SMITHSONIAN INSTITUTION
UNITED STATES NATIONAL MUSEUM

PROCEEDINGS

OF THE

UNITED STATES NATIONAL MUSEUM

VOLUME 119
NUMBERS 3538-3553

SMITHSONIAN PRESS
WASHINGTON : 1967

Publications of the United States National Museum

The scientific publications of the United States National Museum include two
series, Proceedings of the United States National Museum and United States
National Museum Bulletins.

In these series are published original articles and monographs dealing with
the collections and work of the Museum and setting forth newly acquired facts
in the fields of anthropology, biology, geology, history, and technology. Copies
of each publication are distributed to libraries and scientific organizations and
to specialists and others interested in the various subjects.

The Proceedings, begun in 1878, are intended for the publication, in separate
form, of shorter papers. These are gathered in volumes, octavo in size, with
the publication date of each paper recorded in the table of contents of the volume.

In the Bulletin series, the first of which was issued in 1875, appear longer,
separate publications consisting of monographs (occasionally in several parts)
and volumes in which are collected works on related subjects. Bulletins are
either octavo or quarto in size, depending on the needs of the presentation.
Since 1902, papers relating to the botanical collections of the Museum have been
published in the Bulletin series under the heading Contributions from the United
States National Herbarium.

Frank A. TaYLor
Director, United States National Museum
Il

CONTENTS

Barnarp, J. Laurens. Benthic Amphipoda of Monterey
Bay, California. 41 pages, 7 figures. Published December
30, 1966.

New species: Protomedeia penates, Centromedon pavor.
New combinations: Lysianassa holmesi, L. oculata, Orchomene deci-
prens.

Capps, Hann W. Review of New World moths of genus Hu-
chromius Guenee with descriptions of two new species (Lepi-
doptera: Crambidae). 9 pages, 8 figures, 1 plate. Pub-
lished December 30, 1966

New species: Huchromius galapagosalis, E. saltalis.

CotuETre, Bruce B., and Knapp, Lesuie W. Catalog of type
specimens of the darters (Pisces, Percidae, Etheostomatini).
88 pages, 5 figures. Published December 30, 1966

Duckxwortu, W. Donatp. Neotropical Microlepidoptera, X:
Systematic position of two taxa erroneously placed in family
Stenomidae (Lepidoptera). 6 pages, 2 figures, 1 plate.
Published December 30, 1966

New genus: Mattea.
New combination: Mattea phoenissa.

Fieip, Winuiam D. Preliminary revision of butterflies of the
genus Calycopis Scudder (Lycaenidae: Theclinae). 48 pages,
34 figures, 6 plates. Published February 1, 1967 .

New species: Calycopis nicolayi, C. lerbela, C. susanna, C. drusilla,
C. anastasia, C. fractunda.

New combinations: Calycopis bactra, C. caulonia, C. janeirica, C.
chacona, C. torqueor, C. xenata, C. isobeon, C. vibulena, C. vitruvia,
C. bellera, C. partunda, C amplia, C. indigo, C. anfracta.

GitBERT, CarTER R. A revision of the hammerhead sharks
(Family Sphyrnidae). 88 pages, 22 figures, 10 Paial Pub-
lished April 27, 1967 : CN aca

Hosss, Horton H., Jr., and Wineow ipeaeieee my new
genus and six see species of entocytherid ostracods (Ostra-
coda, Entocytheridae). 12 pages, 2 figures. Published De-
cember 30, 1966 BE ee ae nicer tae, Xe 5

New genus: Thermastrocythere.

New species: Dactylocythere brachystriz, D. pachysphyrata, Donnald-
soncythere cayugaensis, Entocythere kanawhaensis, Thermastro-
cythere hartt, Uncinocythere stubbsi.

Number

3551

3550

3540

3552

3539

3542

IV CONTENTS

Hopaes, Ronautp W. Revision of Nearctic Gelechiidae, I:
The Lita group (Lepidoptera: Gelechioidea). 66 pages, 246
figures on 31 plates. Published December 30, 1966 .

New genera: Rifseria, Sriferia.

New species: Lita nefrens, L. deoia, L. incicur, L. geniata, L. dialis,
L. pagella, L. thaliae, L. maenadis, L. obnubila, L. veledae, L. recens,
L. sironae, L. jubata, Arla diversella, Neodactylota liguritrizx, N.
basilica, N. egena, Eudactylota diadota, E. abstemia, Friseria
cateta, F. nona, F. caieta.

New combinations: Arla diversella, Eudactylota iobapta, Friseria lac-
ticaput, F. acaciella, F. infracta, Rifseria fuscotaeniaella, Sriferia
prorepta.

Hower, Rottanp O. The freeze-dry preservation of bio-
logical specimens. 24 pages, 9 figures, 4 plates. Published
March 15, 1967

JAMES, mae A. Range nad carintone a ener BE Cane
barus longulus (Decapoda: Astacidae). 24 pages, 2 nee
1 plate. Published December 30, 1966.

New subspecies: Cambarus longulus chasmodactylus.

Kormitev, Nicuotas A. Notes on Aradidae in the U.S. Na-
tional Museum, IV (Hemiptera: Heteroptera). 25 pages,
23 figures. Published December 30, 1966 .

New species: Aradus saileri, A. ovatus, A. barberi, Aneurus pygmaeus,
Anasutus, Proxius (Neoproxius) peruvianus, Kolpodaptera minuta,
K. rugosa, Miorrhynchus angulatus, M. undulatus, Placogenis
clarket, Notapictinus uruguayensis, N. platyceps, N. ornatus.

Osraztsov, Nicuotas 8. Neotropical Microlepidoptera, XI:
Revision of genus Jdolatteria (Lepidoptera: Tortricidae).
12 pages, 3 figures, 8 plates. Published December 30, 1966 .

New species: Jdolattera fasciata, I. mydros, I. cantharopisca.

Prerrers, James A. The lizards of Ecuador, a check list and
key. 49 pages, 5 figures. Published March 8, 1967 .
PrrtTiponE, Martian H. Type-specimens of Polychaetes de-
scribed by Edith and Cyril Berkeley (1923-1964). 23 pages.
Published February 1, 1967
Topp, E. L. Noctuid set of the eeeeeee eae ee
baer 22 pages, 34 figures on 6 plates, 1 map. Pub-
lished December 30, 1966
New species: Eusceptis atriora, E. koehleri, E. lelae, E. robertae.
New combinations: Husceptis obscura, E. effusa, E. splendens, E.
extensa, E. paraguayensis, Tarachidia bruchi.
New status: Husceptis paraguayensis.
WeItTzMAN, Stantey H. Review of South American characid
fishes of subtribe Nannostomina. 56 pages, 12 figures.
Published December 30, 1967

3547

3549

3544

3548

3543

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3553

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3538

Proceedings of

the United States

National Museum
SMITHSONIAN INSTITUTION + WASHINGTON, D.C.

Volume 119 1966 Number 3538

REVIEW OF SOUTH AMERICAN CHARACID FISHES
OF SUBTRIBE NANNOSTOMINA !

By Stantey H. WEITZMAN
Associate Curator, Division of Fishes

Introduction

Fishes of the subtribe Nannostomina, family Characidae, form a
clearly definable group of about nine small South American freshwater
fishes. No member is known over 44.5 mm. in standard length.
They appear restricted in habitat to shaded forest brooks, seepages,
and ponds, and are apparently confined to the Guianas, Rio Orinoco
in Venezuela, and the Amazon Basin of Brazil, Peru, Colombia, and
Bolivia. All of the known species have been imported into Europe
or North America as aquarium fishes. In the aquarium trade they
are known as pencil fishes.

The osteology and relationships of these fishes have been treated
elsewhere (Weitzman 1964), and their intercharacid relationships
will be but briefly mentioned here. They are members of the characid
subfamily Lebiasininae, which consists of two tribes, the Lebiasinini
and the Pyrrhulinini. The Pyrrhulinini comprises two subtribes, the
Pyrrhulinina and the Nannostomina. The Nannostomina appears to
be the most specialized group within the Lebiasininae.

1 This paper is the third and final of three parts based on a Ph. D. dissertation
submitted at Stanford University, Calif. See Weitzman (1962, 1964) for the
two preceding parts.

1

2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

The Lebiasinini and Pyrrhulinina are small to moderate-sized,
predaceous characids with moderate-sized mouths. The tiny mouths
of members of the Nannostomina, however, are adapted for feeding
on very small organisms. They apparently feed primarily on animals
that occur on plants, rocks, and other objects and also to some extent
on slowly moving, free-swimming organisms.

Wickler (1957) has described the breeding behavior of certain of
these fishes in a preliminary fashion, and there is a very large amount
of literature on this subject published by aquarists. However, even
though they are excellent aquarium subjects, no detailed behavioral
study has been published about them. A breeding pair enters a group
of fine-leaved plants or a clump of roots, the male presses its body
against that of the female, curves its anal fin around the female’s
vent, and the female sheds one to a very few eggs. ‘The structure of
the male’s anal fin in several species is modified for aiding fertilization
by guiding sperm over the female’s vent. At this time the male
sheds an undetermined number of sperm and fertilizes the eggs. The
eggs are very slightly adhesive and may become attached to plants or
fall to the bottom. All the species that I have seen alive (all but
Nannostomus digrammus and Nannostomus bifasciatus) lay eggs of
about 0.6 to 0.7 mm. The young mature in 8 to 10 weeks and speci-
mens may live for as long as 4 years. Their lifespan in aquaria is
usually 2 or 3 years.

Since most of Nannostomina can be bred with relative ease, they
would make fine subjects for comparative ethological studies. It
would be interesting to compare the taxonomic implications of such
studies with those derived from the morphological investigation
presented here.

The spotty locality records of the Nannostomina, the frequent
damaged condition of the specimens, and the paucity of specimens
from many localities have precluded satisfactory use of statistical
procedures in analyzing geographical population differences or in
recording changes in body proportions correlated with growth. Col-
lectors frequently have packed these small, delicate fishes with
specimens of other, larger species with the result that the nannosto-
minans have been squashed out of shape, making valid measurements
impossible. Only specimens in reasonably good condition were
measured. For this reason, counts should be considered more re-
liable than measurements in the data presented below. The data
available indicate in several instances that many geographically
distinct populations of these fishes probably exist. Some of these
differences are pointed out in the discussion under each species. It
would be foolish to designate these different population samples as
subspecies in view of the inadequate number of specimens and data

NO. 3538 CHARACID FISHES—WEITZMAN 3

at hand. The primary function served by this paper is to point out
our present state of knowledge concerning these fishes and to indicate
problems for future study.

Counts and measurements were taken from the left side of each
specimen. In the descriptions the initial values are the arithmetical
means, and the values included in parentheses are the extremes of all
specimens cited in the material examined. Measurements were taken
from adult specimens only.

The following straight-line measurements were made from the
anteriormost part of the fleshy tip of the upper jaw to some point
posterior on the body surface. Standard length (SL) was measured
to the posterior end of the hypural fan. Predorsal length was meas-
ured to the anterior base of the first dorsal fin ray. Preanal length
was taken to the anterior base of the most anterior, visible anal fin
ray. Head length was measured to the most posterior border of the
opercular bone, not the fleshy flap which was often damaged.

Hye length is the greatest distance between the posterior border of
the first infraorbital and the anterior border of the fifth infraorbital
bone. Depth was measured from the anterior dorsal fin base ver-
tically to the median profile of the belly. Least depth of caudal
peduncle was measured vertically. Length of caudal peduncle was
measured from the posterior base of the last anal fin ray to the posterior
end of the hypural fan. Interorbital width is the least width between
the lateral supraorbital borders of the frontal bones.

Vertebral counts were taken from radiographs and include the
vertebrae forming the pars sustentaculum of the Weberian apparatus
and the ultimate vertebra with its urostyle.

Fin counts: The ultimate (posterior) ray of the dorsal and anal
fins was counted as one when one ray was associated with the last
pterygiophore and as two when there were two entirely separate rays
associated with the last pterygiophore. Counts were least variable
when this method was employed. All anterior rudiments of fin rays
were counted. Unbranched rays (except the most posterior ray of
the dorsal, anal, pectoral, or pelvic fins) are designated by lowercase
roman and branched rays by Arabic numerals. The principal caudal
fin ray count includes all rays associated with the hypural elements
and may be taken without recourse to examining the hypural fan by
counting all branched rays and adding two. The count of the upper
lobe is given first, followed by a bar (/) and then the count of the rays
of the lower caudal lobe.

Gill-raker counts in these small fishes can be made most accurately
on alizarin-stained specimens and all such counts were confined to
such specimens. The counts are of the bony supports of gill rakers.
Scale counts in a lateral series refer to all scales in a median lateral

4. PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

series extending to the posterior midbase of the hypuralfan. Vertical
scale rows are counted between the anterior parts of the dorsal and
anal fin bases.

Although the Nannostomina as a whole are easily definable and
the species readily separable, their segregation into generic groups
having a clear phylogenetic basis has not proven easy. However,
some indication of their possible phylogenetic history is provided by
their morphology, and morphological definitions of two generic groups
is possible. Of the characters that seem to have generic and specific
significance, the following appear to be most important.

Anal fin: The anal fin of males of this subtribe is used as an accessory
sexual organ to guide the sperm toward the female’s vent. During
the spawning act, the anterior, posterior, and distal parts of the fin
are cupped to form a bowl that partially covers the female’s vent,
apparently directing the sperm toward the eggs as they are laid.
Correlated with this function is a previously overlooked morphological
fact. The individual anal fin rays of the males of several species are
widened in the sagittal plane (figs. 4 and 6). This modification is also
found to a certain extent in some members of the subtribe Pyrrhu-
linina, the closest living relatives of the Nannostomina. This feature
seems to have more significance at the specific than at the generic level.

Infraorbital bones: The usefulness of these structures also has
been overlooked by previous authors. There are two types of infra-
orbital bone arrangement in the subtribe. In one they are more
elongate, and both the first and second elements have a bony infra-
orbital canal. In the other type, the first two infraorbitals are
short and the second is without a bony canal. The closest relatives
of the Nannostomina, the Pyrrhulinina, have a canal in both the first
and second infraorbital bones; its absence in the second infraorbital
of some Nannostomina is possibly a neotenic specialization. This
character is considered here to have generic significance because it
is consistent and correlated with a definite difference in snout and
head shape.

Color: The melanophore patterns occurring in this subtribe can
be separated conveniently into three categories.

The first of these consists of long, dark, horizontal stripes present
in almost all species. The term stripe is restricted in this account to
the elongate pigmented areas that extend in a horizontal direction
on the sides of these fishes. There are several of these stripes as
follows: Primary stripe, the main midside stripe usually extending
from the snout to the eye, across the operculum to the lower part of
the caudal peduncle and onto the caudal fin. Secondary stripe, the
dark stripe above the primary stripe. It usually extends from the
nape or the top of the head to the upper portion of the caudal peduncle.

NO. 3538 CHARACID FISHES—WEITZMAN 5

It often blends with the darkly pigmented back. Tertiary stripe,
the lowermost stripe often extending from the lower jaw, across the
lower part of the operculum, below the base of the pectoral fin to the
origin of the anal fin. Additional stripes occur on some species,
such as Poecilobrycon eques. Live specimens suddenly illuminated
at night may show the stripes only faintly oralmost not at all. During
the day these same stripes are darkly pigmented.

The second category consists of oblique bands. The anterior of
these lies along the sides a short distance anterior to the dorsal fin,
while the posterior lies along the midside a short distance posterior
to the dorsal fin. These oblique bands are usually absent or extremely
pale in living specimens during the daylight hours; however, in the
absence of light the oblique bands become very dark. Females of
Nannostomus beckfordi while spawning often will show faintly the
oblique bands. The oblique bands appear in some species, such as
N. beckfordi, to be mostly areas of the primary horizontal stripe which
fail to become pale under the influence of darkness. However, even
in this species these bands are a little more than this because some
of the area of the oblique band occurs above the primary horizontal
stripe. In such species as Poecilobrycon eques, much of the area of
the oblique bands is above the primary horizontal stripe.

The oblique bands have been called ‘‘night paint” by Hoedeman
(1950) because they are usually present only at night. Specimens
of N. beckfordi preserved at night in ten percent formalin retain their
‘night pigment”’ pattern. In these specimens the oblique bands are
quite dark and their horizontal stripes are faded and sometimes
indistinct.

In specimens preserved in daylight, oblique bands are present often
but are usually very pale. Horizontal stripes are almost always
present and distinct except on faded specimens or specimens pre-
served at night. Care should be used in determining the presence
or absence of oblique bands from preserved specimens because if, as
is usually the case, the specimens were preserved during daylight
hours, the pigment of the melanophores of the oblique bands may
be so contracted that the bands do not show.

Permanent blotches constitute the third category. These are pres-
ent in only one species, Nannostomus espet, and differ in position and
quality from all other dark markings seen in the other species. They
are permanent, not disappearing in daylight or darkness, although as
with other pigment, they may fade slightly at night. The borders
of the blotches are darker than the rest of the blotch. This is not
true of the oblique bands. There are five such permanent blotches
along the midsides on N. espei, the first above the pectoral fin, the
second below the origin of the dorsal fin, the third midway between

6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

the origin of the pelvic fin and the anal fin, the fourth above the
anal fin origin, and the fifth in the lower half of the caudal fin root.

Caudal fin rays and body swimming position: The normal principal
caudal ray count is 10/9 in characids, and this is true of all Nanno-
stomina examined by me. However, in two species, two of the rays
normally in the upper lobe of the caudal fin have their distal ends
in the lower lobe. The proximal ends of these two rays retain the
normal position on the hypural fan and thus originate in the upper
lobe. The two species having this arrangement swim at an inclined
angle, head up. Steindachner (1876) first noticed that the lower
caudal fin lobe is larger in some Nannostomina, and Hoedeman (1950)
noted that these unequal lobes are correlated with the swimming
position. Hoedeman did not describe accurately the morphological
arrangement of the caudal fin rays. He stated that there is a differ-
ence in the shape of the swim bladder correlated with the swimming
position. This is possibly true, but investigation of this feature has
been postponed pending a histological examination of the swim bladder
and its musculature. A comparative histological examination of the
semicircular canals and associated structures probably would show
also interesting morphological differences associated with the oblique
swimming position. Braemer and Braemer (1958) have begun to
investigate some of the morphological differences in the arrangement
of statoliths in characids, including that in Poecilobrycon eques, one
of the oblique swimmers.

Other characters have been reported to have generic significance
in Nannostomina. For example, EKigenmann (1909) based Poecilo-
brycon mainly on the presence of an adipose fin in the type species,
P. harrisoni. However, additional specimens have shown that
members of this species may or may not have an adipose fin, and the
character is of no generic significance in this group. Steindachner
(1876) noted that the adipose fin is variably present in Poecilobrycon
eques. Hoedeman (1950) attempted to use the shape of the teeth
as a generic character, but their shape often varies from individual to
individual as well as among different geographical populations of the
same species.

The characters most useful in separating species are color pattern,
scale counts, gill-raker counts, numbers of vertebrae and teeth, and
body proportions. Preliminary observations of living specimens in
aquaria indicate that behavior also may be of considerable aid in
understanding specific and generic relationships.

The synonymy used here is selected, and no attempt has been made
to include all references to these fishes that have appeared in the

NO. 3538 CHARACID FISHES—WEITZMAN 7

very extensive aquarium literature. Many references to the aquarium
literature concerning these fishes can be obtained from Hoedeman
(1950).

The following abbreviations are used:

AM Zoological Museum, Amsterdam
ANSP Academy of Natural Sciences, Philadelphia
CAS California Academy of Sciences

CAS (LUM) Specimens formerly at the Indiana University Museum and now
at the California Academy of Sciences but still bearing Indiana
University Museum numbers

CNHM Chicago Natural History Museum
MCZ Museum of Comparative Zoology, Harvard
USNM United States National Museum

For loan of specimens I am indebted to Dr. George S. Myers of
Stanford University, W. I. Follett of the California Academy of
Sciences, Loren P. Woods of the Chicago Natural History Museum,
Dr. Giles Mead of the Museum of Comparative Zoology, Harvard
University and Dr. James Bohlke of the Academy of Natural Sciences
of Philadelphia. I am indebted greatly to Dr. George S. Myers for
numerous discussions and for critically reading the original manu-
script. In addition the following people have read the manuscript
at various stages, all offering useful advice and aid: Drs. Paul Erlich,
Myra Keen and Warren Freihofer of Stanford University, Mrs.
Lillian Dempster of the California Academy of Sciences, and Drs.
Robert H. Gibbs, Jr., and Victor G. Springer of the U.S. National
Museum.

I am indebted also to Gen. Thomas D. White for forwarding living
specimens of Nannostomus marginatus from eastern Colombia so that
I could record their life colors. All drawings are by the author with
the exception of figure 6, the anal fin of a male Nannostomus digrammus
prepared by Dr. Margaret Bradbury.

The work was begun and largely completed at the Department of
Biological Sciences, Stanford University. Additional work was done
at the Department of Anatomy, Stanford University, and the U.S.
National Museum, Smithsonian Institution.

Note on figures: Figures 1, 2, 4, 5, 7, 8, 9, 11, and 12 show male
specimens with the antorbital and first three orbital bones and the
third set (counting from the anterior end of the jaw base) of upper
(premaxillary) and lower (maxillary) teeth. The enlarged scale (or
scales) is from the third longitudinal scale row of the left side, just
below the dorsal fin except where noted in the text. These parts are
all drawn from alizarin specimens.

PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Key to Genera and Species of Subtribe Nannostomina

Second infraorbital bone without bony canal passing in its substance.
Genus Nannostomus. 2
Second infraorbital bone with well-developed bony infraorbital canal passing
in its substance .. . . . . . « Genus Poecilobrycon. 7
Very indistinct primary horiaoutall siting present. Secondary or tertiary
horizontal stripes absent. Five dark permanent blotches along sides (in
life these blotches not fading during exposure of fish to daylight).
Nannostomus espei (p. 9)
Primary horizontal stripe present, very well developed. Some species with
secondary and tertiary horizontal stripes present. One or two pale oblique
bands on sides (in life these bands ordinarily present ail when fish kept
in dark). Permanent blotches absent .... . Ree del)
Primary horizontal stripe present, secondary stripe present or absent, tertiary
horizontal stripe absent. Teeth in single row on premaxillary, 5 to d
(usually 6). Anal fin rays of males slightly to greatly modified, flattene6
in sagittal plane. .. . Sy iiee ets eee
Primary, secondary, and tactiany horiontal SeuBes a aeecapelit Teeth
in one row on premaxillary, 7 to 8 (usually 8). Anal fin rays of males
completely unmodified in appearance .. . The OO
Perforated scales in lateral line 2 to 4. Adipose fin absent. Saal fin rays
iii,9. Anal fin rays of males relatively little modified, not greatly flattened
and expanded in*sagittalplame 7. ceictis ete tomes fe) ete: ore ap te eee
No perforated lateral line scales. Adipose fin present. Anal fin rays iii,8.
Anal fin rays of males greatly modified, flattened, and expanded in sagittal
planes vies . . Nannostomus digrammus (p. 22)
Secondary Horizontal nine eben Gil rakers 9 to 10+17 to 18.
Nannostomus bifasciatus (p. 19)
Secondary horizontal stripe present. Gill rakers 8+14.
Nannostomus beckfordi (p. 12)
Scales 26 to 27 in lateral series. Total vertebrae (including those of Weberian
apparatus) 35 to 36. Adipose fin present, either well developed or rudi-
mentary. About 11 teeth in second tooth row of dentary. Snout in eye
1.0. Least depth of caudal peduncle in standard length 10.0 to 10.4.
Nannostomus trifasciatus (p. 29)
Scales 21 to 23 in lateral series. Total vertebrae 31 to 33. Adipose fin
absent. Teeth in the second row of dentary 6 to 7. Snout in eye 1.3 to
1.4. Least depth of caudal peduncle in standard length 7.8 to 8.5.
Nannostomus marginatus (p. 32)
Principal caudal fin rays 10/9, their distribution in lower and upper caudal
lobes normal, 10 ending in upper lobe, 9 ending in lower lobe. Total
vertebrae 38 to 39. Length of caudal peduncle in standard length 5.3
to 6.1 . . . (Subgenus Poecilobrycon). Poecilobrycon harrisoni (p. 38)
Principal caudal fin rays 10/9, but 8 rays end in upper lobe and 11 in lower
lobe of caudal fin. Total vertebrae 33 to 34. Length of caudal peduncle
in standard length 6.0 to 7.5... . . . . (Subgenus Nannobrycon). 8
Scales 28 to 30 in lateral series. Perforated scales in lateral line 2 to 5.
Adipose fin present. Gill rakers 9+14. Inner tooth row of dentary
absent; teeth in outer dentary tooth row 6 to 7.
Poecilobrycon unifasciatus (p. 42)

NO. 3538 CHARACID FISHES—WEITZMAN 9

Scales 24 to 25 in lateral series. No perforated lateral line scales. Adipose
fin present or absent. Gill rakers 164-24. Teeth in inner dentary row 12;

?

teeth in outer dentaryrow9 .... . . . Poecilobrycon eques (p. 47)

Genus Nannostomus Giinther

Nannostomus Giinther, 1872, p. 146 (type species: Nannostomus beckfordi Giinther,
1872, by monotypy).

