North Horn beds. The variegated clays of the Paleocene series re-
semble those in the lower portion of the North Horn formation but
are usually not so thick and appear to be more gaudily colored and
with conspicuous white channel sands. The upper portion of the
Joes Valley member, above both of the fossil levels, is not so markedly
variegated and includes a greater quantity of buff sandstone, with
thicker zones of more uniformly colored sandy clay, ending abruptly
beneath the Flagstaff limestone. The thickness of the Joes Valley
member was not measured, but it clearly amounts to several hundred
feet. Apparently, however, it is not so thick as the lower portion of
the North Horn.
The Flagstaff limestone, overlying the Joes Valley member, contains
numerous fresh-water shells, but it has produced no vertebrate re-
mains. Its age is not certainly determined, but it may be within
the limits of the Paleocene. Overlying the limestone in various
places in Dragon Canyon is a series of soft clays that on weathered
surfaces show brick red alternating with much lighter colors. Inter-
bedded with the clay are occasional thin beds of limestone. This
material is designated on the map as post-Flagstaff. No fossils
were found in these beds.
Structure—Dragon Canyon is essentially part of a graben that
extends a considerable distance north and south. The downdropped
block is highly faulted and amounts simply to a zone of faulting in
which the slices are all depressed below the relatively undisturbed
PALEOCENE MAMMALS OF CENTRAL UTAH—GAZIN 7
masses to the east and west. The principal fault along the east
side of the zone has had displacement exceeding 2,000 feet in places,
as indicated by the extent to which the Flagstaff limestone has been
depressed. To the west across Dragon Canyon this displacement has
been taken up along three principal surfaces of faulting, but with minor
fractures along which displacement has been in an opposite direction.
Throughout most of the region the rocks are nearly level lying,
but within the depressed zone the sediments are noticeably dis-
turbed, particularly adjacent to the faults, where strong drag folding
was observed. Certain of the slices, particularly the most easterly
block, are depressed northward, and this together with the effect of
drag along the bounding faults has in these cases resulted in an
average northeasterly dip to the various deposits. The slice on which
localities 2 and 3 are shown has been raised relative to both blocks
immediately adjacent; hence the sediments are more nearly level, but
with a noticeable downward drag adjacent to the westerly fault in
the vicinity of locality 3. On the other hand, a very strong upward
drag is apparent along the westerly margin of the two westerly slices,
near locality 4.
Fossil. localities—Four localities have been indicated on the map.
These show the general location of the principal occurrences of fossil
vertebrates with the exception of a locality for Paleocene mammals
on Wagon Road Ridge some distance to the north of the area shown
on the map, and of several sites around North Horn Mountain, which
cannot be shown on the map, from which dinosaur remains have been
recovered.
Those that have been indicated are as follows: (1) A locality in
Cretaceous rock on North Horn Mountain where the greater part
of a sauropod dinosaur was discovered in 1937, near the line between
sections:3 and 4, T. 19 S., R. 6 E. (2) The original Paleocene
locality in Dragon Canyon from which most of the Dragon collection
was obtained; NW}, sec. 8, T. 19 S., R. 6 E. (3) A Cretaceous
locality in the lower part of Dragon Canyon, which produced the
unique fossil lizard collection; S14 sec. 17, T. 19 S., R. 6 E. (4)
The new Paleocene locality where mammalian fossils were discovered
at two distinct levels; W% sec. 7, T.19S., R.6 E.
8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
SYSTEMATIC DESCRIPTION OF THE MATERIAL
MULTITUBERCULATA
Genus TAENIOLABIS Cope
TAENIOLABIS species
The genus Zaenzolabis is apparently represented in the collection
from the Wagonroad horizon by the posterior half of a first lower
molar, U.S.N.M. No. 16172 (fig. 2, a). In size and appearance the speci-
men closely resembles this portion of M, in Taeniolabis tadensis from
the Puerco of New Mexico. The form present in the Wagonroad hori-
zon may represent this species, but in the absence of better material,
showing at least something of the cusp formula, no specific reference
is made.
Although our knowledge of the history or development of the
Taeniolabididae is very incomplete, the presence of T'aeniolabis and
the absence of Catopsalis in the Wagonroad fauna are significant in
indicating a relationship to the Puercan stage.
In the structure of the molars Catopsalis would appear to be an-
cestral to Taeniolabis, but since their known positions in time are
the reverse the two must be regarded as representing separate phyla,
and that having the less specialized molars surviving here longer, or
reaching this region at a later date.
Genus CATOPSALIS Cope
CATOPSALIS UTAHENSIS Gazin
Catopsalis utahensis GAzIn, 1939b, p. 275.
The type of Catopsalis utahensis, U.S.N.M. No. 15757, frotn the
Dragon horizon, as represented at the principal Dragon Canyon locality
(loc. 2 in fig. 1), consists of a single first lower molar (fig. 2, b).
The specimen exhibits the simple type of pattern seen in Catopsalis
from the Torrejon rather than the more specialized dental structure of
the Puerco Zaeniolabis. It differs from M, in specimens of Catopsalis
known from the Torrejon of the San Juan Basin in having the cusp
formula 6:4. In the type of Catopsalis foliatus it is 5:4, and in the
type of C. fissidens the formula is 6:5, or better. Moreover, the
tooth is relatively wider than in either of the Torrejon specimens.
Catopsalis calgariensis from the Paskapoo was described by Russell
from a second lower molar; hence no satisfactory comparison with
the type of C. wtahensis is possible.
From additional material of this form collected in 1939 it is seen
that the lower molars are distinctly wider than in either C. fissedens
or @. foliatus. In an M, (fig. 2, ¢), No. 16185, from the upper or
Dragon horizon at the new locality (loc. 4 in fig. 1), slightly more
PEATE 3
PROCEEDINGS, VOL. 91
NATIONAL MUSEUM
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PALEOCENE MAMMALS OF CENTRAL UTAH—GAZIN 9
worn than the type, the posterointernal cusp is further divided for a
part of its height so that the inner row has five cusps instead of four.
Wear has obscured the posterior portion of the outer row so that
it is uncertain as to whether there were five or six cusps, and the
formula may be 5:5 or 6:5. The tooth is slightly larger than the type
of C. utahensis.
The posterior portion of another M,, No. 16211, shows a cusp divi-
sion suggestive of the formula 7:5 or possibly 6:5. The latter
tooth portion is about the size of the type and comes from the original
Dragon Canyon locality.
An incomplete tooth portion, No. 16210, which has only four cusps
preserved, is relatively large and may be the anterior portion of M,,
in which case it approaches in size small specimens of Z'aeniolabis.
However, it may be the posterior portion of an M, of C. utahensis.
ty S
mAs iS
Ficure 2.—a, Taentolabis sp., lower molar portion (U.S.N.M. No. 16172), occlusal view,
Wagonroad Paleocene, Utah; b, Catopsalis utahensis Gazin, M, (U.S.N.M. No. 15757),
type specimen, occlusal view, Dragon Paleocene, Utah, c, C. utahensis, M; (U.S.N.M.
No. 16185), occlusal view, Dragon Paleocene, Utah. All X 2.
A right lower jaw, No. 16209, in the Dragon collection has both
M, and M, but unfortunately the teeth are checked and partially
obscured by an ironlike matrix.
Material of Catopsalis is particularly rare, there being but about
three known specimens outside of the material herein described, and
one of these, an M,, the type of Catopsalis calgariensis from the
Paskapoo, has been lost, although a cast of it is in the collections
of the American Museum of Natural History. The other two, the
types of C. foliatus and C. fissidens, are lower dentitions from the
Torrejon. The material of C. utahensis though more than doubling
the number of specimens representing Catopsalis does not seem to
present any significant evidence as to the ancestral stages in the
development of this genus. It is interesting to note, however, that
C. utahensis, especially as represented by No. 16185 and No. 16210,
appears somewhat less distinctly removed from 7aeniolabis than do
the Torrejon forms.
The anteroposterior and transverse diameters of the type, No.
15757, are 12 (approximately) and 6.5 mm., respectively. In No.
16185 these diameters are 13 and 7.3 mm., respectively.
302662—4 12
10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
Genus PTILODUS Cope
PTILODUS FERRONENSIS,! new species
Type—Fragment of right ramus of mandible with P,, U.S.N.M.
No. 16176.
Horizon and locality—Dvragon Paleocene, Dragon Canyon, Emery
County, Utah.
Specific characters—Near Ptilodus mediaevus in size. P, in type
longer, with crest less elevated posteriorly. About 12 serrations, as
indicated by ridges on lateral surface of tooth. Notch between an-
terior and posterior roots not so acute and buccal wall of crown not
extending down root portion so far. P* in referred material rela-
tively shorter and wider and P? slightly
wider than in P. mediaevus. Cusps in P?
and P? less elevated and less distinct.
Outer row of cusps on referred M1? less
developed posteriorly.
Description.—Included in the material
representing Ptilodus ferronensis are five
lower jaw fragments with P,, a maxillary
fragment with P* and P?, and an incomplete,
isolated M*. P, in No. 16176 (fig. 3), the
type of P. ferronensis, is a little longer than
in Ptilodus mediaevus and has the posterior
Ficure 3.—Pulodus ferro- portion of the crest a little less elevated.
pe! new species: Jaw "he notch between the anterior and posterior
ragment with P, (U.S.N. B
M. No. 16176), type speci- Toots is not so acute, as viewed from the
men, fateral and occlusal Outer surface, and the buccal wall of the
views, X 3, Dragon Paleo- tooth does not extend so far down on the
cene, Utah. roots in the type. ‘The notch between the
roots of P, in No. 16225, referred to P. ferronensis, does not appear
to be so obtuse. The number of serrations on the crown of P, in the
type is about 12, as indicated in part by the ridges on the lateral sur-
face of the tooth, apparently less by a similar method of counting
than in certain specimens of P. mediaevus examined, although 12 is
the median figure given by Simpson for the Torrejon form.
Pt and P? in No. 16212 compare favorably in size with Ptilodus
mediaevus (Amer. Mus. Nat. Hist. Nos. 3033 and 16533), but P? is
relatively shorter and wider than in the Torrejon material, and P?,
though incomplete posteriorly, is a little wider than in Amer. Mus.
Nat. Hist. No. 3033. The cusps of these two teeth in the Utah speci-
men are not so markedly separated and are less elevated than in
the Torrejon material.
1 Named from Ferron Canyon in Emery and Sanpete Counties, Utah.
PALEOCENE MAMMALS OF CENTRAL UTAH—GAZIN Hl
An incomplete M? in the collection, No. 16216, shows the outer row
of cusps less developed posteriorly than in Amer. Mus. Nat. Hist.
No. 3033 from the Torrejon.
The length of P, in the type, No. 16176, of Ptilodus ferronensis
is 9mm. In No. 16212 P is 3.3 mm. long and 2.8 wide, and P? is
3.5 mm. wide.
INSECTIVORA
Genus APHRONORUS Simpson
APHRONORUS SIMPSONI? Gazin
Aphronorus simpsoni GAZIN, 1988, p. 2738.
About 19 specimens, consisting of isolated teeth or jaw fragments
with one to four teeth, from the Dragon level are considered to
represent, Aphronorus. All but
three of these, upper premolars,
are lower jaw remains. The up-
per molar earlier (Gazin, 1939b,
p. 275) thought to be of Aphro-
norus simpsoni is now cited here-
in as pantolestid (b).
Aphronorus simpsoni is close
in size to A. fraudator from the
Crazy Mountain Fort Union but
differs from this species in cer-
tain relative proportions, which
are outside the limits given by
Simpson for the middle Paleo- Figure 4.—Aphronorus simpsoni Gazin:
ne Left ramus of mandible with Py-M3 (U.
cene form. The ramus, No. 15539 S.N.M. No. 15539), type specimen, later-
(fig. eae made the type, is slight- al and occlusal views, X 4, Dragon Pa-
ly deeper than in the several leocene, Utah.
Fort Union specimens that the
writer examined, a difference more noticeable in the posterior
portion. Also, the posterior molars are relatively larger, par-
ticularly M;, which is larger than in any of the Fort Union
specimens examined. However, the teeth are relatively slender.
This is most noticeable in P,, which combines the greatest length
with the least width given by Simpson for A. fraudator. Moreover,
the posterior wall or shear of the trigonid in the molars is not so
distinctly transverse, but directed slightly more forward externally.
In P, the shear is more nearly transverse though somewhat irregular
as a slight ridge extends down the posterior wall of the metaconid
and unites with the hypoconid crest.
2 Named for Dr. G. G. Simpson.
12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
TABLE 1.— Measurements (in millimeters) of lower teeth of Aphronorus simpsoni
Measurement Py M, M, M;
Anteroposterior diameter)/2U_ 220! J isu Se ee = 3.8 | 3.0 3.0 3.2
Transverse diameters. 62 2) as ee ee se eee 2.0 Dail Delt De
Pantolestid (a), genus and species undetermined
A maxillary portion (fig. 5), No. 16184, with M? and M®, represents
a pantolestid insectivore near Bessoecetor. The teeth are relatively
wide transversely, M? being about one-fourth wider than in Bes-
soecetor thomsoni. Anteroposteriorly the tooth is about the same, or
possibly as much as a sixth greater than in B. thomsoni. The hypo-
cone is markedly lingual in position and the anteroexternal angle,
though partially broken away, is seen to be much heavier than in M?
of the Scarritt Quarry form. The anterior wall of M’ shows a
somewhat heavier cingulum and the posterior wall does not show
so acute a notch adjacent to the metaconule. M®,
though poorly preserved, appears to be anteropos-
teriorly compressed. Both teeth are much more re-
duced anteroposteriorly than in Aphronorus, and
the external styles are directed more as in Bessoe-
cetor. Moreover, the teeth are much smaller than in
Ficure 5.—Panto- Palaeosinopa senior, also recorded from the Scarritt
Oe wees Quarry in the Crazy Mountain field of Montana.
Maxillary por- & lower jaw portion, No. 16219, with M; pre-
tion with M2 and served may belong to this form. M, has the trigonid
M? (U.S.N.M. structure much as in Bessoecetor, or even Aphro-
No. 16184), oc- »oprys, but the talonid is more reduced than in B.
Beat oes diluculi, somewhat as in B. thomsom. However, its
cone eat size is such as to suggest a relationship to the form
represented by the upper molars described above,
and the reduced talonid is quite in accord with the anteroposterior
reduction of M® in No. 16184.
The anteroposterior and greatest transverse diameters of M?, No.
16184, are 2.5 (approximately) and 4.9 mm., respectively; of Ms, No.
16219, 2.7 and 2.0 mm.
Pantolestid (b), genus and species undetermined
A single upper molar, No. 15791, provisionally referred to Aphro-
norus simpsoni, seems on further consideration to represent not
Aphronorus but a pantolestid type closer to Bessoecetor. The tooth
is about a third smaller than the M? in the form herein described as
pantolestid (a), hence somewhat smaller than either of the first
PALEOCENE MAMMALS OF CENTRAL UTAH—GAZIN ie
two molars in B. thomsoni or in B. diluculi. However, this tooth
more closely resembles Bessoecetor in its proportions and outline than
it does the larger Dragon pantolestid (a).
DRACONTOLESTES,; new genus
Type—Dracontolestes aphantus, new species.
Generic characters.—Lingual cusps of lower molars slightly more
elevated than outer. Trigonid moderately elevated. Paraconid crest
extends to a markedly lingual point. Talonid basin closed lingually.
Crest extending forward from hypoconid joins trigonid at a distinctly
lateral point. No external cingulum on M, and M3.
DRACONTOLESTES APHANTUS,‘ new species
Type.—Leit ramus of mandible, U.S.N.M. No. 16180, with M; and
part of M,.
Horizon and locality —Dragon Paleocene, Dragon Canyon, Emery
County, Utah.
Specific characters—Much smaller
than known species of Hudaemonema
and Llpidophorus. Teeth about the
size of those in Aphronorus simpsont.
Description—The lower jaw with
M; and part of M., No. 16180 (fig. 6),
represents a species in the Dragon
level which is clearly mixodectid, but
cannot be certainly referred to any
of the known genera. The form is
much smaller than species of Hudae-
monema and Elpidophorus, being dis- Ficure 6.—Dracontolestes aphantus,
tinctly smaller than Llpidophorus mi- new genus and species: Left ramus
nor from the Crazy Mountain Fort of mandible with M2-Ms (U.S.N.M.
WON No. 16180), type specimen, lateral
i : 3 and occlusal views, 4, Dragon
The inner cusps are only slightly _pajeocene, Utah.
more elevated than the outer, such as
observed in some material of Hudaemonema cuspidata, not so
accentuated in this respect as in Hlpidophorus. ‘The trigonid por-
tion is elevated with respect to the talonid but the cusps in general,
though sharp, are not so elevated as in Hudaemonema cuspidata.
The crest carrying the paraconid extends lingually even more than
in Elpidophorus patratus somewhat as in Elpidophorus minor,
not extending forward or so median in position as characteristic of
Eudaemonema. Moreover, the talonid basin is closed lingually by
§ dpaxwy, dragon + Agoris, thief.
4&pavros, obscure.
14 PROCEEDINGS OF THE NATIONAL MUSEUM YOU, 91
the crest extending forward from the entoconid to the metaconid,
and the crest forward from the hypoconid joins the posterior wall
of the trigonid at a point distinctly more external than in either of
the above genera. The talonid basin is well excavated and in M;
is not so restricted by the fiexure of the outer wall anterior to the
hypoconid as in Elpidophorus patratus. The hypoconulid in
M.,, though weak, is placed almost as close to the entoconid as in the
Crazy Mountain forms. It may be further noted that the two
molars do not show evidence of an external cingulum such as exists
in Elpidophorus material.
The anteroposterior diameter of M; in No. 16180 is 3.5 mm. The
transverse diameters of M, and M; are 2.3 and 2.0 mm., respectively.
This new form is possibly closest to the Hlpidophorus line but
differs most notably in the less accentuated elevation of the inner
cusps and in the more widely basined talonids. The differences from
Fudaemonema that are significant, though not striking, in determin-
ing the relationship of this form lie principally in the position of
the paraconid and in the distinctly closed talonid basins. The lateral
position of the crest joming the hypoconid with the trigonid wall
is distinctive with respect to both.
Specimen No. 15719, which includes an incomplete lower molar,
earlier described (Gazin, 1939b, p. 276) as belonging possibly to a
primate, closely resembles M, in the above described type, so that
in the absence of additional material demonstrating more certainly
the presence of a primate in the fauna this specimen is referred to
Dracontolestes aphantus.
Mixodectid (a), genus and species undetermined
A jaw fragment, No. 16220, with a single molar is seen to rep-
resent a second mixodectid type of insectivore in the Dragon fauna.
The tooth is almost as large as in Hudaemonema cuspidata and appar-
ently a little larger than in ELlpidophorus minor. The protoconid
and metaconid are broken, and although the inner of the two may
possibly have been the larger, in the talonid the entoconid is not
higher than the hypoconid, suggesting Hudaemonema rather than
Elpidophorus, and the talonid basin opens internally with almost,
but not quite, as broad an opening as in specimens of Hudaemonema.
The tooth also lacks the distinct external cingulum seen in material
of Elpidophorus. Wowever, the paraconid is markedly internal in
position, and not so low or projecting so forward as in Hudaemo-
nema cuspidata. The paraconid is placed somewhat as appears
to be the case in M, of Elpidophorus minor. The tooth, though a
little shorter, is relatively wider than in Hudaemonema cuspidata,
suggesting Hpidophorus in this respect, but is slightly lower crowned
than in either. It may be further noted that the hypoconulid, rising
PALEOCENE MAMMALS OF CENTRAL UTAH—GAZIN 15
from a slight posterior cingulum, does not appear to be placed quite
so far internally, and the outer walls of the protoconid and hypo-
conid are not so nearly vertical as in Hudaemonema and Elpidophorus,
but seem to be more sloping, causing at least the talonid basin to
appear slightly narrower with respect to the width of the tooth.
The anteroposterior and transverse diameters of the lower molar,
No. 16220, are 3.4 and 2.9 mm., respectively.
Mixodectid ? (b), genus and species undetermined
A maxillary portion, No. 16200, with an upper molar, possibly
M? (fig. 7), and part of the next succeeding tooth may represent a
small mixodectid in the Wagonroad fauna. The molar shows a
well-developed shelflike cingulum external to the par-
acone and metacone and acute external styles. The
hypocone is markedly lingual in position and a cin-
gulum is continuous around the inner portion of the
protocone, not including the hypocone but appar-
ently terminating posteriorly and upward between
the protocone and hypocone. A posterior cingulum
extends laterally from the hypocone. The lingual poupe 7.—Mixo-
position of the hypocone suggests a relationship to — dectid? (b): Max-
Huda: monema, inasmuch as in £/pidophorus the hy- __ illaryportionwith
pocone is not placed so far inward. The cingular shelf ©? Upper molar
on the outer side of the tooth seems more prominent PR ARONA
than in either of the Crazy Mountain forms. No. 16200), oc-
The occurrence of this small form in the Wagon- clusal view, X 4,
road fauna is of interest, being unlike anything in = Wagonroad_ Pa-
the Puerco and if found to represent a mixodectid — eocene Utah.
it is the earliest known.
The tooth measures about 3.3 and 5.4 mm., anteroposteriorly and
transversely.
TAENIODONTA
Genus CONORYCTELLA * Gazin
CONORYCTELLA DRAGONENSIS ¢ Gazin
Conoryctella dragonensis GAZIN, 1939b, p. 276.
A conoryctid type of taeniodont is recognized in the Dragon collec-
tions by a maxillary portion with three teeth, P* to M’, and a
lower jaw fragment with a single molar obtained in 1938, and two
additional lower molars found in 1939.
The upper teeth, No. 15704, made the type of Conoryctella dragon-
ensis (fig. 8), are seen, as previously described, to be a little smaller
5 Conoryctes+ ella, a small conoryetid.
® Named for Dragon Canyon.
16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
than in Conoryctes comma but distinctly larger than in Onychodectes
tisonensis. The Dragon form is about intermediate between these two
species in degree of hypsodonty. P* is not so nearly molariform
as in C. comma and has the lingual portion more compressed antero-
posteriorly. The protocone and deuterocone are prominent conical
cusps, and the tritocone, though damaged, is seen to be but weakly
developed as compared to the other two cusps. The lingual portion
of this tooth does not appear crescentic; nevertheless, a low crest
or cingulum extends along the posterior portion between the deutero-
cone and tritocone.
The paracone and metacone in the first two molars, as far as pre-
served, are seen to be conical and low and are separated from the
outer margin of the teeth by
a heavy cingulum. The meso-
style, though present, is not so
strongly developed as in @.
comma. It is absent in O. téso-
nensis, The anteroexternal and
posteroexternal angles of the
teeth are more rounded than in
O. tisonensis and do not exhibit
styles at these points as in the
Puerco form.
The anteroposterior diam-
eters of the upper teeth, P* to
M?, are approximately 7.5, 8.2,
and 7.4 mm., respectively. Any
Ficure 8.—Conoryctella dragonensis Gazin: transverse measurements would
Maxillary portion with P4-M? (U.S.N.M. ba highly aubineae
No. 15704), type specimen, lateral and oc- Buy 3 Me
clusal views, X 2, Dragon Paleocene, Utah. The lower Jaw fragment, No.
15722, with a molar tooth, ap-
parently M,, may represent Conoryctella dragonensis, although it is
from an individual somewhat smaller than the type. The tooth is about
intermediate between O. tisonensis and C. comma in hypsodonty but
apparently a little nearer O. tisonensis in size. The trigonid of the
tooth possesses a moderately developed paraconid situated much as
in M, of QO. tisonensis. 'The heel or talonid, though partially ob-
scured by matrix, is relatively broad, appears to be deeply basined
and to have a somewhat cuspidate crest, approaching the condition
seen in C. comma.
The two lower molars, No. 16173, added to the collection in 1939,
exhibit an arrangement of the cusps around the margin of the talonid
very much as in Onychodectes, without the greater number of acces-
sory cuspules seen in Conoryctes. The teeth are relatively a little
PALEOCENE MAMMALS OF CENTRAL UTAH:
GAZIN LZ
wider and the heel more basined than in Onychodectes and as with
other material known of the form the two teeth are intermediate be-
tween O. tisonensis and C. comma in size and hypsodonty.
The Dragon lower teeth do not exhibit the basal accessory cuspule
anteroexternal to the hypocone characterizing Onychodectes rarus.
Stylinodont, near Psittacotherium
A single incisor tooth, No. 16204, apparently lower, seems most
certainly to belong to a stylinodont type of taeniodont. The tooth
is moderately worn but shows evidence of a conical labial portion
and a marked lingual shelf, and exhibits a heavy, transversely flattened
root. The tooth is about intermediate in size between corresponding
teeth in the types of the Puerco and Torrejon species, Wortmania
otariidens and Psittacotherium multifragum. 'The lingual shelf seems
more extended than in Wortmania but is not so prominent or so
broadened as in Pstttacotheriwm, and the enamel does not extend
down the labial wall of the tooth for so great a distance as in the
latter genus.
The occurrence of a stylinodont in the Dragon fauna was to be
expected since this family is represented in both the Puerco and
Torrejon stages; in fact the line appears to be continuous through the
Paleocene, and into Eocene time where it is represented by the
genera E'ctoganus and Stylinodon.
CARNIVORA
Genus PROTOGONODON Scott
PROTOGONODON? SPIEKERI7™ Gazin
Protogonodon? spiekeri GAzIN, 1988, p. 274.
The species Protogonodon? spiekert was described from a right
lower jaw portion with M,, M., and part of M; in the Dragon collec-
tion obtained in 1937. Subsequent material includes a lower jaw
portion with M, and isolated portions of lower molars. Upper
jaw material, including an M? and a maxillary portion with part of
M®* and the root portion of M?, was referred to this species, but the
recognition of a second species, Protogonodon biatheles, from lower-
jaw material obtained from the Dragon horizon in 1939 makes doubt-
ful the reference of these upper teeth to P.? spiekeri, in the absence
of any association between upper and lower teeth.
The lower molars of Protogonodon? spiekeri, as represented by the
type, No. 15538 (fig. 9), correspond closely in size to those of P.
7 Named for Dr. Edmund M., Spieker,
302662—41——_3
18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
pentacus from the Puerco but exhibit more rugose enamel. The para-
conid, which is preserved in only the first two molars, is more lingual
in position and not so distinct from the metaconid. The cusps around
the talonid, however, though low, are somewhat more distinct from
those adjacent than in P. pentacus, with less development of a crest
and basin. The trigonid portions of the teeth are somewhat more
elevated with respect to the talonids than is usual in P. pentacus.
Tn the reduction and position of the paraconid and in the rugosity
of the enamel the Dragon form makes a definite approach toward the
condition seen in the Torrejon specimens referred to Claenodon cor-
rugatus (C. ferow). The paraconid in M,, and perhaps M,, of P.?
spickeri is better developed and more distinctly separated from the
metaconid than in C. corrugatus although it is placed nearly as far
Ficure 9.—Protogonodon? spiekert Gazin: Right ramus of mandible with My, Mg, and
part of M; (U.S.N.M. No. 15538), type specimen, lateral and occlusal views, X 114,
Dragon Paleocene, Utah.
lingually as in the Torrejon material. The union or ridge between
the protoconid and metaconid is simple and not double as frequently
seen in the more coarsely rugose teeth of Claenodon corrugatus. On
the talonid the hypoconulid is more distinct from the entoconid,
whereas in (. corrugatus these two form a more conspicuous ridge,
which usually continues with the cingulum around the hypoconid.
The cusps in general are lower and more distinct than in Claenodon,
with a less distinctly basined talonid, with fewer accessory cuspules,
and a finer quality of rugosity.
M, in the type, though incomplete, is much less elongate than in
CO. corrugatus, as indicated by the spacing of the metaconid, entoconid,
and hypoconulid.
PALEOCENE MAMMALS OF CENTRAL UTAH—GAZIN 19
The maxillary fragment, No. 15541, tentatively referred to Proto-
gonodon? spiekeri, shows no important characters other than a rela-
tively great difference in size between M? and M°. The isolated M?
is complete and shows a slight development of a mesostyle, not
nearly so prominent, however, as in Deuterogonodon montanus, and
the slight hypocone is not nearly so lingual in position.
In most respects, especially in the character of the trigonid of the
lower molars, P.? spiekeri stands in a relation nearly intermediate
between Protogonodon and Claenodon, with perhaps a slightly greater
resemblance to Protogonodon. It is distinct from Deuterogonodon
montanus, as represented by the paratype, in the lowness of the cusps,
the far less developed crest and basin of the talonid, and in the
relatively greater importance of the entoconid.
The anteroposterior diameters of the first and second lower molars
are 10 and 11 mm., respectively. The transverse diameters are 8 and
9.3 mm.
PROTOGONODON BIATHELES, new species
Type.—Portions of both rami of the mandible with M, and M,,
U.S.N.M. No. 16181.
Horizon and locality——Dragon Paleocene, Dragon Canyon, Emery
County, Utah.
Specific characters——M, and M, slightly larger than Protogono-
don? spiekert. Paraconid median in position. Talonid relatively
wide. Teeth slightly rugose.
Description —¥ragments of both rami of the mandible, No. 16181
(fig. 10), with M, and M,, found in a mass of barite crystals to-
gether with well-worn upper teeth of Des-
matoclaenus paracreodus in the Dragon hori-
zon, appear to represent a species of Proto-
gonodon distinct from P.? spiekeri. The
molars are only slightly larger than those in
P.? spiekeri, but in contrast with this form
the paraconid is much more median in posi-
tion, even in comparison with Protogonodon
pentacus. The trigonid portion is relatively
narrow, and the talonid, especially of M., io idee ern eee
is markedly wider and more basined thanin ons of mandible with
either P.? spiekert or P. pentacus. This M, and M2 (U.S.N.M.
specialization is directly opposite to that seen —No. 16181), type specimen,
in Protogonodon kimbetovius where the tal- _!8teral and occlusal views,
a : < 1%, Dragon Paleocene,
onid is relatively narrow. The enamel of Utah.
the teeth is very slightly rugose, much less so
in the talonid basin in comparison with P.? spiekeri, although the
teeth appear to be about as unworn as in the type of the latter.
Frcoure 10.— Protogonodon
20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
Considerable doubt attaches to the assignment of any upper teeth
to this species. Those tentatively assigned to P.? spickeri may be-
long to P. biatheles; however, the reduced size of M; suggested in
the type of P.? spiekeri indicates allocation of the preserved third
upper molars to that species.
PROTOGONODON? species
A maxillary portion, No. 16193 (fig. 11), including M® and a
much damaged M?, in the Wagonroad collection strongly resembles
material in the Dragon collections referred to Protogonodon? spiek-
ert. M® is rounded and the cingulum, which
appears to extend entirely around the tooth, is
rugose, whereas the central basin is smooth.
Arising from the cingulum is a hypocone about
as in the Dragon M®* but between the paracone
and metacone and separate from the cingulum a
Ficure 11.—Protogon- :
ee ee aire. much-worn accessory cuspule or mesostyle is de-
llary portion with Veloped to an extent approaching that in Deu-
M3 and part of M2. terogonodon montanus. In M?, No. 15733, re-
(U.S.N.M. No. ferred to P.? spiekert from the Dragon horizon,
ee tes cea there is a slight cuspule in this position.
atte . H ee The anteroposterior and transverse diameters
Utah. of M°, No. 16193, are about 9.5 and 10.5 mm.,
respectively.
Other incomplete portions of teeth in the collections from the
Wagonroad horizon probably represent the same form as No. 16193,
or a closely related type. All show evidence of a moderately heavy
cingulum but none of the upper tooth fragments exhibit a mesostyle
as in No. 16193.
A single last lower molar, No. 16344, in the small collection from
the original Wagon Road Ridge locality (the equivalence of which
to either the Wagonroad or Dragon horizons is uncertain) may repre-
sent a species of Protogonodon. The elevation of the trigonid sug-
gests Hoconodon but differs from that form in having the paraconid so
nearly median in position.
Genus OXYCLAENUS Cope
OXYCLAENUS PEARCEI,§ new species
Type—Portions of right and left rami of the mandible with M,
and M,, U.S.N.M. No. 16186.
Horizon and locality—Dragon Paleocene, Dragon Canyon, Emery
County, Utah.
8 Named for Franklin Pearce, in recognition of his field assistance.
PALEOCENE MAMMALS OF CENTRAL UTAH—GAZIN Zi
Specific characters—Size near Oxyclaenus simplex. Talonid of
M, relatively wide. Paraconid directed forward and more distinct
from protoconid and metaconid. M, unreduced.
Description.—Several lower jaw fragments from the Dragon hori-
zon represent a species of Oxyclaenus near O. simplex. M, in the
type specimen, No. 16186 (fig. 12), from the upper or Dragon level
at the new locality in the western part of the canyon is about the
same size as the single lower
molar belonging with the type
of O. simplex, being smaller and
not so high crowned as in Oay-
claenus cuspidatus. It differs
from O. simplex principally in
having a wider talonid portion
and a narrower trigonid, some-
what as in Lowolophus but with
the talonid basin more open in-
ternally; however, the teeth are
relatively slender and exhibit a
well-defined external cingulum as
in Oxyclaenus. The paraconid is
directed more forward than in
Oxyclaenus and separated from :
both the protoconid and metaco- Ficurr 12.—Oxyclaenus pearcei, new spe-
nid by a more distinct notch. cies: Right ramus of mandible with
M; in the type exhibits a trigo- eee ee
saa s StyP . Se specimen, lateral and occlusal views;
nid portion much as in M,, but X 3, Dragon Paleocene, Utah.
the tooth is fully as large as M,,
not showing the reduction seen in Puerco specimens referred to O.
stmplew (Amer. Mus. Nat. Hist. No. 16347) and O. cuspidatus (Amer.
Mus. Nat. Hist. No. 16346).
An upper molar fragment, No. 15736, which includes only the in-
ner portion may represent this form, and is characterized by a
prominent lingually placed hypocone and an equivalent protostyle
symmetrically placed.
The anteroposterior diameters of M, and M; in No. 16186 are
5.7 and 6.0 mm., respectively. The transverse diameters are 4.1 and
3.5 mm.
OXYCLAENUS species
A single upper molar, No. 16217, in the material from the Wagon-
road level, is seen to correspond closely to M* in the type of Oxy-
claenus simplew and may possibly represent O. pearcei, the species
described from the Dragon horizon. The tooth differs from M? of
O. simplex only in being slightly narrower transversely and in hav-
22 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
ing cusps, which appear to be somewhat more acute, although this
tooth in the type of O. simplex is rather well worn. The tooth,
No. 16217, measures 4.8 mm. anteroposteriorly across the styles and
5.2 mm. transversely.
Oxyclaenid?
An isolated upper molar, possibly M?, No. 15546, in the 1937 col-
lection from the Dragon level, may be from an oxyclaenid type of
carnivore. The tooth is too large to belong to Owyclaenus pearcet
and differs somewhat from the Oxyclaenus type of tooth. Although
exhibiting a parastyle, the external angles are not so acute as in
either Oxyclaenus or Chriacus. 'The hypocone is more lingual than
in Oxyclaenus and a slight protostyle is present at the lingual ex-
tremity of the anterior cingulum. The hypocone, however, is not
developed as in Chriacus, the cusps in general are more nearly
conical, and the cingulum does not extend entirely across the lingual
wall of the protocone. Moreover, the protoconule and metaconule
are more distinctly separated from the outer cusps than in any of
the oxyclaenid material examined.
Some resemblance is seen between this tooth and M? in the con-
dylarth Dracoclaenus griphus, with which it corresponds closely in
size, but there is no mesostyle, the hypocone is more lingual in posi-
tion, there is a slight protostyle, and, as in comparison with the
oxyclaenids, the protoconule and metaconule are too widely separated
from the paracone and metacone, respectively.
The anteroposterior diameter of the tooth is about 6.2 mm. and
the transverse diameter 7.6 mm.
Genus TRICENTES Cope
TRICENTES ELASSUS,® new species
Type.—Upper molar, M1, U.S.N.M. No. 16178.
Horizon and locality—Dragon Paleocene, Dragon Canyon, Emery
County, Utah.
Specific characters.—A little smaller than T’ricentes subtrigonus.
Cusps and outer angles of upper molars somewhat more acute. Cingu-
lum does not extend around lingual wall of protocone on M’.
Description —At least three isolated upper molars and a lower
molar in the Dragon collection are recognized as belonging to 7’7v-
centes. The upper molars are a little smaller than in material referred
to Tricentes crassicolidens and about a fifth smaller than in, the type
of Tricentes subtrigonus; however, certain specimens from the Tor-
rejon are nearly as small as the Dragon form. The outer angles of
the upper molars are somewhat more acute, and the cusps in general
§’eXaoowv, small, in allusion to its size.
PALEOCENE MAMMALS OF CENTRAL UTAH—GAZIN 23
have a weaker, less inflated appearance. The posterior portion of
the external cingulum of M’, Nos. 16178 (fig. 13) and 15783, rises
forward on the protocone much as in the Torrejon material of
Tricentes, but the inner cingulum does not extend around the pro-
tocone as is common, though not invari-
able, in Tricentes subtrigonus. In M’,
No. 16179 (fig. 13), the cingulum appears
to be continuous around the protecone.
The enamel is weakly rugose on both M+
and M?, but there is no indication of a
mesostyle on the cingulum or between the Ficure 13.—Tricentes elassus,
paracone and metacone on these teeth. new species: M? (U.S.N.M.
A maxillary portion with M® and an in- No. 16178), poe pence fon
complete M*, No. 16206, may represent ine ieee -
Tricentes elassus. The teeth are a little Brasin Dace f
smaller than in 7’. subtrigonus but other-
wise show no importance differences. The enamel is somewhat more
smooth than in the type but the teeth are well worn. The inner por-
tion of M? shows a slightly heavier cingulum around the protocone
than in the isolated M? described above.
The lower molar, No. 16215, in the collection shows no important
differences from material of J7'ricentes subtrigonus except that the
paraconid is perhaps a little more lingual in position.
The anteroposterior and transverse diameters of the type, M’,
are 5.1 and 5.6 mm., respectively.
Genus GONIACODON Cope
GONIACODON? species
An upper molar, U.S.N.M. No. 16207, closely resembles Mt in
Goniacodon levisanus, equaling in size this tooth in individuals hav-
ing somewhat smaller teeth than the average in the known material.
The only apparent distinction lies in the extension of the cingulum
on the anterior wall of the tooth to a more lingual point than in
Goniacodon levisanus. The anteroexternal and posteroexternal styles
are broken off so that the direction or extent of these angles cannot
be determined. The tooth is not greatly different from M? in Claeno-
don procyonoides, but the resemblance between the Utah specimen
and M?* in G. levisanus is more striking.
An isolated upper premolar, No. 16208, resembles P* in Goniacodon
levisanus so closely that it may well belong to the same form as
that represented by the molar. The principal cusp is broken down,
but the deuteroconid portion is preserved and corresponds closely to
that in G. levisanus, except in being a little more restricted antero-
posteriorly. The outer portion of the tooth is somewhat distorted,
24 PROCEEDINGS OF THE NATIONAL MUSEUM | VOL, 91
but it appears as if this portion may not have extended so far antero-
posteriorly as in G. levisanus.
The anteroposterior diameter of the upper molar, No. 16207, cannot
be measured, but the transverse diameter is about 9 mm.
Genus DIDYMICTIS Cope
DIDYMICTIS? species
A fourth lower premolar, U.S.N.M. No. 15763, apparently repre-
sents the genus Didymictis. The tooth is only slightly smaller than
in Didymictis haydenianus from the Torrejon but does not have the
first cuspule posterior to the large cusp so distinctly set off from this
primary cusp. The cuspules of the talonid are more nearly in the
median line of the tooth than was observed in D. haydenianus. The
tooth is distinctly larger than in D. microlestes from the Crazy
Mountain locality in the Fort Union of Montana.
An isolated fourth upper premolar may possibly belong to Didy-
miciis but is too small to belong to the form represented by the
lower tooth. Moreover, the deuterocone portion does not extend
forward so markedly as in the Torrejon material of Didymictis, a
condition suggestive of Jctidopappus, but the posterior cusp, though
prominent, is not developed into so nearly a shearing blade as in
either Didymictis or Ictidopappus.
A fragment of the trigonid portion of a lower molar collected
during the 1939 season may represent Didymictis, but it adds little
or nothing to our information regarding the form occurring in the
Dragon.
CONDYLARTHRA
Genus DRACOCLAENUS” Gazin
DRACOCLAENUS GRIPHUS " Gazin
Dracoclaenus griphus GAzin, 1939b, p. 281.
The material in the Dragon collection representing Dracoclaenus
griphus most closely resembles that of the Torrejon form Protoselene
opisthacus but differs from it in several respects. <A relatively large
number of specimens, though fragmentary, are referred to this form
and four of these are figured (fig. 14).
P* (fig. 14, d) in specimen No. 15705 is larger and more inflated
than in P. opisthacus, although there is much variation in P* of
material referred to P. opisthacus, such as between Amer. Mus. Nat.
Hist. Nos. 16614 and 3285. In size of P* D. griphus approaches
Mioclaenus turgidus, but with less reduction of the cingulum and no
£ 10 §paxwy dragon-+claenus.
1 Griphus, an enigma.
PALEOCENE MAMMALS OF CENTRAL UTAH—GAZIN 25
“metaconule” such as usually is present in U/. turgidus. The tritocone
of P* in Dracoclaenus griphus is almost indistinct from the primary
cusp, whereas this tooth in P. opisthacus (Amer. Mus. Nat. Hist.
No. 16614) exhibits a division of the main outer cusp into a promi-
Ficure 14.—Dracoclaenus griphus Gazin: a, M! and M? (U.S.N.M. No. 15789), type
specimen, lateral and occlusal views; b, M? and M3? (U.S.N.M. No. 16182), lateral and
occlusal views; c, right ramus of mandible with M,; and Mz (U.S.N.M. No. 15773), lateral
and occlusal views; d, P4 and part of M! (U.S.N.M. No. 15705), lateral and occlusal views.
X 3. Dragon Paleocene, Utah.
nent protocone and a lesser tritocone placed close together. The
anteroexternal and posteroexternal styles are more prominent on P*
of the Dragon form, and a slightly better developed cingulum, though
discontinuous, is indicated on the outer surface.
302662—41——-4
26 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
The upper molars (type, fig. 14, a), M* and M?, in No. 15789
resemble closely those in Protoselene opisthacus, but the difference
in size between these teeth is more noticeable than in the Torrejon
form, with M? distinctly larger than in P. opisthacus. The external
cingulum is more prominent and more markedly crescentic about
both the paracone and metacone. The mesostyle is well developed
as in certain specimens of P. opisthacus but more conical and dis-
tinctly separated from the crest which extends between the paracone
and metacone. In P. opisthacus the mesostyle extends outward as
a spur or projection from this crest.
Additional material obtained in 1939 includes several more isolated
teeth, but in particular two maxillary portions: No. 16203 with M+
and M? and No. 16182 with M? and M® (fig. 14, 6). The newly ac-
quired upper teeth show Dracoclaenus griphus to run somewhat larger
than P. opisthacus. The two forms are most nearly alike in M’, but
the posterior upper molars show less resemblance. To the greater
size of M? is further added a much better development of the parastyle
than in P. opisthacus. M®*, not hitherto known, is seen to be more
like M? than in P. opisthacus. This tooth is relatively larger than in
the Torrejon form and, although approaching a triangular outline,
shows a more distinct hypocone and much better developed proto-
conule and metaconule.
A somewhat distinctive upper dentition from the Wagon Road
Ridge locality, including P*-M?, No. 15703, resembles the type in most
characters of the molars but has a weaker hypocone on both molars
and a very weak metaconule on M*. The anteroexternal angle of M?
extends forward even somewhat more, suggestive of the oxyclaenids,
but has the mesostyle, particularly in M’, as in No. 15789. The
external cingulum is not so crescentic around the outer cusps, the
outer wall being more nearly straight. P* is similar but a little
smaller than in Nos. 15705 and 15780. This specimen, No. 15703, may
represent a distinct species of Dracoclaenus or may possibly be an
oxyclaenid, close in size to Oxyclaenus simplex; however, P* and M’
more closely resemble the Dracoclaenus material.
The lower jaw portion, No. 15778 (fig. 14, ¢), considered by com-
parison to represent Dracoclaenus griphus, also resembles material
of Protoselene. It corresponds closely in size to P. opisthacus but has
the paraconid on M, and M, more internal in position, and in
M, it is not placed so low and is less reduced than in P. opisthacus.
The talonid basin is apparently not so deep and is narrower between
the hypoconid and entoconid. A slight accessory cusp is present on
the anterior crest of the entoconid nearly as prominent as in P.
opisthacus.
PALEOCENE MAMMALS OF CENTRAL UTAH—GAZIN 2
An M;, No. 15752, in the collection, possibly belonging to this:
form, does not so closely resemble P. opisthacus. The paraconid,
though low, is placed more internal than is usual in the Torrejon
form. Moreover, the entoconid is not so simple as usual in P. opis-
thacus, exhibiting three small cusps in this position, and the hypoconu-
lid is more distinctly separated from the hypoconid.
TABLE 2.—Measurements (in millimeters) of upper and lower teeth of
Dracoclaenus griphus
U.S.N.M. No.—
Measurement 15705 15789 (type) 16182 15773
P4 M! | M? M2?
Anteroposterior diameter__...____.___--____-- Dad
Transverse diameter id
OXYTOMODON * new genus
Type.—Oxytomodon perissum, new species.
Generic characters—Lower teeth slender with cusps high and
distinct. Paraconid on M, and M; lingual in position and close to
metaconid. Cingula absent or weakly developed
and no crest from paraconid to lingual surface
as in Oxyacodon. Hypoconulid less developed.
M, unreduced.
y
liz
VV
\\ en
g >> SS 6
of Vial
OXYTOMODON PERISSUM,'* new species
Type—Left M, and M;, U.S.N.M. No. 16183.
Horizon and locality—Dragon Paleocene,
Dragon Canyon, Emery County, Utah.
Specific characters—Near Oxyacodon pris-
cilla in size. an 2 C
avs : e 15.—Oxyt
Description—A jaw fragment, No. 16183 ee a ee ee
. se SSUM,
(fig. 15), with M, and M, and three additional aeabe ected Ubearcede
specimens, which include only M;, represent in of left ramus of man-
the Dragon fauna a hyopsodont condylarth dible with M2 and M3
near Oxyacodon. Oxytomodon perissum is (U.S.N.M. No. 16183),
7 Ox don priscilla in size, but the para- Pe, See
oe LYACOGON prescet rae ae pa ‘ and occlusal views, X
conid on the lower molars is lingual in position, 4, Dragon Paleocene,
close to the metaconid, and does not exhibit a Utah.
12’o9fbréyos, Sharp + ’odovs, tooth.
13 repisods, UNNECessary or superfluous, in allusion to the considerable variety of small condylarths.
28 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
crest extending from the paraconid down to a weak inner cingulum
around the metaconid as in Oxyacodon. The form resembles Oxya-
codon and differs from Ellipsodon in having relatively high, distinct
cusps, and M, is unreduced in size. However, the hypoconulid is not
so well developed as in the lower molars of Oayacodon, and in M,
it is more reduced and less distinctly separated from the entoconid.
The teeth are slenderer than in O. priscilla and show no marked
cingula on either the lingual or buccal surfaces, except for one of the
third molars, No. 15542, which has a slight cingulum on the outer
surface.
Litomylus dissentaneus from the Crazy Mountain Fort Union ex-
hibits characters close to those seen in Oxytomodon perissum, par-
ticularly in the sharpness of the cusps, but the paraconid in the
lower molars of Z. dissentaneus is much reduced and median in
position.
The anteroposterior and transverse diameters of M, in No. 16183
are 3.5 and 2.7 mm., respectively. The transverse diameter of M;
is 2.4 mm.
Genus ELLIPSODON Scott
ELLIPSODON SHEPHERDI"™ Gazin
Ellipsodon shepherdi GAzIn, 1939b, p. 283.
Ellipsodon shepherdi is comparatively well represented in the
Dragon fauna. The collection now includes about 55 specimens com-
prised of isolated teeth and lower jaw and maxillary portions having
one or more teeth.
This species, as indicated by the type lower jaw (fig. 16, a), is
slightly smaller than Ellipsodon lemuroides, and the molars, M, and
M;, are relatively narrower. M,; is reduced to about the same extent
as in /. lemuroides, more reduced than in the smaller forms, /.
aequidens, EF’. acolytus, and EF. aquilonius, but less reduced than in
the Puerco species, /. priscus, and possibly somewhat less reduced
than in the genotype, Z. inaeqguidens. The paraconid of the last
two lower molars is more distinct in the Dragon form than in any
of the previously known species of Ellipsodon, much better developed
and more lingually placed than in #. aequidens, but only slightly
more prominent than in /. aquilonius. The talonids of M, and M;
are more distinctly basined than in Torrejon material referred to
E. inaequidens, but less distinctly basined than in /. aguilonius from
Montana; also, the talonid on M, is better developed than in the
Puerco form £. priscus. Moreover, the talonid of M, in LZ. shepherdi
does not exhibit so prominent a hypoconulid as in 2. aequidens, but
shows a more distinct entoconid than in Z. inaequidens.
14 Named for Harold Shepherd, in recognition of his field assistance.
PALEOCENE MAMMALS OF CENTRAL UTAH—GAZIN 29
Additional lower jaw material of 2’. shepherdi collected in 1939
includes two specimens, No. 16289 and No. 16303, in which Py, is
preserved in association with the molars, rendering more certain
the reference of several isolated lower premolars to this species. P,
is seen to be comparable in size to that in Z. lemuroides but showing
a distinct metaconid, a slight paraconid, and two cusps at the
posterior margin of the talonid. These are variably developed in
the premolars referred to 2’. shepherdi, but more distinct that in £.
lemuroides and other species from the San Juan Basin. The meta-
conid is better developed than in specimens of the smaller /'. aguilo-
nius but not to the extent seen in Litaletes disjunctus, nor is the
Figure 16.—Ellipsodon shepherdi Gazin: a, Portion of right ramus of mandible with
M.-M3; (U.S.N.M. No. 15721), type specimen, lateral and occlusal views; ), right maxillary
portion with P*M? (U.S.N.M. No. 15790), lateral and occlusal views. XX 3. Dragon
Paleocene, Utah.
paraconid of P, so well defined as in Zitaletes. The moderately
enlarged P, and the brachydont condition of the teeth, combined with
the reduced size of M;, indicate a closer relationship to certain of
the species regarded as Hllipsodon than to Litaletes disjunctus.
The upper teeth, P* to M? in the maxilla, No. 15790 (fig. 16, 5),
referred to Ellipsodon shepherdi are relatively smaller than in the
type lower jaw and approach somewhat closer to /. acolytus than
to #. lemuroides in size; however, this difference within the Dragon
material may not be greater than can be accounted for by individual
variation.
P* shows a cusp in the position that would be occupied by the
metaconule in the molars. This is absent in the somewhat smaller
P* of the Puerco form, #. priscus, but was observed in certain
specimens of the later material. P* is noticeably larger than in /.
aequidens, and M* and M? are relatively longer.
30 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
An Ms, if properly referred, indicates this tooth to be more reduced
than in /#. lemuroides and much more reduced than in Z. acolytus,
FE. aequidens, and FH’. aquilonius.
The upper cheek teeth do not closely resemble those in the genotype,
FE. inaequidens. The upper teeth in the latter exhibit smooth crests
running to the protocone and weak or undeveloped cingula.
TABLE 3.—Measurements (in millimeters) of upper teeth (U.S.N.M. No. 15790)
and lower teeth (U.S.N.M. No. 15721) of Ellipsodon shepherdi
Measurement P4 Mi! M2 Ma M3
FATITELODOSLOLIOM CLAM CLO nee see tee ee ae Se re ait 3.9 3.6 4.4 3.8
*Dransverse! GiamMeten sashes ae serosa eee See ye eee eee e 4.5 4.9 15.8 4 2.9
1 Greatest transverse diameter.
ELLIPSODON? STERNBERGI* Gazin
Ellipsodon? sternbergi Gazin, 1939b, p. 284.
A species nearly intermediate in size between Hilipsodon lemur-
cides and Mioclaenus turgidus is represented by several fragmentary
specimens from the Dragon horizon, including a jaw portion, No.
15755, with M, and a part of M,,
which was made the type of Hllip-
sodon sternbergi (fig. 17). Mz, is
much larger and broader than in
other species of H7lipsodon,; how-
ever, it apparently shows no cren-
ulation of the crest around the
posterointernal margin of the talo-
nid as seen in many, though not
all, of the lower dentitions of HM.
turgidus. M, is a little larger
than in Lllipsodon shepherdi and
somewhat more rounded, being
nearly oval in shape. The paraco-
Ficure 17.—Ellipsodon sternbergi Gazin: nid is lacking on Ma, with only a
Portion of right ramus of mandible with low crest extending across the
Mz and part of M, (U.S.N.M. No. :
ASTER) Nts oe apeGa a ierasatibaa cece! front of the tooth, connecting the
sal views, X 3, Dragon Paleocene, Utah. PYrotoconid and metaconid. Though
reduced, the paraconid is present
in all specimens of Afioclaenus turgidus in which M, was
observed.
Among the specimens referred to 2. sternbergi is a jaw portion,
No. 16339, having both M, and M, preserved. M, is but little larger
© Named for George F. Sternberg, in recognition of his field assistance.
GAZIN 31
PALEOCENE MAMMALS OF CENTRAL UTAE:
than the corresponding tooth in 2. shepherdi and closely resembles
it in form. M, is considerably larger than /. shepherdi and 1s
further characterized by having the talonid basin more restricted
anteroposteriorly than was noted in other species. The paraconid
is present on M,, though not markedly developed. This cusp seems
even less developed on M, in another referred specimen, No. 15769,
in which only this tooth is preserved.
A few upper teeth may be referred questionably to this species,
but these closely resemble upper teeth in /’. shepherdi except for a
somewhat greater transverse diameter and a more prominent proto-
cone. The protocone, however, is not so broad as in M? of Jepsenia
mantiensis. The reduced extent of the talonid basin of M. in Z.
sternbergi is opposed to the enlargement of the protocone in M? of
J. mantiensis, although both of these teeth are large relative to
other teeth in the series.
There is no certainty that this form represents the genus /J/ipso-
don, particularly since the premolars are not known. It is possible
that a small species of Mioclaenus is represented. Moreover, the
distinctions between EZ’. sternbergi and Jepsenia mantiensis are not
entirely satisfactory, being based for the most part on inference.
The transverse diameter of the second lower molar in the type
is about 5 mm. The anteroposterior and transverse diameters of
the third lower molar are 4.4 and 3.3 mm., respectively.
ELLIPSODON? species (a)
A lower jaw, U.S.N.M. No. 15781, from the Dragon horizon is
unusual in that the two teeth preserved, M, and M,, have rather
blunt cusps, a flattened talonid, and a relatively undepressed area
between the three cusps of the moderately elevated trigonid. It
resembles somewhat specimens from the Torrejon that have been
referred to Hilipsodon inaequidens but with the paraconid more
distinctly set off, although this cusp is subdued as are the other
cusps of the teeth. This may represent an unusual condition in
E’. shepherdi but probably represents a distinct form whose affinities
are uncertain.
ELLIPSODON? species (b)
A small hyopsodont is represented in the Wagonroad horizon by
a portion of an upper molar, a second lower molar, and two third
lower molars. The upper molar portion, No. 16282, is larger than in
Ellipsodon shepherdi and has a relatively more expanded protocone
portion, somewhat as in Jepsenia mantiensis but with no evidence of
a hypocone or protostyle although the tooth is noticeably worn.
M2, No. 16284, is almost identical in size with this tooth in the
type of #. shepherdi but differs from it somewhat in that the tri-
SZ PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
gonid portion appears slightly less inflated anteroposteriorly, per-
mitting a somewhat longer talonid basin, suggestive of Litaletes
disjunctus but with less acute cusps. M, also resembles that in
Dragon material referred to Jepsenia mantiensis but is distinctly
narrower and with somewhat better defined cusps on the crest of
the talonid. The third molars, Nos. 16283 and 16285, which may
also belong to the same type of condylarth, are reduced in size
with respect to the second molar described above but not to the
extent shown in £. shepherdi. The talonid basin is more excavated
than in £’. shepherdi and the hypoconulid is better defined, approach-
ing the condition seen in Lztaletes, quite opposed to the reduction
seen in Hllipsodon priscus. Ms, is appreciably smaller and lower
crowned than in Litaletes disjunctus, and the entoconid is not dis-
tinct as it is in the Crazy Mountain form.
The Wagonroad form, if all the above material can be regarded
as representing the same type, appears to be a hyopsodont close to
or within the genus Hllipsodon, but clearly distinct from the Dragon
EL. shepherdi and the nearly contemporaneous /. priscus from the
Puerco.
The second lower molar, No. 16284, has an anteroposterior diameter
of 4.6 mm. and a transverse diameter of 3.9 mm. M,, No. 16285,
is 4.2 and 3.0 mm., respectively.
Genus JEPSENIA * Gazin
JEPSENIA MANTIENSIS ” Gazin
Jepsenia mantiensis GAZIN, 1939b, p. 285.
Jepsenia mantiensis, from the Dragon horizon, makes the closest
approach to Litaletes disjunctus of the various hyopsodont condyl-
arths with which comparisons have been made. The upper molar
series designated as the type, No. 15747 (fig. 18), is only slightly more
robust than in the Montana form. Mt? has about the relative propor-
tions of that in Z. disjunctus and shows a distinct hypocone about
as in that form. However, the lingual portion of M? is more ex-
panded anteroposteriorly, and the hypocone on this tooth is weaker
and represented only by the abrupt termination lingually of the
posterior cingulum. Also, the midportion of the posterior cmgulum
on both M? and M? is not deflected upward toward the root portion
of the teeth so much as in LZ. disjunctus. The cusps in the upper
molars have a more nearly conical appearance, especially the pro-
toconule and metaconule. Moreover, the protoconule and metaconule
are distinctly better developed. A parastyle and mesostyle are pres-
ent, more noticeable in M?, although the cingulum is not so extended
16 Named for Dr. Glenn L. Jepsen.
17 Named for the Manti National Forest.
PALEOCENE MAMMALS OF CENTRAL UTAH—GAZIN 33
at the anteroexternal portion of the molars. M*® is relatively smaller
than in LZ. disjunctus and the metacone, though distinct, is not so
well developed, and the cingulum is less prominent and is discon-
tinuous around the lingual and buccal surfaces of the tooth.
An M? with material numbered 15544 shows more acute antero-
external and posteroexternal styles, no mesostyle, a lower protocone
than in LZ. disjunctus, protoconule and metaconule relatively weak
as in LZ, disjunctus, but the hypocone is
much more lingual in position and is
nearly matched by a protostyle on the
anterolingual portion of the tooth, with
the cingulum almost but not quite contin-
uous around the inner margin of the pro-
tocone. Mt? in this material, though lack-
ing a mesostyle, corresponds closely to that
in the type of Jepsenia mantiensis. It is
possible that the two molars, which were
found close together, belong to the same
individual and may represent a type dis-
tinct from the foregoing.
Several isolated jaw fragments with
single molars, one with M, and part of
M,, and several with portions or all of M, Ficure 18.—Jepsenia mantiensts
and M;, are presumed to represent Jep- Snes eae por
senia mantiensis. The lower teeth in gen- Ra bs re ae se
ae rete : oO. ), type specimen,
eral show a distinct paraconid in a lin- faite ral anioccluealenew sco:
gual position and a basined talonid with Dragon Paleocene, Utah.
a strong hypoconid, a moderate entoconid,
and a weak hypoconulid which is the dorsal termination of a slight pos-
terior cingulum rising from the posteroexternal portion of the tooth.
The trigonid portion is not greatly different from that in Z. disjunctus,
but with less acute cusps. The entoconid on the heel of M, and of
M, is less developed, and the small cuspule anterior to the entoconid
is less evident than in Litaletes. Mz, is about the size of that in
Ellipsodon? sternbergi but is narrower and shows a distinct para-
conid, not, however, so distinct as in ZL. shepherdi. M, in E£.? stern-
bergi is distinctly wider than in the material referred to Jepsenia
mantiensis but the talonid basin is relatively smaller.
TABLE 4.—Measurements (in millimeters) of upper teeth of Jepsenia mantiensis
Measurement M! M23 M3
AT CCT ODOSLOFIO“ GIAIIGLen hen een dn. 2 se Pe 4.5 4.4 3
PETADSVOISe CIAIMOLOR Sao eee ee ee ee he ie et 5.4 6.4 14.6
1 Greatest transverse diameter.
34 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
DESMATOCLAENUS,® new genus
Perhaps one of the most interesting discoveries made by the
1939 expedition is the finding in both the Dragon and Wagonroad
levels of a new J'etraclaenodon-like form which nearly bridges the
gap between Z'etraclaenodon and forms of Protogonodon. Desmato-
claenus is so nearly intermediate that its assignment to the condy-
larths rather than to the creodonts is entirely arbitrary.
Type.—Desmatoclaenus hermaeus, new species.
Generic characters——P* with prominent deuterocone and no indi-
cation of tritocone. P* intermediate between Protogonodon and Tet-
raclaenodon. Anteroexternal portion of M? projects outward more
than in Protogonodon. External cingulum discontinuous across par-
acone in M* and M?, and there is no mesostyle between the outer
cusps of these teeth. Hypocone, protoconule, and metaconule less
developed than in Tetraclaenodon. Hypocone not so lingual in po-
sition as in Protogonodon. M® relatively small with prominent cin-
gulum about protocone and without evidence of a hypocone. P,
nearly as in Tertaclaenodon but relatively small. Lower molars with
lingually placed paraconid much better defined than in Tetraclae-
nodon, and talonid basin not so broad as in Protogonodon. M,; with
cuspidate entoconid-hypoconulid crest.
DESMATOCLAENUS HERMAEUS ” new species
Type.—Greater portion of upper and lower dentition, U.S.N.M.
No. 16202.
Horizon and locality—Wagonroad Paleocene, Dragon Canyon,
Emery County, Utah.
Specific characters—Size near Protogonodon protogonioides,
shghtly smaller than Tetraclaenodon puercensis.
Description—The specimen comprising the best material is an
assortment of 14 more or less complete upper and lower teeth, clearly
from one individual, No. 16202 (fig. 19), found in the Wagonroad
horizon. The inclusion in the material of upper and lower premo-
lars was extremely fortunate in that the approach to T'etraclaenodon
is more distinctly shown.
P’, though incomplete anteriorly, is much like that in Tetraclae-
nodon, with the principal cusp somewhat flattened transversely and
exhibiting a sloping posterior crest but with no indication of a
tritocone—the principal cusp is higher and more conical in Proteg-
onodon. The deuterocone, a distinct cuspule almost as well devel-
oped as in Tertaclaenodon, is placed somewhat farther forward than
in this form, about in the position occupied by a suggestion of a
18 dégua, a chain or link + claenus.
19*épuaiov, a lucky find.
PALEOCENE MAMMALS OF CENTRAL UTAH—GAZIN 35
deuterocone in P* of Protogonodon. The posterointernal cingulum
is better developed than in Protogonodon, but not so shelflike as in
Tetraclaenodon.
P* is somewhat more worn but shows the principal cusp to be
slightly less conical than in Protogonodon with a more distinct pos-
terior crest. The presence or absence of a tritocone cannot be deter-
mined because of wear, but if present it was not developed to the
extent seen in Tetraclaenodon. The deuterocone portion is restricted
anteroposteriorly more than in J'eéraclaenodon, approaching Pro-
togonodon, but a cingulum not seen in Protogonodon is developed
along the anterior and posterior walls of this cusp, separate from
the shelf or crest joining the deuterocone to the outer extremities of
the tooth. The cimgulum and shelf are not developed to the extent
seen in Tetraclaenodon, nor is there certain evidence of a protoconule
or metaconule on the crest; however, wear may have obliterated an
incipient development of these. The parastyle, as in Tetraclaenodon,
is directed more externally than in Protogonodon.
Figure 19.—Desmatoclaenus hermaeus, new genus and species: Left upper dentition,
including P’, P4, M2, M3, and right lower dentition, including P3-M,, Mz, and part of Mg
U.S.N.M. No. 16202), type specimen, occlusal views, X 2, Wagonroad Paleocene, Utah.
M’ is not preserved in the material of this individual but is
included in a maxilla of another and larger specimen, which pre-
sumably represents a distinct species and is described elsewhere. .
M? is rather well worn but was evidently low cusped and had a
weak hypocone as compared with this tooth in Protogonodon and
in contrast to the marked development of the cusp in Zetraclaenodon.
However, this cusp is located directly posterior to the protocone
as in Tetraclaenodon, occupying a position in the flexure between
the protocone and metaconule, and not so lingual in position as
noted in Protogonodon. The protoconule and metaconule appear to
be less developed relative to the primary cusps than in Tetraclae-
modon, in which the six principal cusps approach equality. In
Protogonodon the protocone is more prominent and somewhat over-
36 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
shadows the protoconule and metaconule. The anterior portion of
the tooth is relatively wide and projects outward somewhat as in
Tetraclaenodon and shows a prominent parastyle. The external
cingulum is much weaker than in Protogonodon and is peculiar in
being discontinuous across the postero-external portion of the para-
cone; however, there is no mesostyle such as observed in TJ'etraclae-
nodon and the cingulum is perhaps a little better developed postero-
external to the metacone than in Tetraclaenodon.
MS is relatively small as in 7'etraclaenodon, more reduced than in
Protogonodon, but the cingulum is continuous around the inner wall
of the protocone as in the latter and there appears to be little or
no evidence of a distinct hypocone.
The lower teeth of the type are from both rami and between
them include a representation of the series from P; to Ms. Although
rather well worn, many characters can be ascertained showing, as
with the upper dentition, the structural position that this form
holds between Protogonodon and Tetraclaenodon.
P;, though incomplete posteriorly, is seen to be small and narrow,
comparable in this respect to Protogonodon, but with a more gently
sloping posterior crest.
P,, though slender and relatively small, shows a marked resem-
blance to Z'etraclaenodon. The parastylid is high, prominent, and
deflected inward from the anterior crest of the protoconid about
as in Tetraclaenodon. The tooth is well worn, but from the outline
of the occluding surface there is little doubt that a pronounced
metaconid was present. The heel structure is nearly as in 7Jetra-
claenodon but with less anteroposterior extent and a less distinct
entoconid.
M, is too worn to show any important characters but as in the
succeeding tooth shows the talonid to be less widely basined than in
Protogonodon.
In M, the trigonid portion exhibits a more prominent paraconid
than in Zetraclaenodon, which is perhaps not so close to the meta-
conid, but as in the latter it is distinctly lingual in position and is
joined by an arcuate crest to the anterior slope of the protoconid,
forming a somewhat more distinct but anteroposteriorly restricted
trigonid basin than in Protogonodon pentacus.
M; is relatively small as in Tetraclaenodon but with a much better
developed paraconid. The trigonid is anteroposteriorly shortened
and the paraconid more lingual in position than in Protogonodon.
The talonid basin is relatively simple, with the entoconid and hy-
poconulid not actually distinct but forming a slightly cuspate crest.
Remarks.—The intermediate position of Desmatoclaenus between
Protogonodon and Tetraclaenodon suggests that Tetraclaenodon may
PALEOCENE MAMMALS OF CENTRAL UTAH—GAZIN Bi
have arisen from Protogonodon through Desmatoclaenus. This may
well be the case but the larger known forms such as P. pentacus or
even P. stenognathus are probably not in the line. It is conceivable
that a small form such as P. protogonioides, whose teeth are closer
to Desmatoclaenus than are those of P. pentacus (especially P*),
may have given rise to Desmatoclaenus, assuming a somewhat earlier
stage for the Puerco of the San Juan Basin.
Taste 5.—Measurements (in millimeters) of upper and lower teeth of
Desmatoclaenus hermaeus (U.S. N. M. No. 16202)
Measurement Pi|M?2!|M3| P3 Pi!|]Mi|/Maj]Ms
Anteroposterior diameter ==" 2.2 22-52--28 522-22 _ |e el Bhat Os in] a ses 7.1 7A | ceeeaee 8.5
MTAnS Verse Gualn Olek - sesso a2a2 2 eee es eee GSO TAS On) USA hazed ASS GN Be|) Te 7 5.8
1 Greatest transverse diameter.
DESMATOCLAENUS PARACREODUS,°° new species
Type.—Right maxillary portion, U.S.N.M. No. 16201, with M'*-M:®.
Horizon and locality—Wagonroad Paleocene, Dragon Canyon,
Emery County, Utah.
Specific characters —Larger than Desmatoclaenus hermaeus. Lin-
gual portion of upper molars more inflated. M® relatively larger.
Hypocone better developed.
Description.—A second and somewhat larger species is indicated
by material apparently from both the Wagonroad and Dragon hori.
zons. The specimen selected as the type, No. 16201 (fig. 20, a),
was obtained from the Wagonroad level and includes M! to M*. The
teeth are much like those in Desmatoclaenus hermaeus in most char-
acters of the molars, but the lingual portions of these teeth have a
more inflated appearance and M® is relatively larger. Although
slightly damaged at the posterointernal angle, M* shows better evi-
dence for a hypocone than in D. hermacus. The upper molars make
an approach toward the conditions seen in Protogonodon stenog-
nathus, but the differences, as in D. hermaeus, are in the direction of
Tetraclaenodon.
A maxillary portion, No. 16177 (fig. 20, 6), with M? and M? from
the Dragon horizon corresponds closely to the type of D. paracreodus,
but the teeth being less worn show characters not seen in the type.
The external cingulum is weaker than in Protogonodon, and, as in
the types of D. hermaeus and D. paracreodus, the cingulum is inter-
rupted along the posteroexternal portion of the paracone in M2, and
the anteroexternal portion of both teeth projects outward promi-
20 rapa, near + xpéas, flesh + ’odots, tooth, in allusion to its resemblance to the carnivore Protogonodon.
38 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
nently. This portion of M? is slightly damaged, but the anterior
cingulum becomes weil developed laterally, suggesting a conspicuous
parastyle as in Z’etraclaenodon. The cusps are all low and conical
in M? and the lingual portion, as in the type, is somewhat inflated
anteroposteriorly, with no cingulum around the inner portion. The
hypocone is weak and situated posterior to the protocone. In the
early stage of wear represented by this specimen the protocone is
seen to be divided, with a slight cuspule immediately adjacent and
posterior to the principal cusp. This may have been the case in M?
of the type of D. hermaeus, as indicated by the outline of the worn
surface of occlusion.
HIN
ce
ii ey
Ficure 20.—Desmatoclaenus paracreodus, new species: a, M!-M3 (U.S.N.M. No. 16201)
type specimen, occlusal view; 6, M?-M? (U.S.N.M. No. 16177), occlusal view; c, lower
molar (U.S.N.M. No. 16196), lateral and occlusal views; d, lower molar (U.S.N.M.
No. 16194), lateral and occlusal views. 2. a, c, d, Wagonroad Paleocene, Utah;
b, Dragon Paleocene, Utah.
M? of the Dragon specimen is somewhat distorted, but the cingulum
is better developed than in M?. The outer cusps are perhaps more
compressed anteroposteriorly and the protocone seems relatively prom-
inent. On both molars the enamel is relatively smooth, except for
a noticeable rugosity around the lingual wall of the protocone near
its peak.
Several isolated lower molars, including No. 16194 (fig. 20, @)
and No. 16196 (fig. 20, ¢), from the Wagonroad level are referred to
this species, being comparable to those of D. hermaeus in structure
PALEOCENE MAMMALS OF CENTRAL UTAH—GAZIN 39
but are appreciably larger, even than in 7’etraclaenodon, being about
the size of those in Protogonodon stenognathus. The trigonids of
these teeth show the paraconids to be entirely lingual in position, as
in Zetraclaenodon, but better developed and perhaps not so close to
the metaconid. The paraconid is more lingual and not so far forward
as in Protogonodon material, and the crest from the paraconid to
the anterior wall of the protoconid is higher, closing the trigonid
basin anteriorly. Moreover, the talonid portion of the lower molars
is relatively narrower than in Protogonodon pentacus with the basin
restricted transversely, being more nearly comparable to the form of
the talonid in the first two lower molars of Yetraclaenodon. A
relatively narrow talonid was noted in the lower molars of the large
Protogonodon kimbetovius.
A jaw portion with M,, No. 16218, and an isolated portion of a
lower molar in the collections from the Dragon level are considered to
belong to D. paracreodus. These closely resemble the lower teeth
from the Wagonroad level referred to D. paracreodus.
TABLE 6.—Measurements (in millimeters) of upper teeth of Desmatoclaenus
paracreodus (U. 8S. N. M. No. 16201)
Measurement | M! | M? | M3
eee aa
ATTHOLODOSLOFION GIANG (Obs cern: =) 2 ee keene ee eRe 2 ae a ee 8.4 8.1 6. 2
MEALS VEESCLIAIIOLED eee See aoe. | sees DOR eee oe ee ee 10.5 12 19.9
! Greatest transverse diameter.
Genus ECTOCONUS Cope
ECTOCONUS SYMBOLUS,” new species
Type.—Right maxillary portion, U.S.N.M. No. 16189, with M1’,
M?, and part of P*.
Horizon and locality——-Wagonroad Paleocene, Dragon Canyon,
Emery County, Utah.
Specific characters —Molars smaller than in E’ctoconus ditrigonus.
Premolars relatively larger. No “protoconule” on Pt. Protostyle on
upper molars weak. Parastyle on M? weak. Parastylid absent or
weakly developed on lower molars.
Description.—Several specimens from the Wagonroad horizon, in-
cluding maxillae and jaws with two teeth each, are found to represent
a new species of Ectoconus. The molar teeth are seen to be dis-
tinctly smaller than in #. ditrigonus, hence much smaller than in
EL. majusculus. The premolars, however, are relatively larger and the
anterior lower premolars, as indicated in referred specimens, are
actually larger than in Z. ditrigonus.
*1 ¢bpBodor, clue, in allusion to its importance in determining the age of the Wagonroad horizon.
40 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
The upper molars, No. 16189 (fig. 21, ), of which only M* and M?
are known, closely resemble those in #. ditrigonus in structural de-
tails, but with perhaps a somewhat weaker protostyle. ‘The postero-
external portion of M‘ shows the cuspate condition characterizing
upper molars in Y'ctoconus. The mesostyle, metastyle, and the large
cusp external to the metacone are developed to about the same extent as
in LZ. ditrigonus ; however, the parastyle on M? appears weaker than in
E.. ditrigonus. P*, No. 16188 (fig. 21, ¢), is of about the same width,
or perhaps slightly wider transversely than M’, and differs from that
Ficure 21.—Ectoconus symbolus, new species: a, Portion of left ramus of mandible with
M.-M; (U.S.N.M. No. 16190), lateral and occlusal views; b, M! and portions of P4
and M? (U.S.N.M. No. 16189), type specimen, occlusal view; c, P4-M! (U.S.N.M. No.
16188), occlusal view. XX 1%. Wagonroad Paleocene, Utah.
in #. ditrigonus in the absence of an accessory cusp anteroexternal
to the deuterocone, in about the position occupied by the protoconule
in the molars.
The lower jaw material consists of three specimens which together
give a representation of the dentition from P, to M;, except for M,.
The premolars are relatively large, particularly P., No. 16213, but
become relatively narrower posteriorly than in #. ditrigonus. The
molars, No. 16190 (fig. 21, a), are smaller and relatively narrower
than in £2. ditrigonus, and there is but the slightest suggestion of a
second paraconid or parastylid; however, the presence of this cuspule
is not invariable in /’. ditrigonus. M, and M; in Hctoconus symbolus
are otherwise similar to those in /. ditrigonus in having low blunt
cusps and a heavy external cingulum.
PALEOCENE MAMMALS OF CENTRAL UTAH—GAZIN 41
TasLe 7.—Measurements (in millimeters) of upper and lower teeth of Ectoconus
symbolus
| U.S.N.M. No.—
Measurement 16188 16189 (type) 16190
wAmteronosterion diameter -2-0--==22.---..--2hsbanaces 6.8 8.4 8.1 18.7 9.6 10.6
Transversprainmeters 5-2". 0) te te 110 10.0 10.5 | 112.5 8.8 8.2
1 Approximate.
Tt on Yh,
SSE 20, A ee
35
iri
io
Th ( \
call TIM
WAN
Ficure 22—Carsioptychus hamaxitus, new species: a, Maxillary portion with two premolars
(U.S.N.M. No. 16198), lateral and occlusal views; b, left maxillary portion with M! and
M? (U.S.N.M. No. 16197), type specimen, lateral and occlusal views; c, portion of left
ramus of mandible with M2 and Ms; (U.S.N.M. No. 16195), lateral and occlusal views
1%. Wagonroad Paleocene, Utah.
42 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
Genus CARSIOPTYCHUS Simpson
CARSIOPTYCHUS HAMAXITUS,~ new species
Type.—Left maxillary portion, U.S.N.M. No. 16197, with M' and
M?.
Horizon and locality—Wagonroad Paleocene, Dragon Canyon,
Emery County, Utah.
Specijfie characters —Teeth smaller than in Carsioptychus coarcta-
tus. Premolars slightly smaller with respect to molars than in C.
coarctatus and upper teeth relatively a little narrower transversely
than in the Puerco form. Lower premolars with slightly better
developed anterior stylid.
Description.—Several specimens, including upper and lower teeth,
from the Wagonroad level represent a small species of Carsioptychua.
Though the teeth are small as compared to those in Carsioptychus
coarctatus, the form is slightly more progressive toward Periptychus
than is the Puerco species, but not so advanced as Periptychus gil-
more? from the Dragon. The premolars are relatively smaller than in
C. coarctatus and the upper molars, No. 16197 (fig. 22, 6), and pre-
molars, No. 16198 (fig. 22, a), are relatively narrower transversely.
Moreover, the lower premolars show a slightly more advanced stage in
the development of an anterior stylid. The lower molars (fig. 22, ¢)
appear to be developed much as in @. coarctatus, and as in that species
show no evidence of the seventh cuspule, near the center of the tooth,
characterizing Periptychus carinidens, but seen only on Mz; of P.
gilmoret.
Tasty 8.—Measurements (in millimeters) of upper and lower teeth of
Carsioptychus hamaxitus
U.S.N.M. No.—
Measurement 16198 | 16197 (type) 16195
| P3? | Pi? Mi! M2 Ma M;
ANITOFiON CIAMeter 223. keer: Paar RL ata AE ee del eel 10.8 17.8 8.2 8.0 9.5
Mransverse: diameters a= eee seek ee ae 11.8 | 13: 5 11.4 11.8 17.8 Tee
|
1 Approximate.
2 The transverse diameter of the upper teeth is taken from the external cingulum to the base of the ename
lingually and at right angles to the direction of the tooth row.
22 quatiros,ecarriage road or wagon road, from the name of the horizon in which it was found
and the name of the ridge, at the lower end of which the locality occurs.
PALEOCENE MAMMALS OF CENTRAL UTAH—GAZIN 43
Genus PERIPTYCHUS Cope
PERIPTYCHUS GILMOREI *° Gazin
Periptychus gilmorei GAzIN, 1988, p. 275.
The large periptychid, P. gilmorei, in the Dragon fauna is rather
well represented in the collection, the best specimen being the type,
No. 15537, and including portions of right and left maxillae with 14
teeth in all (fig. 23). Specimen No. 16228, obtained in 1939, includes
portions of both maxillae with P*-M* and a portion of the left
ramus of the mandible with P.—M;, the lower teeth being partially
embedded in barite. The lower dentition is best represented in speci-
men No. 15689 (fig. 24), which includes portions of right and left
rami, exhibiting M,-M; and P,-M,, respectively.
Ficure 23.—Periptychus gilmoreit Gazin: Right upper dentition including P?-M3 (U.S.N.M.
No. 15537), type specimen, lateral and occlusal views, X 114, Dragon Paleocene, Utah.