DiaGnosts.—Among the Lebiasininae the following character ap-
pears unique for this genus: Second infraorbital bone without bony
canal for infraorbital branch of laterosensory canal; in addition, snout
short, 1.0 to 1.4 in eye diameter. Correlated with the short snout,
Nannostomus has a shorter, broader ethmoid and vomer, a shorter
antorbital and infraorbital, and a shorter mesopterygoid and nasal
bone than Poecilobrycon. An adipose fin may be either present or
absent. Hither the primary or tertiary, or both primary and tertiary
horizontal stripes may be present or absent. The anal fin rays of the
males may be greatly modified or scarcely modified as an accessory
organ. The body is short to fairly elongate, the depth being 3.5 to 5.0
in the standard length.

The name Nannostomus is derived from the Greek vavoc, meaning
little or dwarf, and crowa, meaning mouth.

Nannostomus espei (Meinken)
FIGurE 1

Poecilobrycon espet Meinken, 1956, p. 31 (original description; no type locality;
types in collection of Meinken and at Stanford University, lectotype SU
51593).

Ties espei—Bohlke, 1956, p. 2 (description; unnamed creek emptying
into the Paruma River, a tributary of the Kamarang River, which empties
into the Mazaruni River, British Guiana).—Sterba and Tucker, 1962, p. 213
(description and aquarium notes).

DiaGnosis.—Secondary and tertiary horizontal stripes absent; pri-
mary horizontal stripe poorly developed; permanent blotches on sides
of living specimens (blotches do not disappear during daytime);
scales (fig. 1) very much in outline like scales of members of subtribe
Pyrrhulinina (see description on page 12). Nannostomus espei differs
from all members of subtribe Nannostomina except Nannostomus
marginatus by having fewer than 23 scales in lateral series.

Description.—Body elongate, sides flattened, and body compressed
posteriorly. Greatest depth between posterior tip of appressed pec-
torals and anterior part of base of dorsal fin. Standard length of
largest specimen examined, 28.1 mm. Greatest body depth 4.3 in all
specimens; least depth of caudal peduncle 9.6 (9.4 to 10.0); length of
caudal peduncle of both sexes 6.4 (6.1 to 7.0); snout tip to origin of
dorsal 1.9 (1.9 to 2.0); snout tip to origin of anal 1.3 in all specimens.

10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Head elongate, snout obtuse in vertical and horizontal profiles.
Top of head very slightly convex between eyes. Head 3.6 (3.5 to 3.7);
eye in head 3.0 (2.9 to 3.0); snout in eye about 1.0 in all specimens
examined; least width of bony interorbital in greatest eye diameter
1.0 or very slightly less in all specimens examined.

Premaxillary with 7 teeth. Each tooth except posterior 1 or 2
with 5 cusps. Posterior 2 teeth usually quadricuspid. Sometimes
most anteromedial premaxillary tooth with only 4 cusps. Usually with
largest cusp just medial to farthest posterolateral cusp. Maxillary
with single, uni- to quadricuspid tooth. Teeth in anterior row of
each dentary 7 to 8. Anterior 5 or 6 teeth hexacuspid, third from
anteromedial cusp largest. Posteriormost dentary tooth usually tri-
cuspid. Second dentary tooth row with 8 to 11 unicuspid teeth. Gill
rakers 10+17 on each first gill arch in one alizarin preparation,
SU 50252.

Dorsal fin rays ii,8; anal iii,9; pectoral 1,10; pelvics ui,7, in all
specimens examined. Adipose fin present, well developed in all
specimens. Caudal fin with principal rays 10/9; distal tips of all
ray elements of first (upper) 10 principal rays within upper caudal
lobe; remaining 9 principal rays end in lower lobe.

Male’s anal fin modified (fig. 1); rays somewhat expanded and
more similar to highly modified anal rays of Nannostomus digrammus
than any other species of Nannostomina.

Radial grooves of scales in third horizontal scale series below
dorsal fin shown in figure 1. Posterior field of each scale with 2 to 3
radial grooves; dorsal and ventral fields each with 1 radial groove;
anterior field with 2 to 4 radial grooves. Dorsal and ventral anterior
corners of scales in this species more acute than in any other species
of Nannostomina. Acute corners very similar to those found in sub-
tribe Pyrrhulinina. Scales in lateral series 22 in all specimens ex-
amined. Perforated lateral-line scales on each side 1 or 2. Scales in
median predorsal series 10 in all specimens examined. Total number
of vertebrae 33 (18 precaudals and 15 caudals) in all specimens.

CoLor IN ALconou.—Preserved specimens (SU 50214) have the
following color: Top of head light to dark brown. Primary horizontal
stripe from tip of snout across maxillary and first obrital bones onto
anterior part of iris of eye; then from posterior part of iris to edge of
opercular bone. Posterior to eye, primary horizontal stripe somewhat
diffuse and hard to distinguish. Primary horizontal stripe behind
operculum represented as very diffuse, slightly darkened area ex-
tending posteriorly to caudal peduncle, through third and fourth
longitudinal scale rows. (In live specimens kept in an aquarium with
dark brown peatmoss bottom and subdued light, horizontal stripe
became much darker but remained diffuse.) Large permanent blotches

No. 3638 CHARACID FISHES—WEITZMAN i

5 on sides with slightly irregular margins as shown in figure 1. Borders
of blotches (very different from oblique bands described in other
species) slightly darker than their centers. Very pale stripe above
blotches and above very light and diffuse primary horizontal stripe.
Dorsum brown above pale stripe. Network of dark lines that follow
scale edges superimposed on entire body pigment pattern. Belly
white, all fins hyaline.

Cotor IN LirE.—Life colors of an aquarium specimen as follows:
Five large permanent blotches dark brown to almost black. Body
color very light brown with slight greenish cast. This ground color
darker on back. Abdominal region silvery. Light green, somewhat
iridescent stripe extending along sides just above dorsal borders of
permanent blotches. This stripe reflecting pale gold. Fins translucent
except for few dark spots on anal fin.

Ficure 1.—Nannostomus espei, male, SL 27.6 mm., SU 50214 (see text for discussion
of scales).

Remarxs.—The original description of this species was inadequate,
and no types were designated. Also because the type locality was
unknown, none was cited. Bohlke (1956) has located and published
data on the collecting site of the original aquarium importation of
this species, but did not designate a restricted locality—this is done
here. The ‘‘Testricted locality” is that given by Bohlke (1956, p. 1).
This locality, along the Paruma River, is apparently in the vicinity of
lat. 5°50’ to 55’ N., long. 61°05’ to 10’ W. At my request, Herr
Meinken has sent to me three of the specimens used in his original
description. One of these (SU 51593), an adult female 26.5 in SL,
is here designated as the lectotype. The two other specimens, paralec-
totypes, one an adult female 28.5 mm. and one subadult female 21.0
mm. in SL, now bear the number SU 51592.

12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

The color pattern of Nannostomus espei greatly resembles that of
Pyrrhulina vittata Regan and Pyrrhulina spilota Weitzman. Many of
the scales of Nannostomus espei are more like those in the genera
Pyrrhulina, Copeina, and Copella than those of other species of
Nannostomus or Poecilobrycon. In figure 1, the scale to the left is
from the third longitudinal scale row, just below the dorsal fin. It is
typically pyrrhulininan in outline. That to the right is from the
second longitudinal row near the adipose fin and is typically nan-
nostominan in outline. The phylogenetic significance of these facts
is not understood, especially since this species has the typical osteology
of Nannostomus. It seems possible that this species has retained a few
primitive features that were present in its presumably somewhat more
Pyrrhulina-like ancestors. This species also has a body form which
suggests that of the subtribe Pyrrhulinina more than the body form
of any other species of Nannostomus or Poecilobrycon. 'The body form
of Nannostomus and Poecilobrycon is usually more or less smoothly
fusiform, whereas that of Pyrrhulina is not, the profile showing a
slight notch behind the dorsal fin and the belly being slightly flat.
Nannostomus espet has the dorsal notch of Pyrrhulina. This body
shape in espei is subtle and considerable experience with live and
preserved specimens was necessary before it was detected.

SPECIMENS EXAMINED.—Five (one alizarin preparation), ANSP 73873, SL 21.5-
28.1 mm., British Guiana, an unnamed creek, tributary to the Paruma River, a
tributary of the Mazaruni River via the Kamarang River, Pakaraima Mountain
region of western British Guiana, lat. 5°50’—55’ N., long. 61°05’-10’ W., 1955,
Louis Chung.—One alizarin preparation, SU 50252, SL 25.0 mm., aquarium speci-
mens, locality unknown but probably British Guiana.—Seven, SU 50214, SL 25.5-
27.0 mm., aquarium specimens, locality unknown but probably British Guiana.—
One, SU 51593 (lectotype) and two, SU 51592, paralectotypes, for locality see
discussion on page 11.

Nannostomus beckfordi Giinther
Figures 2, 3, 4

Nannostomus beckfordi Giimther, 1872, p. 146 (original description; type locality:
Goedverwating, a plantation on the coast of Demerara, British Guiana;
holotype in British Museum [Natural History]).—Eigenmann and Eigen-
mann, 1891, p. 49 (listed) —Higenmann, 1912, p. 281 (copied description).—
Hoedeman, 1950, p. 16 (description; no specimens listed).—Boeseman, 1954,
p. 18 (Surinam).

Nannostomus anomalus Steindachner, 1876, p. 129 (original description; type
locality: mouth of Rio Negro, Brazil; types in Vienna Museum).—Higenmann
and Higenmann, 1891, p. 49 (listed) —Eigenmann, 1910, p. 427 (listed).—
Meinken, 1931, p. 554 (description from life; figure) —Innes, 1935 and later
editions, p. 153 (description from life; photograph).—Puyo, 1949, p. 117
(description; Cayenne Island, French Guiana).—Boeseman, 1952, p. 184
(Surinam); 1953, p. 16 (Surinam).—Axelrod and Schultz, 1955, p. 239
(description and aquarium notes).

NO. 3538 CHARACID FISHES—WEITZMAN 13

Nannostomus minimus Eigenmann, 1909, p. 42 (original description; type locality:
Erukin, British Guiana; holotype in Chicago Natural History Museum);
1910, p. 427 (listed) ; 1912, p. 282, pl. 36, fig. 5 (description).

Nannostomus simplex Higenmann, 1909, p. 42 (original description; type locality:
Lama Stop-Off, British Guiana; holotype in Chicago Natural History
Museum); 1910, p. 427 (listed) ; 1912, p. 283, pl. 36, fig. 6 (description).

Nannostomus beckfordit.—Eigenmann, 1910, p. 427 (listed, name emended).

Nannostomus beckfordi surinamt Hoedeman, 1954a, p. 84 (original description;
type locality: Berg en Dal at Surinam River, Surinam; holotype in Zoological
Museum of Amsterdam).

Nannostomus aripirangensis Meinken, 1931, p. 553 (original description; figure;
type locality: Brazil, State of Para, Aripiranga Island NE. of Belém do Par4;
types destroyed during World War II, Meinken in litt.) —Axelrod and Schultz,
1955, p. 240 (copied description and aquarium notes).—Sterba and Tucker,
1962, p. 210 (description and aquarium notes).

Nannostomus beckfordi aripirangensis.—Hoedeman, 1950, p. 18 (copied descrip-
tion); 1954a, p. 84 (listed).

Nannostomus beckfordi anomalus.—Sterba and Tucker, 1962, p. 210 (description
and aquarium notes).

Diaqnosis.—This species possesses few characters not found in
at least some of the other members of the genus Nannostomus and is
therefore difficult to diagnose by any single characteristic. Perhaps
the best single diagnostic character complex is the pattern of horizontal
stripes.

Secondary and tertiary horizontal stripes absent; well-developed
primary horizontal stripe present; few (2-6) perforated lateral line
scales present; 9 to 10+17 to 18 gill rakers; iii,9 anal fin rays; and 23
to 26 scales in a lateral series.

Description.—Body slender, cylindrical except in region of
moderately compressed caudal peduncle. Greatest body depth
slightly anterior to dorsal fin origin. Standard length of largest
specimen 29.6mm. Greatest body depth 4.3 [3.5 in egg-filled females]
to 5.0 [in apparently starving specimens]; least depth of caudal
peduncle 9.9 (8.8-10.8); length of caudal peduncle of population
sample from Obidos (CM 19648) 5.2 (4.9-5.6), of population from
Uruaré Brook (SU 50257) 6.0 (5.5-6.3); length of caudal peduncle
of all specimens measured 5.8 [a rather meaningless figure since the
measurements are biased toward a large number of specimens from
Uruardé Brook, SU 50257] (4.9-6.3); snout tip to origin of dorsal fin
1.85 (1.62-1.99) ; snout tip to origin of anal fin 1.30 (1.24-1.35).

Head conic; snout blunt, obtuse in vertical and horizontal profile.
Head 3.8 (3.5-4.0); eye in head 2.8 (2.5-3.2); snout in eye 1.24
(1.05-1.39) ; least width of bony interorbit in greatest eye diameter 1.1
(1.04-1.19).

Premaxillary with 6 teeth; anterior teeth quinquecuspid, lateral
teeth quinque- to septemcuspid. [There is some geographical varia-
tion in the prominence of the individual cusps; for example specimens

14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

from Baduel, French Guiana, SU 50259, and specimens from Uruard
Brook, SU 50257, have all the cusps of the premaxillary teeth about
equal (fig. 2). Specimens from Obidos have the second cusp from
posterolateral side much enlarged, quite similar to the upper left tooth
shown in the figure of Poecilobrycon eques (fig. 12)]. Maxillary with
single uni- to octocuspid compressed tooth, cusps subequal. Dentary
with 5 to 6 teeth (almost always 6 in all population samples examined).
Teeth in anterior dentary tooth row quinque- to sexcuspid (usually
sexcuspid). Specimens from Uruaré Brook with subequal cusps;
however, specimens from Obidos with centralmost cusp enlarged in
anterior 4 to 5 teeth of dentary. Teeth 2 to 6 in inner tooth row of
dentary (seen in alizarin preparations only). Gill rakers 9 to 10+-17
to 18 (seen in alizarin specimens only).

Ficure 2.—Nannostomus beckfordi, male, SL 30.0 mm., SU 50261 (both scales are from
third longitudinal row, below dorsal fin).

Dorsal fin 11,8; anal fin iii,9; pectoral i,10 to i,12; pelvic fin ii,7.
Adipose fin absent in all specimens examined. Principal caudal fin
rays 10/9; distal tips of all elements of first 10 rays contained in upper
caudal lobe, distal tips of remaining 9 principal rays in lower lobe.
Male with anal fin (figs. 2 and 4) somewhat modified in same manner
as extremely modified anal fin of N. digrammus. Anal fin of female
(fig. 3) unmodified. [Observation of living male beckfordi shows
that while courting and spawning, the male often cups the fin into a
spoon-shaped bowl and that, while spawning, the male places his fin
partially over the female’s vent.]

For radial grooves of scales in median scale row below dorsal fin,
see figure 2. Posterior field with 2 to 6 radial grooves; dorsal and
ventral fields with 1 or 2 radial grooves; and anterior field with 3 to

—

NO. 3538 CHARACID FISHES—WEITZMAN 15

7 radial grooves. Scales in lateral series 24 (23-26) ; perforated lateral
line scales 2 to 6 (usually 4 in population from Uruard Brook). Speci-
mens often damaged so heavily that scales impossible to count ac-
curately. In all instances, intact specimens from other localities
with only 4 to 6 perforated scales. Scales in median series anterior
to dorsal fin 10-11. Specimens from Uruaré Brook with following
total vertebral counts: 6 specimens with 34; 19 with 35 vertebrae.
Specimens from Obidos: 3 specimens with 33, 31 specimens with 34
vertebrae, 8 specimens with 35 and 2 specimens with 36 vertebrae.
Most specimens from other localities with 35 vertebrae. Usually
18 to 20 precaudal vertebrae and 15 to 16 caudal vertebrae, always in
combination to produce total of 33 to 36 vertebrae. Counts of verte-
brae undoubtedly will prove useful in distinguishing and determining
subspecies.

hn
ZEN EOE.
ay ip

Se
ves
Ly

LS
So
——

Ficure 3.—Nannostomus beckfordi, female, SL 29.5 mm., SU 50261.

CoLor IN ALCOHOL.—Preserved specimens from Belém, State of
Pard, Brazil (figs. 2 and 3) have the following color: Top of head
light to dark brown. Primary horizontal stripe extending from tip
of snout and tip of lower jaw across maxillary and first orbital bone
onto eye. Posterior to eye, primary stripe continuing to posterior
edge of opercular bone, but absent or extremely pale on fleshy oper-
cular flap. Beginning on cleithrum and underneath opercular flap,
primary horizontal stripe continuing posteriorly over lower midsides
onto caudal fin, its course reaching posterior root of anal fin, covering
entire lower half of caudal peduncle and continuing onto caudal fin
as a wedge with its apex at posterior junction of upper and lower
caudal fin lobes. In mature males primary horizontal stripe covering
entire fourth horizontal scale row and adjacent parts of third and
fifth scale rows. In mature females and immature specimens of both

16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

sexes lower portion of third scale row often only partly pigmented.
No evidence of secondary or tertiary horizontal stripes.

Very pale stripe extending above primary horizontal stripe from eye
to end of third scale row; this stripe often obscured by light brown
color extending ventrally from dorsum. Females, and sometimes
males, with scatterings of melanophores forming oblique bands in area
of sixth to eighth vertical scale rows and over thirteenth to fifteenth
vertical scale rows. Sides pale below primary horizontal stripe,
shading to white on belly. Pectoral and pelvic fins translucent but
pelvic fins sometimes with scattering of brownish melanophores giving
entire fin light brown appearance. Dorsal fin translucent except for
fairly dense accumulation of melanophores along anteriodistal portion.
Anal fin hyaline except for some accumulation of melanophores in
distal region of posteriormost rays.

Cotor In LiFE.—(The color recorded here is from breeding male
aquarium specimens from an unknown locality. These specimens
appeared, when preserved, exactly like the specimens described above
from Belém.) Dorsal fin, pectoral fins, and pelvic fins mostly hyaline
without red except for small amount at base of pelvic fins. Primary
horizontal stripe dark bluish black. Narrow silvery blue to golden
stripe above primary horizontal stripe, beginning at posterior border
of eye and extending to posterior end of caudal peduncle. Above this,
beginning between third to sixth vertical scale row, with narrow red
stripe, varying in intensity from deep crimson red to pale rose. This
red stripe usually absent in nonbreeding males. Intense crimson spot
above and below primary horizontal stripe at base of each caudal lobe.
Intense red stripe often spreading completely over anal fin below pri-
mary horizontal stripe, and usually extending from sixth or seventh
vertical scale row posteriorly to above anal fin in breeding males.
Nonbreeding males often without red. Distal one-quarter to one-
third of pelvic fins intense silvery blue. Some specimens with this
color on distal tips of third, fourth, and fifth anal fin rays. Mature
females and immature specimens of both sexes often with small amount
of silvery blue at distalmost tips of pelvic rays. Dorsum pale brown
above silvery and red stripes with each exposed scale edge outlined
in darker brown. Lateral surface of snout above primary horizontal
stripe pale brown. Top of head brown. Bright metallic-red spot
frequently present near anterior narial opening. Iris silvery with
reddish suffusion dorsally. Belly and throat regions white.

Remarxs.—Hoedeman (1950, pp. 16-18) seems to have been
correct in assuming Nannostomus anomalus Steindachner to be a
synonym of Nannostomus beckfordi, a polymorphic species. Hoedeman
pointed out that the only difference between beckfordi and anomalus
recorded in the literature is that beckfordi has a black spot on the

nnn © 4 a ee ee

NO. 3538 CHARACID FISHES—WEITZMAN 7

lower half of the gill cover. Hoedeman noted that at night there is a
dark spot on the lower half of the gill cover of living specimens of a
fish, apparently without known locality. He considered this fish to
be beckfordi. These specimens apparently also fit the description of
anomalus. I have been unable to confirm the presence of such a
spot in living specimens available to me. The live specimens I have
seen are without locality but are like specimens known to have come
from near Belém, Para, Brazil. Professor George S. Myers kindly
examined the holotype of beckfordi in the British Museum and noted
that the black spot on the lower half of the gill cover is a dirty black
smudge, does not consist of melanophores, and occurs on the left side
only. Since the spot appears to be an artifact, anomalus is here
considered a synonym of beckfordi.

Meinken (1931) described Nannostomus aripirangensis from
Aripiranga Island near Belém, Parad, Brazil. Herr Meinken (in litt.)
has stated that the types of aripirangensis were destroyed during
World War IJ, and no one reexamined them subsequent to Meinken’s
original account. Hoedeman (1950, p. 118) was possibly correct in
assuming aripirangensis to be a subspecies of beckfordi. I have
examined two specimens of Nannostomus from Aripiranga Island that
were sent to Dr. George S. Myers prior to 1934 by Arthur Rachow,
a prominent German aquarist. These specimens are said to be
representatives of aripirangensis and are identical to specimens of
beckfordi from around Belém. However, Meinken (1954, and in
personal communications) has insisted that aripirangensis represents
a distinct species. JI have seen many living aquarium specimens of
Nannostomus without locality data which I consider to be color vari-
ants (possibly geographical races or subspecies) of beckfordi. In
appearance, one of these groups of specimens was like specimens of
beckfordi collected at Belém, Pard, and Aripiranga Island and had,
when breeding, some colors suggesting those described for aripiran-
gensis by Meinken. Aripiranga Island, the type locality of aripiran-
gensis, is just a few miles from the city of Belém, Para.

Meinken showed in his drawings a small tertiary horizontal stripe,
a stripe which I have never seen in any specimens that I could refer
to beckfordt. Meinken’s dorsal fin count of 11 rays should be con-
sidered with caution and some doubt, for all Nannostomus and Poect-
lobrycon so far examined by me have had a dorsal fin ray count of
1,8 or a total of 10 rays, not 11. Meinken’s count may have been
taken from an abnormal or rare variant specimen. Meinken has
stated in a personal communication that anomalus (here beckfordi)
and arvpirangensis have interbred in aquaria and that in this manner
the living stock of aripirangensis has been lost in Germany. This
indicates at least a close genetic relationship. Recently Vorderwinkler

219-936—66——2

18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 119

(1957), in an aquarium publication, has recorded and published pho-
tographs of two forms of Nannostomus, one of which is stated to be
anomalus, the other aripirangensis; both of these photographs appear
to be of beckfordi, showing two of its several geographical color vari-
ants. Neither has a color pattern exactly like that described for the
original aripirangensis.

Until thorough collections of Nannostomus can be made on the
Island of Aripiranga, the problem of the distinctness of aripirangensis
cannot be settled; however, I tentatively prefer to consider aripiran-
gensis one of the several geographically distinct populations of
beckfordi.

IS
SIH TN
Le LAS TSI %
LO ORS SEE
QO *S
RN Qy eS a

Ficure 4.—Anal fin of male Nannostomus beckfordi, SL 31.0 mm., SU 50261.

Nannostomus beckfordi appears to be a polytypic species with a
wide geographical range, known from the Guianas and for a con-
siderable distance (about 700 or more air miles) up the Amazon
River. Adequate collections and studies will undoubtedly show that
this species consists of several distinct populations, some of which
may prove to be subspecies. Studies of color variation, variation in
certain body proportions (especially head depth compared to body
length and caudal peduncle length), vertebral counts, and arrange-
ment of tooth cusps will probably be of great value in studying sub-
specific population differences.

SPECIMENS EXAMINED.—Three, SU 50256, SL 22.0-22.7 mm., Brazil, State of
Pard, Belém, May 18, 1924, Carl Ternetz.—Four, SU 50262, SL 22.5-27.5 mm.,

Brazil, State of Pard, brook into TApajos near Santarém, Aug. 2, 1924, Carl
Ternetz.—Forty-four, SU 50257, SL 16.0-29.0 mm., Brazil, State of Parad, south

NO. 3538 CHARACID FISHES—WEITZMAN 19

bank of Rio Amazonas, Rio Uruard, about lat. 1°54’ S. and long. 53°27’ W.,
June 26, 1924, Carl Ternetz.—Six, SU 50258, SL 22.0-26.5 mm., alizarin prepara-
tions, Brazil, State of Parad, south bank of Rio Amazonas, Rio Uruard4 about lat.
1°54’ S. and long. 53°27’ W., June 26, 1924, Carl Ternetz (from SU 50257) .—
Five, SU 50261, SL 20.5-29.6 mm., Brazil, State of Pard, Belém (Utinga Forest
Reserve), September 1944, George S. Myers.—Two, SU 18469, SL 20.5-22.5 mm.,
no locality, aquarium specimens sent to Dr. George S. Myers as specimens of
Nannostomus aripirangensis.—Two, USNM 94207, SL 30.0-31.0 mm., Brazil,
state of Pard, Aripiranga Island, sent to Dr. George 8. Myers prior to 1934 by
Arthur Rachow.—Four, SU 50269, SL 19.0-25.0 mm., Brazil, State of Para,
Igarapé Uruar4, tributary of Furo Maic4, about 10 miles southeast of Santarém,
about lat. 2°30-55’ S. and long. 54°35-41’ W., June 23, 1924, Carl Ternetz.—
Two, SU 50260, SL 25.0 mm. (both specimens), Brazil, State of Parad, Lagoa do
Maicd, at the junction of the Igarapé Maicé and the Furo do Ituqui, about lat.
2°27’ S. and long. 54°40’ W., Apr. 2, 1924, Carl Ternetz.—Three, SU 50255,
SL 22.0-24.0 mm., Brazil, State of Pard, Santarém, brook into Rio Tapajos,
June 30, 1924, Carl Ternetz.—Nine, SU 50259, SL 21.0-25.4 mm., French Guiana,
Baduel, on the Ile de Cayenne, July 26, 1947.—Forty-three, CM 19648, SL
20.0-27.0 mm., Brazil, State of Pard, Obidos, 1856, Col. Bentos, Thayer Expedi-
tion.—One, CAS (LUM) 11692, SL 20.3 mm., paratype of Nannostomus simplez ,
British Guiana, Lama Stop-Off, Mahaica River, 1908, Carl H. Eigenmann.

Nannostomus bifasciatus Hoedeman

Nannostomus bifasciatus Hoedeman, 1953, p. x30.11.311 (original description,
in Dutch; type locality: Berg en Dal at Surinam River, Surinam; holotype
in Amsterdam Museum); 1954a, p. 85 (description in English) ; 1954b, p. 77
(description from life).—Sterba and Tucker, 1962, p. 211 (description and
aquarium notes).

Dracnosis.—At least a few perforated lateral-line scales present;
about 8+14 gill rakers; iii,9 anal fin rays, secondary horizontal stripe
present, scales in a lateral series about 25.

Norr.—This species is known from only four specimens and the
original author has kindly sent three of them for examination. The
fish was an aquarium import from Surinam to the Netherlands and
the three available specimens are in poor condition, two having been
dried at one time and all with torn, broken fins and broken jaws with
missing teeth. Unfortunately the original description was inadequate
and inaccurate, and it is impossible completely and competently to
describe or illustrate the fish from the specimens at hand.

Description.—Body elongate; sides somewhat flattened and body
compressed posteriorly. Greatest depth just anterior to origin of
dorsal fin. Standard length of longest specimen examined 28.3 mm.
Greatest body depth 5.4 (5.3-5.6) [females with ripe eggs not at hand
but undoubtedly deeper bodied (see illustrations of live specimens
in Hoedeman 1954a and Nieuwenhuizen 1954)]; least depth of caudal
peduncle 11.6 (11.4-11.9); length of caudal peduncle 5.6 (5.5-5.6)
in females, male damaged. Snout tip to origin of dorsal fin 1.9
(1.9-1.9) ; snout tip to origin of anal fin 1.3 (1.3-1.3).

20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Head conic and snout obtuse in both vertical and horizontal pro-
files, top of head gently convex between eyes. Head 3.8 (3.8-3.8);
eye in head 3.1 (3.0-3.2); snout in eye 1.3 (1.2—1.3); least width of
long interorbital in greatest eye diameter 1.3 (1.3-1.3).

Premaxillary with 7 teeth; anterior and lateral teeth septemcuspid,
third from anterior or median cusp largest. Maxillary with 1 quadri-
cuspid tooth. Dentary with 6 teeth, each usually with 7 cusps. Third
cusp from medial or anterior end of tooth largest; sometimes central
cusp largest. Inner row of teeth damaged in all specimens but appears
to be over 10 simple conic teeth. Gull rakers 8+14 (one alizarin,
specimen).

Dorsal fin 1,8; anal fin 1,9. Adipose fin absent. Caudal fin with
principal rays 10/9.

Anal fin of male in good condition, somewhat modified. First
ray small and completely hidden by a basal scale. Second ray enlarged
and its anterior edge expanded and carinate. Entire fin larger than
that of females and individual rays thickened laterally and somewhat
enlarged in an anterior and posterior plane. In folded position, rays
take an alternate position to each other with regard to median plane.
However, anal fin not nearly as modified as that of Nannostomus
digrammus.

Radial grooves of scales belonging to that part of third horizontal
scale row below dorsal fin having posterior field with 1 to 3, usually
2 radial grooves; dorsal and ventral fields with 1 and anterior field
with 3 to usually 4 radial grooves. Scales in lateral series 25 in
single countable specimen. This specimen also with 3 perforated
lateral-line scales in single row behind cleithrum. Scales in a median
series before dorsal fin 10 in 2 female specimens. Total number of
vertebrae 36 (19 precaudal and 17 caudal).

Co.or IN ALcoHOL.—T op of head and back light brown. Dorsum
at base of dorsal fin dark brown. Primary horizontal stripe dark
brown, extending from snout tip and lower jaw onto anterior surface
of eye, across lower border of fifth orbital bone and upper border of
fourth orbital bone to over upper limb of preopercle, finally terminat-
ing at posterior border of bony opercle. On body, primary horizontal
stripe beginning anteriorly underneath fleshy opercular flap and
extending through middle of fourth scale row, covering lower and
upper exposed portions of third and fifth scale rows respectively.
Primary stripe widening posteriorly until over anal fin covering
entire fourth scale row and upper half of fifth scale row. Primary
horizontal stripe covering entire lower half of caudal peduncle and
entering lower lobe of caudal fin. Its pattern over lower lobe of
caudal fin could not be determined due to damage. Below primary
horizontal stripe, sides and belly pale brown (probably silvery in

re

EES

SS SS

NO. 3538 CHARACID FISHES—WEITZMAN DF

well-preserved specimens) with few irregularly scattered melano-
phores. No evidence of tertiary stripe. Single broad pale brown
stripe (probably silvery in well-preserved specimens) above primary
horizontal stripe extending from eye to caudal fin base. Two areas of
large scattered melanophores occurring on pale stripe, forming oblique
bands; anterior band covering 4 scales anterior to a point below
anterior origin of dorsal fin, posterior band covering another 4 scales
and center of this area located at a point dorsal to vent. Just dorsal
to pale stripe a narrow secondary stripe arising a short distance above
termination of upper opercular opening and extending posteriorly
through center of second scale row to below dorsal fin, here becoming
wider, denser, and rising to top of second scale row, there merging
with dark pigment of dorsum behind dorsal fin and above pale stripe
on caudal peduncle. Dorsum anterior to dorsal fin pale brown, lighter
than secondary stripe. Top of head and snout with scattered large
and small melanophores forming an area darker than dorsum just
anterior to dorsal fin. All fins colorless except for a few scattered
melanophores and dark pigment on lower caudal lobe. Pelvic fins
appear artificially stained gray.

Cotor IN Lire.—Life colors are taken from the original English
description. Primary horizontal stripe ‘‘brilliant’”? black, extending
onto caudal fin rays and onto lower caudal lobe. Typical wedge-shape
distribution of pigment on lower caudal fin lobe apparently not
present but precise distribution of black pigment not stated. Dorsal
and ventral region dusky, back darker. Faint “reddish flush” on
base of caudal and anal fins. Pelvic tipped with “‘ice blue,” especially
in males. Sides above and below primary horizontal stripe white
except for narrow ‘“‘golden-red streak” above primary stripe. (Hoede-
man stated that this streak is diffuse at times except on snout. From
this I would assume that there is probably a golden or red metallic
spot at the anterior narial openings as in so many other nannostomi-
nans.) Secondary horizontal stripe present but tertiary stripe
absent.

REMARKS.—Concerning his new species, N. bifasciatus, Hoedeman
(1954a) stated: ‘‘Phylogenetically the new species seems close to the
stem of the tribe [i.e., his tribe Nannostomidi], near the presumed
Pre-Nannostomid, taking an intermediate position between Nanno-
stomus beckfordi and Nannobrycon eques.” Hoedeman’s opinion was
apparently based on the following observation: ‘The oblique swim-
ming position of Nannobrycon though less pronounced is sometimes
observed in the present species [bifasciatus] also, especially while
nipping planktonic food from the plants.”” He also mentioned that
the black pigment of the caudal fin extends on the lower caudal
fin lobe somewhat as in Poecilobrycon eques and remarked that the

22, PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

adipose fin of bifasciatus is placed more like that of members of
Nannostomus than that of Poecilobrycon harrisoni, which has the
adipose fin posterior to the anal fin, not over it as in Nannostomus.
Hoedeman remarked that bifasciatus has a vestigal adipose fin;
however, I find no trace of one in his 3 specimens. In addition I
fail to find any generic significance in the placement of the adipose
fin in nannostominans. It is somewhat more posterior in Nanno-
stomus trifasciatus and Poecilobrycon harrisoni but more anteriorly
placed in all other species of nannostominans which possess this
fin. Concerning the swimming position, I can make little comment
because I have not seen living specimens of bifasciatus. However,
it should be remarked that Poecilobrycon harrisoni normally swims
horizontally like all living species of Nannostomus that I have seen
and that all species of Nannostomus will occasionally swim slightly
obliquely when feeding. The color claimed for the caudal fin may
be like that of Poecilobrycon eques, but I cannot determine this from
the specimens at hand. The precise significance of this color pattern
if present would be difficult to determine. From the key in the
present paper it would seem that bifasciatus is closest to beckfordi
in several characters and in fact may be rather closely related to it.
I fail to find that Hoedeman presented sufficient evidence to indicate
either that difasciatus is a relatively primitive nannostominan or
that it is intermediate between Nannostomus and Nannobrycon. The
osteology of bifasciatus is typically that of Nannostomus. Weitzman
(1964) noted that pyrrhulininans and nannostominans undoubtedly
had a common ancestor. Presumably a relatively primitive nanno-
stominan would have some characters suggesting pyrrhulininans.
The only nannostominan that does this is Nannostomus espei, and
even in this species the few such characters are difficult to evaluate.
SPECIMENS EXAMINED.—Three, paratypes, AM 100513, SL 27.2-28.3 mm.

(only 2 specimens could be measured as snout of 1 damaged), Surinam, Berg en Dal
at Surinam River.

Nannostomus digrammus Fowler
Fiaures 5, 6

Nannostomus digrammus Fowler, 1913, p. 525 (original description; type locality:
Rio Madeira, about 200 miles east of long. 62°20’ W., Brazil;? holotype in
Academy of Natural Sciences of Philadelphia), 1950, p. 261 (listed).

Poecilobrycon diagrammus.—Hoedeman, 1954a, p. 84 (copied description).

Poecilobrycon digrammus.—Hoedeman, 1954b, p. 71 (listed).

Nannostomus beckfordi beckfordi.—Sterba and Tucker (not Giinther), 1962, p. 209,
(aquarium description of “gold anomalus’” but counts from Giinther’s
original description of beckfordi; see remarks below).

2 Bohlke (1955, pp. 8-12) doubts the quoted locality and presents reasons why
it may be considered as being 20 miles east of long. 62°20’ W.

NO. 3538 CHARACID FISHES—WEITZMAN 23

Dragnosis.—This species may be distinguished from all other
known species of Nannostomus and Poecilobrycon by the greatly
modified anal fin of the males (fig. 6) and an anal fin count of iii,8 in
both sexes. In addition, the following combination of characters will
distinguish this species from all others: Primary and secondary
horizontal stripes well developed; tertiary horizontal stripe absent
or only weakly developed; no perforated lateral-line scales; and
adipose fin present.

Description.—Body elongate, rather cylindrical in its anterior
half but compressed posteriorly. Greatest depth a short distance
anterior to origin of dorsal fin. Standard length of largest specimen
with locality data 25.0 mm. (an aquarium specimen measured, SU
18471, SL 27.6 mm.). Greatest body depth 4.7 (4.5-5.0); least
depth of caudal peduncle 11.1 (10.3-11.5); length of caudal peduncle,
males 5.9 (5.6—6.3), females 5.1 (4.9-5.4); snout tip to origin of dorsal
fin 1.8 (1.7-1.9); snout tip to origin of anal fin 1.3 (1.29-1.34).

Ficure 5.—Nannostomus digrammus, male, SL 24.5 mm., SU 50248,

Head conic but somewhat flattened dorsoventrally; snout obtuse
in vertical and horizontal profiles. Top of head between eyes rather
convex. Head 3.7 (3.5-3.8); eye in head 2.8 (2.5-3.0); snout in eye
1.2 (1.1-1.3); least width of bony interorbital in greatest eye diameter
1.2 (1.05-1.26).

Premaxillary with 6 teeth in anterior row, all quadricuspid; posterior
1 or 2 teeth of this row quinque- to sexcuspid, cusps being subequal.
Central cusps of each tooth largest and longest. Maxillary with 1
quinque- to sexcuspid tooth. Dentary with 5 to 6 (usually 6) teeth
in outer row, anterior 4 teeth with 6 cusps, posterior 2 teeth with
4 to 5 cusps, cusps subequal. Teeth in inner or second tooth row of

24 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

dentary 1 to 4 in the alizarin specimens. Gill rakers 7+-14 (alizarin
specimens).

Dorsal fin 1,8; anal fin iii,8; pectoral 1,8 to 1,10; pelvics i1,7. Adi-
pose fin present on all specimens, its base above scale row that runs
diagonally to posterior base of anal fin. Caudal fin with principal
rays10/9. Distal tips of all elements of first 10 rays contained in upper
caudal lobe. Lower lobe containing all distal tips of remaining 9
principal rays.

Highly modified anal fin of male Nannostomus digrammus (fig. 6)
with an interesting structure. Unbranched anterior rays 3, followed
by 7 once dichotomus branched rays; terminal or last ray unbranched.
Last ray not divided to its base and with its own separate pterygio-
phore, or radial series. Except for anteriormost small ray hidden be-
low scale at anterior base of anal fin, each of first 6 rays greatly ex-
panded anteriorly and posteriorly. These rays arranged as follows in
collapsed fin: Second ray (first large ray of anal fin) partially enclosed
by anterior edges of ray halves of third anal ray. Thus appressed
posterior borders of ray halves of second ray enclosed by 2 diverging
anterior borders of third anal finray. Anterior edge of fourth anal ray
(including both ray halves) lying to left of posterior border of third
anal ray. Anterior border of fifth anal ray lying to right of posterior
border of third anal ray and to right of entire middle region of fourth
anal ray. Sixth anal ray lying to left of posterior border of fifth ray
and will even slip to left of posterior border of fourth ray. Anterior
edge of seventh anal ray lying to right of posterior half of sixth anal
ray and posterior border of seventh ray lying to right of anterior
border of eighth ray. Anterior border of ninth ray lying to right of
posterior border of eighth ray, but tenth and eleventh rays with usual
position in being just posterior to rays anterior to them. Fourth ray
with its left lateral face convex, its right lateral face concave. Fifth
ray, converse of fourth, and sixth ray with convex and concave faces
facing same sides as fourthray. This alternation continuing to eighth
ray. Ninth through eleventh anal rays with little if any convexity or
concavity.

Inclinator muscles extending bilaterally ventrally far beyond base of
anal fin rays and inserting on lepidotrichs to about one-half length of
tenth and eleventh anal fin rays. Similarly inclinator muscle extend-
ing ventrally beyond scales and inserting along basal one-fifth of
second anal fin ray (fig. 6).

Many male nannostominans and some male pyrrhulininans have
anal fins that are modified in a manner similar to that noted here for
Nannostomus digrammus. WHowever, none have their anal fin rays
modified as greatly as those of digrammus. Species of nannostominans
and pyrrhulininans that I have seen breeding in-aquaria all have been

NO. 3538 CHARACID FISHES—WEITZMAN 25

capable, to a greater or lesser degree, of twisting the anal fin into a cup-
shaped organ that is placed with the concave side almost over the fe-
male’s vent during the act of laying an egg. Wickler (1957) has re-
corded some of the behavior of various species of nannostominans but
has not noted the precise use of the anal fins. Although I have not
| seen the anal fin of the male Nannostomus digrammus in use, its morph-
| ology indicates that it undoubtedly changes greatly in shape during
_ the spawning act.

Ficure 6.—Anal fin of male Nannostomus digrammus, SL 19.6 mm., SU 50251.

Radial grooves of scales belonging to third horizontal scale series and
lying below dorsal fin shown in figure 5. Posterior field with 1 or 2
radial grooves; dorsal and ventral fields each with 1 and anterior field
with 3 or 4 radial grooves. Scales in lateral series 25 to 26; no per-
forated lateral-line scales; scales in median series anterior to dorsal fin
10 to 11. Total number of vertebrae 34 to 35, 18 to 19 precaudals and
15 to 16 caudals, but always in a combination to produce a total count
of 34 to 35 vertebrae.

Cotor In ALcoHoL.—Top of head black. Primary horizontal stripe
black, extending from snout tip and lower jaw onto anterior surface
of eye, then from posterior surface of eye across lower border of fifth
orbital bone and upper border of fourth orbital, then continuing over
upper arm of preopercle onto opercle ending at about posterior bony

26 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

border of opercle. On body, primary horizontal stripe begins anteriorly
underneath fleshy opercular flap and extends through approximately
lower half of fourth scale row and upper third of fifth scale row.
Primary stripe widening posteriorly until, over anal fin, there covering
entire fourth, and one-half of fifth scale rows. Posterior to this,
primary horizontal stripe covering almost entire lower half of caudal
peduncle and continuing as an acute wedge posteriorly over caudal fin,
ending at base of caudal fork. Below primary horizontal stripe,
sides and belly silvery with few irregularly scattered melanophores.
Broad silvery stirpe extending from eye to caudal base above primary
horizontal stripe. Silvery stripe with oblique band, about 3 scales
long, located just anterior to dorsal fin. This area spotted with large,
dark melanophores. Few small melanophores scattered over entire
silvery stripe. Narrow, dark secondary horizontal stripe just dorsal
to silvery lateral stripe, beginning at upper posterior edge of operculum
and ending on upper part of caudal peduncle posterior to adipose fin.
Dorsum pale brown above secondary horizontal stripe. Areas of head
and eye not covered by dense black pigment, silvery with few scattered
melanophores. Dorsal, pelvic, pectoral, and anal fins colorless.
Caudal fin, in addition to having narrow wedge of primary horizontal
stripe, with fairly dense aggregation of melanophores along first 3
upper principal rays of upper lobe. Some melanophores scattered
over entire fin. Color in life not known with certainty.

Remarks.—Nannostomus digrammus was known previously only
from the four specimens reported in the original description. However,
this species has been imported as an aquarium fish. Several years ago
a prominent aquarist, Mr. Frederick Stoye, sent a specimen of a
nannostominan to Dr. George S. Myers at Stanford University.
This specimen, labeled as the ‘gold anomalus,” proved to be a large
male digrammus.

All the specimens belonging to the original type lot of digrammus
were females, and the holotype illustrated by Fowler shows the anal
fin of a female.

Sterba and Tucker (1963, p. 209) believed the so-called “gold
anomalus” to be Giinther’s original Nannostomus beckfordi. However
Stoye’s specimen mentioned above would seem to show that the “gold
anomalus” of aquarists is N. digrammus. Aquarists’ records of this
fish cited by Sterba and Tucker (1963) are difficult to compare.
Perhaps more than one species of fish is involved. However, live
specimens of Nannostomus beckfordi from British Guiana are compar-
atively pale in color, never exhibiting the colors described for the
“gold anomalus.”

SPECIMENS EXAMINED.—Two, paratypes, ANSP 39190 and 39191 SL 18.0-19.0
mm., Brazil, State of Amazonas, Rio Madiera, about 200 miles east of long.

NO. 3538 CHARACID FISHES—WEITZMAN HT

62°20’ W., September 1912, Edgar A. Smith.—Eleven, MCA 19797, plus 2 ali-
zarin specimens SU 50249, SL 16.3-19.4 mm., Brazil, State of Amazonas, Mandos,
1865, Louis Agassiz.—Two (male figured), SU 50248, SL 24.1-24.5 mm., Brazil,
Parad, Igarapé do Sapucud into Lagéa Sapucud (This lake drains into the Rio
Trombetas. The Trombetas flows into the Amazon from the north about 25
miles west of Obidos.), June 7, 1924, Carl Ternetz.—Fifteen, SU 50250, three
alizarin preparations SU 50251, SL 18.0-22.8 mm., Brazil, State of Amazonas,
Igrapé do Mai Joana, a tributary of the Rio Negro near Mandos, Dec. 25, 1924,
Carl Ternetz.—One, SU 18471, SL 27.6 mm., (a large adult male), an aquarium
specimen sent to Dr. George S. Myers by Mr. Frederick Stoye.—Hight, SU 51024,
SL 14.8-16.0 mm., British Guiana, Rupununi District, Nappi River, Rio Branco
drainage, about lat. 30°30’ N., long. 59°32’ W., Sept. 29, 1957, Rosemary Lowe
(McConnell).—One, SU 51025, SL 15.5 mm., British Guiana, Manawarin River,
a tributary of the Moruka River about lat. 7°30’ N., long. 59°11’ W., July 16,
1957, Rosemary Lowe (McConnell).—One, SU 51026, SL 16.0 mm., British
Guiana, Kumarow Creek pool into the Rupununi River, a tributary of the
Essequibo River, Dec. 10, 1957, Rosemary Lowe (McConnell).—One, SU 50448,
SL 17.2 mm., British Guiana, pool near Takatu River, Rio Branco drainage,
Emprensa, Rupununi District, Dec. 8, 1957, Rosemary Lowe (McConnell).—
One, SU 50450, SL 19.5 mm., British Guiana, Kumarow Creek Pool which flows
into the Rupununi River in southern savannas of the Essequibo River Drainage,
Dec. 10, 1957, Rosemary Lowe (McConnell).