Periptychus gilmorei is intermediate between Carsioptychus coare-
tatus from the Puerco and Periptychus carinidens from the Torrejon
in almost all characters of the upper dentition. The teeth are
relatively wide transversely as compared with their length, and the
premolars are only slightly larger than the molars. The premolars
show the inner crescent developed almost as much as in Periptychus
carinidens, but the deuterocone portion is more constricted antero-
posteriorly, although not so much as in Carsioptychus coarctatus.
Moreover, P* is much more like that in Periptychus than the simple
condition observed in several specimens of Carsioptychus.
The molar teeth show a distinct resemblance to those in Carsiop-
tychus, and in addition to their being relatively wide transversely
they show a more distinct external cingulum than in Periptychus.
% Named for C. W. Gilmore, whose party discovered the first Dragon Canyon locality.
44 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
The hypocone and protostyle have a somewhat more lingual position,
and the lingual walls of the molars (and premolars as well) appear
to be more gently sloping than in Periptychus. The cusps and
cuspules are somewhat less widely spaced than in P. carinidens, par-
ticularly the protoconule and metaconule, which are located very close
to the protocone.
De
Ro [iN i a
Ceo
Ficure 24.—Periptychus gilmorei Gazin: Left ramus of mandible, PygM3 (U.S.N.M.
No. 15689) (M3; and posterior portion of jaw fragment restored from right ramus), lateral
and occlusal views, X 1%, Dragon Paleocene, Utah.
An additional feature seen in the type of Periptychus gilmorei,
but probably of no importance, as it was not observed in No. 16226,
is the very slight development of a “protostyle” and “hypocone” on
P*. This was not observed in any of the Puerco or Torrejon material.
Also, the third molar, on the right side only, is peculiar in that the
lingual wall exhibits a cuspule median to the protocone, between the
protostyle and hypocone.
The lower teeth of Periptychus gilmorei, as represented by speci-
men No. 15689, are also nearly intermediate in most respects between
Carsioptychus coarctatus and Periptychus carinidens. The protoconid
of P, is not directed posteriorly so markedly as in C’. coarctatus, and a
small anterointernal cusp is present, this being prominent in P. carini-
dens but usually absent in C. coarctatus. On the posterointernal por-
tion of the tooth there is a small cusp; the talonid, however, is not
developed so much as in P. carinidens. The extent to which a meta-
PALEOCENE MAMMALS OF CENTRAL UTAH—GAZIN AD5
conid has become distinct from the protoconid cannot be exactly
determined, owing to wear, but it is clearly not separated to the ex-
tent seen in P. carinidens.
The lower molars are wider than in the Carsioptychus material at.
hand but not so wide as is common in Torrejon material of Peripty-
chus. These teeth show a slight cingulum around the external side,
which was not observed in material of the other forms. The small
seventh cusp located about in the center of the crown of the lower
molars of Periptychus carinidens is not present in the first two molars
of P. gilmorei but is weakly developed in M;. This cusp is not known
in Carsioptychus.
Taste 9.—Measurements (in millimeters) of upper teeth (U.S.N.M. No. 15587,
type) and lower teeth (U.S.N.M. No. 15689) of Periptychus gilmorei
Measurement p2 Ps Pt M! | M? | M3 Pi Mi | M2 | M3;
Anteroposterior diameter________-__-__- PU 6)) 11.7 | 10.5.1), 920) 9851} 838) |-11 10.3 | 10 11.5
Transverse diameter !1___.______._-_.-_- 12.7 | 14.6 | 14.0 | 14.2 | 141 | 111 9.6 8.7 9.7 9
1 The transverse diameter of the upper teeth is taken from the external cingulum to the base of the enamel
lingually and at right angles to the direction of tooth row.
Genus ANISONCHUS Cope
ANISONCHUS DRACUS™ Gazin
Anisonchus dracus GAzIN, 1939b, p. 278.
The larger of the two species of Anisonchus is represented in
the Dragon collection by three maxillary portions with one to four
teeth apiece and five lower jaw fragments with one or two molars
each. The type, No. 15745, is a maxillary fragment with P* to M®
preserved (fig. 25).
The upper teeth in No. 15745 are clearly of an Anisonchus type
and are intermediate in observed characters between A. gillianus
and A. sectorius of the Puerco and Torrejon, respectively; com-
parable in this respect to Periptychus gilmorei in its relationship
to the two developmental stages occurring in the San Juan Basin,
noticeably in the relation of the length to the width of the tooth
crowns.
The Dragon form approaches A. sectorius in size but retains rela-
tively wider teeth transversely, and longitudinally a little shorter,
and the cusp pattern is not so restricted transversely. The upper
teeth appear also to have a longer, more gradually sloping lingual
wall, with a somewhat more lingually placed hypocone column. The
4 dpaxwy, dragon, from Dragon Canyon.
46 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
lingual portion of P* seems more constricted anteroposteriorly and
apparently has a less conspicuously developed lingual crescent.
A, gillianus has teeth relatively wide transversely, the length of
the tooth row shorter, and the hypocone is placed more lingually
with respect to the metacone, and to a certain extent with respect
to the protocone, than in A.
sectorius,
The lower jaw fragments
exhibit teeth comparable in
size to those in A. sectorius
and show no significant differ-
ences from them, nor are dif-
ferences evident in the pre-
served material which would
serve to clearly distinguish
the Dragon form from A.
gillianus. However, the crest
connecting the hypoconid to
the trigonid appears distinctly
lower than that connecting the
entoconid to the metaconid.
This condition was noted in
Ficure 25.—Anisonchus dracus Gazin: Left gn M,.of A: gillianus but not
maxillary portion with P4-M3 (U.S.N.M. in other specimens of either
No, 15745), type specimen, lateral and occlusal pre 7 ple
views, X 3. Dragon Paleocene, Utah. Be eC eae Ore:
Moreover, the hypoconulid
does not project backward in the molars referred to Anisonchus
dracus quite so far as in M, of A. sectorius, a condition approxi-
mated in M, of A. gillianus, though possibly of doubtful significance.
Taste 10.—MVeasurements (in millimeters) of upper teeth (U.S.N.M. No. 15745,
type) and lower teeth (U.S.N.M. No. 16249) of Anisonchus dracus
Measurement Ps M! M? M3 Mi Me
FATILELODOSLELIODO Ameer aie ee ee eee 5? 4.4? 4.8 4? 5.2 5.2
Mransverse: Gliameter. sate Soeein: we ee ee eee ee Cee ee 6. 6? Tulse ere: 3.6 3.9
ANISONCHUS ONOSTUS * Gazin
Anisonchus onostus GaziIn, 1939b, p. 280.
The smaller of the two species of Anisonchus in the Dragon fauna
is represented by the type, No. 15788 (fig. 26), which is a lower
jaw portion with M, and M., and to the species is tentatively referred
an upper premolar and a lower jaw fragment with the teeth P,,
M,, and part of M, much worn.
2 Onostus, despicable, in allusion to its size.
PALEOCENE MAMMALS: OF CENTRAL UTAH—GAZIN 47
Anisonchus onostus is distinctly smaller than A. dracus, being
very near the Puerco form, A. gillianus, in size but with the cusps
on the talonid of both M, and M, slightly more widely spaced, though
having the cut characterizing the anisonchines. This spacing of
the cusps gives the teeth a wider
appearance, whereas actually they
are a trifle narrower than those in
several specimens of A. gillianus with
which comparisons were made. The
teeth also appear somewhat lower
crowned than those of A. gillianus
exhibiting about the same wear.
The anteroposterior diameters of
the first and second lower molars
are 4.3 and 4.1 mm., respectively.
The transverse diameters are 2.9 and
3.2 mm.
ANISONCHUS OLIGISTUS,* new species
Type—tLeft maxillary portion
with M* and M? associated portion
of left ramus of mandible with M,
and M., U.S.N.M. No. 16192.
Ficure 26.—Anisonchus onostus Gazin:
Portion of left ramus of mandible
with M,-M2 (U.S.N.M. No. 15788),
type specimen, lateral and occlusal
views, X 3, Dragon Paleocene, Utah.
Horizon and locality—Wagonroad Paleocene, Dragon Canyon,
Emery County, Utah.
Figure 27.—Anisonchus oligistus, new species: Left maxillary portion with M!-M?;
portion of left ramus of mandible with M;—Mg2 (U.S.N.M. No. 16192), type specimen,
lateral and occlusal views, X 3, Wagonroad Paleocene, Utah.
Specific characters—Upper and lower molars smaller than in
Anisonchus gillianus and relatively narrower transversely. Upper
2f’odiyoTos, least, in allusion to size of teeth.
4S PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
molars more nearly triangular in occlusal view. Talonid basin of
lower molars slightly less constricted anteriorly.
Description—Anisonchus is represented in the Wagonroad collec-
tion by a maxillary portion and a lower jaw fragment found to-
gether and both having the first two molars preserved, U.S.N.M.
No. 16192 (fig. 27), which has been made the type of Anisonchus
oligistus. Six other specimens are referred to this species. These
include two maxillary fragments, with M?—M* and P*-M? somewhat
damaged, two lower jaw fragments each with the greater portions
of two molars, and two isolated premolars.
Anisonchus oligistus is apparently the smallest species known
of this genus, having both upper and lower molar teeth a little
smaller and relatively narrower transversely than in material of A.
gillianus from the Puerco. The lower teeth are also smaller and
more slender than in the type of Anisonchus onostus from the Dragon
level.
The upper molars appear for the most part very much like those
in other species of Anisonchus, but are somewhat more nearly trian-
gular in outline, as viewed from below, with the lingual portion a
little more constricted anteroposteriorly and the hypocone column
distinctly lingual, though not so markedly lingual as in Haploconus.
The anterior cingulum extends to a markedly lingual point but does
not exhibit a distinct protostyle.
The lower molars in addition to their slenderness show relatively
high trigonids, and the cusps appear to be more acute than in
A, gillianus. Moreover, the paraconid may be slightly more external
in position. The talonid appears deeply basined in the type, and
the crest extending forward from the hypoconid joins the posterior
wall of the trigonid at a position which appears to be slightly more
external. This is not so obvious in the type, but noticeable in the
two referred lower jaws. As a result the talonid basin in the
referred specimens appears somewhat less constricted anteriorly.
TABLE 11.—Measurements (in millimeters) of upper and lower teeth of
Anisonchus oligistus (U.S.N.M. No. 16192)
Measurement M! M2 Mi M2
Amfteropostenior Giameters:2— eee he ee a A eee 3.9 3.7 SoS eet. eee
Transverse dismoten esas te set ee a ee ae ee 5.1 6.0 2.8 2.9
) The transverse diameter of the upper teeth is taken from the external cingulum to the base of the enamel
lingually and at right angles to the direction of the tooth row.
PALEOCENE MAMMALS OF CENTRAL UTAH—GAZIN 49
Genus HAPLOCONUS Cope
HAPLOCONUS INOPINATUS 2 Gazin
Haploconus inopinatus GAzin, 1939b, p. 280.
A second genus of anisonchine periptychids is represented in the
Dragon fauna by several fragmentary specimens, including a maxil-
lary portion with M' and most of M’, No. 15760, which has been
made the type of Haploconus inopinatus (fig. 28). The form appar-
ently represents Haploconus as indicated by the prominent lingual
position of the hypocone. It is close in size to the
Torrejon material referred to Haploconus angus-
tus but with the teeth relatively wider transversely
and with M? much wider than M'. A difference
in width between M' and M? was noted in certain
specimens of Haploconus referred to H. angustus,
but apparently the difference is not so marked as
in #7. inopinatus.
The two upper molars in the type show a slight
development of a metaconule, but most noticeable
is the distinct protostyle that characterizes teeth
in Haploconus corniculatus. H. inopinatus is much
smaller than the type of H. corniculatus, and in
the latter the upper molars appear to be relatively
Figure 28.—Haploco-
nus inopinatus Gaz-
as well as actually much longer anteroposteriorly
than in the Dragon form.
The anteroposterior diameter of the first upper
molar in the type is 43 mm. The greatest trans-
verse diameters of the first and second upper
molars are 6.1 and 7.1 mm., respectively.
A second maxillary portion, No. 16256, is re-
ferred to H. inopinatus; however, the two molars
it exhibits are not well preserved and add little to
in: Left maxillary
portion with Mi! and
the greater part of
M? (U.S.N.M. No.
15760), type speci-
men, lateral and oc-
clusal views, X 3,
Dragon Paleocene,
Utah.
our knowledge of this form, An isolated upper premolar, apparently
P*, No. 16254, may well belong to /aploconus, closely resembling this
tooth in HZ. angustus, but a little smaller and with the lingual portion,
though broad, somewhat less inflated anteroposteriorly.
A lower jaw portion, U.S.N.M. No. 15744, with M, and M, poorly
preserved, and partially obscured by ironlike matrix, appears to
represent Haploconus in the absence of a paraconid and in the blade-
like form of the protoconid on M;. It corresponds closely in size
to the type of Haploconus angustus, but with M, narrower, particu-
larly the anterior portion, and M, possibly wider than in the Torrejon
form.
* Inopinatus, unexpected.
50 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
A second lower jaw portion, No. 16255, collected in 1939, has P,
and the greater portion of M,and M, preserved. P, is a little shorter
than in most specimens of 7. angustus, though relatively as wide and
appears inflated as characteristic of this genus. The two molar
portions show no important distinguishing characters. These two
teeth have the cingulum rather prominent external to the protoconid,
but distinctly weak on Py. In No. 15744 the cingulum is not evi-
dent. However, in H. angustus the development of the cingulum
appears to be highly variable, and when present is apt to be most
noticeable on the anterior portion of the tooth and about the hypocone.
In 1940 several isolated teeth were found near one another at a level
about 30 or 40 feet higher than that of the Dragon fauna at the
old Dragon Canyon locality. These include P*, a right and left P,,
portions of two anterior lower molars, and the greater part of M3.
The talonid portions of the various lower molars are to be compared
with those of Haploconus rather than any other known form. One
of the molars, however, has most of the trigonid preserved, and this
exhibits a small paraconid. It is also significant that the two lower
premolars have a moderately developed paraconid and are antero-
posteriorly elongate and slender, approaching the form seen in
Anisonchus, quite unlike the premolar exhibited in No. 16255 referred
to H. tnopinatus. The form represented by these teeth is clearly
distinct from that represented by No. 16255, but I hesitate to describe
it as distinct because, first, there is no certainty as to which of the
types of lower teeth should be referred to H. inopinatus, and secondly,
there is no real assurance that the isolated teeth discussed above are
from one animal, although it seems probable that they are. ,
HAPLOCONUS? ELACHISTUS,” new species
Type.—Left maxillary portion with M? and part of M?, and lower
jaw fragments, U.S.N.M. No. 16191.
Horizon and. locality—Wagonroad Paleocene, Dragon Canyon,
Emery County, Utah.
Specific characters.—Size near that of Conacodon cophater, smaller
than either Haploconus angustus or Haploconus inopinatus. Teeth
relatively a little shorter anteroposteriorly than in H. inopinatus.
Difference between transverse diameters of M? and M? relatively not
so great. Protostyle weak. Lower molars and P, with slight
paraconid.
Description.—Representing Haploconus? elachistus are several iso-
lated teeth and a few jaw and maxillary portions with one or two
teeth. No. 16191, a maxillary portion with M? and part of M’, and
38’ehdxtorTos, Smallest or least, in allusion to size.
PALEOCENE MAMMALS OF CENTRAL UTAH—GAZIN oa
some lower jaw fragments with incomplete teeth and found associ-
ated, is made the type (fig. 29). The teeth are close in size to those
of the nearly contemporaneous Conacodon cophater but more closely
resemble those of species of Haploconus. The form is distinctly
smaller than either Haploconus angustus from the Torrejon or Hap-
loconus inopinatus from the Dragon horizon.
M! and M? resemble these teeth in H. inopinatus, but in addition
to their smaller size do not show so marked a difference between
their transverse diameters as in H. inopinatus; moreover, the upper
molars are relatively a little shorter antero-
posteriorly. The protocone is distinctly
lingual in position, approaching, but not
reaching, the condition seen in Conacodon
cophater. There is a slight protostyle at the
lingual termination of the anterior cingulum,
not so well developed as in H. inopinatus, nor
does the anterior cingulum extend so far
lingually as in @. cophater. In the latter
form the anterior cingulum quite joins the
protocone lingually in M? and M’*. #.?
elachistus also differs noticeably from C.
cophater in the weakness of the external
cingulum. As in later forms of Haploconus,
the external cingulum in H.? elachistus does
not extend across the paracone.
The anteroposterior diameter of M? in the
type is 3.6 mm. The transverse diameter
; elachistus, new species: Por-
from the external cingulum to the base of aoe TeEE eaccilfe eaten
the enamel lingually and at right angles to and part of M! (U.S.N.M.
the direction of the tooth row is about 6.1 No. 16191), type specimen,
mm. lateral and occlusal views,
The lower teeth are much like those in ote Paleo:
Haploconus angustus, except for their peda
smaller proportions. However, the various lower molars referred
to H.? elachistus exhibit a slight, medianly placed paraconid. This is
also true of P, in No. 16548, although P,; in the same specimen,
though not entire, shows no evidence of a paraconid. It is interesting
to note that slight paraconids were observed on the lower molars of
a Torrejon specimen, U.S.N.M. No. 5886, referred to Haploconus cor-
niculatus, as well as on one of the Dragon specimens. The paraconids
of the lower molars of H.? elachistus, however, are not developed as
seen in M' of Conacodon cophater, nor is the talonid portion so com-
pressed anteroposteriorly, and the entoconid, though very well defined,
is not placed so far lingually.
FicurE 29.—Haploconus?
52 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
The presence of a form apparently representing Haploconus in
beds nearly as old as Puerco is interesting in extending downward
the known range of Haploconus and tending to a rather limited
extent to break down certain of the characters separating Haploconus
and Conacodon. Conacodon possesses specialized dental structures
which apparently did not give rise to those seen in Haploconus, but
this earlier form of Haploconus, as represented in the Wagonroad
fauna, shows a less marked separation from the Puerco Conacodon
than do the Torrejon species.
LITERATURE CITED
GazIn, CHARLES LEWIs.
1938. A Paleocene mammalian fauna from central Utah. Journ. Washing-
ton Acad. Sci., vol. 28, pp. 271-277, figs. 1-3.
1939a. Ancient mammals of Utah. Explorations and Field-Work of the
Smithsonian Institution in 1938, pp. 25-28, figs. 22-25.
1939b. A further contribution to the Dragon Paleocene fauna of central
Utah. Journ. Washington Acad. Sci., vol. 29, pp. 273-286, figs.
1-10.
1940. The third expedition to central Utah in search of dinosaurs and
extinct mammals. Explorations and Field-Work of the Smith-
sonian Institution in 1939, pp. 5-8, figs. 5-8.
1941. Trailing extinct animals in central Utah and the Bridger Basin of
Wyoming. Explorations and Field-Work of the Smithsonian Insti-
tution in 1940, pp. 5-8, fig. 6-10.
GILMORE, CHARLES WHITNEY.
1938. Fossil hunting in Utah and Arizona. Explorations and Field-Work
of the Smithsonian Institution in 1937, pp. 1-4, figs. 1-4.
1940. New fossil lizards from the upper Cretaceous of Utah. Smithsonian
Mise. Coll., vol. 99, No. 16, pp. 1-3, figs. 1-2.
GRANGER, WALTER, and SIMPSON, GEORGE GAYLORD.
1929. A revision of the Tertiary Multituberculata. Bull. Amer. Mus. Nat.
Hist., vol. 56, pp. 601-676, figs. 1-34.
JEPSEN, GLENN LOWELL.
1930. Stratigraphy and paleontology of the Paleocene of northeast Park
County, Wyoming. Proc. Amer. Philos. Soe., vol. 69, pp. 463-528,
figs. 1-4, pls. 1-10.
1940. Paleocene faunas of the Polecat Bench formation, Park County,
Wyoming. Part 1. Proc. Amer. Philos. Soc., vol. 83, No. 2, pp.
217-340, figs. 1-22, pls. 1-5.
MATTHEW, WILLIAM DILLER.
1937. Paleocene faunas of the San Juan Basin, New Mexico. Trans. Amer.
Philos. Soc., new ser., vol. 30, pp. i-viii, 1-510, figs. 1-85, pls. 1-65.
RUSSELL, Loris SHANO.
1926. A new species of the genus Catopsalis Cope from the Paskapoo for-
mation of Alberta. Amer. Journ. Sci., vol. 12, pp. 230-234, 1 fig.
SIMPSON, GEORGE GAYLORD.
1986a. Census of Paleocene mammals. Amer. Mus. Nov., No. 848, pp.
1-15.
1936b. Additions to the Puerco fauna, lower Paleocene. Amer. Mus. Nov.,
No. 849, pp. 1-11, figs. 1-6.
PALEOCENE MAMMALS OF CENTRAL UTAH—GAZIN a3
Simpson, GEORGE GAYLoRp—Continued.
1936c. A new fauna from the Fort Union of Montana. Amer. Mus. Noy., No.
878, pp. 1-27, figs. 1-16.
1937a. Additions to the upper Paleocene fauna of the Crazy Mountain
field. Amer. Mus. Nov., No. 940, pp. 1-15, figs. 1-4.
1937b. The Fort Union of the Crazy Mountain field, Montana, and its
mammalian faunas. U. S. Nat. Mus. Bull. 169, pp. i-x, 1-287,
figs. 1-80, pls. 1-10.
SPIEKER, EDMUND MAUTE.
1931. The Wasatch Plateau coal field, Utah. U. S. Geol. Surv. Bull. 819,
pp. i-vi, 1-210, figs. 1-11, pls. 1-83.
SPIEKER, EDMUND MAUvTE, and REESE, JOHN BERNARD, Jr.
1925. Cretaceous and Tertiary formations of the Wasatch Plateau, Utah.
Bull. Geol. Soc. Amer., vol. 36, No. 3, pp. 435-454, figs. 1-3.
US GOVERNMENT PRINTING OFFICE; 1941
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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM
fr
oes Y
2
issued
SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
Vol. 91 Washington : 1941 No. 3122
A NEW FOSSIL CROCODILIAN FROM COLOMBIA
By Caarues C. Moox
Fossim remains of a gigantic crocodilian were collected by Brother
Ariste (Dr. Maurice Rollot) between Neiva and the River Baché
(Colombia) in 1920. The level is not recorded. Dr. J. B. Reeside,
Jr., reports on the basis of invertebrates from nearby localities that
the horizon is probably Lower Cretaceous. These remains consist
of six fairly well preserved vertebrae, with parts of ribs, portions
of maxillary and dentary bones interlocked, several isolated pieces
from the posterior portions of the right and left rami of the lower
jaw, and some fragments. The maxillary portion includes part of
the alveolar series and was evidently situated a short distance
posterior to the maxillo-premaxillary suture. These now constitute
No. 10889 of the collections of the United States National Museum.
I wish to thank C. W. Gilmore, of that institution, for the privilege
of describing this material.
The incomplete nature of this material makes determination of the
relationships extremely difficult if not impossible. Several facts,
however, may be noted. The vertebrae correspond in general char-
acters and somewhat in size with the vertebra described by Gervais
as Dinosuchus terror. The indicated horizon is somewhat lower than
the level of this form, which Gervais notes as “lower Tertiary or
Cretaceous.”
Comparison with the types of Purusaurus brasiliensis Rodriguez
and Brachygnathosuchus braziliensis Mook shows clearly that the
form described has no close relation with either. These species, while
gigantic, have relatively short and broad lower jaws, with large
alveoli, while the form described has relatively long and slender lower
jaws and posterior teeth, at least, of relatively small size.
* Contributions to the Osteology, Affinities, and Distribution of the Crocodilia, No. 35.
406805—41 55
56 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 01
In view of these facts the material described is referred to a new
species of the genus Dinosuchus Gervais (non Holland), which may
be called Dinosuchus neivensis, named for the city of Neiva near
which it was found.
Genus DINOSUCHUS Gervais, 1876
Generic characters—As Gervais never separated the generic char-
acters from those of the species D. terror, the following designation
may be given: Size gigantic, vertebrae procoelian and massively
constructed.
Relationships—The genera Dinosuchus Gervais, Purusaurus
Rodriguez, and Brachygnathosuchus Mook have been treated quite
differently by recent authors. Nopesa, in 1924, considered
Brachygnathosuchus to be a synonym of Purusaurus, and Dinosuchus
to be independent. Because of the latter interpretation he proposed
the name Phobosuchus for Holland’s Deznosuchus. Mook, in 1934,
considered Purusaurus to be a synonym of Dinosuchus, and
Brachygnathosuchus to be independent. Patterson, in 1936, consid-
ered Brachygnathosuchus to be a synonym of Dinosuchus, and
Purusaurus to be a synonym of Caiman of Spix.
At the present time it appears most consistent with the incom-
pletely known characters of these forms and with their geologic
levels to consider the Cretaceous Dinosuchus to be valid and inde-
pendent, and to consider the upper Miocene or lower Pliocene
Purusaurus and Brachygnathosuchus to be valid and to be closely
related to Caiman.
DINOSUCHUS NEIVENSIS, new species
PLATES 4-9
Specific characters —External mandibular foramen unusually large
in proportion to the size of the jaw elements surrounding it, jaw
relatively long and slender, posterior teeth relatively small and close
together.
Description of material—F¥ ive maxillary alveoli are visible on this
specimen. The first is large and is slightly longer than it is broad.
The second is larger than the first. Its external border is incomplete;
consequently its proportions ave difficult to determine. The last
three alveoli are approximately equal to the first in size; they appear
to be subcircular, although their borders are not entirely visible.
Badly mutilated stumps of teeth are visible in these alveoli.
he anterior and posterior ends of the lower jaw section that is
attached to the portion of the maxillary noted above exhibit sections
of alveoli 12 cm. deep and fragments of teeth of corresponding size.
Another section of the right ramus was located much farther back
than the one noted above. The anterior end of the right external
A NEW FOSSIL CROCODILIAN—MOOK 57
mandibular foramen is located at the posterior end of this section
and the posterior end of the alveolar row at the center of the superior
border locates the position of the section in the ramus. Four alveoli
with bases of teeth are clearly visible, and a fifth or last is somewhat
obscure. These alveoli are much smaller than those of the maxil-
lary section noted above, and their height, as indicated by the
anterior surface of the section, is less than half that of the anterior
mandibular teeth. The mandibular cavity, now indicated by matrix,
was large, the bony substance being thin.
The left ramus is represented by a larger section, about 48 cm. long
and composed of two pieces that make clean-cut contacts with each
other. This section is entirely posterior to the alveolar row and in-
cludes the external mandibular foramen, of which the superior
boundary is incomplete. The posterior end of this section is near the
posterior end of the ramus immediately anterior to the glenoid surface.
The sutures separating the elements of which this part of the jaw is
composed are indistinct, the dentary, angular, and surangular bones
being almost indistinguishable from one another.
The external mandibular foramen is unusually long and is not very
high. The exact relation between length and height cannot be made
out because of the incomplete superior border. On comparing the
length of this opening with that of an 84-cm. ramus of Crocodylus
acutus, and assuming that the proportions between the total length and
the length of the foramen are the same in that species and the form now
described, we estimate that the total length of the ramus would be 280
em., or about 9 feet. Comparison with a 32-cm. ramus of Caiman
crocodilus indicates a total length of 172 cm., or about 524 feet, which
is more likely.
One of the vertebral units is composed of the intercentrum of the
atlas, most of the axis, and the proximal portions of the atlas and axis
ribs in natural positions. The atlas intercentrum is a broad, flat bone,
much more distinctly bifurcated posteriorly than in C. acutus. The
atlas ribs attach to the bifurcations and their axes of breadth lie below
the axis and the, axis ribs. The atlas ribs are single headed, of course,
and are considerably thickened where they attach to the atlas inter-
centrum.
The characters of the axis are not particularly distinctive except for
the size and strength of the processes to which the ribs are attached.
The ribs themselves are distinctly two-headed, the upper element, or
tuberculum, being slightly larger than the lower one, or capitulum.
The shaft is slender and is situated on edge, at right angles to the posi-
tion in which the atlas ribs are situated.
Six other vertebrae are preserved, but none of them is complete.
Two of these united together, with a fragment of a third, are cervicals,
58 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
probably 4 and 5. The spines and the postzygapophyses are not pre-
served. The prezygapophyses, diapophyses, and parapophyses are
incompletely preserved. The centrum of the first vertebra of the pair
is incomplete. That of the second is complete and is moderately long,
rather low vertically and narrow posteriorly but broad anteriorly, ap-
parently convex posteriorly, but the degree of convexity cannot be
made out. The prezygapophyses and diapophyses of this vertebra
are incomplete, but enough of them is preserved to indicate that they
were very stout. There is a very small median hypapophysial keel
near the anterior end of the centrum. On the whole the vertebrae
appear small for the size of the mandible. The capitular and tuber-
cular ends of the left rib of the anterior of the two vertebrae are pre-
served ; they are very stout, especially the tubercular process.
MEASUREMENTS (IN MILLIMETERS)
Length of two large contact pieces of left ramus of mandible_-____~ = eS)
Maximum *hereht of isames sae S02 sh See eae Eee oe ee See vale WN Een NALEGA
juength of external mandibularsonramens as. .s0. 2 eae eee 265
Height OL GSA Ses 2 ee eS ee ee ee ee 56
Length over four posterior alveoli, right ramus of mandible______________ 82
Height of maxillary and dentary fragments in place with each other____ 211
Height of anterior mandibular tooth shown in end of this fragment______ 94
Breadth acrosssatlas centrum posterior end = ae mae seer eee eee 88
eng thi of tatlalsi«e@en Gren sees = I a a a a ee 7
Breadth across: rightvatlas) ribvat proximally endea 49
Breadth /acrossuleft atlas Tibvat proximal vend] sss eee eee AT
Breadth across tuberculum end capitulum of right axis rib__--_________ 43
Breadth across tuberculum end capitulum of left axis rib_-_------________~ 746
Breadth across) 2xdS) CENCEUIM ypPOSter1LOT CnC see ae ee ee ee ee 60
WencuhvornttniCe) cervical centrum 2 ena se eee eee 83
Breadth of fifth(?) cervical centrum anterior end__------_____________ 103
Breadth of fifth(?) cervical centrum posterior end___--_-_______-_____-__ *70
Breadth of fifth(?) cervical vertebra across prezygapophyses_____--_-__-_ OT
* Estimate.
LITERATURE CITED
GERVAIS, PAUL.
1876. Crocodile gigantesque fossile au Brésil. Journ. Zool., vol. 5, pp.
233-236, 1 pl.
Mook, CHARLES CRAIG.
1921. Brachygnathosuchus braziliensis, a new fossil crocodilian from Brazil.
Bull. Amer. Mus. Nat. Hist., vol. 44, pp. 48-49, 4 figs.
1934. The evolution and classification of the Crocodilia. Journ. Geol., vol.
42, pp. 295-304, 1 fig.
Noprcsa, FRANZ BARON.
1924. Uber die Namen einiger brasilianischer fossiler Krokodile. Ceniralbl.
Min., Geol. und Pal., 1924, p. 378.
PATTERSON, BRYAN.
1936. Caiman latirostris from the Pleistocene of Argentina, and a sum-
mary of South American Cenozoie Crocodilia. Herpetologica, vol.
1, pp. 48-54, 1 pl.
RODRIGUES, JOAO BARBOSA.
1892. Les reptiles fossiles de lo vallée de l’Amazona. Vellosia, yol. 2
(1885-88), pp. 41-56, 16 pls.
U.S. GOVERNMENT PRINTING OFFICE 1941
U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 91 PLATE 4
DINOSUCHUS NEIVENSIS, NEW SPECIES.
Type (U.S. N. M. No. 10889): Parts of left premaxillary and dentary bones, external view. One-half
natural size.
U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 91 PLATE 5
DINOSUCHUS NEIVENSIS, NEW SPECIES.
1. Type (U.S. N. M. No. 10889): Central portion of right dentary bone, superior view. One-half natural size.
2. Same, external view. One-half natural size.
PLATE 6
PROCEEDINGS. VOL. 91
U. S. NATIONAL MUSEUM
“OZIS [Pinqeu pry
2uQ)
“MO!
A [Bute] x9 ‘a[qipuru jo snued de] Jo uon4od AOMISOd >(6880T “ON (WSN 'S ‘) ada 7,
“SAIDAdS MAN ‘SISNSAIFN SNHONSONIG
U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 91 PLATE 7
DINOSUCHUS NEIVENSIS, NEW SPECIES.
1. Type (U.S. N. M. No. 10889): Parts of atlas and axis vertebrae and of atlas and axis ribs, lateral
view, left side. One-half natural size.
2. Same, inferior view. One-half natural size.
U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 91 PLATE 8
DINOSUCHUS NEIVENSIS, NEW SPECIES.
1. Type (U.S. N. M. No. 10889): Cervical vertebrae, probably fifth and sixth, lateral view, left side.
One-half natural size.
2. Dorsal vertebrae, probably fifth and sixth, lateral view, left side. One-half natural size.
U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 91 PLATE 9
DINOSUCHUS NEIVENSIS, NEW SPECIES.
Type (U.S. N. M. No. 10889): Vertebrae, position in series uncertain, lateral view, left side. One-half
natural size.
PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM
SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
Vol. 91 Washington: 1941 No. 3123
THE NORTH AMERICAN MOTHS OF THE GENUS
ARACHNIS, WITH ONE NEW SPECIES
By J. F. Gates Ciarke
Tus study of the genus Arachnis (family Arctiidae) was undertaken
to determine the exact relationship of the new species described to
the known species, and, in order to accomplish this, characters for
all species in the group needed to be critically reviewed and evaluated.
The species of this group are extremely plastic and readily produce
forms and races apparently constant in coloration. These may be
confined to small islands within the range of the species or may occur
along with the typical race.
The lack of sufficient material has probably prevented a proper
evaluation of characters in one or two instances, but it seems apparent
that at least one species, picta, has given rise to numerous varieties
and races that are so distinct superficially that they appear to be
separate species. The case of midas, for example, is striking. This
so-called species, although easily distinguishable from picta on color-
ation, can be separated from it morphologically only by the shape of
the uncus. As pointed out later, médas is represented only by the
unique type, and the distinguishing character of the genitalia might
well be only one of several variations. Since the matter of coloration
seems to be of little importance in the separation of species, m2das, like
citra, may be nothing more than a form or race of picta.
The genus appears to be best represented in the southwestern part
of the United States, but its distribution ranges into Mexico and to
the Midwest and Florida. It is in the Rocky Mountain region that
408320—41 59
60 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
the predisposition to variation is greatest, more stability being apparent
to the east and west beyond the intermountain area.
The larvae are probably rather general feeders, a character common
to many arctiids, but only a few have been reared.
Dr. J. A. Comstock, director of science in the Los Angeles Museum,
kindly sent material for study, for which thanks are due.
A diagnosis of this well-known genus is not included, but descrip-
tions of the genitalia follow.
The drawings for this paper were made by Mrs. Eleanor A. Carlin,
of the Bureau of Entomology and Plant Quarantine.
Genus ARACHNIS Geyer
Arachnis Geyer, in Hiibner, Zutriige exotischer Schmetterlinge, vol. 5, p. 28,
1837.
Male genitalia.—Harpe broadly attached at base, long, slender, al-
ways with inward lateral projection. Anellus semicylindrical, some-
times concave laterally. Aedeagus long, stout, dorsoventrally curved ;
vesica with numerous minute scobinations. Vinculum with well-
developed winglike lateral expansion. Tegumen with well-developed
dorsal flange.
Female genitalia—Ostium large, extending well beyond ventral
surface of genital plate. Ductus bursae strongly sclerotized, some-
what depressed, concave ventrally. Ductus seminalis greatly enlarged,
membranous or partly sclerotized, and entering at confluence of ductus
bursae and bursa copulatrix. Bursa copulatrix with two small,
round, scobinate signa. Occasionally a third signum is weakly
developed. Dorsal glands well developed, with several branches.
Remarks.—The lateral projection of the harpe does not seem to
represent a clasper or an ampulla, but rather no more than an out-
growth of the ventral margin.
KEYS TO THE SPECIES OF ARACHNIS
Coloration
ie ind wangewathyellowishyercoulds Colo tee er 2
Hind wing with red or reddish ground color__-__---------------~---~--- 3
2. Thorax with conspicuous white posteromedian dorsal spot ; dark
Markings OftOre wingaSlate== === === ae zuni Neumoegen (p. 69)
Thorax without white posteromedian dorsal spot; dark mark-
ings of fore wing gray_-—_------—_- midas Barnes and Lindsey (p. 69)
3. Fore wing with white or whitish ground color___----------------~-------~ 5
Fore wing with ground color otherwise_————__-_-—*__-_ = 4
4. Fore wing with yellowish ground color.
picta citra Neumoegen and Dyar (p. 67)
Fore wing with cerise ground color___--_-~- apachea, new species (p. 68)
5: Abdomen with laterals row. of Oranee (SOUS === =e 6
Abdomen with lateral row of gray to blackish spots____-____-------------- Uf
MOTHS OF THE GENUS ARACHNIS—CLARKE 61
6. Hind wing almost wholly overlaid with blackish fuscous; dark
markings of fore wing dark slate gray, sharply contrasted
against white ground color__________-___~_ aulaea pompeia Druce (p. 63)
Hind wing with dark markings lighter and less abundant; dark
markings of fore wing lighter and not so sharply contrasted
With ywhitishyeround colors: == = see se ees aulaea Geyer (p. 62)
7. Fore wing with at least basal half of underside entirely shaded
Sd a na CTSNet ee 2 hls ee eh 8
Fore wing with basal half or two-thirds of underside of costa
Oniyeshaded {wilt ONG ee =. tA aes ee a 10
8. Gray markings of fore wing strongly outlined with black________________ 9
10.
Gray markings of fore wing without black outlines.
picta insularis Clarke (p. 66)
. Hind wing of male with outer band of gray spots broken but
strongly defined ; female with outer band entire or, if broken,
EhengonlysOncCe. 22= = 25 ee ee ee a picta Packard (p. 63)
Hind wing of male with outer band consisting of three or four
small spots; female with outer band consisting of four spots,
apical pair sometimes fused.
picta verna Barnes and McDunnough (p. 65)
Hind wing of male semihyaline; female with basal band, on
underside, connected to base by a narrow gray line.
picta maia Ottolengui (p. 66)
Hind wing of male not semihyaline; female with basal band, on
underside, connected to base by conspicuous gray triangle.
picta hampsoni Dyar (p. 66)
Male genitalia
. Uneus flattened, with prominent dorsal ridge; lateral projection
of harpe as narrow as, or narrower than, distal part of harpe
beyondmta( plat. es 4) ee eee aulaea Geyer (p. 62)
Uncus conical, without dorsal ridge; lateral projection wider
than distal part of harpe beyond it (pl. 10, fig. 3; pl. 12,
PSA Cie Ol) ae ee ew a ee ee ee 2
Distal portion of harpe greatly dilated (pl. 10, fig. 3).
apachea, new species (p. 68)
Distal portion of harpe not greatly dilated (pl. 12, figs. 7, 8) _-_-_-___________ 3
. Lateral projection of harpe bent toward base; distal end narrow,
somewhat compressed (pl. 11, fig. 5) _--____-_-_____ zuni Neumoegen (p. 69)
Lateral projection of harpe not bent toward base; distal end
SWVC ST eee es ae sen en cee ed it ee ae ee ee ee ee 4
Uncus short, stocky, evenly curved (pl. 12, fig. 7c)_-_-- picta Packard (p. 63)
Uncus long, slender, angulate (pl. 12, fig. 8c).
midas Barnes and Lindsey (p. 69)
Female genitalia*
a uetus seminalis at least partly: sclerotized=—2=+ 2-3 * fee Zr
Ductus seminalis wholly membranous________-_______ picta Packard (p. 63)
Median fleshy protuberance of ostium with broad, sickle-shaped,
sclerotized area on each side (pl. 10, fig. 2) __-_______ aulaea Geyer (p. 62)
Median fleshy protuberance of ostium without such area (pl. 11, fig. 6).
zuni Neumoegen (p. 69)
+The females of midas and apachea are unknown to me.
62 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
ARACHNIS AULAEA Geyer
PLATE 10, FieuREs 2—2a; PLatE 11, FicurEes 4-40
Arachnis aulaea Geyer, in Hiibner, Zutriige exotischer Schmetterlinge, vol. 5,
p. 28, figs. 918, 914, 1837 —CremeEns, Proc. Acad. Nat. Sci. Philadelphia, 1860,
p. 526.—WALKER, List of the specimens of lepidopterous insects in the collec-
tion of the British Museum, vol. 31 (Suppl. 1), p. 300, 1864.—Srrercy, Ilus-
trations of the Zygaenidae and Bombycidae of North America, vol. 1, p. 85,
1873.—Drvce, Biologia Centrali-Americana, Heterocera, vol. 1, p. 98, 1884.—
SmitTH, List of the Lepidoptera of Boreal America, No. 1118, 1891.—Kirgpy,
A synonymic catalogue of the Lepidoptera Heterocera (moths), vol. 1, p. 218,
1892.—Druce, Ann. Mag. Nat. Hist., ser. 6, vol. 18, p. 174, 1894.—OrroLENGuI,
Ent. News, vol. 7, p. 126, pl. 4, 1896.—Drucr, Biologia Centrali-Americana,
Heterocera, vol. 2, p. 377, 1897—HA4Amepson, Catalogue of the Arctiadae
(Arctianae) and Agaristidae in the collection of the British Museum, vol. 3,
pp. 389, 390, 391, fig. 163, 1901 [biology].—Barnrs and McDuNNnovuen, Check
list of the Lepidoptera of Boreal America, No. 967, 1917.—SrTrAnp, Lepi-
dopterorum catalogus, pt. 22, p. 278, 1919.—Sr1Tz, Die Gross-Schmetterlinge
der Erde, vol. 6, p. 314, pl. 40b, 1919.—Barnes and BENJAmIN, Pan-Pac. Ent.,
vol. 3, p. 17, 1926.—McDunnouaH, Check list of the Lepidoptera of Canada
and the United States of America (Part 1, Macrolepidoptera), No. 1080, 1938.
Ecpantheria aulaea (Geyer) Botspuvat, Ann. Soc. Ent. Belg., vol. 12, p. 78, 1869.—
OBERTHUR, Etudes d’Entomologie, vol. 6, p. 111, pl. 19, figs. 4, 7, 1881.—
BuRMEISTER, Ann. Mus. Publ. Buenos Aires, vol. 3, p. 31, 1883.
Ecpantheria aulea ScHaus (misspelling for aulaea), Papilio, vol. 3, p. 188, 1883
[larva].
Arachnis aulea (Schaus) H. Epwarps, U. 8. Nat. Mus. Bull. 35, p. 61, 1889.—
ScHaus, Ent. Amer., vol. 5, p. 190, 1889—NrUMOEGEN and DyAr, Journ. New
York Ent. Soe., vol. 1, p. 178, 1893.—Dyar and Dot1, Ent. News, vol. 4, p. 312,
1893 [larva].—Dyar, Can. Ent., vol. 26, p. 307, 1894 [larva]; Proc. Ent. Soe.
Washington, vol. 14, p. 55, 1912.
Ecpantheria incarnata WALKER, List of the specimens of lepidopterous insects in
the collection of the British Museum, vol. 3, p. 690, 1855.—BurMEIstTer, Ann.
Mus. Publ. Buenos Aires, vol. 3, p. 31, 1883 [as synonym of H. aulaea].
Arachnis incarnata SmitH, List of the Lepidoptera of Boreal America, No. 1118,
1891.—Kuirpy, A synonymic catalogue of the Lepidoptera Heterocera (moths),
vol. 1, p. 218, 1892.—_Barnres and McDunnoueH, Check list of the Lepidoptera
of Boreal America, No. 967, 1917.—Srranp, Lepidopterorum catalogus, pt. 22,
p. 278, 1919.—Sr1rz, Die Gross-Schmetterlinge der Erde, vol. 6, p. 314, 1919.—
McDunnouGH, Check list of the Lepidoptera of Canada and the United States
of America (Part 1, Macrolepidoptera), No. 1080, 1938 [as synonym of A.
aulaea].
Male genitalia—Warpe with slender, inward, lateral projection;
cucullus narrow, scarcely wider than lateral projection of harpe,
slightly swollen distally. Anellus with sides parallel. Aedeagus with
well-developed distolateral flap. Vinculum broad, short, truncate.
Uncus broad, flattened, with prominent dorsal ridge extending beyond
end to form terminal point. Flange of tegumen broadly rounded.
Female genitalia—Median protuberance of ostium fleshy, bulbous,
with conspicuous, sickle-shaped, sclerotized area laterally. Ductus
MOTHS OF THE GENUS ARACHNIS—CLARKE 63
seminalis sclerotized for distance almost equal to length of ductus
bursae.
Alar expanse, 38-60 mm.
Distribution —Southwestern part of the United States and Mexico.
Arizona: Huachuca Mountains, 2 (no date or collector) ; Palmerlee, Cochise
County, ¢ (“VIII’; no collector).
New Mexico: “New Mexico,” 2 (no other data).
Texas: “Southern Texas,” ¢ (no other data).
Types—Unknown (aulaea); in the British Museum (incarnata).
Type localities—Mexico (aulaca and inearnata).
Food plants—Numerous (ace. Schaus, 1889).
Remarks.—This species seems to be essentially a Mexican insect,
since the preponderance of specimens before me is from Mexico. The
few records from the United States are scattered and not altogether
reliable.
ARACHNIS AULAEA POMPEIA Druce
PLATE 10, Ficures 1-la
Arachnis pompeia Druck, Ann. Mag. Nat. Hist., ser. 6, vol. 18, p. 174, 1894; Biolo-
gia Centrali-Americana; Heterocera, vol. 2, p. 377, pl. 75, figs. 2, 3, 1897.—
Hampson, Catalogue of the Arctiadae (Arctianae) and Agaristidae in the
collection of the British Museum, vol. 8, pp. 389, 390, 1901.—Srranp, Lepi-
dopterorum catalogus, pt. 22, p. 279, 1919.—Srrrz, Die Gross-Schmetterlinge
der Erde, vol. 6, p. 315, 1919—BARrNEs and BENJAMIN, Pan-Pac. Ent., vol. 3,
p. 17, 1926—McDunnoueH, Check list of the Lepidoptera of Canada and the
United States of America (Part 1, Macrolepidoptera), No. 1081, 1938.
Arachnis aulaea Hotianp [not Geyer], The moth book, p. 124, pl. 16, fig. 1, 1903.—
Barnes and McDunnovueH, Contr. Nat. Hist. Lepid. North Amer., vol. 1, No. 4,
p. 7, pl. 2, fig. 1, 1912.
Alar expanse, 47-52 mm.
Type.—tIn the British Museum.
Type locality—Mexico, near Durango City.
Remarks.—The racial status of pompeia (known from the female
only) is doubtful, and the genitalia indicate that it may be no more
than a form of aulaea occurring along with the typical race. This
form can be distinguished from aulaea by the darker and move con-
trasting markings.
The specimen figured by Barnes and McDunnough? as aulaea is in
the U. S. National Museum. This specimen is pompeia and was mis-
identified by Barnes and McDunnough.
ARACHNIS PICTA Packard
PLATE 12, Fiaures 7—7c, 9-9a
Arachnis picta PACKARD, Proc. Ent. Soc. Philadelphia, vol. 3, p. 126, 1864—
WALKER, List of the specimens of lepidopterous insects in the collection of
the British Museum, vol. 85 (Suppl. 5), p. 1912, 1866.—Srrercu, Illustrations
7Contr. Nat. Hist. Lepid. North Amer., vol. 1, No. 4, p. 7, pl. 2, 1912.
64. PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
of the Zygaenidae and Bombycidae of North America, vol. i, ps 83, Dbl. 3;
fig. 6, 1873.—OxsertHuR, Etudes d’Entomologie, vol. 6, p. 112, pl. 19, figs. 5, 8,
1881.—Drucer, Biologia Centrali-Americana, Heterocera, vol. 1, p. 98, 1884.—
H. Epwarps, U. 8S. Nat. Mus. Bull. 35, p. 61, 1889 [food plant].—
Dyar, Ent. Amer., vol. 6, p. 73, 1890 [larva, cocoon, pupa].—SMITH,
List of the Lepidoptera of Boreal America, No. 1117, 1891.—Kirgsy, A syno-
nymic catalogue of the Lepidoptera Heterocera (moths), vol. 1, p. 218, 1892.—
NEUMOEGEN and Dyar, Journ. New York Ent. Soc., vol. 1, pp. 178, 179, 1893.—
Orrotencul, Ent. News, vol. 7, p. 124, pl. 4, 1896.—Hampson, Catalogue of
the Arctiadae (Arctianae) and Agaristidae in the collection of the British
Museum, vol. 3, pp. 389, 392, 1901.—Dvyar, U. S. Nat. Mus. Bull. 52, No. 857,
1903.—SmirH, Check list of the Lepidoptera of Boreal America, No. 36,
1903.—HoLLanp, The moth book, p. 124, pl. 16, fig. 2, 1903.—Barnes and
McDunnovucnH, Check list of the Lepidoptera of Boreal America, No. 968,
1917; Contr. Nat. Hist. Lepid. North Amer., vol. 4, p. 90, 1918.—STRAND,
Lepidopterorum catalogus, pt. 22, p. 279, 1919.—Sr1rz, Die Gross-Schmetter-
linge der Erde, vol. 6, p. 315, pl. 40b, 1919.—Essia, Insects of western North
America, pp. 581, 583, 678, 1926 [parasites of, larva, food plants]—McDun-
NouGH, Check list of the Lepidoptera of Canada and the United States of
America (Part 1, Macrolepidoptera), No. 1082, 1938.
Ecpantheria picta (Packard) BurMetstrer, Ann. Mus. Publ. Buenos Aires, vol. 3,
p. 31, 1888 (as synonym of LH. aulaea).
Male genitalia—Lateral process of harpe with posterior edge
smooth, much broader than portion of harpe beyond it; distal end
fleshy, slightly dilated apically. Anellus strongly concave laterally.
Aedeagus with poorly developed distolateral flap; scobinations of the
vesica weak. Vinculum narrowly rounded. Uncus short, stocky,
evenly curved.
Female genitalia—Median protuberance of ostium broad, flat-
tened, without sickle-shaped sclerotized lateral area. Ductus seminalis
membranous.
Alar expanse, 33-62 mm.
Distribution—Southern part of the United States northward to
Illinois, Utah, and northern California and southward into Mexico,
Arizona: Oak Creek Canyon, 2 (6,000 feet, July, F. H. Snow); Prescott, 9?
(“VII,” collector not given).
California: Alameda County, 2¢ ¢, 2 (September, October; no collector) ; Los
Angeles, 6,2 292 (25—-X-1889, H. G. Dyar No. 4084; 26—-X-—1889, H. G. Dyar
Nos. 4190, 4208) ; Los Angeles County, ¢ (no date or collector) ; Sacramento,
@ (no date or collector); San Diego, ¢ (16—X—1909, George H. Field),
2992 (14-X-22; 10-X-23; no collector); San Francisco County, 2¢ 6,
2292 (September and October; no collector) ; several males and females
labeled “Middle California” and “Southern California.”
Colorado: @ (no date; “Bruce’’).
Florida: Palm Beach, ¢ (4-II-1890, H. G. Dyar No. 4552).
Illinois: Quincy, 2 (no date; Poling).
New Mexico: Jemez Springs, 2 (no date or collector).
Utah: 2 (no other data).
MOTHS OF THE GENUS ARACHNIS—CLARKE 65
Type.—tIn the Museum of Comparative Zoology, Cambridge, Mass.
Type locality—San Francisco, Calif.
Food plants —Alfalfa, clover, geranium, lupine, J/alva, rose, sage-
brush, ete.
Remarks.—The genitalia of picta and its varieties show consider-
able variation, but no characters present are sufficiently stable to
enable the absolute separation of one from the other by the use of these
organs. The typical subspecies (picta picta) shows the most con-
sistent form. The lateral projection of the harpe of this subspecies
is usually much thicker than in the others and the posterior edge of
the projection is comparatively smooth. In the other subspecies the
lateral projection varies in thickness and is usually roughened on the
posterior edge.
In addition to the material listed under distribution I have before
me two specimens from Avalon, Santa Catalina Island, Calif.
(2-X-1931, 11-X-1931, Don Meadows), which appear to be an island
race of picta. The gray markings are very light and coalesced and
not sharply defined. The thorax, head, and fore wing have a powdered
appearance. Until more material comes to hand and it is possible
to determine the constancy of this form I am leaving it unnamed.
This race falls between picta and verna in my key.
These specimens were sent to me by Dr. J. A. Comstock, of the Los
Angeles Museum.
ARACHNIS PICTA VERNA Barnes and McDunnough
Arachnis picta verna BARNES and McDuNNovuaH, Contr. Nat. Hist. Lepid. North
Amer., vol. 4, p. 90, pl. 18, figs. 5, 6, 1918—McDuNnnouaH, Check list of the
Lepidoptera of Canada and the United States of America (Part 1, Macrolepi-
doptera), No. 1082c, 1938.
Alar expanse, 45-73 mm.
Distribution —Middle California to Utah.
California: Three Rivers, Tulare County, 3 ¢ 4,6 9 @ (no dates or collector).
Utah: Dividend, 3 ¢6¢, 2 (August and September dates; Tom Spalding) ;
Eureka, 6 $46, 3 992 (August and September dates, 1910 to 1921, Tom
Spalding) ; Provo, ¢, 2 (20-IX—1908; 25-VIII-1908, Tom Spalding).
Type.—tIn the U. S. National Museum.
Type locality —Three Rivers, Tulare County, Calif.
Remarks.—This variety averages slightly larger than typical picta
and has more of the whitish or pale-gray ground color showing, thus
appearing considerably lighter. The dark markings of the hind wing
are reduced in verna.
While this race is at present known only from two rather small areas
it may be found throughout much of the area between California and
the Rocky Mountains, even though this particular species appears to
produce rather restricted races.
66 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
In addition to the specimens listed above, I have before me one
other from Logan Canyon, Utah (August 16, 1939, G. F. Knowlton
No. 34), which appears to belong here. This specimen, however, lacks
the usual median dorsal black line of the abdomen, and the hind wing
is more cerise, with the dark spots greatly reduced.
' ARACHNIS PICTA INSULARIS Clarke
Arachnis picta insularis CLARKE, Bull. Southern California Acad. Sci., vol.
39, p. 187, 1941 [egg, food plant].
Alar expanse, 34-54 mm.
Type.—tIn the U. S. National Museum.
Type locality.—Anacapa Island, Calif.
Food plant.—Plantago (laboratory).
Remarks.—This subspecies is known only from the type locality.
ARACHNIS PICTA MAIA Ottolengui
Arachnis maia, OTTOLENGUI, Ent. News, vol. 7, p. 125, pl. 4, 1896.
Arachnis picta. maia HAmpson, Catalogue of the Arctiadae (Arctianae) and
Agaristidae in the collection of the British Museum, vol. 3, p. 392, 1901.—
Dyar, U. S. Nat. Mus. Bull. 52, No. 857a, 1903.—Smiru, Check list of the
Lepidoptera of Boreal America, No. 946a, 1903.—BarNnes and McDuNNoUGH,
Check list of the Lepidoptera of Boreal America, No. 968a, 1917; Contr. Nat.
Hist. Lepid. North Amer., vol. 4, p. 90, pl. 18, figs. 7, 8, 1918—StTranp, Lepi-
dopterorum catalogus, pt. 22, p. 279, 1919.—Sr1rz, Die Gross-Schmetterlinge
der Erde, vol. 6, p. 815, 1919.—BarNnkEs and LINDSEY, Ent. News, vol. 32, p.
297, 1921.—McDunnovueH, Check list of the Lepidoptera of Canada and the
United States of America (Part 1, Macrolepidoptera), No. 1082a, 1938.
Alar expanse, 44-58 mm.
Distribution—Southern Rocky Mountain region.
Colorado: Chaffee County, ¢, 9 (no date; Bruce) ; Glenwood Springs, ¢ (August
1894 ; W. Barnes) ; Salida, ¢, 2 9 2 (no date or collector) ;11 6 6 (“Colo.”
Bruce).
New Mexico: Las Vegas, ¢ (’89, H. Meske).
Type.—tin the U. 8S. National Museum.
Type locality.—Las Vegas, N. Mex.’
Remarks.—Males of this race are easily distinguishable from picta
by their coloration, but the females are distinguishable only by the
key character, which, although probably rather constant, might fail
to separate the two in borderline cases.
ARACHNIS PICTA HAMPSONI Dyar
Arachnis picta hampsoni DyAr, U. S. Nat. Mus. Bull. 52, No. 857c, 1903.—SmirH,
Check list of the Lepidoptera of Boreal America, No. 946c, 19038.—BaARNES
and McDunnoveH, Check list of the Lepidoptera of Boreal America, No.
968e, 1917; Contr. Nat. Hist. Lepid. North Amer., vol. 4, p. 90, 1918.—StTrAnp,
8 See “Errata,” Ent. News, vol. 7, p. 160, 1896.
MOTHS OF THE GENUS ARACHNIS—CLARKE 67
Lepidopterorum catalogus, pt. 22, p. 279, 1919.—Srrrz, Die Gross-Schmetter-
linge der Erde, vol. 6, p. 815, 1919—McDunnoueu, Check list of the Lepidop-
tera of Canada and the United States of America (Part 1, Macrolepidoptera),
No. 1082d, 1938.
Alar expanse, 45-65 mm.
Distribution.—Southwestern part of the United States.
Arizona: Flagstaff, ¢ (July; no other data) ; Huachuca Mountains, Q@ (no
date or collector) ; Mojave County, 2¢ 6 (August 8-16; no collector) ; Para-
dise, Cochise County, ¢, @ (August; no collector) ; Cochise County, ¢,
8292 (26-VI-1917; 31—-VII-1917; no collector); Phoenix, ¢ (no date or
collector); Prescott, 7¢ 6, 422 (July and August dates; no collector) ;
Yavapai County, 36 6,222 (August; O. Buchholz).
California: Los Angeles, 2¢ 4, 422 (October; V. M. Owen); San Diego,
11¢ 4,522 (September, October, 1921; no collector).
New Mexico: Jemez Springs, ¢, 2 (no date or collector).
Neotype.—In the U. S. National Museum.
Type locality—Jemez Springs, N. Mex.
Remarks.—This race was described by Hampson‘ as “Subsp. 2” of
picta but was not named. Dyar® named this race hampsoni but did
not designate a type. I now designate a male specimen from Jemez
Springs, N. Mex., in the U. S. National Museum, as neotype, since
New Mexico is the first locality cited by Hampson.
ARACHNIS PICTA CITRA Neumoegen and Dyar
Arachnis picta citra NEUMOEGEN and DyAr, Ent. News, vol. 4, p. 140, 1893 ; Journ.
New York Ent. Soc., vol. 1, p. 179, 1893.—OrroLreneul, Ent. News, vol. 7, p. 124,
126, pl. 4, 1896—HaAmpson, Catalogue of the Arctiadae (Arctianae) and
Agaristidae in the collection of the British Museum, vol. 2, p. 393, 1901.—
Dyar, U. S. Nat. Mus. Bull. 52, No. 857b, 1903—SmitruH, Check list of the
Lepidoptera of Boreal America, No. 946b, 19038.—BarRNEs ang McDUNNOUGH,
Check list of the Lepidoptera of Boreal America, No. 968b, 1917; Contr. Nat.
Hist. Lepid. North Amer., vol. 4, p. 90, 1918.—StTranp, Lepidopterorum cata-
logus, pt. 22, p. 279, 1919.—Srrrz, Die Gross-Schmetterlinge der Erde, vol. 6,
p. 315, pl. 40b, 1919.—McDunnoueH, Check list of the Lepidoptera of Canada
and the United States of America (Part 1, Macrolepidoptera), No. 1082b,
1988.
Alar expanse, 46-74 mm.
Distribution—Southwestern part of the United States.
California: ¢ (no other data).
Colorado: Glenwood Springs, 25¢ 6, 162 9 (August and September dates, W.
Barnes) ; 56 ¢, 722 (‘‘Colo.” Bruce).
Utah: Cisco, ¢ (16-VIII-1939, G. F. Knowlton and F. C. Harmston).
Type—tn the U. S. National Museum.
Type locality—Western Colorado.
4Hampson, G. F., Catalogue of the Arctiadae (Arctianae) and Agaristidae in the collec-
tion of the British Museum, vol. 3, p. 392, 1901.
5 Dyar, H. G., U. S. Nat. Mus. Bull. 52, No. 857¢, 1903.
68 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
Remarks.—The single male from the Oberthur collection labeled
“California” is probably mislabeled. The preponderance of specimens
from Colorado and the single specimen from Cisco, Utah, indicate that
the population of this variety is restricted in distribution to the
mountainous area centering about Colorado.
ARACHNIS APACHEA, new species
Pirate 10, Figures 3-8¢
Antenna with basal segment cerise anteriorly, buff posteriorly ;
shaft blackish fuscous; basal two-fifths cream colored above and
faintly annulated with cerise; outer three-fifths overlaid with pale
eray above. Labial palpus whitish ochreous; basal segment with a
conspicuous black spot exteriorly ; second segment bright carmine out-
wardly and above; third segment carmine-tipped above. Face gray,
broadly edged with black. Head pink with a black median spot
posteriorly. Collar pale pink, darker outwardly and edged with black
beneath; on each side a conspicuous black-edged gray spot surrounded
by a narrow, attenuated, cream-colored area. Thorax cerise; mesially
a narrow, longitudinal, ochreous line; on each side a longitudinal,
dorsal, black-edged, gray stripe; tegula pink, edged with cerise and
containing a large, elongate, triangular, black-edged, gray spot. Fore
wing cerise with veins faintly buff; costa narrowly edged with buff;
along costa five conspicuous, irregular, black-edged, gray spots; ex-
tending across wing from these costal spots, five rows of irregular,
black-edged, gray spots and dashes; on costa, at apex, an oval gray
spot narrowly edged inwardly with black; along termen, between
veins 3 and 8, a series of elongate, U-shaped, black-edged, gray
dashes; at tornus a conspicuous, round, black-edged gray spot; cilia
consisting of alternating buff and gray dashes; the underside more
or less suffused with orange-ochreous, the markings less conspicuous
and, except for the inner ones, sooty black; the two basal costal spots
black. Hind wing semihyaline, cerise; costa rather broadly edged
with pale ochreous and with two narrow, poorly defined, fuscous,
transverse dashes about middle; on outer margin, at end of vein 1b,
a small but conspicuous black spot; on the underside, the costa marked
with conspicuous, black-edged, gray dashes. Legs creamy white,
overlaid with cerise and pink and variously marked with black-edged
gray spots; tarsi annulated with black. Abdomen cerise above with
a faint, longitudinal median, black basal dash; beneath pink and buff
mixed. Anal tuft ochreous beneath mixed with black scales; above,
marked with an elongate, median, black, triangular dash.
Male genitalia—Harpe with moderately broad, inward projection,
roughened on posterior edge; distal end of harpe greatly dilated.
Anellus strongly concave laterally. Aedeagus with small distolateral
MOTHS OF THE GENUS ARACHNIS—CLARKE 69
flap. Vinculum broadly rounded. Uncus stout, conical. Flange of
tegumen broadly rounded.
Alar expanse, 54-55 mm.
Type.—U. 8S. N. M. No. 54258.
Type locality —Phantom Ranch, Grand Canyon, Ariz.
Food plant—Unknown.
Remarks.—Described from the type male (12-IX—1938) and one
male paratype (Roaring Springs, Grand Canyon, “VIII-1938”) both
collected and submitted by Louis Schellbach, assistant park naturalist.
This is one of the most brilliantly colored species of the genus and
can be distinguished easily from all others by the concolorous
ground of the fore and hind wings. It appears to be most nearly
related to picta.
ARACHNIS MIDAS Barnes and Lindsey
PLATH 12, FigurRES 8-8c
Arachnis midas BARNES and LInDsEy, Ent. News, vol. 32, p. 297, 1921.—McDun-
novueH, Check list of the Lepidoptera of Canada and the United States of
America (Part 1, Macrolepidoptera), No. 1083, 1988.
Male genitalia.—Lateral projection of harpe not bent toward base,
broader than distal end of harpe beyond it and roughened on posterior
edge; distal end of harpe swollen. Anellus narrower distally than
proximally. Aedeagus with well-developed distolateral flap. Vin-
culum moderately narrow, rounded. Uncus elongate, angular.
Flange of tegumen broad.
Alar expanse, 55 mm.
Distribution —Known only from the type locality.
Type—tIn the U. S. National Museum.
Type locality —KEureka, Utah.
Food plant.—Unknown.
Remarks.—The genitalia of this species are strikingly similar to
those of several of the varieties of picta but are at once distin-
guished by the elongate and angulate uncus, as shown in the figure.
I believe this to be another color form of picta but am retaining the
specific name for the present because it is represented by the unique
type only, which does not offer sufficient evidence for a change. The
distolateral flap of the aedeagus is especially typical of picta.
ARACHNIS ZUNI Neumoegen
PLATE 11, Figures 5-5b, 6-6a
Arachnis zuni NEUMOEGEN, Ent. Amer., vol. 6, p. 173, 1890.—Smiru, List of the
Lepidoptera of Boreal America, No. 1119, 1891.—Kirpy, A synonymic cata-
logue of the Lepidoptera Heterocera (moths), vol. 1, p. 219, 1892.—NruMOEGEN
and Dyar, Journ. New York Ent. Sce., vol. 1, p. 178. ‘i179, 1893.—Drucr, Bi-
70 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
ologia Centrali-Americana, Heterocera, vol. 2, p. 378, pl. 75, figs. 5, 8, 1897.—
Hampson, Catalogue of the Arctiadae (Arctianae) and Agaristidae in the
collection of the British Museum, vol. 3, pp. 389, 393, pl. 47, fig. 15, 1901.—
CocKERELL, Ent. News, vol. 12, p. 209, 1901 [egg].—Dvyar, U. S. Nat. Mus. Bull.
52, No. 858, 1903.—SMiru, Check list of the Lepidoptera of Boreal America,
No. 947, 1903.—Hotianp, The moth book, p. 124, pl. 16, fig. 3, 1903.—BarnEs
and McDunnoueH, Check list of the Lepidoptera of Boreal America, No.
969, 1917.—BonNIWELL, The Lepidopterist, vol. 2, p. 85, 1918.—StTranp, Lepi-
dopterorum catalogus, pt. 22, p. 279, 1919—Sr1rz, Die Gross-Schmetterlinge
der Erde, vol. 6, p. 315, pl. 40c, 1919—BArnrEs and LINpsEy, Ent. News, vol.
32, p. 297, 1921—McDunnoueH, Check list of the Lepidoptera of Canada and
the United States of America (Part 1, Macrolepidoptera), No. 1084, 1988.
Male genitalia—Lateral projection of harpe broader than portion
of harpe beyond it, bent toward base; distal end of harpe not greatly
dilated, somewhat compressed, slightly excurved. Anellus long, nar-
rower distally than proximally. Aedeagus with broad, flattened, dis-
tolateral flap. Vinculum narrow, bluntly pointed, with long, narrow,
lateral, winglike expansion. Uncus conical, elongate with apex nar-
rowly flattened.
Female genitalia——Median fleshy protuberance of ostium flattened,
broad, with shallow indentation on posterior margin; lateral area
membranous. Ductus seminalis weakly sclerotized anterior to its
junction with the ductus bursae and bursa copulatrix.
Alar expanse, 43-70 mm.
Distribution.—Southwestern part of the United States and Mexico.
Arizona: Chiracahua Mountains, 2 6 ¢, 2 92 (June 12 to 26, H. G. Hubbard).
New Mexico: High Rolls, 12 ¢¢, 9 9@ (various dates; no collector) ; Las
Cruces, ¢ (no date; T. D. A. Cockerell) ; Las Vegas, 2 (no date or collector).
Type—tIn the U. 8. National Museum.
Type locality —Las Vegas, N. Mex.
Food plant—Virginia creeper.
Remarks.—This species is easily distinguishable from any other in
the genus by the peculiar slate-colored markings of the fore wing and
the yellow ground color of the hind wing.
A single specimen in the U. S. National Museum from Mexico City,
Mexico, if correctly labeled, suggests that zwni has a much wider dis-
tribution than the above records from the United States indicate.
U. S. GOVERNMENT PRINTING OFFICE: 1941
U. S. NATIONAL MUSEUM
PROCEEDINGS, VOL. 91 PLATE 10
1 pornpeia
3 apachea
1-la. Arachnis aulaea pompeia Druce: 1, Ventral view of female genitalia; 1a, dorsal view of glands.
2-2a. Arachnis aulaea Geyer: 2, Ventral view of female genitalia; 2a, dorsal aspect of glands entering
intersegmental membrane.
3-3¢. Arachnis apachea, new species: 3, Ventral view of male genitalia with aedeagus removed; 3a,
ventral aspect of anellus; 34, lateral view of aedeagus
; 3c, dorsal view of male genitalia
with aedeagus removed and showing flange.
U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 91. PLATE 11
oS \f “6a
6 ZUM J
4-46. Arachnis aulaea Geyer: 4, Ventral view of male genitalia with aedeagus removed showing
flange of tegumen; 42, anellus, ventral view; 44, lateral aspect of aedeagus showing disto-
lateral flap.
S-6a. Arachnis zuni Neumoegen: 5, Ventral view of male genitalia with aedeagus removed; 5a,
ventral view of anellus: 54, aedeagus, lateral view; 6, ventral view of female genitalia; 6c,
dorsal glands.
U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 91 PLATE 12
8IMAAS
~I
Te, 9-9a. Arachnis picta Packard: 7, Ventral aspect of male genitalia with aedeagus removed;
7a, anellus, ventral view; 74, aedeagus, lateral view; 7c, lateral aspect of male genitalia
showing uncus and flange; 9, ventral view of female genitalia; 92, dorsal glands.
8-8c. drachnis midas Barnes and Lindsey: 8, Ventral view of male genitalia with aedeagus
removed; 8a, ventral view of anellus; 84, aedeagus, lateral aspect; 8c, lateral view of
uncus.
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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM
issued
SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
Vol. 91 Washington : 1941 No. 3124
SOME LITTLE-KNOWN FOSSIL LIZARDS FROM THE
OLIGOCENE OF WYOMING
By Cuaries W. Grtmore
Amone a small collection of Oligocene fossil remains acquired for
the United States National Museum in 1931, from George F. Stern-
berg, were two lizard specimens that contribute to a better understand-
ing of the cranial anatomy of the genera Aciprion and Eaostinus.
These specimens were found in a small badland area of the Brule
formation that is bisected by U. S. Highway 20, about 8 miles east
of Douglas, Converse County, Wyo. A detailed description of them
follows. The illustrations were prepared by Sydney Prentice.
Family IGUANIDAE
Genus ACIPRION Cope
ACIPRION FORMOSUM Cope
Fiacures 30, 31
An almost complete skull with both dentaries (U.S.N.M. No. 16566)
of Aciprion formosum Cope gives for the first time a comprehensive
knowledge of the cranium in this little-known genus and species.
Skull.—The skull is complete except for part of the right jugal
and fragments of the squamosal of the same side. The anterior
half of the palate has been disarranged and some of the elements
aremissing. ‘The lower jaws both lack their posterior portions.
Most of the sutural contacts are discernible and so make it possible
clearly to depict the cranial details as shown in the illustrations. In
406806—41 71
72 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
size and general structure the fossil skull displays many resemblances
to the living lizard Crotaphytus. The dentitions of these two forms
likewise are very similar.
Viewed from the side (see fig. 31) the profile of the skull at the
junction of the parietal and frontal is depressed, as contrasted with
the usual convex profile of most of the Iguanidae. From the tip of
the nose to the posterior end of the squamosal the skull has a greatest
length of 27 mm.; the greatest breadth across the jugals is 14.6 mm.
The premaxillary has a long spine that is relatively wider than in
Crotaphytus. Its posterior end is notably different in being broadly
rounded as contrasted with the narrow, sharply pointed extremity
in the extant genus. The nasals are short and wide, being shortened
Ficure 30.—Skull of Aciprion formosum Cope (U.S.N.M. No. 16566), superior view: f,
Frontal; ju, jugal; /a, lachrymal; mx, maxillary; na, nasal; p, parietal; pmx, premaxil-
lary; pof, postorbital; prf, prefrontal; soc, supraocciptal; sg, squamosal. About three
times natural size.
by the large size and partly vertical position of the nostril openings.
The frontal is single and relatively wide between the orbits. The
pineal foramen is on the frontoparietal suture. The prefrontal is
large, but without a preocular boss, which forms such a prominent
projection on the Crotaphytus skull. The postifrontal is absent, a
condition noted by Cope? in Crotaphytus. Its place is taken by a
widening of the frontal on each posterior-external angle. The
postorbital is large, uniting inferiorly with the jugal and posteriorly
with the squamosal. The dorsal surface of the parietal is relatively
narrower between the supratemporal fossa and between the divergent
posterior process than in Crotaphytus. 'The left squamosal is miss-
ing, and only a small part of the right one is present. In the ilustra-
1 Cope, E. D., Ann. Rep. U. 8S. Nat. Mus. for 1898, p. 246, 1900.
FOSSIL LIZARDS FROM WYOMING—GILMORE 73
tions it has been restored following modern iguanids. The lachrymal
is very small and in line with the jugal. The large jugal is without
a posteriorly directed spur. Only the right quadrate is present, and
it is so damaged that its detailed structure is obscured. As depicted
in figure 31 it may be too short. It appears to have a nearly straight
external border. The top of the supraoccipital is not wholly beneath
the overlying parietal but is visible from above as shown in figure
30. A low obtuse vertical ridge extends upward from the top of the
foramen magnum. The supraoccipital is fully coalesced with the
exoccipital. The occipital condyle is plain and without evidence of
participation of the exoccipitals.
The basioccipital and sphenoid surfaces are confluent. Basiptery-
goid processes are large, with spatulate ends directed strongly for-
Figure 31.—Skull and lower jaw of Aciprion formosum Cope (U. S. N. M. No. 16566),
viewed from the left side: co, Coronoid; d, dentary; f, frontal; ju, jugal; gu, quadrate;
la, lachrymal; mx, maxillary; na, nasal; p, parietal; pmx, premaxillary; poc, paraoccipital;
pof, postorbital; prf, prefrontal; sg, squamosal. About three times natural size.
ward. There is no evidence of teeth on the pterygoids. The other
palatal elements are so badly disarranged as to furnish no reliable
information regarding the true structure of the palate.
Lower jaw.—The mandible in specimen U.S.N.M. No. 16566 is rep-
resented by the right dentary, with full dentition posterior of the
coronoid process and the greater portion of the left dentary lacking
most of the teeth. These contribute but little new information, and
since the lower jaw has been described in a previous publication there
is no reason to repeat it here. The dentary carries 25 closely set teeth
in the complete series. In the restoration of the missing part of the
ramus in figure 31, the very complete ramus forming part of the type
of Aciprion majus was used as a guide.
Dentition.—The dentition is pleurodont, the dental formula being
premaxillary 6, maxillary 20, dentary 25. The teeth are closely
placed, cylindric with compressed crowns. The latter support a large
74 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
median and two small lateral cusps. These lateral cusps are most
prominently developed on the teeth of the posterior two-thirds of
both upper and lower series. From this point forward the teeth
gradually diminish in size, and the lateral cusps become smaller and
smaller, disappearing altogether on the first few teeth that have simple
pointed crowns. Upper and lower teeth appear indistinguishable.
Crowns in lower jaw project farther above the alveolar border than
in the maxillary.
Specimen U.S.N.M. No. 16566 in total number of teeth in maxillary
and dentary is in perfect accord with the type of Aciprion majus
Gilmore, but its smaller size clearly shows it to pertain to the earlier
described Aciprion formosum Cope.
femarks.—In 1928° this genus was referred to the family
Iguanidae on rather meager evidence, but after a study of these new
materials the propriety of that assignment now seems assured. The
resemblances found in skull structure and character of dentition to
those of extant members of the family leave little doubt as to the
correctness of this family assignment.
Measurements of Skull, U. S. N. M. No. 16566
mm.
Greatest lenzgthvof sskullE Overs ihe ook eee en ee ee 27.0
Greatestalene tig ofa skeleton cl cll emer ame ne re 22.3
Greatest width Oe SUL GA CrOSS cir LS eee ree eee meg 14.6
Greatest width: parietals at: centers] 22 = sen ee ee ee ee 3.5
Greatest ensthitrontals petween vorbis. eee a eee eee ee 2.3
Grealestelenn gta se eee eae Oo a ea nea oe cre LCL ee ae ee Ea 3.6
Greatest length tron Gales aes se i SENSE ale, EL Po a EEL EEOC ee le 7.0
Greatest) demath parietal eee Be Leh i Te API i UN ee A Wal
Greatest; width voceipitall condyle ns sees ee ae ean eee Eh aes BSL ge 1.2
Genus EXOSTINUS Cope
EXOSTINUS SERRATUS Cope
FIaguRE 32
An anterior portion of a skull and a left dentary (U.S.N.M. No.
16565) is clearly identified as pertaining to H'wvostinus serratus Cope.
It is the first specimen found that displays the complete structure and
osseous scutellation of this part of the cranium, and thus it contributes
to a better understanding of this little-known species.
The entire outer surfaces of the premaxillary, nasal, and maxil-
lary bones, with the exception of a smooth narrow band parallel to
the dentigerous border, is covered by the characteristic osseous promi-
nences, as shown in figure 32. These are coalesced to the underlying
skull elements and thus hide all trace of the cranial sutures. For that
2 Gilmore, C. W., Mem. Nat. Acad. Sci., vol. 22, p. 18, 1928.
FOSSIL LIZARDS FROM WYOMING—GILMORE 19
reason the extent of the underlying skull bones cannot be accurately
determined. The maxillary of the left side is complete and from end
to end has a length of 8.5 mm. The complete dental series of the
maxillary consists of 12 pleurodont, subcylindric teeth. The
premaxillary has eight teeth in the complete series, as in Peltosaurus.
The spine of the premaxillary is ornamented with three longitudi-
nal rows of osseous tubercles, the central row having the largest ossifi-
cations. The nasal region is covered with tubercles of varying sizes
and without definite arrangement. Those above the prefrontal are
the largest tubercles on this portion of the skull and form a distinct
row along the orbital border. Although the frontals are missing in
this specimen, it is quite evident that the prefrontal strongly laps
this bone and that its posterior termination reaches nearly to the cen-
ter of the orbit.
Ficure 32.—Anterior part of the skull of Exostinus serratus Cope (U.S.N.M .No. 16565),
viewed from left side: d, Dentary; f, prefrontal; mx, maxillary; na, nasal; pmx, pre-
maxillary. About three and one-half times natural size.
The type * on which this genus and species is based consists of the
frontals, left zygomatic, and a portion of the dentary with a few
teeth. The frontals are also covered with bony tubercles, a series
along each supraorbital border, longitudinal at the front, quadrate
at the back. A single median row separates them. On the posterior
end of the frontals, they are arranged in three transverse rows of 5,
4, and 3 tubercles, respectively. On the zygomatic there are two lon-
gitudinal rows of flat quadrangular tubercles.
The incomplete dentary carries 14 teeth, and it appears that two or
more may be missing from the posterior end of the series. In the
article cited I stated that “the upper teeth [are] similar to the lower”;
this is true only so far as both are pleurodont, with subcylindric
shafts and simple crowns. The lower are more robust than the up-
per and their crowns project farther beyond the parapet of the jaw,
as Clearly shown in figure 32. In this specimen there are nine teeth
§ Gilmore, C. W., Mem. Nat. Acad. Sci., vol. 22, p. 22, pl. 25, figs. 4-6, 1928.
76 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
in 5 mm., whereas in the type dentary eight teeth occupy a similar
space. The teeth of both upper and lower series decrease in size
toward the front, and the transversely compressed crowns of the
lateral teeth change to simple, rounded, sharp-pointed teeth in front.
The dental formula of Hwostinus serratus may now be stated as
follows:
Maxillary 14++premaxillary 8__ 36_
dentary 14* 28"
This genus and species were tentatively referred in my 1928 review
of the lizards of North America to the family Iguanidae. Although
this new material contributes but scant information on this important
question, the subequal size of the pleurodont teeth, the constantly
long cylindrical shafts, and the gradual change taking place between
the lateral and anterior teeth are all features in accord with its as-
signment to the Iguanidae. The osseous ornamentation of the skull
is highly suggestive of the horny tubercular ornamentation of the
Phrynosoma skull. For the present, therefore, Hxostinus will be
regarded as an extinct representative of the [guanidae.
U.S. GOVERNMENT PRINTING OFFICE: 1948
PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM
issued
SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
Vol. 91 Washington: 1941 No. 3125
NEW SPECIES OF HYDROIDS, MOSTLY FROM THE AT-
LANTIC OCEAN, IN THE UNITED STATES NATIONAL
MUSEUM
By C. McLean FRrAsrr
A paper that might be called a progress report, including the de-
scription of new species from the first portion of a large United States
National Museum collection of hydroids, mostly from the North At-
lantic, was published in 1940.1. The examination of the remainder of
this collection has been completed, and the present paper serves to
report further on the new species in the collection. The whole of the
material has yielded more than 1,200 distribution records for 173
species.
Although most of the material was obtained from the North Atlantic,
it happens that out of the 15 species here considered only 10 were
obtained in the Atlantic. The other five came from the west coast of
America, from Bering Sea to Panama. Two of the most interesting
species in the collection were together in the same vial from Thistle
Ledge, Stephens Pass, not far from Juneau, Alaska. For one of these
species it appears to be necessary to introduce not only a new genus
but also a new family (see p. 78). The other species, Lampra uvularis,
belongs to a genus not previously reported from the Pacific coast of
North America. One from Bering Sea, one from near the Golden
Gate, Calif., and one from near Panama make up the other three
species.
+ Fraser, C. McLean, Seven new species and one new genus of hydroids, mostly from the
Atlantic Ocean. Proc. U. S. Nat. Mus., vol. 88, pp. 575-580, 1940.
406740411 i
78 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
Of the 15 species considered 14 are described as new, and for the
fifteenth the gonosome is described and figured for the first time. As
indicated, one new genus and one new family are described.
The whole collection, therefore, has provided one new family, two
new genera, 21 new species, and the gonosome of two species, of which
the trophosome had been previously described.
I must again express my appreciation of the courtesy shown by the
United States National Museum in providing the opportunity to ex-
emine this material, and my appreciation of the contribution that
Miss Ursula Dale has made in drawing the figures used in illustration.
SYMPLECTANEIDAE, new family
Trophosome.—Zooids without chitinous perisarc, with capitate ten-
tacles, arranged in series over the surface of the body of the hydranth,
each series of three or more fused throughout much of their length
to form a bractlike structure.
Gonosome.—Gonophores producing sporosacs borne on the body
of the hydranth,
SYMPLECTANEA, new genus
Trophosome.—Zooids solitary, without chitinous perisarc; the capi-
tate tentacles in series, graded in length, the longest tentacle medially
placed in the series and the others growing shorter as they appear
farther from the median.
Gonosome.—Gonophores in the form of sporosacs in the axil of a
series of tentacles.
SYMPLECTANEA BRACTEATA, new species
PLATE 13, FicuRE 1
Trophosome.—Solitary zooids grow from a broad base, with stubby
processes projecting from the central portion; largest specimens 33
mm. in length; hydrocaulus 1.6 mm. in diameter, hydranth 2.0 to 4.0
mm., the hydranth making up one-third of the length. No chitinous
covering in any part and no annulations. The hydranth is provided
with numerous tentacles in series, scattered over the whole surface;
the series consists of 3, 5, or 7 tentacles in a row, fused into one bractlike
structure; the median tentacle may be 1 mm. long, the next two, one
on each side, much the same in length, which is less than that of the
median; there is a further recession for the next pair, and the next,
if these are all present. Fusion appears for the greater part of the
length of the lesser tentacle of each pair in succession, always leaving
the capitate portion free. In the younger hydranth the bract makes a
NEW SPECIES OF HYDROIDS—FRASER 79
sharp angle with the body, but when the gonophore develops the
bract is gradually forced outward distally until it is nearly at right
angles to the body.
Gonosome.—The gonophores develop to form sporosacs in the angle
between the tentacular bract and the body of the hydranth; they are
almost spherical, with very short pedicels; ova relatively large and
not numerous.
Type.—vU.S.N.M. No. 43450. Taken by the United States Fisheries
steamer Albatross at station 4253, Thistle Ledge, Stephens Pass,
Alaska, 131 fathoms, July 14, 1903.
Family HYDRACTINIDAE
Genus HYDRACTINIA van Beneden
HYDRACTINIA VALENS, new species
PLATE 138, FIcuRE 2
Trophosome.—Colony growing from a thick, basal coenosare, pro-
vided with short, smooth spines; nutritive zooids large and lusty,
reaching a height of 4.5 mm.; 10 tentacles in rather regular whorls.
Gonosome.—Generative zooids (only female zooids obtained) about
one-half of the length and breadth of the mature nutritive zooids;
tentacles wholly lacking; sporosacs 3-5, forming a whorl at the base
of the proboscis; commonly 6 ova in each sporosac.
Other zooids.——None observed.
Type.—u.S.N.M. No. 43451. Taken by the United States Fisheries
steamer Speedwell at station 284, latitude 42°10’ N., longitude 70°22’
W., southwest of Stellwagens Bank, near Race Point Light, Cape Cod
region, 31 fathoms, August 4, 1879.
Family CORYMORPHIDAE
Genus CORYMORPHA Sars (in part)
CORYMORPHA ADVENTITIA, new species
PLATE 13, Figure 3
Trophosome.—Zooids 20 mm., of which the hydranth is approxi-
mately one-fourth, with adventitious shoots, the longest 0.25 mm..,
passing backward from the main hydrocaulus at various angles, to
serve as accessory means of attachment; the hydrocaulus has much
the same diameter throughout, or this may increase slightly, distally ;
proximal tentacles 20-24 in one whorl, distal tentacles very numerous
in several irregular whorls.
80 PROCEEDINGS OF THE NATIONAL MUSEUM VOU. 91
Gonosome.—Gonophores borne on long, unbranched peduncles, at-
tached to the hydranth just distal to the proximal tentacles, each
gonophore with a short pedicel; apparently these gonophores develop
irregularly, as small and large ones are mixed without any evidence
of their appearing in any regular order.
Type.—U.S.N.M. No. 48452. The vial is labeled “U.S. F. C. Str.
Albatross, Panama, Mar. 12, 1891,” but there is no station listed on that
day. The last haul on March 11 was made in latitude 7°33’ N.,
longitude 78°34’20’” W., in 85 fathoms.
Remarks.—The adventitious shoots in these hydroids are so unusual
that it might seem advisable to place the species in a new genus, but,
although each of the three specimens available for examination had
these shoots, it is just possible that they may have developed under
unusual conditions, and as all the other features are definitely like
Corymorpha, it seems better at the present time to place it in this
genus.
Family TUBULARIDAE
Genus LAMPRA Bonnevie
LAMPRA UVULARIS, new species
PLATE 14, Figure 4
Trophosome.—Zooid 22 mm., of which the hydrocaulus is 15 mm.,
straight, without annulations; hydranths large, 7 mm. in diameter;
proximal tentacles 18-20, long and slender; distal tentacles 40-48,
shorter and stiffer in appearance, in four rather indistinctly different
whorls.
Gonosome.—Gonophores growing in eight erect, closely arranged
clusters, looking like compact bunches of grapes or like the cluster
of flowers in the grape hyacinth; each gonophore is spherical, on a
short pedicel, and shows no sign of tentacular processes.
Type —U.S.N.M. No. 43458. Taken by the United States Fisheries
steamer Albatross at station 4253, Thistle Ledge, Stephens Pass,
Alaska, 1381 fathoms, July 14, 1903.
Remarks.—This appears to be the first record of a species of this
genus from the northeastern Pacific. This is not the place to discuss
the systematic position of Lampra, but it may be stated that it cannot
be placed in the Tubularidae (as Bonnevie has placed it*) as this
family has been defined in all my previous papers.
1Bonnevie, Kristine, Zur Systematik der Hydroiden. Zeitschr. Wiss. Zool., vol. 63,
p. 477, 1898.
NEW SPECIES OF HYDROIDS—FRASER Sl
Genus TUBULARIA Linnaeus (in part)
TUBULARIA CRASSA, new species
PLATE 14, Fieurn 5
Trophosome.—tindividual zooids only were obtained: there is
nothing to indicate whether they grow in colonies or not; the pedicels
appear to be complete, but they are but little more than 1 cm. in
length, which, even in the contracted condition, has a diameter almost
equal to the length of the pedicel. There are no annulations, but
there is a definite ridge at the base of the proximal tentacles; proximal
tentacles long and numerous, 32-36; distal tentacles slender, much more
numerous.
Gonosome.—Gonophores grow in rather long, erect racemes when
well developed; these racemes are densely crowded so that the body
of the hydranth is almost entirely hidden; there are no tentacular
processes on the gonophores.
Type.—vU.S.N.M. No. 22746. Taken by the United States Fisheries
steamer Fish Hawk at station 988, latitude 40°49’30’’ N., longitude
70°47’ W., off Marthas Vineyard, 30 fathoms, September 7, 1881.
Family CAMPANULARIDAE
Genus CAMPANULARIA Lamarck
? CAMPANULARIA FASCICULATA, new species
PLATE 15, FIGURE 6.
Trophosome.—Colony 2 cm. in height, with the base of the main
stem and some of the lower branches fascicled. The simple branches
are short; the hydrothecae arising from the fascicled stem have rela-
tively long pedicels, annulated at each end; those from the simple
portion of the stem and from the branches with shorter pedicels, com-
monly annulated throughout. Hydrothecae large, 0.5-0.6 mm. in
length, broadly campanulate; margin with 16 low, rounded teeth;
lines run down the wall of the hydrotheca from the depressions be-
tween the teeth.
Gonosome.—Not observed.
Type.—uU.S.N.M. No. 43454. Taken by the United States Fisheries
steamer Speedwell at station 984, latitude 41°31’ N., longitude 69°28’
W. off Chatham, Cape Cod, 33 fathoms, August 30, 1881.
82 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
Genus OBELIA Peron and Lesueur
? OBELIA RACEMOSA, new species
PLATE 15, FIGURE 7.
Trophosome.—Colony large, with a main axis 25 cm. and a few
large branches almost as large as the main axis; from these small
branches and branchlets are given off that distally are clustered in
rather stiff racemes. The main stem and larger branches are strongly
fascicled and even the secondary branches may be so in the proximal
portion; the primary branches and the larger secondary branches are
annulated only above the nodes, but the distal branchlets and the
pedicels are extensively annulated; the longer ones are annulated
proximally and distally, with a short, smooth portion between, of
greater diameter, so that the branchlet or pedicel seems to bulge defi-
nitely in this portion; the shorter pedicels are annulated throughout.
The hydrothecae, appearing in close clusters, are broadly campanulate,
at least as broad as deep; margin entire. The larger branches and
the main stem are dark brown, the branchlets and pedicels much
lighter.
Gonosome.—Not observed.
Type—U.S.N.M. No. 4883. Western Bank, off Cape Breton Island,
50-65 fathoms, June 7, 1880.
Remarks.—This species bears some resemblance to Obelia plicata
Hincks, but it is a larger, coarser species, the ultimate branches are
more rigid, the hydrothecae are clustered, and the hydrotheca is more
broadly campanulate.
Family CAMPANULINIDAE
Genus EGMUNDELLA Stechow
EGMUNDELLA GRANDIS, new species
PLATE 16, FIGURE 8.
Trophosome.—Zooids growing singly from an irregularly reticulate
stolon to a height of 8 mm.; pedicel straight, rigid, smooth except for
two or three annulations at each end; hydrotheca of the usual turbinate
type, 0.7—-0.8 mm. in height; operculum of 12 segments. Nematophores
very small for this genus, spherical, with a short pedicel, sparingly
scattered over the stolon, and occasionally occurring on the pedicels.
Gonosome.—Not observed.
Type —U.S.N.M. No. 48455. Taken by the United States Fisheries
steamer Fish Hawk at station 897, latitude 37°25’ N., longitude 74°18’
W., off the mouth of Chesapeake Bay, 15714 fathoms, November 16,
1880.
NEW SPECIES OF HYDROIDS—FRASER 83
Genus LOVENELLA Allman
LOVENELLA GRANDIS Nutting
PLATE 16, Figure 9.
Lovenella grandis Nutrina, U. S. Fish Comm. Bull. for 1899, pp. 325-386, figs.
1-105, 1901.
Trophosome.—Stems simple, rather rigid, unbranched, up to 5 cm.
in length, divided into regular, long internodes by single nodes. Hy-
drothecae arise on short pedicels, with a double annulation from a
process a short distance from the distal end of the internode, regularly
alternate; hydrothecae very large, turbinate; margin with 10-12 sinu-
ations from which arise the segments of the operculum.
Gonosome.—(Not previously described.) Gonangium long, 1.5-1.6
mm., but rather slender, arises from the axil of the pedicel, the basal
portion gradually increasing in diameter, but the distal half prac-
tically tubular; pedicel short, with one annulation. Medusa buds were
developing on the blastostyle, but they were not far enough advanced
to show all the characteristics.
Type.—uvU.S.N.M. No. 48460. Taken by the United States Fisheries
steamer Fish Hawk at station 830, near the mouth of the Sakonnet
River, R. I, 1014 fathoms, August 27, 1880.
Remarks.—Nutting described this species from a specimen dredged
from Newport Harbor, off Castle Hill, a location very near the present
one. As far as I am aware, it has not been reported since until now.
Nutting’s specimen had no gonosome.
Family HALECIDAE
Genus HALECIUM Oken
HALECIUM DUBIUM, new species
PLATE 16, FIGURE 10a; PLATE 17, FIGURE 100
Trophosome.—Colony slightly bushy, reaching a height of 3 cm.;
proximal portion fascicled to a limited extent. Nodes not very
strongly marked; internodes long, turning alternately to one side and
to the other, making a zigzag main stem. The hydrophore, with rela-
tively long pedicel, is given off near the distal end of the internode;
this pedicel makes much the same angle with the vertical as the inter-
node of the stem does. The hydrophore may give rise to one or more
other hydrophores as duplications, the pedicels of these varying much
in length; the margin of the hydrophore is slightly flaring. The
branches arise in the same way as the hydrophores, so it would appear
at first glance that the branching is dichotomous, but the branch is
not like the main stem; the proximal portion is like a hydrophore with
84. PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
an elongated pedicel and it may be duplicated in series; then from
the distal end, or near it, of the main pedicel, an internode is given
off that looks like an internode of the main stem, and from this the
branch continues in the same way that the stem does.
Gonosome.—Male gonangia arise from the base of the hydrophore
pedicels, just beyond where they leave the internodes; they are broadly
obovate in the one direction and almost flat in the other; there is a
short but distinct pedicel present; at the distal end the gonangium has
a small, but distinct, semicircular notch.
Type.—v.S.N.M. No. 22922. Taken by the United States Fisheries
steamer Albatross at station 2572, latitude 40°29’ N., longitude 66°04’
W., off Cape Sable, 1,769 fathoms, September 2, 1885.
Remarks.—Ilt is with some misgivings that I describe this as a new
species, since there is so much resemblance to H. telescopicwm Allman,
as described and figured by Allman? and by Jiiderholm,’ and yet the
specimen from which this species is described has not the characteristic
that these authors, and Pictet and Bedot* as well, consider definitely
distinctive, i. e., the number of the reduplications of the hydrophore,
to form a series with many more units than are exhibited in any other
species. One might surmise that this excessive reduplication was due
to some seasonal or environmental condition, were it not that the same
type of structure appeared in such distant locations. The distribution
itself is indeed remarkable. Allman described it originally from off
Port Jackson, NSW., in 30-85 fathoms. Then Pictet and Bedot re-
ported it from the Gulf of Gascogny in 155-180 meters, and later
Jiderholm reported it from the Bering Sea in 131 meters.
Apart from the matter of reduplication, the only other character
that is noticeably different is the gonangium, or rather the semicircular
notch at the distal end of this, and this is quite a minor difference.
The female has not been reported in any instance.
HALECIUM TENSUM, new species
PLATE 17, Fiaure 11
Trophosome.—Colony rather rigid, with a main axis (5 cm.) and a
few irregularly arranged branches, the proximal being almost as long
as the main axis and the others becoming shorter as they get farther
from the base; proximal portion of the main stem and of some of the
branches, fascicled; there is little indication of nodes on stems or
branches. Each portion of a stem or branch that corresponds to an
2 Allman, G. J., Report on the Hydroida. Challenger Expedition, vol. 23, pt. 70, p. 10,
1888.
3 Jiderholm, E., Der Hydroidenfauna des Beeringsmeeres. Archiv fdr Zool., vol. 4, No. 8,
p. 4, 1907.
*Pictet, C., and Bedot, M., Hydraires provenant des Campagnes de L’Hirondelle (1886—
1888), p. 7, 1900.
NEW SPECIES OF HYDROIDS—FRASER 85
internode in the regular type is much elongated, tubular, and slightly
curved outward distally to end in a hydrophore; then from this pedicel
of the hydrophore, a short distance from the distal end, the pedicel
for another hydrophore is given off. These in succession form a series,
alternately curving to one side and the other and thus maintaining a
linear stem or branch. From within each main hydrophore there is
usually another hydrophore developed with a much shorter and some-
what slenderer pedicel. In some cases this hydrophore is duplicated.
The rim of the hydrophore flares but slightly.
Gonosome.—Not observed.
Type—vU.S.N.M. No. 22926. Taken by the United States Fisheries
steamer Fish Hawk at station 940, latitude 39°54’ N., longitude
69°51’30’’ W., off Marthas Vineyard, 134 fathoms, August 4, 1881.
Remarks.—This Halecium has somewhat the same general appear-
ance as H. kiikenthali Marktanner-Turneretscher, but as a colony it
is more rigid and less branched; the internodes, or rather hydrophore
pedicels, are relatively much longer, and, most noticeably, they lack
the annulations that are so conspicuous in H, kiikenthali.
Family LAFOEIDAE
Genus LICTORELLA Allman
LICTORELLA CRASSITHECA, new species
PLATE 18, Figure 12
Trophosome.—Main stem and the proximal portions of some of the
branches fascicled, branching inclined to be pinnate but irregular;
occasionally secondary branches appear. There are no noticeable nodes
in the ultimate branches, but the hydrothecae are given off in regular
alternation. There is a distinct shoulder at the origin of each hy-
drotheca on which the pedicel of the hydrotheca seems to be somewhat
displaced upward or outward; the pedicel is distinct, with one distinct
annulation. The hydrotheca widens quickly at the base and the re-
mainder is nearly cylindrical, except that it shows a slight campanulate
tendency near its margin, which is entire. The width is much greater
relative to the length than in other species. The diaphragm is distinct
but does not reach in far from the wall of the hydrotheca. The nema-
tocysts are scarce; none was observed on the branches and few on the
fascicled stem.
Gonosome.—Not observed.
Type—vU.S.N.M. No. 43456, Gulf of Maine, 17 fathoms, Also taken
at Albatross station 2430, latitude 42°58’30’’ N., longitude 50°50’ W.,
southeast of Sable Island, 179 fathoms, June 23, 1885.
406740—41 2
86 PROCEEDINGS OF THE NATIONAL MUSEUM YOU, 91
Family PLUMULARIDAE
Genus AGLAOPHENIA Lamoroux (modified)
AGLAOPHENIA INCONSTANS, new species
Prats 18, Ficurr 13
* Trophosome.—Colonies varying in appearance; one, 17.5 cm. long,
has no branches, and all the hydrocladia have disappeared from the
stem except for about 2.5 cm. at the distal end, while at the other
extreme a distal fragment of the main stem, 6 cm. long, has six branches,
each replacing a hydrocladium and each regularly bearing hydro-
cladia; the longest branch is 2.0 em, Stems, with the exception of
the proximal portion, and branches are divided into regular, rather
short internodes by definite nodes, each internode bearing a hydro-
cladial process near the distal end; these processes alternate from side
to side but are not nearly in the same plane; two in succession may
form an angle as low as 60°. Hydrocladia short for the size of the
colony, as short as in some of the minute species of this genus, divided
into regular internodes by definite nodes; each hydrotheca occupies
almost all the internode, so that there is little space between two
hydrothecae in succession; distinctly deeper than broad; margin with
nine irregular and irregularly placed teeth ; the median tooth is slender,
sharp-pointed, and strongly retrorse; each of the first lateral pair is
also slender and acute but points outward; between the first and the
second there is a wide and deep sinus; the second is lower and blunter
than the first; the sinus between the second and third is shallower, and
the third tooth is blunter than the second; the next sinus is even less
marked, for the fourth lateral tooth is rather insignificant in size and
in some cases can scarcely be observed. The intrathecal ridge is
prominent, and there is a second one indicated at the base of the
supracalycine nematophore.
The supracalycine nematophores are large, slightly overtopping the
hydrothecal margin; the mesial nematophore is short, not reaching to
the margin of the hydrotheca and not projecting outward very notice-
ably. ‘There are three nematophores on each internode of the stem
or branch; one on the hydrocladial process, one at the base of this
process, and one in the axil, this being larger than either of the others.
Gonosome.—Not observed.
Type—vU.S.N.M. No. 48457. Taken by the United States Fisheries
steamer Albatross at station 3497, latitude 56°18’ N., longitude 169°38’
W., Bering Sea, 86 fathoms, July 17, 1893.
NEW SPECIES OF HYDROIDS—FRASER 87
AGLAOPHENIA TRANSITIONIS, new species
PLATE 18, FicuRE 14
Tropohosome.—Colony with a long, somewhat rigid main axis, 8 cm.,
a limited number of branches given off from the distal half of the
stem; each branch leaves the stem in the same manner as a hydro-
cladium, but after it has given rise to seven or eight hydrothecae it
definitely becomes a branch and gives off hydrocladia similar to those
from the main stem. The hydrocladia are relatively short (maximum
4 mm.) and arise alternately from the face of the stem, so that the
supporting processes form a zigzag row, but slightly curved; divided
into regular short internodes by distinct nodes, so that the hydrothecae
are closely placed; the hydrotheca is little longer than broad and is
stouter distally than proximally, adnate throughout almost the whole
length; margin with 11 teeth; the median tooth is erect or very slightly
retrorse, sharp, smaller than the tooth on each side; the tooth next
to the median on each side is the longest, the second one is the smallest,
and the third, fourth, and fifth are nearly equal; all of them are rather
sharply pointed. There is no definite anterior intrathecal ridge; the
posterior is strongly marked but does not reach far.
The supracalycine nematophores, which do not nearly reach the
margin of the hydrotheca, are strongly curved, so that the opening
points backward: mesial nematophore not prominent, projecting from
the hydrotheca in the distal third of its anterior surface. Of the three
cauline nematophores that on the hydrocladial process and the one
below the insertion of this process are tubular; the one near the axil,
1. e., distal to the process, is triangular and larger than either of the
others.
Gonosome.—Not observed.
Type.—U.S.N.M. No. 43458. Taken by the United States Fisheries
steamer Albatross at station 3150, latitude 37°47’ N., longitude
122°44’10’’ W.., off Golden Gate, Calif., 21 fathoms.
Genus PLUMULARIA Lamarck (in part)
PLUMULARIA POLYNEMA, new species
PLATE 18, Figure 15
Trophosome.—Stem simple, slender (from fragment 83 mm. long),
divided into regular internodes with well-marked nodes, each bearing
a single hydrocladium on a prominent process near the distal end.
All the internodes in the hydrocladium are long, slender, and thecate,
except that in some instances an extra nonthecate internode appears,
making an intermediate internode, with two nematophores, and a
thecate internode that is much shorter than the others, with but one
88 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
prominent median nematophore. The hydrotheca, placed a considera-
ble distance from the distal end but still in the distal half, is nearly
equal in depth and breadth. In some instances, a secondary branch
or hydrocladium is given off in place of the hydrotheca in an internode
of the primary hydrocladium. There are no definite septal ridges in
stem or hydrocladia.
There are two supracalycine nematophores, two mesial nematophores
on the proximal hydrocladial internode and three on each of the
others, two at the axil of the hydrocladium on the cauline internodal
process, and three (sometimes only two observed) on each of the cauline
internodes.
Gonosome.—Not observed.
Type.—vU.S.N.M. No. 43459. Taken by the United States Fisheries
steamer Fish Hawk at station 1092, latitude 39°58’ N., longitude 69°42’
W., off Marthas Vineyard, 202 fathoms, August 11, 1882. Another
lot taken at Fish Hawk station 1088, latitude 39°58’ N., longitude 70°06’
W.., off Marthas Vineyard, 130 fathoms, September 21, 1881.
10.
10.
11.
12.
13.
14,
15.
EXPLANATION OF PLATES
(Unless otherwise specified the magnification is x 20.)
PLATE 13
. Symplectanea bracteata, new genus and species: a, Hydranth, showing ar-
rangement of tentacular bracts and gonophores (X 12); 6b, tentacular
bract and gonophore.
. Hydractinia valens, new species: a, b, Nutritive zooids; c, d, female generative
zooids; e, spines.
. Corymorpha adventitia, new species: a, Zooid, showing adventitious shoots
(xX 8); 0, hydranth, showing tentacle and gonophore arrangement (x 12).
PLATE 14
. Lampra wuvularis, new species: Zooid, showing tentacle and gonophore
arrangement.
. Tubularia crassa, new species: a, Individual zooid (xX 6); 6, a gonophore
cluster.
PLATH 15
. ?Campanularia fasciculata, new species: a, Portion of fascicled stem with
hydrothecae; b, portion of simple stem.
. ?0belia racemosa, new species: Portion of colony showing hydrotheca arrange-
ment.
PLATE 16
. Homundella grandis, new species: a, b, Hydrothecae and nematophores.
. Lovenella grandis Nutting: a, Portion of colony with hydrothecae and
gonangia; 6, a single gonophore.
Halecium dubium, new species: a, Portion of colony showing hydrophore
arrangement,
PLATE 17
Halecium dubium, new species: b, Portion of colony showing gonophore
arrangement.
Halecium tensum, new species: a, Portion of fascicled stem; 6, c, portions
of simple stem.
PLATE 18
Lictorella crassitheca, new species: a, Portion of fascicled stem; b, portion of
simple stem.
Aglaophenia inconstans, new species: a, Portion of hydrocladium showing
hydrothecae ; 0, three hydrothecae (Xx 40).
Aglaophenia transitionis, new species: a, Portion of hydrocladium showing
hydrothecae; 6, three hydrothecae (x 40).
Plumularia polynema, new spcies: a, Portion of colony showing nematophore
arrangement; 6b, portion of colony showing branched hydrocladium.
89
tSrarned Heo
Ca. Oot 9!
Ne fe
0. A ER
Saad
5 REE
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PROCEEDINGS, VOL. 91 PLATE 13
U.S. NATIONAL MUSEUM
NEW HYDROIDS
FOR EXPLANATION OF PLATE SEE PAGE 89
PROCEEDINGS, VOL. 91 PLATE 14
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NEW HYDROIDS
FOR EXPLANATION OF PLATE SEE PAGE 89
PROCEEDINGS, VOL. 91 PLATE 15
U.S. NATIONAL MUSEUM
wy
aN
NEW HYDROIDS
FOR EXPLANATION OF PLATE SEE PAGE 89
PROCEEDINGS, VOL. 91 PLATE 16
U. S. NATIONAL MUSEUM
NEW HYDROIDS
FOR EXPLANATION OF PLATE SEE PAGE 89
PROCEEDINGS, VOL. 81 PLATE 17
U.S. NATIONAL MUSEUM
NEW HYDROIDS
FOR EXPLANATION OF PLATE SEE PAGE 89
U.S. NATIONAL MUSEUM PROCEEDINGS, VOL. 91 PLATE 18
NEW HYDROIDS
FOR EXPLANATION OF PLATE SEE PAGE 89
U.S. GOVERNMENT PRINTING OFFICE: 1941
PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM
issued 4%
SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
Vol. 91 Washington: 1941 No. 3126
THE NEVADA EARLY ORDOVICIAN (POGONIP) SPONGE
FAUNA
By R. 8. Basser
Tue discovery in 1927 by H. G. Clinton and Percy Train, of Man-
hattan, Nev., of a new fossil sponge fauna in Upper Pogonip (Chaz-
yan) strata of that State, characterized by the trilobite Plomerops
barrandezt Billings, was of such interest that I was prevailed upon to
describe it immediately without illustration, so that the many dupli-
cate specimens belonging to their collections could be sent out with
definite specific names to interested students. It is regretted that the
illustration of these new genera and species has been delayed until
the present time, but the literature upon Paleozoic fossil sponges
grows so slowly that apparently there has been no conflict in the
matter of synonymy. Uncertainty as to the exact location of these
sponge-bearing beds, which was quoted as McMonnigal Canyon,
Monitor Range, 10 miles west of Devils Punch Bowl in Monitor
Valley, Nev., had also to be removed.
Dr. Edwin Kirk, in the course of his stratigraphic studies of the
western Paleozoic for the United States Geological Survey in 1928,
visited the type locality for these sponges. This proved to be the
hillside slope above the cabin half a mile south of Ikes Canyon, 4
miles west of Dianas Punch Bow] as registered in 1929 on the Roberts
Mountain quadrangle, Nev., these being the modern names for
McMonnigal Canyon and Devils Punch Bowl, the latter occurring
only 4 miles east of the canyon. Furthermore, the mountain range
in question is now the Toquima Range in the Toiyabe National
406808—41 91
92 PROCEEDINGS OF THE NATIONAL MUSEUM Von, 91
Forest. Then, in the summer of 1939, Drs. Josiah Bridge and G. A.
Cooper had the opportunity of studying the area and obtaining addi-
tional collections besides confirming Dr. Kirk’s location. They report
that outcrops in the canyon itself afford good collections of the
sponges particularly on the north side about two-fifths of a mile
inside the entrance. Here the best fossils are found on a talus slope
50 to 70 feet above the valley floor below the big cliff, although some
may be collected from outcrops in the several ravines.
Associated with these sponges and the trilobite Pliomerops is an
undescribed fauna of Ostracoda, a few stony Bryozoa, crinoid and
cystid remains, trilobites, cephalopods, gastropods, and brachiopods.
Of the last, the following species were described as new by Ulrich
and Cooper in 19361: Aporthophyla typa, Toquimia kirkt, Goniotrema
perplexa, Rhysostrophia nevadensis, and R. occidentalis. 'This part
of the Pogonip limestone seems to be represented elsewhere in North
America in the Table Head formation of Newfoundland and the Oil
Creek formation of Oklahoma.
These Nevada fossil sponges are preserved in a thin-bedded, dense,
clayey limestone composed largely of organic remains and often
weathered enough at the surface to show silicification of the contained
fossils. With further etching by acid the minute spicular structure
of the sponges can be seen to better advantage at their surface, but
farther within where water has not penetrated the spicules have the
same calcareous structure as the rest of the material. In practically
all publications on the order Tetractinellida of the Silicispongiae,
authors describe the spicules as originally siliceous but explain that
when found calcareous the silica has been replaced by lime. Should
that be true, all these early as well as later Paleozoic sponges have
without exception been so replaced, a phenomenon that certainly has
not occurred so uniformly. These sponges undoubtedly follow the
rule of all other Paleozoic fossils that whenever they are buried in a
calcareous siliceous shale or certain clayey limestones the organic
calcite is replaced at the surface by silica, but the original structure
on the interior remains calcium carbonate just as it does in most other
fossils. Associated with these sponges are great numbers of long,
needle-shaped structures, which may be dermal spicules. These are
here illustrated (pl. 21, fig. 7) as a doubtful species of Hyalostelia,
but their relationship, if any, to the associated sponges has not been
discovered.
The original abbreviated descriptions of the following species, with
the exception of one new form, appeared in the Journal of the
Washington Academy of Sciences, volume 17, No. 15, pages 391-394,
1 Journ. Pal., vol. 10, pp. 616-631, 1936.
BASSLER 93
NEVADA FOSSIL SPONGE FAUNA
1927. Bibliographic references to this paper are omitted in the
present one since all descriptions previous to Patellispongia are
printed on page 392, while that genus and Hesperocoelia are described
on page 393, and the Anthaspidellidae on page 394. Again, the
horizon and locality are not mentioned each time because for all
the species it is, as stated before, the Upper Pogonip (Chazyan) lime-
stone, one-half mile south of Ikes Canyon, 4 miles west of Dianas
Punch Bow! on the eastern front of the Toquima Range, Roberts
Mountain quadrangle, Nev. The same assemblage of species occurs
in Ikes Canyon itself, as mentioned before.
All the illustrations of this paper are unretouched photographs,
except that the pore structure has been emphasized enough to make
it visible. The photography of the thin sections proved difficult,
since on enlargement the definite structure of the spicules loses much
of its clearness.
Subclass SILICISPONGIAE: Order TETRACTINELLIDA
Family ARCHAEOSCYPHIDAE Rauff
Archaeoscyphidae RAurr, Paleontographica, vol. 40, p. 288, 1894.
Sponge attached, simple or branching, ranging from narrow
cylindrical to saucer or funnel shaped, turbinate and frondescent
forms with simple or branched cloaca usually of considerable diam-
eter; oscula represented by numerous often closely spaced, small pores
penetrating the spicular tissue of the wall as definite canals and
opening on the outer surface at regular intervals.
With the recognition of five genera in the Nevada Pogonip fauna,
this family, formerly based upon a single species of the genus
Archaeoscyphia, assumes some importance in the early Ordovician
rocks.
Genus ARCHAEOSCYPHIA Hinde, 1889
Archaeocyathus (part) Bitrines, Paleozoic fossils, Geol. Surv. Canada, yol. 1,
p. 354, 1865.
Archaeoscyphia HINDE, Quart. Journ. Geol. Soc. London, vol. 45, p. 142, 1889.—
RavrF, Palaeontographica, vol. 40, p. 238, 1894.
Sponge simple, attached, short but rapidly expanding funnel-shaped,
6 cm. or more high and about 4 cm. wide, with a broad cloaca 3 cm.
in maximum diameter and the outer surface bearing strongly marked,
angular, parallel, transverse ridges. Wall 5 mm. thick, lined on
both the inside and outside by longitudinal rows of closely spaced
pores traversing the spicular skeleton, which consists of minute sili-
ceous spicules of the tetractinellid type with the rays slightly branched
at their extremities and interlocking without forming prominent
nodes,
94 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
The genotype and only species, A. minganensis, is not any too well
known, but judged from the description and illustrations by Billings
and Hinde the type specimens, although not well preserved, appear to
have the characters mentioned above.
ARCHAEOSCYPHIA MINGANENSIS (Billings)
PLATE 23, FiaguRES 3-6
Petraia minganensis BILLINes, Can. Nat. and Geol., vol. 4, p. 346, 1859.
Archaeocydthus minganensis Brines, Paleozoic fossils, Geol. Surv. Canada,
vol. 1, p. 354, figs. 342, 3438, 1865.
Hthmophylium minganense Watcorr, U. 8. Geol. Surv. Bull. 30, p. 77, figs.
6-8, 1886.
Archeoscyphia minganensis Hinprk, Quart. Journ. Geol. Soc. London, vol. 45,
p. 143, pl. 5, figs. 12-14, 1889—Ravurr, Paleontographica, vol. 40, p. 240,
pl. 1, figs. 1-10, 1894 (see for complete bibliography ).—TWENHOFEL, Geol.
Soe. Amer. Special Pap. 11, p. 34, 1988.
In spite of the various researches upon this species and its ref-
erences to several divisions of the animal kingdom, its exact structure
has not yet been confirmed because of a lack of good study material.
However, the several figures on plate 23 copied from Billings and
practically the same as given in all the references, show that Ar-
chaeoscyphia is a sponge possessing the same general type of structure
as the other genera here referred to the family.
Chazyan (Romame formation): Montagne (Big Romaine) Island
(Mingan Islands), St. Lawrence River, Quebec.
Genus NEVADOCOELIA Bassler, 1927
Simple, erect, obconical to oval, pedunculate sponges pierced
throughout their length by a cloaca about one-third the width and
marked on the outer surface by transverse parallel ridges or rows of
nodes. Pores (oscula) of sponge wall small, appearing at the surface
in more or less closely spaced parallel rows and on the interior as
canals arising from the cloaca and bending gradually to the surface
with the intervening spaces composed of the usual spicular structure
characteristic of the family.
Genoty pe.—Nevadocoelia wistae Bassler.
NEVADOCOELIA WISTAE Bassler
Pate 19, FiecurEs 6, 7; PLATE 24, F1GUREs 6, 7
Sponge elongate, cylindrical to oval, arising gradually from a
narrow base to a length of 12 cm. or more and a width of 4 cm.,
with the cloaca about 18 mm. in diameter. Surface marked by un-
NEVADA FOSSIL SPONGE FAUNA-——-BASSLER 95
dulating, more or less parallel, transverse ridges 1 to 144 mm. wide
and 21% mm. apart, with 7 occurring in 38cm. Sponge pores averag-
ing 0.35 mm. in width, separated by about their own diameter and
opening on outer surface in more or less regular longitudinal rows.
In longitudinal section the pores arise at the cloaca and bend gradu-
ally upward at an angle of about 30° to the surface.
Cotypes.—U.S.N.M. No. 79632.
NEVADOCOELIA TRAINI Bassler
PLATE 19, Figures 1-5
General characters as in the preceding species, but the growth
occurs in shorter, broader sponge bodies, averaging 8 cm. long and
4 cm. wide, with the cloaca about 12 mm. in diameter and the surface
marked by sharp nodes instead of parallel transverse ridges. Six
nodes occur on an average in 2 cm., measured transversely. Pore
structure very similar to the preceding species. In the several hun-
dred specimens of this and the preceding species no intermediate
forms were noted, so that the surface ridges and nodes seem to be
good specific characters.
Cotypes.—U.S.N.M. No. 79633.
NEVADOCOELIA GRANDIS Bassler
PLATE 19, FIGuRE 8
Sponge not unlike WV. wistae in growth and external structure but
much larger and with more separated and broader transverse ridges,
4 of which occur in 83cm. The cloaca is about 3 cm. wide, but the
pores piercing the outer surface have the same size and arrangement
as in the genotype. The type specimen, 15 cm. long and 9 cm. wide,
represents only the upper third of the entire sponge, so it might be
only a giant form of WV. wistae, but a smaller complete example (15
cm. long and 7 cm. wide, with cloaca also 3 cm. in width) shows the
transverse ridges equally large and distant from each other.
Holotype.—U.S.N.M. No. 79634.
NEVADOCOELIA PULCHRA Bassler
PLATE 20, Ficures 1-4
Sponge oval, 7 cm. in greatest diameter and more than 11 cm.
high, with the cloaca 1.5 to 3 cm. wide. Outer surface marked by
unusually strong ridges, which grow into wide, ascending, flangelike
expansions 5 mm. wide and distant at least 1 cm. from each other.
Pore arrangement and size as in other species of the genus, with 6
96 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
pores in 4 mm. measured lengthwise and 8 rows in the same space
transversely.
Holotype.—U.S.N.M. No. 79635.
Genus LISSOCOELIA Bassler
Smooth, cylindrical, hollow stems, branching dichotomously usually
in the same plane but at irregular intervals, constitute the growth
in this genus. The smooth surface under the lens shows minute
rounded pores penetrating the spicular tissue as in other members of
the family. These are the openings of the oscula, which in thin
sections are seen to be closely arranged tubes arising from the basal
wall and gently bending to the surface at a low angle. The cloaca
is narrow and extends the full length of the sponge.
Genotype.—Lissocoelia ramosa Bassler.
LISSOCOELIA RAMOSA Bassler
PLATE 19, Figures 9-11; PLate 24, Fiaures 4, 5
Sponge body of smooth hollow stems, usually about 114 em. wide
although increasing to 2 cm. at the place of branching, which occurs
at intervals of 3 cm. or more, often but not always in the same plane,
a complete growth being 10 cm. in diameter. The cloaca throughout
averages 0.5 cm. in width. Surface smooth, marked by minute
rounded pores about 0.20 mm. in diameter, distributed equally
throughout the spicular tissue at distances of 2 to 3 times their own
width. Spicules exceedingly minute but apparently with the same
structure as in the family. Sections show the cloaca varying from
3 to 5 mm. in diameter, with the oscula arising from the basal sponge
wall as narrow parallel tubes bending in a gentle curve to the surface.
This, one of the commonest of the Nevada sponges, is easily recog-
nized by its cylindrical branching stems with the markedly smooth
surface and very minute pore structure.
Cotypes.—U.S.N.M. No. 79636.
Genus CALYCOCOELIA Bassler
Sponge arising from a blunt broad peduncle into a goblet-shaped
body, which increases rapidly in width from below upward and then
opens at the upper surface in a deep excavation representing the
cloaca. Surface smooth but marked by minute, rounded pores, the
oscula arranged closely in rows parallel to the sponge length, these
representing openings of internal regularly arranged canals separ-
ated by a spicular meshwork as in related genera but with the spicules
exceptionally long and narrow rayed.
Genotype and only species.—Calycocoelia typicalis Bassler.
NEVADA FOSSIL SPONGE FAUNA—BASSLER 97
CALYCOCOELIA TYPICALIS Bassler
PuaTE 21, Fiegures 3-5; PLATE 24, Ficure 3
The goblet-shaped form deeply excavated by the wide cloaca of
about 15 mm. diameter and the smooth, minutely porous surface
characterize this species. The type specimen is nearly 7 cm. in
diameter at the top, decreasing to 4 cm. at the pedunculate base.
The pores, arranged in regular, longitudinal, parallel series, measure
about 10 rows in 10 mm.
Holotype.—U.S.N.M. No. 79637.
Genus PATELLISPONGIA Bassler
Sponge as usually found consisting of unilamellar fragments, some-
times of considerable dimensions, but originally probably broad
saucer-shaped expansions attached by a short stem. Under surface
comparatively smooth, covered by a thick dermal tissue pierced by
minute, closely spaced pores, which when weathered usually show a
regular arrangement in rows parallel to the direction of growth.
Passing through the spicular tissue and opening at right angles
at the upper surface these pores reappear as more or less evenly
spaced rounded canals representing the oscula, surrounded by the
usual spicular tissue of the family.
Genotype.—Patellispongia oculata Bassler.
PATELLISPONGIA OCULATA Bassler
PLATE 22, Ficures 1, 2; PLATE 24, Ficures 1, 2
This species forms broad lamellar expansions 12 cm. or more in
diameter and 1 cm. thick attached by a short peduncle. Upper surface
exhibiting numerous rather regularly spaced pores, the openings of
the oscula nearly 1 mm. in diameter with nearly 6 in 10 mm. and
separated by about their own diameter. Under surface smooth
marked by pores 0.4 mm. wide, with 9 in 5 mm. but without any
special arrangement.
Holotype-—U.S.N.M. No. 79638.
PATELLISPONGIA CLINTONI Bassler
PLATE 20, Figures 5-7
Sponge similar to the preceding in growth and other characters, but
the pores on the upper surface are somewhat larger, open on slight
elevations, and (more important from a specific standpoint) from 4
to5mm.apart. The under side of the lamella, as in other species of the
genus, is smooth and shows minute closely spaced pores in the spicular
98 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
tissue, these in the present case being about 0.35 mm. wide and separated
by their own diameter.
Cotypes—U.S.N.M. No. 79689.
PATELLISPONGIA MINUTIPORA Bassler
PLATE 21, FiaurEs 1, 2
Sponge consisting of a thin, expanded, smooth lamella, 12 cm. or
more wide and 4 mm. thick, differing particularly from other members
of the genus in the minuteness and close spacing of the pores on both
sides. At least 15 pores can be counted in 10 mm. on the upper surface,
where they occur at regular intervals and average 0.5 mm. in width.
The basal surface shows pores of about the same dimensions as the
upper but arranged in longitudinal parallel series.
Holotype —U.S.N.M. No. 79640.
PATELLISPONGIA MAGNIPORA, new species
PLATE 21, HIGURE 6
Sponge a unilamellar expansion 10 cm. or more broad, 3 to 8 mm.
thick, with a smooth but minutely porous base and an upper surface
marked by wide, open canals 2 to 2.5 mm. in diameter, irregularly
arranged and spaced at distances several times their width in the usual
spicular tissue. The minute pores of the basal side are closely spaced
in equally closely arranged parallel longitudinal rows with 7 pores in
5 mm. measured longitudinally, each pore about 0.6 mm. wide.
The surface pores or canal openings in the species, represented by
four specimens, are the largest so far noted in the genus, which fact in
addition to their irregular arrangement causes easy recognition.
Holotype—U.S.N.M. No. 99602.
Genus HESPEROCOELIA Bassler
General structure as in Patellispongia except that the broad, thin,
saucer-shaped lamella of that genus is here represented by a flat, undu-
lated frond or convoluted sponge body with pore openings of similar
size and arrangement on each face but penetrated lengthwise by a cloaca
in the form of a narrow to broad, flattened tube or series of tubes, open-
ing along the upper edge in a row of rounded or oval apertures.
Genotype.—Hesperocoelia typicalis Bassler.
HESPEROCOELIA TYPICALIS Bassler
PLATH 22, FicurES 6-8; PLATE 24, F1cuRE 9
Sponge a smooth, flattened, flabellate frond, 5 cm. or more in diameter
and about 6 mm. in thickness, traversed by longitudinal canals of vary-
NEVADA FOSSIL SPONGE FAUNA—BASSLER 99
ing width representing the cloaca, emerging at the surface along the
upper thin edge in a row of narrow openings, each about 3 mm. long
and 1 mm. wide, spaced so that 4 or 5 occur in 20 mm. The usual
openings or oscula in the spicular tissue show on both sides of the
sponge, with an average of 4 pores in 3 mm. measuring longitudinally.
Cotypes—U.S.N.M. No. 79641.
HESPEROCOELIA UNDULATA Bassler
PLATE 22, FIGURES 8-5; PuLate 24, FiaurE 8
This species differs from the preceding in forming undulated, often
convoluted bodies 8 em. or more high and 1 em. thick and in the fact
that the cloacal openings along the upper edge of the sponge are round,
3.5 to 4mm. in diameter with 4 or 5 in 20 mm. Moreover, the small
pores penetrating the spicular tissue are more delicate and closely
spaced.
Cotypes.—U.S.N.M. No. 79642.
Family ANTHASPIDELLIDAE Ulrich and Everett, 1890
Sponges attached, saucer to funnel shaped, often turbinate with
canal system usually consisting of two sets, one radial and one vertical,
crossing each other at right angles. Skeleton of 4-rayed spicules con-
sisting of a rodlike central part and rapidly diverging bifurcations at
each end, uniting to form radial columns, which when connected by
the horizontal central rods form a minutely tubular meshwork.
Genus ANTHASPIDELLA Ulrich and Everett
Anthaspidella ULRicH and Everett, Geological Survey of Illinois, vol. 8, pp. 255,
256, 1890.
Flat to saucer or funnel shaped sponges supported by a short sub-
cylindrical stem with the upper surface showing oscula, each provided
with its own system of radiating channels, all of which, however, merge
into the prevailing structure. Depressed part of each osculum occu-
pied by a few rather large, thin-walled, vertical tubes. Lower surface
of sponge occupied by rounded canal openings in spicular meshwork,
arranged in more or less radiating rows.
Genotype—Anthaspidella mammulata Ulrich and Everett.
ANTHASPIDELLA CLINTONI Bassler
PLATE 23, FIGURE 9
Sponge of large flattened disks, the type specimen a fragment 9 by 11
cm., indicating a diameter of at least 20 cm. for the entire body and
a maximum thickness of 1 cm. Although similar to Anthaspidella
100 PROCEEDINGS OF THE NATIONAL MUSEUM VOU, 91
scutula Ulrich and Everett, from the Black River (Platteville) lime-
stone at Dixon, IIl., in the small size and comparatively close arrange-
ment of the clusters consisting of the oscula and radiating canals, the
present species differs in that the clusters measuring from center to
center are closer (15 mm.) and coarser, and the canals are shorter,
broader, and less regularly arranged.
Holotype—vU.S.N.M. No. 79648.
ANTHASPIDELLA TRAINI Bassler
PLATE 23, FicureEs 7, 8
Sponge suggesting Anthaspidella florifera Ulrich and Everett, a
small saucer-shaped species from the Black River (Platteville) lime-
stone at Dixon, IIl., but differing in that the body is flat, at least 20 cm.
in diameter, 1 cm. thick, and the clusters are coarser and farther apart,
ranging from 25 to 35 mm. distant from center to center. The canals
in each cluster are also fewer, broader, and radiately arranged.
Holotype —U.S.N.M. No. 79644.
Genus STREPTOSOLEN Ulrich and Everett
Streptosolen ULRicH and EVERETT, in Miller, North American geology and
paleontology * * *, pp. 153, 165, 1889.
The very irregular arrangement of the canals that pass through the
sponge mass in every direction is the chief character separating this
genus from Anthaspidella and other members of the family. The
canals in Streptosolen intertwine to such a degree that it is difficult to
separate the two sets.
Genotype.—Streptosolen obconicus Ulrich and Everett.
STREPTOSOLEN OCCIDENTALIS Bassler
Prare 23, FrcureEs 1, 2
Sponge with the form and general structure of the genotype from
the Black River (Platteville) limestone at Dixon, IIl., but differing in
that the canals do not intertwine so much and the central osculum is
much wider and has larger tubes.
Cotypes.—U.S.N.M. No. 79645.
EXPLANATION OF PLATES
[All the specimens figured are from the Upper Pogonip (Chazyan) limestone,
half a mile south of Ikes Canyon, 4 miles west of Dianas Punch Bow], Roberts
Mountain quadrangle, Nev. Unless otherwise stated, the figures are natural
size. ]
PLATE 19
1-5. Nevadocoelia traini Bassler: (1) View of upper two-thirds of type, show-
ing the characteristic sharp nodes of the surface; (2) surface, X 6,
with nodes and pores (oscula) ; (3) cross section of top showing width
of cloaca; (4, 5) two views of three illustrating canals, X 6, their open-
ing at the surface in definite rows, and spicular tissue.
6,7. Nevadocoelia wistae Bassler: The type specimen, illustrating the some-
what closely spaced, narrow, more or less parallel transverse ridges and
view of the surface, X 6, showing pores and spicular structure. (See
also pl. 24, figs. 6, 7.)
8. Nevadocoelia grandis Bassler: Portion of the type illustrating the large
dimensions and the strong, widely-spaced surface ridges.
9-11. Lissoceelia ramosa Bassler: The type (9) a smooth cylindrical branching
stem, with end view (10) showing its hollow nature, and surface, X 6
(11) illustrating spicular structure and minute rounded pores. (See
also pl. 24, figs. 4, 5.)
PLATE 20
1-4. Nevadocoelia pulchra Bassler: Side and top views of the type (1, 2)
illustrating the wide, flangelike, ascending expansions and the central
cloaca, with enlarged views (3, X 6; 4, X 20) exhibiting the rows of
minute pores and spicular structure.
. Paiellispongia clintoni Bassler: The type specimens, parts of saucer-shaped
unilamellate expansions (5, 6) and surface, < 6 (7) showing the pores
widely separated by spicular tissue.
c
PLATE 21
1,2. Patellispongia minutipora Bassler: Upper surface of type, a fragment of
a thin lamella, and surface, < 6, illustrating the minute closely spaced
pores with intervening spicular tissue.
3-5. Calycocoelia typicalis Bassier: (3, 4) Side and top views of this goblet-
shaped sponge with several areas of oscula darkened to show arrange-
ment; (5) surface view, X 20, showing the oscular pores and the spicu-
lar structure. (See also pl. 24, fig. 3.)
. Patellispongia magnipora, new species: Portion of the type, a unilamellate
expansion, and a small portion, X 6, illustrating large, widely spaced
pores in broad areas of spicules.
7. Hyalostelia ? species: View of needle-like rods, some 50 mm. long, occur-
ring in thick layers, accompanying the various species of sponges herein
described.
fon)
101
102 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
PLATE 22
1,2. Patellispongia oculata Bassler: Portion of the type, a broad lamellar
expansion showing upper surface with regularly but widely spaced
pores about 1 mm. in diameter and enlarged view (X 6) with spicular
structure between pores more visible. (See also pl. 24, figs. 1, 2.)
3-5. Hesperocoelia undulata Bassler: Side and edge views of the flat, undulated
frond (3, 4) pierced by a row of oval openings representing the cloaca,
and surface of same, X 6, exhibiting size and distribution of the minute
pores and the intermediate spicular tissue (5). (See also pl. 24, fig. 8.)
6-8. Hesperocoelia typicalis Bassler: The flattened flabellate type specimen (6)
traversed by longitudinal canals representing the cloaca emerging at
the upper thin end (7) and view of surface <* 6 (8) exhibiting pores
and intermediate tissue. (See also pl. 24, fig. 9.)
PLATE 23
1,2. Streptosolen occidentalis Bassler: Side of one of the types exhibiting
irregularly spaced canal openings and top of larger example showing
wider central osculum and tubes larger than in the type species.
3-6. Archaeoscyphia minganensis (Billings) : Drawing of a restored specimen
about one-half natural size (3) and sketches of three forms of spicules,
<x 80 (46). Chazyan (Mingan): Mingan Islands (after Billings,
1865).
7,8. Anthaspideila traini Bassler: The type, an incomplete specimen showing
the large, coarse, rather widely spaced ciusters (7) and spicular struc-
ture exhibited on etched surface, X 20 (8).
9. Anthaspidella clintoni Bassier: Part of surface of type exhibiting com-
paratively small size and close arrangement of the clusters consisting
of oscula and radiating canals.
PLATE 24
(Thin sections, all X 9, with structure emphasized by shading in some cases.)
1,2. Pateilispongia oculata Bassler: Vertical and tangential sections showing
arrangement of canals and spicular meshwork. (See also pl. 22, figs.
Le)
3. Calycocoelia typicalis Bassler: Section crossing canals and spicular mesh-
work, illustrating size and length of spicules. (See also pl. 21, figs. 3-5.)
4,5. Lissocoelia ramosa Bassier: Tangential section near surface where minute
pores and fine spicular structure are best shown and vertical section
through wall with canals and meshwork. (See also pl. 19, figs. 9-11.)
6,7. Nevadocoelia wistae Bassler: Part of vertical section with several canals.
Most of the spicules are cut so as to show in white points (6). Tan-
gential section (7) through pores and spicular mesh, with a sketch
XxX 15. (See also pl. 19, figs. 6, 7.)
8. Hesperocoelia undulata Bassler: Transverse section through cloaca with
canals and usual tissue, with a small sketch of spicular structure, X 20.
(See also pl. 22, figs. 3-5.)
9. Hesperocoelia typicalis Bassler: Cross section through cloaca with canals
and spicular structure; and small portion of the latter X 20. (See also
pl. 22, figs. 6-8.)
U.S. GOVERNMENT PRINTING OFFICE: 1941
U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 91 PLATE 19
EARLY ORDOVICIAN SPONGES.
FOR EXPLANATION OF PLATE SEE PAGE
U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 91 PLATE 20
EARLY ORDOVICIAN SPONGES.
FOR EXPLANATION OF PLATE SEE PAGE 101,
U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 91 PLATE 21
EARLY ORDOVICIAN SPONGES.
FOR EXPLANATION OF PLATE SEE PAGE
U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 91 PLATE 22
EARLY ORDOVICIAN SPONGES.
FOR EXPLANATION OF PLATE SEE PAGE 102.
U.S. NATIONAL MUSEUM PROCEEDINGS, VOL. 91 PLATE 2
EARLY ORDOVICIAN SPONGES.
FOR EXPLANATION OF PLATE SEE PAGE
U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. 91 PLATE 24
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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM
SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
Vol. 91 Washington: 194] No. 3127
THE MEXICAN SUBSPECIES OF THE SNAKE CONIO-
PHANES FISSIDENS
By Hosarr M. Smiru
In the recent revision of the snakes of the genus Coniophanes Cope,
Bailey? tentatively concluded that mainland specimens of fissidens
must remain under that name, pending the accumulation of further
material that would more clearly delimit the geographic races vaguely
indicated by material then available.
Since the appearance of this work many specimens of these reptiles
have been collected from critical areas in Mexico, chiefly for the
National Museum and for the E. H. Taylor-H. M. Smith collection
at the University of Kansas. This new material, combined with that
already available, has been sufficient to demonstrate rather clearly the
existence in Mexico of four distinct races, occupying as many different
geographic and faunal areas and differing from one another in details
of pattern as well as in average scale counts.
I am indebted to Dr. E. H. Taylor and Dyfrig McH. Forbes for
much assistance in the field and for the loan of specimens. The study
was completed, and a portion of the material was collected, during
my tenure of the Walter Rathbone Bacon Traveling Scholarship of
the Smithsonian Institution.
1 Bailey, Joseph, Papers Michigan Acad. Sci., Arts and Lett., vol. 24, pt. 2, pp. 1-48, figs.
1-5, pls. 1-3, 1939.
103
406804—41
104 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
«
CONIOPHANES FISSIDENS FISSIDENS (Giinther)
Coronella fissidens GUNTHER, Catalogue of the colubrine snakes in the collection
of the British Museum, p. 36, 1858 (Mexico).
Diagnosis.—Scales in 21 rows (rarely 19); males with supraanal
ridges; supralabials 8 (rarely 7); ventrals 117 to 132 in females, 111
to 130 in males; caudals 63 to 79 in females, 62 to 84 in males; ventrals
minus caudals 48 to 63 in females, 38 to 56 in males; a relatively large
spot toward each end of ventrals (usually in addition to numerous
smaller, scattered spots) ; belly always spotted; median dark borders
of dorsolateral light stripe not distinct in front of anus; dorsolateral
light stripe visible a considerable length on neck; no spots or irregu-
larities of pattern in dorsal area between lateral stripes.
Discussion.—The limits of variation of this subspecies are established
by Bailey, whose tabulations for specimens from Honduras south to
Panama are here utilized in addition to data derived from specimens
in the National Museum. I have not utilized his tabulations for
specimens from British Honduras, Guatemala, and Mexico, since sev-
eral forms are involved in these countries.
Mexican specimens I have seen are from Teapa, Tabasco (U.S.N.M.
No. 46590), and San Andrés Tuxtla, Veracruz. The latter is probably
very near the northern limit of the range of the subspecies. Both
have higher ventral counts than typical f. fisstdens and accordingly
show a tendency toward f. proterops. Four other specimens examined
are from very near Mexico (Piedras Negras, Guatemala, U.S.N.M.
Nos. 109720-109722, and one specimen, HMS No. 7353, in the KHT-
HMS collection). These are typical and have a regular series of
relatively large dark spots near the ends of the ventrals.
The eight cotypes of f. fisstdens, the scutellation of which is given by
Boulenger,’? almost certainly include proterops as well as that here
defined as fisstdens. So far as available data on these cotypes indicate,
the preponderance of characters are of f. fisstdens as here defined, to
wit: Ventral and caudal counts typical in three, possible in three;
scale rows typical in six; supralabials certainly typical in six. The
characters indicating proterops are: Ventral and caudal counts typical
in two, possible in three; scale rows typical (fide Boulenger) in two;
supralabials possibly typical in two. Accordingly I restrict the name
to the form here defined as f. fisstdens and to that cotype which most
closely corresponds with all characters defining the form.
* Catalogue of the snakes in the British Museum (Natural History), vol. 3, pp. 207, 208,
1896.
MEXICAN CONIOPHANES FISSIDENS—SMITH 105
CONIOPHANES FISSIDENS PROTEROPS Cope
Coniophanes proterops Corr, Proc. Acad. Nat. Sci. Philadelphia, 1860, p. 249
(Orizaba, Veracruz).
Diagnosis —Scales usually in 19 rows, sometimes 21; males with
supraanal ridges; supralabials usually 7, sometimes 8; ventrals 129
to 138 in females, 126 to 133 in males; caudals 59 to 74 in females, 66
to 76 in males; ventrals minus caudals 55 to 79 in females, 51 to 65 in
males; spots on belly very small, scattered; belly sometimes unspotted
(except ends of ventrals, dark as sides of body); median border of
dorsolateral light stripe usually very poorly defined on tail, or whole
dorsal surface light ; dorsolateral light stripe disappearing on anterior
part of neck; median dark stripe, one scale wide, distinct in young
and subadults, which are somewhat orange colored; no spots or irregu-
larities of pattern in dorsal area between lateral stripes.
Discussion.—The form seems well differentiated from f. fissidens.
The scale rows are usually 19 (69 percent, 24 in 36), 21 in fissidens
(two exceptions in 132); and the supralabials are usually 7 on one or
both sides (58 percent, 20 in 36), rarely in fisstdens (one in 132). ‘There
are conspicuous differences in ventral and ventral minus caudal counts,
as shown in the accompanying table.
The most conspicuous pattern difference between proterops and
fissidens is in the disposition of the ventral spots. In the latter there
is a row of relatively large spots on each side of the abdomen, one spot
near the end of each ventral. ‘These spots occur in addition to numer-
ous other, small flecks. In proterops the ventral surface is marked
with numerous tiny black flecks, but there is no regular series of
relatively large spots forming a row on either side of the belly. In
some proterops there are no ventral markings whatever, except on
the ends of the ventrals where the lateral coloration encroaches upon
the ventral surface.
The dorsal pattern as a rule is more subdued in proterops. The
lateral stripes are sometimes scarcely distinguishable, and the dorso-
lateral light stripes are very short or not visible at all. The dorsal
surface of the tail, which in fissidens bears two dorsolateral light stripes
separated by a very well defined median stripe, is nearly uniform light
in proterops asarule. Some proterops, however, do show the median
dark stripe.
The young of proterops are light orange, and the median dark stripe
is very well defined. Larger specimens show but little trace of the
orange coloration, the middorsal stripe is diffused, and the general
coloration much darker and more like that of typical fissidens.
Specimens examined.—Thirty-six, all from the State of Veracruz.
The following localities are represented: Cuautlapan (U.S.N.M. Nos.
109764-109766; EHT-HMS Nos. 5199, 23537-23545); Jalapa
106 PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 91
(U.S.N.M. No. 5285, type) ; Mirador (U.S.N.M. Nos. 6369[3], 12112,
25034, 46452-46453) ; Orizaba (U.S.N.M. Nos. 12117, 30858) ; Potrero
Viejo (U.S.N.M. Nos. 109767-109768; D. McH. Forbes No. 256;
EHT-HMS No. 5528); Tequeyutepec, 7 miles west of Jalapa
(U.S.N.M. Nos. 109769-109773; D. McH. Forbes Nos. 386-388) ; San
José de Gracia (EHT-HMS No. 5529).
CONIOPHANES FISSIDENS DISPERSUS, new subspecies
Holotype—EHT-HMS No. 5531, male, El Limoncito, Guerrero,
collected by E. H. Taylor.
Paratype—EHT-HMS No. 5532, same data.
Diagnosis.—Scales in 19 rows; males with supraanal tubercles; su-
pralabials 8; ventrals 120 and 122 in two males; caudals 81 in one male;
ventrals minus caudals 41 in one male; spots on belly small, scattered,
not forming regular series; middle and posterior part of belly may
be completely unspotted (except ends of ventrals) ; white dorsolateral
stripes on nape short, diffuse; inner border of dorsolateral light stripe
poorly defined on tail, not evident in front of anus; no spots or irregu-
larities of pattern in dorsal area between lateral stripes; latter poorly
defined, diffuse.
Description of holotype-—Rostral much broader than high, portion
visible from above a little less than half length of internasals; latter
two-thirds length of prefrontals; frontal pentagonal, anterior edge
straight, its length (4.8 mm.) greater than its distance from tip of
snout (4.3 mm.), less than maximum length of parietal (5.9 mm.),
subequal to distance of frontal from posterior edge of parietal (not
in median line); sides of frontal parallel; nasal large, divided; a
large loreal; one preocular; two subequal postoculars; temporals
1+2+3, the upper secondary and upper tertiary fused to form an
elongate scale similar to primary temporal; eight supralabials, fourth
and fifth entering orbit, seventh highest and largest, sixth next largest ;
nine infralabials, five in contact with chin shields; mental separated
from anterior chin shields, which are a little longer and larger than
posterior.
Dorsals in 19-19-15 rows, smooth, without pits; ventrals 122; caudals
81; anal divided.
Ground color light grayish brown, somewhat darker near middle of
body and on sides; a broken, dim dark line extending along adjacent
edges of fourth and fifth scale rows, descending posteriorly to middle
of fourth; a broken, scarcely discernible vertebral dark line; a dorso-
lateral area somewhat lighter, well defined only on nape, where it is
white; a small white spot three scales back of tertiary temporals, even
with end of dorsolateral light line, which terminates two scales behind
light nuchal spot; all dorsal scales with a dark edge. A dark stripe
MEXICAN CONIOPHANES FISSIDENS—SMITH 107
through the upper edges of supralabials, white-edged below; re-
mainder of supralabials stippled, and each (except eighth) with a
larger, rounded dark spot near center of light area; chin, infralabial,
and gular regions stippled; some larger black dots on certain infra-
labials. Tail with a dark lateral stripe, black-edged above; area be-
tween these, on dorsal surface, darker near middle; edges of sub-
caudals dark; ends of ventrals dark-spotted; a few small, scattered
spots on anterior ventrals; remainder of ventral surface white.
Variation—The single paratype is a male with 120 ventrals, tail
tip missing. The scales are in 19-19-15 rows. Supralabials 8, infra-
labials 10, one preocular, two postoculars, temporals as described in
type.
Coloration as in type, except dorsolateral light stripes somewhat
more evident; a faint, broken, very fine temporal stripe; small dark
spots irregularly placed near ends of ventrals.
Remarks.—W ith this subspecies the specimen from Carrizal, Micho-
acin (Brit. Mus. No. 1914.1.28.141) is to be associated; also perhaps
another from Cafetal Mirador, Oaxaca (A. M. N. H. No. 19748). These
are mentioned by Bailey (op. cit., p. 23) ; Ihave not seen them, nor are
counts available.
This subspecies resembles proterops in the possession of 19 scale
rows but differs in having higher caudal and lower ventral counts and
eight supralabials. It resembles fisstdens in ventral and caudal counts
but differs by lacking the regular series of spots near ends of ventrals
and by having only 19 scale rows. It resembles punctigularis in belly
coloration and number of ventrals and caudals but differs in dorsal
coloration, ventral minus caudal count, and by having 19 scale rows.
CONIOPHANES FISSIDENS PUNCTIGULARIS Cope
Coniophanes punctigularis Corg, Proc. Acad. Nat. Sci. Philadelphia, 1860, p. 248
(Honduras).—StLevin, Proc. California Acad. Sci., ser. 4, vol. 23, pp. 410-411,
1939.
Dromicus chitalonensis MULiER, Verh, Naturf. Ges. Basel, vol. 6, p. 407, 1876
(Hacienda de Chital6n, near Mazatenango, Guatemala).
Diagnosis —Scales in 21 rows; males with supraanal tubercles;
supralabials 8, rarely 7; ventrals 119 to 130 in females, 116 to 125 in
males; caudals 71 to 85 in females, 80 to 91 in males; ventrals minus
caudals 39 to 54 in females, 31 to 38 in males; spots on belly very small,
scattered; belly sometimes unspotted (except ends of ventrals, dark
as sides of body); median border of dorsolateral light stripes very
distinct on posterior part of body as well as on tail; dorsolateral light
stripes distinct on much of anterior part of body; a series of spots
on each side of middorsal line, about halfway between lateral and
middorsal stripe; spots fused with dorsolateral stripe in all except
108 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
young specimens, but always distinct laterally; body not orange in
young.
Discussion.—This form resembles proterops in belly coloration, lack-
ing the large lateral spots of fissidens, but with small, scattered spots
or belly unspotted. It resembles jfisstdens in number of scale rows,
labials, and in ventral and caudal counts, but it is very different in
dorsal pattern.
In punctigularis a series of rounded dark spots occurs on each side
between the lateral and dorsolateral dark stripes. In young specimens
the spots are free, but in older ones they merge medially with a vague
dark area. Even in the largest specimens the outer edges of the dark
spots are well defined, at least anteriorly, and give a scalloped effect
to the inner edge of the dorsolateral light stripe.
On the tail two light stripes, broader than in fissidens, are separated
by a median dark stripe, its outer border well defined. In fissidens
the inner edge of the light stripe is well defined only at the anus and
on the tail, while in punctigularis it is distinct on the posterior part
of the body as well.
Seven supralabials occur on one side in three specimens. Scale rows
21 in all.
The name chitalonensis certainly applies to this subspecies. How-
ever, punctigularis, proposed 17 years earlier, seems also to refer to
the same form. I have not seen the type, but Bailey (op. cit., p. 16)
says that “The middorsal band is 5 scales wide anteriorly and 3 wide
posteriorly. Anteriorly it is represented by a double row of darker
spots, which are evident posteriorly only as scalloped outer bor-
ders of the band. This type of coloration is frequent in western
Guatemala and Mexico.” Accordingly there can be little doubt that
this specimen (male, with 125 ventrals) is the same as that here rede-
fined. Its locality, however, is rather far removed from the nearest
authentic record in Guatemala. Records show that the collector of
the specimen, Dr. J. L. LeConte (and J. S. Hawkins), actually was
in Honduras, where he was connected for a few months in 1857 with
the Honduras Interoceanic Railway Survey. Faunistically the Pa-
cific coast mountain ranges of El Salvador are known to be much
like those of Guatemala and extreme southeastern Chiapas. It is to
this faunal area the present form is confined. Accordingly, if the
type of punctigularis actually is from Honduras, it must have been
collected on the Pacific side, and in that case very near El Salvador
and probably in the same faunal area as is represented by other speci-
mens. This would account for the difference between the type of
punctigularis and all other Honduras specimens, which are from the
Atlantic coast.
Specimens examined.—Fifty-one, from Tehuantepec (U.S.N.M.
Nos. 30167-30169, 30525) ; Chicharras, Chiapas (U.S.N.M. No. 46448) ;
MEXICAN CONIOPHANES FISSIDENS—SMITH 109
various localities in the vicinity of Escuintla, Chiapas (La Esperanza,
Cruz de Piedra, Salto de Agua, Finca Juarez, U.S.N.M. Nos. 109723-
109763) ; Tonala, Chiapas (EHT-HMS No. 53829). The “Tehuante-
pec” specimens presumably are from extreme eastern Oaxaca, Pacific
slope. Slevin (op. cit.) records 87 specimens from Finca El Ciprés,
Volcan Zunil, Guatemala. Data presented by him are included in the
accompanying table.
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Ficure 33.—Distribution of the Mexican forms of Coniophanes fissidens: Dots, except as
indicated, punctigularis; inverted triangles, fissidens; triangles not inverted, dispersus.
KEY TO THE MEXICAN SUBSPECIES OF CONIOPHANES FISSIDENS
1. A regular row of relatively large dark spots near ends of ventrals, in addition
to other dark flecks that may or may not be present; dorsolateral white
stripe extending posteriorly a considerable distance on neck; inner dark
border of dorsolateral tail stripes absent on posterior part of body, present
ODE Vgr ONSET) Skea eeereeeener nee ce 4 eee ere oe ee fissidens fissidens
No regular row of relatively large dark spots near ends of ventrals; belly with
small flecks of black, or unspotted; dorsolateral white stripes very short
(no more than twice length of head), or absent posteriorly, or, inner dark
border of dorsolateral tail stripes present on posterior part of body_------ 2
2. A series of dark spots between lateral and middorsal light lines; dark border
on inner edge of dorsolateral stripes distinct on body as well as tail; scales
AT eo eT OW Soe ee eee ee Se, Sn ae) ie ee ee f. punctigularis
Color not as described; markings dim, except (in young) a middorsal dark
INR SA a ag a ate te a eae ede 3
3. Ventrals fewer (120 to 122 in known males); caudals more numerous (81,
male); ventral minus caudal index lower (41, male) —--__-~-_ f. dispersus
Ventrals more numerous (126 to 133 in males); caudals fewer (64 to 76 in
males) ; ventral minus caudal index higher (51 to 65 in males).
f. proterops
PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM
SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
Vol. 91 Washington : 194] No. 3128
REPORT ON THE SMITHSONIAN-FIRESTONE EXPEDI-
TION’S COLLECTION OF REPTILES AND AMPHIBIANS
FROM LIBERIA
By Arruour Lovermce
Dr. William M. Mann, director of the National Zoological Park in
Washington and leader of the Smithsonian-Firestone Expedition,
1940, is to be congratulated on finding time to assemble a representa-
tive collection of the Liberian herpetofauna, despite the exacting
duties involved in the capture and care of wild creatures, the securing
of which was the primary purpose of his journey.
This collection, consisting as it does of over 500 specimens repre-
senting 56 species, naturally adds considerably to our knowledge of
the lower vertebrates of the country whose fauna is so imperfectly
known as that of the Liberian Republic. Among the results of a
study of this material, therefore, the following species have had to
be described as new:
Typhiops manni, new species from Harbel.
Hylambates cochranae, new species from Bendaja.
Leptopelis bequaerti, new species from Gbanga, Gibi, ete.
Rana albolabris parkeriana, new name for acutirostris Parker, preoccupied.
(This is the Angolan race of the typical form occurring in Liberia.)
In addition we are able to add the undermentioned to the steadily
growing list of species to be found within the boundaries of the
Republic:
Boaedon lineatus lineatus. Rana longirostris.
Crotaphopeltis duchesnii guineensis. Phrynobatrachus natalensis.
Hylambates leonardi.
113
406739—41——1
114 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
Neusterophis variegatus should be substituted for Natria fuliginoides,
whose admission was based on a misidentification, and Agama a.
africana (Hallowell) substituted for A. a. savatiert Rochebrune.
Certain other species should be regarded as synonyms, viz:
Aliurus Dunn and Dunn, 1940=Hemidactylus Gray, 1827 (not 1825).
Tropidonotus ferox Giinther, 1863=Natriz anoscopus anoscopus (Cope) 1861.
L. liberiensis Ahl, 1929=Leptopelis viridis (Giinther) 1868.
while
Lacerta langi Schmidt is revived as a race of L. echinata Cope.
Helicops gendrii Boulenger as a race of Natrixz anoscopus Cope.
Neusterophis variegatus (Peters) from synonymy of fuliginoides Giimnther.
Boaedon virgatus (Hallowell) is made a race of lineatus Duméril and Bibron.
Rana alleni (Barbour and Loveridge) a race of crassipes Peters.
Phrynobatrachus brongersmai Parker a race of ogoensis Boulenger.
Pertinent data regarding the material are supplied to enable fellow
herpetologists in checking identifications or extending the known
range of variation. I take this opportunity of thanking Dr. Doris
M. Cochran and Dr. W. M. Mann for the privilege of studying this
material now in the United States National Museum at Washington.
As none of the villages mentioned in this paper are to be found in
Stieler’s Atlas, the accompanying sketch map—kindly supplied by
Dr. Leonard P. Schultz, of the National Museum—is intended to
give the approximate positions of places from which specimens were
obtained. Dr. Mann has kindly furnished me with the under-
mentioned information regarding them, together with dates of the
itinerary. It has not been thought necessary to repeat these dates
except where some notes concerning breeding are involved. As re-
gards place names, Dr. Mann points out that no uniformity of spell-
ing is to be found on maps or in his correspondence with government
officials.
Bellyella: Spelling rendered in half a dozen different ways. No two maps of
Liberia locate this village in the same place. Dr. Mann has placed it
approximately in the position as given in the State Department’s map.
March 23-30, 1940.
Bendaja: Also spelled Bandeja, Bendeja, Bandaja, and Bendija. A village 5 or
6 miles from the border. May 14-27, 1940.
Bromley: A mission station on the St. Paul River above Monrovia. June 6-8,
1940.
Cape Mount: A name used locally for the Cape, the Mountain, the County, and
the mission. Robertsport is the chief town and port for the Cape Mount
district. May 7-12 and May 29-31, 1940.
Degain: Also spelled Dagain and Digain, a village where a night was spent on
the journey to Bellyella and return. March 22 and 81, 1940.
Gibi: Also spelled Gebi. The name applied to a low range of mountains whose
highest elevation is 2,042 feet. No collecting was carried out above 900 feet,
however, for Dr. Mann’s party stayed at Managey’s town while in the
vicinity. April 10-16, 1940.
REPTILES AND AMPHIBIANS FROM LIBERIA—LOVERIDGE 115
Harbel: This is the name of the Firestone Plantation, which covers an area of
approximately 25 square miles. It was the expedition’s headquarters from
March 10 to July 17, 1940.
Mombo: May 13 and 28, 1940.
Reputa: Also spelled Wreputa. June 21-26, 1940.
Zorzor: A mission station on the frontier of French Guinea about two days’
march, i. e., approximately 40 miles northwest of Bellyella. The locality
was not visited by members of the expedition, and the only specimens—
tortoises—from this locality were sent in by the missionaries.
TORTOISES
KINIXYS HOMEANA Bell
1827. Kinivys Homeana BELL, Trans. Linn. Soc. London, vol. 15, p. 400, pl. 17,
fig. 2 (West Africa).
444,322 (U.S.N.M. Nos. 109685, 109689-90, 109692-3, 109698-9), Zorzor
A nuchal shield, except in U.S.N.M. No. 109698, which is also
aberrant in possessing 21 marginals (all the rest have 22) and 5
(right) or 7 (left) costals (all the rest have 4); fifth vertebral
descending abruptly in all.
Males, characterized by longer tail and concave plastron, have a
gular suture, which is included in the total length of plastron 614-634
(7-714 in females) times, and an abdominal suture 17-2 (134 in
females) times as long as the pectoral suture. Shell of largest ¢
(U.S.N.M. No. 109689) measures 195 mm. over all; largest ¢?
(U.S.N.M. No. 109685) measures 223 mm.
KINIXYS EROSA (Schweigger)
1802. Tesiudo Denticulata SHAw (not of Linnaeus), General zoology, vol. 3,
pt. 1, p. 59, pl. 18 (‘‘Supposed to be a native of North America”).
1814. Testudo erosa SCHWEIGGER, Prodromi monographiae Cheloniorum, p. 52
(“America septentrionali (Shaw)”).
246,622 (U.S.N.M. Nos. 109687-8, 109691, 109694-7), Zorzor
No nuchal shield; 22 marginals, except in U.S.N.M. No. 109696
which has 24; costals 4; fifth vertebral descending obliquely in all.
Males, characterized by longer tail and concave plastron, have a
gular suture which is included in the total length of plastron 514-534
(634-714 in females) times, and an abdominal suture 214-814 (2-3 in
females) times as long as the pectoral suture. Shell of largest 2
(U.S.N.M. No. 109688) measures 260 mm. over all; largest
(U.S.N.M. No. 109687) measures 242 mm.
116 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
LIZARDS
HEMIDACTYLUS FASCIATUS Gray
1842. Hemidactylus fasciatus Gray, Zool. Misc., 1842, p. 58 (no locality).
1845. Leiwrus ornatus GrAy, Catalogue of the specimens of lizards in the
collection of the British Museum, p. 157 (West Africa) (monotype,
ornatus Gray; not Leiurus Hemprich and Ehrenberg, 1829, in
Arachnida).
1856. Hemidactylus formosus HALLOWELL, Proce. Acad. Nat. Sci. Philadelphia,
1856, p. 148 (Liberia).
1862. Liurus ornatus Corr, in Slack, Handbook of the Museum of the Academy
of Natural Sciences of Philadelphia, p. 32 (monotype, Hemidactylus
ornatus Hallowell; not Liwrus Ehrenberg, 1828, in Arachnida).