Nannostomus trifasciatus Steindachner
Figure 7

Nannostomus trifasciatus Steindachner, 1876, p. 123, pl. 9, fig. 2 (original
description; localities: backwaters and quiet tributaries of the Amazon close
to mouth of the Rio Negro, Brazil; also Amazon at Tabatinga; types in
Vienna museum).—Higenmann and Higenmann, 1891, p. 49 (listed).—
Eigenmann, 1910, p. 427 (listed).—Innes, 1935, p. 155 (description from life
and photograph) and later editions.—Hoedeman, 1950, p. 18 (description) .—
Axelrod and Schultz, 1955, p. 248 (description, aquarium notes).—Sterba
and Tucker, 1962, p. 212 (description and aquarium notes).

?Cyprinodon amazona Eigenmann, 1894, p. 627 (in part; original description;
type locality: Lower Amazonas; types apparently lost).—Garman, 1895,
p. 28 (in part; copied description).—Hubbs, 1926, p. 16 (in part; discussion
of identity).—Fowler, 1954, p. 216 (in part; listed).

Poecilobrycon erythrurus Eigenmann, 1909, p. 44 (original description; type
locality: Rockstone, British Guiana; holotype in Chicago Natural History
Museum).—Eigenmann, 1910, p. 427 (listed).—Eigenmann, 1912, p. 285,
pl. 37, fig. 3 (description).

?Cyprinodon amazonus Eigenmann, 1910, p. 456 (in part; listed).

Poecilobrycon vittatus Ahl, 1933, p. 84, fig. (original description; type locality:
“Parad”; type presumably in Berlin Museum); 1934, p. 124 (a longer, more
useful description; no figure).

Nannostomus trilineatus (lapsus for trifasciatus) Innes, 1933, p. 142 (description
from life).—Ladiges, 1948, p. 30 (description from life).

NOMENCLATURAL NoTE—The identity of Cyprinodon amazona is in
doubt, but it seems most likely the fish is a synonym of either Nanno-
stomus trifasciatus or Nannostomus marginatus. Eigenmann de-

28 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

scribed the fish from 18 specimens collected by Frederick C. Hartt
somewhere along the ‘Lower Amazon.”’? Hubbs (1926), after ex-
amining the specimens, reported that the fish belongs to the genus
Nannostomus and at that time Eigenmann, in personal communication
with Hubbs, concurred in this referral. Neither Hubbs, Eigenmann,
nor anyone else has tried to identify Eigenmann’s syntypes of amazona
with any known species of Nannostomus. Unfortunately, a check of
various American museums and with Dr. Hubbs has not located the
type specimens. From examination of Eigenmann’s original de-
scription it seems probable that two species, Nannostomus marginatus
and N. trifasciatus, were among the original specimens. Measure-
ments and counts include the range of both species, and the color
description could be of either species. It seems likely that the male
mentioned by Eigenmann is NV. marginatus but that some of the other
specimens are most probably JN. trifasciatus. The name trifasciatus
Steindachner 1876 predates amazona Eigenmann 1894; however,
amazona predates marginatus Eigenmann 1909; Nannostomus mar-
ginatus is a well-established name, especially in the aquarium litera-
ture, where it is used frequently. It would serve no useful purpose
to substitute the name amazonus for marginatus, especially since
some of the syntypes at least appear to have been ¢rifasciatus. If
the types of amazona are ever found and some of them prove to be
trifasciatus, one of these should be selected as the lectotype and
Cyprinodon amazona placed in the synonymy of Nannostomus
trifasciatus.

DraGnosts.—This species, like Nannostomus beckfordi, lacks obvious
unique characters, but it may be readily diagnosed by the following
combination of characters: Anal fin rays completely unmodified; gill
rakers 9+13; scales in a lateral series 26-27; perforated lateral-line
scales 3 to 5; one maxillary tooth present; primary, secondary, and
tertiary horizontal stripes present; 11 teeth in second tooth row of
dentary.

DescripTion.—Body elongate, somewhat cylindrical through an-
terior half, but sides flattened and body compressed posteriorly.
Greatest depth lying between posterior tip of appressed pectorals
and origin of dorsal fin. Standard length of largest specimen 28.2
mm. (a larger aquarium specimen, a male, measured at 32.5 mm.).
Greatest body depth 4.6 (4.5-4.7); least depth of caudal peduncle
10.2 (10.0-10.4); length of caudal peduncle (including both sexes)
5.8 (5.6-6.1); snout tip to origin of dorsal 1.9 (1.8-2.0); snout tip to
origin of anal 1.28 (1.27-1.30).

Head elongate, rather depressed, especially in snout region; top of
head only slightly convex between eyes. Snout obtuse, especially
so in dorsal profile. Head 3.6 (3.4-3.9) [Peruvian specimens 3.73.9,

NO. 3538 CHARACID FISHES—WEITZMAN 29

British Guiana specimens 3.4-3.6]; eye in head 3.0 (2.9-3.1); snout
in eye 1.0 in all specimens examined; least width of bony interorbital
in greatest eye diameter 1.0 in all specimens.

Premaxillary with 7 quinquecuspid teeth, lateroposterior cusp
largest. Maxillary having 1 tooth with 4 or 5 cusps. Dentary with
8 sexcuspid teeth, cusps subequal and central cusps longest; 11 teeth
in second row of dentary in alizarin specimens. Gill rakers 9+ 13.

Dorsal fin rays ii,8; anal iii,9; pectoral i,9 to i,10; pelvics iii,7.
Adipose fin present in all specimens examined but reduced in size in
some. Caudal fin with principal rays 10/9; distal tips of all elements
of first 10 principal rays in upper caudal lobe. Lower caudal lobe
with all elements of remaining 9 principal rays.

Figure 7.—Nannostomus trifasciatus, male, SL 27.5 mm., SU 36899,

Anal fins of male and female appear alike in this species; rays of
male’s anal fin not modified. Inclinator muscles appear slightly, if
at all, more developed in male than in female.

Radial grooves of scales belonging to third horizontal scale series
and lying below dorsal fin are shown in figure 7. Posterior field with
1 or 2 radial grooves. Dorsal and ventral fields each with 1 radial
groove and anterior field with 2 to 4 radial grooves. Scales in a
lateral series 26 to 27; 3 to 5 lateral-line scales and 10 scales in median
series anterior to dorsal fin. Total number of vertebrae 35 to 36,
19 precaudal and 16 or 17 caudal vertebrae.

Cotor IN ALcoHoL.—Top of head light to dark brown. Primary
horizontal stripe extending on head from snout tip and lower jaw
posteriorly over first orbital bone across eye and operculum, termi-
nating at posterior border of operculum. On body, primary horizontal
stripe extending from under fleshy flap of opercle through fourth scale

30 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

row and partially on adjoining third and fifth scale rows; expanding
over anal fin to include one-half of third scale row and on caudal
peduncle extending downward to all but the lowermost vertical scale
row. Primary horizontal stripe ending on caudal peduncle in Peru-
vian specimens but extending posteriorly onto caudal fin as a very
narrow wedge on most specimens from British Guiana.

Above primary horizontal stripe, pale stripe extending from eye to
caudal peduncle. Melanophores scattered over entire surface of this
stripe being thicker and heavier in region of fifth through eleventh
and sixteenth to twentieth vertical scale rows. These areas consti-
tute oblique bands. Secondary horizontal stripe dorsal to silvery
lateral stripe, and extending from posterior upper end of gill cover
through second row of scales onto caudal peduncle. Secondary hori-
zontal stripe darker and wider at posterior margin of each scale in
second row. Back above secondary horizontal stripe dark, rather
densely covered with melanophores. Sides below primary horizontal
stripe lightcolored with few scattered melanophores. On sixth scale
row tertiary horizontal stripe extending from below operculum along
sides of belly, where extending over or interrupted by pelvic fins, and
then continuing to posterior end of base of anal fin. Tertiary hori-
zontal stripe narrow and sometimes absent or restricted to region
behind pelvic fins. Portion of tertiary horizontal stripe on head
along surface of third orbital bone. Pectoral and pelvic fins without
dense aggregations of melanophores. Sometimes rather weak aggre-
gation of melanophores present along uppermost and lowermost
principal caudal fin rays. Dorsal fin dusky on its anterior distal
portion and first and second unbranched rays frequently dark. Anal
fin of male and sometimes female dusky along its distal and anterior
regions. Sometimes this duskiness not evident.

Cotor IN LiFE.—The color description given below agrees well
with that given by Eigenmann (1909, p. 45) for live specimens of
Poecilobrycon erythrurus from British Guiana. The specimens used
here were collected in British Guiana by Louis Chung in 1955 or 1956
but are without definite locality. Midanterior region of dorsal fin with
single bright red spot. Red spots on base of upper and lower lobes
of caudal fins and bright red spots on anterior base of anal fin and
proximal region of pelvic fin. In some specimens these red spots
almost absent. Location of these spots well shown in color photograph
by Innes (1953, p. 192). Photograph in Axelrod and Schultz (1955, p.
248) shows color of this species well. Primary horizontal stripe metal-
lic pale green-gold. Caudal fin posterior to metallic pale green-gold
stripe with few streaks of white pigment. Thin red line extending
lengthwise through center of metallic pale green-gold stripe. Fre-
quently this red line broken into series of streaks, reduced in length or

NO. 3538 CHARACID FISHES—WEITZMAN 31

absent. Laterodorsal and anterior surface of snout anterior to nostrils
bright red. Also single red line in upper part of iris. Remainder of
iris pale gold except where primary horizontal stripe extends through
eye. Distal region of pelvics silvery blue, especially in male. In some
specimens this color almost absent. Belly and region below primary
horizontal stripe silvery white.

RemarKks.—Poecilobrycon vittatus of Ah] (1933) was described very
poorly and the type locality is indefinite. Ahl, following Eigenmann,
seemed confused about the identity of Steindachner’s trifasciatus and
eques, even though, for their time, Steindachner’s figures and descrip-
tions of these two species were excellent. Ahl believed his vittatus was
related to Eigenmann’s erythrurus, a form which perhaps could be
recognized at most as a subspecies of trifasciatus. For the most part,
Ahl’s description of vittatus seems to fit the characters listed here for
trifasciatus, and I concur with Hoedeman (1954) in considering vittatus
a synonym of trifasciatus.

Kigenmann (1909, pp. 44-45) described Poecilobrycon erythrurus
and noted that it was most closely allied to his own P. marginatus
(now considered to be Nannostomus marginatus) and Steindachner’s
P. trifasciatus. Later Eigenmann (1912, p. 284) considered Poecilo-
brycon auratus to be synonymous with Steindachner’s Nannostomus
trifasciatus. He apparently had mentally transposed Steindachner’s
trifasciatus and eques. This probably caused Eigenmann to consider
that his erythrurus (at most a subspecies of trifasciatus) was not tri-
fasciatus and therefore that it was a distinct species. In any event,
examination of some of Eigenmann’s specimens of erythrurus and of
specimens of trifasciatus from near the type locality of the latter
indicates that the two forms are closely related and can be considered
members of a widespread, somewhat polymorphic species. Adequate
study of many samples from populations throughout the range of the
species probably will show that there are several distinct races, some
of which undoubtedly will be considered as subspecies. Schultz in
Axelrod and Schultz (1955, p. 248) considered Poecilobrycon auratus
Eigenmann a synonym of Nannostomus trifasciatus. Examination of
type specimens of auratus shows it to be a synonym of Poecilobrycon
eques (Steindachner) (p. 50).

SPECIMENS EXAMINED.—Three, SU 36900 (SL 24.5 mm.), SU 36999 (SL 27.0
mm.), SU 50221 (SL 27.0 mm.), Peru, Shansho Cano, Pévas [Pébas] District, July
28, Oct. 20, 1936 (date for third specimen unknown), W. G. Scherer.—One, SU
50222, SL 33.0 mm., Peru, Rio Ampiyacu, Pévas [Pébas] District, July 23, 1944,
W. G. Scherer.—Two (paratypes of Poecilobrycon erythrurus Eigenmann), CNHM
52974, SL 26.5-28.2 mm., British Guiana, Rockstone River, 1908, Carl H. Eigen-
mann.—Two (paratypes of P. erythrurus), CAS(IUM) 11693, SL 23.0-28.6 mm.,
British Guiana, Rockstone sandbank, Essequibo River, 1908, Carl H. Eigen-
mann.—One (paratype of P. erythrurus), CAS(IUM) 11694, SL 24.0 mm., British

32 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Guiana, Gluck Island, Essequibo River, 1908, Carl H. Eigenmann.—One (para-
type of P. erythrurus), CAS(IUM) 11695, SL 25.5 mm., British Guiana, Rupununi
River, Twoca Pan, 1908, Carl H. Eigenmann.—One, CNH M 50162, SL 27.0 mm.,
British Guiana, Rockstone, Essequibo River, Mar. 1937, E. R. Blake.—Five,
SU 50223 (one alizarin preparation), SL 25.0-30.0 mm., British Guiana, no other
data.—Two, SU 18472 (one alizarin preparation), SL 31.5-32.0 mm., aquarium
specimens said to be from ‘‘the Amazon,” sent to Dr. G. 8. Myers by Frederick
Stoye, no other data.—Seven, SU 50220, SL 29.5-32.5 mm., aquarium specimens
probably from British Guiana, 1955.—Two, SU 50434, SL 21.0-22.1 mm., British
Guiana, Moreby Creek, Rupununi River near Karanambo, Essequibo drainage,
Sept. 17, 1957, Rosemary Lowe (McConnell) .—Two, SU 50437, SL 23.2-24.8 mm.,
British Guiana, Karanambo, “‘Grass Pond,’”’ Rupununi River, Essequibo drainage,
Sept. 14, 1957, Rosemary Lowe (McConnell).—Two, SU 50432, SL 24.0-26.0
mm., British Guiana, ‘Crane Pond Creek,’? Karanambo Rupununi River, Esse-
quibo drainage, Sept. 14, 1957, Rosemary Lowe (McConnell).—Two, SU 50440,
SL 19.0-27.0 mm., British Guiana, Rupununi District, Nappi River and other
Tocati drainage creeks, Rio Branco drainage, Sept. 20, 1957, Rosemary Lowe
(McConnell).

Nannostomus marginatus Eigenmann

Figure 8

?Cyprinodon amazona Kigenmann, 1894, p. 627 (in part; original description; type
locality: Lower Amazonas; types apparently lost)—Garman, 1895, p. 28
(in part; copied description).—Hubbs, 1926, p. 16 (in part; discussion of
identity).—Fowler, 1954, p. 216 (in part; listed).

Nannostomus marginatus EKigenmann, 1909, p. 41 (original description; type
locality: British Guiana, Maduni Creek, Mahaica River; holotype in Chicago
Natural History Museum).—Eigenmann, 1910, p. 427 (listed) ; 1912, p. 281,
pl. 37, fig. 4 (description).—Innes, 1935, p. 154 (description from life and
photograph; see also later editions)—Hoedeman, 1950, p. 20 (description;
figures).—Boeseman, 1952, p. 184 (Surinam); 1953, p. 16 (Surinam); 1954,
p. 18 (Surinam).—Axelrod and Schultz, 1955, p. 241 (description and
aquarium notes).—Sterba and Tucker, 1963, p. 211 (description and aquarium
notes).

?Cyprinodon amazonus Eigenmann, 1910, p. 456 (in part; listed).

Nannostomus marginatus picturatus Hoedeman, 1954a, p. 87 (original description;
type locality: ‘‘A ditch near Zonderij II, Surinam’; types in Amsterdam
Museum).

DiaGnosis.—This species is the deepest bodied of all nannosto-
minans; only egg-filled females of Nannostomus beckfordi were found to
have a body depth as great as the most slender specimens of marginatus.
The body depth of marginatus is 3.5 to 4.0. Their caudal peduncle is
the deepest of all nannostominans, 7.5 to 8.5 in standard length. In
addition, marginatus may be distinguished from all other nannosto-
minans by the following combination of characters: Primary, second-
ary, and tertiary horizontal stripes all present; maxillary tooth rarely
present; 21 to 23 scales in a lateral series; 7-++12 gill rakers; 31 to 33
vertebrae; male anal fin with unmodified fin rays.

DescripTion.—Body elongate, robust; sides flattened and body

NO. 3538 CHARACID FISHES—WEITZMAN 33

compressed posteriorly. Greatest depth between posterior tip of
appressed pectorals and anterior base of dorsal fin. Standard length
of largest specimen examined 25.5 mm. Greatest body depth 3.8
(3.5-4.0) ; least depth of caudal peduncle 8.0 (7.5-8.5) ; length of caudal
peduncle (including both sexes) 5.7 (5.3-6.2); snout tip to origin of
dorsal fin 1.8 (1.8-1.9); snout tip to origin of anal 1.3 (1.2—1.3).

Head and snout obtuse in vertical and horizontal profiles, top of
head convex between eyes. Head 3.4 (3.2-3.6); eye in head 2.8 (2.7—
2.9); snout in eye 1.3 (1.3-1.4); least width of bony interorbital in
ereatest eye diameter 1.0 (1.0—-1.1).

Premaxillary with 6 to 7 teeth, each quinque- to tricuspid, sometimes
most anterior tooth with only 4 cusps; lateral cusps largest. In large
specimens, largest cusp just medial to lateralmost cusp; cusps subequal
in some specimens. Only 1 specimen, SU 50219 from Brazil, with any
maxillary teeth; this specimen with 1 conical tooth on 1 side only.
Dentary with 7 to 8 quinque- or sexcuspid teeth in first outer row.
Most posterior tooth in dentary conical, bicuspid, or tricuspid. Speci-
mens from Brazil (except specimens from Obidos) having dentary with
quinquecuspid teeth, central cusp being largest. Specimens from
British Guiana with lower jaw teeth having either 6 subequal cusps or
5 cusps with a large central cusp. Smallest Brazilian specimen with
lower jaw teeth having almost subequal cusps, while some of largest
specimens from British Guiana with very large central cusps on
quinquecuspid teeth. Specimens from Obidos with 6 subequal cusps
on lower jaw teeth and 5 subequal cusps on upper jaw teeth. Teeth
6 to 7 in second tooth row of dentary. Gill rakers 7+ 12 in all alizarin
preparations.

Dorsal fin rays ii,8; anal iii,9; pectoral 1,10 to 1,13; pelvics 1,7.
Adipose fin absent in all specimens examined. Caudal fin with princi-
pal rays 10/9; distal tips of all ray elements of first 10 principal rays in
upper caudal lobe. Lower caudal lobe containing all elements of
remaining 9 principal rays.

Anal fin of male and female alike. Rays of male’s anal fin not
modified. Inclinator muscle appears slightly, if at all, more developed
in male than in female.

Radial grooves of scales in third horizontal scale series lying below
dorsal fin shown in figure 8. Posterior field with 1 to 3 radial grooves.
Dorsal and ventral fields each with 1 radial groove and anterior field
with 2 to 4 radial grooves. Scales 21 to 23 in lateral series in all
specimens examined and 3 to 5 lateral-line scales (usually 4); 10 scales
in median predorsal series. Total number of vertebrae 31 to 33. Of
35 specimens from localities in Brazil, 32 with 32 vertebrae, 2 with 31
vertebrae and 1 with 33 vertebrae. Of 15 specimens from localities
in British Guiana 9 specimens with 31 vertebrae, 1 with 33 vertebrae

219-936—66——3

34 PROCEEDINGS OF THE NATIONAL MUSEUM you. 119

and 5 with 32 vertebrae. Precaudal vertebrae 17 to 19 and caudal
vertebrae 13 to 14, never in a combination over 33 or under 31 total
vertebrae.

CoLor IN ALcoHOL.—(Specimens from British Guiana). Dorsum
of head light to dark brown. Primarily horizontal stripe beginning
on lower part of snout tip and tip of lower jaw, extending to pupil of
eye, then passing over posterior part of iris, across fourth and fifth
orbitals to posterior edge of operculum. Primary horizontal stripe
then continuing from under fleshy opercular flap posteriorly over lower
portion of third scale row and upper half of fourth scale row to termi-
nate as narrow black wedge on caudal fin. On caudal peduncle,
primary horizontal stripe expanding downward to cover all but
lowermost ventral scale row.

Ficure 8.—Nannostomus marginatus, male, SL 19.5 mm., CAS(IUM) 11700 (see text for
discussion of teeth).

Pale silvery stripe extending from eye onto caudal fin above primary
horizontal stripe. Melanophores scattered over entire surface of this
silvery stripe, thickest and heaviest in region of third through about
eighth and thirteenth or fourteenth to sixteenth vertical scale rows.
These darker areas representing oblique bands, one anterior, one pos-
terior to dorsal fin. Silvery band most pale over hypural fan and on
caudal fin between wedge-shaped portions of primary and secondary
horizontal stripes. Secondary horizontal stripe dorsal to pale lateral
stripe. Secondary stripe extending above opercular bone through
upper part of second lateral scale row onto caudal fin, terminating on
this fin as an acute wedge. Dorsum dark above secondary horizontal
stripe, rather densely covered with melanophores, especially behind
dorsal fin. Below primary horizontal stripe, sides white to very pale

NO. 3538 CHARACID FISHES—WEITZMAN 35

brown with few scattered melanophores. ‘Tertiary horizontal stripe
extending along sides of belly through sixth lateral scale row and across
head over preopercle and third orbital bone to angle of jaw. Some-
times this stripe reduced in region of pectoral fins. Pectoral fin
translucent; pelvic fin same except for thin line of melanophores along
anterior 2 rays in some specimens. Dorsal fin dusky at its anterior
base, its first two unbranched rays dark. Almost entire lower half
of anal fin dark brown to black; its distal edge and upper edges also
dark.

Coor IN LIFE.—The color of aquarium specimens which are from
an unknown locality, but which agree with Eigenmann’s (1909, p. 42)
brief color description of his material from British Guiana, is as
follows: Midanterior region of dorsal fin with one red spot, ventral
region of anal fin red; medial half of pelvic fins red. Color photo in
Axelrod and Schultz (1955, p. 241) shows position of this red pigment
well. A short red stripe along midupper border of primary horizontal
stripe (which is black in life) ; half of this stripe on primary horizontal
stripe and half on pale golden stripe above black horizontal stripe.
This red stripe extending from about sixth to ninth vertical scale
rows. <A few streaks of white pigment at posterior end of pale gold
band above primary horizontal stripe. This white pigment mostly
confined to caudal fin rays between wedges of primary and secondary
horizontal stripes. Lateral surface of the snout pale gold. Iris pale
gold where melanin absent. Belly and region below primary hori-
zontal stripe silvery white. Distal anterior parts of pelvics white.

Aquarium specimens which agree well with specimens recorded
below from the lower Amazon have the following color: Red of fins
almost absent except on dorsal fin. Red stripe on side of body absent.
Secondary and tertiary horizontal stripes somewhat broader and more
sharply delineated. Stripe above primary horizontal stripe not as
golden as in presumed British Guiana specimens. Belly and area
below primary horizontal stripe brownish white. White spot on
caudal fin between wedges of primary and secondary horizontal
stripes more intense.