1940. Aliwrus ornatus DUNN and DuNN, Copeia, 1940, p. 71 (substitute name for
Liurus Cope, preoccupied by Liurus Ehrenberg, 1828, in Arachnida).
é (U.S.N.M. No. 108631), Harbel
Midbody rows of dorsal tubercles 25; lamellae under first toe 8,
under fourth toe 11; femoral pores 20+19; subcaudals more than
half the width of tail. Total length 182 (85+97) mm.
The recently proposed generic name of Adiwrus Dunn and Dunn
becomes a synonym of Hemidactylus, as will be seen from the some-
what complicated synonymy given above. I have been unable to
examine Slack’s rare Handbook and so quote the citation from it as
given by Dunn and Dunn.
AGAMA AGAMA AFRICANA (Hallowell)
1844, Tropidolepis Africanus HALLOWELL, Proc. Acad. Nat. Sci. Philadelphia,
1844, p. 171 (Liberia).
1845. Calotes versicolor HALLOWELL (not of Daudin), Proc. Acad. Nat. Sci.
Philadelphia, 1845, p. 247 (Liberia).
1884. Agama savatieri ROCHEBRUNE, Faune de la Sénégambie, Rept., p. 89, pl. 11,
figs. 1, 2 (Bathurst, Gambia) (restricted).
2¢¢4,2 92 (U.S.N.M. Nos. 109291-4), Bellyella
2 (U.S.N.M. No. 109580), Bendaja
Midbody scale rows 60-64; preanal pores 12-14. Larger 3
measures 112 mm. from snout to anus, tail truncate.
Heretofore (1936, p. 54) I have used savatiert for agamas of the
extreme west, which have fewer midbody scale rows than typical A.
a. agama ot the Cameroons, but Hallowell’s name africana, which was
overlooked by Boulenger and all subsequent herpetologists, is un-
doubtedly an Agama and has a wide margin of priority over any
other name that is applicable.
LACERTA ECHINATA ECHINATA Cope
1862. Lacerta (Zootoca) echinata Corn, Proce. Acad. Nat. Sci. Philadelphia,
1862, p. 189 (West Africa).
2 (U.S.N.M. No. 109632), Harbel
REPTILES AND AMPHIBIANS FROM LIBERIA—LOVERIDGE 117
Midbody scale rows 37; parietal present; gular granules between
chin shields and collar 30; femoral pores 12+12. Total length 342
(97+245) mm.
Trinomials are used on account of ZL. e. langi Schmidt (1919) of
the eastern Congo, which Boulenger (1920, p. 382) unjustifiably
synonymized with echinata. The type of the latter (U.S.N.M.
No. 5995) almost certainly came from Liberia, for it was described
at the same time as Cophoscincus dura, whose type (U.S.N.M. No.
5996) was also said to be from West Africa yet is known only from
Liberia.
MABUYA BLANDINGII (Hallowell)
1844. Huprepes Blandingii HALLOWELL, Proc. Acad. Nat. Sci. Philadelphia, 1844,
p. 58 (Liberia).
1857. Huprepes frenatus HALLOWELL, Proc. Acad. Nat. Sci. Philadelphia, 1857,
p. 50 (Liberia).
9 (U.S.N.M. Nos. 109024-32), Gibi
1 (U.S.N.M. No. 109295), Bromley
1 (U.S.N.M. No. 109581), Bendaja
1 (U.S.N.M. No. 109638), Harbel
Midbody scale rows 30-34; dorsals with 3 (in young) to 5, and
rarely even 7, keels; supranasals separated in five specimens, in con-
tact in eight; prefrontals separated in four examples, in contact in
nine; supraoculars 4; supraciliaries 83-6. The largest,a 9 (U.S.N.M.
No. 109295), measures only 182 (741108) mm. In its oviducts (April
10-16) are 4 eggs, measuring 12 by 7 mm., but without embryos.
A good deal of variation is displayed in the matter of coloration.
The dark brown lateral band is faintly edged above with white in a
young skink, below by a sharply defined white band in four speci-
mens, by a series of white flecks, or altogether lacking, in others.
Below, pure white.
COPHOSCINCOPUS DURUS (Cope)
1862. Tiliqua dura Corn, Proc. Acad. Nat. Sci. Philadelphia, 1862, p. 190 (Western
Africa).
1884. Cophoscincus simulans VAILLANT, Bull. Soc. Philom. Paris, ser. 7, vol. 8,
p. 170 (Couacrou, Ivory Coast).
1 (U.S.N.M. No. 109674), Gibi
The type of this interesting, though common, Liberian skink is in
the National collection (U.S.N.M. No. 5996). In all probability it
came from Liberia, as the only record of its occurrence elsewhere is
that of Vaillant, whose type locality I have failed to trace, unless it
be Kurako or Kurukoro, north of Ganta, in what is now French
Guinea. Owing to an unfortunate accident, the Gibi specimen is too
dried to be of much taxonomic value.
118 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
CHAMAELEO GRACILIS GRACILIS Hallowell
1842. Chamaeleo gracilis HALLOWELL, Journ. Acad. Nat. Sci. Philadelphia, vol. 8,
p. 324, pl. 18 (Liberia).
286,38 22 (U.S.N.M. Nos. 109019-23), Gibi
Males with tarsal spurs. Larger ¢ measures 186 (96+90) mm.;
largest @ measures 280 (132+148) mm. Trinomials are used on
account of C. g. etiennei Schmidt (1919) of Banana, Belgian Congo.
SNAKES
TYPHLOPS MANNI, new species
Type—U.S.N.M. No. 109634, from Harbel, Republic of Liberia,
March 10-July 17, 1940.
Diagnosis.—Agrees with 7. p. punctatus (including its Liberian
synonyms of léberiensis Hallowell, nigrolineatus Hallowell, and inter-
media Jan) in possessing 26 midbody scalerows. It differs from both
T. p. punctatus and 7. leucostictus in its broad and trilobate snout, lat-
eral nostrils, absence of an ocular, and in its midbody diameter being
included in its total length 40 times (instead of 24-86 times in punctatus,
45 in leucostictus). In addition, it differs from Jeucostictus in possess-
ing 26 (instead of 22) midbody scalerows, a preocular as wide as a
nasal, incompletely divided nasal, and—though probably of little sig-
nificance—completely hidden eyes.
Description—Snout prominent, trilobate as seen from above,
rounded, without obtuse horizontal edge; rostral half the width of the
head ; nasal swollen, semidivided, the suture extending from the second
labial to the nostril, which is lateral; preocular present, as broad as the
nasal, much broader than either of the small scales which might be
termed an ocular, the lower in contact with the third labial; eyes
hidden; only 3 upper labials. Midbody scalerows 26. Diameter of
body included 40 times in total length, tail broader than long, ending
in an obtuse spine.
Coloration.—About to slough. Above, silvery gray, base of each
scale with a transverse brown spot. Below, yellowish gray, with a
few fine black flecks.
Measurements —Total length 348 (838+5) mm.; diameter at mid-
body 8.5 mm.
NATRIX ANOSCOPUS ANOSCOPUS (Cope)
1861. Tropidonotus anoscopus Corr, Proc. Acad. Nat. Sci. Philadelphia, 1861,
p. 299 [“Cuba” (error, probably Liberia) ].
1863. Tropidonotus ferow Giinruer, Ann. Mag. Nat. Hist., ser. 3, vol. 12, p. 355,
pl. 6, fig. F (Fernando Po).
REPTILES AND AMPHIBIANS FROM LIBERIA—LOVERIDGE 119
229 (U.S.N.M. Nos. 109297-8), Gibi
8 64,2 92 (U.S.N.M. Nos 109585-9), Bendaja
ZOO TS (U.S.N.M. Nos. 109636-8), Harbel
SWE eyul (M.C.Z. Nos. 22505-8), Paiata (G. M. Allen)
Midbody scalerows 23-25; ventrals 138-146; anal divided; sub-
caudals 65-88 ; internasals 2, rarely single; preoculars 1, rarely 2; post-
oculars 2, rarely 1, 8, or 4; suboculars 2-4; temporals 1+2 or 1173;
upper labials 9, rarely 8 or 10; lower labials 9-12. Males may be dis-
tinguished readily by the presence of papillalike rugosities on the
sublinguals. Largest ¢ (M.C.Z. No. 22505) measures 626 (470+
156) mm.; largest 9 (M.C.Z. No. 22506) measures 629 (4734156)
mm.
Coloration: Above, blackish or grayish, uniform or with a series
of transverse crossbars which may be interrupted dorsally, or obso-
lescent as a dorsal bar and vertical lateral stripes. Below, white or
gray, uniform or more usually with the base of each ventral shield
black, rarely (U.S.N.M. No. 109636) with a longitudinal series of
black spots.
The stomach of one snake held a toad (Bufo regularis maculatus)
and remains of a frog (ana sp.), that of another a fish (Hem-
chromis fasciatus). One Bendaja reptile was heavily infested with
anisakine nematodes.
The name ferox, by which this water snake has been known until
now, must be referred to the synonymy of anoscopus.. Angel (1938,
p. 71), after examination of the types of Helicops gendrii Boulenger,
referred them to the synonymy of ferox, for he found they bore a sutural
scar on the posterior portion of their single internasal. This is exactly
the position in two of the present series (U.S.N.M. Nos. 109297, 109586) ,
which have only single internasals. However, Boulenger (1893, p.
241) confused two forms under the name of ferowx in his Catalogue.
Both of these forms, while normally possessing a pair of internasals,
may at times have them fused into a single shield; they can be separated
structurally and geographically as follows:
Midbody scale rows 21-25, normally 23; ventrals 188-148 (23 exam-
ples) ; range: Liberia east to Cameroons and Fernando Po_----~ a. anoscopus
Midbody scale rows 23-27, normally 25; ventrals 146-159 (10 exam-
ples) ; range: Sierra Leone and French Guinea__-_------------~-- a. gendrii
The possibility of retaining ferow as an insular third subspecies with
21 scalerows is rendered impossible by the recording of a snake with
21 rows from Atakpame, Togo, on the mainland.
1Dr, E. R. Dunn informs me that he reached the same conclusion, after examination of
Cope’s type.
120 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
NEUSTEROPHIS VARIEGATUS (Peters)
1861. Mizodon variegatus Peters, Monatsb. Akad. Wiss. Berlin, 1861, p. 358
(Pel, Gold Coast).
6 (U.S.N.M. No. 109058), Gibi
@ (U.S.N.M. No. 109307), Bromley
62 (U.S.N.M. Nos. 109588-4), Bendaja
Midbody scale rows 15; ventrals 124-131; anal divided; subcaudals
75-78; labials 8, the fourth and fifth entering the orbit; preoculars
1-2. Larger ¢ measures 279 (186+93) mm.; larger @ measures
237 from snout to anus, tail truncate.
Bogert (1940, p. 83) advances sound reasons for separation of the
smooth-scaled African “Matrix” under Giinther’s (1858) name of
Neusterophis. I take this opportunity of correcting the misidentifi-
cation of three Paiata, Liberia, snakes reported as Vatrix fuliginoides
by Barbour and Loveridge (1930, p. 772), and my (1936, p. 21) mis-
taken action of synonymizing variegatus with fuliginoides on account
of their occurrence together at Bitye, Ja River, French Cameroons.
The two species are very closely related, practically identical in
markings, yet cannot be regarded as races of one species for their
ranges are largely coextensive, though vwariegatus extends farther
westward to Liberia and French Guinea. This means that fuliqi-
noides should be deleted from the Liberian list.
BOAEDON LINEATUS VIRGATUS (Hallowell)
1854. Coclopeltis virgata HaLLowELL, Proc. Acad. Nat. Sci. Philadelphia, 1854,
p. 98 (Liberia).
6 (U.S.N.M. No. 109592), Bendaja
Midbody scale rows 23; ventrals 223; anal entire; subcaudals 50;
labials 8, the fourth and fifth entering the orbit; preoculars 2; post-
oculars 2; temporals 1+2 (R) and 1+1 (L).
Trinomials are used because this extreme western form differs from
the nominate species only in the reduced number of midbody scale
rows, which, however, is almost constant for Liberia, becoming rarer
as one proceeds eastward and unknown east of the Belgian Congo.
The region of overlap is very extensive.
BOAEDON LINEATUS LINEATUS Duméril and Bibron
1854. Boacdon lineatus Dum*érm and Brpsron, Erpétologie générale, vol. 7,
p. 363 (Gold Coast).
1
6 (U.S.N.M. No. 109673), Mombo
REPTILES AND AMPHIBIANS FROM LIBERIA—LOVERIDGE 121]
Midbody scale rows 27; ventrals 197; anal entire; subcaudals 63;
labials 8, the fourth and fifth entering the orbit; preoculars 2; post-
oculars 2; temporals 1+1.
This house snake so closely resembles the foregoing in color pat-
tern and squamation that there can be no doubt of their close relation-
ship. It constitutes, however, the first recorded occurrence of
lineatus in Liberia.
BOAEDON OLIVACEUS (Duméril)
1856. Holurophis olivaceus A. DumERIn, Rev. Mag. Zool., ser. 2, vol. 8, p. 466
(Gaboon).
6 (U.S.N.M. No. 109590), Bendaja
é (U.S.N.M. No. 109639), Harbel
Midbody scale rows 25-27 ; ventrals 208-218; anal entire; subcaudals
39-40, single; labials 8, the fourth and fifth entering the orbit.
Larger ¢ measures 99 (67+32) mm.
HAPSIDOPHRYS LINEATA Fischer
1856. Hapsidophrys lineatus FiscHer, Abh. Nat. Ver. Hamburg, vol. 3, p. 111,
pl. 2, fig. 5 (Elmine, Gold Coast).
6 (U.S.N.M. No. 109594), Bendaja
Midbody scale rows 15; ventrals 165; anal entire; subcaudals ?
(tail truncate) ; upper labials 8, the fourth and fifth entering the
orbit; preocular 1; postoculars 2.
RHAMNOPHIS AETHIOPISSA AETHIOPISSA Giinther
1862. Rhamnophis aethiopissa GUntuer, Ann. Mag. Nat. Hist., ser. 3, vol. 9,
p. 129, pl. 10 (West Africa).
6 (U.S.N.M. No. 109593), Harbel
Midbody scale rows 16 (for certain); ventrals 165; anal divided;
subcaudals 114*; upper labials 8, fourth and fifth entering the orbit;
preocular 1; postoculars 2, the lower in contact with three upper
labials. Total length 1185* (805+380*) mm., tail tip truncate.
GRAYIA SMYTHIT (Leach)
1818. Coluber Smythii Leacw, in Tuckey, Narrative of an expedition to explore
the river Zaire, App., p. 409 (Embomma, i. e. Boma, Belgian Congo).
1854. Coronella triangularis Hatntowett, Proe. Acad. Nat. Sci. Philadelphia,
1854, p. 100 (Liberia).
2 (U.S.N.M. No. 109582), Pendaja
6 (U.S.N.M. No. 109640), Aarbel
406739—41——2
122 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
Midbody scale rows 17; ventrals 152-161; anal divided; subcaudals
91-99; labials 7-8, fourth or fifth entering the orbit; temporals 2+3,
the lower anterior longer than its distance from the loreal. Larger,
the 2, measures only 312 (228 +84) mm.
BOIGA BLANDINGII (Hallowell)
1844. Dipsas Blandingii Hattowert, Proce. Acad. Nat. Sci. Philadelphia, 1844,
p. 170 (Liberia).
1856. Dipsas valida Fiscurer, Abh. Nat. Ver. Hamburg, vol. 3, p. 87, pl. 3,
fig. 4 (Edina, Grand Bassa County, Liberia).
1856. Dipsas globiceps FiscHmr, ibid., p. 89, pl. 3, fig. 6 (Edina, Grand Bassa
County, Liberia).
é (U.S.N.M. No. 109306), Bromley
Midbody scale rows 21; ventrals 264; anal entire; subcaudals 135;
labials 8, the third, fourth, and fifth entering the orbit. Total length
1,415 (1,085+330) mm.
CROTAPHOPELTIS DUCHESNII GUINEENSIS (Chabanaud)
1920. Leptodira guineensis CHABANAUD, Bull. Com. Etudes Hist. Sci. Afrique
Occ. Francaise, 1920, p. 491 (Dieke, Nzerekore region, French Guinea).
2 (U.S.N.M. No. 109645), Harbel
Midbody scale rows 17; ventrals 223; anal entire; subcaudals 111;
labials 8, third, fourth, and fifth entering the orbit; loreal sharply
distinct, not entering orbit. Total length 733 (547+186) mm. In
its oviducts (March 10-July 17) are 4 eggs, each measuring about
27 by 8 mm.
This species is the genotype of Dipsoglyphophis Barbour and
Amaral, 1927, and I should prefer to recognize this name for the
group of chunky-headed, attenuate, long-tailed, arboreal snakes (as
distinct from the moderate, short-tailed, terrestrial species like C. h.
hotamboeia) rather than force them into the genus Dipsadoboa
(which differs in possessing large vertebrals and single subcaudals)
as has been advocated by Bogert (1940, p. 65). Admittedly they
occupy an intermediate position between C. h. hotamboeia and D.
unicolor, but in head shape C. shrevei conforms to hotamboeia rather
than to its long-tailed relatives. Nor can I agree with the synonymiz-
ing of guineensis with duchesnii Boulenger, from which it differs in
several particulars, so that I should have preferred to let it remain
as a full species until more material is available, but I compromise in
reviving it to subspecific rank.
REPTILES AND AMPHIBIANS FROM LIBERIA—LOVERIDGE 123
MIODON ACANTHIAS (Reinhardt)
1860. Urobelus acanthias REINHARDT, Vidensk. Medd. Kjgbenhayn, 1860, p. 229,
pl. 3 (Guinea).
@ (U.S.N.M. No. 109057), Gibi
Midbody scale rows 15; ventrals 212; anal entire; subcaudals 17;
labials 7, the third and fourth entering the orbit. Total length 533
(505+28) mm. In its oviducts (April 10-16) are about 4 eggs, each
measuring circa 22 by 7 mm.
APARALLACTUS MODESTUS (Giinther)
1859. Elapops modestus GUNTHER, Ann. Mag. Nat. Hist., ser. 3, vol. 4, p. 161,
pl. 4, fig. C (West Africa).
1860. Periaspis plumbeatra Corr, Proc. Acad. Nat. Sci. Philadelphia, 1860,
p. 242 (Liberia).
@ (U.S.N.M. No. 109635), Harbel
Midbody scale rows 15; ventrals 154; anal entire; subcaudals 36;
labials 7, the third and fourth entering the orbit. Total length
465 (407+58) mm.
DENDROASPIS VIRIDIS (Hallowell)
1844. Leptophis viridis Hattowrtt, Proc. Acad. Nat. Sci. Philadelphia, 1844,
p. 172 (Liberia).
1852. Dinophis Hammondii Hattowett, Proc. Acad. Nat. Sci. Philadelphia,
1852, p. 203 (Liberia).
6 (U.S.N.M. No. 109675), Harbel
Midbody scale rows 13; ventrals 220; anal divided; subcaudals
114; labials 8, the fourth entering the orbit; upper temporal sep-
arated from its fellow by three scales. Total length 2,040
(1,520+520) mm.
CAUSUS RHOMBEATUS (Lichtenstein)
1823. Sepedon rhombeatus LIcHTENSTEIN, Verzeichniss der Doubletten des
zoologischen Museums . . . zu Berlin, p. 106 (no locality).
1842. Distichurus Maculatus HAtLOwELL, Journ. Acad. Nat. Sci. Philadelphia,
vol. 8, p. 387, pl. 19 (Liberia).
é (U.S.N.M. No. 109591), Bendaja
4 92 (U.S.N.M. Nos. 109641-4), Harbel
Midbody scale rows 19; ventrals 128-141; anal entire; subcaudals
17-20; labials 6; suboculars 1-2. Total length of 3, 497 (452+ 45)
mm.; of largest 2, 400 (865+35) mm.
BITIS GABONICA (Duméril and Bibron)
1854. Echidna Gabonica Dumérit~ and Brsron, Erpétologie générale, vol. 7, p.
1428, pl. 80b (Gaboon).
$2 (U.S.N.M. Nos. 109683-4), Harbel
124 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
Midbody scale rows 36; ventrals 128-130; anal entire; subcaudals
26-28; labials 14. Both specimens are young.
BITIS NASICORNIS (Shaw)
1802. Coluber Nasicornis SuHaw, Nat. Misc., vol. 3, pl. 94 (interior of Africa)
(from the master of a Guinea vessel).
466,44 22 (U.S.N.M. Nos. 109018, 109676-82), Gibi
6 (U.S.N.M. No. 109308), Bromley
Midbody scale rows 80-38; ventrals 121-129; anal entire; sub-
caudals 15-26; labials 14-17. Largest ¢ (U.S.N.M. No. 109678)
measures 651 (570+81) mm.; largest 9 (U.S.N.M. No. 109680)
measures 955 (870+85) mm.
FROGS AND TOADS
XENOPUS TROPICALIS (Gray)
1864. Silurana tropicalis Gray, Ann. Mag. Nat. Hist., ser. 3, vol. 14, p. 316
(Lagos, Nigeria).
3 (U.S.N.M. Nos. 109571-3), Bendaja
These frogs have the vestigial lower eyelid and, though less well
defined on account of their dry condition, snout and chin beset by
pustules characterizing the species as redefined by Parker (1936a,
p. 157).
BUFO REGULARIS MACULATUS Hallowell
1850. Bufo cinereus HALLOWELL (not Schneider), Proc. Acad. Nat. Sci. Philadel-
phia, 1850, p. 169 (Liberia).
1854. Bufo maculatus HALLOWELL, Proc. Acad. Nat. Sci. Philadelphia, 1854, p. 101
(new name for cinereus, preoccupied).
5292 (U.S.N.M. Nos. 109286-90), Bellyella
1 yng. (U.S.N.M. No. 109305), Bromley
6 yng. (U.S.N.M. Nos. 109554—-6, 109666-8), Bendaja
864,12 (US.N.M. Nos. 109622-30), Harbel
1 yng. (U.S.N.M. No. 11314), Reputa
The series maintains the small size of this western form, the very
largest 6 3 (U.S.N.M. No. 109623, etc.) measuring 53 mm., the largest
gravid 2 (U.S.N.M. No. 109286) measuring 67 mm., the youngest
toad (U.S.N.M. No. 109556) 15mm. One of these toads was recovered
from the stomach of a water snake (WVatrix a. anoscopus).
BUFO CAMERUNENSIS CAMERUNENSIS Parker
1936. Bufo camerunensis camerunensis PARKER, Proc. Zool. Soe. London, 1986,
p. 153 (Oban, Calabar, Nigeria).
@ (U.S.N.M. No. 109285), Bellyella
REPTILES AND AMPHIBIANS FROM LIBERIA—LOVERIDGE 125
This somewhat dried individual has been compared with a paratype
of camerunensis, a species long confused with /atifrons Boulenger.
First recorded from Liberia by Parker (1936c, p.97). Length 76 mm.
HYLAMBATES COCHRANAE, new species
Cotypes.—U.S.N.M. Nos. 109569-70, being anadult ¢ and gravid 9
from Bendaja, Republic of Liberia, collected by William M. Mann,
May 14-27, 1940.
Diagnosis ——Color pattern somewhat resembling that of Kassina
senegalensis from which it differs in the possession of well-developed
digital disks. Intermediate in position between HW. cassinoides
Boulenger of McCarthy Island, Gambia (with topotypes of which
they have been compared), and H. leonardi Boulenger of Fernando
Po (with Liberian and Congo examples of which they have been
compared). It differs from cassinoides in having rather smaller,
rounded (instead of subtriangular) disks, broader habit, larger size,
and in the thighs and concealed surfaces of tibia and foot exhibiting
contrasted black marblings on a white (? red in life) ground. It
differs from leonardi in possessing smaller disks; shorter hind limb,
which reaches axilla instead of eye; smaller size; and pure white
(instead of black) breast and belly, ete.
One wonders if the frogs from French Guinea, referred to Cassina
weali of southeast Africa (!) by Chabanaud (1921, p. 460), might
not in reality represent this undescribed tree frog.
Description.—V omerine teeth in two oblique groups situated some-
what posteriorly between the choanae (poorly developed in the @
cotype). Head broader than long; snout rounded, shorter than the
diameter of the eye, interorbital space broader than an upper eyelid;
tympanum rather more than half the diameter of the eye; fingers
long, free, with small rounded disks which are half the size of the
tympanum; toes half-webbed, their disks smaller than those of the
fingers; inner metatarsal tubercle small, rounded, feebly prominent,
tibiotarsal articulation of the adpressed hind limb reaches only to the
axilla. Skin smooth (or rugosely warty due to immersion in strong
alcoho!) above; granular on belly and under the thighs. Length of
6 36 mm.; length of 2 mm.
Color in alcohol.—Above, blackish, everywhere with numerous
large, oval, light-edged dark spots (as in maculatus) ; groin, thighs,
posterior side of tibia, and upper part of foot, marbled with white
(? red in life). Below, throat of 3 black, that of ¢ white with dusky
freckles around its labial border; breast and belly of both sexes white
with brown vermiculations along the flanks; limbs white (? red)
marbled with black; soles chiefly brown mottled with white.
126 _ PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
iow 8'N ow
Gbangé
Degain%
iow
Ficure 34.—Liseria. This map was modified by Dr. Leonard P. Schultz from a map of I
made to the Geographical Institute of Harv.
REPTILES AND AMPHIBIANS FROM LIBERIA—LOVERIDGE 127
ited by the Institute of Geographical Exploration, Harvard University. Acknowledgment is
sity for permission to trace the river systems.
128 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 81
HYLAMBATES LEONARDI Boulenger
1906. Hylambates leonardi BouLencer, Ann. Mus. Stor. Nat. Genova, ser. 3, vol. 2,
p. 167, pl. 2, fig. 8 (Punta Frailes, Fernando Po, and N’Djole, French
Congo).
66 (U.S.N.M. Nos. 109567-8), Bendaja
In view of this record involving a westward extension of the range
of nearly a thousand miles, one might have supposed that they would
represent a western race for they differ in several details from the
description. Their vomerine teeth are between (not behind the level
of) the choanae, though situated somewhat posteriorly; the head is
distinctly (not slightly) broader than long; the snout is two-thirds
(not equal to) the diameter of the eye; the interorbital space is broader
than (not as broad as) an upper eyelid; the tympanum is two-thirds
(not two-fifths) the diameter of the eye.
In all these respects, however, they agree with a series (M.C.Z. Nos.
21681-8) of females and young from Djamba, Belgian Congo (det. de
Witte), and all with the striking color pattern as figured by
Boulenger. There is considerable variation in the amount of white
(? red, ? yellow) in the groin and elsewhere. These males exhibit the
black vocal sacs flanking the central gular disk which is common to
males of their allies of the genus Xassina. Boulenger gave 54 mm. as
the length, probably of his cotype 2 and not of the ¢ for the Bendaja
é é measure 45-47 mm.
LEPTOPELIS VIRIDIS (Ginther)
1868. Hylambates viridis GinTHER, Proc. Zool. Soc. London, 1868, p. 487 (West
Africa).
1929. Leptopelis liberiensis ABL, Sitz. Ges. Naturf. Freunde Berlin, 1929, p. 194
(Liberia).
24,4 22 (U.S.N.M. Nos. 109530, 109557-61), Bendaja
6 @ (U.S.N.M. Nos. 109620-1), Harbel
Parker (1936c, p. 95) has cleared up the confusion that has long
centered about West African frogs of this group and referred hyloides
Boulenger, nanus Ahl, and togoensis Ah] to the synonymy. To these
I would now add Uiberiensts Ahl, described as having a very faint rudi-
ment of web between the fingers but in all other respects agreeing with
viridis, which most authors agree to consider as having “fingers free.”
Males are distinguishable by their dark throats. Length of ¢ ¢ 30-34
mm., of ¢ 2 36-46 mm.
LEPTOPELIS BEQUAERTI, new species
Correction—In 1930, Barbour and Loveridge referred certain Li-
berian frogs to tessmanni Nieden (of Makomo, Spanish Guinea). In
REPTILES AND AMPHIBIANS FROM LIBERIA—LOVERIDGE 129
the absence of topotypic material of Nieden’s frog, the Mount Coffee
(p. 785) specimens may still be considered to represent tessmanni, but
the frogs from Gbanga and Du River (p. 782) that I thought to be
young tessmanni must be considered specifically distinct on account of
the less developed webbing on both hand and foot. I take pleasure in
naming the new species after its collector, Dr. J. Bequaert, who has
done so much to advance our knowledge of African zoology in many
fields.
Type—M.C.Z. No. 12000, a female from Gbanga, Republic of
Liberia, collected by Joseph Bequaert, September 1926.
Paratypes—Young ? (M.C.Z. No. 12001), Gbanga, Liberia (J.
Bequaert); ¢ and juv. (M.C.Z. Nos. 12002-3), Plantation No. 3, Du
River, Liberia (G. M. Allen); @ (U.S.N.M. No. 109051), Gibi, Liberia
(W. M. Mann).
Diagnosis.—Digits with a mere rudiment of web; toes with a single
joint free of web on the first, second, and third, two joints free on the
fourth, a single or only half a joint free on the fifth.
In contrast the Mount Coffee frog (M.C.Z. No. 15939) has only the
first finger with a rudiment of web, the second has one joint free, the
third two joints, the fourth one and a half joints; of its toes only the
first has a single joint free, the second, third and fifth are webbed to
the disks on at least one side, the fourth has one and a half (right) to
2 (left) joints free of web. It isa ¢ of larger size, viz, 50 mm.
Description—Vomerine teeth in two small groups between the
choanae. Head as broad (or slightly broader than) long; snout
roundish, half to two-thirds the diameter of the eye; interorbital space
slightly broader than (or as broad as) an upper eyelid; tympanum
two-thirds the diameter of the eye; fingers rather long with a mere
rudiment of web, their disks as large as the tympanum; toes two-
thirds webbed, one joint free of web on the first, second, and third
toes, two joints free on fourth, half (or one) joint free on fifth, the
disks a little smaller than those on the fingers; inner metatarsal tuber-
cle large, oval, strongly compressed; the tibiotarsal articulation of the
adpressed hind limb reaches the nostril (or eye). Skin of dorsum
shagreened and with small scattered warts; on the throat, belly, and
under the thighs, granular; males with a callous pad on the breast
in the region of the axilla,
Coloration—Above, pale brown, a dark, triangular, interorbital
marking, its apex directed posteriorly and often confluent with a more
or less distinct hourglass pattern on the back; a dark line from the
eye passes over the tympanum to the base of the forearm (and may be
continued on the flank as a series of dashes); flanks marbled with
brown; forearm, thighs, and to some extent the foot, crossbarred with
dark brown; from disk of outer finger to elbow, from disk of outer toe
130 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
to heel, and above anus a narrow white line. Below, creamy white
sparsely mottled with brown (or uniform) ; limbs brownish merging
into purplish brown on palms and soles.
Measurements.—Length from snout to anus of type 2, 28 mm.; of
paratype 2 2 from Gbanga and Gibi, 29 and 33 mm., respectively ;
of paratype 3, 29 mm.; of a juvenile, with rudiment of tail still
visible, 15 mm.
MEGALIXALUS FULVOVITTATUS (Cope)
1860. Hyperolius fulvovittatus Corr, Proe. Acad. Nat. Sci. Philadelphia, 1860,
p. 517 (Liberia).
1876. Hyperolius vittiger Perrrs, Monatsb. Akad. Wiss. Berlin, 1876, p. 122
(Liberia).
17 6,5 22 (U.S.N.M. Nos. 109534-53, 109664-5), Bendaja
222 (U.S.N.M. No. 109618-9), Harbel
2 (U.S.N.M. No. 109672), Reputa
6 (U.S.N.M. No. 113818), Cape Mount
All possess the characteristic chocolate-brown dorsal stripes.
Length of ¢ 6, 23-27 mm., average 24 mm.; length of ? 92, 24
27 mm., average 26 mm., being somewhat less than that of the
enormous series from Ganta, Liberia, reported on elsewhere (Love-
ridge, 1938, p. 66).
MEGALIXALUS PLATYCEPS (Boulenger)
1900. Rappia platyceps BouteNceR, Proc. Zool. Soe. London, 1900, p. 444, pl. 27,
fig. 4 (Benito River, French Congo).
6 (U.S.N.M. No. 109533), Bendaja
é6 (U.S.N.M. Nos. 109616-7), Harbel
A broad vertebral band or hourglass pattern, dorsal spinosities,
and vertical pupil present in all. Length of ¢ 4, 26-29 mm. See
remarks in Loveridge (1938, p. 66).
HYPEROLIUS CONCOLOR (Hallowell)
1844. Ixalus concolor Hattowertt, Proc. Acad. Nat. Sci. Philadelphia, 1844,
p. 60 (Liberia).
3 99 (U.S.N.M. Nos. 109531-2, 109655), Bendaja
2 (U.S.N.M. No. 109669), Reputa
Q (U.S.N.M. No. 110447), Harbel
Fourth and first toe with one phalange free of web, third scarcely
(riggenbachi) or fully (concolor) webbed, second and fifth fully
webbed to disks. Three subadult frogs (U.S.N.M. Nos. 109531-2,
109655) are typically riggenbachi Nieden in their dorsal markings,
but Mertens (1938, p. 27) considers this to be the juvenile stage of
concolor, stating that a riggenbachi, which he captured in the Cam-
eroons, transformed in his vivarium to a typical, uniform concolor.
REPTILES AND AMPHIBIANS FROM LIBERIA—LOVERIDGE 131
One of the two adult females still retains its dorsal coloring of vivid
green merging into yellow on the flanks, upper arm, and thighs.
Below, uniform white. The subadults measure 23-28 mm., adult 2 9,
38-42 mm. Both the latter are gravid, having been taken between
March 10-July 17 and June 21-26, respectively.
HYPEROLIUS PLEUROTAENIUS (Boulenger)
1906. Rappia pleurotaenia BouteNncrr, Ann. Mag. Nat. Hist., ser. 7, vol. 17,
p. 322 (Zima, French Cameroons).
2 (U.S.N.M. No. 109040), Gibi
Q (U.S.N.M. No. 109670), Reputa
Fourth toe with one phalange free of web, remaining toes webbed
to their disks; tibiotarsal articulation of the adpressed hind limb
reaches posterior border of eye. Coloration precisely like that shown
on colored plate in Barbour and Loveridge (1930, pl. 465, fig. 4).
Length of 2 ¢, 29-37 mm.
HYPEROLIUS PICTURATUS Peters
1875. Hyperolius picturatus Perers, Monatsb. Akad. Wiss. Berlin, 1875, p. 206,
pl. 2, fig. 2 (Boutry, Ashanti, Gold Coast).
3 22 (U.S.N.M. Nos. 109309, 109653, 109659), Bendaja
222 (U.S.N.M. Nos. 109609, 110448), Harbel
Fourth and fifth toe with one phalange free of web, second, third,
and fifth webbed almost, or entirely, to their disks; tibiotarsal articula-
tion of the adpressed hind limb reaches to between eye and nostril.
Above, pale gray to chocolate-brown, uniform, or with a few black
flecks; upper lip and flank with characteristic dark, or black, speckling,
marbling or vermiculation. Length of 2? 2, 27-30 mm. ALI five are
gravid, having been taken between May 14-27 and March 10—July
17, respectively.
HYPEROLIUS OCELLATUS Giinther
1858. Hyperolius ocellatus GUNTHER, Catalogue of the Batrachia Salientia in the
collection of the British Museum, p. 88, pl. 7, fig. B (Fernando Po and
Angola).
38 $46,166 92 (U.S.N.M. Nos. 109310-453, 109462, 109518-29, 109654,
2 (U.S.N.M. No. 110487), Bendaja
Fourth toe with one phalange free of web, remaining toes webbed to
their disks; tibiotarsal articulation of the adpressed hind limb
reaches the eye or nostril. Coloration of the largest and smallest
frogs is as follows: Above, pinkish white, minutely speckled with
brown dots, a few large brown blotches (formed of a concentration
of the smaller dots) on back and limbs; a brown canthal streak from
nostril to eye. Below, white. Two 23-mm. frogs in the Harbel series
are not typical but are so fresh as to have retained certain fugitive
182 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
colors. Above, pale greenish yellow minutely speckled with brown
on back and limbs; a brown canthal streak is overlaid by a blood-red
band, which continues on from eye to groin as an undulating line;
hands and feet blood red. Below, transparently white. Length of
9 9,2226mm. The largest is gravid, having been taken at Harbel
between March 10 and July 17.
HYPEROLIUS FUSCIVENTRIS Peters
1876. Hyperolius fusciventris PeTers, Monatsb. Akad. Wiss. Berlin, 1876, p. 122
(Liberia).
2 (U.S.N.M. No. 109041), Gibi
3 64,166 22 (U.S.N.M. Nos. 109310453, 109462, 109518-29, 109654,
109656-8, 110438-46), Bendaja
24 292 (U.S.N.M. Nos. 109595-607, 109646-9, 110451~7),
Harbel
2 (U.S.N.M. No. 109671), Reputa
Fourth and first toe with one phalange free of web, second and third
with one-half or one phalange free, fifth webbed to disk; tibiotarsal
articulation of the adpressed hind limb reaches the eye or nostril.
Coloration, ? 2 : Above, blue-gray (green in life), an irregular, un-
dulating, often broken, black line extends from below commissure of
mouth over forearm to groin, sometimes continued over thigh to meet
its fellow below anus, similar wavy lines present on anterior and
posterior aspect of fore limb, tibia, and foot. Below, pale to lead
gray, almost black. A few individuals (U.S.N.M. Nos. 109380,
109394, 109404, 109518-23, 109605) lack the lateral line in varying de-
grees and may be classed as underpigmented and overpigmented as
they vary from very pale gray to dark plumbeous above, and all are
paler below, the pallid specimens being actually white. One might
suppose that they were young concolor but for the fact that most of
them are gravid and usually carry, in the shape of scattered specks,
some traces of the whereabouts of the typical markings. The ¢ ¢
differ slightly. Above, pale gray; a black canthal stripe present or
absent; a few conspicuous black spots on flanks; a light dorsolateral
line (as in ademetz?, which see) from posterior border of eye towards
groin, just discernible in one frog. Below, white. Length of ¢ ¢,
21-22 mm.; of 50 2 9 , 22-28 mm., average 25 mm.
HYPEROLIUS ? ADEMETZI Ahl
1931. Hyperolius ademetzi ABL, Mitt. Zool. Mus. Berlin, vol. 17, p. 37 (Bamenda,
British Cameroons).
50 66,18 22 (U.S.N.M. Nos. 10945461, 109463-517, 109660-3, 11315),
Bendaja
9 $4 (U.S.N.M. Nos. 109610-5, 109650-1, 110458), Harbel
Fourth and first toe with one phalange free of web, second and
REPTILES AND AMPHIBIANS FROM LIBERIA—LOVERIDGE 133
third with one-half a phalange free, or webbed to disk like fifth;
tibiotarsal articulation of the adpressed hind limb reaches the eye or
nostril. A light dorsolateral line almost always present. Every
specimen has a more or less conspicuous, dark-edged, silvery, naso-
lateral stripe; the density of pigmentation on the dorsum varies con-
siderably, reaching its maximum in U. S. N. M. No. 109469, in which
even the gular disk and throat are stippled, though as colorless as
the belly in most of the series. Seven males (U.S.N.M. Nos. 109454,
109456, 109459, 109461, 109481, 109610, 109612), though indistinguish-
able as to color and pattern, may be males of some other dimorphic
and slightly larger species, possibly picturatus, for they measure
24-29 mm., average 26 mm. Length of 50 6 3, 19-23 mm., average
22 mm., length of 18 presumed ¢ @ (assumed to be so as without
gular disks, but certainly young, and possibly including some young
males among them) 16-23 mm., average 19 mm.
These frogs are conspecific with the 143 ¢ ¢ and 5 @ 9 previously
reported upon, which I (1988, p. 69) treated in the same way for
reasons stated at that time. In view of the preponderance of male
ademetzi in both collections, paralleled by the predominance of female
fusciventris, one might be tempted to assume that we were dealing
with a single species exhibiting sexual dichromatism. A careful ex-
amination of the earlier material, however, lends no weight to such
an assumption, and one must conclude that the ademetzi males are
assembling to summon their females at a time when the fusciventris
females are ovulating. The majority of fusciventris females collected
by Dr. Mann are distended with ova, but none of the ademetzi
females appears gravid.
HYPEROLIUS ? FESTIVUS Barbour and Loveridge
1927. Hyperolius festivus Barsour and Lovertpce, Proce. New England Zool.
Club, vol. 10, p. 17 (Firestone Plantation No. 3, Du River, Republic of
Liberia).
? 6 (U.S.N.M. No. 109052), Gibi
Above, brown, but lips and flanks white, instead of dark; the
absence of the dark hourglass pattern on the dorsum is of no im-
portance, as it is absent in a @ paratype of festivus. This rather
dried specimen has no gular disk but a baggy singing pouch. Length,
26 mm. It seems probable that festivus, as well as bawmanni Ahl,
of which we have a Togo cotype, will probably have to be synonymized
with acutirostris Peters, of Cameroons, of which we have no typical
material.
SYNOPSIS OF THE SPECIES OF RANA IN LIBERIA
In connection with this work I came across a specimen (M.C.Z. No.
24461) of Rana longirostris Peters that constitutes the first Liberian
134 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
record of this Gold Coast (type locality, Keta) species of which
aequiplicata Werner, of the Cameroons and Congo, is a synonym
according to Nieden, who made direct comparison of the types.
The specimen comes from the Firestone Plantation No. 3, on the Du
River, Liberia, where it occurs together with A. maccarthyensis
(M.C.Z. Nos. 24462-38), R. o. gribinguiensis (M.C.Z. Nos. 24458-60),
R. o. oxyrhynchus (M.C.Z. Nos. 24455-7), and &. m. mascareniensis
(M.C.Z. Nos. 11927-3831), the latter having been erroneously recorded
under the name b7bronii in 1930.
In view of the remarkable similarity of all these frogs and the
consequent difficulty of distinguishing them, it seemed advisable to
draw up the following key after a careful examination of Dr. Mann’s
material together with that in the Museum of Comparative Zodlogy.
1. A conspicuous transverse fold connects posterior edges of upper
eyelids}: toes twebbed)tocipse Nabi ter OD US te eee ee ee er 2
No transverse fold across crown of head; habit more or less
Slender sik ee UE ote Ne MiB EUs anes Bee Be EE Lae DG a 0 oe 3
2. Vomerine teeth in two oblique rows, anteriorly touching inner
posterior edge of choanae, posteriorly converging; snout acumi-
nate, as long as, or almost as long as, orbital diameter; tym-
panum sharply distinct, large, its diameter almost that of
orbit; tips of toes not dilated, at most thickened_______________ occipitalis
Vomerine teeth (absent in young) in two round groups between,
but posterior to an imaginary line connecting hind edges of,
choanae; snout rounded, once and a half as long as orbital di-
ameter ; tympanum indistinct, small, its diameter about a third
that of orbit; tips of toes dilated into distinct, though small,
GiSkS 2234 TANS BNE) ES a A es Se oe Le ce. alleni
3. Vomerine teeth in two oblique rows between, though not in contact
with, choanae; tips of fingers and toes dilated into distinct disks.
Fourth toe with only 1 phalange free of web (or which may
be continued up it as a narrow seam to disk), remaining
toes webbed to base of their disks; tibiotarsal articula-
tion of adpressed hind limb reaches eye or just beyond
end of snout; vocal sacs of ¢ internal, but a glandular
Swelling present at base of forearm in ¢ @____________ a. albolabris
Vomerine teeth in two rows projecting inward from anterior
edges of choanae; tips of fingers and toes simple, not dilated______________ 4
4. An inner and an outer metatarsal tubercle, latter connected by
a series of minor tubercles with first subarticular tubercle of
fourth toe.
Fourth toe with 2 (rarely 3) phalanges free of web, first
toe with 1% (rarely 1 or 2), second with 1 (rarely 114),
third with 1 (rarely 2), fifth with %4 (rarely 1 or 114)
phalanges free of web; tibiotarsal articulation of ad-
pressed hind limb reaches nostril or well beyond end of
snout; vocal sac of ¢ external, its aperture extending
posteriorly toward lower insertion of forearm______ maccarthyensis
An inner metatarsal tubercle only, no minor tubercles on basal
phalange of ‘fourth toe lo te) SEA SA ae a Se 5
REPTILES AND AMPHIBIANS FROM LIBERIA—LOVERIDGE 135
5. Fourth toe with only 1 phalange free of web (though sometimes
second represented only by a narrow seam in o. gribinguiensis),
FEE TEU VV FIED CON EO eLearn ae rere a
Hourth'toe with:2 or more phalanges freeiof web-==-- =. = ee ik
6. First, second, and third toes with 4% a phalange free of web;
tibiotarsal articulation of adpressed hind limb reaches end of
snout or far beyond adult, 29" 50-58 mime === - > longirostris
First, second, and third toes webbed to tips; tibiotarsal articula-
tion of adpressed hind limb reaches well beyond or far beyond
end of snout; inhabits rain forest; size larger, adult 9 2 58-74
STON ew Ra os Sn ee ee o. gribinguiensis
7. Fifth toe webbed to tip; vocal sac of ¢ external, its aperture ex-
tending posteriorly toward lower insertion of forearm.
Fourth toe with 2 phalanges free of web; first, second, and
third toes with 1 phalange free of web; tibiotarsal articu-
lation of adpressed hind limb reaches nostril or just be-
yond end of snout; adult 2 9 53-55 mm-____-________ o. oxyrhynchus
Fifth toe with 1 or more phalanges free of web______--~--_-__=__-_________ 8
8. Fourth toe with 2% phalanges free of web, first, second, and
third toes with 1 (or rather more than 1) phalange free, fifth
with only 1 free; tibiotarsal articulation of adpressed hind
limb reaches nosiril or just beyond end of snout; vocal sac of
é external, its aperture extending posteriorly toward upper
INSertiOn, Of fOreaArM = See sees oo nen eee m. mascareniensis
Fourth toe with 3 phalanges free of web, first, third, and fifth
with 2, second with 1144 phalanges free of web; tibiotarsal ar-
ticulation of adpressed hind limb reaches well beyond or far
beyond end of snout; vocal sac of ¢ external, its aperture ex-
tending posteriorly toward lower insertion of forearm____________. bibronii
RANA OCCIPITALIS Giinther
1858. Rana occipitalis GUNTHER, Catalogue of the Batrachia Salientia in the col-
lection of the British Museum, p. 130, pl. 11 (Gambia) (restricted).
6,2 29 (U.S.N.M. Nos. 109299-301), Bromley
3 646,2 92 (U.S.N.M. Nos. 109574-8), Bendaja
Characters as in foregoing synopsis. Length of ¢ ¢, 80-93 mm.;
of 2 9, 82-94 mm.
RANA CRASSIPES ALLENI (Barbour and Loveridge)
1927. Pseudorenopus alleni Barsour and Lovermer, Proc. New England Zodl.
Club, vol. 10, p. 14 (Firestone Plantation No. 3, Du River, Liberia).
Yng. and ¢ (U.S.N.M. Nos. 109050, 11313), Gibi
Characters as in foregoing synopsis. Length of ¢,65mm. Parker
(1931, p. 493) has accidentally reversed the character of snout length
in relation to that of occipitalis. It seems best to regard alleni as the
western race of crassipes as suggested by Parker.
136 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 01
RANA ALBOLABRIS ALBOLABRIS Hallowell
1856. Rana albolabris HALLowett, Proc. Acad. Nat. Sci. Philadelphia, 1856,
p. 153 (West Africa).
244,222 (U.S.N.M. Nos. 109053-6), Gibi
@ (U.S.N.M. No. 109304), Bromley
Characters as in foregoing synopsis. Length of ¢ ¢,42-45 mm; of
9 9,38-51mm. As &. a. acutirostris Parker (1936b, p. 141) is pre-
occupied by 2. acutirostris Fatio (1872), I take pleasure in renaming
the former, of which we have a paratype, parkerzana, after its describer.
RANA MACCARTHYENSIS Andersson
1937. Rana maccarthyensis ANDERSSON, Arkiv Zool., vol. 29A, No. 16, p. 9, figs. 3-4
(Maccarthy Island, Gambia).
2 (U.S.N.M. No. 109038), Gibi
3646,3 22 (US.N.M. Nos. 109265-70), Bellyella
Characters as in foregoing synopsis. Itisa ? and3 é 6 (U.S.N.M.
Nos. 109267-70) that have such long hind limbs as to necessitate ex-
panding the description in this respect. Length of 3 ¢,4243 mm.; of
9 9, 51-64 mm.
RANA OXYRHYNCHUS OXYRHYNCHUS Smith
1849. Rana oxyrhynchus A. SMITH, Illustrations of the zoology of South Africa,
Rept., pl. 77, figs. 2, 2a—c (Kaffirland and the region of Port Natal, South
Africa).
é (U.S. N. M. No. 109039), Gibi
Q (U.S.N.M. No. 109302), Bromley
juv. (U.S.N.M. No. 109652), Bendaja
Characters as in foregoing synopsis, except that the 23-mm. juvenile
has rather more extensive webbing and should perhaps be referred to
fi. 0, gribinguiensis Angel, which occurs in the rain-forest areas of
Liberia. Length of 3,41 mm.; of ?, 53 mm.
RANA MASCARENIENSIS MASCARENIENSIS Duméril and Bibron
1841. Rana mascareniensis DuMERIL and Brsron, Erpétologie générale, vol. 8,
p. 350 (Madagascar; Mauritius; Seychelles).
466,1 2 (U.S.N.M. Nos. 109033-7), Gibi
9 66,4 92 (U.S.N.M. Nos. 109271-83), Bellyella
2 (U.S.N.M. No. 109303), Bromley
6 (U.S.N.M. No. 109579), Bendaja
Characters as in foregoing synopsis. Length of adult 3 6, 46-55
mm.;ofadult 9 9,54-64mm. While possessing the short hind limbs
of the typical form, the Bellyella frogs reach the large size of the rain-
forest race venusta Werner.
REPTILES AND AMPHIBIANS FROM LIBERIA—LOVERIDGE 137
ARTHROLEPTIS POECILONOTUS Peters
1863. Arthroleptis poecilonotus Prrers, Monatsb. Akad. Wiss. Berlin, 1863,
p. 446 (Boutry, Ashanti, Gold Coast).
(U.S.N.M. Nos. 109048, 109049), Gibi
(U.S.N.M. No. 109284), Bromley
(U.S.N.M. Nos. 110459-60), Harbel
(U.S.N.M. No. 110461), Bellyella
(U.S.N.M. No. 11821), Reputa
(U.S.N.M. No. 111320), Degain
A single metatarsal tubercle; tibiotarsal articulation of the ad-
pressed hind limb reaches to between eye and nostril in all. Length
of adults, 25-27 mm.; of juveniles, 12-16 mm.
Bee be bo
he
ARTHROLEPTIS CALCARATUS (Peters)
1863. Hemimantis calcaratus PrerErs, Monatsb. Akad. Wiss. Berlin, 1863, p.
452 (Boutry, Ashanti, Gold Coast).
Juv. (U.S.N.M. No. 111822), Gibi
Two metatarsal and a tarsal tubercle; tibiotarsal articulation of
the adpressed hind limb reaches nostril; upper eyelid with a small
wart in lieu of the elongate tubercle characteristic of the adult, with
whose coloring it is in fairly close agreement though the spotting
on throat and breast is even more pronounced. Length of juv.,
12 mm.
ARTHROLEPTIS WERNERI Nieden
1910. Arthroleptis werneri Nrzpen, Arch. Naturg., vol. 76, pt. 1, p. 242 (Banjo
district and Bamenda, British Cameroons).
9 (U.S.N.M. No. 111319), Gibi
Two metatarsal and a tarsal tubercle, of which the inner is equi-
distant from the outer as from the tarsal tubercle; tibiotarsal articula-
tion of the adpressed hind limb reaches the posterior border of the
eye; upper eyelid warty; snout slightly longer than the orbit. Throat
and lower flanks finely vermiculate. Length of 2,20 mm. Gravid
when taken April 10-16.
As stated by Parker (1936c, p. 93) the identification of Liberian
frogs with werneri should be regarded as tentative until direct com-
parison has been made with Cameroons material.
PHRYNOBATRACHUS NATALENSIS (Smith)
1849. Stenorhynchus natalensis A. SmirH, Dlustrations of the zoology of South
Africa, Rept., App., p. 24 (Natal, South Africa).
Q (U.S.N.M. No. 109042), Gibi
Three phalanges of the fourth toe free of web, first and second
with 1, third and fifth with 2 phalanges free; tibiotarsal articulation
138 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
of the adpressed hind limb reaches the eye. Length of ¢, 36 mm.
This frog, which constitutes the first record of the species from
Liberia, has been compared carefully with specimens from the Natal
border; it appears to be specifically identical with the juvenile
(M.C.Z. No. 11984) from Suahkoko, Liberia, referred to francisci
Boulenger by Barbour and Loveridge (1930, p. 779).
PHRYNOBATRACHUS LIBERIENSIS Barbour and Loveridge
1927. Phrynobatrachus liberiensis BArBour and Loverincr, Proc. New England
Zool. Club, vol. 10, p. 14 (Gbanga, Liberia).
Hgr. (U.S.N.M. No. 111316), Degain
Three phalanges of the fourth toe free of web, first and second
toe narrowly webbed to the disk on one side only, third and fifth with
2 phalanges free; tibiotarsal articulation of the adpressed hind limb
reaches just beyond end of snout. Length of hgr., 24 mm.
PHRYNOBATRACHUS PLICATUS (Giinther)
1858. Hyperolius plicatus GUNTHER, Catalogue of the Batrachia Salientia in the
collection of the British Museum, p. 88, pl. 7, fig. C (Coast of Guinea).
Juv. (U.S.N.M. No. 111317), Mombo
Two phalanges of the fourth toe free of web, third phalange with
a narrow margin only; remaining toes webbed to their disks though
second and third toes only narrowly on one side; tibiotarsal articula-
tion of the adpressed hind limb reaches well beyond tip of snout;
characteristic dorsal glandular folds present. Length of juv., 18 mm.
PHRYNOBATRACHUS OGOENSIS BRONGERSMAI Parker
1936. Phrynobatrachus brongersmai ParKeR, Zool. Meded., vol. 19, p. 90
(Grand Cape Mount, Liberia).
2 (U.S.N.M. Nos. 109048, 110462), Gibi
@ (U.S.N.M. No. 109562), Bendaja
6 (U.S.N.M. No. 110463), Reputa
Two phalanges of the fourth toe free of web, first and second with
half or 1, third and fifth with 1 phalange free; tibiotarsal articula-
tion of the adpressed hind limb reached the eye (in gravid ¢) or
beyond end of snout (in three ? ¢ 3). Length of ¢ 16-17 mm.;
of 2,26 mm. The latter gravid when taken between May 14-97.
The ¢ has a vocal sac, though this is one of the three characters
used by Parker to distinguish the Liberian frog from the slightly
smaller ogoensis Boulenger, to which Barbour and Loveridge (1930,
p. 780) referred certain Liberian frogs. The latter are certainly
conspecific with the present material.
REPTILES AND AMPHIBIANS FROM LIBERIA—LOVERIDGE 139
PHRYNOBATRACHUS LATIFRONS Ahl
1924, Phrynobatrachus latifrons Anu, Zool. Anz., vol. 60, p. 272 (Dodo, French
Cameroons).
3 (U.S.N.M. Nos. 109045-7), Gibi
4 (U.S.N.M. Nos. 109563-6), Bendaja
Two phalanges of the fourth toe free of web, remaining toes webbed
to their disks at least on one side, but less fully than in allené for the
web is deeply incised between digits; tibiotarsal articulation of the
adpressed hind limb reaches the eye or nostril. Lengths 22-25 mm.
Females from both localities gravid when taken between April 10-16
and May 14-27 respectively.
PHRYNOBATRACHUS ALLENI Parker
1936. Phrynobatrachus alleni Parker, Zool. Meded., vol. 19, p. 91 (Firestone
Plantation No. 3, Du River, Liberia).
1 (U.S.N.M. No. 109044), Gibi
Two phalanges of the fourth toe free of web, remaining toes fully
webbed to their disks, at least on one side; tibiotarsal articulation
of the adpressed hind limb reaches end of snout. Length, 28 mm.
LITERATURE CITED
ANGEL, FERNAND.
1933. Les serpents de l'Afrique Occidentale Francaise, 246 pp., 83 figs.
Paris.
Barpour, THOMAS, and LOVERIDGE, ARTHUR.
1930. Reptiles and amphibians from Liberia. In Strong, R.: Report of
the Harvard-African Expedition upon the African Republic of
Liberia and the Belgian Congo, vol. 2, pp. 769-786, 2 pls.
Cambridge, Mass.
Bocert, CHARLES MITCHILL.
1940. Herpetological results of the Vernay Angola Expedition. Bull. Amer.
Mus. Nat. Hist., vol. 77, art. 1, pp. 1-107, 18 figs., 1 pl.
BOULENGER, GEORGE ALBERT.
1893. Catalogue of snakes in the British Museum (Natural History), vol.
1, xiii-++-448 pp., 26 figs., 28 pls. London.
1920. Monograph of the Lacertidae, vol. 1, x-+352 pp. London.
CHABANAUD, PAUL.
1921. Contribution & V’étude de la faune herpétologique de Jl Afrique
Occidentale. Deuxieme note. Bull. Com. Etudes Hist. Sci. Afrique
Occidentale Frangaise, pp. 445-472, map, 4 pls.
LOVERIDGE, ARTHUR.
1936. African reptiles and amphibians in Field Museum of Natural History.
Field Mus. Nat. Hist. Zool. Ser., vol. 22, pp. 1-111.
1938. On a collection of reptiles and amphibians from Liberia. Proc. New
England Zo6l. Club, vol. 17, pp. 49-74.
MERTENS, ROBERT.
1938. Herpetologische Hrgebnisse einer Reise nach Kamerun. Abh. Senck.
Naturf. Ges., No. 442, pp. 1-52, 10 pls.
PARKER, HAMPTON WILDMAN.
1931. Some new and rare frogs from West Africa. Ann. Mag. Nat. Hist.,
ser. 10, vol. 7, pp. 492-498. .
1936a. The amphibians of the Mamfe Division, Cameroons: (1) Zoogeography
and systematics. Proc. Zool. Soe. London, 1986, pp. 135-163,
8 figs., 1 pl.
1936b. Dr. Karl Jordan’s expedition to Southwest Africa and Angola.
Herpetological collections. Nov. Zool., vol. 40, pp. 115-146, 2 figs.
1936e. Amphibians from Liberia and the Gold Coast. Zool. Meded., vol. 19,
pp. 87-102.
140
U.S. GOVERNMENT PRINTING OFFICE: 1941
PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM
‘ a by the
issued /
SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
Vol. 91 Washington: 1941 No. 3129
NOTES ON SOME CRAYFISHES FROM ALABAMA CAVES,
WITH THE DESCRIPTION OF A NEW SPECIES AND A
NEW SUBSPECIES
By Renpett Ryoapes
From Dr. Alvin R. Cahn, formerly chief of the Biological Re-
adjustment Division of the Tennessee Valley Authority, I received a
small collection of crayfishes that he had collected in Shelta Cavern
and Belgreen Cave, in northern Alabama. Those from Shelta Cavern
had been tentatively determined as Cambarus pellucidus, but in order
to establish their status definitely it was necessary to secure more
material. Early the next year I obtained a male, form I, from this
same cavern and later additional material from Leslie Hubricht, of
the Missouri Botanical Garden. With his aid I have been enabled
to study a complete series of this particular crayfish, which is here
described as a new subspecies of Cambarus pellucidus Tellkampf
(1844). The crayfishes from Belgreen Cave are described as a new
species of Cambarus. This particular species is interesting because
it shows affinities to both cave and surface forms.
The types and allotypes of the new forms have been deposited in
the United States National Museum; paratypes are in the Alabama
Museum of Natural History, the Academy of Natural Sciences of
Philadelphia, the collection of Leslie Hubricht, and my own
collection.
I am indebted to Dr. Cahn and Mr. Hubricht for the bulk of the
material reported on in this paper. Grateful acknowledgment is
also made to Dr. Walter B. Jones, director of the Alabama Depart-
408589—41 141
142 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 91
ment of Conservation and director of the Alabama Museum of Nat-
ural History, who has generously provided me with material from
several caves in that State. Dr. Allan F. Archer, director of re-
search, Alabama Department of Conservation, has assisted both in
collecting the material and the data. I wish to express my thanks
to Dr. A. H. Wiebe, chief of the Biological Readjustment Division,
Forestry Relations Department, Tennessee Valley Authority, who
has been most cooperative during the course of this study.
Genus CAMBARUS Erichson (1846)
Subgenus FAXONIUS Ortmann (1905)
CAMBARUS (FAXONIUS) PELLUCIDUS AUSTRALIS, new subspecies
Male I—Body white, digestive tract dark. Rostrum with mar-
gins only slightly converging. Marginal spines short and acute.
Acumen long and slender. Upper surface of rostrum moderately
concave. Postorbital ridges with short acute spines. Sides of cara-
pace minutely granular. Cervical groove unbroken in front of five
or six lateral spines on each side. Spininess usually reduced from
typical (. pellucidus. Antennae as long as the body. Antennal
scale broadest anterior to the middle, with inner margin gently
rounded. Apical spine short; half the length of that of typical
C. pellucidus. Dorsal surface of chelipeds with small tubercles.
Tips of fingers sparingly setose. Hooks on the third walking legs
prominent, globose, and recurved. Hooks on the fourth walking
legs lacking. Gonopods reaching to the coxopodites of the third
walking legs. Rami short and nearly equal in length. Outer ramus,
with corneus tip, curved tightly around the inner ramus. Inner
ramus straight with slightly recurved slender fleshy tip. Setose
along the ventral line.
Male IJ —Hooks on the third walking legs recurved and rounded
but reduced in size. Gonopods with fleshy tips reaching to the
coxopodites of the third walking legs. Inner ramus a little more
inflated.
Female——Annulus ventralis contrasting sharply with that of typi-
eal (. pelluctdus in that the large central hemispherical tubercle has
its greatest height on the anterior wall. The tubercle recedes pos-
teriorly and levels out to form a narrow flat border for the full
width of the annulus. A shallow median furrow marks the posterior
slope and becomes deeper and sinuate with a sharp curve to the
observer’s right in the posterior margin.
Variations—I have placed in this subspecies a crayfish from sev-
eral caves in northern Alabama on the basis of identical genitalia.
However, there are slight variations from cave to cave. The num-
CRAYFISHES FROM ALABAMA CAVES—-RHOADES 143
\
F
Figure 35.—Cambarus pellucidus australis, new subspecies: 4, Gonopod, male, form 1B
outer view; E, gonopod, male, form I, inner view; C, dorsal view of carapace; D, annulus
ventralis; B, gonopod, male, form II, outer view; F, gonopod, male, form II, inner view.
144 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
ber of lateral spines of the carapace varies from two to nine. The
spines are not necessarily paired. A specimen may have four spines
on the left side and nine on the right side. The areolae of the speci-
mens from Cave Spring Cave range from 33.3 to 36.6 percent of the
total length of the carapace. The Shelta material measures 38.5
to 40.5 percent. The crayfishes of this subspecies from other caves
in this region range from 36.1 to 39.5 percent. The blind cray-
fishes of the Mammoth Cave region also vary from cave to cave.
The areola of typical C. pellucidus is 36 to 41 percent. Shelta Cav-
ern and Huntsville Spring Cave specimens are similar in having
very short apical spines of the antennal scale. Other caves yield
specimens with long apical spine similar to C. pellucidus.
In spite of the variations listed above, the marked similarity
of these varieties causes me to place them all in the subspecies
C. pellucidus australis.
No doubt C. pellucidus australis of the South bears the same
affinity to C. pellucidus pellucidus as does the Cambarus pellucidus
testii Hay (1893) of the North.
Ecology and distribution—This crayfish is found throughout the
caverns of the limestone region in northern Alabama. According to
Dr. Walter B. Jones the presence of crayfishes in caves seems to be
correlated with the presence of blind fishes and aquatic insects. In
caves without connections with the surface, food chains develop
among the animals present. Mr. Hubricht suggests that bat guano
may provide some food for crayfishes.
Dr. Jones writes, “Shelta Cavern is a rather large cave with sev-
eral underground streams and rather large underground lakes. I
have never seen muddy waters in Shelta Cavern. There is scarcely
any outside trash entering the passages.
“Cave Spring Cave is a typical underground stream although there
are some rooms scattered about here and there. That cave is 3,050
feet long, or longer, and the water is quite cold. At times the stream
is muddy and completely fills many parts of the passage. In fact,
one cannot go very far back into it in wet seasons. The crayfish
fauna is rather abundant, and I could easily have taken a gallon of
specimens. Cave Spring Cave, as does Shelta Cavern, has white fish.
“Huntsville Spring Cave is about 84 mile long with a low ceiling
and a deep-channeled stream. It is reached by a vertical manhole in
a street near the center of the city. The roof and the floor are irregu-
lar. The stream is spring-fed and permanent, having an average
flow of 39,000,000 gallons a day. The cave is located under the city
of Huntsville and is full of narrow passages, crevices, and loose rock.
No fish have been found there.
CRAYFISHES FROM ALABAMA CAVES—-RHOADES 145
“Saddler Springs Cave is a typical underground stream that has
no connection with the surface. Apparently there has never been
the slightest bit of sediment or trash in the cave. Stalactites are
like crystal, and the floor of the stream is neatly carved out of
limestone rock with scarcely any sand or gravel anywhere in the
place. The crayfish fauna is somewhat limited, as are the other
faunas.
“McFarlen Cave is some 700 feet long and of varying width. The
entrance is archlike and of easy access. The stream is located in
back of the cave, and is spring-fed. It is my impression that there
is no permanent water in the front portion of the cave. The water
level may have been higher in former times. Boulders are to be
found on the floor of the cave. No fish have been taken there.
“Saltpeter Cave, in the Clear Creek area of Jackson County, is
located under a high bluff near the foot of a mountain. It is of the
fissure type. It is 1,895 feet long and most of its length is in the
zone of total darkness. The floor of the cave has a stream, evidently
permanent, and fed by several springs.”
Type locality.—Shelta Cavern, SEY,NE1, sec. 27, T.3 S., R. 1 W.,
north of Huntsville, Madison County, Ala.
Material examined—Two males II, 1937, Alvin R. Cahn coll.
(one paratype, U.S.N.M. No. 79365) ; 1 male I, March 1, 1938, Alvin
R. Cahn coll. (holotype, U.S.N.M. No. 79363); 1 male II, 2 females,
August 5, 1939, Leslie Hubricht coll. (one female is the allotype,
U.S.N.M. No. 79364) ; 3 males II, 3 females, 5 young, September 28,
1940, Walter B. Jones coll.
Additional records ——Cave Spring Cave, NW14,NE1, sec. 10, T. 5
S., R. 2 E., near New Hope, Madison County, Ala., September 26,
1939, Walter B. Jones (1 male II, 3 females, 5 young); December 1,
1939, Walter B. Jones (4 males II, 6 females).
Huntsville Spring Cave, SEYSW, sec. 36, T. 3 S., R. 1 W,,
Huntsville, Madison County, Ala., October 6, 1939, Walter B. Jones
(1 female).
Saddler Springs Cave, SE1,NE}{ sec. 3, T. 4, R. 1 E., Monte Sano
State Park, Madison County, Ala. June 14, 1940, Walter B. Jones
(1 male I, 4 males IJ, 3 females).
McFarlen Cave, SW14NW, sec. 22, T. 3, R. 3 E., near Garth,
Jackson County, Ala., February 29, 1940, Walter B. Jones (2 males I,
1 male II, 1 female).
Saltpeter Cave, NWY,SWY, sec. 16, T. 3, R. 3 E., Jackson County,
Ala., June 9, 1940, Walter B. Jones (1 male I, 6 males IT, 4 females).
146 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
Subgenus CAMBARUS Erichson (1846)
CAMBARUS (CAMBARUS) CAHNI, new species
Male I—Unknown.
Male II1—Body white, digestive tract dark. Rostrum of mod-
erate length, sides converging and sharply elevated. Marginal spines
small and often reduced to angles. Acumen rather short and broad.
Broad median carina reaching to a line drawn between the post-
orbital spines. Carapace slender, rounded, and minutely granular
on the sides. Cervical groove sinuate but unbroken, on the sides
above small rounded tubercles. Lines of the areola not clearly de-
fined. Length of areola varying from 36.7 to 40.5 percent of the
length of the carapace. Width accommodating three rows of widely
spaced dots. Epistoma oval, with small acute terminal spine.
Lateral margins sharply elevated. Antennae reaching to the telson
or beyond. Antennal scale triangular, broadest anterior to the
middle. Apical spine short. Chelae rather smooth, two or three
rows of low tubercles on the inner margin of the palm. Dots dis-
tributed evenly over the hand but tending to form furrows on the
dorsal surface of the fingers, two on the immovable finger and three
on the movable finger. Fingers two to three times the length of the
inner margin of the palm and twice as long as the width of the
palm. Merus with prominent furrow in the dorsal surface. Sharp
spine on inner surface with 0 to 8 small accessory spines. Carpus
with usual biserial row of spines down the ventral. Outer series
much exceeded by the inner. Hooks on the third walking legs rather
sharp and recurved. Gonopods thick, with fleshy tips recurved at
right angles with the shank. Inner ramus with tips slightly out-
curved as well as recurved. Setae on the ventral line.
Female—Chelae slightly shorter. Annulus broadly ovate. Cen-
tral and posterior regions elevated. Anterior wall somewhat de-
pressed. Fossa anterior and shallow. Median furrow curved to
form a small blunt hook to the observer’s left in a central position.
Affinities —C. cahni is intermediate between the “Section of C.
hamulatus” and the “Section of C. extraneus” (Ortmann, 1931, pp.
95-96). However, the cave modifications place it in the former
section. The carapace is subcylindrical, the chelae are long and
subcylindrical, and the eyes are greatly reduced, though not to the
extent found in C. hamulatus. The gonopods are recurved and the
lateral spines are present on the rostrum. I believe this crayfish has
sufficient constant and peculiar characteristics to give it the status
of a distinct species.
I possess a female crayfish from Saddler Springs Cave that is
lightly pigmented on the carapace and the dorsum of the abdomen.
It bears close resemblance to C. cahni in the subcylindrical carapace
CRAYFISHES FROM ALABAMA CAVES—-RHOADES 147
and body proportions. However, the eyes are normal, the antennae
are shorter, and the antennal scale is much narrower. The sides of
the rostrum converge more strongly, and there is no trace of a
median carina. The lateral spines of the carapace are small and
acute. The annulus ventralis is bisected by a deep median furrow
which curves strongly to the observer’s left to form a large blunt
Ficure 36.—Cambarus cahni, new species: 4, Dorsal view of carapace; B, annulus ventralis;
C, gonopod, male, form II, outer view; D, gonopod, male, form II, inner view.
lobe. I do not place this record with (@. cahni since the specimen
at hand bears greater affinity to the “Section of C. extraneus” than
to the “Section of C. hamulatus.”
Distribution —C. cahni is known only from the type locality, but
it will probably be found distributed over the limestone cave region
of northern Alabama where cave ecology is suitable. Belgreen Cave
is a small cave with a very deep underground stream. The stream
becomes muddy and almost fills the cavern in wet seasons.
148 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
Type locality—Belgreen Cave, NW14SWY, sec. 12, T. 7 S., R. 18
W., Franklin County, Ala.
Material examined.—Five males I1; 4 females, May 24, 1937, Alvin
R. Cahn coll. (one male is the holotype, U.S.N.M. No. 80031; 1 female
is the allotype, U.S.N.M. No. 80082.)
I take pleasure in naming the species for my friend Dr. Alvin
Robert Cahn, its collector.
CAMBARUS (CAMBARUS) HAMULATUS (Cope and Packard (1881))
The species is well known from Nickajack Cave and Wine House
Cave, Marion County, Tenn. An additional record, a female taken
with C. pellucidus australis from Shelta Cavern, Huntsville, Madi-
son County, Ala., March 1, 1938, by Alvin R. Cahn, is here con-
tributed. The sides of the rostrum of this specimen are more
convergent than typical and the lateral spines are very short. The
annulus ventralis is identical with the annuli of the Nickajack
female.
BIBLIOGRAPHY
Corr, EDwArRD DRINKER, and PACKARD, ALPHEUS SPRING.
1881. The fauna of Nickajack Cave. Amer. Nat., vol. 15, pp. 872-882.
Hay, WILLIAM PERRY.
1893. Observations on the blind crayfishes of Indiana, with a description
of a new subspecies; Cambarus pellucidus testiit. Proc. U. S. Nat. Mus.,
vol. 16, pp. 2838-286, 2 pls.
1902. Observations on the crustacean fauna of Nickajack Cave, Tenn.,
and vicinity. Proc. U. S. Nat. Mus., vol. 25, pp 417-439, 10 figs, 1 map.
ORTMANN, ARNOLD EDWARD.
1931. Crawfishes of the southern Appalachians and Cumberland Plateau.
Ann. Carnegie Mus., vol. 20, No. 2, pp. 61-160.
U. S. GOVERNMENT PRINTING OFFICE: 1941
PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM
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SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
Vol. 91 Washington : 1941] No. 3130
NOTES ON THE SNAKE GENUS TRIMORPHODON
By Hozgart M. Suir
Tuere are 13 forms definable at present in the colubrid genus
Trimorphodon. 'These very readily fall into two groups of six or
seven forms each, one characterized by presence of large, V-shaped
marks on head and neck (biscutatus group), the other characterized
by a transverse, light nuchal collar of varying width (wpsilon group).
The forms contained in the discutatus group are béscutatus biscutatus,
b. quadruplex, paucimaculatus, lyrophanes, lambda, and vanden-
burght. The members of the upsilon group are latifascia, fascio-
lata, upsilon, collaris, tau, vilkinsonii, and forbest. These two groups
form natural assemblages that certainly are of subgeneric rank.
In Trimorphodon, as in many other genera of snakes, evolution
has produced but few morphological innovations, and those that have
been produced are evident almost universally in terminal species that
appear to have been recently differentiated from a generalized stock.
Evolution in this genus has been evidenced chiefly in pattern; this is
the basic medium of speciation. Accordingly, differences in species
are to be sought primarily in the pattern, only secondarily in mor-
phology. Likewise, relationships and direction of evolution must be
traced through pattern changes, not by morphological variations.
Fortunately many of the steps in pattern evolution are shown or
indicated by species yet extant. The most important steps of all,
however—those that link the two radically different head and neck
patterns of the two groups—are lacking completely, and are not even
408590—41——1 149
150 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
indicated by variants of the several species. Only by sheer guess-
work can the process of divergence of these two types from some
common prototype be imagined.
Evolution within each group is relatively clear, and follows amaz-
ingly parallel trends.
In the bdiscutatus group are two closely related sections, of which
quadruplex is the most primitive of one, paucimaculatus of the other.
Of these two species, the latter exemplifies a more primitive pattern,
but both have large blotches and identical ventral counts, and they
differ from each other only in subcaudal counts and in extent of
subdivision of the blotches. In paucimaculatus the spots are very
broad but are divided only across the middle by a light streak or
spot; in guadruplex they are also divided medially, but the light
streak has completely split each blotch, and each of the resulting
spots is again split medially, so that superficially guadruplex very
strongly appears to have double the normal complement of blotches
of the group.
Modification of the pattern of guadruplea resulted in the develop-
ment of biscutatus. This form differs from guadrupleax only in its
pattern, which appears to have been produced by suppression of the
alternate blotches of guadruplex. That this was the procedure is
indicated by the fact that (1) the primary blotches in the northern
form are widely separated and number about half as many as in
quadruple; (2) the spaces between the blotches in bzscutatus are
frequently occupied by narrow, interrupted dark bands, which oc-
casionally are of the same shape as the primary blotches (more or
less H-shaped, light-centered) ; and (3) these “secondary” bands (sup-
pressed primary blotches), if enlarged to the size and character of
the primary bands, would reproduce the pattern of guadruplez.
The same process apparently has been followed in the section in-
cluding paucimaculatus, with the production of lyrophanes, lambda,
and vandenburghi. The most primitive pattern type among the de-
rivatives of paucimaculatus is, curiously enough, that of vanden-
burght (structurally the most highly modified species of the genus),
which represents a phase intermediate between paucimaculatus and
lyrophanes. To explain, the first step beyond the pattern type of
the former is the production of quadruple blotches, or, in other words,
double the usual number of primary blotches (as in quadruplez).
The next step is suppression of alternate blotches; in vandenburghi
about half have been suppressed (and accordingly the number of
biotches is distinctly higher than in paucimaculatus). In lyrophanes
nearly all alternating blotches have been suppressed, and secondary
bands are made evident betwen the primary blotches; sometimes one
or two of the alternate blotches are not completely suppressed but
THE SNAKE GENUS TRIMORPHODON—SMITH 151
remain evident as very small blotches. In lambda the process of
suppression is complete; the secondary bands are scarcely evident.
Obviously this succession of pattern types (paucimaculatus to
vandenburghi to lyrophanes to lambda) is not to be considered as an
indication of a similar succession in species evolution, for the
morphology here shows otherwise. Certainly /yrophanes and vanden-
burghi have been isolated for a long period from paucimaculatus,
since in them has been developed a spineless (i. e., very minute
lambda
vandenburghi
vilkinsonit
forbes
lyrophanes
ups ‘lon
paucimaculatus
fasciolata
collaris
biscutatus
latifescia
fou
quadruplex
Figure 37.—Diagram of the possible phylogeny of Trimorphodon.
spines) hemipenis. For some reason pattern change in vandenburghi
ceased or greatly slowed, and perhaps through its influence lyro-
phanes did not reach the stage of complete suppression of alternate
blotches that characterizes /ambda. The latter, of course, did not have
the retarding influence of vandenburghi,; and presumably its genetic
(and geographic) differentiation from paucimaculatus was made
complete at an early date—very likely at the time the lyrophanes-
vandenburghi stock was isolated.
152 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
This accounts for the déscutatus group. The record is not so clear
for the wpsilon group, which has members with more highly modified
patterns than the former but (with one exception) without special
morphological peculiarities. In this group two primitive forms are
still living—Jatifascia and fasciolata—of which the former has per-
haps the most primitive pattern. Both of these species have very
large, few blotches. In distribution they are peripheral to the cen-
tral plateau of Mexico. In relation to other members of the group
these two stand in much the same position as paucimaculatus and
quadruplex do in relation to other members of the biscutatus group.
However, it is difficult tc reconstruct so plausibly the process by
which other members of the wpsilon group were derived from
latifascia and fasciolata, suffice it to remark that their patterns may
have evolved by a splitting and suppression process much like that
which occurred in the dzscutatus group.
Evolution within the upsilon group is made most apparent by
changes in the head pattern. The two most primitive types have
none, or only a poorly indicated interocular light bar. The least
modification in other species is found in ¢aw, in which the inter-
ocular light bar is generally complete, and an indentation of the dark
head color along the parietal suture is evident. 7’. collaris repro-
duces this head pattern, and with ¢aw delimits an extensive geo-
graphic range completely peripheral to the central plateau. Since
increase in number of blotches seems to be the trend in the wpsilon
group, collaris with few, broad blotches is conceived to be more
primitive than tau. It is noteworthy that the opposite extreme
(from collarts) in number of blotches in taw occurs in Michoacan,
which 1s also the farthest extreme from collaris geographically.
The central-plateau species, wpsilon, was obviously derived from
tau or its near ancestor, as its head pattern, with a Y-shaped parietal
mark, is clearly derived from that of tau. In number of blotches
it remains very similar to the latter.
The end form in the upstlon group is vilkinsonii, in which are
apparent the extremes in reduction of head pattern and of body
blotches. The latter is not evidenced by trends in other species of
the group, although it is generally the case that multiplication in
number of blotches is followed by a decrease in their size. The sim-
ple 38-spot head pattern of juvenile vilkinsonii, however, is the end
result of the general trend, observed in other species, toward enlarge-
ment of the light areas of the head and consequent reduction in size
of the dark areas.
The body pattern of vilkinsonii is highly suggestive of the pat-
tern of Lampropeltis leonis, which is fairly certainly known to have
been derived by suppression of alternate blotches. This similarity at
THE SNAKE GENUS TRIMORPHODON—SMITH 153
least suggests the possibility that vi/kinsonii’s pattern was produced
in the same manner. The multiplicity of blotches in certain central
(Guanajuato?) specimens of wpsélon is an apparent step in this di-
rection. Possibly specimens from areas between Zacatecas and Chi-
huahua would show whether such a course may have been pursued
in the evolution of vilkinsoniz.
In view of the fact that several morphological changes took place
in the biseutatus group, with differences apparent in subcaudals,
hemipenis, and anal plate, it is remarkable that only one species in
the upsilon group possesses morphological characters sufficiently dif-
Ficure 38.—Distribution of the species of Trimorphodon. Inverted triangles, tau; triangles
not inverted, b. biscutatus (unless otherwise indicated); dots, unless otherwise indicated,
upsilon; vertical cross hatching, lambda; horizontal cross hatching, lyrophanes; diagonal
cross hatching, vandenburghi.
ferent from the group norm to identify it. This species (forbes?)
is very much like wpséon in pattern, and its apparently recent devel-
opment tempts a chronological association with the development of
the species in the other group with a single anal (vandenburghi).
With respect to pattern, it is noteworthy that, curiously, the end
form in neither group has undergone sufficient morphological dif-
ferentiation that it may thereby be distinguished from the members
of the group to which it belongs.
The relative age of the two groups is difficult to determine. One
group (biscutatus) appears to be of lowland origin, while the other
154 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
appears to be of highland origin. Accordingly, the fact that the
biscutatus group may have a Central American, or at least a more
southerly, origin does not necessarily mean that the wpsilon group
is a derivative of it, since it occurs toward the north in the general
direction of migration of the béscutatus group. In fact, the connec-
tion between the two groups is so remote that, were morphological
characters available, they would better be separated as different
genera.
KEY TO THE GENUS TRIMORPHODON
i Large V-shaped marks"on) headsand!na pes) ee ee eee 2
No such marks; a transverse nuchal collar (or whole neck light, as in
TUN SONU) a SEY a SDE AN AE eats dae, ee ee 7
2 VANS) entries oe np 3o es lh Te ee a vandenburghi
Ama) yi Gedo 6 ie I Bos Iie ab Aedes oe Rl Fb Ge SARC FI ea 3
3. Light V-shaped mark, which involves parietals, not confluent posterolaterally
10.
ale
with light color (or white) of ventral surface, but cut off by the continua-
tion posteriorly of the black band that on the head precedes the light band.
lyrophanes
Light mark extending posterolaterally direct to ventral surface, or at least
not cut off laterally by the preceding dark band_---___________________ 4
. Ventrals less than 245; blotches on body relatively numerous (maximum 34),
about as broad as long, not connected laterally in pairs (nor such a connec-
ELON STM ATCA TEA Vise Lee SR IEE Ea RN ea watt ee ae lambda
Ventrals more than 245; blotches on body numerous or few, but if the
former, connected laterally in pairs (or such a connection indicated) ___ 5
. Blotches on body numerous (about 338), connected laterally in pairs.
biscutatus quadruplex
iBlotchesvonsbody. less) mim en ouse (2a KOT 1 CSS) eee eee ee ee 6
. Blotches more than twice as long as spaces between; no evidence middorsally
of secondary bands orsblotchessa = eee paucimaculatus
Blotches less than twice as long as spaces between; usually secondary bands
or blotches present middorsally on some part of body.
biscutatus biscutatus
. Anterior dorsal blotch covering 15 or more scale lengths on middorsal line,
involving seven or more ventrals; blotches usually gray or black______ 8
Anterior dorsal blotch covering 18 or fewer scale lengths middorsally, in-
VOLVAN Pet OWED tia Tay SEVEN VCMT Tee Geese ease emia cae eae a 9
. Blotches little narrower laterally than dorsally, much broader on belly than
TNT ST) COS ee secre re Fe ee OS CE a ee fasciolata
Blotches much narrower laterally than dorsally, on belly equal to or narrower
than’: white minterspaces2 S220 eae ek i CRE cea latifascia
. Blotches very narrow, a third length of interspaces; anterior border of first
dorsal blotch 9 or 10 scales behind parietal_____.___________ vilkinsonii
Blotches broader, little if any narrower than spaces between; anterior
border of first dorsal blotch farther forward, not more than six seales
DehingdS parle teers sea a ee Se a eae ee eee eee 10
Fifth and six labials entering orbit; anterior loreal split, an upper and lower ;
talliwhite sunmarkeds tbelonyeecss 2820 Meals ole ee peer ee eee ee forbesi
Fourth and fifth labials entering orbit; anterior loreal single; tail marked
below: OF Mts os eel et ie ee ee 3 Te ee ale!
Bands: on; bod y-aGes sewers ee oe ee ee Ee es Sse eee collaris
IBANASON DOGY 2aTOT TO TC ee ee ns ees ane me sc ea 12
THE SNAKE GENUS TRIMORPHODON—SMITH 155
12. A roughly Y-shaped mark on parietals, the arms forking just behind frontal,
the mark usually enclosed by dark color posteriorly ; belly with some, sub-
caudal surface with numerous dark marks; blotches on body 28 to 82.
upsilon
No similar mark on head; dark color of head sharply truncate near posterior
edge of parietals, with a narrow or broad, light indentation along parietal
TEU ee eee eee ee ee ee ae eee ae ee tau
This study was completed, and a number of specimens on which it
is based was collected, during my tenure of a Walter Rathbone Bacon
Traveling Scholarship of the Smithsonian Institution. I am much
indebted to Dr. E. H. Taylor and L. M. Klauber for the loan of
numerous important specimens and for invaluable advice and criti-
cism, without which the study would have been impossible.
Genus TRIMORPHODON Cope
TRIMORPHODON PAUCIMACULATUS Taylor
Trimorphodon paucimaculatus Taytor, Kansas Univ. Sci. Bull, vol. 24, pp.
527-529, pl. 46, fig. 1, 19386 (19387) (Mazatlan, Sinaloa) ; ibid., vol. 25, p. 360,
pl. 35, fig. 3, 1938 (1989).—Ktavuser, Trans. San Diego Soc. Nat. Hist., vol.
9, p. 185, 1940.
Diagnosis —Large V-shaped marks on head, these not continued on
neck but disappearing laterally just behind head; hemipenis long,
with a middle belt of enlarged spines; ventrals 251 to 253, caudals
76 to 84; anal entire; blotches on body 20 to 25, a little more than
twice as long as spaces between; secondary bands reduced to small
lateral spots, not extending dorsally; tail blotches 10 to 18.
Specimens examined.—Two, including type.
Locality records—Mazatlin and Presidio, Sinaloa; San Blas,
Nayarit (U.S. N. M. No. 46617).
Remarks—The San Blas specimen is in very poor condition but
can be seen to have very broad blotches; it has 84 caudals.
This species, I believe, possesses the pattern of the ancestral type
of lambda, lyrophanes, and vandenburghi, which IT interpret as being
direct derivatives of it. It is, moreover, near the ancestral type of
the whole group, since it is a little less specialized, in pattern, than
the direct ancestor (quadruplex) of the other member of the group
(biscutatus).
TRIMORPHODON LAMBDA Cope
Trimorphodon lambda Corr, Proc. Amer. Philos. Soc., vol. 28, pp. 286-287, 1886
(Guaymas, Sonora).—Taytor, Kansas Uniy. Sci. Bull., vol. 25, pp. 360-361,
pl. 35, fig. 4, 1988 (1939).
Trimorphodon lyrophanes Ktauser, Trans. San Diego Soc. Nat. Hist., vol. 9,
pp. 181-187 (part), 1940.
Diagnosis —Large V-shaped marks on head, these not continued on
neck but disappearing laterally just behind head; hemipenis long,
156 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
with a middle belt of spines; ventrals 243 or less, caudals 86 or less;
anal entire; spots on body 34 or less.
Specimens ewamined.—Twelve, including type.
Locality records.—Various localities in California, Nevada, Utah,
Arizona, and Sonora (Klauber, op. cit., p. 187).
Remarks.—The hemipenis of a specimen from Guaymas, Sonora
(EHT-HMS No. 4572) is more than 16 caudals long (a portion
everted, dried); three large flounces cover the length of about 11
caudals; an area of spines covers a length of about five caudals,
proximal to area of flounces.
Another specimen from Telegraph Pass, Summit of Gila Moun-
tains, Yuma County, Ariz. (L. M. Klauber, No. 25488) has a hemi-
penis 22 caudals long; three large flounces, extending to the thirteenth
caudal from base, passing through an area of enlarged spines covering
the length of three caudals; remainder ridged, with tiny spicules.
The spinous area in this specimen includes the proximal ends of the
flounces, from the fourteenth to the sixteenth caudal inclusive; in
other words, the spines begin about seven caudals from the distal tip.
This is different from the condition in the Guaymas specimen, but
there seems to be a similar variation in position of the spinous area
in other species.
TRIMORPHODON LYROPHANES (Cope)
Lycodon lyrophanes Corr, Proc. Acad. Nat. Sci. Philadelphia, vol. 12, p. 348,
1860 (Cape San Lucas, Baja California).
Trimorphodon lyrophanes Corr, Proc. Acad. Nat. Sci. Philadelphia, vol. 13, p.
297, 1861.—Taytor, Kansas Univ. Sci. Bull., vol. 25, p. 363, 1938 (1939).—
KLAUBER, Trans. San Diego Soc. Nat. Hist., vol. 9, pp. 181-187 (part), pl.
%, fig. 2, 1940.
Diagnosis —Large V-shaped marks on head, these continued onto
neck, not terminating laterally behind head; hemipenis relatively
short, without enlarged spines; ventrals less than 243; anal divided.
Specimens examined.—N ine.
Locality records.—Various localities in Baja California: Cape San
Lucas, San José del Cabo, Santa Anita, Miraflores, Sierra San
Lazaro, Todos Santos, La Paz, Santa Rosalia, San Ignacio (Klauber,
loc. cit.).
Remarks.—The present species differs most markedly from lambda
in the character of the hemipenis, which is spineless (2. e., withont
enlarged spines) and shorter in lyrophanes (as in vandenburght),
while in /ambda it is longer and with spines (as in all other Z’rimor-
phodon). Three hemipenes dissected in situ on specimens from
Baja California agree well with the description of the extruded
hemipenis of vandenburghi given by Klauber (op. cit., p. 170), with
the exception that there are but three large flounces (instead of four;
THE SNAKE GENUS TRIMORPHODON—SMITH 57;
an additional, smaller, terminal flounce is not readily discernible in
noneverted hemipenes). In addition it may be observed that the
hemipenis is 16 to 20 caudals long (in situ) and that the flounces are
relatively small, near the tip, and cover a length equal to the length
of four or five caudals.
Specimens examined show constant differences in head and neck
pattern from lambda. In lyrophanes the dark, V-shaped mark (which
extends nearly or quite to a line even with posterior border of orbits)
extends posteriorly onto the neck, without a break; in lambda it
extends posterolaterally and terminates a little posterior to the labials,
about even with a line drawn back from the lip. The light band
posterior to this dark band in /yrophanes continues onto the neck and
terminates with a large neck blotch, or else its arms unite posteriorly
and may pierce the neck blotch posteriorly; in Jambda this mark
extends posterolaterally and usually unites with the white of the
ventral surface.
A difference in the character of the dorsal blotches in lyrophanes
and lambda is evident to the eye but is not well suited to measurement.
The blotches are narrower and longer in lyrophanes, and fairly well
severed from their lateral extensions; they are broader and shorter in
lambda, and their lateral extensions are not so strongly differentiated.
TRIMORPHODON VANDENBURGHI Klauber
Trimorphodon vandenburghi KLAUBER, Bull. Zool. Soc. San Diego, No. 1,
pp. 17-18, fig. 8, 1924 (Wildwood Ranch, 1,520 feet, 5 miles southwest of
Ramona, San Diego County, Calif.) ; Trans. San Diego Soe. Nat. Hist., vol.
5, pp. 183-194, pls. 22, 23, 1928; vol. 9, pp. 169-180, pl. 7, fig. 1, 1940.
Diagnosis.—Large V-shaped marks on head, these usually not con-
tinued on neck; hemipenis short, without spines; ventrals 244 or less;
anal entire.
Specimens examined.—One.
Locality records——Numerous localities in southern California (see
Klauber, op. cit., 1940).
Remarks.—A single hemipenis examined in situ agrees with the
description given by Klauber (op. cit., 1940, p. 170), with the excep-
tion that only three flounces are discernible (instead of four). In
addition, the hemipenis is 14 caudals long and the flounces are rela-
tively small, as in lyrophanes.
This very distinct species appears to be directly related to lyro-
phanes. Its chief difference from the latter—the entire anal—is an
amazing development in a genus with so few structural variations.
TRIMORPHODON BISCUTATUS QUADRUPLEX, new subspecies
Holotype.—U. S. N. M. No. 89476, female, Esteli, Nicaragua, col-
lected by J. H. Ivy in 1932.
408590—41——_2
158 PROCEEDINGS OF THE NATIONAL MUSEUM Vou. 91
Paratypes.—U. 8. N. M. No. 5569, Realejo, Nicaragua; No. 6805,
Guatemala; No. 32274, San Juan, Nicaragua.
Diagnosis —A member of the biscutatus group, with large V-
shaped marks on head; dark blotches completely divided, each of
practically all the resulting sections again partially split medially;
counted separately, blotches 83 (pairs numbering 17); ventrals 251
to 263; total counts 334 to 347.
Description of holotype.—Supralabials 9-9, fourth and fifth enter-
ing orbit, third smallest, fifth (or sixth) largest; three preoculars,
upper largest, in contact with frontal; three large loreals, the
smallest lowermost and directly above third supralabial; three sub-
equal postoculars; three anterior temporals; infralabials 13-13, 4-5
in contact with chin shields.
Dorsals in 25-26-17 rows, with two apical pits, those on posterior
third of body convex or bluntly keeled; ventrals 261; anal divided;
caudals 82.
Maxilla with 11 teeth, the last two grooved, offset from others,
shghtly shorter than longest anterior teeth, preceded by a short
diastema; other teeth separated from each other by equal spaces,
decreasing in size posteriorly; anterior smaller than succeeding
teeth, which are the largest of maxilla; tooth preceding fangs half
length of latter.
Hemipenis (of No. 32274) 25 caudals long; flounces three, large,
covering a length equal to between seven and eight caudals; about
70 enlarged spines in a small area (length of four caudals) proximal
to flounces.
General color gray; a dark-brown, black-edged bar extending
across top of head a little in front of eyes, anterior edge of frontal
about in its middle; this followed by a light bar which extends
diagonally onto sides of head, reaching labial border at eighth and
ninth labials; this followed by a broad, V-shaped black mark, termi-
nating laterally even with mouth, split by a longitudinal white line
on middorsum; this followed by a somewhat narrower V-shaped
light mark, extending laterally to ventral surface; following this, a
similar V-shaped dark mark, but this prolonged posteriorly and
uniting with first blotch, enclosing medially a long, broad, light line;
this blotch is the first of a series of 33 brownish-gray, dark-edged
blotches, many of which are joined in pairs, most with a light, broad,
transverse median area which nearly divides them; sides of body
with a series of small, irregular spots, one placed between alternating
spots (i. e., between the pairs) ; ventral surface stippled, a little more
posteriorly than anteriorly; ends of about every other or every third,
occasionally of two adjacent ventrals dark brown; chin and gular
region immaculate; ventral surface of tail a little more heavily
stippled than body.
THE SNAKE GENUS TRIMORPHODON—SMITH 159
Variation—The paratypes available are in such poor condition
that the number of blotches cannot be counted, but they are of the
same nature as in the type. The scale characters of Nos. 5569, 6805,
and 32274, respectively, are: Scale rows 25-25-17, ?, 23-25-?; ventrals
255, ?, ?; caudals 92 (4), 93 (6), 90 ( 8); supralabials 9-9, preocu-
lars 3-3, postoculars 3-3, in all; infralabials 13-14, 13-14, 12-13;
loreals 3-8, 2-3, 2-3; preoculars separated from frontal on one side
im one.
Comparisons.—The present form differs from béscutatus solely in
the extent of subdivision of the blotches, which in this are very com-
plex, consisting of two halves (each of which appears like the pri-
mary blotches of biscutatus), which again are partially divided.
For practical purposes of separation from biscutatus, the blotches
may be considered separately, whereby the number secured is much
greater than the number of primary blotches in bescutatus.
TRIMORPHODON BISCUTATUS BISCUTATUS (Duméril and Bibron)
Dipsas biscutata DUMERIL and Brsron, Erpétologie générale, vol. 7, p. 1158, 1854
(Mexico).
Trimorphodon major Corr, Proc. Amer. Philos. Soc., vol. 11, p. 153, 1869 (Tehuan-
tepec).
Diagnosis —Large V-shaped marks on head; dorsal blotches 18 to 23
on body, separated from one another by a distance at least a little
greater than half their own length (usually equal or greater) ; a sec-
ondary, transverse, broken, narrow, black band between each pair of
primary blotches (rarely reduced to lateral spots; in this case the
primary blotches do not close the large space between the primary
blotches) ; ventrals 251 to 275; caudals 81 to 102; total counts 343 to
376.
Specimens eaamined.—T wenty-four.
Locality records.—Acceptable records are from the Isthmus of
Tehuantepec (Santa Efigenia, El Barrio, Tres Cruces, Tehuantepec,
Cerro Guengola, La Concepcién) in the State of Oaxaca; Tonala and
San Ricardo in Chiapas; Huajintlan, Morelos; Agua del Obispo, Or-
ganos, Acapulco, and La Crucita, Guerrero; and Hda. El Sabino and
10 miles north of Tafetan, Michoacan.
Remarks—A. specimen from Tehuantepec has a hemipenis 24
caudals long; flounces 3, large, covering a length equal to about 7
caudals; area of spines covering a length of 4 or 5 caudals.
As pointed out by Taylor; northern specimens have higher average
ventral and caudal counts than southern specimens. Present speci-
mens are insufficient, however, to show whether the differences are
significant and practically recognizable. The counts are given in
table 1.
? Kansas Univ. Sci. Bull., vol. 24, pp. 358-360, 1939.
160 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
TasLE 1.—Scale counts of Trimorphodon biscutatus biscutatus
No. Sex Venirals Caudals Totals State
23619 a 260 100 360 | Michoacan.
5339 a 267 95 362 Do.
5338 a 269 101 370 Do.
110410 a 275 100 375 | Guerrero.
5508 a 260i) espe ee |e ipa eae Do.
21404 o 275 85 360 Do.
4588 a 270 100 370 Do.
5145 oO 275 101 376 Do.
5146 o' 274 99 373 Do.
5147 a 272 99 371 Do.
5148 o Die 102 374 Do
30406 o 252 91 343 | Oaxaca.
30427 J 260 94 354 Do.
30428 etd ee re RARE OO) fe Res Seerra Do.
30429 o 260 85 345 Do.
46547 ot 260 85 345 Do
110404 o! 263 81 344 Do
110405 o 251 94 345 Do
110406 oJ 255 94 349 Do
110407 o 255 96 351 Do
110409 J DOS ee eee ata ea yee Chiapas
110403 Q 271 88 359 | Oaxaca
110408 2 269 90 359 Oo.
4589 9 261 85 346 | Chiapas.
TRIMORPHODON LATIFASCIA Peters
Trimorphodon biscutata latifascia PETERS, Monatsb. Akad. Wiss. Berlin, 1869,
p. 877 (Puebla).
Trimorphodon latifascia TAytLor, Kansas Univ. Sci. Bull., vol. 25, pp. 364-365
(part), pl. 36, fig. 2, 1938 (1939); vol. 26, p. 479, pl. 52, 1940.
Diagnosis.—A light, transverse nuchal collar; hemipenis long, with
a median belt of spines; blotches very long, 18 to 15 on body, 5 to 7
on tail, the first covering 15 or more scale lengths middorsally; num-
ber of ventrals involved by each dark band slightly more to half
number involved by adjacent light areas.
Specimens examined.—Ten.
Locality records—‘Puebla” (perhaps the region of Matamoras) ;
12 miles south of Puente de Ixtla, Morelos; Huajintlén, Morelos;
between Cuernavaca and Tepoztlin, Morelos.
Remarks.—Hemipenis (HHT-HMS No. 5540, Huajintlan, Morelos)
28 caudals long (in situ), with three large flounces extending 10 caudal
lengths proximally, followed by an area of enlarged spines about
three caudals long; remainder with longitudinal ridges surmounted
by tiny spines.
TRIMORPHODON FASCIOLATA, new species
Holotype.—U.S.N.M. No. 110400, male, from near Zararacua Falls,
6 kilometers southeast of Uruapan, Michoacan.
THE SNAKE GENUS TRIMORPHODON—SMITH 161
Diagnosis —A member of the wpsilon group, having a transverse,
light nuchal collar; dorsal bands few (18 in type), little narrower on
sides than on middorsal line, and much longer ventrally than light
spaces between; ventrals 219, caudals 76, scale rows 23, in type; no
interocular light bar.
Description of holotype-—Supralabials 8 or 9, fourth and fifth
entering orbit on one side, third also on other; two large loreals and
on one side a third small loreal at posterolateral border of second
loreal; preoculars 2 or 3, upper somewhat the largest and in contact
with frontal; three postoculars, median somewhat the smallest; three
anterior temporals, followed by three secondary temporals on one side,
four on other; 12 infralabials, six in contact with chin shields, five
with anterior pair; posterior chin shields separated medially, nar-
rower and shorter than, and about two-thirds the size of anterior
chin shields.
Scales in 21-23-15 rows, smooth, with paired apical pits; scales
above anus slightly convex; ventrals 219; caudals 76; anal divided.
Maxilla with 10 teeth, in four groups; three anterior teeth, the
anterior smallest of the three and subequal in size to ungrooved teeth
in other groups, the posterior somewhat larger than second, which is
very nearly as large as posterior grooved teeth; one tooth in second
group, about size of first tooth, separated from other teeth on either
side by a short but very evident diastema; four teeth follow, smallest
of the maxilla, very slightly decreasing in size; two posterior teeth
enlarged, offset, separated by a distinct diastema (subequal in length
to other diastemata) from preceding teeth.
Hemipenis long (25 caudals), slender (not everted) ; proximal third
with numerous ridges capped by very minute, scarcely discernible
spines; adjacent sixth with about 50 small spines, which extend to
the middle of the hemipenis; distal half without spines, ridged, with
three large flounces, which have tiny papillae on their free edges; distal
half with tiny papillae; tip with somewhat larger papillae, apparently
not bifurcate; sulcus single.
Top of head dark, with numerous tiny light flecks, no trace of reg-
ular markings except a median, V-shaped mark posteriorly, apex for-
ward; sides of head more light than dark, top of head more dark
than light; nuchal collar white, with some dark stippling, its pos-
terior border nearly straight, somewhat concave, a little more than
two scale lengths behind parietals medially ; anterior border of nuchal
collar vague, grading into darker color of head, especially laterally.
Thirteen very broad, dark cross bands on body, four on tail; first
five bands covering 19 to 21 scale lengths medially, remaining bands
decreasing in length posteriorly; first five bands covering 15 to 18
scale lengths on first scale row, remaining bands fewer, but all bands
162 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 9t
covering about three-fourths as many scale lengths laterally as on
middorsal line; each dark band with a narrow, broken, transverse
white line dividing it into two halves; spaces between bands white,
covering one and one-half to two and one-half scale lengths medially,
all except the anterior three and nuchal collar enclosing laterally a
small dark spot, which involves two scales of the first scale row and
the end of the ventral scale between them; dark bands encroaching
on ventral surface, the median and posterior completely encircling
body, although with numerous light flecks on midventral surface;
numerous dark flecks on venter between posterior bands; ventral
surface of tail irregularly mottled with light and dark; chin
immaculate.
Comparisons.—This species most closely approaches Jlatifascia
Peters, as defined by the specimens reported by Taylor.? One of
these is described as having the first four bands covering 19, 15, 16,
i6 scales (first five covering 19 to 21 in fasciolata), but they are
distinctly narrower laterally, involving 7 to 9 ventrals, while the
white areas between involve 9 or 10 (dark bands involve 13 to 17,
light bands 6 ventrals in fasciolata).
TRIMORPHODON UPSILON Cope
Trimorphodon upsilon Corr, Proc. Amer. Philos. Soc., vol. 11, p. 152, 1869 (Guada-
lajara; type, U.S.N.M. No. 31858).—Taytor, Kansas Univ. Sci. Bull., vol.
25, pp. 865-366, pl. 35, fig. 2, 1988 (1939).
Diagnosis —A light, transverse nuchal collar; head largely dark,
but with a light interocular bar and a Y-shaped light mark on parietal
region, the arms of which fork immediately behind frontal; 23 to 32
body blotches, 11 to 15 tail blotches.
Specimens examined.—Twelve.
Locality records——Known from the central, southern, and north-
western plateau region. Recorded from the States of Chihuahua
(Batopilas), Durango (Ventanas); Guanajuato; Hidalgo (Zacual-
tipan; 10 km. north of Jacala) ; Jalisco (Cumbre de los Arrastrados;
Guadalajara; Magdalena) ; Michoacan (Tacicuaro) ; Nayarit (Sierra
de Nayarit) ; Zacatecas (San Juan Capistrano).
Remarks.—The dorsal bands of a specimen observed in life (from
Magdalena, Jalisco) were reddish brown; the color and general char-
acter of the rhombs resembled to some extent those of certain Lam-
propeltis.
The ventral surface in this species is distinctly marked with ir-
regular black spots; the subcaudal surface is more heavily blotched
than the belly. A single exception is a somewhat faded, soft speci- |
—
2 Kansas Univ. Sci. Bull., vol. 25, pp. 864-365, pl. 36, fig. 2, 1939; vol. 26, p. 479, pl. 52,
1940,
'
i
'
THE SNAKE GENUS TRIMORPHODON—SMITH 163
men evidently preserved just before shedding, so the color is greatly
obscured (No. 12419, Guadalajara) ; another specimen, nearly perfect,
from the same locality, has the whole ventral surface very heavily
pigmented. In this respect upsilon differs from typical specimens
of tau, collaris, forbesi, and vilkinsonii and agrees with fasciolata
and latifascia.
The hemipenis of a specimen from “Mexico” (with 30 body
blotches) is 26 caudals long; three large flounces, covering the length
of eight caudals; area of spines covering the length of four caudals.
In general there appears to be an increase in number of body
blotches toward the east. Western specimens (three from Guada-
lajara, and Magdalena, Jalisco) have the fewest (23, 24, 25), while
specimens from eastern localities (Guanajuato, Hidalgo, Zacatecas)
have 27 to 32.
TRIMORPHODON FORBESI, new species
Holotype-—U.S.N.M. No. 110402, male, from San Diego (about
5 miles south of Tehuacin), Puebla, collected by Dyfrig McH.
Forbes.
Diagnosis —A transverse nuchal collar, heavily suffused dorsally
with dark pigment, so that the first dorsal band is more or less con-
fluent with the dark head color; belly very light, dark markings
dim; no markings on ventral surface of tail; bands on body 21, the
first five covering 13, 8, 9, 10, 10 scale lengths, respectively; nine
supralabials, fifth and sixth entering orbit; anterior loreal divided;
a large light area on head, including posterior portions of supraocular
and frontal, and more than half (anterior) the parietals, indented
posteriorly by a dark area, which reaches nearly to the posterior tip
of frontal.
Description of holotype.—F¥rontal as high as wide, portion visible
from above a little longer than its distance from prefrontals, as
long as internasals; latter two-fifths as large as prefrontals; length
of frontal equal to its distance from tip of snout; nasal completely
divided, anterior section somewhat smaller than posterior; anterior
loreal wedged between internasals and prefrontals, divided into an
upper and lower part; a large posterior loreal; on one side a small
subloreal, making a total of three loreals on one side, four on other;
three preoculars; three postoculars, middle smallest, lowest largest;
temporals 8-4-5; supralabials nine, fifth and sixth entering eye, fourth
smallest, sixth perhaps largest; infralabials 12, five in contact with
anterior chin shields, two with posterior; first infralabial largest;
anterior chin shields twice size of posterior.
Dorsal scales smooth, with two apical pits, in 28-23-16 rows;
supra-anal scales convex; ventrals 213; anal divided; caudals 77.
Total length 818 mm.; tail 150 mm.
164 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
Hemipenis 23 caudals long; three large flounces, covering eight
caudal lengths; area of spines covering four caudal lengths.
Color.—Dorsal color very light brownish gray, lighter in vertebral
region; 21 rhombs on body, 11 on tail; rhombs light brown, with a
slightly reddish tinge; a narrow black border on each rhomb, the
borders not extending below about the third scale row; rhombs ex-
tending to ventral scales; first five rhombs covering 13, 8, 9, 10, 10
scale lengths, last five 6, 5, 6, 6, 7 scale lengths, respectively (on mid-
dorsum) ; spaces between rhombs about equal to three scale lengths
middorsally; on first scale row rhombs cover only one or two scale
lengths; a series of very small, lateral spots alternating with the
rhombs, these involving the lower part of the first scale row and
the ends of the ventrals, each spot covering an area about equal to
the size of three lateral scales.
Ventral surface-of body nearly white; lateral spots encroaching
upon venter, but very subdued, as are all other dorsal markings where
they reach the venter; ventral surface of tail white, immaculate.
General tone of head color gray-brown; snout light gray, stippled;
this color extending in a wide band along the prefrontal suture to
frontal; latter band with a black border extending a little anterior
to middle of prefrontals, posteriorly continuing onto corner of frontat
and then curving onto supraocular; area enclosed by these dark
borders on the frontal is dark, confluent with a dark interocular bar,
which is black-edged posteriorly, passes through the middle of the
supraocular and occupies the same position as the usual interocular
light bar; posterior to this a narrowly black-edged, extensive light
area, which occupies the posterior half of frontal, posterior portion
of supraoculars, and anterior half of parietals; this light area
notched posteriorly, the dark edge curving sharply forward nearly to
tip of frontal; posterior and lateral to this is a darkly suffused area,
which medially extends to the anterior border of the first dorsal
rhomb; nuchal light collar present; its posterior border nearly
straight (anterior edge of first rhomb), but the collar itself very
dim, due to the dark dorsal suffusion; sides of head gray; posterior
supralabial region suffused with pink.
Remarks.—One of the most remarkable features of this snake is the
peculiar head pattern, which is, in general, much like that figured
for tau (Taylor, op. cit., 1940, fig. 8), except that the dark area of
the frontal and parietals is light, although just as distinctly out-
lined; the dorsal nuchal area, light in taw (and in all other members
of the upsilon group) is dark in forbesi,; the interocular light bar,
characteristic of the entire group, is dark in forbesi.
It appears that a pattern reversal has taken place; whether it is
an anomaly in the single type or is characteristic of the species can-
THE SNAKE GENUS TRIMORPHODON—SMITH 165
not now be stated. It is remarkable that the reversal of pattern
begins anteriorly precisely at the frontal-prefrontal suture; anterior
to this suture the head pattern is normal, with a light snout and a
light, longitudinal median line with darker sides; posterior to this
suture the light color is very sharply changed to dark, and vice versa,
with the exception of the black borders, which outline the markings
and which remain constant.
While the head pattern of forbest is very different from that of
other species of the group, it cannot be considered in differentiation
of the species from wpsilon, since there is a strong possibility it may
be anomalous. There are numerous other unique characters in
forbest. No specimens of other species of Zrimorphodon of the
upstilon group have the anterior loreal divided; and no other of that
group has the fifth and sixth labials entering the eye. These charac-
ters, combined with a faintly marked belly and white, unmarked
subcaudal surface (wpsilon has the belly, and especially the tail,
distinctly dark-mottled) ; number of rhombs (fewer than in wpsilon
and taw with a minimum of 23, and more than in collaris with 16) ;
narrow black borders of the rhombs (broad in tau, possibly in
collaris) ; length of rhombs on middorsal line (as long as in codlaris,
longer than in tau or upsilon) ; all define a species very different from
any other of the upsz/on group.
The closest relative of forbesi, I believe, is wpsilon,; the general
appearance of the dorsal rhombs is much the same. The elimination
of the ventral markings and lightening of the dorsal markings may
be compared with the same tendency in other deserticolous reptiles
which develop a faded pattern. The remarkable changes in cephalic
scutellation bring to mind a somewhat similar, recent change in van-
denburghi of the other (déscutatus) group, in which a single anal is
developed. Neither of these two species is otherwise greatly (although
somewhat) different from its closest relative.
The type is from a semiarid region. So far as known wpsilon is
restricted to more humid areas.
TRIMORPHODON COLLARIS Cope
Trimorphodon collaris Corr, Journ. Acad. Nat. Sci. Philadelphia, ser. 2, vol. 8,
p. 181, 1875 (“Orizaba’’).—SumicHRast, La Naturaleza, vol. 6, p. 14, 1882.
Trimorphodon latifascia TAytor (part), Kansas Univ. Sci. Bull, vol. 25, pp.
364-365, 1938 (1939).
Diagnosis.—A light, tranverse nuchal collar; an interocular light
bar; snout light; 16 bands on body, the longest covering 13 scale
lengths middorsally, eight on venter ; spaces between blotches covering
four and one-half to six scale lengths middorsally.
Specimens examined.—The only one known, the type (U.S. N. M.
No. 26499).
166 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
Locality records.—Described from “Orizaba,” but doubt is cast
upon this locality by the presence of two different labels (in the same
handwriting) with the type, both stating “Tehuantepec” as the lo-
cality, Sumichrast, the collector, states that “the typical individuals
came from Tuxpango, near Orizaba” (doe. ecét.).
Remarks.—In the absence of well-differentiated scale characters in
the group, color differences must be relied upon to distinguish various
species, It is true the type of cod/arés has nine labials, as do /atifaseia
and fasevolata, but this of itself means very little, since occasional
specimens of wpsiZon also have nine. I have considered collaris dis-
tinct from Zatifaseia because (1) the bands are considerably smaller on
the middorsal line (13 seale lengths, maximum), and the intervening
spaces cover four and one-half to six scale lengths; and (2) there are
distinet head markings, including sharp differentiation of head pat-
tern from nuchal collar, latter encroaching upon parietals, interocular
heht bar evident, a light bar evident along internasal and prefrontal
suture, and snout white. These characters place it in the section with
upsilon.
Essentially the only difference between this and wps/on is the small
number of blotches (16) on body. The minimum in upsilon is 23
(specimen from Quadalajara, Jalisco, type locality).
This is the only specimen of the genus that has ever been taken
on Atlantic slopes, at least in Mexico.
TRIMORPHODON TAU Cope
Trimorphodon tau Cork, Proc, Amer. Philos. Soe., vol. 11, p. 152, 1869 (“Tehuan-
tepec,” in error).—Sumicwrast, La Naturaleza, vol. 6, p. 14. 1882.—Taytor,
Kans. Univ. Sci. Bull, vol. 25, pp. 865-866, pl. 35, fig. 2, 19388 (1989) ; vol.
26, pp. 464-477, pl. 51, fig. 8, 1940.
Diagnosis. —A light nuchal collar; an interocular light bar indi-
cated; an indentation posteriorly of black head cap, but no Y-shaped
head mark behind frontal.
Specimens examined.—Five.
Locality records —Quidtepec (U. S. N. M. No. 30338, type), San
Felipe (KHT-HMS No. 5507), and Oaxaca (EHT-HMS No. 5506),
all in the State of Oaxaca; 7 miles east of Chilpancingo, Guerrero
(EHT-HMS No. 23417); and between Morelia and Hidalgo,
Michoacin (EHT-HMS No. 21402).
Remarks.—The type locality of this species is not Tehuantepec, as
stated by Cope, since Sumichrast (oc. cit.) states, “I found the
type of this species near Quidtepec, between Tehuacin and Oaxaca.”
The primary difference between few and wpsi/on is in head pattern.
In the former the dark head color is abruptly truncate near the
posterior tips of the parietals, and a light indentation (broad or nar-
row) is visible along the parietal suture. In wpsilon the dark head
THE SNAKE GENUS TRIMORPHODON—SMITH 167
color is not so abruptly truncate posteriorly, terminating posterior
to the parietals; and the light, midparietal indentation of tay is re-
placed by a narrow, Y-shaped mark, the arms of which follow near
the posterior sutures of the frontal, and sometimes reach to the outer
edge of the supraoculars, where they join with the tips of the inter-
ocular light bar.
Variation in body pattern in tau is so great that no contrast of the
species as a whole with upsilon is possible. The variants of tau ap-
pear to be segregated geographically but are represented by so few
specimens that the apparent differential characters of the three popu-
lations indicated may not be well founded.
The range of tau is apparently the periphery of the central Mexican
plateau. The extreme southern records near Oaxaca city, in the iso-
lated mountains of central Guerrero, and in the mountains at the ex-
treme edge of the plateau in Michoacan all indicate such a peripheral
distribution. AJ] three loci represented by specimens, however, are so
far removed from each other that the peculiarities of each popula-
tion (two of which are represented by single specimens) may prove
to have special significance: that is, at least three subspecies may
exist in tau:
1, Oaxaca specimens (3). Dorsal blotches 23 to 26; tail bands
9 to 10; belly very little pigmented ; subcaudal surface nearly uniform
white; interocular band complete; nuchal blotch two to three scale
lengths behind parietal; body blotches (except two nuchal ones) in-
volving three or fewer scales in first row, average two.
2. Guerrero specimen (1). Dorsal body blotches 22; tail blotches
8; belly heavily pigmented, the dorsal bands visible (not sharply
defined) ; subcaudal surface very strongly mottled; interocular band
reduced to a round spot in middle of frontal; nuchal blotch five
scale lengths behind parietal; body blotches (except two nuchal)
involving two to six scales in outer row, average five.
3. MicnosacAn specimen (1). Dorsal body blotches 24; tail bands
11; belly with some dark spots, poorly defined; subcaudal surface
moderately pigmented ; interocular band complete; nuchal blotch one
scale length behind parietal; body blotches not well defined on outer
scale rows, involving two or three scales on outer row where visible.
TRIMORPHODON VILKINSONII Cope
Trimorphodom vilkinsonii Core, Proc. Amer. Philos. Soc., vol. 23, pp. 285-286,
1886 (Chihuahua).—Tayiorz, Kansas Univ. Sci. Bull., vol. 25, pp. 361-363,
fig. 1, pL 28, 1928 (1929).—Ktaverr, Trans. San Diego Soc. Nat. Hist., vol. 9,
pp. 187-189, 1940.
Diagnosis —A broad, light area on neck, between dark areas on
head and first body blotch; bands a third length of interspaces; dark
head area only three spots in young.
168 PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 91
Specimens examined —One, the type, U. S. N. M. No. 14268.
Locality records.—Chihuahua and El Paso, Tex.
Remarks.—This species obviously is a close relative of wpsilon, from
which it differs chiefly in the narrowness of the dark bands, which
are a third as broad as the spaces between them.
U. §. GOVERNMENT PRINTING OFFICE: 1941
PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM
SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
Vol. 91 Washington: 1942 No. 313]
CATALOG OF HUMAN CRANIA IN THE UNITED STATES
NATIONAL MUSEUM COLLECTIONS: ESKIMO IN
GENERAL
By Autres HrpuicKa
INTRODUCTION
In 1924 the United States National Museum published the first
of its catalogs of crania. This included the measurements of 245
Eskimo skulls from one locality (St. Lawrence Island), with four small
series of other Alaskan skulls, which at that time was the total from
these peoples or localities in the Museum’s possession. Since then,
under the auspices of the Smithsonian Institution, no less than 18
expeditions to Alaska have been made, 10 of them conducted by the
author. These expeditions covered all the more important parts of
the coast, the main rivers, and the principal islands. Their purpose
was to study the living Eskimo, to collect skeletal remains over all
the once-inhabited territory, and to excavate old sites, which every-
where in Alaska yield both cultural and_ skeletal materials.
Most of the results of researches on the living Eskimo have been
published, as have those on much of the skeletal material collected
before 19301; but today the Eskimo material alone comprises more
than 2,200 crania, 2,100 of which are adult, mostly in an excellent state
of preservation and in a large proportion of cases accompanied by the
rest of the skeleton. The whole constitutes an exceedingly precious
series, data on which will be of basic importance. These data are
1 Hrdlitka, Ales, in Smithsonian Exploration Pamphlets, 1926-39; Anthropological survey in Alaska, 46th
Ann. Rep. Bur. Amer. Ethnol., 374 pp., 1930; The Eskimo of the Kuskokwim, Amer. Journ. Phys. Anthrop.,
vol. 18, pp. 93-135, 1933.
169
416178—42— —1
170 PROCEEDINGS OF THE NATIONAL MUSEUM you. 91
assembled in the present publication, and no pains have been spared
to make them thoroughly reliable. All the measurements were made
by the author, using the same tested instruments and identical methods
throughout. Several parts of the series have been sexed two or even
three times; some, for the sake of accuracy, were completely remeasured
and some determinations have been added to those of the first Catalog.?
It has been necessary to make only inconsequential changes in the
earlier figures, so far as they went.
The methods used are given in the author’s ‘Practical Anthro-
pometry,’”’ * but for the sake of completeness they are repeated
herewith:
THE VAULT
Maximum length of the skull, or greatest anteroposterior diameter.—
From the center of the glabella to the most distant normal point of
the occiput.
Maximum breadth of the vault, or the greatest transverse diameter. —
Above the supramastoid crests (posterior roots of the zygomae).
Height of the vault: The basibreqmatic height—The linear distance
from the midpoint on the anterior edge of the foramen magnum
(basion) to bregma.
FACE
Menton-nasion height.—Total morphological facial height on the
skull. The distance from menton to nasion, with the lower jaw in
place and the teeth in normal apposition.
Alveolar point—nasion height—The upper facial height on the
skull. The distance from the upper alveolar point to nasion.
Maximum bizygomatic diameter.—The greatest bizygomatic breadth.
BASE
Endobasion—nasion diameter.—Distance between the endobasion and
nasion.
Endobasion-subnasal point diameter —Distance between endobasion
and the left subnasal point.
Endobasion—prealveolar point diameter.—Distance between endo-
basion and the prealveolar point.
Angles of facial and alveolar prognathism.—The most satisfactory
way of obtaining these angles is to chart, with the aid of the sliding
compass, the three measurements together with the nasal and naso-
alveolar heights, and measure the angles directly by a transparent
(celluloid) goniometer. For the naso-alveolar height for this purpose
it suffices to take the difference between the nasal and nasion-alveolar
point measurements.
2 Catalogue of human crania in the United States National Museum collections: The Eskimo, Alaska and
related Indians, northeastern Asiatics. Proc. U.S. Nat. Mus., vol. 63, art. 12, 51 pp., 1924.
3 Wistar Institute, Philadelphia, 1939. ‘
CATALOG OF HUMAN CRANIA—HRDLICKA 171
ORBITS
Orbital height —The maximum height between normal inferior and
superior borders, exclusive of any notches. Seldom perfectly vertical,
though near.
Orbital breadth.—The greatest breadth of the drbital lumen, from
the lacrimal point. It is only incidentally at exact right angle with
the height, though always near.
NOSE
Nasal height—Height from midpoint of line connecting lowest
parts of the borders of the two nasal notches, to nasion.
Nasal breadth—The maximum breadth of the nasal cavity.
UPPER ALVEOLAR PROCESS
The length of the arch is its anteroposterior diameter, in the median
line, from the prealveolar point to the midpoint of a line connecting
the posterior limits of the arch. These limits are the posterior
tuberosity of the arch on either side, or, when this is not developed,
the alveolo-palatine suture.
The breadth of the upper alveolar process is its breadth maximum,
obtained by applying the branches of the sliding compass, symme-
trically, to the greatest bulge of the process above the molar teeth.
SKULL CAPACITY
See pages 135-138 of ‘‘ Practical Anthropometry.”
THE LOWER JAW
Height at the symphysis —The height from the lowest median point
of the jaw, at the symphysis, to the lower alveolar point; the lower
alveolar point being the tip of the process of the bone between the
median incisors.
THE GROUPS AND TERRITORY INCLUDED
In addition to the Eskimo proper, it would be important to include
in this catalog measurements of crania of Alaskan peoples who, on
account of linguistic affinities, were hitherto classed with the Eskimo
but who now, with the present available skeletal remains, are recog-
nized as quite different. Furthermore, satisfactory data can now be
provided on two extinct groups of southwestern Alaska and on addi-
tional Alaskan Indians, all of which will permit for the first time a
definite view of both the older and the more recent population of
Alaska, which is one of the basic desiderata of American anthropology.
Unfortunately the costs forbid, so that the data on the non-Eskimo
Alaskan people and those on the Siberians must be left over for future
172 PROCEEDINGS OF THE NATIONAL MUSEUM Vou, 91
publication; but a few remarks concerning these groups will be useful
in these connections,
Of the physically non-Eskimo peoples of the coast and islands of
Alaska there are now known four groups, and it seems probable that
no Other larger units will be discovered. These are the Aleuts and the
Kodiak Island Koniags, with the Pre-Aleuts and Pre-Koniags unearthed
in our excavations; and there are the people of the eastern third of the
Alaska Peninsula, who are a mixture of the Eskimo and the Aleut.
As for the Indians, some additions are now possible from southern
and southeastern Alaska.
The statement that no further large ethnic unit is likely to be
discovered in Alaska should not be taken to mean that no other con-
tingents have ever passed through or along the Territory. It means
that no trace of occupancy by any such group has been discovered in
our general and intensive survey of the region. This survey covered
all the more important parts of the coasts, rivers, and islands, and it is
unlikely that evidence of occupancy by an additional physical or
cultural group was missed; the same applies to evidence of any really
ancient occupation. But the present shores of rivers, coasts, and
islands are far from where they were three, four, or more thousands
of years ago. Alaska is a land of living geology, with erosion every-
where very active. Banks and shores are constantly being cut or
undermined, and the silts and debris build new bars, shallows, islands,
and eventually flats. Yet man at all times in these parts has been
obliged to live close to the sea or on the banks of the larger streams,
and such settlements in the course of time have all had to be abandoned,
or else be ultimately cut away. These matters were discussed, with
some examples, in the report ‘Anthropological Survey in Alaska,”
already cited. What chance, under such circumstances, would there
be of a survival of evidence of any ancient human occupation? More-
over, as long as the road “‘toward the sun” was free, man would hardly
stop in the inhospitable Far North for any permanent or long-lasting
settlement. Thus, the absence of evidence in Alaska of human groups
other than those here mentioned cannot be a negation of the proba-
bility of other, older contingents of man having passed through; it
only emphasizes the fact that there is little possibility of their being
discovered.
The Eskimo territory, as is well known, spreads from Greenland and
Labrador in the east to the Alaska Peninsula in the west, skirting ev-
erywhere the seashores. The linguistic and cultural similarities over all
this region indicate that the spread of the group must be relatively
recent, and the close physical likenesses sustain this opinion. There
are some dialectic differences, but they do not show satisfactory lines
of demarcation. From place to place the Eskimo differ somewhat in
stature and even in head form, but with one exception there is no
CATALOG OF HUMAN CRANIA—HRDLICKA iva
possibility of subdividing them into distinct types. The arrangement
of the data given herein must therefore be merely geographical.
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CATALOG OF HUMAN CRANIA—HRDLICKA 419
ABSTRACT AND NOTES ON THE ESKIMO CRANIA
The preceding records relate to 11 larger geographical groups of the
true Eskimo, extending over almost their whole habitat. They in-
clude only fullbloods, i. e., unmixed with the white or the Negro.
Any specimen doubtful in this respect was excluded, but there were
very few such specimens. The material was almost wholly collected
by scientific workers.
The measurements show some local differences, but a close basic
similarity is evident throughout. Here, plainly, is a single physical
strain of the human family, differing only, as any other large and
widely scattered strain would, in secondary peculiarities. There is
nevertheless a possibility that the ancestry of the group was not
homogeneous but that it consisted of two related yet separate strains,
one with shorter head and face and one with longer; but it may be
possible also that such local differentiations as the group presents
were realized within itself, through inherent variability and segrega-
tion. Whatever may be the truth in this respect, it seems certain
that the mixture or changes took place not on the American continent
but well back in the original habitat of the people, which doubtless
was Arctic Asia.
Before proceeding with the results as shown by the preceding data
it will be useful to give what is now known of the statures of the people
in the various regions, and the mean bicondylar length of the femur.
Where the stature is not known or not known well, the length of the
femur gives a very good basis by which to gage the relative values of
the various measurements. This femoral length moreover gives an
excellent means for estimating the stature where it is not known
definitely. In all the Eskimo groups where both the stature and the
femoral length are known with some reliability the latter is close to
26 percent of the former. The available data follow; they are by
no means all that could be desired, yet they have a value.
420 PROCEEDINGS OF THE NATIONAL MUSEUM vOL. 91
Eskimo: Stature and mean ' bicondylar length of the femur
Male | Female
Group | ! 4
Length Ratio Length Ratio
Stature | of femur | (F=100) | St8ture | of femur | (F=100)
Western rivers and coasts and North- (202) (157) (36) (168) 9
east Boring Sea: ac 0, dene A es { wi7| 4209 } #5.0 { 151.0| 39.31 \ 26.0
: 66) (57)
Seward (peninsula 222202 eee eee { agen \ Been. ea =| eee { 39.82 } ha Pies We
(63) (80) (48) (51)
Sua Mawrence Island) = 23x28. Seyler! { 163.3 42.49 \ 26.0 { 151.3 38, 82 \ 25.7
: (13) (39) (13)
Point Hope... ---------------------------- { 166.5 43. 43 \ 26.1 |---------- { 40.55 \ -=-n an =--
: 33) (25)
Telogs nearvBarrowses ts ee ea | ee { yas \ PURGS TS Aer tere: { } ee aes sf
. 40. 31
: if (51) (8) (28)
Barrow region. ___---.---.---------------- \ 161.5 | 2 (42.45) } (26.3) { 153.6 | (40, a (26. 1)
North and northeast Alaska_______________ { ieee, } Ne 8 ANE eee aed { iene, } en ot Sef eat oe
(86) (3) = (62)
Greenland (all) .-------------------------- { 159.0 | (40.80) \ (26.7) { 153.0 } ~--2 2-22 -|------=---
1 Mean of the 2 sides.
2 Inadequate numbers.
THE CRANIAL INDEX
The importance and stability of the cranial index and the corre-
sponding cephalic index have been much overrated; nevertheless the
index is always of interest and help in racial studies. It is, of course,
only the percental relation of the cranial breadth and length, has no
bearing on the size of the skull, and must always be considered with
the height of the vault, which may completely change its significance.
The values of this index in the Eskimo, it was seen in the
General Abstract, were 70.3 to 77.4 in the males and 70.4 to 78.6 in
the females. As a rule it is somewhat higher in the females than in
the males, though in some of the groups the differences are small. It
does not, it will be seen below, harmonize wholly with territorial
sequence, and it presents one striking peculiarity, in the old “‘igloo”’
people near Barrow. It shows the highest values along the great
western Alaskan rivers, along the coasts and on most of the islands
of the Bering Sea, and at Point Hope; also in Hudson Bay and in
Smith Sound, which are not given in the abstract!; it is lowest in
the old “igloos’’ near Barrow, partly about Barrow itself, in Green-
land, and on most of the Seward Peninsula. Its means are abstracted
in the following table:
176.3 and 76.2; see detailed tables; and detailed data in author’s Anthropological Survey in Alaska,
46th Ann. Rep., Bur. Amer. Ethnol., pp. 259-260, 1930.
CATALOG OF HUMAN CRANIA—HRDLICKA ADN
Cranial index, in detail, by locality groups, west to east
Group Male | Female | Cua Ee Ano Male Female
Nushagak River___-_-------- (13) 78.9 | (20) 79.1 | St. Lawrence Island, Gam-]| (5) 72.8 |_..______-
Kuskokwim River, Upper-_-_-| (27) 75.2 | (86) 77.7 bell, Early
Kuskokwim River, Lower_._| (30) 78.3 | (21) 79.3 pee Lawrence Island and Pu- |(229) 76.9 |(249) 77.4
Mrkonun iver no (41) 76.9 | (63) 77.8
Bion sikgems te EE eS ee te (5) 78.6 | (7) 82.7 ines Tslandiess sees see (5) 78.8 | (6) 77.0
MAGEE ake ees eels (4) 78.5 | (4) 80.6 || Northeast Siberia_.___-___-_._| (18) 76.4 | (18) 77.6
Hoopen Baye: = 22-222. 255. (15) 78.9 Oe Wes) | Phot, Hope 2a saes senor ee (163) 75.6 |(118) 76.1
Nunivak Island_.-.--------- (46) 75.0 | (70) 76.3 || Old Igloos, near Barrow__---- (52) 70.3 | (44) 70.6
Nelson-lsland == s6t2 22.2222: (9) 77.2 | (17) 78.7 || Barrow (Utkiavik) -____.____- (33) 72.9 | (46) 74.0
St. Michael Island____---_--_- (8) 75.9 | (6) 75.5 || Piginik (near Barrow)_---_-- (2) 7388) | kan
Unalakleet__-___- GO etwas (9)) 765" || Point Barrow. ---3- 2 (49) 73.9 | (52) 74.4
Norton Bay_ =H CO) ) eacOi CLL aa OcOln || PINGRCE ake < 2/23 i eee eee (26) 74.9 | (28) 75.1
Golovin Bay-------- _| (16) 72.4 | (15) 73.6 || Northern groups (west of (5) 74.0 | (16) 74.0
Rooky Point__--_- -| (18) 74.3 | (27) 74.9 Hudson Bay)
Capes Derby and Nome____-| (5) 73.4 (6) 73.8 || Hudson Bay and Strait_____- (5) 75.1 (2) 77.5
Sledge slain eae (5) 71.7 (9) 74.4 || Southampton Island__ -| (10) 74.1 (4) 75.2
epelike wets oe eee ee (7) 75.1 | (16) 75.6 || Northeastern groups (west of (16) 73.1 | (17) 73.3
Por Clsrence®=< _--=— .--- =. (12) 74.5 | (13) 75.4 Greenland and Labrador)
MUGIGS St Vi a ee CO 2580 22) rood eS mith Sound ss 222 sees (7) 75:8.) (2) 76.7
IMetInTAvIK.: Deen 8 oa Ss (15) 73.5 | (26) 73.1 |} Greenland (mainiy north- | (52) 71.6 | (47) 71.9
Shishmareves. (2h 2. eke (17) 74.0 | (15) 74.8 west).
The above data are of considerable interest. Notwithstanding the
inadequacy of the numbers of specimens in many of the series, cer-
tain facts are quite evident. The cranial index differs regionally, and
the differences apparently are not insignificant. There are repre-
sented in the Eskimo, it seems, two related yet unequal strains, one
considerably to extremely dolichocranic, the other mesocranic. The
presence of the dolichoid variety in the earliest strata discovered so
far near Gambell, St. Lawrence Island, suggests that this strain might
have been the earlier; but the distribution of the two forms would
seem to incline to the opposite conclusion. The narrow type is found
in its greatest purity in the old “‘igloos’’ near Barrow,’ where the
mean cranial index in both sexes does not reach even 71 and indi-
vidually falls as low as 62; but it is also manifest in Greenland (and
Labrador *), more or less in the more eastern of the northern groups,
and in most localities on the Seward Peninsula. The mesocranic
strain, on the other hand, reaches in a large are from northeastern
Asia to the Alaska Peninsula, but it occurs also quite pure at Point
Hope, and it is probably somewhat mixed with the more oblong
type at the old settlement of Nixerak near Point Barrow, in some
localities about the Hudson Bay, and in Smith Sound. It is quite
probable that both the variants developed in prehistoric times, under
some territorial segregation, in the same stock, but the evidence
indicates that they were separate when they came to America, and
that while the broader-headed strain spread essentially southwest-
ward, the narrower extended mainly northward and then northeast-
ward.
Both the extreme narrow and the broader type are in all visual
and most other metric aspects true Eskimo and cannot be separated
as distinct racial components.
2 For details of these finds see Hrdlitka, ‘‘Anthropological Survey in Alaska,’’ p. 318.
In 34 male skulls 7/.8—Stewart.
422 PROCEEDINGS OF THE NATIONAL MUSEUM vou, 91
THE MEAN HEIGHT INDEX OF THE SKULL
The mean height index is the percental relation of the basion-bregma
height of the vault to the mean of its length and breadth. The use of
this mean is preferable to that of either of the single measurements,
because these stand in close compensation with each other and have
therefore but little if any individuality. The mean values of this
index in human groups range from approximately 76 to 88, in individ-
uals they reach both lower and higher. The Eskimo values are given
in the following table:
Eskimo: Mean height index in detail, by locality groups, west to east
Group Male Female Group Male Female
Nushagak River_..__._.*__. (13) 83.4 | (20) 84.1 || Gambell, early--_._-__._____ (5) 8$3.|Gn] pears
Kuskokwim River (30) 83.5 | (20) 82.3 || St. Lawrence Island and Punuk (206) 84.4 |(216) 84.5
Wukon River----_.2-= ____| (41) 84.9 (63) 83575||*Diomedeisland == 22ers (5) 83.9 (6) 84.7
Doplake se ote ea _.--| (5) 82.1 | (7) 82.0 || Northeastern Siberia----___- (17) 83.6 | (18) 84.7
Mam tra kee eet ae (6) 82.7 (6) 8254:)|) Point Hopes see 2228 Sa (160) 86.3 |(115) 85.2
HOODEH Bay nse ee ee (15) 84.1 (9) 83.8 || Old Teo near Barrow---_- (51) 85.8 e 86.3
INunivak island 23222" 2089 (46) 83.2 | (70) 83.4 || Barrow (Utkiavik)-_________ (25) 83.3 | (37) 83.3
INeISontslind eee ea en os (9) 82:0) (G6) $250 || Point Barrow222 22 ea (47) 84.7 | (52) 83.4
St. Michael Island__-________ (8) 86.2 (@)-84500)) Nixerak.Giss sl se eee (26) 83.5 | (24) 84.2
Urialakleet see = See (7) 84.0 (9) 83.4 || Northern groups (west of
INTOREOnED Byer eee eo 8 (6) 85.3 | (10) 82.7 Huidson; Bay) esses nese (5) s 1} (16) 82.3
Golovin Bay 222 pee (16) 85.7 | (15) 83.9 || Hudson Bay and Strait______ (CS) ASB eng eee
Rockyreomt 2.2 SL eee (18) 84.3 | (27) 84.6 || Southampton Island___-_____ (10) 85. 1 (4) 85.1
SledgevIsland=- =. oe (5) 85.8 (9) 83.6 || Northeastern groups (west of
Kovieruk2 ets) oe pre ee 2S (7) 84.9 | (16) 84.9 Greenland and Labrador)-} (16) 84.9 | (17) 84.0
IPOnbLOISTeEN Ces sees ee eee Gm) S8he57 83) P5145 Smithy Sounmdueese a eunis (7) 84.4 | (2) 85.8
Wisless ae 7e bail te the S92) (20) 86.4 | (22) 84.6 || Greenland (mainly north-
Metlatavikes = ore is (15) 86.9 | (24) 83.7 WESt) sso ee eee (52) 85.4 | (47) 84.8
Shishmiarev2= 2222 Balke se (16) 84.4 | (14) 84.5
The means of the index range only from 82 to 86.9, or approxi-
mately 5 points, and this would probably be reduced were the series
more adequate. With such a widespread habitat and such differ-
ences in the cranial index, this range is small. Moreover, but little
correlation is evident in the two indexes. The relatively broad-
headed southwestern Alaska groups are on the whole somewhat lower
than the rest, but this does not hold true for all the contingents nor
for those of similar type beyond that region. Among the narrow
headed the index in most is above its general mean, but here too there
are exceptions.
In general the mean height index of the Eskimo skull may be said
to range from somewhat submedium to above medium, with most of
the groups in the latter class. It is low in no part of their territory,
nor is it exceptionally high. It would not be a reliable means of
distinguishing the type of the skull as indicated by the cranial index.
CRANIAL MODULE (Ae)
The cranial module, or mean diameter, is a highly convenient and
valuable means of expressing the size of the skull; and it bears close
CATALOG OF HUMAN CRANIA—HRDLICKA 423
relation, though this differs in the two sexes,‘ to the size of the brain.
Throughout the habitat of the Eskimo the module shows good propor-
tions and a considerable similarity. There are some differences, but
these would probably diminish were all the localities represented
adequately. The female—male relation of the module, in the larger
groups, is also much alike. The details are given in the next table.
Eskimo: Cranial Module
Module Module
Group Tha ca ee ue Group == ue
Male | Female} tion Male | Female} tion
ye, (13) | (20) z (5)
Nushagak River---___- ss { 15.04] 14.55 } 96.7 || Gambell, early----------- { 15.11 } hese eles 2 ae
Nis 57) | (56)
Kuskowim River________ ( 95.4 || St. Lawrence and Pu- |f (206) (216)
\ Ma Mea) j nuk Islands { 15.42 | 14.83 \ 96.2
Yukon River-...----..-- 11634] 14°73 |} 98-9] (5)
TE. 2H | 22) lh os Pp aemete teen) ea 15.38 | 15.07 |t-------
gia nay ey Northeastern Siberia____- teins inte \ 95.4
2 ESET Eee 15.3] 1200 | 99/5. (160) | (115)
ees £5] 29 |} 93.9 Se are 15.43 | 1477 |f 958
Nelson Island________-.-- 15.59 | 14.64 : : (51) (43)
) (8) (6) Old igloos (near Barrow). 15.50 | 14. \ 94.7
ST aan { 15{30 | 14,72 |f---—-- eee (25) | _ (37)
Unalakl (7) |) Barrow (Utkiavik) ------ {1843 | aor |} 95.0
eee {15s | raged [fe : (47) | _ (52)
Reus £5] 29 | ee ae 144 | 14.75 |} 95
eae asnn--ae=- 15.48 Wea BO ace ttre (26) | (24)
ee nae. 10s [G9 | G9 1) gs.¢ Se ee Tat aac: 15-43 | 1480 |} 96-5
eee Vo reo Northern groups (west of { (5) (16) } ‘tae
ocky Point_.._..._____- { ie ae \ 96.2 || | Hudson Bay) 15.68 14, 63
Sledge Island-_._________ { sete chtaa ne \ ee gd Hudson Bay and Straits. { 15. 55 | (14. 57) \ -------
Kovieruk__......-.--_-_- { 5Q 29) } eins Southampton Island.._-- { eer | 15°13 } soatnee
(11) (13) Northeastern groups
OI as {15.47 | 1.71 |} 4 |] “ GwestotGroonlandand eucel eal nae
Wales { (20) (23) } 96.6 Labrador) :
eee eee 15.47 | 14.93 ate (7) (2)
Wetinia ik {09 | 2D Nh oy Bun ente cunts -2t---7e7 15,81 |(15.15) |f-------
pat nee ne ve ; Greenland ee { (52) (47) \ 94.5
. northwestern) ----_-_-_- 15. 52 14. 66 :
Sbhishmiaroys=- 3. =) os. -* { 15.24 | 14.77 \ 96.9
General Female : Male mean = approx. 95.6.
CRANIAL CAPACITY
The cranial capacity was taken by the method described in my
Anthropometry.’ Though I am convinced that this is the best
method devised so far, it is still laborious, time-consuming, and not
ideally satisfactory. It would almost seem desirable to replace the
measurement by that of the mean diameter, were it not for the fact
that it is a great and often a deciding factor in the sexing of the
skull. This is due to the fact that the relation of the capacity to
the module is in general markedly less in the female than in the male
cranium. There are exceptions, but they are infrequent. In the
males the capacity in cubic centimeters is near the module expressed
‘ See Hrdlitka, Practical Anthropometry, Wistar Institute, 1939.
5 Wistar Inst., 1920, 1939.
424 PROCEEDINGS OF THE NATIONAL MUSEUM vou. 91
in four figures; in the female it is farther from it. The difference in
the female may amount to as much as 200 units, which appears
never to be equaled in the male.
The cranial-capacity data on the Eskimo are given in the following
table. They show much similarity, which would doubtless be even
more striking were all the series fully adequate and equal in number.
There are, unfortunately, not yet enough data for racial com-
parisons.
Eskimo: Cranial capacity
Capacity in ce. Capacity module
F:-M relation
Group — relation
Male Female Male Female
B (18) (21)
Nerina See, eu eae eee \ 1 sm0.0 | 1,341.0 \ 90.2} 99.1 93.1
- 17 1
TITRE 24a Bt a a le eM: 1 4,480,0 | aann.0 |p 89-2. 97.6 91.2
Muniveaktiianae se okt) edie eins) So as 1 sue Ne aia ale) S0ea mene 90.8
Nelson Island.--___-_---------------- eg) ceca { 4 bosto Me asase alps oene)| aca 91.1
Sue Midhelislende ae a \ a adnior lie tases nity S525) | en 87.8
Sledge istaadh 2 WAN geese ee ete Seales eee a 90.0
- (19) )
yaleamereeers. be Ml Ler enw tee ee ae a) 472.0 | 1,3010 } 92.5) 95.8 91.2
15 20
IVIStintea Rts ee tS loci ier el CPN \gveroniela aunties 1ise8: |anumeeeae 91.8
St. Lawrence Island and Punuk____-- { ieee (a saa \ 91.1 95.0 90.0
"(126) |” (84)
Paint Eopes ee es Mente Bas Net wn Nbpee A) { Berea Ae \ 89.2 95.7 89.1
Southampton Island:.........02:2)) ween ition.) { ieee \ ne EASE Re Uda 95.8 roar
Greenland (mainl (45) (38)
: Venorthwestern)= 2. os: eS ee 1,527.0 | 1,295.0 84.8 98.4 88.3
Goeneralibiskimo means) ).\- se ee ee ea eT { 1 oe 1 Cah \ 89.0 97.5 90.5
It is regrettable that up to the present time we do not have similar
data on the White people, at least. There are fairly numerous data
on the capacity of the White and other crania, but they have been
taken by several differing methods and the results are not strictly com-
parable either with the records presented here or one with another.
What is plain from the above figures is that the Eskimo cranial
capacity, and hence the size of the brain, is by no means inferior to
the Whites, particularly when we consider that in general the Eskimo
are of decidedly lower stature than the Whites.
The female-male relation in the dimension is less than that in sta-
ture. Thus on St. Lawrence Island the relation in stature between
63 adult nonsenile males and females is 92.7, which is about identical
with that in Old White Americans (92.9); that in the Eskimo capacity
CATALOG OF HUMAN CRANIA—HRDLICKA 425
is but 89.0. Either the capacity in the males is relatively submedium,
or that in the females relatively above medium. There are indications
that would seem to sustain the latter deduction, but a real conclusion
is not yet possible
The relation in the two sexes between the capacity and the mean
cranial diameter is of special interest. This relation in all the groups,
and in man in general is distinctly lower in the females than in the
males. I have pointed this out on several occasions. The reason for
this is not yet clear, but it is doubtless connected with differences in
the two sexes in the development of certain parts of the brain—the
contents of the fossae and perhaps of the base of the brain—in the
two sexes.
FACIAL INDICES
The facial dimensions of the Eskimo are among the largest known in
human groups. and both the indices are rather high, indicating a
relatively high face. There are individual Eskimo crania, especially
on St. Lawrence Island, in which the face is very high indeed; but
there are also others in which the facial height relative to the great
breadth of the skull is moderate. The indices in our different groups
follow:
Eskimo: Facial indices
Facial Facial
index, Upper index, Upper
total tota
Group Group ae on Zz
Fe- Fe- Fe- Fe-
Male mine Male OND Male real Male asin
i (8)| (16)} (10) | (16) : (7 2)| (13 12
Nushagak River----.--- tore 91.0) 55.1 | 54.6 || Shishmarev___-.----.__- {sa a 0.7} red oy Z
Lower Kuskokwim-- -- ree oF oe ye) Gambell, early___-______ (aay Tera aa weet
s 3 (20)} (20)} (20) (22) || St. Lawrence and Pu- (41)| (55)| (198 ~ (200)
Upper Kuskokwim_-__- Nowe 91.6| 56.2 | 54.2 nuk Islands__.--_-_--- {ud 3 : : eas
Yukon River-.........-- Jarre | teen ee yilieern |) Diomede Island 0. 8 ta 1 aitigg>
(16)| (10)| (19) | (14) aa 3 3) (14 1
West Coast. ...-...---.- ist 8 89.0 35.0 of 6 Northeastern Siberia___- {3%} 1th Bel Bee
se 24)| (26)| (4 51 4 (27)| (22)! (138 98
Nunivak Island...-...-. ae 88. 2| 54.6 | 54.8 || Point Hope------------- as bral ae
Nelson Island._________- fase GO) 5) | 24 || old igloos near Barrow. feeee Ce ee
St. Michael Island _____- {censy en AO on Barrow (Utkiavik) ------|------]------ Moe ee)
(3)} (6) | (7 : 36 :
Unalakleet_-----.--...-- (988) (Oh, 4) TA 68, : Point Barrow-----------|------|--.-.. Wee a
4)| 5 ; 16
Norton Bay_--.--------. (954) ot as 65 . Nixerak.-_--------------|------ | woaoo- Baa ie
; 4 5 il Northern groups (west 5 11
Golovin Bay_--..------- {ott 86.3| 56.3 | 54.7 of Hudson Bay)______- } == 2-|------ yee es a
Rocky Point_.._.-_---- {95> | 4's! be'y | 56°=. || Hudson Bay and Straits. fone Sarena a Gan
(4)} (3)|_ (5) (7) || Southampton Island_.--|f (7)|_..___| (1 (
Sledge Island__-_-_------ {g0°2 (60.0) 60.2 BB, : ale AE {er yr Ba (61.6)
: or eastern groups
Kovieruk __-_.-..--.----|------ ‘Gey (51.9)} 54.0 (west of Greenland ae (5)} (12) | _(11)
Part Caceaicc (5) (10) (7) and Labrador__-______- : 86.6) 53.7 | 53.9
treqenm anes 91.9 |--____| 63.7 | 52.4 || goin sound j (6)} (2) (2) (2)
Wales {3 (17)} (17) | (22) Ree Ree 82.4 |(84.9)} 52.0 |(51.5)
Gar aw para ane ea To 89.7 a ry a Grodlid ey { (13)| _(4)} (48) | (40)
f Western) eee ee 87.2 85.0} 54.2 | 54.
Pcp aby Anon a=} -s—o= { (93. 2) 55.3 | 56.7 :
<p LLL CE MEET MEI Gaim ICUEE nnn niet nnn neon tener ne SS
416178—42——_17
426 PROCEEDINGS OF THE NATIONAL MUSEUM you. 91
FACIAL ANGLES
The method of taking the facial angles has been explained. One
measures the total facial protrusion, the other measures that of the
alveolar portion, which is somewhat independent. Both these angles
in the Eskimo show but a moderate protrusion of the face—more
than in the whites, about as much as in the Indian, decidely less than
in the Negro, the Melanesian, and the Australian. Direct racial
comparisons, regrettably, are not yet possible.
The total angle, it is seen, is much alike in the two sexes, but the
alveolar slant is appreciably greater in the females (narrower angle)
of nearly all the groups.
Eskimo: Facial angles
Angles, Alveolar Angles, Alveolar
facial facial
Group meen nee Group MWe hee Pad ey
Male| Fe- | Male| Fe- Male} Fe- | Male} Fe-
male male male male
Nushagak River-____----- (9)} (16)} _(9)} (16) |) Shishmarev-____________ { (13)} (13)} (13)} (43)
69.3} 69.7] 55.4) 53.8 67.9} 68.7| 54.2) 56.1
Lower Kuskokwim- ---- 8) en mnan Gambell, Early-_--_____- any EEN: a pa
, F : : : St. Lawrence and : ous
Upper Kuskokwim- ---- (19)} (20)} (9)} (20) (184)} (184)} (184)| (184)
ne { 68.3| 67.7| 56.1| 51.6 Punuk Island__.___-_- { 67.6| 68.0| 57. 7| 54.5
NVAUKONURLVelass= 2 soe (33)| (51)| (83)] (51) |; Diomede Island__-_-__---- (4) (5) (4) (5)
{ 69.7] 67.9) 55.9) 52.8 68.0) 69.1 i etal
iWresti@oaste sss sees (19); (44)} (19)| (14) |} Northeastern Siberia_.__|f (12)] (15)| (12)} (15)
68.4) 67.5} 55.7) 54.1 66.9) 66.8) 55.8) 53.4
Nunivak Island_.______- { (41)| (45)| (41)] (45) Point Hopes sens a2 sae2 { (128)] (95)} (128)} (95)
68.0} 67.5} 58.0} 55.0 69.9] 69.8 6.0) 55.3
Nelson Island_____------ (7)| (18) (7)| (43) || Old Igloos, near Barrow-|f (39)| (31)| (39)] (81)
66.0} 66.5} 53.0} 50.0 70.0} 69.9] 56.2} 55.2
St. Michael Island - -.___- { (7) (3) (7) (3) Barrow (Utkiavik)--___- (11)} (6)} (1)} (6)
69. 0|(71..0)} 56. 5} (57. 0) 70.0) 71.3} 58.8) 55.6
Unalakleets 2 2o cst eee (6) (6) (6) (6) Point Barrowee- seen (36)| (37)| (86)} (37)
68.8) 67.4) 59.3) 55.2 69.0} 69.0] 55.9) 55.0
Norton Bay----------..- Neo 3 a fn INU ere keene ee Gs) Ga Gs) 2)
: : ; : : Northern groups (west . ; aS 3
Baas Geo Greal $6.3] 52.6. || of Hudson Bay)... 6 o| O&O| B50 SLO
Rocky, EoMts este (13)} (19)} (18)] (19) || Hudson Bay and Straits_ { (5) (2) (5) (2)
68.1) 67.6] 55.9} 54.0 8.9] (71.7)| 54. 0} (55. 0)
Sledge Island__--______-- (5) (6) (5 (6) || Southampton Island --__ { (9) 3) (9) 3)
71.1] 68.6} 60.1} 52.1 69. 3](69.7)| 53. 9} (54. 0)
AKO VICK Keenan soe oe a eee ea ip wilees (12) || Northeastern groups
70.1 54. 4 (west of Greenland (12)} (12)} (412)] (12)
Port Clarence__-_-_----- oo (6) G0) Prey and Labrador)_______- 70.0} 69.5} 67.0} 51.7
9.2; 70. : i
Winlesaece ee 2 Pele at { aa S (17 12) Smith/Sound sss sess = Ao en { 1m Bee AL.
a 56. 3 : . é
: P Greenland (mainly 7
Motlatavite 2.0.82. kee (11)} (8)} (o)} (18) (46)} (40)} (46)| (40
{ae 67.5| 54.4] 54.0 northwestern) ...----_- { bor6 69.9] 55.2] 53.0)
THE ORBITS
The orbits of the Eskimo skulls are absolutely large. The orbital
index is fairly high, but not extraordinary. ‘The index in the females,
as usual, is somewhat higher than that in males (approximately as
101.5is to 100). The variation according to locality is very moderate.
CATALOG OF HUMAN CRANIA—HRDLICKA 427
Eskimo: Orbital index
Orbital index Orbital index
Group aaa Group eae
Male Female Male - Female
Nushagak River_____-.-__ (11) 88.7 (17) 92.2 || St. Lawrence and Punuk
Lower Kuskokwim___-_-___ (28) 88.9 (17) 89.1 Tslandes== see he set (211) 91.4 |(204) 91.7
Upper Kuskokwim___-____ (23) 88.7 (31) 89.2 |} Diomede Island BAe see ie (5) 89.0 (5) 90.5
NWakon iver =-—2 02) 2 2-= (40) 91.3 (57) 93.1 || Northeast Siberia_________ (18) 91.0 (18) 90.9
Vike (Ol S jee (22) 91.3 (8) Pole orlivPomnt Hopes e.—-- a e (149) 89.7 | (106) 90.5
Nunivak Island_________- (42) 89.2 (59) 90.9 || Old igloos, near Barrow__| (47) 90.2 (33) 92.1
NWelsompisiand: 235 55s) f (9) 92.0 (15) 91.9 |} Barrow (Utkiavik) -_-___- (18) 87.5 (19) 91.5
St. Michael Island________ (8) 88.3 (5) 95.5 || Point Barrow. -_--------- (48) 89.9 (42) 90.8
Orrinkieepeeee von soe (6) 88.3 (9) 90857 || Nikerakc# Mee eer oe (22) 89.9 (16) 90.2
INorton| Bays 22885 8 (6) 91.8 (9) 92.2 || Northern groups (west of
GoloviniBay_ 2 --22-2--- (16) 89.6 (14) 90.8 Hudson Bay)---_-_--- See (5) 90.5} (16) 91.3
Rocky: Romb. 5 2-2-2 - = 2 (17) 89.1 (24) 91.2 || Hudson Bay and Straits_- (5) 90.0 | (2) (94.7)
Sledge Island____________- (5) 89.0 (7) 89.8 || Southampton Islanad______| (10) 90.0 (3) (93.7)
OWAGH Re ee ee ae (6) 89.9 (16) 90.5 || Northeast groups (west of |
Port pctaerice pe Re (10) 88.6 (10) 89.0 Greenland and Labra-
bVVinles Stee cee ee (20) 89.7 (23) 89.2 Gor) Geet wee eee ee (15) 88.8 (13) 91.1
Metlatavik SEES ee ee (12) 92.3 (21)79229) |i SnithSound= 42) (7) 86.7 (2) (88. 6)
Shishmarev- --.—-..---.-- (15) 91.6 (15) 89.1 || Greenland (mainly north-
Gambell, early. _......_-- (EST |= =a ees WES) 2 ten (49) 91.7 (43) 91.8
NASAL INDEX
The nasal index in the Eskimo is decidedly low. It averages.
in general, approximately 43.8 in the males and 45.3 in the females
and presents much similarity all over the territory occupied by these
people. Nevertheless, in general it is higher (nose broader) in the
southwestern and St. Lawrence Island contingents than it is farther
north. Because of their relatively lower nasal height it is larger in
the females than in the males, in the proportion of approximately
103): 100:
Eskimo: Nasal index
Group Male Female Group Male Female
Nushagak River_-.-_.------- (12) 44.8 | (18) 47.6 || St. Lawrence and Punuk
Lower Kuskokwim River---_| (29) 44.1 | (17) 48.4 Slang: oS 35ers cae =|(220) 45.3 |(214) 46.6
Upper Kuskokwim Shee. (26) 45.7 | (31) 48.4 || Diomede Island __ 5) 44.6 (5) 44.0
Yukon River_____- (41) 44.2 | (62) 46.1 || Northeast Siberia (18) 45.7 | (18) 48.0
West Coast _._-_--- (22) 44.3 | (20) 46.3 || Point Hope_-_-_-_- bee cess 68). 44.7 CD) 6452
Nunivak Island_ (44) 48.8 | (63) 46.4 || Old igloos, near Barrow_____| (52) 43.4 | (39) 45.2
Nelson Island______ (9) 43.0 | (14) 46.3 || Barrow (Utkiavik)_--__-__- (17) 44.6 | (23) 43.3
St. Michael Island___ (8) 42.1 (6) 43:8 || Point Barrow_.--_-2_-_-__- (46) 42.2 | (46) 44.9
Onsigkipare eee ee (6) 42.3 (S) R442 Nixerak.< eee ie: soe ee (22) 43.9 | (18) 44.2
Norbonub ay i: eaves! Sar (6) 42.9 | (10) 44.7 || North groups (west of
Golovin Baye oe er Ss (16) 42.8 | (13) 45.8 Hudson! Bay) hee 22 se (5) 42.6 | (15) 41.3
Rocky Romts =e eo (17) 48.0 | (25) 44.3 || Hudson Bay and Straits____| (5) 45.3 (2) (43.9)
Sledge Island=.* 22 3-6 2 222 (5) 42.3 (7) 44.3 || Southampton Island________ (10) 42.6 (8) (43. 8)
WO VIClU Kees Sas see sae (7) 45.9 | (15) 45.8 || Northeast groups (west of :
Port iGltarance:< 22) 2:5) 4. Fe (11) 43.8 (9) 47.7 Greenland and Labra-
Bis pain seb fis ete (20) 44.6 | (23) 46.1 GOT) be see (16) 48.4 | (18) 44.3
Minilatavik. ts 2 eee Se (42), 4352) (22) 442) | Smith,Sounde_ 22 2-8 2 (7) 39.7 | (2) (48.9)
Shishmarev____.______-_-----| (15) 44.2 | (15) 46.4 || Greenland (mainly north-
Gambell early.) 222 552 eS (Bb) 546795 | oe ees WeSt) =. a2 eee 2b ee | (51) 43.3 | (45) 43.9
UPPER ALVEOLAR ARCH
This index in the Eskimo is but moderate, owing largely to the
considerable breadth of the arch. It averages approximately 84.4 in
the males and 84 in the females, and the range of the means of the
428 PROCEEDINGS OF THE NATIONAL MUSEUM you. 91
different localities is small. As with the orbital and nasal indices, it
is higher in the females than in the males, owing mainly to a slightly
greater relative breadth of the female arch; but the excess is slight,
the female-male proportion being approximately 100.7:100. The
point of principal interest in this connection is the large absolute size
of the arch.