Living specimens collected by White, Reynolds, and Wulff (see list
of specimens below) from Colombia near Tres Esquinas have the
following color: Proximal two-thirds and often entire length of fourth
to sixth dorsal fin rays bright orange red. First 3 dorsal rays black,
seventh to tenth rays and adjoining tissue hyaline. Middle rays of
caudal fin red between dark caudal wedges (this region occupied by
intense white in living specimens presumably from lower Amazon).
Pelvic fins red except for anterior black border. Anal fin from its
middle and posterior base out to one-third or three-quarters of its
distal surface varying from silvery to golden yellow. Anterior base

36 PROCEEDINGS OF THE NATIONAL MUSEUM yoL. 119

of pelvic fin and distal portion of its anterior 4 to 5 rays bright red.
Distal tips of second to about fourth pelvic rays black. Belly silvery;
area below primary horizontal stripe bright gold. Entire length above
primary horizontal stripe with narrow iridescent gold or sometimes
silvery-red stripe. Above this with a greenish dark-brown area, same
color as top of head and dorsum. Primary, secondary, and horizontal
stripes black. Iridescent golden to pale reddish spot near anterior
nostril.

Remarks.—Nannostomus marginatus has been reported only from
the Guianas. It now appears to be a widespread polymorphic species
found in the Guianas and in at least several widely separated areas in
the Amazon Basin. Hoedeman’s subspecies picturatus from Surinam
is described very inadequately and some of the description appears to
be in error. For example he reported a difference in caudal fin ray
counts between the British Guiana and Surinam populations. He
stated the caudal rays to be 10+10 in specimens from British Guiana
and 8 to 9+9 to 8 in specimens from Surinam. All specimens of nan-
nostominans and indeed all characids that I have examined, except for
a few obviously abnormal specimens of the genera Hyphessobrycon and
Moenkausia, have had a principal caudal fin ray count of 10/9 (equals
10+9). JI am not sure whether Hoedeman counted principal caudal
fin rays or only unbranched rays. However, he reported the wrong
count for either method. It should be noted that in counting the prin-
cipal caudal rays of two of Hoedeman’s types of N. bifasciatus, the
typical count of 10/9 was found. Hoedeman (1954a) reported 8(7)-++
8(7). Despite the probable errors in some of Hoedeman’s counts, and
possibly also his measurements, probably it will be found eventually
that his subspecies picturatus is valid. Also the populations noted
here from lower and upper Amazon Basin very probably will, when
adequate material becomes available, prove to be sufficiently different
to merit subspecific recognition. An example of the difference in
teeth of different populations can be seen in figure 8. The two
opposing teeth to the right are from British Guiana specimens [CAS
(LUM) 11700] while those to the left are from Lagéa Grande, Brazil
(SU 50219).

SPECIMENS EXAMINED.—Four, paratypes, CNHM 52776, SL 17.0-18.5 mm.,
Brutish Guiana, Maduni Creek, Mahaica River, 1908, Carl H. Eigenmann:—
Two, CNHM 54932, SL 19.4-20.3 mm., British Guiana, Hubabu Creek, Demerara
River, Oct. 1, 1910, Max Ellis —One, paratype, SU 21941, SL 17.6 mm., British
Guiana, Cane Grove Corner, Mahaica River, 1908, Carl H. Eigenmann.—Two,
paratypes, CAS(IUM) 11697, SL 16.0-16.4 mm., British Guiana, Lama Stop-Off,
Mahaica River, 1908, Carl H. Eigenmann.—One, paratype, CAS(IUM) 11698,
SL (damaged), British Guiana, Crab Falls, Essequibo River, 1908, Carl H.
Eigenmann.—One, paratype, CAS(IUM) 11699, SL 19.5 mm., British Guiana,
Rockstone sandbank, Essequibo River, 1908, Carl H. Eigenmann.—Two, para-

NO. 3538 CHARACID FISHES—WEITZMAN 37

types, CAS(IUM) 11700, SL 19.5-21.8 mm., British Guiana, Gluck Island,
Essequibo River, 1908, Eigenmann.—One, paratype, CAS(IUM) 11701, SL
(damaged), British Guiana, Christianburg Canal, Demerara River, 1908, Eigen-
mann.—Five, paratypes, CAS(IUM) 11702 (one alizarin preparation), SL 17.5-
18.0 mm., British Guiana, Cane Grove Corner, Mahaica River, 1908, Eigenmann.—
Thirty-two, SU 50219 (three alizarin preparations), SL 15.0-22.8 mm., Brazil,
State of Pard, Lagéa Grande, Igrapé do Meritysdl (probably Lagéa Grande do
Javafy about 34 miles northwest of Santarém), Aug. 20, 1924, Carl Ternetz.—
Four, SU 50218, SL 16.4-19.5 mm., Brazil, State of Pard, Igrapé Uruard4, tribu-
tary of the Furo Maica about 10 miles southeast of Santarém, June 26, 1924,
Carl Ternetz—Two, MCZ 19645, SL 23.5-25.4 mm., Brazil, State of Pard,
Obidos, 1865, Col. Bentos, Thayer Expedition—Two, SU 50217, SL 20.0-
22.0 mm., Brazil, State of Parad, Lagéa Grande (probably Lagéa Grande do
Javafy about 34 miles northwest of Santarém), a stream into Ajamurf beach,
July 17, 1924, Carl Ternetz.—One, SU 50555, SL 18.0 mm., Colombia, Caquet4
Province, swampy pond about one-fourth mile inland from Rio Orteguaza, across
the river and slightly below Tres Esquinas, lat. 0°54’ N. and long. 75°15’ W.,
Feb. 8, 1958, 8-10 a.m., Gen. T. D. White, Col. J. N. Reynolds, Lee Wulff, and
Dr. George 8. Myers.—One, SU 50669, SL 16.5 mm., Colombia, Caquet4 Prov-
ince, small stream across Rio Orteguaza from Tres Esquinas, lat. 0°45’ N., long.
75°15’ W., Feb. 13, 1958, Gen. T. D. White, Col. J. N. Reynolds, Lee Wulff and
Dr. George S. Myers.—Twenty-six, SU 50628, SL 16.0-22.1 mm., Colombia,
Caquetd4 Province, small forest tributaries of Rio Orteguaza on road from Tres
Esquinas to Solano, lat. 0°45’ N., long. 75°15’ W., Feb. 12, 1958, Gen. T. D.
White, Col. J. N. Reynolds and Lee Wulff.—Six, SU 50457, SL 18.0-18.4 mm.,
British Guiana, Ishalton, southern Rupununi District, Rupununi River drainage,
tributary of Essequibo River, Dec. 12, 1957, Rosemary Lowe (McConnell).—
Two, SU 50445, SL 17.5-19.6 mm., British Guiana, Atkinson Field Creek, near
Hyde Park, tributary of Demerara River, Nov. 11, 1957, Rosemary Lowe
(McConnell).—Eight, SU 50451, SL 16.0-17.0 mm., British Guiana, Kumarow
Creek pool, tributary of Rupununi River in southern savannas, Essequibo River
drainage, Dec. 10, 1957, Rosemary Lowe (McConnell).—Twenty-six, SU 50452,
SL 16.2-18.7 mm., British Guiana, creek approximately 5 mi. south of Lumidpau,
southern Rupununi savannas (flows into Rupununi River), Essequibo drainage,
Dec. 11, 1957, Rosemary Lowe (McConnell).—Twenty-five, SU 50441, SL 15.7-
19.5 mm., British Guiana, Rupununi District, Nappi River and other Tacatt
drainage creeks, Rio Branco drainage, Sept. 20, 1957, Rosemary Lowe (McCon-
nell).—Five, SU 50428, SL 16.5-19.0 mm., British Guiana, Manawarin River,
tributary of the Moruka River about lat. 7°30’ N. and long. 59°00’ W., July 16,
1957, Rosemary Lowe (McConnell).

Genus Poecilobrycon Figenmann

Poecilobrycon Eigenmann, 1909, p. 43 (type species Poecilobrycon harrisoni
Kigenmann, 1909, by original designation).

Archicheir Kigenmann, 1909, p. 46 (type species Archicheir minutus Eigenmann,
1909, by monotypy).

Nannobrycon Hoedeman, 1950, p. 22 (type species Nannostomus eques Steindachner,
1876, by original designation).

Dracnosis.—The following two characters appear unique for this

genus: Second infraorbital bone with a bony canal for infraorbital
branch of latersensory canal. In addition, snout long, 0.8 to 1.2

38 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

in eye diameter. The members of this genus have, correlated with the
long snout, a longer, narrower ethmoid and prevomer, a longer ant-
orbital and infraorbital, and a longer mesopterygoid and nasal bone
than do members of the genus Nannostomus. Primary and secondary
horizontal stripes are present in all species, but a tertiary horizontal
stripe is absent in 2 species. The anal fin rays may or may not be
modified in males. The body is elongate in all species, the depth
being 4.5 to 5.7 in standard length.

The name Poecilobrycon is derived from the generic names Poeciia
and Brycon, the implied meaning being a Poecilia-like Brycon, i.e., a
Brycon-like fish or characid with characters of the poeciliid fishes.
This name probably occurred to Eigenmann because of the small mouth
and numerous tooth cusps in Poecilobrycon. However, in detail, the
mouth of nannostominans bears little resemblance to that of any
poeciliid fish.

Subgenus Poecilobrycon Kigenmann

DiaGnosis.—As in all members of Nannostomus, 10 principal rays
of upper 4 hypural bones all enter upper caudal lobe. Lower lobe of
caudal fin with dark pigment only at its base. Normal resting and
swimming position horizontal. See under the subgenus Nanno-
brycon for a discussion of this character.

Poecilobrycon harrisoni Eigenmann
Fiaures 9, 10

Poecilobrycon harrisoni Kigenmann, 1909, p. 43 (original description; type locality:
British Guiana, Canal at Christianburg, Demerara River; holotype in Chicago
Natural History Museum); 1910, p. 427 (listed); 1912, p. 284, pl. 37, fig. 1
(description; British Guiana) —Hoedeman, 1950, p. 22 (copied descrip-
tion); 1954b, p. 89 (description; British Guiana).—Weitzman, 1964, p. 136
(osteology and relationships).

Archicheir minutus Eigenmann, 1909, p. 46 (original description; type locality:
British Guiana, Canal at Christianburg, Demerara River; holotype in Chicago
Natural History Museum); 1910, p. 427 (listed); 1912, p. 287, pl. 37, fig. 5
(description; British Guiana).

?Nannostomus kumuni Ladiges, 1948, p. 30 (name only listed, nomen nudum).

?Nannostomus cumuni Arnold, 1950, p. 63 (original description, from life, no type
specimens).

Diacnosis.—This species may be distinguished from all other
nannostominans by its greater number of vertebrae, 38 to 39, con-
trasted to 31 to 36. It may be recognized further by the following
combination of characters: Second tooth row of dentary with 12 to
15 teeth; 7+13 gill rakers; iii,9 anal fin rays; 27 to 30 scales in lateral
series; maxillary tooth present; fairly long caudal peduncle, 5.3-6.1
in standard length.

NO. 3538 CHARACID FISHES—WEITZMAN 39

DescripTION.—Body elongate, slender, sides only slightly flattened
anteriorly and caudal peduncle compressed. Greatest depth lying
just anterior to anterior base of dorsal and anal fins. Standard
length of largest specimen 44.5 mm. One old aquarium specimen
reached a standard length of 44.0 mm. after living 2% years. Greatest
body depth 5.4 (4.9-5.7); least depth of caudal peduncle 10.9 (10.4—
12.1); length of caudal peduncle (including both sexes) 5.7 (5.3-6.1);
snout tip to origin of dorsal fin about 1.9 in all adult specimens; snout
tip to origin of anal fin about 1.3 in all adult specimens.

Head elongate and somewhat depressed. Top of head very slightly
convex between eyes. Snout rounded in horizontal and vertical
profiles. Head 4.0 (3.8-4.1); eye in head 3.2 (3.0-3.2); snout in eye
0.9 (0.8-1.0) ; least width of bony interorbital in greatest eye diameter
1am 1021.2),

Ficure 9.—Poecilobrycon harrisoni male, SL 38.5 mm., SU 50243.

Premaxillary with 7 quadri- to septemcuspid teeth. Most with 5
or 6 cusps; cusps subequal or posterolateral cusps most prominent.
Maxillary with 1 conical, bicuspid, or tricuspid tooth. Dentary with
7 to 8 (usually 8) teeth in first row. Posterior 2 teeth with 3 to 5
subequal cusps, remainder of teeth in first row with 6 subequal cusps.
Second dentary tooth row of about 12 to 15 small conical teeth. Gill
rakers 7-+-13 in alizarin preparations.

Dorsal fin rays 1,8; anal i1,9; pectoral i,10 to i,11; pelvic fin rays
ii,7. Adipose fin varying from well developed to rudimentary or
absent. Caudal fin with principal rays 10/9; distal tips of all ray
elements of first 10 principal rays included in upper lobe. Lower
caudal lobe with all elements of remaining 9 principal rays.

Radial grooves of scales in median series below dorsal fin shown in
figure 9. Posterior field with 1 to 4, dorsal and ventral fields 1, and
anterior field 2 to 5 or 6 radial grooves. Scales in a lateral series 27

40 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

to 30 (usually 29). lLateral-line scales 3 to 5, usually 4. Scales in
a median series in front of dorsal fin 11 to 13, usually 12. Total
number of vertebrae 38 to 39 with 21 precaudal and 17 to 18 caudal
vertebrae.

CoLor IN ALCOoHOL.—Specimens from British Guiana, SU 50446,
have the following colors: (The paratypes are coated rather heavily,
apparently with lead oxide from a metal number tag, but agree well
with the description below.) Top of head light to dark brown.
Primary horizontal stripe extending from snout tip and lower jaw
posteriorly over maxillary, first and second orbitals, and onto eye.
From posterior part of eye, primary horizontal stripe extending to
posterior edge of opercular bone. On body, primary horizontal
stripe continuing from under fleshy opercular flap posteriorly through
fourth scale row and part of fifth to caudal peduncle and caudal fin;
on caudal peduncle extending over ventral body surface. On caudal
fin, primary horizontal stripe not taking form of a narrow wedge but
expanding to cover upper part of lower caudal lobe.

Above primary horizontal stripe a pale stripe extending from snout
tip to caudal base. Dark area on sixth through eight vertical scale
rows. Aggregation of melanophores on pale stripe vertically above
anal fin. Secondary horizontal stripe apparently absent; entire
dorsum above middle of second horizontal scale row light brown.
Belly silvery, but posterior to pelvic fins in males becoming dusky.
Pectoral fins translucent. Dorsal fin slightly dusky anterodistally
but otherwise translucent and pelvics clear except for dusky distal
band. Upper lobe of caudal fin slightly dusky. Anal fin translucent;
in some specimens last ray black.

Cotor IN LiFE.—The colors of aquarium specimens, SU 50246,
from Demerara River, British Guiana, are as follows: Dorsal and
pectoral fins hyaline. Pelvic fin of male with proximal one-third to
three-fourths of fourth to ninth pelvic rays orange red. These fins
hyaline in female. Distal one-third of third through fourth pelvic
rays and adjacent membrane silvery blue in both sexes. Anal fin
of female hyaline with occasionally one spot of silvery blue on distal
one-fourth of fourth anal finray. Male also with silvery blue disposed
in same manner but proximal one-third to one-half of all anal fin
rays bright red. Proximal portions of caudal fin lobes red above and
below extension of primary horizontal stripe on fin rays. These
areas silvery yellow with faint indications of red infemales. Primary
horizontal stripe black. Above this stripe a narrow silvery band and
above this body shades to light or sometimes brown on dorsum.
Metallic-red spot near anterior narial opening. Belly and sides
below primary horizontal stripe white.

Remarxs.—So far as is known Poecilobrycon harrisoni is restricted

NO. 3538 CHARACID FISHES—WEITZMAN 41

to British Guiana. All the aquarium specimens that I have seen
over the past 10 years have had approximately the same color pattern.
Thus it appears that aquarium fish collectors are collecting all their
specimens from the same general area. Very probably this is from
the vicinity of Georgetown in tributaries of the Demerara River,
British Guiana, where the fishes are known to occur and where col-
lectors are known to have been active in the last several years. In
many respects, the color and breeding habits of Poecilobrycon harrisoni
are like those of Nannostomus beckfordi suggesting that the two may
have been derived from a common ancestor and may be related more
closely than their morphology indicates.

The type of Archicheir minutus is the young of Poecilobrycon
harrisont. ‘The type was studied at the Chicago Natural History
Museum and found identical with young specimens of harrisoni
spawned and raised in the author’s aquarium.

SPECIMENS EXAMINED.—T wo, paratypes, CAS(IUM) 11709, SL 36.5-41.5 mm.,
British Guiana, Canal at Christianburg, Demerara River, 1908, C. H. Eigen-
mann.—Four, SU 50243, SL 34.0-38.0 mm., British Guiana, Georgetown.—
Twelve, SU 50245 (one alizarin preparation), SL 31.0-37.5 mm., British Guiana,
no other data.—Three, SU 50244, SL 13.3-44.5 mm., aquarium specimens,
Aug. 21, 1956, 8S. H. Weitzman.—Eight, SU 50246, SL 9.5-37.5 mm., aquarium
specimens from Demerara River, British Guiana, and their young aquarium
spawned and raised, 8. H. Weitzman.—Seven, SU 50446, SL 17.0-44.5 mm.,
British Guiana, Atkinson Field Creek, near Hyde Park, tributary of the Demerara
River, Nov. 11, 1957, Rosemary Lowe (McConnell).

Subgenus Nannobrycon Hoedeman

Draanosis.—Rays of caudal fin arranged unlike those of all other
nannostominans; 10 rays of upper 4 hypural plates not all entering
upper caudal lobe. Lowermost 2 of these rays enter lower caudal
lobe, giving that lobe greater depth than upper lobe. Lower caudal
lobe usually with dark pigment extending well onto fin, sometimes
over its entire surface. Normal resting position of living members of
this subgenus oblique, snout upwards. Angle of this oblique position
is from about 45° to about 75° or 80°.

Remarxks.—Hoedeman (1950) proposed Nannobrycon as a generic
name for Poecilobrycon eques (Steindachner). In it he included only
this species, believing P. wnifasciatus to be a synonym of P. eques.
Both species of this subgenus have the same peculiar caudal fin
structure and behavior mentioned above. Hoedeman (1950) noted
that the shape of the swim bladder of the members of this subgenus is
different from those of all other nannostominans that he examined.
However, he did not examine the swim bladder of Poecilobrycon
harrisoni, as he had no specimens. The swim bladder differences that
Hoedeman noted were crudely presented and are not suitable to
distinguish harrisonit from eques and unifasciatus. However, the

219-936—66——4

42 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

normal resting position of members of the subgenus Nannobrycon is so
different from that of the single known member of the subgenus
Poecilobrycon, and indeed from all other known nannostominans, that
there is undoubtedly a difference in shape correlated with this swim-
ming position. Critical morphological examination of the organs of
equilibrium, the swim bladder, and musculature of the caudal fin of
all nannostominans will possibly show sufficient differences for
Nannobrycon to be retained as a full genus. Braemer and Braemer
(1958) have started a project studying the behavior and position of
the utricular statolith with reference to Poecilobrycon eques and other
characids.

Figure 10.—Anal fin of Poecilobrycon harrisoni, SL 38.5 mm., SU 50243.

Poecilobrycon unifasciatus (Steindachner)
Figure 11

Nannostomus unifasciatus Steindachner, 1876, p. 127, pl. 9 (original description;
localities: Mouth of the Rio Negro and Teffé on the Solimoens; types in
Vienna Museum).—Eigenmann and Eigenmann, 1891, p. 49 (listed); 1910,
p. 427 (listed)—Sterba and Tucker, 1962, p. 214 (description and aquarium
notes; photographs are of P. harrisoni; also part of color description seems
to refer to this species).

Poecilobrycon ocellatus Eigenmann, 1909, p. 45 (original description; type locality:
British Guiana, Wismar, Demerara River, holotype in Chicago Natural
History Museum) ; 1910, p. 427 (listed) ; 1912, p. 286, pl. 37, fig. 4 (description).

Poecilobrycon unifasciatus—Innes, 1935, p. 158 (description from life and photo-
graph).—Axelrod and Schultz, 1955, p. 249 (description and aquarium notes).

Poecilobrycon eques (in part) —Hoedeman, 1950, p. 23.

Nannostomus ocellatus—Sterba and Tucker, 1962, p.214 (description and aquarium
notes).

Diaanosis.—This species may be distinguished from all other
nannostominans examined by the complete absence of a second row

No. 3538 CHARACID FISHES—WEITZMAN 43

of teeth in the dentary; in addition, it may be distinguished by the
following combination of characters: Adipose fin present; 33 to 34
vertebrae; 2 to 5 perforated scales in the lateral line; iii,9 anal fin
rays; 9+14 gill rakers and 28 to 30 scales in a lateral series.

DescripTion.—Body elongate, slender, and somewhat cylindrical
anteriorly, but sides somewhat flattened; compressed posteriorly.
Greatest depth anterior to dorsal fin and posterior to distal tips of
appressed pectoral fins. Standard length of largest specimen 38.3
mm. Greatest body depth 5.0 (4.7-5.3); least depth of caudal
peduncle 10.4 (10.0-11.2); length of caudal peduncle (including both
sexes) in specimens listed below from Brazil 6.9 (6.7—7.5) and from
British Guiana 6.2 (6.0-6.6); snout tip to origin of dorsal fin 1.9
(1.8-1.9) ; snout tip to origin of anal fin 1.2 (1.2-1.3).

Head elongate, somewhat depressed, especially over snout. Top
of head slightly convex between eyes. Snout rounded in horizontal
profile. Head 3.9 (3.9-4.0); eye in head, Brazilian specimens 3.0
(2.8-3.2), British Guiana specimens 3.4 (3.2-3.4); snout in eye 1.0
(0.9-1.2); least width of bony interorbital in greatest eye diameter
1.1 in all specimens from Brazil and 1.0 in all specimens from British
Guiana.

Premaxillary with 6 to 7 quinquecuspid teeth. Posteriomost lateral
tooth sometimes with only 3 to 4 cusps. Each maxillary with 1
quadricuspid tooth. Dentary with 6 or 7 quinque- or sexcuspid
teeth, cusps subequal. Second row of dentary absent in all alizarin
specimens. Gill rakers 9+ 14.

Dorsal fin rays 11,8; anal 1,9; pectoral i,9 to i,11; pelvics ii,7.
Adipose fin present in all specimens examined. Caudal fin with
principal rays 10/9. Distal tips of 8 principal rays in upper lobe and
11 in lower lobe.

Anal fin of males slightly modified. Male specimens examined from
Venezuela and Brazil with anal fin shape as shown in insert in figure 11.
This shape approaches that of anal fin of Poecilobrycon eques but fin of
P. unifasciatus not as large. Second undivided ray expanded anteriorly
and posteriorly, giving this ray a flattened appearance. Posterior
margin of this ray fits into recess formed by 2 ray halves of expanded
and flattened third unbranched ray behind. First branched ray with
slight recess for the enlarged third ray. Inclinator muscle only slightly
developed and beyond basal scales attached to ninth ray only.
Specimens from British Guiana with anal fin of male less developed
than specimens from Brazil and Venezuela. Anal fin of specimen
from British Guiana resembles that of entire specimen in figure 11.
Second and third unbranched and first and second branched rays only
slightly enlarged. Anal fin of female from British Guiana, Brazil, and
Venezuela with rays not at all enlarged. Male specimens from Brazil

44 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

and Venezuela shorter than about 24 mm. in standard length with
anal fins much like those of males from British Guiana. Possibly
anal fin develops its pointed posterior border only in males longer than
about 29 mm. in standard length.

Radial grooves of scales in that part of third horizontal scale series
below dorsal fin rather variable but frequently resemble scales in
figure 11. Posterior field with 1 to 3 (most frequently 2) radial
erooves, dorsal and ventral fields with 1 and anterior embedded field
with 2 to 6 radial grooves. Scales in a lateral series 28 to 30 (usually
29) and 2 to 5 (usually 4) scales in lateral line. Scales in a median
series before the dorsal fin 11. Total number of vertebrae 33 to 34
with 21 precaudal and 12 to 13 caudal vertebrae. British Guiana

Figure 11.—Poecilobrycon unifasciatus, male, SL 33.4 mm., CAS(IUM) 11703 (see dis-

cussion for remarks on insert drawings of anal and caudal fins).

specimens all with 34 vertebrae and all specimens recorded below from
Tgarapé do Mai Joana, near Mandos, with 33 vertebrae. Of specimens
examined from Teffé, both with 33 vertebrae.

CoLor IN ALCoHOL.—(Specimens from British Guiana) Dorsum of
head light brown. Primary horizontal stripe beginning on snout and
lower jaw, extending posteriorly over maxillary and first orbital bone
onto eye. Posteriorly from iris this stripe continuing across part of
fourth and fifth orbitals to across opercular bone, not present on
fleshy opercular flap and extending through fourth scale row to end
of caudal peduncle as shown in figure 11. On caudal peduncle primary
horizontal stripe expanding ventrally to include entire ventral area.
On caudal fin primary horizontal stripe covering entire lower lobe
except for crescent near lower posterior margin and central clear area

NO. 3538 CHARACID FISHES—WEITZMAN 45

as shown in figure 11. Upper lobe dusky near its basal, lower portion,
and this dusky region with dark spot (ocellus) with a translucent area
around it. Secondary horizontal stripe appearing absent and entire
back light brown. Belly bright silvery with a dark spot in front of
vent and a very small dark spot in front of pelvic fin base. Head below
primary horizontal stripe bright silver. Pectoral, dorsal, and anal
fins translucent. Anal fin with single dark spot in its basal and central
regions. Preserved specimens from Brazil and Venezuela not differing
in color except in caudal fin. Specimens from these areas lack an
ocellus, except one from Venezuela with an imperfect ocellus. Also
markings on lower lobe of caudal fin somewhat different. These well
shown in figure accompanying Steindachner’s original description.
Preserved specimens from Igarapé do Mai Joana at Manfos, with
primary horizontal stripe extending onto caudal fin as a narrow wedge,
and distal region of caudal fin dusky to dark (see upper right insert in
fig. 11). Steindachner’s figure shows primary horizontal stripe ex-
tending onto caudal fin as slightly widening band. However, all
specimens that I have examined from Brazil and Venezuela with a
caudal fin color pattern about as that shown in the upper right insert
in figure 11.

CoLor IN LirE.—Aquarium specimens from an unknown locality
(but agreeing well with color of preserved specimens from British
Guiana) had the following color pattern: Dorsal and pectoral fins
hyaline; pelvic fin also hyaline except for second and longest ray,
which is silvery blue on distal one-third to one-half of its length.
Anal fin hyaline except for central black blotch and silvery white distal
tips of fourth to seventh rays. Caudal fin variously colored, the upper
lobe hyaline; area dorsal to black ocellus white, and below it creamy
yellow to rich orange. Nonmelanistic areas on the lower caudal fin
lobe creamy white to yellow suffused with varying shades of orange.
Belly and throat regions white. Area below primary horizontal stripe
white. Silvery white stripe especially distinct behind dorsal fin above
black primary horizontal stripe. Above silvery stripe, dorsum and
top of head light to medium brown. Bright metallic-orange spot near
the anterior narial opening.

Remarxs.—Poecilobrycon unifasciatus was known previously only
from the mouth of the Rio Negro and Teffé and (under the name
Poecilobrycon ocellatus) from British Guiana. The records listed
below considerably enlarge its known range up the Rio Negro to the
headwaters of the Rio Orinoco in Venezuela. Hoedeman (1950)
apparently did not have specimens of the true unifasciatus, or if he
did, he confused them with P. eques. Hoedeman stated that some of
the young from parents with the typical color pattern of eques look

46 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

like typical unifasciatus. Poecilobrycon eques have spawned in my
aquaria and I have raised the resulting young. Also I have seen
young of eques raised by other aquarists. In none of these have I
seen any specimens that look like wnifasciatus. The description of
eques given by Hoedeman is hard to follow and seems somewhat
contused and inconsistent. However, his main thesis seems to be
that the differences noted by Steindachner between eques and wni-
fasciatus are those of age and sex. In any event, the descriptions
given here should serve to distinguish eques and wnifasciatus and put
an end to further confusion. From the key and descriptions, it can
be seen that the species are quite distinct in their morphology.

SPECIMENS EXAMINED.—Thirteen, SU 50263 (three alizarin preparations, SU
50264), SL 27.0-34.8 mm., Brazil, State of Amazonas, Igarapé do Mai Joana
into Rio Negro near Mandos, Dec. 24, 1924, Carl Ternetz.—One, MCZ 19769,
SL 31.5 mm., Brazil, State of Amazonas, Lake Hyanuary, near Mandos, 1865-
1866, Thayer Expedition, Mr. Navez.—One, MCZ 19935, SL 28.0 mm., Brazil,
Jutahy, 1865-1866, Thayer Expedition, James, Thayer and Tailisman.—Two,
MCZ 19787, SL 29.0-38.3 mm., Brazil, Lake José Asst, 1865-1866, Thayer
Expedition, Louis Agassiz.—Fifteen, MCZ 19591, SL 25.0-33.0 mm., Brazil,
State of Amazonas, Teffé, 1865-1866, Thayer Expedition, Louis Agassiz—Two,
SU 50266, SL 23.0-27.2 mm., Venezuela, Rio Orinoco, Cafio de Quiribana,
opposite Pan de Aztcar, near Caicara, May 7, 1925, Carl Ternetz.—Five, SU
50265, SL 26.0-33.0 mm., Venezuela, Rio Orinoco, creeks at high water into
Laguna San Raphael, Caicara, Apr. 28, 1925, Carl Ternetz.—Two (paratypes
of P. ocellatus Eigenmann), CAS(IUM) 11703, SL 31.2-32.8 mm., British Guiana,
Demerara River, Wismar, 1908, Carl H. Eigenmann.—Ten (paratypes of P.
ocellatus Eigenmann), CAS(IUM) 11704 (one specimen as alizarin preparation),
SL 29.6-34.0 mm., British Guiana, Rockstone sandbank, Essequibo River, 1908,
C. H. Eigenmann.—Two (paratypes of P. ocellatus Eigenmann), SU 21963, SL
31.8-33.5 mm., British Guiana, Rockstone, Essequibo River, 1908, Carl H.
Eigenmann.—Three (paratypes of P. ocellatus EKigenmann), CAS(IUM) 11705,
SL 29.2-37.1 mm. and three (paratypes of P. ocellatus Eigenmann) CNHM
52782, SL 32.0-34.0 mm., British Guiana, Gluck Island, Essequibo River, 1908,
Carl H. Eigenmann.—Three (paratypes of P. ocellatus Eigenmann), CAS(IUM)
11706, SL 28.2-29.2 mm., British Guiana, Rupununi, Essequibo drainage, 1908,
C. H. Eigenmann.—One (paratype of P. ocellatus Eigenmann), CAS(IUM)
11708, SL 30.0-33.5 mm., British Guiana, Konawaruk, Potaro River, 1908,
C. H. Eigenmann.—Two, SU 19049, SL 31.0-34.4 mm., British Guiana, no
other data—Three, SU 19048, SL 27.0-28.0 mm., British Guiana, no other
data.—One, SU 25613, SL 29.5 mm., aquarium specimen, no other data.—T wo,
SU 50267, SL 30.1-30.5 mm., aquarium specimens, no other data.—Two, SU
50268, SL 27.8-30.4 mm., alizarin preparations, aquarium specimens, no other
data.—One, SU 50438, SL 30.6 mm., British Guiana, Karanambo, ‘‘Grass Pond,”
Rupununi River, Essequibo drainage, Sept. 19, 1957, Rosemary Lowe (McCon-
nell).—Three, SU 50435, SL 23.2-31.4 mm., British Guiana, Moreby Creek,
Rupununi River near Karanambo, Essequibo drainage, Sept. 17, 1957, Rosemary
Lowe (McConnell).

No. 3638 CHARACID FISHES—WEITZMAN AT

Poecilobrycon eques (Steindachner)
Ficure 12

Nannostomus eques Steindachner, 1876, p. 126, pl. 9, fig. 3 (original description;
type locality: Peruvian Amazon above Tabatinga; types in Vienna Mu-
seum).—LHigenmann and Eigenmann, 1891, p. 49 (listed) —Sterba and
Tucker, 1962, p. 215 (description and aquarium notes).

Poecilobrycon auratus Kigenmann, 1909, p. 43 (original description; type locality:
British Guiana, canal at Christianburg; holotype in Chicago Natural History
Museum); 1910, p. 427 (listed).—Innes, 1935, p. 156 (description from life
and photograph).

Poecilobrycon trifasciatus.—(not Steindachner) Higenmann, 1912, p. 284, pl. 37,
fig. 2 (description).

Poecilobrycon eques.—Eigenmann, 1910, p. 427 (listed) —Eigenmann and Allen,
1942, p. 288 (listed).

Nannobrycon eques.—(in part) Hoedeman, 1950, p. 23 (description; photograph) ;
1954b, p. 82 (description; photograph); 1954b, p. 82 (description; photo-
graph).

Diaenosis.—The unique feature of this species is its five lateral
horizontal stripes. In addition it may be distinguished from all
other nannostominans by the following combination of characters:
No perforated lateral-line scales; 24 to 25 scales in lateral series;
33 to 34 vertebrae; 11,9 anal fin rays; 16-+24 gill rakers; 12 teeth in the
second tooth row of dentary.

Description.—Body elongate, somewhat cylindrical through an-
terior half, but sides flattened and compressed posteriorly. Greatest
depth occurs between posterior tip of appressed pectorals and anterior
base of dorsal fn. SL of largest specimen 35.0 mm. (largest aquarium
specimen examined, a female, with SL of 37.0 mm.). Greatest body
depth 4.5 (4.0-5.1); least depth of caudal peduncle 10.0 (10.0-11.0);
length of caudal peduncle (including both sexes) 6.7 (6.0-7.1); snout
tip to origin of dorsal fin 1.7 (1.7-1.8); snout tip to origin of anal
fin about 1.3 in all specimens examined.

Head elongate, rather acute in vertical profile with tip of snout
obtuse. Snout region somewhat depressed. Top of head very
slightly convex between eyes. Head 3.5 (3.5-3.7); eye in head 3.3
(3.0-3.5); snout in eye 0.9 (0.8-1.0); least width of bony interorbital
in greatest eye diameter 1.0 (0.9-1.0).

Premaxillary with 8 to 9 teeth, each quadri- to quinquecuspid,
cusp next to posterolateral one largest. Maxillary with 1 quadri- or
quinquescuspid tooth, cusps subequal. Dentary with 7 to 9 quinque-
to sexcuspid teeth. Posterior 1 or 2 teeth usually with 3 cusps.
Either central cusp or 2 posterolateral cusps largest (fig. 12). About
12 teeth in second dentary tooth row. Gill rakers 16+ 24.

Dorsal fin rays 1,8; anal iii,9; pectoral i,9 to i,11; pelvics ii,7.
Adipose fin present or absent, seemingly not correlated with geo-

48 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

graphical origin. Caudal fin with principal rays 10/9; distal tips of
8 principal rays in upper lobe and 11 in lower lobe.

Anal fin of males enlarged and modified, but not as greatly as in
Nannostomus digrammus. Second undivided ray expanded anteriorly
and posteriorly, giving this ray a flattened appearance. Posterior
margin of this ray fits into recess formed by 2 ray halves of expanded
and flattened third unbranched ray behind. First, second, and third
branched rays slightly flattened and expanded, first to greatest extent.
These rays slightly ribbed. Rest of anal fin rays almost normal,
being only slightly heavier than usual type of characid fin ray. In-
clinator muscle attached to ninth ray only.

Figure 12.—Poecilobrycon eques, male, SL 29.0 mm., SU 50299.

Radial grooves of scales in third horizontal scale series below
dorsal fin quite variable but general pattern shown in figure 12.
Posterior field may have 1 to 4 radial grooves, anterior field 2 to 6
radial grooves and dorsal and ventral fields one radial groove each.
Scales in a lateral series 24 to 25, no lateral-line scales and 11 or
12 predorsal scales. ‘Total number of vertebrae 33 to 34, usually
18 to 19 precaudal and 15 caudal vertebrae.

CoLtor IN ALcoHoL.—Top of head light brown. Nape, areas
immediately above nares and eyes, central parts of frontals, and
tip of snout, dark brown. Primary horizontal stripe beginning at
tip of snout and lower jaw, extending posteriorly over first infra-
orbital to pupil of eye. Then extending from posterior edge of
pupil to posterior edge of opercular bone. On opercle width of
this stripe somewhat variable. On body, primary horizontal stripe
beginning under fleshy opercular flap and extending posteriorly
through entire fourth and fifth scale rows. In some specimens upper
two-thirds of fourth scale row free of heavy pigment. Over anal fin

NO. 3538 CHARACID FISHES—WEITZMAN 49

primary horizontal stripe extending dorsally to cover lower half of
third scale row. On caudal peduncle primary horizontal stripe
extending ventrally to cover entire lower half of caudal peduncle
and then posteriorly to include entire lower lobe of caudal fin. On
fifth row of scales primary horizontal stripe frequently confined to
center of each scale, producing spotted appearance. In some speci-
mens this shows well, in others pigment expanded and covering entire
fifth row of scales, making primary horizontal stripe appear solid.
Above primary horizontal stripe a broad pale-yellowish stripe covering
lower two-thirds of horizontal scale row. This stripe beginning on
snout above primary horizontal stripe and extending posteriorly
through caudal peduncle to caudal fin. Weak scattering of melano-
phores over this band. Region of posterior part of sixth to posterior
edge of eighth vertical scale rows with a dense aggregation of melano-
phores extending from middle of first to middle of sixth horizontal
scale rows. These are oblique bands, and they appear faintly to
strongly in all preserved specimens examined. A dense aggrega-
tion of melanophores occurs over anal fin. Entire dorsum light
brown and secondary horizontal stripe consists of dense aggregation
of melanophores through center of each scale in second horizontal
scale row. Fourth narrow horizontal stripe extending through first
horizontal scale row. Third horizontal stripe present as a series of
dark spots along sixth horizontal scale row, each dot being in center
of scale. Belly and upper parts of head clear yellowish white. In
some specimens third horizontal stripe present on head as a thin
dark stripe on third infraorbital bone. Adipose fin, when present,
frequently black. Dorsal, pectoral, and pelvic fins hyaline. Anal
fin almost entirely black except for clear area in its midbasal region.
Second undivided ray also hyaline.

CoLor IN LIFE.—Aquarium specimens from an unknown locality
had the following color pattern: Pectoral, dorsal, and upper lobe of
caudal fins hyaline. Anal fin dark brown or black except for clear
area mentioned above; this area bright orange red. Large second ray
of anal fin silvery blue for about one-third its proximal length. Pelvic
fins hyaline except for small amount of silvery blue pigment along
large second fin ray. Lower lobe of caudal fin dark brown or black,
variously and sparsely mottled with pale gold or orange. Some
specimens with slight suggestion of an ocellus as in specimens of
unifasciatus from British Guiana. Bright pale-gold blotch in region
of caudal fin between two caudal lobes. Bright silvery stripe extend-
ing from snout tip to caudal peduncle above primary horizontal
stripe. Dorsum light to medium brown except where covered by
darker stripes described above. Pale orange or yellow metallic spot
in region of nares. Belly and throat white.

50 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Remarxs.—As noted above under Poecilobrycon unifasciatus,
Hoedeman (1950) confused wnifasciatus and eques. However, as the
two descriptions presented here show, these two names are applied to
quite distinct species. As noted on page 28, under the remarks about
Nannostomus trifasciatus, Eigenmann (1909 and 1912) was apparently
confused in regard to the identity of Steindachner’s trifasciatus and
eques. This probably led to the description of auratus, a synonym of
eques. In any event, comparisons of specimens of eques (including
those described as auratus) from British Guiana, widely separated
parts of the Amazon and Rio Negro Basins, and from the headwaters
of the Rio Orinoco, show that this species is remarkably constant in
its color pattern throughout its range, more so than any other widely
distributed nannostominan.

SPECIMENS EXAMINED.—One, SU 36898, SL 31.0 mm., Peru, Pévas [Pébas]
District, Shansho Cafio, Sept. 20, 1936, W. G. Scherer.—Two, SU 50236, SL 23.0-
33.5 mm., Peru, Rfo Ampiyacu, near Pévas [Pébas], Aug. 16, 1940, W. G. Scherer.
—Four, SU 50237, SL 21.8-27.0 mm., Peru, Pévas [Pébas], 1940, W. G. Scherer.—
Two, SU 50226, SL 26.2 mm., Peru, Yaguasyact, July 7, 1941, W. G. Scherer.—
Five, SU 50229, plus two alizarin preparations SU 50247, SL 28.4-30.5 mm.,
Brazil, State of Amazonas, Sao Gabriel, Rio Negro, rock pools below rapids,
Feb. 1, 1925, Carl Ternetz.—One, SU 50227, SL 29.2 mm., Venezuela, mouth
of the Casiquiare, Feb. 24, 1925, Carl Ternetz.—Two, SU 50228, SL 31.5-32.9
mm., Venezuela, Cafio de Quiribana, near Caicara, May 2, 1925, Carl Ternetz.—
One, SU 50233, SL 31.4 mm., Brazil, State of Pard, Igarapé do Sapucta into
Lagéa Sapucta (this lake drains into the Rio Trombetas, a tributary of the Amazon
River), June 7, 1924, Carl Ternetz.—Three, SU 50230, SL 25.4-25.7 mm., Brazil,
State of Pard, Cabeceira Maturaca into Lagéa Grande, Nov. 1924, Carl Ternetz.—
Four, SU 50231, SL 24.38-25.0 mm., Brazil, State of Amazonas, Igarapé do Mai
Joana, into Rio Negro near Mandos, Dec. 25, 1924, Carl Ternetz.—Three, SU
50232, SL 26.5-26.6 mm., Brazil, ““Pard, Juntil lago Jard,” Apr. 4, 1924, Carl
Ternetz.—Six, MCZ 6312, SL 23.5-27.5 mm., Brazil, State of Amazonas, Lake
Hyanuary, near Mandos, Oct. 28, 1865, Thayer Expedition, Louis Agassiz.—
Three, MCZ 19473, SL 21.5-24.4 mm., Brazil, State of Amazonas, Villa Bella,
1865-1866, Thayer Expedition, Louis Agassiz—Two, MCZ 19594, SL 24.0-24.5
mm., Brazil, State of Amazonas, region of Tabatinga, Sept. 20 to Oct. 20, 1865,
Thayer, Expedition, Bourget.—Four, MCZ 19714, SL 24.6-28.5 mm., Brazil,
State of Amazonas, Lagéa, Saracd, Silves, near Serpa (Itacutiara), 1865-1866,
Thayer Expedition, Stephen Van Renssalaer—Two, MCZ 19861, SL 21.0—-
21.5 mm., Brazil, State of Pard, Santarém, 1865-1866, Thayer Expedition,
Bourget.—Seven, MCZ, 19646, SL 17.5-27.4 mm., Brazil, State of Parad, Obidos,
1865-1866, Thayer Expedition, Col. Bentos—One, MCZ 19765, SL 27.4 mm.,
Brazil, State of Amazonas, Lake Hyanuary, near Mandos, 1865-1866, Thayer
Expedition, M. Navez.—Thirteen, MCZ 19965, SL 24.0-27.5 mm., Brazil, State
of Amazonas, Codajdz, 1865-1866, Thayer Expedition, Bourget.—Four, MCZ
19978, SL 25.4-30.5 mm., Brazil, Juturana, 1865-1866, Thayer Expedition.—
Two, MCZ 19998, SL 25.7-27.0 mm., Brazil, Curupira, 1865 or 1866, Thayer
Expedition, Major Coutinho.—One, SU 21964, SL 24.4 mm., and four CAS(IUM)
11688, SL 21.6-25.5 mm., British Guiana, Konawaruk, Essequibo River, 1908,
Carl H. Eigenmann.—Three, CAS(IUM) 11690, SL 24.9-27.0 mm., British
Guiana, Essequibo River at Rockstone, 1908, C. H. Higenmann.—Five,

i
i
i
|
|
.
:

I
|
\
|
H

NO. 3538 CHARACID FISHES—WEITZMAN 51

CAS(IUM) 11690, SL 19.5-23.7 mm., British Guiana, Essequibo River at Gluck
Island, Sept. 30, 1908, C. H. Higenmann.—Two, SU 50234, SL 25.0-27.0 mm.,
British Guiana, no other data.—Four, SU 50225, SL 29.4-35.4 mm., aquarium
specimens, no other data.—Three, SU 50224, SL 25.8-32.5 mm., aquarium
specimens, alizarin preparations, no other data,

Discussion

In the course of this study three major problems have become ap-
parent. The first of these is that of generic allocation; the second and
really inseparable one is the relationships among the species; and the
third and largest problem, left largely unworked by this study, is geo-
graphical variation and subspeciation. To untangle the nomencla-
tural mixup and mistaken identifications of past authors has presented
some minor difficulties, and the conclusions reached are presented in
the synonymies above.

Generic designation has proved difficult; indeed, I have believed at
times that perhaps it would be best to place all generic and subgeneric
taxa treated here under Nannostomus, following Sterba and Tucker
(1962). However, it seems that to do this would obscure valid differ-
ences and relationships. It appears that the only valid difference be-
tween Nannostomus and Poecilobrycon is the presence or absence of a
sensory tube in the second orbital bone. This difference seems small,
but it proved to be remarkably constant in all specimens examined.
The only other character found to correlate with this was a tendency
for a longer snout in Poecilobrycon than in Nannostomus. However,
measurements show an overlap in snout lengths even though individual
bones of the snout region always seemed broader and relatively shorter
in Nannostomus than Poecilobrycon. Both Poecilobrycon eques and
Poecilobrycon unifasciatus are obviously closely related because of their
caudal fin structure and swimming habits. Their tendency toward
long snouts and their possession of tubed second infraorbital bones
indicate a relationship with Poecilobrycon harrisont. Indeed, the liv-
ing color pattern of wnifasciatus suggests relationships both with harri-
sont and eques. These facts made it seem desirable to place these
fishes under the genus Poecilobrycon with Nannobrycon as a subgenus
for unifsaciatus and eques. The color pattern of P. harrisoni suggests
relationships with Nannostomus beckfordi, and it is possible that they
had a common ancestor. It thus seems that Poecilobrycon is most
closely related to that section of the Nannostomus containing beckfordi.

A few groupings can be determined in Nannostomus itself. Nannos-
tomus trifasciatus and marginatus are obviously closely related in color
pattern and anal fin structure; indeed, a separate subgenus could be
erected for their reception but I do not believe it advisable. Nanno-
stomus espei appears apart from all other members of this genus in hav-

52 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

ing the shape of certain scales and the color pattern similar to those of
pyrrhulininans. However, a generic or subgeneric name does not seem
advisable for this species. Whether these pyrrhulinian-like characters
of espei indicate an archaic nature is problematical. I am inclined to
believe that they are, especially when one considers that the body
shape of this species is more like that of pyrrhulininans than any other
nannostominan. On the other hand, it lacks the tube in the second
orbital bone that is present in Poecilobrycon and the pyrrhulininans.
Presumably the presence of this tube is a primitive character for nan-
nostominans. It is possible that the loss of the tube in espe? was inde-
pendent of its loss in other species of Nannostomus and that my
association of espet with species of Nannostomus rather than with
Poecilobrycon is artificial. The problem cannot be settled with the
morphological data at hand.

Nannostomus digrammus is likewise apart from other species of Nan-
nostomus in the highly modified anal fin of the male. It appears, on
basis of color pattern and premaxillary teeth, to be more closely
related to beckfordi and bifasciatus than to any other known nanno-
stominan.