Eskimo: Index of the upper alveolar arch
Group Male Female Group Male Female
Nushagak River-_-_.------ (10) 83.2] (15) 84.2 St. Lawrence and Punuk
Lower Kuskowim_-_----_- (21) 82.9] (13) 83.8 Islands!) Vases acre $3.1 |(182) 82.8
Upper Kuskowim_------- (21) 83.5} (24) 86.1 Diomede Island ___ (5) 83.3 (3) (79.8)
Yukon River (32) 82.8] (55) 85.0 Northeast Siberia_- S 84.7 | (15) 84.7
West Coast___-_-_- 1 R (13) 82.5 PointpHopeteee eee ( 84.7 | (93) 84.7
Nunivak Island_-_------- (44) 83.4 | (46) 85.4 Old igloos, near Barrow_-| (39) 84.3 | (83) 85.7
Nelson Island_-_-___--...-- (8) 85.8} (14) 85.3 Barrow (Utkiavik)__--_-- 8) 85.9] (18) 848
St. Michael Island_____--- (7) 82.1 (8) (86:6); |) Point Barrow2-22-=--2--. (33) 86.9 | (33) 87.4
Wnalakleet 2. ise Se (6) 82.6] (7) 84.1 INixerak: 22 see ee eee (11) 87.0 | (13) 3
INOFtOnGB BY =--25-— aoe (4) 84.2 (7) 86.8 Northern groups (west of
Golovin/Bay-)-= = (15) 85.1] (10) 85.7 HudsoniBay)eas- se (4) 86.6 | (11) 84.
Rocky Otis ensue secon (10) 84.1] (21) 85.5 Hudson Bay and Straits__| (5) 86.0 (2) (82.9)
Sledge Islande--2-22-- == (5) 83.3 (6) 86.9 Southampton Island___._| (10) 83.5 (3) (82.9)
USOVAGTU Ks So ees ee (3) (84.0)} (13) 85.3 Northeast groups
Port: Clarentes: 22 ssn =o (9) 83.9 (6) 83.3 (west of Greenland and
Wiles betaedice® Bio (18) 83.8] (21) 84.9 Labrador) Sissel (11) 83.8 | (12) 87.6
WMetiatavikss- 2222 52s (11) 83.9 | (12) 88.2 Smith Sounds ane (7) 81.6 (2) (83.9)
Shishmareayoosee. e (12) 84.9] (11) 83.8 Greenland (mainly north-
; 83.
Gamibell; early. 22222 2-2- (GQ) Res teae. oe eee ‘WESt) 22 hy eee (47) 85.5 | (40) 86.5
JUVENILE ESKIMO CRANIA
For the first time in our studies of the Eskimo, in fact for the first
time in the study of any American group or any other human group
except possibly that of the Whites, it is possible to present data on a
large series of juvenile skulls. From the inception of my work in
Alaska I made it a point to collect all such skulls (and skeletons) in
good state of preservation, and with additions from some of our other
expeditions we have gathered the 80 specimens here reported upon.
An additional similar report will also be possible on juvenile crania
from the Kodiak Island and the Aleutian chain.
The specimens are of different ages, from about 3 months after
birth to 19 years. The ages have been estimated from the denture
and often from other parts of the skeleton. There is not enough in any
age category for satisfactory conclusions, but the data give some clear
indications, and they are supplemented by records on adult skulls
from the same regions. Sex identification has been added only where
very palpable.
Cranial indez—This index is decidedly higher in the young, in
every subdivision; but lower indices occur individually from as early
as the first year.
Mean-height index.—This index, conversely to the cranial, is evi-
dently relatively low at birth and it gradually rises with age, but it may
individually in later childhood reach or even surpass the adult mean.
CATALOG OF HUMAN CRANIA—HRDLICKA 429
The opposed behavior in the young of the two indices indicates that
the growth of the skull during this period is relatively greater in its
length and height than in its breadth. The probable cause of this is
the restraining effect on the breadth of the temporal muscles.
Facial indices.—In the Eskimo infant the face is relatively low and
as a result so are the facial indices; but from the second year the
relative proportions of the face approach those of the adult.
Facial angles.—Facial and alveolar protrusion, low in the infant,
gradually increases with age, the angles correspondingly growing less
obtuse. The cause, of course, is the development of the dental
apparatus.
Orbital index.—In general in juvenile Eskimo skulls this index is
very perceptibly higher than it is in the adults; and there is no definite
regression in it up to adolescence and even later. After that it is
doubtless influenced, especially in the males, by the development of
the supraorbital region.
Nasal index.—The nose in the young is relatively short but grad-
ually grows longer; the nasal index correspondingly is higher at first
but gradually, in general, becomes lower. As in all other characters
there are some individual exceptions.
Dental arch.—The dental arch in the young is defective posteriorly
and so cannot well be compared with that of the adults. Itis especially
short in the infant, giving low index; but from childhood on its relative
dimensions show no clear-cut difference from those in the adult.
General.— The present available data show that the Eskimo infant
is characterized by the following conditions, as contrasted with the
adult:
. Relatively its head is markedly broader;
. The vault is relatively lower;
. The face is relatively shorter, its indices lower;
. Facial protrusion is lesser, facial and alveolar angles more obtuse;
. The orbits are relatively higher, their index higher;
. The nose is relatively lower, its index higher; and
. The dental arch is relatively shorter and its index is lower.
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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM
SMITHSONIAN INSTITUTION
U. S. NATIONAL MUSEUM
Vol. 91 Washington: 1942 No. 3132
THE SPECIES OF AEGLA, ENDEMIC SOUTH AMERICAN
FRESH-WATER CRUSTACEANS?
By Watpo L. Scumirr
Wwe ty distributed throughout the greater part of temperate South
America from about latitude 20°30’ S. (Franca, Sao Paulo, Brazil)
to latitude 40° 28’ S. (Abtao, Llanquihue, Chile) is the unique, ende-
mic genus of fresh-water decapod Crustacea known as Aegla (family
Aeglidae). Its nearest relatives are marine and probably to be found
somewhere among the galatheids (tribe Galatheidea). There are no
fresh-water Crustacea at all like Aegla anywhere else in the world.
Most authorities have believed the genus monotypic—genotype, A.
laevis (Latreille), 1818 (pl. 308, fig. 2). In so doing they certainly
must have considered differences that are at times rather marked
between specimens from widely separated places, or in some instances
from the same locality, as variations of no great importance, or else
were possessed of altogether too little material to be able to evaluate
it properly. Carlos Moreira (1901), at the time a member of the
zoological staff of the Museu Nacional, Rio de Janeiro, Brazil, was
the first to dissent, insisting and, indeed, demonstrating that at least
the species described by Fritz Miiller (1876) as A. odebrechtii was
distinct from A. daevis. For his Brazilian specimens, regrettably,
Moreira employed the name Aegla intermedia, which had been given a
1This paper was first presented as an illustrated address, entitled “Some Remarks on
the Endemic South American Freshwater Crustacean Aegla laevis (Latreille),’ before
Section II, Biological Sciences, of the Eighth American Scientific Congress, Washington,
May 16, 1940. An abstract of this address appears in the Proceedings of that Congress,
vol. 3, p. 491, 1942.
431
432 PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 91
Chilean species by Girard (1855, p. 255) and which species, by the way,
seems never to have been taken again.
On my first visit to South America, in the fall of 1926, under the
auspices of the Walter Rathbone Bacon Scholarship of the Smith-
sonian Institution, I planned to obtain additional specimens of A.
odebrechtii. I thought I was successful at Castro, Parana, Brazil,
but the specimens I got there, however much they may superficially
resemble A. odebrechtii, are another species (castro), named in this
paper.
En route to Castro, I stopped in Rio Negro. Here, with the help
of Carlos Zornig, at whose hotel I stayed, and with baited wicker
fish traps that he provided, I caught several large Aeglas. One of
these is the largest representative of the genus ever to be taken,
measuring approximately 44 mm. in length of carapace and rostrum
together. It is the type of the species parana, which I am naming
for the State in which it was found.
Although I was chiefly interested in procuring marine decapods at
the time, I did not neglect looking for Aeglas as opportunities arose.
In that verdant park, the Prado, at Montevideo, Uruguay, Juan
Tremoleras and I collected a lot of small Aeglas from one of the
smaller watercourses. These, too, proved new, and are named prado
in commemoration of the place and occasion of their capture.
When Dr. Martin Doello-Jurado, director of the Museu Argentino
de Ciencias Naturales, learned of my interest in Aegla, he most gen-
erously took me on an all-day excursion to the delightful resort of
Tigre. Here numbers of smaller specimens of a hitherto unrecog-
nized species were found. This species (wruguayana), however, I
have described from a larger, more fully developed specimen from San
Carlos, Uruguay, belonging to the Field Museum of Natural History
in Chicago. Dr. Doello-Jurado also kindly granted a loan of his
museum’s collection of these crustaceans. Without this great help,
this paper could scarcely have been written, for in that fine collection,
along with representatives of several other species, are the holotypes
of four of the new species herein described: sanlorenzo, jujuyana,
afines, and humahuaca.
At Concepcion, Chile, January 1927, the director of the Concepcion
Museum, Dr. Carlos Oliver Schneider, Carl Junge, and I made a very
successful haul of Aeglas on the outskirts of town. These formed
the basis of A. concepcionensis.
In the course of an examination of the crustacean collections of
the Field Museum, two new species of Aegla were located, one
(papudo) from Papudo, Chile, and one (wruguayana) from San
Carlos, Uruguay, a species already referred to above.
THE SPECIES OF AEGLA:-—SCHMITT 433
The Museum of Comparative Zoology, Cambridge, Mass., through
the kindness of Dr. Fenner A. Chace, Jr., also lent me all their Aeglas
for study. One specimen of a lot from Santiago, Chile, was selected
as the neotype of A. /aevis. There is no certainty that the original
type is extant or in the Paris Museum, where it was believed to have
been deposited. Another specimen, from Talcahuano, Chile, has
been made the type of a new subspecies of A. laevis bearing the
subspecific name tal/cahuano.
From the late Dr. Carl H. Eigenmann, of the University of Indi-
ana, the National Museum received certain Chilean Crustacea, which
included a new species, A. abtao, and several specimens of the long-
lost A. denticulata of Nicolet.
In the type collections of the United States National Museum, in
addition to A. castro, parana, prado, odebrechtii (neotype), con-
cepcionensis, and abtao, there are the types of five other new forms:
A. platensis, franca, odebrechtii paulensis, neuquensis, and
riolimayana.
The late Dr. Florentino Felippone, of Montevideo, contributed
specimens of Aegla from Uruguay to the United States National
Museum collections on several occasions, as did also Alberto Tre-
moleras, of the same city. Finally, I received additional very help-
ful material from Dr. Carlos E. Porter, of Santiago, collected in
part by Dr. A. Santa Cruz, of Concepcion, Chile; from Dr. Carlos
Moreira, of Rio de Janeiro, collected by Dr. G. Kuhlmann at Blu-
menau, Santa Catharina, Brazil; and from Dr. Paulo Sawaya, of the
University of Sao Paulo.
Through the kindness of Henry W. Fowler, of the Academy of
Natural Sciences of Philadelphia, and G. Ayres Coventry, research
associate in charge of Crustacea, I had the opportunity of examining
seven Aeglas (four lots) contained in the Academy’s collections: (1)
Three females collected by “Dr. Wilson” in Chile, which proved to be
A. papudo; (2) two females of A. daevis received years ago from the
Smithsonian Institution, for which regrettably there are no locality
data or any record at the Institution of this particular sending; (3)
a dried specimen of what is unmistakably A. odebrechtui, “du Brésil.
Donni par M. M. Derreaux”; and (4) one of Dana’s Wilkes Exploring
Expedition Aeglas with an original printed Expedition label filled out
presumably by Dana himself—“Aeglea laevis. Chili.”
I am immeasurably indebted to the Walter Rathbone Bacon
Scholarship of the Smithsonian Institution, which enabled me to visit
South America personally to collect some of the specimens upon
which this paper is based and to establish the many helpful contacts
that made it possible to gather the most comprehensive representation
of the genus Aeg/a that has ever been in anyone’s hands for study at
434 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
one time. I am also deeply grateful to the many good friends and
scientific institutions who helped me with specimens, pertinent in-
formation, facilities of various kinds, and assistance in the field and
otherwise. Most, if not all, of these are mentioned either in the
foregoing recapitulation or in the following text.
The manuscript was helpfully criticized and typed by my secre-
tary, Miss Lucile McCain. The drawings are the work of Mrs. Aime
Awl, staff artist to the department of biology of the United States
National Museum. The photographs and prints were made by
Gurney I. Hightower and F. B. Kestner, of the Museum’s photo-
graphic staff. I am also indebted to Dr. Olga Hartman, of the Allan
Hancock Foundation, and Dr. Walter Weymouth, of Stanford
University, for some very helpful suggestions.
HISTORICAL REVIEW
In 1818 (pl. 308, fig. 2) Latreille figured, without description, a
new crustacean to which he gave the name Galathea laevis, perhaps
unaware that his species was from fresh water and that the genus
in which he placed it was exclusively marine. Not more than two
years later Leach (1820 [1821], p. 49) quite correctly observed that
Latreille’s species represented not only a new species, but a new
genus as well. This he named Aegla.
According to Dr. R. A. Philippi (1894, p. 372 [p. 4 of sep.]), and
the late Edwyn C. Reed in a letter to Dr. Mary J. Rathbun dated
June 6, 1895, a crustacean of this type was recognized (but not de-
scribed) as early as 1782 (pp. 206, 347; 1789, p. 182) by Molina in his
“Saggio sulla Storia Naturale del Chile” as Cancer apancora.
So far as I am aware, it was Desmarest (1825, p. 187, pl. 33, fig. 2)
who, without contributing any additional information, introduced the
incorrect spelling of the generic name, Aeglea, which all subsequent
authors, except Dr. Mary J. Rathbun (1910, p. 602), seem to have
followed, even Latreille (1829, p. 84) himself. Miss Rathbun, how-
ever, called attention to the fact that Leach spelled the name Aegla,
not Aeglea.
The figure of Aegla laevis that Desmarest published along with
his brief description is very similar to Latreille’s, yet in some re-
spects it is different enough in the shape of the chelae and in the
addition of orbital spines to have been taken from some other speci-
men. If based on the same specimen, Desmarest’s is the better figure.
Both Leach and Desmarest state that the material upon which their
remarks were based was to be found in the collections of the Paris
Museum. Neither made mention of a locality. There is now no
specimen in that museum that can be definitely linked with either of
these authors, or with Latreille, for that matter, unless, as I am
informed by Dr. Louis Fage, of the Laboratoire de Zoologie (Vers
THE SPECIES OF AEGLA—SCHMITT 435
et Crustacés), Muséu National d’Histoire Naturelle, Paris, it might
be a very old, dried specimen carrying the name A. laevis without
other data.
Griffith (1833, p. 184, pl. 7, fig. 2), who, in his “Animal Kingdom
of Cuvier,” supplied a colored illustration of A. Jaevis, which appears
to be a crudely done, reversed reproduction of Desmarest’s figure,
adds nothing in the way of a locality or specific characters to the
still scanty knowledge of this crustacean.
Tn his classic “Histoire Naturelle des Crustacés,” H. Milne-Edwards
(1837, p. 258) gave a rather extensive discussion of the genus, and a
concise description of the species, which, however, is of no more than
generic value today. Also, he is the first to give the species a home:
“Habite les cétes du Chili.”
The “Disciples Edition” of Cuvier’s “Le Régne Animal” (1837,7 p.
124, pl. 47, fig. 3) has an Aegla in color, together with some details
in black and white, that is quite different from the figures that ante-
dated it. The Paris Museum may have come into possession of better
material of what was taken to be A. laevis, but it is difficult to believe
that this particular drawing could have been based on the original
type, for, in spite of its more natural appearance the lateral margin
of the anterior portion of the carapace is most certainly not accurate,
no matter what the species represented may actually be.
The “aeglée lisse” of these several authors next appears as “Aeglea
laevigata” in H. Milne-Edwards and Lucas’ account (1843 [1844],
p. 34) of the Crustacea of d’Orbigny’s “Voyage dans l’Amérique
Méridionale,” surely an unintentional mistranslation of the French
common name of what was known in the scientific literature of the
day as A. laevis.
It may be that all the foregoing records were based on the same
species, but it was given to Nicolet (1849, p. 200; Atlas, pl. 2, fig. 1)
to add a second and unmistakably new species to the genus, A. dentz-
culata, in Gay’s monumental “Historia Fisica y Politica de Chile.”
His well-characterized and distinctively figured species is readily iden-
tifiable. On the other hand, his description of A. /aevis, which he
unfortunately did not figure, leaves much to be desired. It cannot
be distinguished from any of the species, except A. denticulata, now
known to inhabit Chile. Nicolet’s A. denticulata was so at variance
with what most authors, myself included, thought a species of Aegla
could possibly look like, that it always was believed to have been
4Jn a little note seeking to establish the date of issue of the crustacean plates of Cuvier’s
“Le Régne Animal” (Disciples Edition) I stated (1937, p. 151) that no reference to this
particular edition was to be found in the second volume of Milne-Edwards’ ‘Histoire
Naturelle des Crustacés” (1837). In the course of reviewing the history of Aegla laevis, I
find that I was mistaken and that a number of the Disciples Edition plates are cited in
that volume. This oversight in no way invalidates my contention that the date of the
crustacean plates in the Disciples Edition should be 1837.
436 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
incorrectly figured and described. The most surprising thing about
it, however, is that a so strikingly different Aegla eluded rediscovery
for so long a time. Specimens taken by the late Dr. C. H. Eigen-
mann at Osorno, Chile, in 1919 have at last enabled me to establish
the validity of Nicolet’s species 93 years after its original description.
In April 1889, the United States Exploring Expedition secured a
number of Aeglas “in shallow fresh water streams, [in] Chili, from
beyond the Cuesto del Prado, on the road from Valparaiso to Santi-
ago, sixty miles from the sea; abundant, swimming generally over the
bottom.” Dana (1852, p. 476; Atlas, 1855, pl. 30, fig. 6a-f) de-
termined, redescribed, and figured these specimens as A. laevis, yet
they cannot safely be assigned to any of the known species of the
genus, as the fingers of the chelae as figured are without a lobular
tooth on their prehensile margins; the general appearance of the
palmar crest and the lack of a definite or spiny lobe on the outer
margin of the movable finger near the base suggest A. lacus
talcahwano.
| After the foregoing paragraph had been type-set I had the oppor-
tunity of examining one of Dana’s original specimens as noted above
(p. 483). It is identical with what I have redescribed as true A.
laevis. Except for its somewhat smaller size, 9 mm. less in length
of carapace and rostrum taken together, it might have been the speci-
men figured by Dana. His drawing seems to have been a little hastily
done, for the rostrum is too slender and sharp, and the hands are not
very well drawn. This particular specimen distinctly shows a well-
formed lobular tooth on the prehensile margin of the fixed finger of
each hand and a definite, though small, spined lobe near the base of
the outer margin of each movable finger. |
A third species, A. intermedia, was proposed by Girard (1855,
p. 255) in his report on the Crustacea of the United States Naval
Astronomical Expedition. A discussion of the genus preceded a list-
ing of the two previously described species, A. laevis and A. denticu-
lata, and his description of the new one. This description does not
supply enough detailed information to permit the keying out of his
from the other species of Aegla. I have therefore not dealt with
Girard’s species beyond this brief mention and on page 431 and page
448, footnote. Some day it may be found again at the type locality,
“the upper affluents of the Rio de Maypu, 2,000 feet above the level
of the sea, near Santiago [Chile],” and perhaps be recognized by the
second row of spines on the carpus of the cheliped. Such a second
row of spines occurs in A. denticulata but not in any of the other
known Chilean species, but the marginal toothing of the posterior
portion of the carapace at once sets the two apart. If Girard’s A.
intermedia had possessed such toothing, surely he could not have
failed to see or mention it.
THE SPECIES OF AEGLA—SCHMITT 437
Heller’s report (1868, p. 81) on the Vovara Crustacea has A. laevis
as being represented in the material collected in “Chili.” Up to and
including Heller’s report, Aeg/a had been recorded only from Chile.
The very first records from any other part of South America are
those of von Martens (1868, p. 26; 1869, p. 14). He had specimens
from Rio Grande do Sul, Brazil, Rodersberg, and Porto Alegre, some
of which had been collected as early as 1831. Unless specimens are
extant and in good condition, it will be impossible to determine just
what von Martens, or, indeed, almost every other author cited in this
paper, took to be A. laevis.
The next record from Brazil is that of Fritz Miiller (1876, p. 13).
He described a unique species from the Serra do Mar, between the
headwaters of the Itajahy and the Rio das Marombas, in the State
of Santa Catharina, under the name of A. odebrechtii. His species,
like A. denticulata, by virtue of its illustration and excellent descrip-
tion, was easily recognizable on rediscovery (see Moreira, 1901; p. 489
of this résumé; also p. 481 above).
This same year Lucas (1876, p. ex) announced the discovery of
A. “laevis” in Argentina from the Rio de la Plata. He said that
on the tidal flats of the estuary, which are exposed at low tide, and
where the water is quite fresh because of the great distance from the
sea, this crustacean is found in prodigious numbers under slightly
embedded rocks, shingle, pebbles, remains of shells, and detritus of
all kinds, and that it is much sought after for food by the inhabitants,
with whom, in this part of South America, it occupies the place held
by the crayfish in Europe. Some time later (1891, p. lxxxix), Lucas
received specimens from the Rio Mendoza in the Argentine Cor-
dillera at an elevation of from 1,800 to 2,000 meters.
Scarcely six months thereafter Wierzejski (1892, p. 15 [1893, p.
232, 243]) obtained A. “laevis” from the environs of the city of
Mendoza, in the province of the same name. Wihierzejski’s paper,
perhaps because it was published in Polish, escaped notice until he
(1897, p. 1) furnished a German translation of the portion dealing
with Aegla, in order to correct Nobili’s impression (1896) (below,
p. 438) of being the first to report Aegla from the Argentine. Wier-
zejski’s remarks, in part, are here translated again, this time some-
what freely into English: “Associated with [the fresh-water amphi-
pod] Hyalella inermis in one of the streams discharging from one
of the larger lagunas in the vicinity of [the city of] Mendoza. In
life apparently dark blue; alcoholic specimens are dorsally bluish
gray, ventrally reddish. So far as I can ascertain from the de-
scription of Professor Martens, there are no appreciable differences
between the Argentine form and those from Chile and Brazil which
were described by Milne-Edwards and Dana. The largest specimens
measure 7 cm. in length and 1.7 cm. in width; the natives gather this
438 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
crustacean for culinary purposes. Hitherto, it was known only from
the streams in virgin forests in Chile and Brazil. Martens regarded
it as an endemic South American species.”
In 1892 (p. cevi) Berg corroborated Lucas’ (1876; 1891) observa-
tions on the occurrence of Aegla, and its range from the elevated
regions of the country to the lowlands, from the Cordillera of Men-
doza to the River Plate in the vicinity of Buenos Aires, but there
at a distance from the sea. He reported its presence in Uruguay,
where he said that it is more abundant and is found [at times] close
to the sea coast, as in the rivulets Miguelete and Carrasco, and also
in some localities where the fresh water becomes brackish at sea
level, and that it had also been found at Minas, about 159 kilometers
from Montevideo, in a spring that had been uncovered on a small
mountain in the course of excavating limestone. Berg, who appears
not to have seen these particular specimens, credited the find to Prof.
Arechavaleta, the chemist who examined the water with a view to
its utilization by the city. He regretted that the latter failed to
state whether the organs of sight were developed in these animals
or not. Berg also took occasion to say, on comparison of specimens
from southern Brazil, Chile, Mendoza, Buenos Aires, and Montevideo,
that it was his belief that Fritz Miiller’s A. odebrechtti is the same
as A. laevis.
This same year, Ortmann (1892, p. 246) summarized the distribu-
tion of A. “laevis” and added a new south Brazilian locality record,
Sao Lourenco, and figured the mouth parts.
Not aware that he had been antedated, Nobili (1896, p. 1) thought
he had seen the first Aeglas from the Argentine, from San Lorenzo
(Jujuy), Tala (Tucuman), and the Province of San Luis. He ob-
served that the coloration of the Tala specimens differed from that
of the San Lorenzo and San Luis ones. To some degree, at least,
I believe color of specific significance in this genus. Nobili also
called attention to S. I. Smith’s (1869, p. 31) “List of the Described
Species of Brazilian Podop[h]thalma,” saying that A. laevis had
been omitted. Smith (1869, p. 39) made reference to a Galathea
amplectens of Fabricius (1798, p. 415) but believed that “it is prob-
ably not a true Galathea.” This species in some respects suggests
Aegla. According to Fabricius, the carapace of G. amplectens is
smooth and the rostrum short and emarginate [forming the orbits] ;
but, contrariwise, Fabricius distinctly stated that this species in-
habits the ocean off Brazil and that it is luminous at night. The
latter phenomenon might have been due to bacterial infection and
the reference to a marine habitat in error. However, as this crusta-
cean seems to have come under the scrutiny of Latreille (1803, p.
199), the author of A. Zaevis, as well as that of H. Milne-Edwards
(1837, p. 276), and yet was not identified by either of them with
THE SPECIES OF AEGLA—SCHMITT 439
Aegla, it must be distinct, even if not a true Galathea as Smith
suspected.
Apprised by Wierzejski (1897, p. 1) of the shortcomings of his
earlier note, Nobili (1898, p. 6) hastened to publish an emendation.
In this he pointed out that Wierzejski (1892) himself had been
anticipated by Lucas (1876), and that Berg’s note (1892) appeared
the same year as Wierzejski’s.
Almost on the heels of this note of Nobili’s (1898), not quite three
months later, Berg (1898, p. 7) reprinted verbatim his notes of 6
years before. To these he added references to the remarks of Nobili
(1896) and Wierzejski (1892; 1897), and three new Argentine rec-
ords: the provinces of Salta and Cordoba and Neuquen Territory.
Strictly in agreement with the pronouncements of Wierzejski
(1892) and Berg (1892; 1898), Ortmann (1898, p. 1149), under the
family Aegleidae [now better Aeglidae], tersely stated, “Monotype
Familie, von der Gattung Aeglea Leach gebildet, die einzige Art
(A. laevis Latr. Taf. lxxiv, Fig. 1**) in Siid-Brasilien, Argentinien
und Chile besitzt, wo sie in Siisswasser, besonders in Gebirgsbachen
lebt.” As the figure cited appears to have been copied directly from
Cuvier (1837, pl. 47, fig. 3), quite naturally my comments on the
original (p. 435) apply to Ortmann’s black-and-white reproduction
of it.
Following Cunningham (1870, p. 495), who merely mentions
A. “laevis” as having been “collected in a fresh-water stream in the
neighborhood of Valparaiso,” no further references to Aegla from
Chile appeared in literature so far as I am aware, until that of
Doflein (1901, p. 185). He added a new locality to its range in that
country: Lake Lilanquihue, near Puerto Montt. His A. “laevis” may
be A. abtao.
Carlos Moreira (1901, pp. 21-23, 84) with fresh material that he
had collected in the State of Santa Catharina, Brazil, in his invalu-
able work on the “Crustaceos do Brazil,” fully demonstrated the
distinctness of the A. odebrechtii of Fritz Miiller. At the time, un-
fortunately, he believed it to be synonymous with Girard’s Chilean
A. intermedia.
In spite of Moreira’s able presentation of the case, Ortmann (1902),
in his extremely interesting paper on “The Geographic Distribution of
Freshwater Decapods and Its Bearing upon Ancient Geography,”
continued to insist that the genus was monotypic. This stand, which
also had been emphasized by Berg (1892; 1898), seemed to close the
door on further taxonomic investigations. Most, if not all, subse-
quent work has apparently been undertaken under the impression
that there was only one species of Aegla, for it has been confined
“**A4_ odebrechti F. Miill. is hiervon nicht verschieden.”
440 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
largely to morphologic, parasitological, and biological investigations:
Porter, 1907; Bennati-Mouchet, 1931a, 1931b, 19382a, 1932b; Porter,
1936 *; Perez, 1936.
I should not fail to mention here the modest yet very useful check-
list prepared by the late Hermann Luederwaldt, naturalist to the
Museu Paulista, at the time curator of the invertebrate collections.
In his “Lista dos Crustaceos Superiores (Thoracostraca) do Museu
Paulista que Foram Encontrados no Estado de §. Paulo” (1919, p.
431) under A. intermedia, the species with which A. odebrechtii had
been thought synonymous, he has specimens from “Perus” and “Alto
da Serra,” localities that I have included in the distribution of
A. odebrechtii paulensis (p. 492), and states that the A. Jaevis from
Franca is regarded as an “especie duvidosa.” From undoubted dupli-
cates of this Franca material received from Dr. H. von Ihering in
1915 the type of a new species, A. franca, has been selected. Dr.
von Ihering also sent the National Museum specimens of A. 0. paulen-
sis from Perus.
The foregoing résumé by no means represents a complete bibli-
ography of Aegla. It has been assembled for the purpose of setting
forth its taxonomic history, indicating its distribution and the sources
of my information. More has been done on its parasites than is indi-
cated by the works cited above. The genus and its supposedly unique
species are usually, if only briefly, referred to in the more compre-
hensive general zoological and carcinological texts.
ZOOGEOGRAPHIC NOTES
°
The recorded occurrences of the several species of Aegla, despite
the present additions thereto, are altogether too few to admit of
more than brief mention of the intriguing speculations that are sug-
gested by their geographic distribution. When this is plotted it
appears that each of the major tributaries of the largest rivers
possesses its own peculiar species (as exemplified in part of A. franca,
A. castro, and A. parana). Although in some cases several tribu-
taries, where near enough together, may have the same species in
common (A. platensis* and A. uruguayana*), other localities of
® Porter remarks that Aegla “laevis” has often been collected in the Chilean provinces
Valparaiso, Aconcagua, Coquimbo, and Atacama and records the recent accession of a
specimen from the Rio Maipo, at Santiago.
‘This species is found in the State of Rio Grande do Sul, Brazil; Uruguay ; and Buenos
Aires, Argentina. However, I cannot explain its existence in a locality as far removed
as Tucuman, Argentina. A confirmation of this occurrence is needed, as well as collections
from the vast stretch of country between Tucuman and the eastern seaboard.
®This species seems to be rather generally distributed in the River Plate region and
more particularly on both sides of the Rio Uruguay and some little distance up the Paranda.
For this species we have one tentative record from San Luis, Argentina between 400 and
500 miles to the westward of Buenos Aires. As with A. platensis (footnote 4), collections
from the intervening stretch of territory, from which we have seen no Aeglas at all, are
much to be desired.
THE SPECIES OF AEGLA—SCHMITT 441
80 70 f 60 50 40
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KEY TO
DISTRIBUTION OF AEGLA
a Q parana @ odebrechhis
@® sanlorenzo @ 0. pavlensis
@ platensis ® 72evqvensis
@ vurvgvayana © Ofinis
© prado ® Aumahvaca
© castro (© conceocionensis
@ franca @ laevis
so- ® /ywuyana © / falcahvano
© aeniiculata © abtao
( paoudo © os/imayana
\ | ! / /
100 90 60 o 40 30 20
Ficure 40.—Distribution of Aegla., This map is based on material that has actually been
studied in preparation of this account of the genus. So far as collecting stations are
definitely known, they have been accurately plotted; otherwise, their positions are approxi-
mations only. The actual locality at which the Rio Grande do Sul, Brazil, specimens of
A. platensis were taken is not known; it is also unknown for A. uruguayana from the
Province of San Luis, Argentina. As indicated, three species, 4. platensis, prado, and
uruguayana, have been found at Montevideo or in its immediate vicinity; at Buenos
Aires both 4. platensis and uruguayana occur. As some doubt attaches to the origin of
our A. affinis material, its occurrence has not been plotted (cf. p. 498, “Holotype”)s
442 PROCEEDINGS OF THE NATIONAL MUSEUM Vou, 91
even lesser extent may support more than one species (i. e., Buenos
Aires and adjacent region, two species: A. platensis and A. wru-
guayana; and Montevideo and vicinity, three species: A. platensis,
A. uruguayana, and A. prado). (Fig. 40.)
The presence of two or more species in one locality, as in Buenos
Aires and its environs and perhaps also Montevideo, may have
resulted from the tremendous floods to which at least the lower
reaches of the several rivers that converge to form the Rio de la
Plata are subject. Such an agency would serve to bring together in
the same region species that otherwise might exist at some distance
from one another.
Generally speaking, most of the species seem rather circumscribed
in their distribution (but it must be remembered that the number of
records we have for any one species is still very small). If this
is so, the Aeglas may be very responsive to their immediate environ-
ment, very plastic forms, or else the species are very “young.”
The climatic extremes encountered by Aegla in its geographic
range are considerable (K6ppen, 1930, fig. 41). These, too, may
have a marked effect not only on the distribution of the species but
on their actual development or evolution. Two species that may
be a living demonstration of the effects of climatic conditions, which,
after all, are but a part of the environment of a species, are A.
jujuyana and A. hwmahwaca. So far as we know now the two are
scarcely more than 70 miles apart at their point of nearest approach,
yet, on the basis of precipitation figures alone, they are a vastly
greater distance apart. At Jujuy, Province of Jujuy, Argentina,
the type locality for A. jujwyana, as much as 29.26 inches of rain
falls during the year, with some rain in each of the twelve months;
at Humahuaca, in the same province, the type locality for A.
humahuaca, on the other hand, the total yearly rainfall, 6.11 inches,
is less than that of the wettest month of the year at Jujuy (January,
with 6.65 inches), while five months (May to September) are wholly
without appreciable precipitation (Reed, undated MS.; see footnote,
p. 500).
If it is true that the least differentiated, least spiny or ornamented
species stands nearest the ancestral Aegla, then perhaps our A.
jujuyana is least removed from it in an evolutionary sense. This
would place the center of distribution somewhere in the northwestern
part of Argentina (Province of Jujuy), which is at variance with
Ortmann’s belief (1902, p. 389) that Aegla was originally indigenous
to Chile and subsequently extended into northern Argentina and
southern Brazil, or perhaps in the reverse direction.
A. jujuyana lacks or has not yet developed the palmar crest that
is so characteristic of almost every other species of Aegla; its rostrum
THE SPECIES OF AEGLA—SCHMITT 443
is somewhat intermediate between the flatter, troughed (Pacific or
Andean) type*® present in species found on the east and west slopes
of the Andes and the more spinelike, ridge-roofed (Atlantic type)
rostra of the species of the great region more or less immediately
tributary to the River Plate.
Of special interest in this connection is the fact that we meet also
with the so-called Pacific or Andean type of rostrum in the Serra
do Mar bordering the Atlantic coast of Brazil, in Santa Catharina
(A. odebrechtii) and in Sao Paulo (A. odebrechtii paulensis). This
discontinuous distribution of the forms with the Pacific or Andean
type of rostrum may be apparent only.
From the center in Argentina at or in Jujuy it may be that the
forms or variants with the Pacific type of rostrum spread out west-
ward to the Andes and beyond to Chile and eastward to the Serra do
Mar of Brazil, while down the vast Argentine Rio Parana drainage
area and across to at least the lower reaches of the Rio Uruguay to
Rio Grande do Sul, and to Parané, migrated those that developed
what I have called the Atlantic type of rostrum. Not fitting in with
this speculative scheme of things is A. franca, from Franca, Sao Paulo,
Brazil, also a species with what might be called the more intermediate
type of rostrum found in A. jujuyana. It could be a northeastern
offshoot of the original or ancestral jujuyana stock, or else a reversion
to the ancestral condition of a Brazilian form with the Pacific type of
rostrum.
The marine origin of Aegla appears indisputable, and therefore it
is of more than passing interest that the general region in which
A. jujuyana is centered has geologically had a long-continued marine
history, with marine deposits antedating the Devonian, up through
the Carboniferous (Berry, 1922). Since Cretaceous time that part
of South America seems to have been wholly continental and its
waters no longer marine. Undoubtedly the elevation of the land
above the sea was gradual, or at least long enough drawn out to
allow the ancient forebears of the Aeglas of today to adapt them-
selves to progressively less saline and increasingly fresher waters.
Although there are a few very fragmentary crustacean remains
said to be decapod in the Permian, the first unquestionable fossil
decapods, already well differentiated into groups or tribes, families,
genera, and species, are Triassic (Zittel, 1913, p. 760; Glaessner,
1929, pp. 404, 462). Galathea first appears in the Upper Cretaceous.
Pseudogalathea from the Lower Carboniferous of Scotland, however,
has been assigned to the “schizopoda” by paleontologists (Zittel,
1913, p. 757).
®A more detailed description of these types of rostra will be found on p. 448 of the notes
on “characters used in diagnostic key and specific descriptions,’ and in the key itself,
pp. 451 and 454.
444 PROCEEDINGS OF THE NATIONAL MUSEUM Vow. 91
Ortmann (1902, p. 341) in his discussion of the geographic distribu-
tion of fresh-water Crustacea and its bearing upon ancient geography
stated that “the presence of the genus Parastacus on both slopes of the
Cordilleras (even the identical species is found in one case on both
sides, and in this respect Aeglea agrees with Parastacus) points to a
time when the Cordilleras had not yet attaimed their present eleva-
tion. As von Ihering [1907, 1911] has shown, for many groups of
animals this chain forms a very sharp barrier, and it does not seem
probable that these freshwater Crustaceans are able to cross these
high snow and ice covered mountains.” Although this may well
have occurred, it is not very necessary to presuppose that Aegla
reached its continent-wide distribution before the Andes attained
their present elevation, for, in spite of the height of this great
mountain range and the rigors of the climate investing its summits,
there certainly are passes, particularly in the lake region of Chile
and Argentina, through which Crustacea such as Parastacus and
Aegla might have made their way in times past, if not present.
There must be a pass of this sort above the headwaters of the Rio
Petrohue and Lago Todos Santos, where are to be found “on top
of the pass of Perez two small streams, one flowing toward the
Pacific, the other toward the Atlantic * * * (Kigenmann, 1928,
p. 25). Today one can go by bus, automobile, motor boat, and
steamer from Chile to Argentina by way of Petrohue, Lago Todos
Santos, Peulla (elevation 190 meters), Casa Pangue, Chile (elevation
320 meters), Lago Frias, Argentina, to Puerto Blest on Lago Nahuel
Huapi, Argentina (elevation 756 meters).
Insofar as they apply to the same geographic area, I am most
anxious to have an opportunity of checking Dr. Eigenmann’s findings
(1928, especially references given in the partial bibliography on p. 2)
based on his studies of the fresh-water fish fauna, its distribution,
and origin, against that of Acgla. But before that can be done,
vastly more Aegla material than has yet been collected would have
to be assembled.
There seems to be a relation of sorts between our rostral types
and such of the “environment complexes in which the sum total of
the natural conditions are about equal” of Haseman (1912, pl. 15).
The forms with the ridge-roofed, Atlantic type of rostrum more or
less occupy Haseman’s “Uruguay-Rio Grande do Sul” area plus
some additional territory to the south and west, while the Andean
or Pacific type, along with the intermediates, A. jujuyana and A.
franca, occupies his “West Andean,” “Patagonian,” and “Alto Parana
and its affluents” areas. As the forms with Andean type of rostrum
are found in each of the last-named “environmental complexes” of
Haseman, they must have something in common, be it geologic his-
tory, environment, or something else.
THE SPECIES OF AEGLA—SCHMITT 445
In an endeavor better to evaluate the specific characters of A.
odebrechtii paulensis, I besought Drs. Paulo Sawaya and Ernesto
Marcus, of the University of Sao Paulo, for further material of this
subspecies. Although it was not possible for them to obtain it, I did
receive some illuminating information regarding the waters of Alto
da Serra, the type locality, in a letter from Dr. Marcus:
“Alto da Serra is a mountain pass, 38 km. from Sao Paulo and 22
km. from Santos by rail, where the high-road and the railway, after
having climbed the very steep coast-slope of the Serra do Mar, reach
the level of the highland of Sao Paulo. The brooks of Alto da Serra
chiefly fall in cascades down the coast-slope to the narrow lowland
of Santos, but some of them also enter the system of the Tieté River
that springs in the Serra do Mar, 15 km. distant from the sea, and
flows westward through the city of Sao Paulo and the interior of
the state. The mouth of the Tieté in the Parana is 650 km. distant
from the coast.”
Our neotype of A. odebrechtii is labeled as from Santa Catherina
without particulars, but more recent specimens most helpfully pro-
vided by Dr. Carlos Moriera, through the kindness of his good friend
Dr. G. Kuhlmann, of Blumenau, are from that place in the State
of Santa Catharina. One cannot ascertain from which particular
watershed, Atlantic slope or westward slope of the Serra do Mar,
Fritz Miiller’s original specimens were taken.
From what Dr. Marcus had to say about Alto do Serra and from
what we now know of the occurrence of A. odebrechtz at Blumenau,
it may be that the forms with the Andean type of rostrum in east-
central Brazil are confined to watercourses draining into the Atlantic
Ocean direct.
We need not only a great deal of additional material from all
parts of the country but, along with it, much more complete locality
and environmental data than has been available heretofore before
we can hope to elucidate the distributional and taxonomic problems
that have been raised by this manifestly preliminary study.
CHARACTERS USED IN DIAGNOSTIC KEY AND SPECIFIC
DESCRIPTIONS (Fig. 41)
It is little wonder that the genus Aeg/a has been considered mono-
typic by so many authorities. In a general way and in many par-
ticulars all Aeglas bear a very close resemblance to one another, but
there is diversity of form of the cheliped, shape and armature of
the orbit, proportion of the carapace and rostrum, relative develop-
ment of the anterolateral spines, hepatic lobes, cardiac area, and areola,
revealing differences of a kind that can no longer be explained merely
as variations of a single species.
446 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 91
In his studies on the North American crayfishes of the genus Cam-
barus, Dr. Herman A. Hagen remarked, according to Faxon (1885,
p. 17): “If the reader is unable to determine * * * the speci-
mens in his hands * * * through lack of males, the fault lies,
* * * not in the principle of classification, but in the scantiness
of his material. A species involves two sexes; and until the species
is known, it avails little to attempt the determination of a specimen
in this difficult genus.” 7
Aegla, likewise, is a difficult genus. Certain forms represent un-
questionably distinct species; others have been proposed with some
hesitation; two have been rated merely subspecies.
For the present, at least, it has been necessary to confine specific
descriptions and diagnostic key characters to as fully developed male
specimens as it has been possible to obtain, for in the females the
specific characters do not seem to come to full fruition, and with
only females at hand it may be difficult or perhaps at times impossible
to identify them as to species.
In Aegla, the female, in some respects at least, is definitely the
weaker sex, and, even if attaining as large a size, it 1s never so
distinctively developed specifically as the corresponding male. This
is particularly true of the hands, or chelae. In either sex these are
sufficiently asymmetrical to be referred to as the major and the
minor chela. The larger chela may be either the right or the left
one, but it is usually the left hand, with comparatively few excep-
tions, that is the larger. The chelae in the female are undersized
and underdeveloped, more of the pattern of the minor chela of the
male, which, in turn, might be described as being more or less
feminine in appearance. The hands or chelae of the males, more
especially the larger one, tend to become more and more swollen as
the animals get older and larger.
The prehensile margins of the fingers are furnished with a close-
set pavement or palisade of corneous scales; this armature is not
otherwise mentioned, although the presence or absence of a large,
usually conspicuous, “lobular” tooth is mentioned in the descrip-
tions of certain species and in the diagnostic key. A tooth of this
nature occurs on the prehensile margin of the fixed finger of the
major chela of most species, usually on the corresponding finger
of the minor chela also; often the movable finger has a somewhat
similar tooth opposed to one on the fixed finger. In three species
the prehensile margin of the fixed finger is without such a lobular
tooth: A. sanlorenzo, A. jujuyana, and A. humahuaca.
4 Specimens studied should be of reasonable size and development. It is difficult to deal
with specimens of less than 20 mm. in length of carapace and rostrum together and, indeed,
even slightly larger individuals are often none too well developed, even though male.
THE SPECIES OF AEGLA—SCHMITT 447
On the outer margin of the movable finger of a number of species
near the base there is a definite projecting lobe or angle, usually
spined, and, when present, spined in younger specimens if not in
the fully developed adults (as in A. platensis) ; sometimes the lobe
is reduced in size or suppressed and no more than suggested by some
small spinulation at the place occupied by it in other species, or
there may be no lobe, angle, or spinulation present at all, the finger
being perfectly smooth and rounded off, as in A. laevis talcahuano.
The carpus of the chelipeds is armed on the inner margin with a
row of strong spines, but in this series I do not include the spine that
---------------- rostrum
Bee APES ee rostral carina
ee orbital sinus
pee ee ee orbital spine
Ph AE et extra-orbital sinus
Jil ferro anterolateral spine
7 X---}---------- epigastric prominence
pn 1 Sey SESS = ant. margin protogastric lobe
Sinica) posta ee first P
ao second}hepatic lobe
eae third
REQ cervical groove
Eo
--=
Se -
L_-
--- gastric area
---internal
Ficure 41.—Diagram of 4egla carapace, illustrating some of the terms used in describing
species.
may arm what I have called the carpal lobe at the anterior inner
angle of the carpus. This angle or lobe may be scarcely more than
bluntly rounded off and scabrous, sometimes it is more acute and
apically spinulated or furnished with a sharp denticle or small corn-
eous spine or two, and it may, as in A. rioliémayana, carry a slender,
clean-cut, sharp, corneous-tipped spine of good size, about as large
and conspicuous as the penultimate spine of the series arming the
inner margin of the carpus. The carpal lobe is not always so well
developed or so well armed in the female as in the male Aegla; the
descriptions given are based on male specimens only.
More or less parallel to and above the inner spined margin of the
carpus there is in most species a definite carpal ridge, usually more
or less nodulated, with the nodulations more or less scabrous; on
each nodulation there is generally a row of small, corneous scales,
435661—42—2
448 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
which are arranged more or less transversely in the distal half of
the ridge and somewhat or at times almost wholly longitudinally at
the proximal end of the ridge. In some species the nodulations be-
come tuberculiform, and in still others, such as A. denticulata and
A. castro,’ actually replaced by sharp-pointed conical spines.
On the middorsal line of the carpus in a number of species there
is a suggestion of a second though much less well formed ridge in
the shape of an irregular, more or less scattered, longitudinal row
of scabrosities somewhat larger than the others that may roughen
the surface of the carpus; in A. parana there is a middorsal row of
small sharp spines, few in number.
The upper longitudinal margin of the merus of the cheliped may
be very sharply and conspicuously spined, or else tuberculated with
apices of tubercles scabrous, or virtually unarmed as in A. jujuyana
and A. hwmahuaca.
The anterior margin of the merus may be perfectly smooth and
evenly rounded off (A. hwmahuaca), actually spined, or more or less
finely denticulate; in other species it will have middorsally a more
or less definite swelling, nodule, lobe, or even tubercle (A. odebrechitiz)
which may anteriorly be minutely spinulated whether the rest of the
anterior margin is similarly armed in part or not. As with many
of the other morphologic features of Aegla, there seems to be con-
siderable variation in the degree of development exhibited by this
lobe, so that its specific importance, in the light of our limited
knowledge of the members of the genus, cannot be satisfactorily
determined.
The basis and ischium of the chelipeds are fused to form one joint.
Below, toward its proximal end, there are three transverse, more or
less impressed lines. The anterior and posterior lines mark muscle
attachments; the middle one constitutes “‘a fracture plane’ at which
separation of the limb takes place in [this and] many [other Crus-
tacea] Reptantia” (Calman, 1909, p. 278). In describing the arma-
ture of the “inner margin of the ventral surface of the ischium”
only that portion of the ischium proper, or of the fused joint, basis-
ischium, beyond or distal to the anterior of these three lines of
demarcation is referred to.
There seem to be two principal types of rostra to which the various
species of Aegla may be referred. The first of these I shall call the
ridge-roofed (Atlantic) type. In this the dorsal surface rises from
5A. intermedia, described by Girard (1855, p. 255) but not yet rediscovered, is described
as having two rows of spines on the carpus, its only recognizable or rather distinctive
character that it shares with A. denticulata and A. castro. The second of these certainly
does not occur in Chile and so could not be confused with Girard’s species, from which
A. denticulata is at once set off by the longitudinal keeling of its carapace and the con-
spicuous saw-teeth arming the lateral margin of the posterior portion of the carapace.
THE SPECIES OF AEGLA—SCHMITT 449
the lateral margins to form a very definite, rather sharp carina ex-
tending straight forward to the anterior extremity of the rostrum,
which is distally not, or at most only slightly, bent upwards. At
about the level of the corneae the dorsal carina of this type of ros-
trum almost always attains a greater height or elevation above the
lateral margin than the ventral keel has depth below the margin.
Above the level of the lateral margins the rostrum in cross section
is definitely triangular, like the roof of a ridge-roofed or gable-ended
house. The sides of this roof run straight down from the ridge
or carina to the lateral margin either side at about a 45° and often
steeper angle (that is, at about the middle of the free portion of
the rostrum or between that point and the level of the anterior margin
of the corneae). At most these lateral slopes in this first group may
be slightly concave; they are, however, never particularly depressed or
flattened down, excavate, or longitudinally grooved or troughed.
The other type of rostrum (Pacific or Andean) is fairly flat from
side to side and not as a rule at all like the so-called ridge-roofed
type, although some species assigned to it (A? section of the key,
p. 454) may have rather a sharp rostral carina (i. e., A. riolimayana).
In general, rostra of this type in cross section form more of a flat
longitudinally corrugated roof than a steep-sided ridged roof, inas-
much as the sides of the roof either side of the median carina are
usually more or less depressed below the lateral margins, and ex-
cavate or longitudinally troughed. The height to which the rostral
carina rises above the lateral margins, at about the level of the
corneae, is usually appreciably less than the depth to which the ven-
tral keel extends below the lateral margins. As a general rule, the
dorsal carina tends to fade out or disappear as a carina before
attaining the distal extremity of the rostrum, which is generally more
or less definitely recurved or bent upward.
A few species seem to have rostra of an intermediate or transitional
type that may not have been altogether satisfactorily placed in our
key. However, such species have been assigned to that primary
group, A’ or A*, to which they appeared to be most closely related,
all characters considered. A. jujuyana and A. franca have been
assigned to section A* of the diagnostic key, and A. affinis to section
A*, This last-named species, in the unique specimen at hand, has the
dorsal rostral carina somewhat higher at the level of the corneae
than the ventral keel is deep, yet its basally broad and flattened
rostrum is certainly indicative of a nearer relation to the A* than to
the A’ species. A. jujuyana and A. humahuaca fall into opposed
primary sections of our key on the basis of the character of the
rostrum; nevertheless, there is in some respects a tantalizing resem-
blance between the two that suggests a suspiciously close kinship.
450 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
Width of the orbital and extraorbital sinuses, where referred to,
has been measured in line with the tips of the orbital spines, from
the extremity of the spine to the rostral margin, and from the spine to
the inner margin or slope of the anterolateral spine. The orbital
spine (or spinule) is the actual spine or spinule marking the outer
or distal end of the orbital margin, without reference to scabrosities,
denticles, scales, or tiny, often microscopic, spinules that may arm or
persist on the orbital margin of some species. In most species the
outermost of such a series of orbital scabrosities becomes so developed
as unmistakably to become the orbital spine or spinule.
The length of the anterolateral spines in relation to the eyestalks
is perhaps not a very reliable character, owing possibly to differences
in contraction as a result of preservation, yet in a species like A.
sanlorenzo the anterolateral spines exceed the eyestalks, while in
A. abtao and A. riolimayana, for example, they generally fall short
of the posterior margin of the cornea.
I have not been able so far to “pin down” the relative proportions
of the areolations of the carapace in a way to permit their satis-
factory use in specific description. The areola itself is rather elongate
in some species, very squat in a number of others, and quite different
in the relation of its posterior lateral margins to the lateral furrows
or suture lines of the cardiac area, in at least two nearly related
species, A. abtao and A. riolimayana.
Most of the species of Aegla exhibit a tendency toward smoothness
and bluntness, even to the suppressing of spines in the older, more
developed specimens. In A. parana quite the reverse is true; there
seems to be an accentuation of the spininess of this species, the fully
developed adults are very spiny or at least more prickly appearing
than any other one of 20 species or subspecies described.
Aegla parana is the only Aegla having the ambulatory legs strongly
spined above and, with A. sanlorenzo and perhaps also A. prado, the
only species having reasonably strong spines below near the anterior
end of the ambulatory merus. Only one ambulatory leg, the first
on the left side, has been figured for each of the species dealt with
in this paper, chiefly to show the proportions not as yet clearly
proved to be of specific value.
In the majority of the Aeglas the sternal plate between the chelipeds
carries no particular armature; in a few species a very definite, often
corneous-tipped tubercle or low conical spine is to be found on the
median line toward its anterior end; the anterolateral angles of this
particular plate are sometimes markedly produced or even spiniform.
Except for the contours of the epimera of the second (in lateral
view, apparent first) abdominal somite no particularly noticeable
specific differences have been observed in the abdomen or the tail
THE SPECIES OF AEGLA—SCHMITT 451
fan. In the female the abdomen as a whole is relatively shorter and
broader than in the male, and the median dorsal area of the respective
abdominal somites is relatively wider. This character usually enables
one superficially to distinguish males from females. The sexes, how-
ever, are definitely distinguished by means of their genital apertures.
In the female these open on the coxopodites of the third (antepenulti-
mate) pair of legs, and in the male on the fifth (last) pair of legs.
In mature males the vas deferens on each side is externally produced
as a thin-walled tube.
KEY TO THE SPECIES OF AEGLA
At. Rostrum definitely ridge-roofed,® triangular in cross section; above, with
lateral slopes of “roof” running down often at nearly a 45° angle from
median carina to lateral margins (these lateral slopes are not distinctly
troughed or excavate either side of the median carina as are practically
all the relatively flattened rostra of the species under A’, p. 454; at most the
lateral rostral slopes may be slightly concave) ; rostrum sometimes show-
ing a slight upward inclination toward tip, but usually straight and
not recurved; rostral carina and scales with which it may be furnished
running straight and usually definitely to anterior extremity; front of
species belonging to this section of key generally wide or at least moder-
ately wide, rarely somewhat narrowed (as in jujuyana and franca) ;
orbital spines well developed; sinus (extraorbital) between orbital and
anterolateral spines generally of good size, wide or moderately wide, rarely
small (jujuyana, franca, and occasional specimens of prado).
B*. Hands, though they may become somewhat thick and swollen, never taking
on markedly inflated or subglobular appearance of jujuyana (B’, p. 453) ;
inner margin of palm always more or less crested, and when crest is low
and little developed armed with at least one sharp spine at anterior end;
lobular tooth on at least fixed finger usually present and well developed ;
rarely is this tooth not definitely present, or obsolescent, as in A. san-
lorenzo (p. 452) (lobular tooth on fixed finger is also lacking in jujuyana,
B?, p. 453, and humahuaca, under section A’, p. 456 of this key) ; dorsal
anterior angle of epimeron of second (in lateral view, apparent first)
abdominal somite almost always armed with a spine (sometimes not in
franca).
Cc’. Front generally very wide, extraorbital sinus at least half, usually more
than half of, to nearly subequal to orbital sinus (somewhat nar-
rower than other species in this section is sanlorenzo) ; orbital spines
a prominent feature of frontal margin.
D‘. Posterior margin of ventral surface of first ambulatory merus armed
with at least one conspicuous strong spine near distal end about ona
level with proximal border of articular membrane”; inner margin
of ventral surface of ischium of chelipeds armed with two fairly
long, well-developed, acute, corneous-tipped spines, one near distal
end of joint, the other near proximal end (spines of this size and
°In lateral view at the level of the anterior margin of the cornea, the height of the
rostrum, or its carina, above the lateral margin of the rostrum is usually much greater
than the depth of the rostrum below the lateral margin.
104. prado has a spine of moderate size in this position and A. castro a quite small one
or two, but both are species with the front only moderately wide, O°, p. 453, this key.
452
PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
prominence are not found in any species of Aegla other than the two
grouped here under D*) ; movable finger without definite or real lobe
on outer margin near base, even though margin of finger may some-
times be spiny; epimeron of second (in lateral view, apparent
first) abdominal somite with anteroventral border more or less
deeply concave; anterior dorsal angle produced to form a sharp spine
of good size, ventral angle also produced, narrow, extremity may
be blunted, sometimes sharply spined like anterior dorsal angle (in
certain large specimens of paranda).
E*. Merus of ambulatory legs armed on upper margin with several,
usually a full series of, strong, well-developed spines; carpus with
ridge above spined inner margin, also well spined, and with a
second longitudinal row of normally three sharp spines on mid-
dorsal surface, sometimes posterior two spines of this series much
reduced or wanting; a well-developed lobular tooth at least on
fixed finger of either chela; movable finger without a definite lobe
or projection on outer margin near base, sometimes, but not often,
a spine or several spinules in this position, not to be unexpected
in this otherwise very spiny species; outer margins of hands
spined; inner margin of palm forming a comparatively low ridge
(palmar crest), which is serrate, serrations spined; sometimes
inner margin or ridge fairly straight and serrulate.
parana (p. 458)
FE’. Upper margin of ambulatory merus not armed with a number of
strong spines, at most scabrous or small spinulated; carpus with
ridge above inner spined margin not spined, scabrous-nodulated,
and without a longitudinal row of spines on middorsal surface;
prehensile margins of fingers slightly sinuous but without lobular
tooth on either fixed or movable finger; no lobe on outer margin
of fixed finger near base; outer margins of hands scabrous but
not spined as in preceding species; inner margin of palm scarcely
crested, broadly rounded off, rising anteriorly in a low keel
(palmar crest) to form a single short, sharp spine.
sanlorenzo (p. 461)
D*. No noticeably strong spine near distal end of ventral posterior margin
of ambulatory merus, at most a relatively small spine, tubercle or
scale in this position; epimeron of second (in lateral view, apparent
first) abdominal somite with anteroventral border more or less
straight, at most only slightly concave; ventral angle rounded off;
fixed finger at least with a definite lobular tooth of good size one
prehensile margin.
BE’. Normally only first hepatic lobe well defined and anteriorly spined,
second and third lobes scarcely more than indicated (occasionally
one of other lobes fairly well marked on one or the other side of
carapace) ; movable finger definitely with a lobe on outer margin
near base; in most specimens, especially those of medium and
small size, the lobe furnished with a small spine or sharp scale,
in many of the larger specimens, such as the type, the lobe fre-
quently unarmed, but always distinctly present and more or less
angled; ischium of chelipeds with a not particularly prominent
tubercle (not spine), which is furnished with a corneous apex or
seale, at distal end of inner margin of ventral surface.
platensis (p. 464)
THE SPECIES OF AEGLA—SCHMITT 453
E?. All three hepatie lobes well marked in specimens of fair size; in
mature or adult specimens anterolateral angles of at least first
two and usually all three lobes acute and sharply spined; movable
finger without a lobe on outer margin near base; ischium of
chelipeds with a conspicuous sharp fairly slender spine at distal
end of inner margin of ventral surface____ uruguayana (p. 467)
C*. Front only moderately wide, extraorbital sinus less than, half width of
orbital sinus, often only one-third or less than one-third its width;
a well-developed orbital spine intervenes between the two sinuses;
fixed finger at least with a definite lobular tooth of good size on
prehensile margin; movable finger definitely and normally with a
spined lobe on outer margin near base; anteroventral border of
epimeron of second (in lateral view, apparent first) abdominal somite
generally just about straight, may at times be very slightly concave.
D’*. All three hepatic lobes well marked, their anterolateral angles acute
and spined, each forming a decided offset in lateral margin
(forming three steps, as it were before the cervical groove) ;
anterior margins of protogastrie lobes acute-angled, more sharply
peaked perhaps than in any other species of Aegla.
prado™ (p. 470)
D*. The three hepatic lobes plainly indicated but only the first well
marked and acutely spined at its anterolateral angle alone, form-
ing a distinct offset in the lateral margin of the anterior margin
of the carapace (before the cervical groove); anterior margins
of protogastric lobes more or less rounded off, or broadly
obtuse-angled.
E*. Carpus of cheliped with ridge parallel to and above inner spined
margin armed with conical tubercles, of which the greater part
take the form of acute-tipped conical spines; orbital spines
well set off from anterolateral; posteriorly dorsal margin of
rostrum merges with general surface of carapace on a level with
protogastric lobes; palmar crest somewhat approaching sub-
disciform, impressed, with upturned margins, reminiscent of
odebrechtiin (ps 455 below) s2222a2e_ eaten ten castro (p. 473)
E*. Ridge above inner spined margin of carpus armed with neither
spines nor acute conical tubercles, but scabrous, being furnished
with more or less transverse rows of small corneous scales;
orbital spine small and placed fairly close to anterolateral;
posteriorly dorsal margin of rostral carina ending between
protogastrie lobes well below general level of carapace behind
this point; palmar crest not subdisciform, narrow, longitudinally
somewhat troughed or excavate____------_-_--_ franca (p. 476)
B’. Hands very thick and inflated and, though scabrous, smooth appearing,
as they are rounded off in all directions; inner margin of palm neither
crested nor spined, thick and broadly rounded off; fingers with lobular
tooth not at all, or at most only very obscurely, indicated; fixed finger
of large hand very short and stubby looking (more so perhaps than
in any other species of Aegla), no lobe on outer margin of movable
finger near base; dorsal anterior angle of epimeron of second (in
lateral view, apparent first) abdominal somite (based on the very few
11The median line of A. prado is usually more or less definitely angled the full length
of the carapace, in effect carrying the carination of the rostrum back to the posterior border
of the carapace in the form of a prominent ridge; carination of this sort is found only in
this species and in A. denticulata under A, B, this key, p. 454, in which it is very
pronounced.
454
PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
specimens of this species available) at least spined or with corneous
granule or denticle on one or the other side of body in two specimens,
in a third specimen, however, on both sides; anteroventral border of
epimeron slightly coneave to fairly straight-________ jujuyana (p. 478)
A*. Rostrum more or less transversely flattened”; longitudinally troughed or
excavate either side of the median carina, often conspicuously so; rostral
margins often thickened and appearing more or less raised or upturned ;
rostral extremity often noticeably recurved, though sometimes straight
or only slightly upturned; rostral carina sometimes fading out ante-
riorly before reaching tip of rostrum, sometimes also merging or fusing
with anterior extremity of rostrum to the more or less complete oblitera-
tion of carina and the scales with which it may be furnished, corneous
scales sometimes continued to tip of rostrum as a feeble, scattered line of
scales only; front of species in this section of key relatively narrow, at
least in appearance, as compared with species of A’ section, p. 451;
orbital spines usually small, often placed rather close to and sometimes
apparently even up the inner slope or margin of anterolateral spine,
or wanting altogether.
B*. Carapace prominently keeled or carinated for its entire length; rostral
carina anteriorly fading out in distal third of free portion of rostrum,
merging with its thickened distal extremity; lateral margin of pos-
terior portion of carapace (behind cervical groove) conspicuously
serrate, sharply notched, and armed with prominent sawteeth or
flattened triangular spinelike teeth; orbital spine of good size; extra-
orbital sinus well formed, a prominent feature of the front, though
moderately narrow, being perhaps no more than one-fourth width of
orbital sinus; anterolateral spines attaining one-third to one-fourth
length of cornea; palmar crest thick, conspicuously spined; movable
finger with a sharply spined acute lobe on outer margin near base;
dorsal anterior angle of second (in lateral view, apparent first)
epimeron produced to form an acute corneous tipped spine.
denticulata (p. 480)
B*. Except for rostral carina, which may run backward as far as level of
anterior margins of protogastric lobes, carapace not noticeably if at
all keeled; lateral margins of posterior portion of carapace (behind
cervical groove) at most small spinulate or small corneous spined and
not at all toothed except perhaps for notch at lateral extremity of
cervical groove and at end of suture line immediately behind lateral
terminus of cervical groove.
Cc’. Anterior third, or even nearly half in some cases, of upper surface of
free portion of rostrum gently excavate or concave from side to side
with usually no more than trace of forward extension of rostral
carina or scales with which its carina is furnished; distal portion
of rostrum typically and usually strongly and more or less abruptly
recurved; rostral outline moderately broad triangular, carina short
but well marked, furnished with a single row of irregularly alter-
nating corneous scales; orbital spine may or may not be developed;
nearly always, however, a slight, sometimes abrupt, but always
narrow offset between outer end of orbital margin and inner slope
or margin of anterolateral spine; this offset about as often without
In lateral view at the level of the anterior margin of the cornea, the dorsal height of
the rostrum, or its carina, above the lateral margin of the rostrum is usually much less
than the depth of the rostrum below the lateral margin.
THE SPECIES OF AEGLA—SCHMITT 455
as with a small corneous scale or spinule (present in type), which
may represent or take the place of an orbital spine; offset usually
with slight notch or incision next to anterolateral spine; palmar
erest thick, almost obsolescent, upper surface not impressed or ex-
eavate. Dorsal anterior angle of epimeron of second (in lateral view,
apparent first) abdominal somite normally and usually rounded off
and unarmed; very rarely does one find a corneous scale or denticle
or two or even a small spinule, and then usually on epimeron of
oneysidevonly 225 see hse aie es ee a papuda (p. 483)
O°. Not even distal third of rostrum concave from side to side without
noticeable intervention of dorsal carina; rostrum carinated virtually
to its distal extremity or else anterior fourth or so of free portion of
rostrum so thickened that rostral carina and any longitudinal
troughing that dorsal surface of rostrum may otherwise have either
side of carina becomes more or less completely obliterated in this
terminal fourth of rostrum.
D*, Dorsal anterior angle of epimeron of second (in lateral view,
apparent first) abdominal somite rounded off and unarmed.
E*, Margins of palmar crest appreciably and noticeably upturned, upper
surface of crest definitely impressed or excavate; crest some-
what or quite subdisciform; movable finger with a definite
though sometimes small, but always spined or spiny lobe or
projection on outer margin near base; hands more or less sub-
ovoid in outline; rostrum normally not exceeding eyestalks by
as much as length of cornea; rostral carina not even faintly
traceable behind anterior margins of protogastric lobes; orbital
spine and extraorbital sinus definitely present; latter always
distinct though sometimes small.
F", Palmar erest conspicuously large and expanded, subdisciform,
noticeably excavate, much as if it had been impressed or
pinched out while soft with the ball of one’s thumb; margin
of crest obscurely serrate at best, scabrous and small spinulose ;
rostral carina furnished with an irregularly alternating dou-
ble (in very small part, at times triple) row of small corneous
SCH] OG e eS ake Soe Re ae ae re odebrechtii (p. 487)
F°’. Palmar crest only moderately large or expanded and, though
somewhat rounded off, not particularly subdisciform, more or
less longitudinally troughed; margin of palmar crest definitely
serrate; rostral carina almost smooth and naked appearing
on top, at most sparsely and well-nigh microscopically scaled
where it appears scaled______--_ odebrechtii paulensis (p. 490)
E*. Margins of palmar crest not noticeably upturned, crest at best only
slightly or narrowly and very shallowly, if at all, troughed or
excavate, not particularly impressed looking; crest more sub-
rectangular in outline than subdisciform; at most only a slight
lobe or projection on outer margin of movable finger near base;
margin of finger rough-spinulose and usually with a few larger
spinules on a slight elevation near base of finger, better seen in
smaller than in larger specimens.
F*. Definite orbital spine or spinule present, set off from anterolateral
spine by a small, narrow sinus or notch; palmar crest thinning
out to its outer margin, which is sharply though not deeply
saw-toothed, and sharply small-spinulose, not troughed or ex-
456 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
cavate; hands more or less subovoid in outline; rostrum plainly
troughed either side of well-defined, narrow, median carina.
neuquensis (p. 498)
F*. No orbital spine or spinule as such (in unique type specimen),
outer margin of orbit merges with inner slope or margin of
anterolateral spine without appreciable offset; small spinules
or spiniform scales on orbital margin tending to run up onto
sides of base of anterolateral spine; outer margin of palmar
erest fairly thick, rough-Sscabrous and somewhat lumpily
toothed; upper surface of crest longitudinally and narrowly,
slightly troughed; hands more or less elongate-subrectangular ;
rostrum only very shallowly and more or less obsolescently
troughed either side of rather biunt and rounded-off median
Carina’ Tai Sa Sint iy Bae Pie Amey teat ea affinis (p. 495)
D*. Dorsal anterior angle of epimeron of second abdominal somite armed
with a small spine or spinule (very rarely is angle armed with two
little spines or spinules).
E*. Fingers lacking lobular tooth characteristic of most species of Aegla,
fixed finger at most with only slight sinuosity on prehensile
margin; no lobe on outer margin of movable finger near base,
although a few larger corneous scales or small spinules sometimes
occur there; palmar crest low and thick, very broadly tri-
angular in cross section, dorsal surface at most very shallowly
and obsolescently excavate, more scabrous than spinulated, though
slightly marked serrations of blunt crest may be spinule tipped;
rostrum triangular, thick-looking, only shallowly troughed either
side of blunt, proximally more or leSs swollen median carina.
humahuaca (p. 498)
E’. Fixed finger at least with a well-developed lobular tooth on its
prehensile margin.
F", Rostrum more or less lingulate (tending to be tongue-shaped rather
than sharply triangular), lateral margins often more or less
subparallel in midsection of free portion of rostrum; rostrum
in lateral view noticeably bent downward, distally recurved;
orbital spine or spinule present or not present, when present
frequently much reduced, often no extraorbital sinus or notch
(small extraorbital sinus and orbital spine or spinule perhaps
always present in A. laevis talcahuano).
G*. Typically no orbital spine, normally outer end of orbital margin
merging with inner slope or margin of anterolateral spine
with little or no demarcation; sometimes a slight sinuosity
developing, or a more or less insignificant oblique offset;
rarely ever a real offset, notch, or projection with an orbital
spinule on one or the other side at all like the condition
found in either of the two species immediately following;
rostrum broadly lingulate, more or less triangularly so, but
never sharply triangular as in abtao and riolimayana (F”, G’,
and G’ below) ; movable finger with a distinct, usually spined
or spinulated lobe on outer margin near base; palmar crest
not particularly prominent, posterior margin of crest usually
noticeably upturned, troughed or excavate with upturned and
broadly and shallowly serrate margins.
concepcionensis (p. 501)
THE SPECIES OF AEGLA—SCHMITT 457
G. Orbital spine, or the orbital spinule usually taking its place,
generally present on one or both sides of front; extraorbital
sinus well formed but narrow or reduced to a mere notch
between orbital spinule and anterolateral spine; in the
absence of a real orbital spine or spinule (as in about half
the representatives of A. laevis) virtually always a well-
marked, often abrupt, sometimes nearly right-angled offset
between inner slope or margin of anterolateral spine and
outer end of orbital margin; rostrum somewhat narrowly
lingulate, subparallelism of margins of midsection often
rather pronounced.”
H*. Movable finger with distinct and usually spined or spinu-
lated lobe on outer margin near base; palmar crest only
somewhat excavate or impressed with upturned and dis-
tinctly serrate spine or sharp-scaled tipped margins.
laevis (p. 504)
H’, No lobe on outer margin of movable finger near base; palmar
erest noticeably excavate, impressed, or longitudinally
troughed, margins upturned and more or less entire, obso-
lescently if at all serrate (remotely somewhat reminiscent
of the palmar crest in odebrechtii).
laevis taleahuano (p. 508)
F*. Rostrum distinctly and sharply triangular, lateral margins taper-
ing from base to tip (in no part at all subparallel), rostrum
in lateral view running about or nearly straight forward, with
only slight if any upward inclination distally (neither upcurved
nor recurved) ; orbital spine or spinule and extraorbital sinus,
though sometimes small or narrow, always definitely present.
G*. Rostrum moderately broad and, though sharply triangular,
rather broadly so, gradually and not particularly narrowed
distally; rostral carina dorsally furnished with two more or
less distinct rows of corneous seales for greater part of
length, anterior to middle of free portion of rostrum two
rows OF Scales running together to form a single sometimes
somewhat scattered row, which continues about to the
anterior extremity ; areola widening behind____ abtao (p. 510)
G’. Rostrum narrowly and sharply acuminate (stilletolike) ; rostral
earina sharp crested for greater part of its length and fur-
nished with a single at times slightly wavering row of
corneous scales, which in some specimens tends to become a
double row of more or less closely juxtaposed scales a little
before distal extremity of rostrum; areola narrowing
POSteriOr] y26 = Sis _ SUES PETE) aL riolimayana (p. 513)
143 The rostra of the two species falling within this section of the key, in general, so far as
the specimens I have seen are concerned, look somewhat amorphous, as if they had been
partially melted and then solidified.
458 PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 91
Family AEGLIDAE
Genus AEGLA Leach
AEGLA PARANA, new species
Fieures 42, 48; PLATE 25, A
Description.—A large species attaining a length of carapace and
rostrum together of at least 44 mm.
Carapace slightly convex anteriorly and laterally, medially quite
flattened; front very wide. Rostrum long, slender-spinelike, sharply
carinated, ridge-roofed, triangular in cross section, exceeding eyestalk
by two to three times the length of the cornea; crest of rostral carina
furnished with a closely juxtaposed double row of good-sized corneous
scales about to level of corneae, anterior to which the row becomes
single with scales often closely set, sometimes a bit separated from one
another; posteriorly the carina proper ends just before the anterior
margins of protogastric lobes, larger scales of carina often stop at
level of epigastric prominences. Epigastric prominences well marked,
though low tubercular, furnished with one or more, usually several,
corneous scales, individually about the size of the scales on the
rostrum; anterior margin of protogastric lobes, though only slightly
raised, distinctly marked, in part at least, by a short row of sizable
corneous scales, of which the apical one is larger and heavier than
the others. Areola relatively long and narrow, lateral sutures of
cardiac area markedly converging behind.
Orbits fairly wide, moderately deep, separated from the wide extra-
orbital sinus either side by a conspicuous strong yet slender spine;
the extraorbital sinus exceeds half the width of the orbital one, often
about equal to three-fourths of its width.
Anterolateral spine long, strong, acuminate, reaching to middle of
cornea or beyond, sometimes nearly as long as eye. Anterolateral
angle of first hepatic lobe sharply and strongly spined, spine more
or less exserted, second lobe may also be spined, or, like the third,
carry a good-sized corneous scale. Angle on lateral margin behind
cervical groove spiniform and armed with one, usually several,
smaller, sharp, corneous-tipped spines on its posterior slope; angle
behind notch which follows the preceding angle also spined; entire
lateral margin of posterior portion of carapace (behind cervical
groove) conspicuously armed with a continuous fringe of sharp
spines; other species may have the corresponding margin more or
less small-spinulose or scabrous, but in none (except A. denticulata)
is it as strongly and well spined as in this one.
Large hand more or less subquadrate, thick, but not inflated or
particularly swollen looking, moderately rough scabrous, armed on
THE SPECIES OF AEGLA—SCHMITT 459
é a
Ficure 42.—Aegla parana, new species, male holotype: a, Dorsal view; b, lateral view of
anterior portion; ¢, sternum of third and fourth thoracic somites; d, inner ventral margin
of ischium of left cheliped; ¢, lateral view of second abdominal epimeron. a, b, natural
size; c—e, twice natural size.
Ficure 43.—Aegla parana, new species, male paratype: Chelipeds, showing variation in
shape of hands and palmar crest. Natural size.
460 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
outer margin (of both hands) with a number of spinuliform scales,
or sharply pointed, short spines. Movable finger with no true lobe
on outer margin near base, at best a low, scabrous thickening, but
so slightly developed that it in no sense can be considered a lobe
such as is found in certain other species of Aegla; both fingers with
a stout lobular tooth. Palmar crest a comparatively low ridge,
broadly serrate, angles of serrations sharp-spined ; sometimes (fig. 43)
serrations are virtually obliterated so that free edge of crest is nearly
straight, and furnished with some corneous, perhaps pointed, scales
and a fair-sized spinule anteriorly and posteriorly.
Carpus sharply and strongly spined on inner margin, ridge above
this row of spines also sharply and strongly spined; apparently there
is an additional longitudinal row of spines running along the median
line of the dorsal surface of the carpus; this normally seems to be
armed with three good spines, sometimes one or both of the posterior
spines may be reduced to a stout scale, or a short-conical spinule.
Dorsal longitudinal margin of merus of cheliped armed with a row
of large, sharp, well-developed spines; at middle of anterior margin
of merus a strong spine about as large as anterior spine of dorsal
longitudinal margin. The inner margin of the ventral surface of
the ischium of the cheliped is armed with at least two fairly long,
strong, more or less subequal corneous-tipped spines; among the
Aeglas two ischial spines of this size and prominence are found only
in this species and A. sanlorenzo (see also last paragraph under
“Remarks,” A. castro, p. 475).
Meri of ambulatory legs likewise normally armed with a series of
strong spines along upper margin; sometimes the series is not quite
so large and regular as in the type, yet enough of it is present
to distinguish this species from all other Aeglas by this feature alone;
near distal end of posterior margin of ventral surface of merus, at
the level of the posterior end of the articular membrane of the joint,
there is a strong spine, behind this there may be a second smaller
one, and at the extreme anterior end a small spine or two.
Anterior dorsal angle of epimeron of second (in lateral view, ap-
parent first) abdominal somite produced into a long, sharp spine
strongly buttressed behind by a conspicuous ridge or angle running
obliquely longitudinally back on the epimeron; anterior margin be-
low this spine deeply concave, ventral angle narrowly produced, sub-
acute and often, as in the type, tipped by a strong corneous spine.
Color—A. parana is very beautifully marked. The general body
or ground color is a dark, almost black, bottle green; in one instance
a dark grass green with faint suggestions or touches of parrot green;
sometimes bister X olive-green to a blackish bister with raw-umber
higher portions.
THE SPECIES OF AEGLA—SCHMITT 461
The chelipeds and chela for the greater part have the same general
color as the rest of the body, except that as much as the distal half
of the fingers may be a bright French or a dark turquoise blue; the
dark grass-green specimen has marine or indigo blue on the fingers
of the left hand and royal purple on the right; one other specimen
has the greater part of the hand Indian purple with prune purple
distally on the fingers.
The most proximal portions of the chelipeds and ambulatory legs,
more or less hidden by the lateral margins of the carapace, take on
a dirty cream-buff to clay color; the under parts of the body are
similarly colored, except that the sternum sometimes is a Mars brown,
and the outer surface of the turned-under abdominal somites and
telson are often faintly tinged with a greenish, bluish, or purplish
color much like a poorly dyed, plain-colored Easter egg. The ambu-
latory legs, usually greenish like the body, are sometimes flushed with
purple or blue, especially the under side of the dactyls; in other
specimens they may be an almost buff or dirty cream-buff; in two
cases it was noted that the articulating membranes are brightly col-
ored ferruginous in one, coral red in the other. Distally, the third
maxillipeds at least occasionally are faintly tinged with blue, or the
last joints even take on a turquoise blue color. The antennal flagella
are usually colored like the carapace. (For colors see Ridgway, 1886.)
Holotype.—A large male, U.S.N.M. No. 80016, the largest of several
collected at Rio Negro, October 21, 1925, in a wicker fishpot kindly
baited and provided by Carlos Zornig, of the Hotel Zornig. This is
the largest individual Aegla I have ever seen. It measures a full 44
mm. in length of carapace and rostrum together and 75 mm. from tip
of rostrum to posterior margin of telson extending abdomen as much
as possible without breaking; from telson margin of extended abdo-
men over extended chelipeds, 108 mm.
Distribution.—The species so far has been collected only at Rio
Negro, Parana, Brazil, where I secured a modest number of speci-
mens by means of the fishpot and also a cast net used by a local
fisherman at night over a brief period from October 12 to 14 and
again on October 21 and 22. On the early morning of the 14th the
air temperature was 58° F., while the water near the bank at about
a foot below the surface registered 64° F.
AEGLA SANLORENZO, new species
Ficure 44; PLaAte 25, B
Description—The unique type male is a specimen of just about
29 mm. in length of carapace and rostrum taken together. The arms
are broken and the right, minor hand is shattered; only the first left
leg is complete, though detached. In the accompanying drawing the
specimen is “restored.”
462 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
Carapace slightly to moderately convex, front wide. Rostrum
moderately long, spinelike, triangular in cross section, exceeding
eyestalks by about three times the length of the cornea; rostral carina
sharply ridged, furnished with a double row of light corneous scales
closely juxtaposed and more or less alternating up to a little anterior
to the level of the posterior margin of the orbits, where the scales
form a single, closely set row of scales which extends to the anterior
extremity of the rostrum.
é a
Ficure 44.—Aegla sanlorenzo, new species, male holotype: a, Dorsal view; d, lateral view of
anterior portion; c, sternum of third and fourth thoracic somites; d, inner ventral margin
of ischium of left cheliped; e, lateral view of second abdominal epimeron. a, d, natural
size; c—e, twice natural size.
Epigastric prominences low, with few small beadlike scales on sum-
mit; anterior margins of protogastric lobes forming an acute angle
outlined by a closely-set row of light-colored scales; similar scales
scattered elsewhere over carapace. Areola moderately wide, fairly
long.
Orbits only moderately wide, moderately deep, separated from the
fairly wide extraorbital sinus by a well-developed orbital spine; ex-
traorbital sinus about one-half the width of orbital.
Anterolateral spine long and slender, sharply spiniform, exceeding
the cornea. First hepatic lobe sharply spined anteriorly, spine end-
ing in a slender corneous tip and appreciably exserted; second and
third hepatic lobes set off by not very prominent, rather weak notches,
margins corneous-granulated or scaled.
THE SPECIES OF AEGLA—SCHMITT 463
Larger hand very smooth appearing, but under glass finely granu-
lated (or minutely scaled like the carapace), more or less subrec-
tangular, gently convex, rising to an apparent median longitudinal
angle extending from a little distance behind the posterior margin
of the sinus between the fingers to the posterior margin of the palm;
inner margin of palm can scarcely be said to be crested, it is broadly
rounded off but rises at a little distance before the anterior border
to form a conspicuous, though short, acutely corneous-tipped spine;
the smaller hand of this unique specimen is crushed but seems to have
the same conformation as the larger one. There is no lobe on the
outer margin of the movable finger near the base; the prehensile
margins of both fingers are slightly sinuous, but neither reveals any
trace of the large lobular tooth found in most species of Aegla.
Carpus of cheliped granulated like hand; ridge above spined inner
margin more or less obsolescent, at least not very prominent, lobe at
anterior angle produced to form a strong prominent spine. Dorsal
longitudinal margin of merus strongly and sharply spined above;
anterior margin unarmed, finely scabrous. Inner margin of the ven-
tral surface of ischium is armed with a pair of well-developed strong
spines; only on the left (figured type) ischium does a small acute
spine intervene between the two large spines; on the right the inner
margin of the joint is uninterrupted.
Merus of first ambulatory leg scabrous above; armed with an an-
teriorly directed spine on the posterior border of the ventral surface
a little behind the level of the posterior margin of the articular mem-
brane; there is also a small corneous point or spine close to the anterior
end of the ventral margin.
Anterior dorsal angle of epimeron of second (in lateral view, ap-
parent first) abdominal somite produced into a slender, sharp spine;
margin of the epimeron below this spine deeply concave; ventral
angle strongly and narrowly produced, though bluntly rounded off
at its extremity.
Holotype.—The unique male specimen collected by Dr. Carlos Speg-
gazzini in the Rio San Lorenzo, Salta, Argentina (M.A.C.N.™ No.
7099) ; length of carapace and rostrum taken together, 29 mm.
Remarks.—This species is certainly more nearly related to A. parana
than to A. uwruguayana, which it superficially resembles. The strong
ventral spine on the ambulatory legs and the shape of the epimeron
of the second abdominal somite point in the direction of A. parana;
moreover, the inner ventral border of the ischium of the cheliped,
like that of A. parana, is armed with a strong hooked spine at the
anterior end as well as at the posterior end of the joint but, unlike
14 Museo Argentino de Ciencias Naturales.
435661—42 3
464 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
it, it may have a small tubercular or nodular projection intervening
between the anterior and posterior spine.
The hands, however, appear to resemble more closely those of
A. uruguayana in most particulars: Low or no crest, anterior sharp
spine on inner margin, and no lobe on outer margin of movable finger.
The palm of A. sanlorenzo is relatively shorter than that of A. wru-
guayana, and the fingers lack the lobular teeth present in the last-
named species.
AEGLA PLATENSIS, new species
FIGuRES 45, 46; PLATE 25, C
Aegla laevis R. von IHERING, Atlas da fauna do Brasil, pl. 4, fig. 17,’° 1917.
Description.—A large species, attaining a length of carapace and
rostrum together of about 39 mm.
Carapace, though gently convex, more or less flattened, front very
wide. Rostrum an elongate ridge-roofed, narrowly triangular spine,
exceeding eyestalks by about twice the length of the cornea; rostral
carina somewhat blunt, only fairly sharp ridged, furnished with
three to five rows of cornified, sometimes almost microscopic punctae,
except very close to anterior extremity of rostrum, where there is an
irregular, short, single row of larger corneous scales; carina runs back
as far as the anterior margins of the protogastric lobes, neither proto-
gastric lobes nor epigastric prominences at all well marked. Anterior
margins of protogastric lobes broadly obtuse angled, not at all tuber-
culiform at apex of angle. Areola widens noticeably behind.
Orbital sinus wide, but only a little longer and a little wider ap-
pearing than extraorbital sinus, orbital spine well developed. An-
terolateral spines large and conspicuous, reaching nearly or about
to middle of cornea. Anterolateral angle of first hepatic lobe is
produced into a prominent, sharply acute spine; second and third
hepatic lobes may be indicated, but are not at all well marked; if
spinulated, spinules no larger than spinules found elsewhere on lateral
margin of anterior portion of carapace; occasional specimens may
have a small notch marking the second hepatic lobe on one or the
other side of the carapace, perhaps never on both sides.
Hands large, broadly ovate, much flattened as compared with most
species of Aegla. Movable finger more or less cylindrical, rather
slender in well-developed specimens, and arched, making a consider-
able gape between the fixed and movable fingers; movable finger with
1 This figure of Rudolfo von Ihering is original and is undoubtedly based on one of a
lot of specimens collected by his father Hermann von Ihering, in the State of Rio Grande
do Sul (collector’s No. 619). The Rio Grande do Sul record given under “Distribution”
of A. platensis below is also based on a specimen from that lot of material, presented to
the U. S. National Museum by Dr. H. von Ihering in 1915. A comparison of this specimen
and the figure convinces me that A. platensis is the species represented.
THE SPECIES OF AEGLA—SCHMITT 465
a noticeable lobe at base, blunt angled in the largest specimens (and
in the type) but usually sharp angled and anteriorly spined at least
in specimens up to 33 mm. in length of carapace and rostrum taken
together. Upper margin of palms somewhat compressed, forming a
low ridge (palmar crest), most developed at its posterior angle, or
“heel”; margin of crest more or less irregular, angulations armed
with small, sharp, corneous spines or spinules, sometimes corneous
é a
Ficure 45.—Aegla platensis, new species, male holotype: a, Dorsal view; b, lateral view of
anterior portion; c, sternum of third and fourth thoracic somites; d, inner ventral margin
of ischium of left cheliped; ¢, lateral view of second abdominal epimeron. 4, }, natural
size; c-e, twice natural size.
spinulate at anterior angle, border of posterior angle, or heel, some-
what upturned, forming a very slight, short, very shallow trough
between border of “heel” and margin of palm proper. In young
specimens the margins of the crest may be quite spiny, but this
condition is not carried over into the more developed, adult stages.
Ridge of carpus of cheliped above inner spined margin somewhat
lumpy and obliquely scabrous ridged, but not spined; anterior inter-
nal lobe or angle of carpus produced into a short, stout, conical spine.
Upper longitudinal margin of merus with a strong, moderately stout
to slender spine at anterior end; anterior margin with only a slight,
466 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
denticulated convexity on margin in line with spine at anterior end
of dorsal longitudinal ridge. Inner margin of ventral surface of
ischium not spined, at most with only a low swelling at anterior end,
and perhaps a very slight convexity at posterior end.
Anterior dorsal angle of epimeron of second (in lateral view, ap-
parent first) abdominal somite produced to form an acute corneous-
tipped spine buttressed behind by a blunt ridge or thickening of
epimeron; anterior margin below spine more or less straight, at most
only slightly concave; ventral angle rounded off.
Ficure 46.—Aegla platensis, new species, male paratype. Natural size.
Holotype —The largest male, U.S.N.M. No. 80018, from a lot of
2 males and 2 females collected at “Isla Flores” [? Tigre, Buenos
Aires, Argentina] by Dr. W. E. Safford, U.S. N., at the time attached
to the U. S. S. Mohican, May 4, 1887. This specimen measures
slightly over 38 mm. in length of carapace and rostrum together;
the largest female is 33.5 mm. long.
Remarks.—This species and the next are in many respects very
similar. They differ, however, in a number of particulars. The
movable finger in this species has a lobe on the outer margin near
the base; no such lobe seems ever to be developed in any specimen
of A. uruguayana, male or female; moreover, in case of doubt, the
presence of a well-developed sharp spine at the anterior end of the
inner border of the ventral surface of the ischium of the cheliped
THE SPECIES OF AEGLA—SCHMITT 467
will always distinguish A. wruguayana from A. platensis, even in
very small juvenile specimens.
In well-developed females of A. platensis the hands are flatter
than in the males, and also somewhat narrower; the fingers are much
less strong, and more slender.
The sternal plate between the chelipeds carries a low, blunt keel,
which anteriorly may at times be raised a bit or project forward as
a low, ventrally keeled, conical tubercle; there is some suggestion of
similar keeling on the following sternum between the first pair of
ambulatory legs, which, though elevated about as much as the pre-
ceding keel, forms a very broad, low swelling, larger and broader
at the anterior end than at the posterior.
A. uruguayana has a low median swelling on the anterior half of
the sternum between the chelipeds, a little peaked at the forward
end, but not appearing so keeled as in A. platensis,; often in specimens
of medium size this swelling or projection takes on the form of a
stout, conical, corneous-tipped spine inclined obliquely forward.
Distribution.—In addition to the type lot, I have seen various speci-
mens from the vicinity of Buenos Aires and from Tigre nearby,
where Dr. Martin Doello-Jurado, director of the Museo Argentino,
most kindly took me collecting one day; from the Prado and the
Arroyo Miguelete, Montevideo, and Bahia de Colonia, Uruguay; Rio
Grande do Sul, Brazil; and one specimen that appears to be this
species from Tucuman, Argentina.
AEGLA URUGUAYANA, new species
FIGURE 47; PLATE 25, D
Description.—A species of good size, attaining a length of carapace
and rostrum together of 33 mm.
Carapace moderately convex, well areolated, front wide. Rostrum
long, slender, and sharply acuminate, above lateral margins distinctly
triangular in cross section; rostrum in the type specimen exceeds
the eyestalks by 114 to nearly 2 times the length of the cornea (in
very small specimens rostrum may be only little longer than eye-
stalks) ; rostral carina prominent, multiscaled, scales intermingled,
plainly marked backward to a little behind the level of the anterior
margin of the protogastric lobes. Epigastric prominences just low
swellings situated on the forward slope of the carapace between the
orbital margin and the much higher lying anterior margins of the
protogastric lobes; the anterior margins sharply marked by a row
of five or six light corneous beadlike scales. Areola of good size.
Orbits very wide and shallow, distinctly set off from extraorbital
sinus by an orbital spine of good size, extraorbital sinus about three-
fifths as wide as the orbital sinus.
468 PROCEEDINGS OF THE NATIONAL MUSEUM You, 91
Anterolateral spines of carapace scarcely reach posterior margin
of cornea, in some specimens a little beyond this level. Anterolateral
angles of all three hepatic lobes well marked, at least the first (in the
type all three) sharply acute and spined; first spine long and slender
and appreciably exserted; the second about half the length of the
first; the third in the type as much reduced again.
Large hand quite smooth appearing, only very finely scabrous,
elongate, subrectangular, upper surface gently convex, with pair of
faint yet discernible low obsolescent ridges converging from each of
é a
Ficure 47.—Aegla uruguayana, new species, male holotype: a, Dorsal view; b, lateral view
of anterior portion; c, sternum of third and fourth thoracic somites; d, inner ventral
margin of ischium of left cheliped; ¢, lateral view of second abdominal epimeron. 4, b,
natural size; c-e, twice natural size.
the posterior upper angles of the palm to meet and become one about
the middle of the length of the palm, shortly thereafter to fade out
before reaching the posterior margin of the sinus between the fingers. ©
No lobe on outer margin of movable finger near base; tooth on fixed
finger well developed. Virtually no palmar crest, inner margin of
palm more or less obsolescently and rather broadly carinated, carina
armed anteriorly with a sharp corneous spine.
Carpus of cheliped with acutely spined lobe at anterior-internal
angle. Dorsal margin of merus armed with a longitudinal row of
strong spines; at anterior end this row of spines appears to turn
THE SPECIES OF AEGLA—SCHMITT 469
inward *° for inside and often a little in advance of the anteriormost
of the longitudinal series there is one and sometimes two or more
almost equally strong, though usually somewhat more slender, spines
in an oblique row (the second spine of this row is always smaller
than the first and if there are additional spines they are in turn
smaller than the second one); anterior margin of merus with small
rounded lobe or tubercle. The inner margin of the ventral surface
of the ischium is armed with a well-developed spine anteriorly and
only a low swelling or slight nodulation at the posterior end.
Anterior dorsal angle of the epimeron of the second (in lateral
view, the apparent first) abdominal somite much produced, ending
in a sharp corneous spine; anterior margin of this epimeron below
the spine slightly concave and nearly vertical in direction; ventral
angle very little less than a right angle, apically rounded off.
Holotype.—The only large specimen, a male, in a lot of 2 males
and 4 females, of which the rest are all under 14 mm. in length of
carapace and rostrum taken together. This measurement in the holo-
type about equals 33.3 mm. These specimens were obtained by the
Captain Marshall Field Brazilian Expedition of the Field Museum,
October 20, 1936, 14 kilometers northeast of San Carlos, Uruguay,
Karl P. Schmidt collector, and are in collections of the Field Museum.
The holotype carries Field Museum number 2287; paratypes, 2288.
Remarks—This species is characterized by its long, slender ros-
trum, triangular in cross section, or, as one might say, ridge-roofed
rostrum; the only slightly convex, more or less subrectangular, virtu-
ally uncrested hands; and by the distinctly marked hepatic lobes of
which the anterolateral angles of at least the first two and often all
three are spined. (See also “Remarks” under A. platensis and A.
prado.)
Distribution.—This species seems to be widely distributed on both
sides of the River Plate, definitely eastward as far as Punta del Este,
Uruguay ; south and westward to Buenos Aires, Isla Flores, Belgrano,
and Lujan, Province of Buenos Aires, Argentina; north and west-
ward to Paysandu, Uruguay; and Concordia and Parana, Entre Rios,
Argentina. One specimen, a small male, one of the Aeglas examined
by Nobili, from San Luis, Argentina, received from the Turin Mu-
seum, seems to be near, if not identical with, this species. It is,
however, rather far removed from the above indicated range of A.
uruguayana. This may be due to the lack of collections from the
intervening region, or perhaps even to the lack of development of
the specific characters in this small specimen.
I have seen specimens from the above-mentioned range-determining
localities and also from Paso de la Arena, Arroyo Miguelete (very
16 A somewhat similar condition occurs in A. affinis, p. 495.
470 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
small specimen, determination doubtful), St. Lucia, River San José,
Rosario, from near Carmelo, Nueva Palmira, and Frey Bentos, Uru-
guay; and Arroyo El Gato, Guateguaycht, Entre Rios, Argentina.
One small lot examined (M. C. Z. No. 10478) was labeled Maldonado,
Brazil (I believe that this should be Maldonado, Uruguay).
AEGLA PRADO, new species
Ficures 48, 49; PLATE 26, A, B
Description.—A small to moderate-sized species. One of the larg-
est specimens I have seen measures about 25.5 mm. in length of
earapace and rostrum taken together.
Carapace usually very convex, more so than in any nearly related
species; front fairly wide, narrower than in A. platensis. Rostrum
sharp, spinelike, ridge-roofed, exceeding eyes by at least twice the
length of the cornea; the rostral carina is furnished with several lon-
gitudinal rows of irregularly placed, tiny corneous scales; the carina
is continued backward past the anterior margins of the protogastric
lobes, at the level of which it widens out to form a low, blunt ridge
that may be more or less readily traced to the posterior margin of the
carapace; it is interrupted only by the cervical groove; this ridging
or transverse angling of the median line is not so prominently devel-
oped in all the specimens at hand, yet it is a conspicuous feature in
a very considerable number of the larger representatives of the species.
Though otherwise quite distinct this was the first species I personally
encountered in South America that had any real resemblance to
Nicolet’s prominently keeled Chilean A. denticulata.
Epigastric prominences are low to obsolescent swellings; anterior
margins of protogastric lobes sharply acute-angled, apex raised up and
almost small-tuberculiform, more prominently so in the smaller than
in the larger specimens.
Orbits of good size, much larger than extra-orbital sinuses, which
are relatively moderate to small in size; orbital spine small, standing
fairly close to anterolateral spine.
Anterolateral spines well-developed, reaching not quite to middle
of cornea. All three hepatic lobes well marked and corneous spined,
and each well set off from the others, so that the lateral margin of
the anterior portion of the carapace narrows stepwise from the cervi-
cal groove to the anterolateral spine.
Hands very swollen looking, more or less broadly ovate. Movable
finger with a plainly marked, generally small-spined lobe on outer
margin near base. No particular crest developed on inner margin of
palm, and no such posterior angle or “heel” as in A. platensis; how-
ever, there is a noticeable spine or two (sometimes more, and then
THE SPECIES OF AEGLA—SCHMITT 471
Ficure 48.—Aegla prado, new species, male holotype: a, Dorsal view; b, lateral view of
anterior portion; c, sternum of third and fourth thoracic somites; d, inner ventral margin
of ischium of left cheliped; e, lateral view of second abdominal epimeron. a, b, natural
size; c—e, twice natural size.
Ficure 49.—Aegla prado, new species, male paratype: a, Dorsal view; 4, merus of first right
ambulatory leg. This specimen has avery prominently ridgedcarapace. The hands are
less typical, the larger has perhaps been recently regenerated; likewise, the first left
ambulatory leg is certainly relatively feebler than the other legs of this same specimen and
lacks the ventral meral spines present on the first right ambulatory leg and both of the
first pair of ambulatories of the type. a, natural size; b, twice natural size.
472 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
smaller spines) in line on the inner margin of the palm a little
behind its anterior border; outer margin of hand somewhat small-
spinulose, occasionally with a larger spinule or spine.
Anterior internal lobe or angle of carpus of cheliped forming a
stout, acute, conical spine. Upper margin of merus with a straight,
longitudinal row of sharp spines, no inward turn at anterior end
as in A. uruguayana; anterior margin of merus scabrous or small
denticulate. Ischium below on inner margin armed with a well-
developed sharp spine at anterior end, a prominent feature even
in quite small specimens; at posterior end a low conical tubercle or
nodule, often with acute corneous tip (in only one of well-developed
males was there a fairly sharp spine at the posterior end of the
ischial border in addition to the much stronger spine at the anterior
end).
The first ambulatory merus has a spine of fair size developed on
the posterior ventral margin at about the level of the proximal margin
of the articulating membrane, besides the smaller spine at the distal
end of this same margin. With respect to this ventral meral spine,
A. prado reveals kinship to A. parana and A. sanlorenzo, though quite
different from them in a number of other respects, particularly in its
smaller extraorbital sinuses, and therefore only moderately wide
front.
Anterior dorsal angle of epimeron of second (in lateral view, ap-
parent first) abdominal somite spined, anteroventral border almost
straight to very slightly concave, ventral angle rounded off.
Sternal plate between chelipeds carries a median, corneous, spine-
tipped, conical tubercle; even in very small specimens this sternal
spine is of good size, well formed, and sharply acuminate.
Holotype.—One of the larger males of a sizeable lot of specimens,
U.S.N.M. No. 80017, collected in a small tributary of the Arroyo
Miguelete in the Prado, Montevideo, by the late Dr. Juan Tremoleras
and myself, December 1, 1925. This specimen, the second largest
male, is 25 mm. in length of carapace and rostrum taken together; the
largest male, is 25.5 mm., the largest female 21 mm. long; included
in the material are a considerable number of juveniles between 10
and 15 mm. long. These Aeglas were plentiful under the grass and
vegetable debris that carpeted this very shallow stream, perhaps be-
cause of the numerous fragments of picnic lunch, bits of bread and
meat scraps, that had been thrown into the water. The water tem-
perature was between 28° and 29° C.
Remarks.—This species and A. uruguayana are much alike in gen-
eral appearance, though very probably not in color in life. Most
specimens of the latter that I have seen are very light colored in
alcohol; A. prado, on the other hand, is quite dark, even the specimens
that I collected 17 years ago.
THE SPECIES OF AEGLA—SCHMITT 473
The stepwise arrangement of the well-marked hepatic lobes and
the frequently strongly ridged carapace tend to set this species
apart from those that are most closely related to it. As in A.
uruguayana, there is a sharp to spinous tipped tubercle on the anterior
sternite, but in the present species it is larger, usually sharper, and
more erect, forming roughly an angle of about 45° with the general
surface of the sternite. The ventral inner ischial borders of the
chelipeds are similarly armed in the two species, but in A. prado the
posterior tubercle is more prominent, larger, higher, and more pointed,
occasionally quite spinelike; in small specimens it is already sharp-
pointed and readily hooks or engages a needle drawn backward along
the ischial border; in small as well as large A. wrwguayana posteriorly
there is but a small low tuberclelike swelling or small nodulation
which often is relatively inconspicuous.
Distribution.—A. prado, so far as at present known, has been found
only in watercourses in and about the city of Montevideo. Dr.
Florentino Felippone, long a valued correspondent of the United
States National Museum, collected 2 males and 2 females of this
species in the Miguelete on December 6, 1922, along with four smaller
specimens of A. platensis. More recently, Alberto Tremoleras, son
of the late Dr. Juan Tremoleras, of Montevideo, Uruguay, kindly
collected for us a lot of 19 females in Arroyo Malvin, January 21,
1936, about 2 kilometers from its mouth. Of these, 16 were oviger-
ous specimens. He noted on the label, “fresh water, partly stagnant.”
AEGLA CASTRO, new species
FIGURE 50; PLATE 26, F
Description—A small species of which the largest specimen I
have seen measures 28.5 mm. in length of carapace and rostrum
together.
Carapace moderately convex. Rostrum an elongate, triangular,
ridge-roofed spine, exceeding eyestalks by about 114 times the length
of the cornea; rostral carina well defined, furnished with about two
rows of more or less alternating, often closely set, small, corneous
scales; the rostral carina posteriorly merges with the general surface
of the carapace on a level with the protogastric lobes.
Epigastric prominences somewhat rounded, blunt tuberclelike;
anterior margin of protogastric lobes forms a conspicuous obtusely
angled ridge or elevation which at its apex may be slightly scabrous.
Orbit wide, orbital spines well set off from anterolateral spines by
a small to moderately wide extraorbital sinus.
Anterolateral spine of carapace fairly slender, reaching to middle
of cornea or beyond. All three hepatic lobes usually plainly indi-
474 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
cated; only the first has its anterolateral angle spined, and forms
an offset in the general trend of the anterolateral margin of the
anterior portion of the carapace.
Large hand moderately inflated, somewhat elongated. Movable
finger carries a well-formed, often small spiny lobe on outer margin
near base. Palmar crest fairly large, conspicuous, somewhat sub-
disciform, distinctly shallowly impressed or excavate with upturned,
more or less serrate, and definitely sharply spinulose margins; outer
margin of hand finely spinulose.
é a
Ficure 50.—Aegla castro, new species, male holotype: a, Dorsal view; b, lateral view of
anterior portion; c, sternum of third and fourth thoracic somites; d, inner ventral margin
of ischium of left cheliped; ¢, lateral view of second abdominal epimeron. a, b, natural
size; c—e, twice natural size.
Ridge above inner spined margin of carpus armed with conical
tubercles, of which the greater part are more properly acute conical
spines; the anterior internal lobe or angle of the carpus is broadly
conical and tipped with a small sharp corneous spine; upper margin
of merus armed with slender spines, of which the most anterior and
sometimes the largest is situated directly on the anterior margin of
the merus, the next spine may be slightly larger or slightly smaller
than the anteriormost spine. Inner margin of ventral surface of
ischium also has a strong conical spine at anterior end, and generally,
in addition, a smaller one of variable size and acuity at the posterior
end, and a much smaller one or two in between.
Meri of ambulatory legs with a small spinule or two near anterior
end of lower outer margin on level with posterior portion of articular
membrane or behind it, perhaps to some degree comparable to the
similarly placed but relatively ever so much larger, conspicuous
THE SPECIES OF AEGLA—SCHMITT 475
spines in A. parana and A. sanlorenzo, and not quite so prominent
one in A. prado.
Anterior dorsal angle of epimeron of second (in lateral view, ap-
parent first) abdominal somite produced and well spined, anteroven-
tral margin about straight; the ventral angle is rounded off.
Holotype.—The largest and best-developed male of a lot of nearly
200 specimens about equally divided between males and females taken
from the Rio [apé in the State of Parana, Brazil, October 1925. The
type, U.S.N.M. No. 80020, measures 28.5 mm. in length of carapace
and rostrum.
Remarks.—The somewhat subdisciform palmar crest of this species
is so strongly reminiscent of that of A. odebrechtii (p. 487) that when I
first found this species in the field I thought I had found the species
described by Fritz Miiller, but the spined dorsal anterior epimeral
angles of our species at once set it apart from his odebrechtii, in
which these angles are rounded off and not spined. Moreover, the
rostral carina and the spined carpal ridge of A. castro are very dif-
ferent. The palmar crest is also very similar to that of A. odebrechtii
paulensis, from which, however, our species may be distinguished by
the same characters that separate it from A. odebrechtit.
In the primarily 2-spined inner ischial margin the present species
has something in common with A. parana, sanlorenzo, and prado,
and also, as suggested above, in the armature of the ventral margin
of the first ambulatory merus. In the first two of these species the
posterior of the two ischial spines is about or nearly equal to the
anterior one; the first and third species appear to have no intervening
conical spines or nodules. On the other hand, in A. castro and in
A. sanlorenzo there usually seems to be an intervening nodule, or
small spine or two. In both A. prado and castro the posterior ischial
spine, even if well developed, is noticeably smaller than, often only a
fraction of the size of, the anterior one.
Color.—In life, a rather uniform very dark olive all over, with
occasional suggestion of olive-green; suture lines a little muddy or
grayish owing to dirt held there; antennae colored like carapace;
antennules brownish gray, in part clay color. Prehensile margins
of fingers of chelae dark orange-chrome, lighter below flushing the
movable finger with color, with a bright spot at the articulation.
Distal half of ambulatory dactyls saturn red to light orange-chrome
suffusing the dark greenish basal half of the dactyls at the juncture
of the two colors. Under parts generally dirty white, central portion
of sternum sometimes with a faint touch of blue (?cerulean blue),
under side of ambulatory propodi and carpi and outer margin of
hands and maxillipeds dirty chromium green (for colors see Ridgway,
1886).
476 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
When turned over these specimens righted themselves very handily,
a faculty not so apparent in the larger parana specimens collected
at Rio Negro, Parana, Brazil. Small specimens would “freeze”
when taken hold of by one leg, but not the larger individuals.
Distribution.—So far collected only in the general region about
the town of Castré6, Parana, Brazil, chiefly in the Rio Iap6 near the
town, and for some distance up and down stream. In obtaining the
considerable series of specimens I brought back with me, I was
most helpfully assisted by the Harry Preston Midkiffs, of the Insti-
tuto Christaio, by Camille Cunha and several of his nephews, and by
Werner Nickol, Conrado Pusch, Amacleto Baptista, and a friend of
theirs who took me on an all-day automobile trip to the Hacienda
Marumby, where we obtained additional material. Air and water
temperatures there were about 68° F. At Castro on October 20 at
about 9:30 a. m. the air was 72° F., water 66° F.
AEGLA FRANCA, new species
FIGURE 51; PLATE 26, D
Aeglea laevis (especie duvidosa) LUEDERWALDT, Rev. Mus. Paulista, vol. 11, p. 431
(sep., p. 5), 1919.
Description—A small species; the largest so far seen attains a
length of carapace and rostrum together of 24 mm.
Carapace moderately convex, front relatively narrow. Rostrum
moderately broad, ridge-roofed, lateral slopes of “roof” may be
slightly concave; exceeds eyes by very little more than the length of
the cornea; carinated to tip, carina furnished with a few irregular,
fairly closely set rows of small corneous scales; posteriorly the
dorsal margin or carina of the rostrum ends in a depression between
and appreciably below the general level of the protogastric lobes of
the carapace; front relatively narrow.
Epigastric prominences not at all well marked, obsolescent; ante-
rior margins of protogastric lobes, on the other hand, are very
prominent, acute angled, and almost tuberculiform apically (some-
what as in A. prado).
Orbital sinus of moderate size; orbital spine small and set close
to anterolateral spine, making extraorbital sinus appear very small,
more a small U-shaped notch than a sinus.
Anterolateral spine appears to be fairly short, yet it reaches at
least to level of middle of cornea, often beyond. Anterolateral angle
of first hepatic lobe acute, corneous-spine tipped, second and third
lobes fairly well marked, scabrous or minutely spinulated, but not
spined.
Large hand only moderately inflated, moderately broad. Movable
finger has a small but definite spined lobe on outer margin near base.
THE SPECIES OF AEGLA—SCHMITT 477
Palmar crest low, narrow; obscurely and irregularly serrate,
spinulose or small spined, margin very slightly upturned. Ridge on
carpus of cheliped above spined, inner margin furnished with small,
more or less transverse scabrous ridges; anterior internal lobe of
carpus subacute with several spinules on its margins besides the
small apical one; upper longitudinal margin of merus with a single
row of sharp spines of which the first is much the longer; on the
anterior margin of the joint in line with the upper marginal row
of meral spines is a low scabrous tubercle. Inner margin of ischium
beneath with a sharp conical spine at anterior end, another usually
slightly smaller one near the posterior end, and one or two much
smaller ones in the interval between the first two.
é a
Ficure 51.—Aegla franca, new species, male holotype: a, Dorsal view; }, lateral view of
anterior portion; ¢c, sternum of third and fourth thoracic somites; d, inner ventral margin
of ischium of left cheliped; ¢, lateral view of second abdominal epimeron. a, b, natural
size; c-e, twice natural size.
Anterior dorsal angle of epimeron of second abdominal somite more
or less blunt-angled; usually, but not always, with one or more tiny
hyaline or corneous spinules or granules at apex of angle; anterior
margin below angle about straight.
Holotype—The largest of 10 males from Franca, State of Sao
Paulo, Brazil, collected in October 1910, by E. Garbe (No. 622),
received some years ago as a gift from the late Dr. Hermann von
Ihering. The type, U.S.N.M. No. 80019, measures 24 mm. long
(carapace and rostrum).
Remarks.—This species and the one following have relatively nar-
row fronts as compared with the several preceding species (A* sec-
tion of diagnostic key). This character and the reduced extra-
orbital sinuses are suggestive of the species that follow (A? section
of key), yet, in general, the more or less ridge-roofed type of rostrum
478 PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 91
and the fact that the rostral carina goes straight through to the tip
of the rostrum seem to identify this species with the A’ rather than
the A® group.
In a measure, perhaps, A. franca and A. jujuyana are to be re-
garded as transition forms lying between those having a ridge-roofed
rostrum and those in which the rostrum is longitudinally more or
less troughed or excavate either side of the median carina.
Certainly A. jujuyana, next dealt with, is very closely related to
A. humachuaca, with which it might have been grouped except for
its sharply carinated rostrum, which for this reason appears to be
more or less definitely ridge-roofed, as the broader, flatter, blunt-
ridged rostrum of A. hwmachuaca decidedly is not. Moreover, the
latter possesses a definite palmar crest of which there is no trace in
A. jujuyana.
Distribution—So far known only from the type locality.
AEGLA JUJUYANA, new species
FIGURE 52; PLATE 26, E
Description—aA_ species of moderate size, attaining a length of
carapace and rostrum together of about 29 mm. Otherwise I have
seen but two small specimens of 18 and 18.5 mm., respectively.
Carapace moderately convex. Rostrum fairly wide-triangular,
scarcely exceeding eyes by the length of the cornea; median carina
sharply crested to the anterior extremity, giving rostrum a definitely
ridge-roofed appearance, particularly in the anterior half or third
of its free portion, even though the lateral slopes of the dorsal sur-
face of the rostrum toward the base of the rostrum are somewhat
concave; rostral carina for whole or greater part of its extent with
a single row of good-sized corneous scales, at least on that portion
of the rostrum lying anterior to the posterior margins of the orbits;
posteriorly the carina scarcely runs back to the anterior margin of
the protogastric lobes; these are low, anteriorly blunt and scarcely
marked except for the few corneous scales outlining them anteriorly.
Epigastric prominences also low, scarcely better developed than the
anterior margin of the protogastric lobes.
Orbital sinus of moderate width; orbital spine small, placed well
up on inner margin or slope of anterolateral spine and set off from it
by a small blunted-V-shaped sinus.
Anterolateral spines, though fairly short, appear moderately slen-
der, reaching at least to middle of cornea or beyond. Anterolateral
angle of first hepatic lobe acute and tipped with a small, sometimes
acute corneous scale; second and third lobes indicated, somewhat
scabrous, second usually a little better marked than the third.
THE SPECIES OF AEGLA—SCHMITT 479
Large hand short, stout, inflated, and smooth appearing; short
fingers gaping, without the usual characteristic lobular tooth of an
Aegla on prehensile margins (there is perhaps a very faint indica-
tion of an obsolescent lobular tooth on the movable finger of the minor
chela) ; no lobe or trace of one on outer margin of movable finger
near base; no trace of a ridge, however faint, on upper surface of
palm. No palmar crest, dorsal margin of palm broad, thick and
rounded off. Ridge on carpus of cheliped above spined inner margin
en
é a
Ficure 52.—Aegla jujuyana, new species, male holotype: a, Dorsal view; b, lateral view of
anterior portion; c, sternum of third and fourth thoracic somites; d, inner ventral margin
of ischium of left cheliped; ¢, lateral view of second abdominal epimeron. a, }, natural
size; c—e, twice natural size.
low and more or less obsolescent (it may be faintly traced for about
two-thirds the length of the carpus), at most only slightly scabrous;
anterior internal lobe of carpus subacute, flattened-conical, armed
with two or three small corneous scales, of which the apical one is the
larger; dorsal ridge of merus of cheliped furnished only with a longi-
tudinal row of small, low, not very conspicuous, scabrous swellings;
anterior margin merely slightly scabrous. Inner margin of ischium
armed with two stout, low, conical, corneous scale-tipped tubercles,
one anterior, one posterior; there may be one or two irregularities,
obsolescent tubercles, or nodules on the inner margin between these
spines.
First ambulatory legs with a small sharp spine or acutely pointed
tubercle near anterior end of ventral margin of merus about opposite
435661—42——-4
480 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
the middle of the length of the articular membrane and a stouter
low-conical one on inner side of ischium near “apex” of ventral face
of this joint.
Anterior dorsal angle of epimeron of second (in lateral view,
apparent first) abdominal somite may be blunt or rounded off, or
armed with a tiny corneous spinule; the anterior margin below the
anterior angle or spinule is very slightly concave. In the largest
of three specimens, the male type, there is a definite small spine on
the left side and none on the right; the other two specimens are quite
small, the larger of these has a corneous spine on the right side and an
almost imperceptible corneous scale or tiny granule on the left; the
smaller has neither scale nor spine on either side.
Holotype—The largest of three male specimens measuring about
29 mm. in length of carapace and rostrum together, collected by An-
tonio Pozzi and Angel Gatta, Rio Chico, Jujuy, 1925 (M.A.C.N. No.
16237).
Remarks.—See under A. franca, above, and A. hwmahuaca, below.
Distribution.—Known only from the type locality.
AEGLA DENTICULATA Nicolet
FIGURE 53; PLATE 26, C
Aeglea denticulata NicoLeT, in Gay, Historia fisica y politica de Chile, Zool.,
vol. 3, p. 200, 1849; Atlas, Crustaceos, pl. 2, fig. 1, 1854.—Girarp, Report
of the U. 8S. Naval Astronomical Expedition to the Southern Hemisphere,
vol. 2, p. 255, 1855 (listed only).
Aegla denticulata, RATHBUN, Proc. U. S. Nat. Mus., vol. 38, p. 602, 1910 (listed
only).
Description.—A. distinctive, well-marked species of good size when
fully grown, attaining a length of carapace and rostrum together of
at least 31 mm. (based on the estimated length of a large specimen
with broken rostrum) ; smallest specimen seen, also a male, 14.5 mm.
Carapace prominently and boldly, but bluntly, keeled for practi-
cally the full length of its median line, interrupted only by the
cervical groove; carapace more ridge-roofed than convex; lateral
margin of posterior portion of carapace behind cervical groove con-
spicuously serrate, first of these saw-teeth just behind cervical groove
larger and broader than anterolateral tooth of carapace, second nearly
equal to first; following teeth of lateral margin decreasing in size
posteriorly to transverse suture line separating the anterior portion
of the branchial region from the posterior; behind this suture line
the margin is scarcely more than small denticulate, almost crenulate
in appearance; the larger teeth or serrations of the lateral margin
are often secondarily toothed or spined on their posterior borders.
Front narrow. Rostrum moderately broad-triangular, scarcely if at
all exceeding eyestalks by as much as the length of the cornea; an-
THE SPECIES OF AEGLA—SCHMITT 481
teriorly the median carina fades out in the distal third of free portion
of rostrum, to become merged in the thickened tip of the rostrum;
there is definite groove or depression either side of the medially raised
portion of the rostrum and its somewhat thickened lateral margins;
the rostral carina, though prominent, has a bluntly rounded-off crest
on which there is a scattering of very fine, almost microscopic scabrosi-
ties. Epigastric prominences low, obsolescent, protogastric lobes
equally poorly developed, causing scarcely more than a break in
reflected light.
7
Figure 53.—Aegla denticulata Nicolet, male neotype: a, Dorsal view; b, lateral view of
anterior portion; c, sternum of third and fourth thoracic somites; d, inner ventral margin
of ischium of left cheliped; ¢, lateral view of second abdominal epimeron. a, b, natural
size; c—e, twice natural size
Orbital sinus fairly narrow, an obtuse-angled V; orbital spine
spiniform, rather high up on inner slope of anterolateral spine; extra-
orbital sinus small, a narrow V-shaped notch. Anterolateral spines
moderately slender-conical, sharply acute, reaching about to or a little
past middle of cornea. Anterolateral angle of first hepatic lobe a
stout, somewhat exserted spine; second and third lobes well marked
by sizable notches, although their anterolateral angles are neither
sharp nor particularly well developed, at most a little scabrous.
Hands, compared with most Aeglas, relatively feeble and under-
developed, small and only lightly convex; prehensile margins of fin-
gers fitting closely together; movable finger with a sharp spinous
lobe on outer margin near base (in the largest specimen this lobe
takes the form of a stout, sharply pointed, conical spine). Upper
margin of palm forming a thick crest conspicuously spined, spines
fairly slender and of good size, usually four spines; sometimes there
is an additional smaller spine inserted near the base of one of the
larger ones.
482 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
Ridge of carpus of cheliped above inner spined margin armed with
four to five sharp spines, occasionally with a few very much smaller
ones in between, sometimes, as in one of the females, these spines
may not be fully developed, for they seem to be represented by
scabrous-tipped tubercles; the spines arming the inner margin of the
carpus are very prominent, long, very strong, particularly the more
anterior, very sharp, and two in number not counting the almost
equally strong spine, which appears to be more properly a part of
the lobe at the upper anterointernal angle of the carpus; in advance
of this particular spine the lobe carries a small, low, but sharp, conical,
and relatively inconspicuous spine. Dorsal margin of merus of
chelipeds armed at anterior end with a large, strong, sharply pointed
spine, followed by perhaps two or three very much smaller ones; a
spine similar to the large spine on the dorsal margin of the merus
but of even larger size arms the anterior margin of the joint; often
this spine has a little sharp spine or spinule on the inner or outer
side of its base.
Inner margin of ventral surface of ischium with a very low, sub-
acute, corneous-tipped cone at anterior end, scarcely developed enough
to be called a spine, followed by three or four more or less equally
spaced little bumps or small nodules which in some cases apically
carry tiny, almost imperceptible, corneous scales.
Anterior dorsal angle of epimeron of second (in lateral view, ap-
parent first) abdominal somite markedly produced, forming an acute
corneous tipped spine which is strongly buttressed behind by a promi-
nent ridge or carina; anterior lateral margin below approximately
straight.
Neotype.—Second largest male measuring slightly over 27 mm. in
length of carapace and rostrum, one of a lot of 10¢ 2° from Orsono,
Chile, collected by the late Dr. C. H. Eigenmann, March 14, 1919
(U.S. N. M. No. 80021).
Remarks.—On the basis of the general character and appearance
of the other species of Aegla described in this paper, Nicolet’s original
description and figure of denticulata scarcely appeared credible; the
rather feeble hands led one to believe he had figured a female; the
dorsal longitudinal keel or ridge running the full length of the cara-
pace seemed an exaggeration; while the large prominent saw-teeth
along the distal moiety of the lateral margin of the posterior portion
of the carapace immediately behind the cervical groove gave the
impression that they were a figment of the imagination. But after
seeing the specimens of A. denticulata collected by Dr. Eigenmann,
here redescribed, I am willing to believe that almost anything in the
way of ornamentation and spining may be possible in the Aeglas.
Nicolet’s apparently crude figure has proved to be a surprisingly
THE SPECIES OF AEGLA—SCHMITT 483
accurate portrayal of the salient characters in nearly every particular,
including the sharply spined epimeral angle and the stout meral spines
of the cheliped, as well as the row of spines on the carpus above the
spined inner margin of this joint; only the middorsal row of scabrosi-
ties of the carpus are a little too prominent in his figure.
A. denticulata is virtually in a class or group apart from all other
Aeglas; only A. prado, which I discovered and described before I
came upon this denticulata material, at all approaches it, and then
only in the keeling of its carapace in certain specimens, and also,
to a slight degree, in the spining of the palmar crest and the inner
margin of the carpus of the chelipeds.
Distribution—As Nicolet says, “found in the republic” of Chile,
but, so far as I know, the only specimens that have been seen since
his time, 1849, are those from Osorno redescribed here.
AEGLA PAPUDO, new species
FIGURE 54; PLATE 27, C
Description.—A. species of moderate size, attaining a length of
carapace and rostrum of at least 26 mm.
Carapace very convex, perhaps more so than any other species of
Aegla, especially across the gastric region. Rostrum more or less
elongate-triangular yet along the middle of its length, in small part
at least, with its lateral margins approximately subparallel; basally
the rostrum is transversely fairly flattened and depressed either side
of rostral carina; the rostrum has a strong downward trend, but its
distal portion is markedly recurved; rostrum extends at least the
length of the cornea or a little more in front of the eyestalk; either
side of its median carina the rostrum is a little troughed or excavate;
the carina extends forward only from one-half to not more than
two-thirds the length of the free portion of the rostrum; beyond the
anterior end of the carina the dorsal surface of the rostrum is gener-
ally for the most part gently concave from side to side and usually,
but not always, without any but a slight trace of the carina or any
corneous scaling in line with that on the carina itself; the corneous
scales on the carina are very dark brown, thick, and almost beadlike ;
the carina runs posteriorly almost to the anterior margin of the pro-
togastric lobes, its dorsal beading, however, extending back only to
about halfway between the epigastric prominences and the anterior
margins of the protogastric lobes; the carinal beading forms a single,
virtually straight, at times slightly wavy row of scales.
The epigastric prominences are subacute-tubercular and topped
with 2 to 6 beadlike scales like those on the rostral carina; one or
two similar beads likewise mark the apices of the acute-angled an-
484 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
terior margins of the protogastric lobes. Areola wide, appearing
very squat.
An orbital spine may be characteristic of this species; the evidence
is not conclusive; the spine is often represented by a small spinule
or acute corneous scale scarcely to be recognized as an orbital “spine”;
about a third of the specimens examined, mostly small, had no spinule
on either side, one-third had a definite spinule present on one or the
other side, while the remaining third had a spinule or correspond-
ingly sharp-pointed scale at the outer end of each orbit; whether
Ficure 54.—Aegla papudo, new species, male holotype: a, Dorsal view; b, lateral view of
anterior portion; c, sternum of third and fourth thoracic somites; d, inner ventral margin
of ischium of left cheliped; e, lateral view of second abdominal epimeron. a, b, natural
size; c—e, twice natural size. The rostrum is more lingulate than is apparent in a (cf.
pl 27"C).
armed with a spinule, scale, or granule or not, there is nearly always
a slight, sometimes abrupt but narrow, often lightly notched or in-
cised offset, usually no wider than the thickened border of the orbit,
between the outer end of the orbital margin and the inner slope or
margin of the anterolateral spine. Each of the three specimens be-
longing to the Philadelphia Academy, referred to in the remarks
appended to the “Distribution” of this species below, shows a definite
though small orbital spine on each side, separated from the corre-
sponding anterolateral spine by a narrow notch.
Anterolateral angle of carapace forming a sharply acute, fairly
slender conical spine, which reaches to and a little past the posterior
border of the cornea, in some cases about to the middle of the cornea.
Anterolateral angle of first hepatic lobe thick, lumpy, and blunted,
with a few corneous, scalelike projections, scarcely to be called
THE SPECIES OF AEGLA—SCHMITT 485
spinules; second and third hepatic lobes evident, but poorly marked.
Large hand stubby, palm thick and heavy looking, much swollen,
almost subglobular in appearance in some specimens, scabrous. Mov-
able finger with a low swelling or rather a small, more or less obso-
lescent spinulose lobe on outer margin near base. Palmar crest low,
outer margin thick and blunt-tubercular; the almost tuberclelike ser-
rations are furnished with short, more or less transverse rows of small,
pointed, corneous scales, few in number. Carpus rough-scabrous,
the only longitudinal ridge being the one above the inner marginal
row of spines; this ridge appears doubled, as it carries two longi-
tudinal series of more or less transverse rows of small, pointed, almost
spinulelike corneous scales. Anterior internal lobe or angle of carpus,
though at times subacute, more usually blunt, generally furnished
with several scattered, more or less subequal, almost spinuliform,
corneous scales; occasionally the apical one is a little larger than
the others. Merus armed above with a longitudinal series of blunt
tubercles topped with one, two, three, or more small, pointed, corne-
ous scales; anterior margin fine denticulate, without lobe or swelling.
Inner margin of ventral surface of ischium armed with three to four
more or less subequal, more or less equispaced, low, but definite and
well formed, conical tubercles or spines with subacute to acute corne-
ous tips, one anterior, one posterior, and one or two in the interspace
between the first two.
Dorsal anterior angle of epimeron of second abdominal somite
normally and usually rounded off and unarmed; very rarely does
one find a corneous scale or denticle or two, or even a small spinule
here and there usually on the epimeron of one side only. The speci-
men selected as the type is, in this respect only, perhaps one of the
most atypical specimens in the entire type lot. It is the largest speci-
men and has two little scales or tiny denticles on the right epimeron
and one tiny “cornule” on the left; the next largest specimen has
nothing of the sort on either dorsal epimeral angle; otherwise, only
four specimens out of the original lot of 20 have any trace of spinule,
denticle, or scale on the right or left epimeron. In about its middle
third the sternite between the bases of the chelipeds of the type and
one other specimen is somewhat swollen or raised up along the median
line, more so anteriorly, where it carries a perhaps adventitious, tiny,
corneous prickle or spinule, than posteriorly. In the next largest
specimen this swelling is much less marked. Also, it is unarmed, as
it is in the rest of the specimens at hand. Most of these have the
median elevation more or less obsolescent, yet have an appreciable,
though not very noticeable, convexity of the underside of the sternite ;
in a few of the smaller specimens it is not evident at all.
486 PROCEEDINGS OF THE NATIONAL MUSEUM Vou, 91
Holotype.—The largest male out of a lot of 14 males and 6 females
(1 ovig.), measuring 26 mm. in length of carapace and rostrum, col-
lected by J. A. Wolfsohn at Papudo, Chile, and received at the Field
Museum on February 3, 1925 (Field Museum No. 2285; paratypes,
2286).
Remarks.—This species, because of its very strongly reflexed,
anteriorly concave, or excavated rostrum, very convex carapace, and
much-swollen hands with low thick palmar crest, stands quite apart
from the other species of Aegla.
Although the dorsal anterior angle of the epimeron of the second
abdominal somite may rarely, and I believe only adventitiously,
carry a small, corneous scale or two, or even a tiny spinule, it does
seem that A. papudo is properly one of the group of species with a
rounded, unarmed dorsal anterior epimeral angle which includes
A. odebrechtii, A. o. paulensis, A. neuquensis, and A. affinis. In
certain other respects A. papudo seems to stand not far from A.
concepcionensis.
The several suture lines that meet to form the anterolateral angles
of the cardiac area of the carapace combine to form a short, quite
longitudinally oriented bar (fig. 54). It holds for every specimen of
A. papudo. Otherwise, I have noticed this state of affairs only in
the unique holotype of A. affinis (p. 496, fig. 58, a). In all other species
this short “bar” is, in contrast to A. papudo and A. affinis, oriented
so as to be very nearly transverse, or at least obliquely transverse.
Distribution.—So far known only from the 20 specimens (14 males,
6 females) of the type lot from Papudo, Chile; 3 males and 1 female
from Talcahuano, Chile (M. C. Z. No. 10480) and 1 male (about
24 mm. long) with only the indication Chile on the label, belonging
to the Museu Paulista, Sao Paulo, Brazil (M. P. No. 1306). I have
also seen a not altogether satisfactorily determinable female specimen
from the Rio Mapocho, near Talaganti, Province of Santiago, Chile,
collected by my good friend Dr. Carlos E. Porter, March 17, 1940, that
seems to be this species.
Recently I had the opportunity of examining the Aeglas belonging
to the Academy of Natural Sciences of Philadelphia. Included in
that collection were three dried female specimens between 30 and 31
mm. in length of carapace and rostrum together, labeled “Aeglea
laevis, Chili, Dr. Wilson” (Acad. Nat. Sci. Phila., no. 484, pt.). All
showed the more or less longitudinal suture lines of A. papudo (and
A. affinis). Their anterior dorsal epimeral angles are rounded off and
show no trace of either corneous scale or spinule. The rostral carina
seems a little more prominent for a greater extent of the rostrum than
is the case in most of the representatives of the species I have seen
so far, the carina having perhaps become accentuated as a result of
THE SPECIES OF AEGLA—SCHMITT 487
the drying out of the specimens. Orbital spines, separated from the
anterolateral spines by narrow notches or incisions, are definitely
present. The palmar crest is typical, low and appearing lumpy.
AEGLA ODEBRECHTII Miiller
FIGURE 55; PLATE 27, A
Aeglea Odebrechtii Fritz MUtirr, Jen. Zeitschr. Naturw., vol. 10 (new ser.,
vol. 3), p. 13, pl. 1, figs. 1-10, 1876.
Aeglea intermedia MoreErra, Arch. Mus. Nac. Rio de Janeiro, vol. 11, pp. 21, 84,
1901.
Description.—A species of moderate size, attaining at least 28 mm.
in length of carapace and rostrum taken together.
é a
Ficure 55.—Aegla odebrechtii Miller, male neotype: a, Dorsal view; }, lateral view of anterior
portion; ¢c, sternum of third and fourth thoracic somites; d, inner ventral margin of ischium
of left cheliped; ¢, lateral view of second abdominal epimeron. a, b, natural size; c-e,
twice natural size.
Carapace with gastric region quite convex. Rostrum relatively
short, not exceeding eyes by more than the length of the cornea,
fairly flat, broadly triangular, and appreciably widely grooved or
excavate either side of the well-marked median carina; distally the
carina tends to fade out before reaching the anterior extremity of
the rostrum; on the carina are two rows of small, more or less alter-
nating, corneous scales fairly close together, so much so that close to
the distal end of the carina the two rows merge to form one irregular
row.
Protogastric lobes not well marked, because of the very appreciable
convexity of the gastric region; epigastric prominences blunt swell-
ings.
488 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
Orbital sinus moderately wide but shallow, orbital spine a small,
acute, corneous projection set close to the anterolateral spine and sep-
arated from it by a much-reduced extraorbital sinus, a small V-shaped
incision or notch. Anterolateral spine relatively short, flattened tri-
angular in the largest specimen, more slender and elongate appearing
in the smaller ones, may reach a little past the level of the posterior
margin of the cornea.
First hepatic lobe, though separated from the anterolateral one by
a conspicuous notch, has its anterolateral angle bluntly rounded off
and its lateral margin small scabrous, as are the margins of the
second and third hepatic lobes, which are only poorly indicated; in
smaller specimens the first hepatic lobe is subacute and tipped with
a corneous scale larger than those elsewhere on the lateral margin.
Hands broadly ovate, more or less flattened, yet gently convex.
Movable finger with a definite lobe on the outer margin near the base;
lobe tipped or furnished with one or more acutely conical corneous
scales (almost very small, short, conical, corneous spines). Palmar
crest the most distinctive feature of this species, large, subdisciform,
and noticeably excavate, much as if it had been impressed or pinched
out while soft with the ball of one’s thumb; margin of crest notice-
ably upturned, more or less obscurely serrate, scabrous to small-
spinulose.
Ridge of carpus of cheliped above spined inner margin well devel-
oped, raised above general level of carpus, and marked with nodular
swellings carrying transverse rows of corneous scales; anterior inter-
nal lobe or angle of carpus low, conical, and furnished with small
corneous scales apically and on its slopes. If one regards the largest
spine of those arming the inner margin of the carpus as the most
anterior of that particular series, we find then in this species on the
inner anterior slope of the base of that first spine a smaller, yet
conspicuous, strong spine located in more or less of a triangular
area delimited by that first spine, the carpal ridge, and the anterior
internal lobe of the carpus. This “inserted” spine may sometimes be
closer to, but not normally fused with, the large first spine of the
series arming the inner margin of the carpus than it is to either the
carpal lobe or the carpal ridge. This spine seems to be represented
in the closely related A. odebrechtii paulensis, immediately following,
by a similar one also placed on the anterior slope of the first spine
of the series arming the inner carpal margin; unlike the independent,
distinct spine of the species proper (s. s. ), it is always much fused with
the first spine (of the inner marginal series), so that usually only its
tip is distinguishable; sometimes it is wholly fused with the first
spine, which, in either case, is a very much thickened spine. In A.
odebrechtii, between the “inserted” spine as it may be designated and
THE SPECIES OF AEGLA—SCHMITT 489
the carpal ridge there may be, also in the larger specimens of the
species, an acute little tubercle armed apically with two or three
sharp, dark-colored corneous scales. Inner margin of ventral sur-
face of ischium armed with four more or less subequal, at times more
or less equispaced, low, but definite and well-formed, conical tubercles
or spines with subacute to acute corneous tips, the anteriormost the
largest, the most posterior second in size, the anterior of the two in
between the first two named, third, and the posterior fourth in size
(this describes the margin of the left ischium of the neotype; the
right is armed like the left except that the two spines in the inter-
space between the anterior and posterior spines are just about equal
in size and placed quite close together in the middle of the inter-
space) ; in the specimen next in size (25 mm.) the anterior spine is
quite appreciably larger than any of the others on this margin of
the ischium.
Anterior dorsal angle of epimeron of second (in lateral view,
apparent first) abdominal somite broadly rounded off, not spined.
Neotype.—The largest male I have seen (U.S.N.M. No. 80022),
28 mm. in length of carapace and rostrum, was collected by Dr.
Carlos Moreira in 1904 in Santa Catharina, Brazil, and later gen-
erously presented by him to the United States National Museum.
Remarks.—More intuitively than he realized, Fritz Miller (1876)
exclaimed, when his first specimen of Aegla odebrechtii came to
hand, “How is it that we find this Pacific crustacean [from the west-
ern slopes of the Andes] in our mountains [here on the Atlantic coast
of Brazil]?” So far as he knew at that time, no representative of the
genus had been discovered outside of Chile, and, in spite of the wide
distribution of the Aeglas here described, his species is the one east
South American form that seems most to resemble those inhabiting
the slopes of the Andes.
Distribution —Aside from the neotype, I have seen just 8 other
specimens, 6 males, of which the largest measured 25 mm. in length
of carapace and rostrum, the next in size 14, and the smallest
1314 mm., and 2 females of 15 and 14 mm., respectively. These speci-
mens were kindly obtained for me by Dr. Carlos Moreira through the
kind offices of his good friend Dr. G. Kuhlmann, Blumenau, Santa
Catharina, Brazil. I am very grateful to both of these estimable
gentlemen for their interest and help in this matter.
An additional, quite typical male belonging to the Academy of
Natural Sciences of Philadelphia (no. 484, pt.), 26 mm. long, cara-
pace including rostrum, and labeled “du Brésil. Donni par M. M.
Derreaux,” has lately come to my attention. It has the characteristic
“inserted” spine easily observable in the neotype (fig. 55, a, and pl.
27, A) ; the ventral inner margin of the ischium of the right cheliped
is likewise armed as in this figured specimen.
490 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 92
AEGLA ODEBRECHTII PAULENSIS, new subspecies
Figure 56; PLATE 27, B
Aeglea intermedia LUEDERWALDT, Rev. Mus. Paulista, vol. 11, p. 431 (sep., p. 5),
1919.
Description—Perhaps only a small species; my material of this
form is limited; the largest specimen at hand, a male, in length of
carapace and rostrum together measures 20 mm.; the male holotype
is just 1 mm. shorter.
Carapace moderately convex, front of moderate width. Rostrum
broad and somewhat stubbily triangular; bluntly carinated nearly
to the anterior extremity, noticeably troughed or excavate either side
of carina, which broadens out and becomes more or less lost in the
general surface of the carapace at a level about halfway between the
level of the epigastric prominences and the anterior borders of the
protogastric lobes, these last take the form of a low, somewhat arc-
uate, blunt elevation or obsolescent ridge; the epigastric prominences
are fairly well developed and nodular or near rounded-tubercular.
Orbital sinus moderately wide, only moderately deep, fairly deep
as compared to A. odebrechtii; orbital spine small; extraorbital
sinus is quite shallow and, though small, is relatively moderately
wide as compared with A. odebrechtii.
Anterolateral spines small, stubby, and only moderately advanced
beyond the orbital spines (in some apparently more or less worn
individuals the orbital spines are nearly on a level with antero-
lateral ones). First hepatic lobe set off from anterolateral lobe by
a fairly wide, relatively good-sized notch; anterolateral angle of the
first hepatic lobes a little produced and subacute, carrying a small
corneous granule or denticle, lateral margin of lobe scabrous; second
and third hepatic lobes, though not much more so, are a little better
marked than in A. odebrechtii.
Large hand relatively of good size, broadly oval, stockily built,
with palm rather thick and swollen toward outer margin. Movable
finger with a small, definite, though not particularly conspicuous,
scabrous lobe on outer margin near base.
Inner margin of palm with a well-developed, impressed or exca-
vate crest, having its outer margin somewhat parallel to the dorsal
margin of the palm proper, not nearly so subdisciform as in A.
odebrechtii; margin of this palmar crest more or less definitely serrate,
serrations marginally scabrous or fine denticulate or corneous gran-
uled, perhaps even small spinulate at or on apices of serrations.
Ridge of carpus of cheliped above spined inner margin more or
less well developed, scabrous-nodular; large anterior spine of series
arming inner margin of carpus may be as large and thick as if it
THE SPECIES OF AEGLA—SCHMITT 491
were formed by the merging of two spines of normal size to form one;
usually most traces of the double nature of this large anterior spine
are lost except as evidenced by its noticeable breadth as in the case
of the spine on the right carpus of the type, which is only most
obscurely 2-pointed; nevertheless, there are instances, as on the left
carpus of the type, that reveal very clearly the double nature of this
thickened first spine with a distinctly twinned or 2-spined extremity;
in the interval between the base of this thickened first spine, the
base of the carpal lobe, and the anterior portion of the carpal ridge,
<—S
Ficure 56.—Aegla odebrechtit paulensis, new subspecies, male holotype: a, Dorsal view;
b, lateral view of anterior portion; c, sternum of third and fourth thoracic somites; d,
inner ventral margin of ischium of left cheliped; e, lateral view of second abdominal
epimeron. a, b, natural size; c—e, twice natural size.
there may be two or three very small, slightly tubercular scabrosities ;
the carpal lobe itself is scabrous, bluntly rounded to subacute. Upper
longitudinal margin of merus armed with at least two strong spines
of good size followed by several smaller ones; in advance of the
anteriormost, the largest spine, on the actual anterior margin of the
merus is a low, lobular, subrectangular ridge, longitudinally oriented.
Armature of the inner margin of the ventral surface of the ischium
very like that of A. odebrechtii, a fair-sized, stout, conical spine at
anterior end with usually two subequal, somewhat smaller ones close
together at posterior end, often a fourth still smaller spine in the
interval between the posterior pair and the anterior spine; only
exceptionally is there only an anterior and one posterior spine or
only one intervening one (as in fig. 56, @).
Anterior dorsal angle of epimeron of second (in lateral view,
apparent first) abdominal somite in general more or less rounded off,
as in A. odebrechtii.
Holotype.—The next to largest male, U.S.N.M. No. 80023, of a lot
of 4 males and 3 females collected by Dr. Doris M. Cochran at Alto
492 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
da Serra do Cubatio, between Santos and Sao Paulo, Brazil, April
26, 1935.
Remarks.—Although this subspecies is decidedly similar to A.
odebrechtit Fritz Miiller, I do not have at hand enough well-developed
specimens to prove either their specific distinctness or identity.
Therefore the specimens I do have have been given subspecific
ranking.
In relation to the eye, the rostrum of the species proper appears
a little longer; also it seems to be relatively a little more recurved
distally ; the rostrum is more nearly straight in the subspecies. The
orbits of the subspecies are definitely wider than in the species proper
and represent perhaps the most noticeable difference between the two
forms. Though not affording a very clear-cut difference, the an-
terolateral spines seem a little longer in the species proper, appearing
to reach a little past the posterior margin of the cornea, while in the
subspecies the anterolateral spine scarcely reaches the cornea. The
anterior margins of the protogastric lobes are definitely elevated in
the subspecies and the epigastric prominences, though low, are con-
spicuously tuberculiform; the reverse is true in the species proper
on both counts.
Next to the orbits, the chelae of the two forms seem to be most
definitely different. In the subspecies they are relatively heavier,
stouter (chunkier, more swollen, or inflated), with appreciably
shorter, broader (stubbier) fixed fingers; the outer margin of the
palm of either hand has a comparatively greater convexity; while
the palmar crest is generally more (more or less) subparallel-sided
trough-shaped than impressed or excavate-subdisciform, and certainly
more definitely serrate in nearly every specimen of the subspecies
than in the species proper.
Ordinarily, the female Aeglas do not exhibit the pronounced
asymmetry found in the male major and minor chelae, but in this
subspecies at least there is such asymmetry that at first glance the
two females with both chelae present (of the three females seen)
were taken to be males.
It is possible that I have set up one form too many in naming
this subspecies.
Distribution.—Other than the specimens from the type locality, I
have seen only a few small individuals, of which the largest was
about 15.5 mm. in length of carapace and rostrum together, which
may represent this subspecies, but I do not feel that I can make more
than tentative determinations of small specimens of forms as closely
related as the two here designated as A. odebrechtii and A. 0. paul-
ensis. One lot of four small specimens received from Dr. Hermann
von Ihering a number of years ago is from the “Rio Juquery, Perus,
THE SPECIES OF AEGLA—SCHMITT 493
Estado Sao Paulo”; another small female, also from Dr. von Ihering,
is labeled simply Alto da Serra, Sao Paulo (Coll. J. Lima, 1908). A
third lot of seven small specimens collected by E. Garbe, from Castro,
Est. Parana, is even more of a puzzle than either of the preceding
lots; the rostra do not seem to be quite typical of paulensis, yet the
specimens cannot be identified with the species A. castro, which
I found so common in the Rio Iapé at Castro, for their unarmed
dorsal epimeral angle precludes the possibility; even much smaller
Castro specimens of my own collecting have this angle unmistakably
spined.
é
Ficure 57.—Aegla neuquensis, new species, male holotype: a, Dorsal view; b, lateral view
of anterior portion; c, sternum of third and fourth thoracic somites; d, inner ventral
margin of ischium of left cheliped; ¢, lateral view of second abdominal epimeron. a, b,
natural size; ¢-e, twice natural size.
AEGLA NEUQUENSIS, new species
FicurRE 57; PLATE 27, E
Description—aA species of moderately to fairly large size, exceed-
ing a length of carapace and rostrum together of at least 30 mm.
(based on the largest specimen seen, a “soft” male with regenerated
but not yet fully developed rostrum).
Carapace moderately convex, front moderate; rostrum flattened
triangular and deeply grooved or excavate either side of median
carina, which tends to fade out toward tip of rostrum which is ap-
preciably reflexed or upturned; rostrum exceeds the eyestalks from
about 114 (in the type) to about 2 times the length of the cornea;
the rostral carina is furnished with a more or less double row (on
494. PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 91
occasion in places apparently three rows) of closely set corneous
scales for at least half the length of the free portion of the rostrum,
beyond the midpoint there is but a single row of the scales, which,
like the carina, tends to fade out or disappear before reaching the
distal extremity of the rostrum (sometimes there is an odd grouping
of a few scales on the dorsum of the extreme tip of the rostrum) ;
the rostral carina is plainly marked backward to the level of the
anterior margins of the protogastric lobes, and in at least the larger
of the specimens at hand, faintly to be seen if only as an interruption
to reflected light halfway back to the cervical groove.
Protogastric lobes poorly indicated; epigastric prominences not
very prominent, obliquely elongated, scabrous swellings.
Orbital sinus moderately wide, in dorsal view appearing not much
wider than deep; orbital spine always present, small but well formed;
extraorbital sinus narrow, a V-shaped notch. Anterolateral spines
relatively small, yet reaching past posterior border of cornea often
about or nearly to middle of its length. Anterolateral angle of first
hepatic lobe produced but not spined, though scabrous or small
spinulated as on lateral margin of lobe; second and third lobes no
more than plainly indicated by shallow emarginations in lateral
margin of anterior portion of carapace.
Asymmetry of hands not very pronounced; large hand of moderate
size, more or less subovoid, only moderately inflated; the hands are
coarsely scabrous, almost tuberculated. Movable finger in the type
does not seem to have a real lobe developed on outer margin near
base, yet there are a few larger spinules on a very slight elevation at
the site of the lobe found in other species; however, in other speci-
mens smaller than the type a slight lobe armed with several sharp
spinules seems definitely present. Palmar crest more or less nar-
rowly subrectangular, fairly thin-edged, serrate or notched, and
spinulose; dorsal surface of crest at most only very slightly concave,
margin of crest not noticeably or appreciably, if at all, bent upward.
Ridge of carpus above spined inner margin carrying practically
a double row of scabrous elevations; between anterior spine, the
largest of the series arming the inner margin of the carpus, and the
carpal ridge there is a short, acute, conical spine nearly subequal
in elevation with the scabrosities of the carpal ridge (this spine seems
to be present in the specimens from the type locality, Arroyo, but
not at all, or only almost imperceptibly indicated in the specimens
from Covunco) ; anterior internal lobe or angle of carpus flattened-
conical, or triangular, armed with one larger, sharp-pointed corneous
denticle, with a smaller one close behind on the posterior slope, and
usually one or more still smaller spiniform scales. Upper longi-
tudinal margin of merus of cheliped with a series of small, more or
THE SPECIES OF AEGLA—SCHMITT 495
less subequal scabrous tubercles, except the first which is quite the
largest; anterior margin of joint medially produced, forming a den-
ticulated lobe; these denticulations are usually carried outward along
the anterior margin of the merus, scarcely ever and perhaps only
adventitiously along anterior margin inside the lobe itself. Inner
margin of ventral surface of ischium armed with from four to six
conical corneous-tipped tubercles or spines, of which the most anterior
and, posterior are more or less subequal and the largest; often the
first spine is twinned (the twin being smaller and on the posterior
slope of the anterior spine proper and included in the four to six
count) ; more rarely is the posterior, or one of the intermediate and
always smaller spines twinned (as in left ischium of type, fig. 57, d).
Anterior dorsal angle of epimeron of second (in lateral view, ap-
parent first) abdominal somite evenly rounded off, anterior margin
below angle straight.
Sternite between bases of chelipeds with anterolateral angles pro-
duced, tuberculiform; on median line near anterior margin of this
sternite there is a low conical elevation topped by a small, usually
acute corneous spinule.
Holotype.—The second largest male, U.S.N.M. No. 80024, of a lot
of 4 males and 1 female, measures 29 mm. in length of carapace and
rostrum; the female measures 20.5 mm.; the smallest male, 17.5.
All were collected at Arroyo, Territory of Neuquen, Argentina, by
John W. Titcomb, November 12, 1903, while conducting a fisheries
survey in that vicinity for the Argentine Government.
Remarks.—This species is certainly closely related to the following,
yet differs from it in several important points. The separate descrip-
tion of the latter seems fully warranted.
Distribution.—In addition to the type lot, I have examined a second
lot of material, 5 males and 1 female, ranging from 18 to 24 mm.
in length of carapace and rostrum together. These specimens were
collected the same day as the type lot, November 12, 1903, by Mr.
Titcomb at Covunco [?] or in the [Rio] Covunco; the original label
is somewhat rubbed and partly illegible, but the date and “Neuquen”
[Territory ?] are unmistakable.
AEGLA AFFINIS, new species
Ficurp 58; PLATE 27, F
Description—I have seen but one specimen of this species, the
unique holotype, a male of fairly large size, measuring in length of
carapace and rostrum 31mm. Most of its legs are broken, and the
chelipeds are detached; in addition there is another loose cheliped of
a specimen of probably the same size.
435661—42—__5
496 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
In general appearance it is much like A. neuquensis; carapace and
rostrum very similar, but front seemingly narrower, extraorbital
sinuses and orbital spines wanting. Rostrum exceeds eyes by not
quite twice the length of cornea; the blunt carina more or less con-
tinued to distal extremity, more nearly approaching the ridge-roofed
condition of rostrum than any of the Aeglas of the A? division of
the diagnostic key; the rostrum otherwise appears quite flat, particu-
larly basally, and fairly well troughed or excavate either side of
median carina; the latter is bluntly rounded off and scabrous, with
rather numerous, closely set, partly imbricate-appearing corneous
Ficure 58.—Aegla affinis, new species, male holotype: a, Dorsal view; b, lateral view of
anterior portion; ¢c, sternum of third and fourth thoracic somites; d, inner ventral margin
of ischium of left cheliped; ¢, lateral view of second abdominal epimeron. a, b, natural
size; c—e, twice natural size.
scales not at all arranged in rows as in A. newquensis; rostral carina
most imperceptibly if at all suggested posterior to obsolescent anterior
marginal indications of protogastric lobes.
Anterolateral spines flattened-triangular in dorsal view, reaching
on the left side nearly to middle of cornea, on right well past middle
of cornea; anterolateral angle of first hepatic lobe somewhat pro-
duced, subacute or rounded off, small spinulose or scabrous; second
and third lobes poorly, obsolescently indicated.
Hands more elongate-subrectangular than subovoid as in A. neu-
quensis, and more coarsely scabrous. Movable finger seems to be
without trace of lobe on outer margin near base, except on minor
chela, where there is a very small corneous spinule or denticle larger
THE SPECIES OF AEGLA—SCHMITT 497
than the scabrosities of the surface of the finger otherwise in the
position normally occupied by the lobe in other species. (Having
so little material, it is impossible to tell whether the lobe is in evi-
dence in small individuals of the species. There is no trace of it
on the movable finger of the loose cheliped.) Palmar crest more or
less subrectangular, thicker appearing than in A. neuquensis and
certainly with thicker, blunter, obscurely crenulate, coarsely scabrous
margin; dorsal surface of crest decidedly more concave (more or less
Jongitudinally troughed) than in A. newquensis, but without giving
the margin of the crest any noticeable bent-up appearance. Carpal
ridge fairly broad and blunt, more or less obscurely scabrous, and
only obscurely double-rowed as in A. neuguensis; spines of inner
margin of carpus thickened and scabrous, between anterior spine
(very much the largest and stoutest of this inner marginal series)
and the carpal ridge is a tuberculiform, scabrous elevation corre-
sponding to the similarly placed spine in typical A. newqguensis; lobe
at anterior inner angle of carpus quite rounded off in general outline,
margined with small, denticuliform, corneous scales. Upper longi-
tudinal margin of merus of cheliped armed with a series of small
scabrous tubercles; this row or series at its anterior end makes prac-
tically a right-angled bend one or two tubercles long, toward the
inside, more or less paralleling anterior margin proper of joint *’;
this is very evident in the meri of the type but not in the additional
loose claw (No. 4186) of this species. No indication, or scarcely any,
of this state of affairs exists in A. neuquensis, there may be a bare
suggestion of it in some specimens in which a tiny, well nigh micro-
scopic corneous scale or prickle may appear on the inner side of the
anterior spine or tubercle of the upper longitudinal margin of the
merus of the cheliped. The anterior margin of the merus of A.
affints, though scabrous or fine denticulate, shows no median lobular
development as is present in A. neuquensis.
Inner margin of ventral surface of ischium armed much as in A.
neuquensis, only cones are smaller, mostly blunter, and on the whole
more nearly subequal throughout, four on right ischium, six on left,
because of a twinning of the posterior spine, and also the one just
anterior to it.
Anterior dorsal angle of epimeron of second (in lateral view, ap-
parent first) abdominal somite rounded off; anterior margin below
angle straight, or very slightly concave.
Anterolateral angles of sternite between bases of chelipeds pro-
duced, tuberculiform; on median line near anterior margin a low
swelling, but with no trace of a corneous spine or denticle arming it.
“A similar condition is found in A. uruguayana, p. 467.
498 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
Holotype-——A single male carrying M.A.C.N. tag No. 9817, con-
tained in a bottle with an unattached left cheliped surely the same
species with an M.A.C.N. tag, No. 4186, affixed, together with
a specimen of each of two other species without tags. Of these last,
one is a female of A. hwmahuaca, 22.0 mm. in length of carapace
and rostrum together, the other a male of A. abtao, of 28.0 mm.
In the catalogs of the Museo Argentino Ciencias Naturales entry No.
4186 reads simply, “Neuquen, Mayo 16, 1898; Sr. Carlos Burmeister” ;
entry No. 9817 concerns specimens of Mytilus chorus Molina received
in exchange from Dr. Carlos S. Reed, 21-V, 1919. As a result, it is
impossible to determine satisfactorily the type locality for the
‘species, and there is no locality at all for the other, untagged, speci-
mens in the same bottle. It is a mixed lot of material, or else a case
of misattached label or labels.
Remarks.—As pointed out under A. papudo above, this is the only
other species in which the several suture lines that meet to form
the anterc:ateral angles of the cardiac area of the carapace combine
to form a short, quite longitudinally oriented bar (fig. 58). In all
other species except these two this short “bar” is oriented so as to
be very nearly transverse, or at least obliquely so.
AEGLA HUMAHUACA, new species
FIGURE 59; PLATE 27, D
Description—A species of moderate size. The largest of five
specimens seen measures about 28 mm. in length of carapace and
rostrum taken together.
Carapace moderately convex, front relatively narrow. Rostrum
rather thick looking, proximally more or less broadly flattened-
triangular, noticeably depressed anteriorly, bent downward, so much
so that in lateral view the rostral extremity is about on or even
slightly below the level of the anterolateral spines; distally the
rostrum becomes somewhat lingulate, slightly parallel sided, low,
and broadly blunt-ridged, scarcely to be called carinated; only very
shallowly excavate either side of median carina; carina marked in
basal half with three or four very irregularly intermingled rows of
corneous scales, becoming distally more or less a single scattered
row, which near tip of rostrum tends to disappear, scarcely or not
distinguishable from the few scattered corneous scales on the dorsum
of the apical portion of the rostrum. Epigastric prominences and
anterior margins of protogastric lobes poorly developed.
Orbital sinus fairly narrow, more or less V-shaped; orbital spine
small, placed well up on inner slope or margin of anterolateral
spine and separated from it by only a small notch. Anterolateral
spine relatively small, short, and flattened-conical. Anterolateral
THE SPECIES OF AEGLA—SCHMITT 499
angle of first hepatic lobe low, scabrous-tubercular; second and third
lobes very poorly marked.
Hands large, oval, moderately inflated, and without usual lobular
tooth on prehensile margin of immovable finger; movable finger like-
wise without such a tooth; there is no lobe on the outer margin of
the movable finger, and the palmar crest, though not prominent, is
distinctly present, thick, low, and in cross section broadly triangular;
dorsal surface of crest faintly, shallowly, or more or less obscurely
excavate; the crest is scabrous with an outline that is more slightly
irregular than obscurely serrate; serrations may be spinule tipped.
é a
Ficure 59.—Aegla humahuaca, new species, male holotype: a, Dorsal view; b, lateral view
of anterior portion; c, sternum of third and fourth thoracic somites; d, inner ventral
margin of ischium of left cheliped; e¢, lateral view of second abdominal epimeron. a, 4,
natural size; c—e, twice natural size.