Both N. beckfordi and N. bifasciatus appear related. They lack
specialized or distinctive characters that set the other species of
Nannostomus apart from them. They both lack an adipose fin; appar-
ently both have at least some perforated lateral-line scales and have
only moderately modified anal fins in the male. Their color patterns
are somewhat different. However, until the color pattern and other
characters of the widely distributed beckfordi can be studied adequately,
it will remain difficult to evaluate its relationship with bifasciatus.

Concerning geographical variation within a species, N. espei, N.
bifasciatus, and P. harrisoni appear to have restricted ranges, and little
or no geographical variation was noted in the specimens utilized in the
present study. Nannostomus beckfordi, digrammus, trifasciatus, margi-
natus, Poecilobrycon unifasciatus, and eques have a wide distribution in
South America. Of these, beckfordi and marginatus were noted to
have the greatest geographical variation. Indeed beckfordi is so vari-
able that future work based on sufficient collections may show good
reasons to recognize many subspecies and perhaps even species of this
group of related populations.

Although some morphological, especially live color, variation cor-
related with geographical distribution was noted in marginatus, the
close relationship of these population samples seems clearer than those
of beckfordi. Some degree of difference was found in specimens of
trifasciatus from Peru and British Guiana and this, it seemed, could
be correlated with color differences in aquarium specimens said to have

NO. 3538 CHARACID FISHES—WEITZMAN 53

come from these respective areas. However, the source of the speci-
mens was never reliable enough for serious consideration.

Insufficient specimens of digrammus in good condition were at hand
to consider geographical variation in color pattern or other characters.

The presence or absence of an ocellus in wnifasciatus seemed geo-
graphically correlated, along with a few other characters. Undoubt-
edly ocellatus and unifasciatus will be recognized as subspecies when
sufficient study material becomes available.

Poecilobrycon eques seems remarkably constant over its large range
from Peru to British Guiana. Both color pattern and morphology of
preserved specimens seem quite stable, and I have not noticed any
differences in live specimens said to be imported from Peru and
British Guiana.

Summary

The characid subtribe Nannostomina (defined by Weitzman, 1964)
contains two genera, Nannostomus and Poecilobrycon. Poecilobrycon
is divided into two subgenera, Poecilobrycon and Nannobrycon. Nan-
nostomus consists of six known species, Poecilobrycon of three, with two
of these in the subgenus Nannobrycon. The species are as follows:

Nannostomus espet (Meinken)
Paruma River, British Guiana.
Nannostomus beckfordi Giinther
British Guiana, south through French Guiana, and Surinam, and in the
Amazon Basin as far west as the Rio Negro.
Nannostomus bifasciatus Hoedeman
Surinam River, Surinam.
Nannostomus digrammus Fowler
Rupununi district of British Guiana and west into Brazil as far as the
Rio Negro.
Nannostomus trifasciatus Steindachner
British Guiana, westward and south into the Amazon Basin as far as the
Pébas district of Peru.
Nannostomus marginatus Kigenmann
British Guiana, southward into the Amazon Basin and as far west as the
Caquetd Province of Colombia.
Poecilobrycon harrisoni Kigenmann
Demerara River, British Guiana.
Poecilobrycon unifasciatus (Steindachner)
British Guiana, Rio Negro in Brazil north to Rio Orinoco in Venezuela.
Poecilobrycon eques (Steindachner)
British Guiana south into Amazon Basin and west to Pébas region of Peru,
Rio Negro north to Caicara in Venezuela.

Literature Cited

AuL, ERNST

1933. Poecilobrycon vittatus. Das Aquarium, p. 184.

1934. Beschreibung einer neuen siidamerikanischen Characiniden—art der
Gattung Poecilobrycon. Zool. Anz., vol. 106, pp. 124-125.

ARNOLD, JOHANN P.

1920. Importriickblick 1948-1949: Ueber fremdlindische Siisswasserfische:
Taschenkalender fiir Aquarien—und Terrarienfreunde, 1950, pp.
63-87. [Not seen.]

AXELROD, HERBERT, and ScuuutTz, LEonarpD P.
1955. Handbook of tropical aquarium fishes, xii+718 pp.
BorsEmMan, M.

1952. <A preliminary list of Surinam fishes not included in Eigenmann’s
enumeration of 1912. Zool. Meded., vol. 31, pp. 178-200.

1953. The fishes, 1. No. 2 in Zoology, part 2 of Scientific results of the
Surinam Expedition 1948-1949. Zool. Meded., vol. 32, pp. 1-24.

1954. Onasmall collection of Surinam fishes. Zool. Meded., vol. 33, pp.
17-24,

BOHLKE, JAMES E.

1955. Studies on fishes of the family Characidae, 9: Notes on the distribution,
variation, and type locality of Gnathocharax steindachneri Fowler.
Notulae Naturae, no. 277, 14 pp.

1956. Studies on fishes of the family Characidae, 12: On the status of
Poecilobrycon espet Meinken, a recently described pencil fish from
headwaters of the Mazaruni in western British Guiana. Notulae
Naturae, no. 290, 7 pp.

BRAEMER, WOLFGANG, and BRAEMER, HELGA

1958. Orientation of fish to gravity. Limnology and Oceanography, vol. 3,

pp. 362-372.
EIGENMANN, Car. H.

1894. Notes on some South American Fishes. Ann. New York Acad.
Sci. 1892-1894, vol. 7, pp. 625-637.

1909. Some new genera and species of fishes from British Guiana. Report
No. 1. zr» Reports on the expedition to British Guiana of the
Indiana University and the Carnegie Museum, 1908. Ann.
Carnegie Mus., vol. 6, pp. 4-54.

1910. Catalogue of the fresh-water fishes of tropical and south temperate
America. Vol. 3, of Zoology in Report of the Princeton University
Expedition to Patagonia, 1896-1899, pp. 375-511.

1912. The fresh-water fishes of British Guiana. Mem. Carnegie Mus.,
vol. 5, xxii++578 pp., 103 pls.

EIGENMANN, Car H., and EIGENMANN, Rosa SMITH

1891. A catalogue of the fresh-water fishes of South America. Proc. U.S.

Nat. Mus., vol. 14, pp. 1-81.
EIGENMANN, Caru H., and ALLEN, WILLIAM R.
1942. Fishes of western South America, xv-+494 pp., 22 pls.

54

NO. 3538 CHARACID FISHES—WEITZMAN 55

Fow.er, Henry W
1913. Fishes from the Madeira River, Brazil. Proc. Acad. Nat. Sci. Phil-
adelphia, vol. 65, pp. 517-579.
1950. Os peixes de agua doce do Brasil (2.%entrega). Arq. Zool. Est. Sao
Paulo, vol. 6, pp. 205-404.
1954. Os peixes de agua doce do Brasil (4.*entrega). Arq. Zool. Est. Sio
Paulo, vol. 9, pp. 1-400.
GARMAN, SAMUEL
1895. The Cyprinodonts. Mem. Mus. Comp. Zool. vol. 19, no. 1, pp. 1-179,
pls. 1-12.
GintHer, ALBERT C. L. G.
1872. On a new genus of characinoid fishes from Demerara. Proc. Zool.
Soc. London, 1872, p. 146.
HoEpEMAN, J. J.
1950. Rediagnosis of the characid-nannostomine fish genera Nannostomus
and Poecilobrycon. Amsterdam Natur., vol. 1, pp. 11-27, pls. 8-9.
1953. De Encyclopaedie voor de aquariumhouder. Leaflet X.30.11.311.
Amsterdam. [Not seen.]
1954a. Notes on the ichthyology of Surinam (Dutch Guiana), 3: A new
species and two new subspecies of Nannostomidi from Surinam
River. Beaufortia, Ser. Misc. Publ. Zool. Mus., Amsterdam, vol.
4, pp. 81-89.
1954b. Aquariumvissen-encyclopaedie, nieuwe importen. Amsterdam,
527 pp.
Husss, Caru L.
1926. Studies of the fishes of the order Cyprinodontes, 6. Misc. Pub. Univ.
Michigan Mus. Zool., no. 16, 86 pp., 1 pl.
InNEs, WILLIAM T.
1933. Nannostomus trilineatus. The Aqarium (Philadelphia), vol. 2, p. 142.
1935. Exotic aquarium fishes, ed. 1, 463+1 pp.
1953. Exotic aquarium fishes, ed. 15, 523 pp.
LapIcEs, WERNER
1948. Importbericht: Poecilobrycon und Nannostomus Arten. Wochenschr.
Aquarien- und Terrarienkunde, Jahrg. 42, vol. 2, pp. 28-31.
MEINKEN, HERMANN
1931. Nannostomus anomalus Steindachner, marginatus Kigenmann und
aripirangensis n. spec. Wochenschr. Aquarien- und Terrarien-
kunde, Jahrg. 28, pp. 553-555.
1954. Nannostomus anomalus Steind. 1876 und Nannostomus aripirangensis
Meinken, 1931. Aquarien und Terrarien Zeitschr., Jahrg. 7, pp. 1-4.
1956. Poecilobrycon espei spec. nov., eine hiibsche Neueinfiihrung. Aqua-
rien und Terrarien Zeitschr., Jahrg. 9, pp. 31-32.
NIEUWENHUIZEN, A. V. D.
1954. Nannostomus bifasciatus Hoedeman, 1953. Het. Aquarium, Jaargang
243, pp. 258-262.
Puyo, JosEpH
1949. Poissons de la Guyane Francaise. Vol. 12 of Fauna de l’Empire
Francaise (Paris), 280 pp.
STEINDACHNER, FRANZ
1876. Ichthyologische Beitrige (V), 2: Ueber einige neue Fischarten,
insbesondere Characinen und Siluroiden aus dem Amazonenstrome.
Sitzb. Akad. Wiss. Wien, Math-Naturw. Classe, vol. 74, pp. 49-240,
15 pls.

56 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

STERBA, GUNTHER
1962. Freshwater fishes of the world, 878 pp. [Translated and revised by
Denys Tucker.]
VORDWINKLER, WILLIAM
1957. Pencilfishes. T.F.H. Publications Inc., New Jersy, 26 pp.
WEITZMAN, STANLEY H.
1962. The osteology of Brycon meeki, a generalized characid fish with an
osteological definition of the family. Stanford Ichth. Bull., vol. 8,
no. 1, pp. 3-77.
1964. Osteology and relationships of South American characid fishes of
subfamilies Lebiasininae and Erythrininae with special reference to
subtribe Nannostomina. Proc. U.S. Nat. Mus., vol. 116, no. 3499,
pp. 127-169.
WICKLER, WOLFGANG
1957. Verhalten von Nannostomus beckfordi ariptrangensis Meinken, mit
einer Besprechung der Schwimmblasenfunktion. Beaufortia Ser.
Misc. Publs. Zool. Mus. Amsterdam, vol. 6, pp. 203-220.

U.S. GOVERNMENT PRINTING OFFICE: 1966

eroceedines of

the United States

National Museum

SMITHSONIAN INSTITUTION + WASHINGTON, D.C.

rT eee
Volume 119 1967 Number 3539

A REVISION OF THE HAMMERHEAD SHARKS
(FAMILY SPHYRNIDAE)

By Carter R. GILBERT

Introduction

The hammerhead sharks, so named because of the unusual lateral
expansion of the head, comprise a moderately large family of sharks
occurring in tropical and temperate waters throughout the world.
They are a rather old group; fossil remains have been found in Eocene
deposits from Virginia (Shelton P. Applegate, pers. comm.). The
family belongs to the relatively speciose and poorly known suborder
Galeoidea, in which are found most of the large and all of the dangerous
kinds of sharks. Since the hammerheads are well defined, and, with
one exception, adequate study material exists for all the species, they
form an ideal taxon for systematic study.

The most recent works of a revisionary nature that have appeared
on the hammerheads (Tortonese, 1950, pp. 1-39; Fraser-Brunner,
1950, pp. 213-219) list eleven and ten species, respectively; however,
the above authors were unable to examine examples of all the species
treated or all the extant types. Consequently, a number of con-
clusions were reached that now appear to be erroneous. The following

' Florida State Museum, University of Florida, Gainesville.

2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119
is a comparison of the systematic arrangements of the family Sphyr-
nidae, as presented by Tortonese and Fraser-Brunner, with a classifi-
cation as determined from the present study:

Proposed

Subgenus Eusphyra Gill, 1861

1. Sphyrna blochii (Cuvier,
1817)
Subgenus Sphyrna Rafin-
esque, 1810
2. Sphyrna mokarran
(Riippell, 1835)

3. Sphyrna zygaena
(Linnaeus, 1758)

4. Sphyrna lewini (Griffith
and Smith, 1834)

5. Sphyrna couardi
Cadenat, 1951
Subgenus Platysqualus
Swainson, 1839
6. Sphyrna tiburo tiburo
(Linnaeus, 1758)
Sphyrna tiburo vespertina
Springer, 1940
7. Sphyrna media
Springer, 1940
8. Sphyrna corona
Springer, 1940
9. Sphyrna tudes
(Valenciennes, 1822)

Tortonese, 1950

Fraser-Brunner, 1950

Eusphyra blochit (Cuvier Sphyrna blochii

1817)

Sphyrna mokarran
(Rippell, 1835)
Sphyrna tudes
(Valenciennes, 1822)
Sphyrna zygaena
(Linnaeus, 1758)
Sphyrna lewint
(Griffith, 1834)
Sphyrna oceanica
(Garman, 1913)
Sphyrna diplana
Springer, 1941

Sphyrna tiburo
(Linnaeus, 1758)
Sphyrna tiburo
(Linnaeus, 1758)
Sphyrna media
Springer, 1940
Sphyrna corona
Springer, 1940
Sphyrna bigelowt
Springer, 1944

(Cuvier, 1817)

Sphyrna tudes
(Valenciennes, 1822)

Sphyrna ligo, new
species

Sphyrna zygaena
(Linnaeus, 1758)

Sphyrna lewint
(Griffith, 1834)

Sphyrna oceanica
(Garman, 1913)

Sphyrna tiburo
(Linnaeus, 1758)
Sphyrna tiburo
(Linnaeus, 1758)
Sphyrna media
Springer, 1940
Sphyrna corona
Springer, 1940
Sphyrna bigelowt
Springer, 1944

Another nominal species, Sphyrna nana, which was recently

described by Sadowsky (1965), is regarded as a synonym of S. media,
for reasons discussed elsewhere in this paper.

Adequate study material exists for all of the above species, with
the exception of Sphyrna couardi. I have not seen an entire specimen
of this species, although I have examined a head; however, the mor-
phology of the head and chondrocranium, together with the patterns
of the Ampullae of Lorenzini, indicate that S. coward: is a valid form.

A number of structures have been studied that have proved to be
useful in the delineation of the species of Sphyrna. All species can
be distinguished on the basis of head shape alone. Head shape ranges
from evenly rounded and spadelike (Sphyrna tiburo tiburo) to ex-
tremely wide and narrow (Sphyrna blochii). Some forms have an

NO. 3539 HAMMERHEAD SHARKS—GILBERT 3

indentation on the median anterior margin of the head, while in
others this indentation is lacking. The presence or absence of an
outer narial and/or inner narial groove and the relative length of the
latter help to distinguish even the smallest individuals of a particular
species.

The various chondrocrania also are distinctive. Shape of both
the anterior and posterior parts of the preorbital process, degree of
development of the ‘‘wings” on the olfactory cartilage, development
of the rostral ‘wings,’ shape of the rostral cartilage, presence or
absence of accessory rostral cartilages, presence or absence of a
rostral fenestra, and shape of the anterior fontanelle all differ from
species to species.

The arrangement of the Ampullae of Lorenzini on the ventral
surface of the head is relatively constant for each species, with the
arrangement of those in the anterior-median area being particularly
diagnostic. These ampullae, which are long canals filled with mucus,
comprise a highly modified part of the lateral-line system, and it has
been suggested that they may be involved in the detection of temper-
ature changes, hydrodynamic pressure changes, tactile stimulation,
and/or electric gradients (Young, 1962, p. 172). For brevity, these
structures are referred to as mucal pores throughout this paper.

Teeth vary according to species, although in several instances the
differences are rather slight. Shape and arrangement of the teeth,
as well as the presence or absence of serrations, are distinctive.

There are differences between the species in size and shape of the
fins and in the positions of these structures relative to each other.
The positions of the fourth and fifth gill slits in relation to the insertion
of the pectoral fin and the relative lengths of the gill slits are important.
Finally, the various species differ in the presence or absence of a
lower precaudal pit and the shape of the upper precaudal pit.

Recent studies by Drs. Victor G. Springer and J. A. F. Garrick
(1964) on carcharhinid sharks indicate that the nature and number of
the vertebrae can be an extremely useful taxonomic tool, often showing
differences not evident from the study of external morphology. Un-
fortunately, the value of vertebral characters was not recognized
while my study was in progress, and consequently only a few counts
were made for the species of Sphyrna.

Other characters that are probably of taxonomic importance in
hammerheads are the morphology of the pelvic claspers and the
electrophoretic patterns of the serum proteins. The latter were not
studied because of the unavailability of fresh specimens and lack of
facilities. Because of the relatively small size and immaturity of
the available specimens, claspers of only three species, Sphyrna tudes,
S. media, and S. tiburo, were examined.

4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 119

Another problem encountered was the scarcity of large specimens;
however, I know of no taxonomically significant characters, with the
exception of the pelvic claspers, that are not evident in the embryos
and young. Although it would be desirable to know the full extent
of the various morphometric changes occurring during growth, such
data probably would not alter the present taxonomic conclusions.

The degree of confidence in the taxonomic conclusions reached in
this paper is dependent largely upon the number of specimens available
and on the extent of geographic coverage. The decision reached
concerning the status of Sphyrna mokarran, therefore, does not rest
on as firm a basis as for S. lewint and S. zygaena because far fewer
specimens of mokarran were seen and the geographic coverage of the
last two species was much better. Only one specimen of Sphyrna
tudes from the Mediterranean was examined, and, as stated previously,
S. couardi was represented only by a head. In the case of S. tudes,
however, only one other specimen of this species outside of the western
Atlantic area is known by me to be present in collections.

Acknowledgments

This study was supported by contracts between the Smithsonian
Institution, Office of Naval Research (ONR no. 1354(09)), and the
Atomic Energy Commission, and was carried out while the author
was a research associate of the U.S. National Museum. The aid
and cooperation of the following people have been invaluable during
the course of this work: Dr. Leonard P. Schultz, U.S. National Mu-
seum, who supervised the project; Drs. J. A. F. Garrick, Victor G.
Springer, Ernest A. Lachner, William Ralph Taylor, Stanley H.
Weitzman, and Mr. Robert H. Kanazawa, U.S. National Museum;
Dr. Shelton P. Applegate, Los Angeles County Museum, for most
of the information on dental variation in Sphyrna lewini and for other
information on shark classification; Madame M. L. Bauchot, Museum
National d’Histoire Naturelle, Paris, France, for loan of the type
specimens of Zygaena tudes; Dr. Bruce B. Collette, U.S. Fish and
Wildlife Service, Washington, D.C., for informing me of a specimen
of Sphyrna couardi he had collected; Dr. Daniel M. Cohen, U.S.
Fish and Wildlife Service, Washington, D.C.; Mr. Stewart Springer,

U.S. Fish and Wildlife Service, Stanford, Calif.; Drs. Giles W. Mead |
and Henry B. Bigelow, Mr. William C. Schroeder, and Mrs. Myvanwy ~
Dick, Museum of Comparative Zoology, Harvard University; Mr. |
Loren P. Woods, the late Mrs. Marion Grey, and Miss Pearl Sonoda, |

Field Museum of Natural History, Chicago, Ill.; Dr. James E. Boéhlke,
Academy of Natural Sciences of Philadelphia; Drs. Carl L. Hubbs,

Robert Wisner, and Richard H. Rosenblatt, Scripps Institution of |
Oceanography, University of California; Dr. Boyd W. Walker and Mr. |

NO. 3539 HAMMERHEAD SHARKS—GILBERT 5

Wayne J. Baldwin, University of California at Los Angeles; the late
Miss Margaret H. Storey, Stanford University; Dr. David K. Caldwell,
Los Angeles County Museum; Mr. William I. Follett and Mrs.
Lillian Dempster, California Academy of Sciences; Dr. Earl 8S.
Herald and Mr. Robert Dempster, Steinhart Aquarium, San Fran-
cisco, Calif.; Drs. Arthur D. Welander and Kelshaw Bonham, Uni-
versity of Washington; Dr. John C. Briggs, University of South
Florida; Drs. Reeve M. Bailey and Robert Rush Miller, University of
Michigan; Mr. Frank Williams, Director, Guinean Trawling Survey,
Lagos, Nigeria; Prof. Enrico Tortonese, Museo Civicio de Storia
Naturale, Genoa, Italy; Prof. M. Benazzi and Mr. Alberto Lanfranchi,
University of Pisa Museum, Pisa, Italy; Dr. Wolfgang Klausewitz,
Senckenberg Museum, Frankfurt-on-Main, West Germany; and to
the authorities of the British Museum of Natural History. Finally,
I would like to express my special appreciation to Mr. Paul Laessle,
staff artist of the Department of Zoology, University of Florida;
Mrs. Dorothea B. Schultz; and Mr. Craig Phillips, who are responsible
for the illustrations and graphs.

Materials and Methods

Specimens examined or recorded are from the following museum
collections: Academy of Natural Sciences of Philadelphia (ANSP);
British Museum (Natural History) (BMNH); California Academy
of Sciences (CAS); Field Museum of Natural History (formerly
Chicago Natural History Museum) (FMNH); Museum of Com-
parative Zoology, Harvard University (MCZ); Museum National
d’Histoire Naturelle (Paris) (IMNHN); Scripps Institution of Oceanog-
raphy (SIO); Stanford University (SU); University of California,
Los Angeles (UCLA); Florida State Museum, University of Florida
(UF); University of Miami, Institute of Marine Science (UMML);
University of Pisa (Italy) (UP); University of Michigan, Museum
of Zoology (UMMZ); United States National Museum (USNM).

Proportional measurements for all specimens were made according
to the methods described by Springer (1964, pp. 562-568). Measure-
ments are expressed in thousandths of total length (TL). Various
external morphological characters are illustrated and labelled in
figures 1 and 2. While the specimen on which figure 1 is based is
not a member of the family Sphyrnidae, most of the characters apply
equally to the hammerhead sharks.

Head-pore and chondrocranial terminologies (figs. 2, 3) are adapted
from Daniel (1922, p. 59) and Tortonese (1950a, p. 9), respectively.

All radiographs of the chondrocrania were made on type M Kodak
Industrial film with a “hard ray”? machine (Miller, 1957, pp. 29-40).

6 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

Additional proportional measurements, other than those found in
the tables, were also taken. These have not been included, however,
since they give no information not already expressed by the measure-
ments in the tables.

TOTAL LENGTH

ANTERIOR (UPPER) MARGIN

i+ LENGTH oF HEAD ——_____» DISTAL MARGIN

'

: 1
'

HEIGHT OF DORSAL FIN
i ANTERIOR (UPPER) MARGIN
+ ORIGIN

;

POSTERIOR (REAR) EXTENSION

A DORSAL

PRECAUDAL PIT
LENGTH OF SECOND DORSAL

DORSAL LOBE

:
{ NICTITATING MEMBRANE
:

SUBTERMINAL NOTCH

LENGTH OF DORSAL LOBE

Jo-" CAUDAL
* (TAIL) FIN

POSTERIOR (INNER) MARGIN ce

1 '
'
GREATEST DEPTH
F ’

‘
'
:

Reed enm op

P i LENGTH OF

: en, /

i NOSTRILS ¢ : See

TOR NARES i

SECOND DORSAL —__——

LEAST DEPTH OF
CAUDAL PEDUNCLE

PELVIC FIN SECONDARY KEEL

——s 6
{ LENGTH OF } YENTRAL LOBE

® .
} SNOUT '
’ '

INSERTION

ees eeeeeeeeeeee

INSERTION LENGTH OF PECTORAL FIN

I
PREORAL LENGTH.