Ridge of carpus of cheliped above inner spined margin not very
prominent, low and broad and in small part only slightly scabrous;
the armature of the inner margin of the carpus is not so definitely
spinelike as in most other Aeglas; here it consists more of spinelike
tubercles, perhaps only the most anterior of the series may be so
designated, as the next three or four are more or less tuberclelike in
appearance; the posterior one or two of these are indeed very low,
blunt, and scabrous; anterior internal lobe or angle of carpus scarcely
more than obtuse angled; this angle is armed with one or more small,
low-conical but more or less sharp-pointed corneous scales; upper
longitudinal margin of merus blunt angled, hardly more than a
scabrous ridge marked or armed with a row of fairly well separated,
short, subacute, corneous scales; the anterior margin of the merus
is finely denticulate, but no lobe or forwardly directed projection
500 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
is developed there. Inner margin of ventral surface of ischium
with an anteriorly corneous spine- or pointed-scale-tipped tubercle
at anterior end, and a lower, likewise corneous spine-tipped tubercle
at posterior end; two slight, at times almost imperceptible undula-
tions, or slight low swellings, may occupy the interspace.
Anterior dorsal angle of epimeron of second (in lateral view, ap-
parent first) abdominal somite rounded off, yet armed on its anterior
margin, to the right, with two corneous spinules or denticles set quite
close together, to the left with one.
Holotype.—The largest of four males from Humahuaca, Jujuy,
Argentina (M. A. C. N. No. 8837) measuring about 28 mm. in length
of carapace and rostrum together; the other three males of the type
lot measure respectively 25.0, 24.5, and 17.5 mm.
Remarks.—This species and A. jujuyana so resemble each other in
general appearance that one cannot escape the conviction that they
may be very closely related in spite of the fact that A. hwmahuaca
possesses a palmar crest and has a very bluntly ridged rostrum, char-
acters definitely differentiating the two. Geographically in the
Province of Jujuy these species are found scarcely more than 70
miles apart, but environmentally, or at least climatologically, they
are far removed one from the other. At Humahuaca the annual
rainfall totals only 6.11 inches'*; five months, May to September,
are without any precipitation whatsoever, while January, the wettest
month, has a rainfall of but 3.27 inches. At Juiuy, on the other
hand, the total is 29.26 inches; no month is wholly without some pre-
cipitation, although this may fall as low as 0.12 inches in August;
the wettest month, January, marks a high of 6.65 inches, more in one
month than Humahuaca receives in a year.
Distribution —Other than the holotype and three paratypes from
Humahuaca, Province of Jujuy, Argentina, I have seen but one other
specimen, a female of 22.0 mm. in length of carapace and rostrum
taken together. This particular specimen was found in a bottle con-
taining two other specimens specifically different, together with a
detached cheliped. One of these specimens was selected as the type
of A. affinis (M. A. C. N. tag No. 9817), the loose cheliped (M. A. C.N,
tag No. 4186) represents the same species; the remaining specimen
proved to be a male Aegla abtao (28.0 mm. in length of carapace and
rostrum). This lot of material certainly contains a mixture or else
one or both of the labels may be misattached. In the catalogs of the
Museo Argentino Ciencias Naturales entry No. 4186 reads simply
14 The figures on precipitation given in this paragraph were taken from W. W. Reed’s
undated, bound, typewritten manuscript, “Distribution of Precipitation over the Earth,”
lent me by the Library of the United States Weather Bureau, through the kindness of
Miss Rose Vickers, librarian.
THE SPECIES OF AEGLA—SCHMITT 501
“Neuquen, Mayo 16, 1898; Sr. Carlos Burmeister”; entry No. 9817
concerns specimens of Mytilus chorus Molina from Chile received in
exchange from Dr. Carlos S. Reed, “21-V, 1919.” There is no telling
whence comes this unlabeled specimen of A. humahuaca.
AEGLA CONCEPCIONENSIS Schmitt
FIGuRE 60; PLATE 28, A
Aeglea concepcionensis ScuMItTT, Rev. Chilena Hist. Nat., vol. 44 (1940), p. 26,
pl. 5, fig. 1, 1942.
Description.—A fairly large species attaining a length of carapace
and rostrum together of at least 33 mm.
é a
Ficure 60.—Aegla concepcionensis, new species, male holotype: a, Dorsal view; b, lateral
view of anterior portion; c, sternum of third and fourth thoracic somites; d, inner ventral
margin of ischium of left cheliped; ¢, lateral view of second abdominal epimeron. a, b,
natural size; c—e, twice natural size.
Carapace moderately convex. Rostrum somewhat elongate-triangu-
lar tongue-shaped, exceeding the eyestalks by not quite twice the
length of the cornea, inclined downward, but anteriorly recurved,
transversely flattened, excavate either side of median carina. Crest
of rostral carina furnished with two rows of tiny corneous scales
situated fairly close together behind the level of the posterior margin
of the orbit and very closely juxtaposed, or at times even inter-
mingled or imbricated anterior to that level, and in the anterior half
502 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
of the free portion of the rostrum apparently becoming a some-
what broken or irregular single line of scales; the more prominently
raised portion of the carina extends backward about to the level
of the epigastric prominences of the carapace, posterior to these the
carina is less prominently marked to between the anterior margins of
the protogastric lobes where the carina fades out. Epigastric prom-
inences blunt-nodular, anterior margins of protogastric lobes scarcely
or poorly marked, obsolescent and not scaled. Areola short and wide,
squat looking.
Orbits of good size, fairly deep, typically without an orbital spine,
and usually with scarcely any or only (rarely) a very slight inter-
ruption or offset in the outward sweep of the orbital margin at the
point where it passes over into the inner margin or slope of the antero-
lateral spine of the carapace; in the very largest specimens, such as
the type, there is more of an offset than in any other specimens of the
species that I have seen; there may be one or a few tiny spinules along
the outermost portion of the orbital margin, but in no sense is any
of them of sufficient consequence to be considered as representing
an orbital spine.
Anterolateral spine of good size, anterior extremity reaching nearly
or about to the level of the middle of the cornea; the dorsal surface
of the anterolateral lobes is much flattened, almost or slightly exca-
vate, giving the impression that the anterolateral spines are inclined
upward to a greater extent than in any other species of Aegla. An-
terolateral angle of first hepatic lobe slightly scabrous and more or
less rounded off; just within and below the angle of the right first
hepatic lobe of the type is a low projection or tubercle, which is
occasionally present in other specimens on one or the other side or
sometimes on both sides; second and third hepatic lobes slightly indi-
cated, in some specimens scarcely so.
The larger hand is of good size, moderately inflated or swollen;
on the upper surface of either palm there is a faint, obsolescent, yet
plainly discernible, low, obliquely longitudinal, narrow swelling run-
ning from near the outer posterolateral angle of the palm to the pos-
terior margin of the sinus between the fingers; this ridge is scabrous
like the rest of the hand, and is more evident in the smaller specimens
than in the very largest ones. On the outer margin of the movable
finger of either hand, near its posterior end, there is a well-defined
lobe or projection, anteriorly angled and carrying there a small spine
or spinule; lobe otherwise scabrous, or very small-spinulose. What
there is of a palmar crest (on inner margin of palm) is broadly
and shallowly serrate, fairly thin-edged and furnished with a scatter-
ing of small spinules; the crest runs back from below the movable
THE SPECIES OF AEGLA—SCHMITT 503
finger to form a higher crest at the posterior end than at the anterior
end; posteriorly the crest is somewhat troughed or excavate with
slightly upturned margin which stands well away, almost at a right
angle, from the inner margin of the palm proper just in advance
of the articulation with the carpus.
Carpus of either cheliped carries two longitudinal ridges, the first
is the usual somewhat nodulated ridge with more or less transverse
short rows of small corneous scales, situated above the spines arming
the inner margin of the carpus; the second, scarcely to be called a
ridge, is on the middorsal surface of the carpus. It consists of an
irregular, scattered row of slight elevations anteriorly scabrous.
Anterointernal angle of carpus of cheliped fairly blunt, scarcely sub-
acute, sparsely small-spinulated. Dorsal longitudinal margin of
merus armed with a row of corneous tipped or blunted, somewhat
conical tubercles which become more conically spinelike as they ap-
proach the distal margin of the joint; the anterior margin of the
merus at its middorsal point shows but a very faint indication of what
might have been an obsolecent swelling with one (on right merus) or
two (on left) small corneous denticles; in smaller specimens there is
more of an evident lobe or small nodular swelling at this point with
finely denticulate anterior margin; outward from this lobe the
anterior margin of the merus is in part more or less denticulated.
Inner margin of ventral surface of ischium with four, five, or six
low swellings or nodulations, of which the anteriormost is usually the
largest, and in occasional specimens somewhat blunt tuberculiform ;
in some others this ischial margin appears no more than a little wavy
behind the anterior nodule or tubercle; only rarely does this seem
to be tipped with a tiny corneous scale.
Anterior dorsal angle of epimeron of second (in lateral view, ap-
parent first) abdominal somite produced to form an acute corneous
spinule-tipped angle.
Holotype.—A large male measuring 33 mm. in length to carapace
and rostrum, U.S.N.M. No. 79078.
In all, I have examined about 30 specimens of this species. Sev-
eral are of good size; the majority, however, are of medium or small
size. All of them I collected January 13 and 14, 1927, near Con-
cepcion, Chile, in company with Dr. Carlos Oliver Schneider and
Carl Junge.
Remarks.—A. concepcionensis keys out near A. laevis; in the “Re-
marks” under the latter (p. 507) the two are compared.
In its lack of an orbital spine, A. concepcionensis stands near
A. papudo, in which such a spine is often not properly or truly de-
veloped, and A. affinis, in which it is lacking (in the unique holo-
504 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 91
type). Of these three species, only A. concepcionensis has the an-
terior epimeral angle definitely acutely produced and spined; in
A. affnis it is rounded off and unarmed; in A. papudo likewise
rounded off and generally unarmed, though the angle may carry a
tiny adventitious corneous scale, spinule, or “cornule.” The hands
of A. papudo and A. concepcionensis are more or less ovoid and swol-
len or inflated, more so in the former than in the latter, while in
A. affinis they are more or less subrectangular, and less swollen, though
rougher, more scabrous, than in either of the others. A. papudo has
the most convex carapace, A. affinis the least, the convexity of the
carapace of A. concepcionensis being intermediate. Further, the sev-
eral suture lines which meet to form the anterolateral angles of the
cardiac area of the carapace run together to form a short, transverse
or obliquely transverse bar in A. concepcionensis, and a more or less
longitudinally oriented bar in A. papudo and A. affinis.
Distribution.—In addition to the type material, I have seen three,
not altogether typical males, between 15.5 and 24.5 mm. in length of
carapace and rostrum together, from Corral, Chile, collected by Dr.
Thomas Barbour (M.C.Z. No. 10481), and two males of 25.5 and
26.5 mm. respectively, collected by Dr. A. Santa-Cruz in the vicinity
of Concepcion, Chile, and presented to the United States National
Museum by our good friend Dr. Carlos E. Porter, of Santiago.
AEGLA LAEVIS (Latreille)
FIcuRE 61; PLATE 28, D
Galathea laevis LATREILLE, Tableau encyclopédique et méthodique . . ., pt. 24,
pl. 308, fig. 2, 1818.
Aegla laevis Leacu, Dictionnaire des sciences naturelles, vol. 18, p. 49, 1821.
Aeglea laevis DEsMAREST,” Considérations générales sur la classe des Crustacés,
p. 178, pl. 38, fig. 2, 1825.
Aegla laevis RATHBUN, Proc. U. S. Nat. Mus., vol. 38, p. 602, 1910 (neither
synonymy, except first two entries, nor distribution, except Chile, applies).
Description.—A species of small to moderate size, the largest speci-
men seen measuring 24.5 mm. in length of carapace and rostrum taken
together.
Carapace moderately or a little better than moderately convex.
Rostrum more or less lingulate (more tongue-shaped than sharply
triangular), lateral margins more or less subparallel in the midsec-
tion of the free portion, exceeding eyes by 114 times to nearly twice
19TInasmuch as nearly ail authors since Desmarest (with the exception of Nicolet,
Girard, Fritz Miiller, and Moreira) have considered the genus monotypic and so have failed
to give specifically recognizable descriptions and illustrations of their material, it is im-
possible to assign correctly the many specimens that have in the past been determined as
Aegla laevis to the species to which they properly belong. What I take to be true Aegla
laevis was never well enough characterized to distinguish it from the now known Chilean
species, or, in fact, from any species of Aegla other than Nicolet’s A. denticulata.
THE SPECIES OF AEGLA—SCHMITT 505
the length of the cornea; in lateral view the rostrum inclines down-
ward, although the distal extremity is again lightly but definitely
recurved ; rostral carina very blunt, often somewhat lumpy and some-
times a bit twisted looking, with an irregular row or two of, at most,
microscopically cornified punctae; otherwise, the carina is in general
quite smooth appearing; distally the carina tends to fade out or
disappear, inasmuch as it becomes indistinguishably merged with the
thickened distal, recurved portion of the rostrum which may take in
as much as or sometimes even slightly more than the distal third of
the free portion of the rostrum; either side of the carina, the dorsal
surface of the rostrum is lightly troughed or excavate; at about the
é
Ficure 61.—Aegla laevis (Latreille), male neotype: a, Dorsal view; ), lateral view of anterior
portion; c, sternum of third and fourth thoracic somites; d, inner ventral margin of ischium
of left cheliped; ¢, lateral view of second abdominal epimeron. a, b, natural size; c-e,
twice natural size.
level of the posterior margin of the orbits the rostral carina attains
its greatest elevation, posteriorly it merges in the general surface
of the carapace before reaching the level of the anterior margins of
the protogastric lobes. The rostrum of this species is more or less
amorphous-looking, much as if in the course of the formative processes
it had congealed or become hardened before taking on a truly defini-
tive form.
Protogastric lobes but poorly indicated; except for the gastric
region, anterior portion of the carapace is very coarsely and closely
punctate, the gastric region is smooth appearing, the punctae being
small and relatively widely separated and in part obsolescent ; anteri-
orly the line of demarcation between the two types of punctae defines
the anterior margins of the protogastric lobes, at which level the cara-
506 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
pace also begins to slope down toward the orbits; epigastric promi-
nences coarsely punctate, not very conspicuous, low swellings.
Extraorbital sinus very small, at times obsolescent and represented
by no more than a definite, usually abrupt, often nearly right-angled
offset between the outer end of the orbital margin and the inner slope
or margin of the anterolateral spine; an orbital spine, or rather
spinule, generally present, usually much reduced in size.
Anterolateral spines relatively small, moderately slender, reaching
at least to middle of cornea and often beyond. Anterolateral angle
of first hepatic lobe fairly well marked, little produced, subacute ap-
pearing, though scabrous, and tipped with a corneous scale or two
of about the size of, or very slightly larger than, the scattering of
similar scales on the lateral margin of this lobe; second and third
lobes set off from the preceding and each other by a short, though
plainly marked and nearly closed, notch or incision.
Larger hand relatively of good size, moderately thick and swollen,
finely scabrous, though appearing smooth and evenly rounded. Moy-
able finger with a small but evident, anteriorly spined lobe on outer
margin near base; outer margin of palmar crest more or less sub-
parallel to upper margin of palm proper, cut into three or four
scabrous-margined shallow serrations; with rare exceptions the an-
terior end of upper margin of palmar crest ends abruptly a little
distance behind dorsal anterior margin of palm posterior to the base
of the movable finger, so that a more or less sharply right angled
notch is formed between anterior end of palmar crest and anterior
dorsal margin of palm (a somewhat similar, though less noticeably
and less well developed notch occurs in the subspecies of A. laevis
described below, in A. neuquensis, perhaps also in A. affinis, in
A. riolimayana, and to some degree in A. abtao though in most if
not all other species of Aegla any comparable notch is scarcely to be
distinguished from the toothing or serration of the palmar crest
itself). The palmar crest of A. Zaevis is fairly thin, and slightly exca-
vate or troughed adjacent to the margin of the palm proper.
Ridge of carpus of cheliped above spined, inner margin more or less
obsolescently nodulated (on the carpus of the minor cheliped of one
male the anterior “nodulations” have taken on a distinctly tubercular
form; ordinarily the nodulations on this ridge are low and little
scabrous) ; anterior internal lobe or angle of carpus obtusely tri-
angular, apically carrying two or three stout, pointed, conical, corne-
ous scales; spines of inner margin stout, conical, and acutely corneous
tipped. Upper longitudinal margin of merus furnished with series
of apically scabrous, raised tuberculiform elevations, of which the
anteriormost is the largest; middorsal point of anterior margin of
merus without node or swelling and otherwise unarmed or unorna-
THE SPECIES OF AEGLA—SCHMITT 507
mented. Inner margin of ventral surface of ischium may have as
many as three or four low swellings, the anteriormost of which is
the larger and somewhat conical tuberculiform with tiny corneous
tip; sometimes second and third swellings, though considerably
smaller, are similarly developed; in the neotype only the ultimate
and penultimate of these swellings are developed; though small, each
is corneous tipped; the ischia of most specimens seem to be armed
as in the neotype.
Anterior dorsal angle of epimeron of second (in lateral view, ap-
parent first) abdominal somite acutely produced and corneous tipped ;
anterior margin below acute anterior dorsal angle straight, or at
most only slightly concave.
Neotype.—A male of 24 mm. in length of carapace and rostrum
taken together, one of a lot of 14 males and 17 females (12 ovig.)
contained in the collections of the Museum of Comparative Zoology
(M. C. Z. No. 10478) collected “dans une riviére prés de St. Iago-de-
Chile,” collector and date unknown.
Remarks.—This species in some respects seems to be very much
like A. concepcionensis, though, so far as I am aware, never attaining
so large a size, but throughout its several characters lack the definite-
ness and distinctness of that species. A. concepcionensis, except
in very rare and obviously not typical instances, lacks anything re-
motely resembling the usually abrupt offset between the orbit proper
and the anterolateral spine of A. /aevis; moreover, the anterolateral
spine of its carapace is stouter and more flattened triangular and the
anterolateral lobe is more of an alate expansion in comparison to the
more slender, more conically circular (in cross section) spine and
more triangular anterolateral lobe of the carapace of A. laevis. The
second and third hepatic lobes of A. laevis are the better marked.
Its rostrum is the more truly lingulate of the two, and is more
bluntly carinated. The rostrum of A. concepcionensis is the nearer
an elongate isosceles triangle in shape. The palmar crest of A.
concepcionensis has nothing like the right-angled notch intervening
between the anterior end of the crest and the anterior margin of the
palm in advance of the crest as in A. daevis; moreover, the palmar
crest of A. concepcionensis is not at al] longitudinally troughed or
excavate in any manner suggestive of that state of affairs in A.
laevis.
A. laevis talcahuano, which follows, differs from both A. /aevis and
A. concepcionensis in that the movable finger is wholly without a
trace of a lobe, spined or not, on its outer margin near the base.
Distribution.—Besides the lot of material from which the neotype
has been selected, I have seen two small ovigerous specimens (19 and
21 mm. long) from the Rio Maipo (M. C. Z. No. 1417) collected by
508 PROCEEDINGS OF THE NATIONAL MUSEUM Vol, 91
Lieutenant Gilliss, of the United States Naval Astronomical Expedi-
tion of 1849-52, and determined by William Stimpson; three small
males (15 to 21 mm. long) and one female (17.5 mm.) from near
Melipilla, Province of Sanitago, Chile, which were collected for me
by Dr. Carlos E. Porter; and two lots of two ovigerous females each,
both belonging to the Museo Argentino and carrying the same cat-
alog number (M. A. C. N. No. 4673) but with no indication other than
that they were collected by F. Silvestri in Chile.
Since the foregoing was first written I have seen three additional
specimens of A. laevis. The most interesting of these is one of Dana’s
original specimens, already referred to (pp. 483, 486). Beyond the re-
marks there it is to be noted that the right-angled notch formed be-
tween the anterior end of the palmar crest and the anterior dorsal
margin of the palm is no better developed than in the subspecies
talcahuano below, and that the armature of the ventral inner margin
of the ischium of the right cheliped closely approximates that of the
figured neotype. The specimen in question is 21 mm. in length, cara-
pace and rostrum taken together, and carries Acad. Nat. Sci. Phila.
no. 486.
The other two (Acad. Nat. Sci. Phila. no. 1243) are both females,
18 and 22 mm. in length of carapace and rostrum, respectively. In
the smaller specimen a small extraorbital sinus and a tiny orbital
spinule are present on the right side; on the left side the offset usually
found on the inner margin of the anterolateral spine in the absence
of an orbital spine or spinule is wanting. The larger specimen has no
orbital spinule on either side, but there is instead an appreciable off-
set to the inner slope or margin of each of the anterolateral spines, a
more abrupt offset on the left than on the right side. The hepatic
lobes are rather well marked for A. daevis,; the anterior dorsal epimeral
angles in both specimens are furnished with a small corneous spinule
or sharp scale. In the larger specimen only, the sternal plate between
the chelipeds carries a low, acute, conical, corneous scale, probably
adventitious.
AEGLA LAEVIS TALCAHUANO, new subspecies
FicurRp 62; PLATE 28, B, C
Description.—Very near A. laevis in all particulars except that the
movable finger is wholly without trace of a lobe, whether spined or
not, on its outer margin near the base; the palmar crest, though low
and very remotely suggestive of the subdisciform crest of odebrechtu
and its subspecies, is much narrower than in either of those forms;
margin of the crest, as compared to A. laevis, is scarcely to be de-
scribed as obsolescently serrate; the notch corresponding to the
sharply defined, approximately right-angled one at the anterior end
of the palmar crest of A. laevis is only obscurely and shallowly
THE SPECIES OF AEGLA—SCHMITT 509
present as a slight emargination at the anterior end of the crest in
the type of our subspecies and to an even less degree in the largest
of the Hassler specimens without locality data; in the latter the
crest, though somewhat scabrous, is virtually entire-margined.
Holotype and material examined.—Of this subspecies I have but
two reasonably well developed specimens. The first to come to my
attention was included in a small lot of A. papudo taken by the
Hassler at 'Talcahuano, Chile (M. C. Z. No. 10480). This specimen
has been made the type of the subspecies; it measures 23.0 mm. in
(4 a
Ficure 62.—Aegla laevis talcahuano, new subspecies, male holotype: a, Dorsal view (the ros-
trum is actually slightly distorted, compare pl. 28, B; it has been symmetrically rendered
here by the artist); b, lateral view of anterior portion; c, sternum of third and fourth tho-
racic somites; d, inner ventral margin of ischium of left cheliped; ¢, lateral view of second
abdominal epimeron. a, b, natural size; c-e, twice natural size.
length of carapace and rostrum taken together. The second specimen
(a shade more than 23.0 mm. long) is the largest of three males also
secured by the Hassler Expedition (M. C. Z. No. 10483). It lacks
locality data; the second and third specimens of this lot are
respectively 17 and 14 mm. long.
Remarks.—It is with some hesitation that I have here proposed
this subspecies of Aegla laevis, for, in the light of my studies on the
several forms of Aeg/a occurring east of the Andes, those from their
western slopes do not seem to be either as well marked or as sharply
defined, except of course A. denticulata and A. papudo. More and
better material from Chile, especially from the vicinity of Santiago,
Talcahuano, and Corral, is much needed to properly evaluate A.
laevis and the forms that stand nearest to it.
Distribution—Known only from the type locality, Talcahuano,
Chile, and the one small lot of Hassler specimens without locality
data.
510 PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 91
AEGLA ABTAO Schmitt
FIGURE 63 ; PLATE 28, F, G
Aeglea abtao ScHMITT, Rey. Chilena Nat., vol. 44 (1940), p. 30, pl. 5, fig. 2,
1942.
Description.—A species of moderate size, attaining a length of
carapace and rostrum together of at least 26 mm.
Carapace moderately convex. Rostrum elongate-triangular, but
not particuarly long, exceeding eyestalks by less than the length of
the cornea, sometimes by no more than half the length of the cornea,
fairly straight, not anteriorly reflexed, sharply triangular, trans-
versely flattened and only moderately troughed or excavate either
side of the median carina. Crest of rostral carina almost fades out
near the distal end of the rostrum, which is scaled much as in A.
concépcionensis; the carina behind the level of the posterior margins
of the orbits furnished with two rows of corneous scales set fairly
close together; a little anterior to the orbital margin the two rows
become somewhat intermingled and even imbricated, so much so in
part that in the anterior half of the free portion they form what may
be described as an irregular single row of scales; in distal third of
free portion this row, like the carina itself, tends to fade out, only
suggested by a few scattered scales; raised portion of carina becomes
broader and blunter posteriorly, extending backward about to the
anterior margin of the protogastric lobes. Epigastric prominences
low and blunt; anterior margins of protogastric lobes not particu-
larly set off from the rest of the carapace, but nevertheless well
marked by a row of thick, closely set corneous scales much larger
than the tiny scales seated in most of the punctae of the anterior
portion of the carapace. Areola moderately broad.
Orbits fairly shallow, orbital sinus set off from the distinct and
well formed though small extraorbital sinus by a not large but well-
developed orbital spine.
Anterior extremity of relatively small anterolateral spine scarcely
falling short of, or scarcely reaching, the posterior margin of the
cornea; anterolateral lobes of carapace not particularly flattened;
the anterolateral spines of this species are among the most reduced
in size of any species of Aegla. First hepatic lobe like rest of lateral
margin of anterior portion of carapace minutely spinulated, a
slightly larger corneous spinule tips the subacute anterolateral angle
of this lobe; second and third lobes indicated by slight notchings of
the lateral margin.
Larger hand of good size, swollen, no low ridge as in A. concep-
cionensis apparent. There is an evident, though reduced lobe on the
outer margin of the movable finger near its base; anteriorly the lobe
is small spined. Palmar crest well formed but not high, sharply
THE SPECIES OF AEGLA—SCHMITT 5 |
serrate, serrations spinulated, small spine-tipped; in thickness crest
tapers more or less evenly from base to margin, dorsal surface not
impressed or excavate. No evident ridging on dorsal surface of
carpus other than the usual transversely scabrous, somewhat nodu-
lated ridge above the spined inner margin of the joint. Antero-
internal angle or lobe of carpus armed with an acute, corneous spine
of good size, almost invariably accompanied by a smaller spine lying
immediately against the posterior border of the larger spine; one
or two additional still smaller spines or spinules may be inserted on
the posterior margin of the carpal lobe. Dorsal longitudinal margin
é a
Ficure 63.—Aegla abtao, new species, male holotype: a, Dorsal view; b, lateral view of
anterior portion; c, sternum of third and fourth thoracic somites; d, inner ventral margin
of ischium of left cheliped; ¢, lateral view of second abdominal epimeron. a, }, natural
size; ce, twice natural size.
of merus of cheliped armed with a row of conical tubercles tipped
with several or a few closely juxtaposed pointed corneous scales; at
middle of anterior margin of merus there is a low but evident ante-
riorly convex and fine denticulate swelling. Inner margin of ventral
surface of ischium with a prominent, stout, conical, acutely corneous
tipped spine at anterior end, a very much lower (squat) and perhaps
a little broader one, also with acute corneous tip, at posterior end; at
anterior third of margin there is a similar about subequal swelling
of the same sort as the posterior one, and between these two some-
times a very slight or merely suggested swelling or nodulation.
Anterior dorsal angle of epimeron of second (in lateral view, ap-
parent first) abdominal somite somewhat produced and armed with
512 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 91
an acute, flattened, corneous spine; anterior margin below spine about
straight; ventral angle rounded off.
Holotype.—The largest of seven specimens (5 males and 2 females),
a male measuring 26.6 mm. in median length of carapace and rostrum
together, U.S.N.M. No. 79079. The smallest specimen is also a male
and measures about 11 mm. in median length of carapace and
rostrum. The specimens were collected by Dr. C. H. Eigenmann
at Abtao, Chile, February 22, 1919.
Remarks.—See under A. riolimayana, “Remarks,” p. 515.
Distribution.—W ith certainty at present known only from the type
locality, Abtao, Chile. An unmistakable representative of the species,
an untagged male of 28.0 mm. in length of carapace and rostrum,
was found along with several other specimens in a bottle of material
borrowed from the Buenos Aires Museum. One was the type of
A. affinis (M.A.C.N. No. 9817), another an unattached cheliped of
the same species (tagged M.A.C.N. No. 4186), and an untagged
female of A. hwmahuaca (22.0 mm. long). In the catalogs of the
Museo Argentino Ciencias Naturales entry No. 4186 reads simply,
“Neuquen, Mayo 16, 1898; Sr. Carlos Burmeister”; entry No. 9187
concerns specimens of Mytilus chorus Molina received in exchange
from Dr. Carlos 8. Reed, 21-V, 1919. The bottle contains a mixture
of things, or else a misattached label or labels, and to the untagged
specimens no locality at all may be safely attached.
Further, I have before me a small male of 19.5 mm. in length of
carapace and rostrum together, also collected by Dr. Eigenmann in
Chile, “Falls of Petrohue,” March 8, 1919. Although this particular
specimen has been only tentatively placed with A. abtao, it is prob-
ably correctly determined; the rostrum seems a bit more slender than
typical A. abtao, the areola perhaps a bit narrower and the ventral
inner margin of the ischium somewhat smoother.
Almost too late for mention, I received a very fine, dried example of
this species from Dr. Carlos A. Porter. It measures 26 mm. in length
of carapace and rostrum and was collected by Dr. Porter himself, in
December 1941, near “El Valean,” Santiago, Chile. The rostrum fits
the description of the type almost exactly; indeed this specimen is a
very close counterpart of the type. The lobular tooth on the fixed
finger of the minor right cheliped is no more in evidence than in the
type (fig. 638, a). However, the conical tubercles on the dorsal longi-
tudinal margin of the merus of the cheliped appear single-spined or
spinule-tipped; the inner ventral margin of the ischium is as in the
type on the right cheliped; on the left one there are two small eleva-
tions of which the anterior is the larger and small spinule-tipped
between the anterior and posterior spines. The anterior dorsal angle
of the epimeron of the second abdominal somite is armed with two
small spines or spinules on the left side, with one only on the right.
THE SPECIES OF AEGLA—SCHMITT 513
AEGLA RIOLIMAYANA, new species
Ficure 64; PLatEe 28, 5
Description.—A species of perhaps moderate size, the largest speci-
men so far seen does not exceed 24.0 mm. in length of carapace and
rostrum together. Stands near the preceding species, A. abtao.
Like A. abtao, our species has the carapace moderately convex;
the rostrum, though basally broad and flattened, distally is narrowly
and sharply triangular, almost stilletolike, straight, and more or less
sharply carinated to the tip (A. riolimayana has the most sharply
acuminate and distally narrowed rostrum of all species included in
the A* section of our diagnostic key) ; the tip of the rostrum extends
beyond the eyestalks by about one-half the length of the cornea; the
rostral carina is armed with a somewhat wavering, virtually single
line of small tiny corneous scales, which get a little larger anteriorly ;
toward the tip these scales sometimes, for a very brief interval, may
form an irregular double row; the dorsal surface of the rostrum is
noticeably depressed or excavate either side between the rostral carina
and the seemingly elevated lateral margins of the rostrum; the rostral
carina runs back about to the level of the anterior margins of the
protogastric lobes which, like the epigastric prominences, are not
particularly well marked.
Orbital sinus relatively wide, orbital spine but a spinule, extra-
orbital sinus small, at times scarcely more than a notch at the base
of the inner slope or margin of the anterolateral spine; the latter
small, conical, scarcely reaching the posterior margin of the cornea.
Anterolateral angle of the first hepatic lobe well marked, though no
more than scabrous with corneous scales no larger than the others
with which the lateral margins of the hepatic lobes are armed;
second and third hepatic lobes scarcely more than sinuosities in the
lateral margin of the forepart of the carapace.
Hand of moderate size, moderately inflated; lobular tooth on fixed
finger relatively small but plainly marked; a definite, though small,
spined lobe on outer margin of movable finger near base. Palmar
crest resembling that of A. Zaevis, outer margin of crest more or less
subparallel to upper margin of palm proper, and cut into three or four
scabrous-margined shallow serrations, anterior angles or apices of
serrations, however, armed with a sharp-pointed scale or spinule;
as in A. laevis there is a more or less definitely right-angled notch
between anterior end of the palmar crest and the anterior dorsal
margin of palm.
Ridge of carpus of cheliped above spined inner margin not prom1-
nent, low and obsolescently nodulated; armed on these low swellings
with a few small corneous denticles or scales; spined inner margin
armed with slender, conical, clean-cut spines, of which the anterior-
514 PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 91
most is longest and most slender; anterior internal lobes of carpus
armed with a single, well-developed, smooth, clean-cut, spine; all
carpal spines with acute corneous tips. Upper longitudinal margin
of merus with a series of sharp corneous spines, of which the anterior-
most is the larger and elevated on a small conical tubercle above the
level of the rest; anterior margin of merus in front of this anterior
spine has a very slightly marked, minutely denticulate lobe; a few
other tiny denticles may also occur along the anterior margin of the
merus. Inner margin of ventral surface of ischium with low, broadly
conical, corneous scale-tipped tubercle at anterior end and a relatively
insignificant, low, nodular swelling at posterior end, margin of ischium
between these two low elevations virtually straight, at most only very
shghtly sinuous.
Ficure 64.—Aegla riolimayana, new species, male holotype: a, Dorsal view; }, lateral view
of anterior portion; ¢c, sternum of third and fourth thoracic somites; d, inner ventral margin
of ischium of left cheliped; ¢, lateral view of second abdominal epimeron. a, b, natural
size; c—e, twice natural size.
Anterior dorsal angle of epimeron of second (in lateral view, ap-
parent first) abdominal somite produced to form an acute corneous
tipped spine; anterior margin below this spine more or less straight.
Holotype.—The largest of five males taken by John W. Titcomb,
November 19, 1903, in the Rio Limay, which forms the boundary line
between the territories of Rio Negro and Neuquen, Argentina. These
specimens were taken not far from the outlet of Lago Nahuel Huapi,
where Mr. Titcomb had obtained other specimens of this species a
few days before. The holotype, U.S.N.M. No. 80025, measures
23.5 mm. in length of carapace and rostrum together.
THE SPECIES OF AEGLA—SCHMITT Sib
Remarks.—This species and A. abtao are related. They are very
similar in appearance and in common have noticeably short antero-
lateral spines, relatively shorter than in other Aeglas, yet on
close examination there appear constant differences in the general
shape of the rostrum, its relative degree of flatness and excavation,
and distal attenuation. The anterolateral angle of the first hepatic
lobe of A. abtao seems always to be acutely armed with a small
spinule or sharply pointed scale, larger than those generally arming
the lateral margin of the forepart of the carapace; in riolimayana
this angle is more or less rounded off, at most subacute, and scabrous
with scales no different from those generally arming the lateral
margins of the hepatic lobes. The anterior internal lobe of the carpus
of the chelipeds seems to be differently armed or spined in the two
species; there seems to be less nodulation of the inner ventral border
of the ischium of the chelipeds in A. rioliémayana than in A. abtao.
The posterior more or less straight portion of the lateral grooves or
furrows of the areola are subparallel in A. abtao; in A. riolimayana
they exhibit a decided convergence posteriorly; the straight sections
of the lateral boundaries of the areola are farther removed from the
lateral suture lines of the cardiac area at their posterior than at
their anterior ends (fig. 64) ; in A. abtao the reverse is true (fig. 63).
Distribution.—All specimens of this species that I have seen are
from the Rio Limay in the vicinity of Lago Nahuel Huapi or from
the lake itself, or from their immediate tributaries. In addition to
the type lot of five males, Mr. Titcomb obtained some 20 specimens,
males and females nearly equally divided, from the outlet of the lake,
November 15, 1903. Of these the largest and smallest males are,
respectively, 24 and about 9 mm. in length of carapace and rostrum
taken together, the largest and smallest females 20.5 and 10.0 mm.,
respectively; two small males (8.5 and 14 mm.) from Arroyo de
Jones, tributary to Lake Nahuel Huapi; and another small male
(21 mm.) from “Victoria Island, Nahuel Huapi,” November 29, 1903.
On November 22, 1926, R. C. Shannon collected one small male
(16.0 mm.) at Correntoso, north end of Lago Nahuel Huapi, which
he presented to the United States National Museum. Otherwise, I
have examined three small specimens belonging to the Museo Argen-
tino, two small females (19.0 and 20.0 mm.) from Lago Nahuel
Huapi, which had been purchased from Emilio Budin (M.A.C.N.
No. 9679), and one male (20.0 mm., M.A.O.N. No. 8388), which
appears to be this species and which carries merely the designation
“Neuquen” [Territory ?].
LITERATURE CITED
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THE SPECIES OF AEGLA—SCHMITT 517
GIRARD, CHARLES.
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THE SPECIES OF AEGLA—SCHMITT 519
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INDEX TO DESCRIPTIONS OF AND DIAGNOSTIC NOTES ON THE
SPECIES AND SUSPECIES OF AEGLA
abtao, 457, 510. laevis taleahuano, 457, 508.
affinis, 456, 495. neuquensis, 456, 493.
eastro, 453, 473. odebrechtii, 455, 487.
concepcionensis, 456, 501. odebrechtii paulensis, 455, 490.
denticulata, 454, 480. papudo, 455, 483.
franca, 453, 476. parana, 452, 458.
humahuacea, 456, 498. platensis, 452, 464.
intermedia, 4386, 448. prado, 458, 470.
jujuyana, 454, 478. riolimayana, 457, 513.
laevigata, 435. sanlorenzo, 452, 461.
laevis, 436, 457, 504. uruguayana, 453, 467.
NOTE
Unless otherwise stated, the photographs shown in the plates that followfare of the male
holotype, approximately natural size.
520
U.S. GOVERNMENT PRINTING OFFICE: 1942
U.S. NATIONAL MUSEUM PROCEEDINGS, VOL. 91
PLATE 25
zo C. 7 latensts; A. uruguayana.
A, Aegla parana; B, A. sanlorenzo; C, A. platensis; D, 4. uruguay
U.S. NATIONAL MUSEUM PROCEEDINGS, VOL. 91 PLATE 26
A, Aegla prado; B, A. prado, male paratype; C, A. denticulata, male neotype;
D, A. franco; E, A. jujuyana; F, A. castro.
U.S. NATIONAL MUSEUM PROCEEDINGS, VOL. 91 PLATE 27
. Tn a wana h ; .
A, Aegla odebrechtii; B, A. 0. paulensis; C, A. papudo, D, A. humahuaca;
E, A. neuquensis; F, A. affinis (the ambulatory leg shown is the inadvert-
ently reversed left leg of text fig. 58).
PLATE 28
PROCEEDINGS, VOL. 91
U.S. NATIONAL MUSEUM
‘rad Aqeied ayew ‘oviqv *p ‘ty Sovjqv wa
‘puvkowuyord Pp ‘sp tadAjoau ayew ‘s1ean) +p Sq fedAqesed ayeu
‘
OUDNYVIIDI *] “PD Sounnynryy staan] *p ‘gq fsisuauotsgaauog vj3ap ‘Vv
INDEX
(New genera, species, ete., are printed in italics.)
abtao, Aegla, 488, 441, 450, 456, 457, 498,
500, 506, 510, 511 (fig.), 512, 518,
515.
Aeglea, 510.
acanthias, Miodon, 123.
Urobelus, 123.
Aciprion, 71.
formosum, 71, 72 (fig.), 73 (fig.).
majus, 74.
acolytus, Ellipsodon, 28, 29, 30.
acutirostris, Rana, 136.
Rana albolabris, 118, 1386.
acutus, Crocodylus, 57.
ademetzi, Hyperolius, 132.
adventitia, Corymorpha, 79, 89.
Aegia, the species of, 481.
abtao, 438, 441, 450, 456, 457, 498,
og 506, 510, 511 (fig.), 512, 513,
15.
affinis, 432, 441, 449, 456, 486,
496 (fig.), 508, 504, 506, 512.
castro, 432, 483, 440, 441, 448,
453, 460, 473, 474 (fig.), 498.
eoncepcionensis, 452, 483, 441,
486, 501 (fig.), 503, 504, 507, 510.
denticulata, 433, 435-487, 441, 448,
458, 454, 458, 470, 480, 481 (fig.),
482, 488, 504, 509.
franca, 433, 440. 441, 448, 444, 449,
451, 453, 476, 477 (fig.), 478, 480.
humahuaca, 432, 441, 442, 446, 448,
449, 456, 478, 480, 498, 499 (fig.),
500, 501, 512.
intermedia, 481. 486, 439, 440, 448.
jujuyana, 432, 441-444, 446, 449, 451,
454, 478, 479 (fig.), 500
laevis, 481, 433-441, 457, 508, 504,
505 (fig.), 506-509, 518.
laevis talcahuano, 433, 436, 441, 447,
456, 457, 507, 508, 509 (fig.).
neuquensis, 433, 441, 456, 486, 498
(fig.), 496, 497, 506.
odebrechtii, 431-488, 437-441, 443,
445, 448, 455, 547, 486, 487 (fig.),
488-492, 508.
odebrechtii paulensis, 433, 440, 441,
443, 445, 455, 475, 486, 488, 490,
491 (fig.), 492, 493.
papudo, 432, 438, 441, 455, 488, 484
(fig.), 486, 498, 508, 504, 509.
parana, 432, 433, 440, 441, 450, 452,
458, 459 (fig.), 463, 472, 475, 476.
platensis, 433, 440-442, 447, 452, 464,
465 (fig.), 467, 469, 470, 473.
495,
451,
456,
Aegla prado, 482, 433, 441, 442, 450, 451,
453, 469, 470, 471 (figs.), 472, 478,
475, 476, 483.
riolimayana, 433, 441, 447, 449, 450,
456, 457, 506, 512, 513, 514 (fig.),
515.
sanlorenzo, 482, 441, 446, 450-452,
460, 461, 462 (fig.), 464, 472, 475.
uruguayana, 432, 440-442, 453, 463,
464, 466, 467, 468 (fig.), 472, 478.
Aeglea, 434, 489, 448, 444.
abtao, 510.
concepcionensis, 501.
denticulata, 480.
intermedia, 487, 490.
laevis, 433, 476, 486, 504.
odebrechtii, 487.
Aeglidae, 481, 458.
aequidens, Ellipsodon, 28, 30.
aethiopissa, Rhamnophis, 121,
Rhamnophis aethiopissa, 121.
affinis, Aegla, 432, 441, 449, 456, 486, 495,
496 (fig.), 508, 504, 506, 512.
africana, Agama agama, 114, 116.
africanus, Tropidolepis, 116.
agama africana, Agama, 114, 116
agama savatieri, Agama, 114, 116.
Agama agama africana, 114, 116.
agama savatieri, 114, 116.
savatieri, 116.
Aglaophenia, 86.
inconstans, 86, 89.
transitionis, 87, 89.
Alabama, crayfishes from caves of, 141.
Alaska, catalog of Eskimo crania in
U. S. National Museum, 169.
albolabris, Rana, 136.
Rana albolabris, 134, 186.
albolabris acutirostris, Rana, 113, 136.
albolabris albolabris, Rana, 184, 136.
albolabris parkeriana, Rana, 113, 136.
Aliurus, 114, 116.
ornatus, 116.
alleni, Phrynobatrachus, 139.
Pseudoxenopus, 1385.
Rana, 114.
Rana crassipes, 134, 135.
Amphibians and reptiles from Liberia,
Smithsonian - Firestone Expedition,
113.
amplectens, Galathea, 438.
angustus, Haploconus, 49, 50, 51.
Anisonchus, 45, 46, 48, 50.
dracus, 3, 45, 46 (fig.), 47.
gillianus, 45-48,
521
Dee PROCEEDINGS OF THE NATIONAL MUSEUM Vou, 91
Anisonchus oligistus, 8, 47 (fig.). biscutata, Dipsas, 159.
onostus, 3, 46, 47 (fig.), 48. biscutata latifascia, Trimorphodon, 160.
sectorius, 45, 46. biscutatus, Trimorphodon, 149-155, 158,
anoscopus, Natrix, 114, 119. 159, 165.
Natrix anoscopus, 114, 118. Trimorphodon biscutatus, 149, 150,
Tropidonotus, 118. 153, 154, 159.
anoscopus gendrii, Natrix, 119. biseutatus quadruplex, Trimorphodon,
Anthaspidella, 99. 149-152, 154, 155, 157.
elintoni, 99, 102. Bitis gabonica, 123.
florifera, 100. nasicornis, 124.
mammuilata, 99. blandingii, Boiga, 122.
traini, 100, 102. Dipsas, 122.
Anthaspidellidae, 99. Euprepes, 117.
apachea, Arachnis, 60, 61, 68. Mabuya, 117.
apancora, Cancer, 484. Boaedon lineatus, 114, 120.
Aparallactus modestus, 123. lineatus lineatus, 118, 120.
aphantus, Dracontolestes, 3, 18 (fig.). olivaceus, 121.
Aphronorus, 11, 12. virgatus, 114.
fraudator, 11. Boiga blandingii, 122.
simpsoni, 3, 11 (fig.), 12, 18. Brachygnathosuchus, 56.
Aporthophyla typa, 92. braziliensis, 55.
aquilonius, Ellipsodon, 28, 29, 30. bracteata, Symplectanea, 78, 89.
Arachnis, North American moths, 59. brasiliensis, Purusaurus, 55.
apachea, 60, 61, 68. braziliensis, Brachygnathosuchus, 55.
aulaea, 61, 62, 63. brongersma!, Phrynobatrachus, 114, 138.
aulaea pompeia, 61, 68. Phrynobatrachus ogoensis, 138.
aulea, 62. Bufo camerunensis camerunensis, 124.
incarnata, 62, 63. cinereus, 124.
maia, 66. latifrons, 125.
midas, 59, 60, 61, 69. maculatus, 124.
picta, 59, 61, 63, 69. regularis maculatus, 119, 124.
picta citra, 60, 67.
picta hampsoni, 61, 66. cahni, Cambarus (Cambarus), 146, 147
picta insularis, 61, 66. (fig.).
picta maia, 61, 66. Caiman, 56.
picta picta, 65. crocedylus, 57.
picta verna, 61, 65. calearatus, Arthroleptis, 137.
pompeia, 63. Hemimantis, 187.
zuni, 60, 61, 69.
Archaeocyathus, 98.
Archaeoscyphia, 93.
calgariensis, Catopsalis, 8, 9.
Calotes versicolor, 116.
Calycocoelia, 96.
minganenis, 94, 102. ees 2
Archaeoscyphidae, 93. ues oe Neg
Archeoscyphia minganensis, 94. (Cambarus) cahni, 146, 147 (fig.).
Arctiidae, 59.
Arctocyonidae, 3.
Arthroleptis calearatus, 137.
poecilonctus, 137.
werneri, 137.
Atlantic Ocean, new hydroids from, 77.
extraneus, 146, 147.
(Cambarus) hamulatus, 146, 147.
pellucidus, 141, 142, 144.
(Faxonius) pellucidus azstralis,
142, 143 (fig.).
aulaea, Arachnis, 61, 62, 63 pellucidus pellucidus, 144.
Ecpantheria, Goes pellucidus testii, 144.
aulaea pompeia, Arachnis, 61, 63. Cambarus, subg., 149. ‘
aulea, Arachnis, 62. camerunensis, Bufo camerunensis, 124.
australis, Cambarus (Faxonius) pelluci- | Campanularia fasciculata, 81, 89.
dus, 142, 148 (fig.). Campanularidae, 81.
Campanulinidae, 82.
Cancer apancora, 434.
barrandei, Pliomerops, 91. carinidens, Periptychus, 42, 43, 44, 45.
Bassler, R. S., on Nevada Early Ordovi- | Carnivora, 3, 17.
cian (Pogonip) sponge fauna, 91. Carsioptychus, 4, 42, 48, 45.
bequaertii, Leptopelis, 113, 128. coarctatus, 42-44.
Bessoecetor, 12. hamasvitus, 3, 41 (fig.), 42.
dilueuli, 12. Cassina weali, 125.
thomsoni, 12. cassinoides, Hylambates, 125.
biatheles, Protogonodon, 3, 17, 19 (fig.). | castro, Aegla, 432, 433, 440, 441, 448, 451.
bibronii, Rana, 184, 185. 453, 460, 478, 474 (fig.), 493.
INDEX
Catopsalis, 8.
calgariensis, 8, 9.
fissidens, 8, 9.
foliatus, 8, 9.
utahensis, 3, 8, 9 (fig.).
Causus rhombeatus, 123.
Caves, crayfishes from in Alabama, 141.
Chamaeleo gracilis, 118.
gracilis etiennei, 118.
gracilis gracilis, 118.
chitalonensis, Dromicus, 107, 108.
chorus, Mytilus, 498, 501, 512.
Chriacus, 22.
cinereus, Bufo, 124.
citra, Arachnis picta, 60, 67.
Claenodon corrugatus, 18.
ferox, 18.
procyonoides, 23.
Clarke, J. F. G., on North American
moths of the genus Arachnis, 59.
clintoni, Anthaspidella, 99, 102.
Patellispongia, 97, 101.
coarctatus, Carsioptychus, 42, 48, 44.
cochranae, Hylambates, 118, 125.
Coelopeltis virgata, 120.
collaris, Trimorphodon,
168, 165.
Colombia, a new fossil crocodilian from,
DD.
Coluber nasicornis, 124,
smythii, 121.
comma, Conoryctes, 16, 17.
Conacodon, 52.
cophater, 50, 51.
concepcionensis, Aegla, 432, 483, 441,
456, 486, 501 (fig.), 508, 504, 507,
510.
Aeglea, 501.
coneolor, Hyperolius, 130, 182.
Ixalus, 130.
Condylarthra, 3, 24.
Coniophanes, 103.
fissidens, Mexican subspecies of, 103.
fissidens dispersus, 106, 109-111.
fissidens fissidens, 104, 105, 107-111.
fissidens proterops, 104, 105, 107-
at:
fissidens punctigularis, 107, 109-111.
proterops, 105.
punctigularis, 107.
Conoryctella, 15.
dragonensis, 3, 15, 16 (fig.).
Conoryctes, 16.
comma, 16, 17.
cophater, Conacodon, 50, 51.
Cophoscincopus durus, EG
Cophoscineus dura, 117.
simulans, 117.
corniculatus, Haploconus, 49, 51.
Coronella fissidens, 104.
triangularis, 121.
corrugatus, Claenodon, 18.
Corymorpha, 79.
adventitia, 79, 89.
Corymorphidae, 79.
Crania, Eskimo, in U. S. National Mu-
seum, 169.
149, 151-154,
523
crassa, Tubularia, 81, 89.
erassicolidens, Tricentes,
crassipes, Rana, 114.
crassipes alleni, Rana, 134, 135.
crassiiheca, Lictorella, 85, 89.
Craytishes from Alabama caves, 141,
crocodylus, Caiman, 57.
Crocodylus acutus, 57.
err duchesnii guineensis, 113,
122%
hotamboeia hotamboeia, 122.
shrevei, 122.
Crotaphytus, 72.
Jrustacea, the species of Aegla, 431.
crayfishes from Alabama caves, 141.
cuspidata, Hudaemonema, 13, 14.
cuspidatus, Oxyclaenus, 21.
Oo
hate
Decapoda, the species of Aegla, 431.
Deinosuchus, 56.
Dendroaspis viridis, 123.
denticulata, Aegla, 433, 485-487, 441, 448,
458, 454, 458, 470, 480, 481 (fig.),
482, 483, 504, 509.
Aeglea, 480.
Testudo, 115.
Desmatoclaenus, 4, 34, 36, 37.
hermaeus, 3, 34, 35 (fig.), 37, 38.
paracreodus, 3, 19, 37, 38 (fig.).
Deuterogonodon montanus, 19, 20.
Didymictis, 3, 24.
haydenianus, 24.
microlestes, 24.
sp., 24.
diluculi, Bessoecetor, 12.
Dinophis hammondii, 123.
Dinosuchus, 56.
neiwensis, 56.
terror, 55, 56.
Dipsadoboa, 122.
unicolor, 122.
Dipsas biscutata, 159.
blandingii, 122.
globiceps, 122.
valida, 122.
Dipsoglyphophis, 122.
disjunctus, Litaletes, 29, 32, 33.
dispersus, Coniophanes fissidens,
109-111.
dissentaneus, Litomylus, 238.
Distichurus maculatus, 123.
ditrigonus, Ectoconus, 39, 40.
Dracoclaenus, 24, 26.
griphus, 3, 22, 24, 25 (fig.).
Dracontolestes, 18.
aphantus, 3, 13 (fig.).
dracus, Anisonchus, 3, 45, 46 (fig.), 47.
dragonensis, Conoryctella, So libs oe
(fig.).
Dromicus chitalonensis, 107, 108.
dubium, Halecium, 83, 89.
duchesnii guineensis, Crotaphopeltis,
113, 122.
dura, Cophoscineus, 117.
Tiliqua, 117.
durus, Cophoscincopus, Te:
106,
524
Marly Ordovician (Pogonip)
fauna from Nevada, 91.
Hehidna gabonica, 1238.
echinata, Lacerta, 114, 117.
Lacerta (Zootoca), 116.
Lacerta echinata, 116.
echinata echinata, Lacerta, 116.
echinata langi, Lacerta, 117.
EKepantheria aulaea, 62.
incarnata, 62, 63.
picta, 64.
Ectoconus, 4, 39, 40.
ditrigonus, 39, 40.
majusculus, 39.
symbolus, 3, 89, 40 (fig.).
Ectoganus, 17.
EHgmundella, 82.
grandis, 82, 89.
elachistus, Haploconus?, 3, 50, 51 (fig.).
Elapops modesius, 128.
elassus, Tricentes, 3, 22, 23 (fig.).
Ellipsodon, 3, 28, 30, 32.
acolytus, 28-80.
aequidens, 28, 30.
aquilonius, 28-80.
inaequidens, 28, 30, 31.
lemuroides, 28-380.
priscus, 28, 29, 32.
shepherdi, 3, 28, 29 (fig.), 31-33.
sp., 31.
sternbergi, 3, 30 (fig.), 33.
Elpidophorus, 138, 14.
minor, 138, 14.
patratus, 18, 14.
Hoconodon, 20.
erosa, Kinixys, 115.
Testudo, 115.
Eskimo, catalog of crania in U. S. Na-
tional Museum, 169.
Ethmophyllum minganense, 94.
etiennei, Chamaeleo gracilis, 118.
Eudaemonema, 18, 14.
cuspidata, 18, 14.
Huprepes blandingii, 117.
frenatus, 117.
Pxostinus, 71, 74.
serratus, 74, 75 (fig.).
extraneus, Cambarus, 146, 147.
fasciatus, Hemichromis, 119.
Hemidactylus, 116.
fasciculata, Campanularia, 81, 89.
fasciolata, Trimorphodon, 149, 151, 152,
154, 160, 162, 163.
Faxonius, subg., 142.
ferox, Claenodon, 18.
Tropidonotus, 114, 118.
ferronensis, Ptilodus, 3, 10 (fig.).
festivus, Hyperolius, 133.
fissidens, Catopsalis, 8, 9.
Coniophanes, Mexican subspecies of,
103.
Coniophanes fissidens, 104, 105, 107-—
ae
Coronella, 104.
PROCEEDINGS OF THE NATIONAL MUSEUM
VOL, 91
sponge | fissidens dispersus, Coniophanes, 106,
109-111.
fissidens fissidens, Coniophanes, 104, 105,
107-111.
fissidens proterops, Coniophanes, 104,
105, 107-111.
fissidens punctigularis,
107, 109-111.
florifera, Anthaspidella, 100.
foliatus, Catopsalis, 8, 9.
fon beet Trimorphodon, 149, 151, 153, 154,
Coniophanes,
nee Aciprion, 71, 72 (fig.), 7
(fig.).
formosus, Hemidactylus, 116.
Fossils, a new ecrocodilian from Colom-
bia, 55.
Nevada Harly Ordovician (Pogonip)
sponge fauna, 91.
Oligocene lizards from Wyoming, 71.
franca, Aegla, 433, 440, 441, 443, 444,
449, 451, 458, 476, 477 (fig.), 478, 480,
francisci, Phrynobatrachus, 1388.
Fraser, ©. McLean, on Atlantie hy-
droids, 77.
fraudator, Aphronorus, 11.
frenatus, Huprepes, 117.
Frogs and toads, Liberian, 124.
fuliginoides, Natrix, 114, 120.
fulvovittatus, Hyperolius, 1380.
Megalixalus, 130.
fusciventris, Hyperolius, 182, 133.
gabonica, Bitis, 123.
Hehidna, 123.
Galathea, 488, 489.
amplectens, 4388.
laevis, 484, 504.
Gazin, C. Lewis, on the mammalian
faunas of the Paleocene of central
Utah, with notes on the geology, 1.
gendrii, Helicops, 114, 119.
Natrix anoscopus, 119.
gillianus, Anisonchus, 45-48.
Gilmore, Charles W., on some little-
known fossil lizards from the Oligo-
cene of Wyoming, 71.
gilmorei, Periptychus, 3, 42, 43 (fig.), 44
(fig.), 45.
globiceps, Dipsas, 122.
Goniacodon, 3, 23.
levisanus, 23, 24.
sp., 23.
Goniotrema perplexa, 92.
gracilis, Chamaeleo, 118.
Chamaeleo gracilis, 118.
gracilis etiennei, Chamaeleo, 118.
gracilis gracilis, Chamaeleo, 118.
grandis, Egmundella, 82, 89.
grandis, Lovenella, 83, 89.
Nevadocoelia, 95, 101.
Grayia smythii, 121.
gribinguiensis, Rana oxyrhynchus, 134,
135, 186.
griphus, Dracoclaenus, 3, 22, 24, 25 (fig. ».
INDEX
guineensis, Crotaphopeltis duchesnii,
113, 122.
Leptodira, 122.
Halecidae, &8.
Halecium, 838.
dubium, 83, 89.
kiikenthali, 85.
telesenpium, 84.
tensum, 84, 89.
hamazitus, Carsioptychus, 3, 41 (fig.),
42,
hammondii, Dinophis, 128.
hampsoni, Arachnis picta, 61, 66.
hamulatus, Cambarus (Cambarus), 146,
147.
Haploconus; 4, 48, 49, 51, 52.
angustus, 49-51,
corniculatus, 49, 51.
elachistus, 3, 50, 51 (fig.).
inopinatus, 3, 49 (fig.), 50, 51.
Hapsidophrys lineata, 12i.
lineatus, 121.
haydenianus, Didymictis, 24.
Helicops gendrii, 114, 119.
Hemichromis fasciatus, 119.
Hemidactylus, 114, 116.
fasciatus, 116.
formosus, 116.
Hemimantis ealecaratus, 137.
hermaeus, Desmatoclaenus, 3, 34, 35
(fig.) , 37, 38.
Hesperocoelia, 93, 98.
typicalis, 98, 102.
undulata, 99, 102.
Holurophis olivaceus, 121.
homeana, Kinixys, 115.
hotamboeia, Crotaphopeltis hotamboeia,
a:
hotamboeia hotamboeia, Crotaphopeltis,
ipa
Hrdlitka, Ales, on catalog of Eskimo
erania in U. 8. National Museum, 169.
humahuaca, Aegla, 432, 441, 442, 446, 448,
449, 456, 478, 480, 498, 499 (fig.), 500,
501, 512.
Hyalella inermis, 487.
Hyalostelia, 92.
Hyé@ractinia, 79.
walens, 79, 89.
Hydractinidae, 79.
Hydroids, new Atlantic, 77.
Hylambates cassinoides, 125.
cochranae, 113, 125.
leonardi, 113, 125, 128.
viridis, 128,
hyloides, Leptopelis, 128.
Hyopsodontidae, 3.
Hyperolius ademetzi, 132.
concolor, 130, 132.
festivus, 133.
fulvovittatus, 130.
fusciventris, 132, 133.
ocellatus, 131.
pleurotaenius, 131,
picturatus, 131, 133.
535587—_43——_2
525
Hyperolius plicatus, 188.
riggenbachi, 130,
vittiger, 130.
Ictidopappus, 24.
Iguanidae, 71.
inaequidens, Ellipsodon, 28, 30, 31.
incarnata, Arachnis, 62, 63.
Eepantheria, 62, 63.
inconstans, Aglaophenia, 86, 89.
inermis, Hyalella, 437.
ROL IaEL, Haploconus, 3, 49 (fig.), 50,
Insectivora, 3, 11.
insularis, Arachnis picta, 61, 66.
intermedia, Aegla, 431, 436, 439, 440, 448.
Aeglea, 487, 490.
Typhlops, 118.
Ixalus concolor, 130.
Jepsenia, 32.
mantiensis, 3, 31, 32, 33 (fig.).
jujuana, Aegla, 482, 441-444, 446, 449,
451, 454, 478, 479 (fig.), 500.
Kassina, 128.
senegalensis, 125.
kimbetovius, Protogonodon, 19, 39.
Kinixys erosa, 115.
homeana, 115.
kirki, Toquimia, 92.
kiikenthali, Halecium, 85.
Lacerta echinata, 114, 117.
(Zootoca) echinata, 116,
echinata echinata, 116.
echinata langi, 117.
langi, 114.
laevis, Aegla, 431, 4383-441, 457, 503, 504,
505 (fig.), 506-509, 513.
Aeglea, 4383, 476, 486, 504.
Galathea, 434, 504.
laevis talcahwano, Aegla, 438, 486, 441,
447, 456, 457, 507, 508, 509 (fig.).
Lafoeidae, 85.
lambda, Trimorphodon, 149-151, 153-157,
Lampra, 80.
wvularis, TT, 80, 89.
Lampropeltis, 162.
leonis, 152.
langi, Lacerta, 114.
Lacerta echinata, 117.
latifascia, Trimorphodon, 149, 151, 152,
154, 160, 162, 165, 166.
Trimorphodon biscutata, 160.
latifrons, Bufo, 125.
Phrynobatrachus, 139.
Leiurus ornatus, 116.
lemuroides, Ellipsodon, 28, 29, 30.
leonardi, Hylambates, 113, 125, 128.
leonis, Lampropeltis, 152.
Leptodira guineensis, 122.
Leptopelis bequaerti, 113, 128.
hyloides, 128.
liberiensis, 114, 128.
526
Leptopelis nanus, 128.
tessmanni, 128, 129.
togoensis, 128.
viridis, 114, 128.
Leptophis viridis, 128.
leucostictus, Typhlops, 118.
levisanus, Goniacodon, 28, 24.
Liberia, reptiles and amphibians from,
118.
liberiensis, Leptopelis, 114, 128.
Phrynobatrachus, 1388.
Typhlops, 118.
Lictorella, 85.
crassitheca, 85, 89.
lineata, Hapsidophrys, 121.
lineatus, Boaedon, 114, 120.
Boaedon lineatus, 113, 120.
Hapsidophrys, 121.
lineatus lineatus, Boaedon, 118, 120.
Lissocoelia, 96.
ramosa, 96, 101, 102.
Litaletes, 29, 32, 33.
disjunctus, 29, 32, 33.
Litomylus dissentaneus, 28.
Liurus, 116.
ornatus, 116.
Lizards, fossil, from Wyoming Oligocene,
cl!
Liberian, 116.
longirostris, Rana, 113, 183, 13865.
Lovenella, 83.
grandis, 83, 89.
Loveridge, Arthur, on reptiles and am-
phibians from Liberia, 113.
Loxolophus, 21.
Lycodon lyrophanes, 156.
lyrophanes, Lycodon, 156.
Trimorphodon, 149-151, 154-157.
Mabuya blandingii, 117.
maccarthyensis, Rana, 184, 136.
maculatus, Bufo, 124.
Bufo regularis, 119, 124.
Distichurus, 1238.
magnipora, Patellispongia, 98, 101.
maia, Arachnis, 66.
Arachnis picta, 61, 66.
major, Trimorphodon, 159.
majus, Aciprion, 74.
majusculus, Hetoconus, 39.
Mammals, fossil, from Paleocene of cen-
tral Utah, 1.
mammulata, Anthaspidella, 99.
Mann, William M., 113.
manni, Typhlops, 118, 118.
mantiensis, Jepsenia, 3, 31, 32, 33 (fig.).
mascareniensis, Rana, 136.
Rana mascareniensis, 134-136.
mascareniensis venusta, Rana, 1386.
mediaeyus, Ptilodus, 10.
Megalixalus fulvovittatus, 180.
platyceps, 180.
Mexico, subspecies of snake Coniophanes
fissidens from, 103.
Miacidae, 3.
microlestes, Didymictis, 24.
midas, Arachnis, 59, 60, 61, 69.
PROCEEDINGS OF THE NATIONAL MUSEUM
VOL, 91
minganense, Ethmophyllum, 94.
minganensis, Archaeoseyphia, 94, 102.
Archeoscyphia, 94.
Petraia, 94.
minor, Elpidophorus, 13, 14.
minutipora, Patellispongia, 98, 101.
Mioclaenus, 31.
turgidus, 24, 30.
Miodon acanthias, 128.
Mixodectidae, 3, 14, 15 (fig.).
Mizodon variegatus, 120.
modestus, Aparaliactus, 128.
Elapops, 123.
montanus, Deuterogonodon, 19, 20.
Mook, Charles C., on a new fossil eroco-
dilian from Colombia, 55.
Moths, North American, of genus Arach-
nis, 59.
multifragum, Psittacotherium, 17.
Multituberculata, 3, 8.
Mytilus chorus, 498, 501, 512.
nanus, Leptopelis, 128.
nasicornis, Bitis, 124.
Coluber, 124.
natalensis, Phrynobatrachus, 113, 137.
Stenorhynchus, 137.
Natrix anoscopus, 114, 119.
anoscopus anoscopus, 114, 118.
anoscopus gendrii, 119.
fuliginoides, 114, 120.
neivensis, Dinosuchus, 56.
neuquensis, Aegia, 483, 441, 456, 486, 493,
(fig.), 496, 497, 506.
Neusterophis variegatus, 114, 120.
nevadensis, Rhysostrophia, 92.
Nevadocoelia, 94.
grandis, 95, 101.
pulchra, 95, 101.
traini, 95, 101.
wistae, 94, 95, 101, 102.
nigrolineatus, Typhlops, 118.
obeonicus, Streptosolen, 100.
Obelia, 82.
plicata, 82.
recemosa, 82, 89.
occidentalis, Rhysostrophia, 92.
Streptosolen, 100, 102.
occipitalis, Rana, 134, 135.
ocellatus, Hyperolius, 131.
oculata, Patellispongia, 97, 102.
odebrechtii, Aegia, 431-483, 487-441, 448,
445, 448, 455, 457, 486, 487 (fig.),
488-492, 508.
Aeglea, 487.
odebrechtii paulensis, Aegla, 483, 440,
441, 443, 445, 455, 475, 486, 488, 490,
491 (fig.), 492, 498.
ogoensis, Phrynobatrachus, 114, 138.
ogoensis brongersmai, Phrynobatrachus,
138.
oligistus, Anisonchus, 3, 47 (fig.).
Oligocene lizards from Wyoming, 71.
olivaceus, Boaedon, 121.
Holurophis, 121.
onostus, Anisonchus, 3, 46, 47 (fig.), 48.
INDEX
Onychodectes, 16, 17.
rarus, 17.
tisonensis, 16, 17.
opisthacus, Protoselene, 24, 25, 26, 27.
ornatus, Aliurus, 116.
Leiurus, 116.
Liurus, 116.
otariidens, Wortmania, 17.
Oxyacodon, 27, 28.
priscilla, 27, 28.
Oxyclaenid, 3, 22.
Oxyclaenus, 3, 20, 22.
cuspidatus, 21.
pearcei, 3, 20, 21 (fig.), 22.
simplex, 21, 22, 26.
sp., 21.
oxyrhynchus, Rana, 136.
Rana oxyrhynchus, 134, 185, 136.
oxyrhynchus gribinguiensis, Rana, 134,
135, 136.
oxyrhynchus oxyrhynchus, Rana, 134,
135, 136.
Oxytomodon, 27.
perissum, 8, 27 (fig.).
Palaeosinopa senior, 12.
Paleocene of central Utah, the mam-
malian faunas of, with notes on the
geology, 1.
Pantolestid, 3, 12 (fig.).
Pantolestidae, 3.
papudo, Aegla, 482, 488, 441, 455, 483,
484 (fig.), 486, 498, 508, 504, 509.
paracreodus, Desmatoclaenus, 3, 19, 37,
38 (fig.).
parana, Aegla, 432, 433, 440, 441, 450, 452,
458, 459 (fig.), 463, 472, 475, 476.
Parastacus, 444.
parkeriana, Rana albolabris, 113, 186.
Patellispongia, 93, 97, 98.
clintoni, 97, 101.
magnipora, 98, 101.
minutipora, 98, 101.
oculata, 97, 102.
patratus, Elpidophorus, 13, 14.
paucimaculatus, Trimorphodon, 149-152,
154, 1565.
paulensis, Aegla odebrechtii, 433, 440,
441, 443, 445, 455, 475, 486, 488, 490,
491 (fig.), 492, 493.
pearcci, Oxyclaenus, 3, 20, 21 (fig.), 22.
pellucidus, Cambarus, 141, 142, 144.
Cambarus pellucidus, 144.
pellucidus australis, Cambarus (Fax-
onius), 142, 143 (fig.).
pellucidus pellucidus, Cambarus, 144.
pellucidus testii, Cambarus, 144.
Peltosaurus, 75.
pentacus, Protogonodon, 18, 36, 37, 39.
Periaspis plumbeatra, 123.
Peripitychidae, 38.
Periptychus, 42-45.
carinidens, 42—45.
gilmorei, 8, 42, 43 (fig.), 44 (fig.),
45.
perissum, Oxytomodon, 8, 27 (fig.).
pe’ vlexa, Goniotrema, 92.
527
Petraia minganensis, 94.
Phenacodontidae, 3.
Phobosuchus, 56.
Phrynobatrachus alleni, 139.
brongersmai, 114, 138.
francisci, 138.
latifrons, 1389.
liberiensis, 1388.
natalensis, 113, 187.
ogoensis, 114, 138.
ogoensis brongersmai, 138,
plicatus, 138.
picta, Arachnis, 59, 61, 63, 69.
Arachnis picta, 65.
Ecpantheria, 64.
picta citra, Arachnis, 60, 67.
picta hampsoni, Arachnis, 61, 66.
picta insularis, Arachnis, 61, 66.
picta maia, Arachnis, 61, 66.
picta picta, Arachnis, 65.
picta verna, Arachnis, 61, 65.
picturatus, Hyperolius, 131, 133.
platensis, Aegla, 433, 440-442, 447, 452,
464, 465 (fig.), 467, 469, 470, 473.
platyceps, Megalixalus, 180.
Rappia, 130.
pleurotaenia, Rappia, 131.
pleurotaenius, Hyperolius, 131.
plicata, Obelia, 82.
plicatus, Hyperolius, 1388.
Phrynobatrachus, 188.
Pliomerops barrandei, 91.
plumbeatra, Periaspis, 123.
Plumularia, 87.
polynema, 87, 89.
Plumularidae, 86.
poecilonotus, Arthroleptis, 137.
Pogonip sponge fauna, Nevada, 91.
polynema, Plumularia, 87, 89.
pompeia, Arachnis, 68.
Arachnis aulaea, 61, 63.
Porifera, fossil, Nevada Early Ordo4
vician (Pogonip), 91.
prado, Aegla, 482, 433, 441, 442, 450, 451,
453, 469, 470, 471 (figs.), 472, 473, 475,
476, 483.
priscilla, Oxyacodon, 27, 28.
priscus, Ellipsodon, 28, 29, 32.
procyonoides, Claenodon, 23.
proterops, Coniophanes, 105.
Coniophanes fissidens,
107-111.
protogonioides, Protogonodon, 34, 37.
Protogonodon, 3, 4, 17, 19, 34-36.
biatheles, 3, 17, 19 (fig.).
kimbetovius, 19, 39.
pentacus, 18, 36, 37, 39.
protogonioides, 34, 37.
sp., 20.
spiekeri, 3, 17, 18 (fig.), 20.
stenognathus, 37, 39.
Protoselene, 26.
opisthacus, 24-27.
Pseudogalathea, 443.
Pseudoxenopus alleni, 135,
Psittacotherium, 3.
multifragum, 17.
104, 105,
528
Ptilodontidae, 3.
Ptilodus, 10.
ferronensis, 3, 10 (fig.).
mediaeyus, 10.
puercensis, Tetraclaenodon, 34.
pulehra, Nevadocoelia, 95, 101.
punctatus, Typhlops punctatus, 118.
punctigularis, Coniophanes, 107.
Coniophanes fissidens, 107, 109-111.
Purusaurus, 56.
brasiliensis, 55.
quadruplex, Trimorphodon biscutatus,
149-152, 154, 155, 157.
racemosa, Obelia, 82, 89.
ramosa, Lissocoelia, 96, 101, 102.
Rana, Liberian, synopsis of, 133.
Rana acutirostris, 136.
albolabris, 136.
albolabris acutirostris, 113, 136.
albolabris albolabris, 1384, 136.
albolebris parkeriana, 113, 136.
alleni, 114.
bibronii, 184, 135.
erassipes, 114.
erassipes alleni, 134, 135.
longirostris, 118, 138, 135.
maccarthyensis, 134, 136.
mascareniensis, 136.
mascareniensis mascareniensis, 134,
135, 136.
mascareniensis venusta, 136.
occipitalis, 134, 135.
oxyrhynchus, 136.
oxyrhynchus gribinguiensis, 134, 135,
136.
oxyrhynchus oxyrhynchus, 134, 135,
136.
Rappia platyceps, 130.
pleurotaenia, 131.
rarus, Onychodectes, 17.
regularis maculatus, Bufo, 119, 124.
Reptiles, notes on snake genus Trimor-
phodon, 149.
Reptiles and amphibians from Liberia,
Smithsonian - Firestone Expedition,
113.
Rhamnophis aethiopissa, 121.
aethiopissa aethiopissa, 121.
Rhoades, Rendell, on crayfishes from
Alabama caves, 141.
rhombeatus, Causus, 123.
Sepedon, 123.
Rhysostrophia nevadensis, 92.
occidentalis, 92.
riggenbachi, Hyperolius, 130.
riolimayana, Aegla, 4388, 441, 447, 449,
450, 456, 457, 506, 512, 5138, 514 (fig.),
515.
sanlorenzo, Aegla, 432, 441, 446, 450-452,
460, 461, 462 (fig.), 464, 472, 475.
savatieri, Agama, 116.
Agama agama, 114, 116.
PROCEEDINGS OF THE NATIONAL MUSEUM
VOL, 91
Schmitt, Waido L., on the species of
Aegla, 431.
sectorius, Anisonchus, 45, 46.
senegalensis, Kassina, 125.
senior, Palaeosinopa, 12.
Sepedon rhombeatus, 123.
serratus, Exostinus, 74, 75 (fig.).
shepherdi, Ellipsodon, 3, 28, 29 (fig.),
31-33.
shrevei, Crotaphopeltis, 122.
Silicispongiae, 93.
Silurana tropicalis, 124.
simplex, Oxyclaenus, 21, 22, 26.
Simpsoni, Aphronorus, 3, 11 (fig.), 12, 13.
simulans, Cophoscineus, 117.
Smith, Hobart M., on Mexican subspecies
of the snake Coniophanes fissi-
dens, 103.
on snake genus Trimorphodon, 149.
Smithsonian-Firestone Expedition to Li-
beria, reptiles and amphibians, 113.
smythii, Coluber, 121.
Grayia, 121.
Snakes, Mexican subspecies of Conio-
phanes fissidens, 103.
notes on genus Trimorphodon, 149.
South America, the species of Aegla
from, 431.
ApIcEGEs Protogonodon ?, 3,17, 18 (fig.),
Sponges, fossil, from Nevada
Ordovician (Pogonip), 91.
stenognathus, Protogonodon, 387, 39.
Stenorhynehus natalensis, 187.
sternbergi, Eliipsodon, 3, 30 (fig.), 33.
Streptosolen, 100.
obconieus, 100.
occidentalis, 100, 102.
Stylinodon, 17.
Stylinodont, 17.
Stylinodontidae, 3.
subtrigonus, Tricentes, Bo:
symbolus, Ectoconus, 3, 39, 40 (fig.).
Symplectanea, 78.
bracteata, 78, 89.
Symplectaneidae, 78.
Early
99
tay
Taeniodonta, 3, 15.
Taeniolabididae, 3.
Taeniolabis, 3, 4, 9.
SDspo omy):
tadensis, 8.
talcahuano, Aegla laevis, 4338, 486, 441,
447, 456, 457, 507, 508, 509 (fig.).
tadensis, Taeniolabis, 8.
tau, Trimorphodon, 149, 151-153, 155,
163-166.
telescopium, Halecium, 84.
tensum, Halecium, 84, 89.
terror, Dinosuchus, 55, 56.
tessmanni, Leptopelis, 128, 129.
testii, Cambarus pellucidus, 144.
Testudo denticulata, 115.
erosa, 115.
Tetraclaenodon, 34-389.
puercensis, 34.
INDEX
Tetractinellida, 93.
thomsoni, Bessoecetor, 12.
Tiliqua dura, 117.
tisonensis, Onychodectes, 16, 17.
Toads and frogs, Liberian, 124.
togoensis, Leptopelis, 128.
Toquimia kirki, 92.
Tortoises, Liberian, 115.
traini, Anthaspidella, 100, 102.
Nevadocoelia, 95, 101.
transitionis, Aglaophenia, 87, 89.
triangularis, Coronella, 121.
Tricentes, 22.
crassicolidens, 22.
elassus, 3, 22, 23 (fig.).
subtrigonus, 22, 23.
Trimorphodon, notes on the genus, 149.
Trimorphodon biscutata latifascia, 160.
biscutatus, 149-155, 158, 159, 165.
bisecutatus biscutatus, 149, 150, 153,
154, 159.
biscutatus quadruplew, 149-152, 154,
155, 157.
collaris, 149, 151-154, 163, 165.
fasciolata, 149, 151, 152, 154, 160,
162, 163.
forbesi, 149, 151, 153, 154, 168.
lambda, 149-151, 153-157.
latifascia, 149, 151, 152, 154, 160,
162, 165, 166.
lyrophanes, 149-151, 154-157.
major, 159.
paucimaculatus, 149-152, 154, 155.
tau, 149, 151-158, 155, 168-166.
upsilon, 149, 151-155, 161, 162, 164—
168
vandenburghi, 149-151, 153-157, 165.
vilkinsonii, 149, 151-154, 163, 167.
tropicalis, Silurana, 124.
Xenopus, 124.
Tropidolepis africanus, 116.
Tropidonotus anoscopus, 118.
ferox, 114, 118.
Tubularia, 81.
crassa, 81, 89.
Tubularidae, 80.
turgidus, Mioclaenus, 24, 30.
typa, Aporthophyla, 92.
{
%
529
Typhlops intermedia, 118.
leucostictus, 118.
liberiensis, 118.
manni, 118, 118.
nigrolineatus, 118.
punctatus punctatus, 118.
typicalis, Calycocoelia, 96, 97, 101, 102.
Hesperocoelia, 98, 102.
undulata, Hesperocoelia, 99, 102.
unicolor, Dipsadoboa, 122.
upsilon, Trimorphodon, 149,
161, 162, 164-168.
Urobelus acanthias, 123.
uruguayand, Aegla, 432, 440-442, 453,
463, 464, 466, 467, 468 (fig.), 472, 473.
utahensis, Catopsalis, 3, 8, 9 (fig.).
uvularis, Lampra, 77, 80, 89.
151-155,
valens, Hydractinia, 79, 89.
valida, Dipsas, 122.
vandenburghi, Trimerphodon, 149-151,
153-157, 165.
variegatus, Mizodon, 120.
Neusterophis, 114, 120.
venusta, Rana mascareniensis, 136,
verna, Arachnis picta, 61, 65.
versicolor, Calotes, 116.
vilkinsonii, Trimorphodon, 149, 151-154,
163, 167.
virgata, Coelopeltis, 120.
virgatus, Boaedon, 114.
viridis, Dendroaspis, 123.
Hylambates, 128.
Leptopelis, 114, 128.
Leptophis, 128.
vittiger, Hyperolius, 130.
weali, Cassina, 125.
werneri, Arthroleptis, 137.
wistae, Nevadocoelia, 94, 95, 101, 102.
Wortmania otariidens, 17.
Wyoming, fossil lizards from Oligocene
of, 71.
Xenopus tropicalis, 124.
zuni, Arachnis, 60, 61, 6¥.
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