\— CONCAVE OR FALCATE EDGE LENGTH OF VENTRAL LOBE

‘STRAIGHT EDGE OF FIN
VENTRAL PRECAUDAL PIT

LENGTH OF BASE OF FIN LENGTH OF CAUDAL PEDUNCLE

PECTORAL FIN

a

CONYEX EDGE

LENGTH
OF SNOUT LABIAL GROOVE

ASYMMETRICAL CAUDAL FIN

DISTANCE BETWEEN NOSTRILS

GILL SLITS
NOSTRILS OR NARES

VENTRAL SIDE OF HEAD NEARLY SYMMETRICAL CAUDAL FIN

Figure 1.—External features of a shark with indications for measurement: a, lateral view
of body; b, ventral view of head and comparison of caudal fins. (Drawings by Craig
Phillips, from Garrick and Schultz, 1963, pp. 5, 6.)

As indicated in plates 7c, p, one of the main chondrocranial features
separating Sphyrna corona and S. media is rostral cartilage length.
This difference has been expressed as a ratio by dividing the rostral
cartilage length by the distance between the tips of the preorbital
cartilages (fig. 17). Values were obtained by measuring the distances

}

NO. 3539 HAMMERHEAD SHARKS—GILBERT ZL

between the points marked by X (rostral cartilage length) and Y
(distance between tips of preorbital cartilage) in figure 3.

In addition, S. corona and S. media may also be distinguished by
difference in snout length. The values have been expressed as a
ratio by dividing snout length by head width; these are compared in
figure 16.

In general, determination of the patterns of the Ampullae of Loren-
zini on the ventral surface of the head becomes progressively easier
with an increase in size of the individual. This is subject to variation,
however, even within the same series of specimens.

MEDIAN INDENTATION
ANTERIOR-MEDIAN PORES

LATERAL INDENTATION
NARIAL OPENING ANTERIOR-LATERAL PORES
SUBNASAL PORES INNER NARIAL GROOVE

SUBOCULAR PORES

\i Zn

OUTER NARIAL GROOVE

MOUTH WIDTH
PREORAL PORES

POSTERIOR-LATERAL PORES
HEAD WIDTH

Ficure 2.—Ventral side of head, showing important morphological features and mucal pore
patterns (terminology of mucal pores adapted from Tortonese, 1950a, p. 9; drawing by
Paul Laessle).

Since a whole specimen of Sphyrna coward: was not available and
the head examined was seen after most of the figures were prepared,
this species has not been included in the species key, in most of the
tables, or in figures 4 and 22. Also, no diagnosis or description is
given under the account of the species.

Information regarding life histories, morphometric changes re-
sulting from growth, and zoégeography are usually included in sep-
arate sections rather than in the individual species accounts. If, in
a species account, there is no discussion of morphological variation, it
may be assumed that no noteworthy variation was discovered. New
names, new combinations of names, and information pertaining to
original descriptions and type material have been included in the
synonymies; if the nomenclatural history of a species is of sufficient
complexity, additional discussion is presented elsewhere in the species
account. More complete synonymies, based on references for the

8 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

western Atlantic region, may be found in Bigelow and Schroeder
(1948).

Localities from which specimens were examined are represented on
the distribution maps by solid symbols; literature records are repre-
sented by open symbols.

In the tabulation of proportions values have been separated, as
much as practicable, by geographic area. In cases where paucity of
specimens does not permit an adequate comparison, and if the speci-
mens do not appear to be different from those from other areas,

ARC RF RN

Figure 3.—Diagram of hammerhead chondrocranium (terminology adapted from Daniel,
1922, p. 59): AF (anterior fontanelle), AMC (anterior mediorbital connection), ARC
(accessory rostral cartilage), IOC (innerorbital cartilage), OC (olfactory cartilage), OW
(olfactory wing), PMC (posterior mediorbital connection), POP (postorbital process),
PRC (preorbital cartilage), PRP (preorbital process), RC (rostral cartilages), RF (rostral
fenestra), RN (rostral node), RW (rostral wing). (Drawing by Paul Laessle).

values have been combined. An exception to this was made in the
case of the single specimen of Sphyrna tudes from the Mediterranean.
This specimen is the holotype, and a comparison with material from
the western Atlantic was desirable.

Classification

The hammerhead sharks have, in the past, been accorded family
status by most of the investigators responsible for the major works on

NO. 3539 HAMMERHEAD SHARKS—GILBERT 9

fish classification (Gill, 1861, pp. 403-412; Jordan, 1923; Berg, 1940),
while others (Regan, 1906, pp. 722-758; Norman, unpubl. ms.) have
regarded this group as a subfamily of the Carcharhinidae. Berg’s
action apparently was based primarily on the studies by White (1937).

Although White (1937) retained the Sphyrnidae as a distinct family,
she gave no convincing reasons supporting this decision. Little was
said about the relationships of the sphyrnids and carcharhinids, and
the only places where they were compared were in the key and in the
discussion of the pelvic claspers. While the pelvic claspers obviously
constitute a valuable taxonomic character for the separation of various
species, their value in determining interspecific relationships has been
subject to question. According to White (1937, p. 94), ‘“They [the
claspers] tend to vary within the order [Galea] . . . so inconsistently
that they are of little value in establishing relationships.” Final
judgment should be reserved until a more intensive study is made of
most or all the species.

The sphyrnids and carcharhinids share a large number of mor-
phological characters. Both have (1) the first dorsal fin anterior to
the pelvic fins; (2) no naso-oral groove; (3) the keels of the dermal
denticles complete and dentate on the posterior edge and situated on a
flat surface; (4) a nictitating membrane; (5) teeth few in number,
never multicuspidate; (6) an asterospondylic type of vertebrae, which
has the appearance of a ‘‘maltese cross”’ when viewed in cross section;
(7) a “scroll type” spiral valve; (8) three rows of valves on the conus
arteriosus; and (9) an ovoviviparous type of reproduction.

The fact that the sphyrnids and carcharhinids have so many basic
characters in common suggests a close relationship; however, most
of these features are found also in other families, some of which are
not intimately related to these two groups. The “maltese cross”
pattern of the vertebrae is as well developed in the family Triakidae
as it is in the carcharhinids and sphyrnids. In addition, the nicti-
tating membrane, which has been thought to be unique to the latter
two families, also occurs in the triakids (V. G. Springer and J. A. F.
Garrick, in litt.).

Recent unpublished studies by Dr. Shelton P. Applegate indicate
that the hammerheads, in addition to the pronounced lateral expansion
of the head, differ from carcharhinid sharks in the following ways:
(1) The vertebrae, in cross section, show a convexity of the uncalcified
canal of the neural and haemal arches that is “‘butterfly-like” in shape,
rather than straight, and (2) in the teeth the lower blade has a smooth,
symmetrical arc, whereas in carcharhinid genera there is a definite
break in the arc on the side away from the crown. I regard these
differences as sufficient justification for continued recognition of the
hammerheads as a distinct family.

10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Genus Sphyrna Rafinesque

Cestracion Klein in Walbaum, 1792, p. 580. (Type species, Squalus zygaena
Linnaeus, by subsequent designation of Gill, Ann. Lyceum Nat. Hist. New
York, vol 7, p. 403, 1861; name inadmissable, as result of International
Commission rulings of 1907 and 1910.)

Sphyrna Rafinesque, 1810, pp. 46, 60. (Type species, Squalus zygaena Linnaeus,
by subsequent designation of Jordan and Gilbert, U.S. Nat. Mus. Bull. 16,

p. 26, 1883.)
Sphyrnias Rafinesque, 1815, p. 93. (Name substituted for Sphyrna Rafinesque,

1810.

ne Blainville, 1816, p. 121. (Type species, Squalus zygaena Linnaeus,
by subsequent designation of Fowler, Bull. Geol. Surv. New Jersey, vol 4,
Da Gigel Ot.)

Zygaena Cuvier, 1817, p. 127. (Type species, Squalus zygaena Linnaeus, by
absolute tautonymy; genus name preoccupied by Zygaena Fabricius, 1775,
for Lepidoptera.)

Zygoena Risso, 1826, p. 125. (Hmended spelling for Zygaena Cuvier, 1817.)

Sphyrnichthys Thienemann, 1828, p. 408. (Substituted for Sphyrna Rafinesque,
1810.

Zygana a alu 1839, p. 318. (Apparently a typographical error; should be
spelled Zygaena.)

Platysqualus Swainson, 1839, p. 318. (Type species, Squalus tiburo Linnaeus, by
original designation.)

Sphyra van der Hoeven, 1855, p. 68. (Emended spelling for Sphyrna Rafinesque,
1810.

Reniceps faa 1861, pp. 4038, 412. (Type species, Squalus tiburo Linnaeus, by
original designation.)

Eusphyra Gill, 1861, pp. 403, 412. (Type species, Zygaena blochiit Cuvier, by
original designation.)

PuyLoGeny.—There has, in the past, been some question as to
whether the genus Sphyrna has had a monophyletic or polyphyletic
origin. Members of the genus are characterized by certain apparently
basic morphometric characters, in addition to the widely expanded
head, which indicate derivation from a single common ancestor. The
most important of these are: No middorsal ridge on the posterior
part of the back; dermal denticles not closely overlapping; teeth
moderately to strongly oblique; labial furrow either small or absent;
body deeply compressed; pectoral fin short. Of the above characters,
the last two may have evolved in direct response to the lateral widening
of the head, and thus should be viewed with caution. ‘The remaining
characters are not likely to have been affected by such a change, and
are thus regarded as fundamental evidence of a close relationship
among the species of Sphyrna. Perhaps the most convincing study
supporting a monophyletic derivation has resulted from the work by
Starr and Fosberg (1957, pp. 292-295, figs. 1 and 2), in which serum
proteins were analyzed using electrophoretic techniques. Work done
by various investigators (the most important and extensive being that
of C. G. Sibley, of Cornell University, on birds) has shown that the

NO. 3539 HAMMERHEAD SHARKS—GILBERT 11

patterns resulting from these analyses may be valuable indicators of
higher relationships in the various groups of vertebrate animals. In
the paper by Starr and Fosberg, analyses were made of the serum
proteins from one specimen of Sphyrna lewini, two specimens of
Sphyrna tiburo, two of Rhizoprionodon terraenovae, and two of Aprion-
odon isodon. The patterns of the latter two species, both of which
are members of the family Carcharhinidae, differed notably from each
other and from the patterns in the two species of Sphyrna. On the
other hand, the patterns in Sphyrna lewini and S. tiburo showed a
marked similarity, thus suggesting a close phylogenetic relationship.
It would be desirable to run similar tests on Sphyrna blochi, which is
the most divergent species of hammerhead and the one most likely
to have evolved independently. Applegate (in litt.), however, informs
me that the teeth in S. blochii are similar to those of other hammer-
heads, although he has not yet studied the vertebrae of this species.
In all likelihood, the family Sphyrnidae has had a monophyletic origin.

Sphyrna tudes, Sphyrna corona, and Sphyrna media show a closer
phylogenetic relationship than any other species of hammerheads
(fig. 4). All possess a number of common morphological features,
some of which are not found in other members of the genus (table 1,
figs. 14-22, and pls. 7 c,p and 8). In addition, the three species
share several other characters which, although not confined to them
alone, further demonstrate a close relationship. The form in which
the various characters most closely approach those found in the
above species is S. tiburo (table 1, figs. 4, 11, 21, and 22, and pls. 7 a, B);
as a result, these four apparently closely related species are grouped
together in the subgenus Platysqualus. The respective lengths of
the anal fin and anal fin base, the relative lengths of the gill slits,
the diameter of the orbit in relation to the distance from the anterior
edge of the orbit to the outer margin of the narial opening, the relative
positions of the orbits and anterior part of the mouth, the presence
of a lower precaudal pit, the appearance of the upper precaudal pit,
and several chondrocranial characters all are similar in these four
species. Finally, the species are comparable in size, all apparently
reaching a maximum size of about 5 feet.

Within the subgenus Platysqualus, Sphyrna media seems to show
a slightly closer relationship to S. tiburo than does S. corona. This
assumption is based, first, on the greater overall similarity of the
chondrocrania (pls. 7 A-p), as well as similarities in appearance of the
anterior-median pore patch (figs. 22e-h).

The fifth species, Sphyrna blochii, is strikingly different from all
other species of Sphyrna in several characters. The extreme expansion
of the head, the presence of an outer narial groove, the greatly decreased
total number of vertebrae, and the marked enlargement of the verte-

12 PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 119

brae situated over the posterior part of the coelom are all unique to
this species, and it is consequently placed in a separate subgenus,
Eusphyra. 8S. blochii is so different from the other hammerheads that
recognition of Husphyra as a genus would not be unwarranted.

Contrary to the relative ease with which the relationships of the
preceding species are discerned, those of Sphyrna zygaena, S. lewini,
S. couardi, and S. mokarran are more difficult. The main feature
indicating a fairly close interrelationship among these species is
similarity in maximum size. All reach a total length of at least 12
feet, more than twice that attained by any other members of the genus.
S. zygaena and S. lewint (and probably S. couardi) have certain
features in common, such as a low second dorsal fin with a long lobe;
a short anal fin base, the length of which is never more than slightly
oreater than the pectoral base; a long slender anterior part of the
preorbital process of the chondrocranium; a deeply falcate anal fin;
and a deep, widely V-shaped upper precaudal pit, with a definite ridge
on the anterior margin. This suggests that they probably are more
closely related to each other than to S. mokarran (fig. 4, table 1,
and figs. 6-10).

The following examples, however, illustrate the difficulties one
encounters in attempting to determine the relationships of S. zygaena,
S. lewini (including S. couardi) and S. mokarran: The overall shape of
the rostral node, anterior-median pore patch, and anterior part of the
preorbital process in the chondrocranium of Sphyrna zygaena are most
similar to those found in S. blochii; S. zygaena and S. mokarran are
the only hammerheads lacking a wing on the olfactory cartilage; S.
lewint and S. mokarran are the only members of the genus that ordi-
narily have a rostral fenestra; S. mokarran and S. blochia have, as do S.
tudes, S. media, S. corona, and S. tiburo, a rather tall second dorsal fin;
S. mokarran has, as do S. tudes, S. media, S. corona, and S. tiburo, a
blunt, clublike knob at the end of the anterior part of the preorbital
process of the chondrocranium; and S. mokarran is unique, among the
hammerheads, in having a deeply falcate pelvic fin.

Despite the difficulties encountered in interpreting the relationships
of the above four species, they appear to be related more closely to one
another than to any other group of species of Sphyrna and thus are
included in the subgenus Sphyrna.

In the preceding discussion, nothing was said regarding head width, |
although this is almost the only basis for previous separations of the |
genus Sphyrna into separate genera and subgenera. For example, —

Tortonese (1950a, pp. 3-6) placed Sphyrna tiburo in the subgenus

Platysqualus and placed Sphyrna blochii in a separate genus, Husphyra. —
Although I have followed both of these allocations (in modified form), |

NO. 3539 HAMMERHEAD SHARKS—GILBERT 13

the differences in head width have had comparatively little to do with
my decision.

The phylogenetic conclusions may be summarized as follows:

1. The nine species of Sphyrna, on the basis of morphological
evidence and electrophoretic analyses of serum proteins, likely evolved
from a single common ancestor.

2. Sphyrna tudes, Sphyrna corona, and Sphyrna media, with
numerous identical morphological characters, some of which are
unique, are regarded as the most closely related members of the
genus Sphyrna.

3. Sphyrna tiburo, with a number of morphological characters
similar to those in S. tudes, S. corona, and S. media, appears to occupy
a phylogenetic position intermediate between these three species and
other members of the genus Sphyrna. Therefore, because of their close
relationship, the four species are placed in the subgenus Platysqualus.

4. Sphyrna blochit possesses several characters that are markedly
different from those found in other hammerheads, such as an ex-
tremely expanded head, an outer narial groove, a reduced number of
vertebrae, and enlarged vertebrae over the posterior part of the
coelom. It is placed in a separate subgenus, Husphyra, although
elevation of this taxon to the generic level might be justified.

5. The interrelationships of Sphyrna zygaena, S. lewini, S. couardi,
and S. mokarran are more difficult to determine, although zygaena,
lewint, and couardi seem to show a closer relationship to each other
than to mokarran. 'The most important feature shared by the four
species is their large maximum size. They are referred to the sub-
genus Sphyrna.

GROWTH CHANGES.—Certain modifications in shape of the head of
sphyrnids occur with increase in size and age, for which there is
nothing comparable in other carcharhinid sharks. In young hammer-
heads, particularly embryos, the anterior margin of the head is more
or less broadly rounded, becoming gradually straighter with increase
in size and age until sometimes, as in Sphyrna mokarran and Sphyrna
tudes, this margin is nearly straight (figs. 7a and 20c). In Sphyrna
blochii, which has the most widely expanded head of any member of
the genus, the lateral extensions of the head in embryonic and early
postembryonic specimens are nearly parallel to the body axis (fig.
5c). With an increase in size of the individual, these structures
become situated nearly perpendicular to the body (fig. 5b). It has
long been thought that such changes in head shape would make
hammerhead taxonomy a difficult problem. Such is not the case.

Those head characters of greatest importance in distinguishing
species of sphyrnids, such as structure of the chondrocranium, nature
of the intranarial and extranarial grooves, and pattern of the mucal

14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

pores on the ventral surface of the head, appear to change very little
throughout life, whereas the shape of the fins may be modified with
an increase in body size. Other postcephalic features seem to remain
fairly constant.

Negative allometric growth occurs in the head of sphyrnids since,
with an increase in total body length, there is a proportional decrease
in the following characters: Width of head, tip of snout to nares, tip
of snout to orbit, tip of snout to symphysis of jaws, diameter of orbit,
and intranarial width.

The shape of the upper precaudal pit, changing with an increase in
size of the individual, undergoes the greatest modification during

S.BLOCHII S.ZYGAENA S.TIBURO TIBURO S.MEDIA S.TUDES

S.MOKARRAN S. LEWINI S.TIBURO VESPERTINA S, CORONA

SUBGENUS PLATYSQUALUS

SUBGENUS SPHYRNA

SUBGENUS EUSPHYRA HYPOTHETICAL PHYLOGENY
OF THE GENUS SPHYRNA

CARCHARHINID ANCESTOR

Ficure 4.—Hypothetical phylogeny of the genus Sphyrna (figure by Paul Laessle). (S.
couardi, which was not included, is closely related to S. lewint.)

early growth and likely reaches its final form before the shark is 1,000
mm. long; this is ulustrated, from specimens of Sphyrna media
measuring 356 and 900 mm. TL, in figures 21f, g.

Other morphometric changes are indicated with an increase in size
(tables 2-9), although more large specimens must be examined before
the validity of these changes can be established. Pronounced modifi-
cations, particularly in length of the trunk between the axil of the
pectoral fin and the insertion of the pelvic fin, have been shown to
occur in squaloid sharks (Garrick, 1960, pp. 546-548). Since most
squaloids do not reach a very large size, however, considerable changes
in body proportions can occur with a relatively small increase in total

NO. 3539 HAMMERHEAD SHARKS—GILBERT 1

body length. The galeoid sharks, on the other hand, usually reach a
much larger size, and thus proportional changes are not likely to be
so pronounced within a comparable size range.

As indicated previously, a straightening of the fins and fin lobes
occurs as the shark approaches maturity. In this process the fins
(particularly the first dorsal) become increasingly more erect and
pointed, whereas the anal fin tends to become somewhat more falcate.
In contrast to these true morphometric changes are the unfolding of
the fins after birth, a phenomenon that might be compared with the
situation in a newly emerged butterfly. Curling of the fins is most
pronounced in embryos of Sphyrna mokarran, although the reason for
this is not clear.

The dermal denticles undergo a noticeable morphological change
with an increase in size and age, showing an increased number of
“teeth”? on the posterior margin. As indicated by Bigelow and
Schroeder (1957, p. 24), however, this does not result from changes in
the original denticle, but instead occurs as the result of one or more
denticle replacements.

In Sphyrna lewini there is a progressive darkening of the tips of the
pectoral fins with increasing size and age, so that in large specimens
these extremities are black. This also may be true to a greater or
lesser degree for S. zygaena and S. mokarran, although I have not yet
substantitated it.

Variation among specimens of the same species is discussed under
the accounts of the individual species.

Lirz HISTORY AND EcoLoGy.—No detailed discussion of hammer-
head ecology is presented here, since most of the available information
has appeared in works by Springer (1939, pp. 9-41; 1940b, pp. 188-
194), Bigelow and Schroeder (1948, pp. 407-449), and Baughman and
Springer (1950, pp. 96-152). The following is a summary of the in-
formation appearing in their papers.

The larger species of Sphyrna (S. zygaena, S. mokarran, S. lewint,
and, presumably, S. cowardi) apparently occur both in inshore and
offshore waters and have even been known to move into brackish
areas. SS. tiburo, in contrast, seems to prefer shallow embayments
and rarely ventures far from land. Little is known about the habitat
of the remaining three species.

Hammerheads swim near the surface of the water. It is doubtful if
any species descends to very great depths, since there have been no
verified captures below a hundred fathoms (Stewart Springer, in litt.).

Sphyrnids feed largely on fish and squid and also may take such
items as shrimp, crabs, stingrays, and sharks, including individuals of
their own species. The larger hammerheads are regarded as danger-

16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

ous, and there are substantiated records of fatal attacks on humans
(Gudger, 1937, pp. 417, 418).

Hammerheads along the eastern North American coast are known
to migrate northward in summer and southward in winter, often
traveling in schools in both inshore and offshore waters (Bigelow and
Schroeder, 1948, pp. 442-444). To the north, Sphyrna zygaena is the
only species involved in such migrations, whereas, to the south, S.
lewini is the predominant species. The fact that the only specimens of
Sphyrna zygaena seen by me from south of the Carolinas (UMML
5639 and UMML 10444, from Miami, Fla.) were taken in winter
seems to confirm this. Presumably similar migrations occur in other
parts of the world.

Although S. media and S. corona have been taken in the same areas
from Panama Bay north to southern Mexico (map 5), in only one of
the collections examined (from Chiapas, Mexico) were the two ac-
tually taken together. Whether this is due to a partial ecological
separation or simply to lack of material is presently not known.

The number of young in a brood appears to be directly proportional
both to size of the individual and to maximum size of the species.
The smallest recorded brood (8) was found in S. tiburo, the largest
(38) in S. mokarran.

ZooGEoGRAPHY.—The present distribution of the hammerhead
sharks indicates, on the whole, a slow rate of evolution. The members
of the subgenus Sphyrna (S. zygaena, S. lewinr, S. couardi, and SV.
mokarran) apparently have undergone no intraspecific differentiation,
despite worldwide distributions (for three of the four species) and
long periods of separation of the various segments of their respective
populations. The five forms included within the subgenus Platy-
squalus (S. tudes, S. corona, S. media, S. tiburo tiburo, and S. tiburo
vespertina) are morphologically similar to one another, and thus pre-
sumably have undergone a more recent evolution. Only in the case
of S. tiburo is it possible to estimate how long it has taken for evolution
to have occurred, because the two subspecies probably differentiated
following the most recent (Pliocene) emergence of the Central Ameri-
can isthmus. However, the presence of individuals of this species
with a head shape more or less intermediate between typical tiburo
and vespertina indicates that the above explanation may not be as
simple as first appears. Possibly the initial separation goes farther
back and the intermediate examples have resulted from a partial
reamalgamation of the stocks.

Insufficient evidence exists to explain the present distribution of S.
tudes, S. corona, and S. media. Evolution of these three species is
almost certainly correlated with the various submergences and emer-
gences of the Central American land mass, although it is impossible

NO. 3539 HAMMERHEAD SHARKS—GILBERT 17

to determine the exact sequence of events. Judging from morpho-
logical evidence, however, S. tudes evolved first, later followed by
differentiation of the other two species. The presence of S. tudes
in both the Mediterranean and western Atlantic is unusual in that
few species display such a distribution pattern. This common occur-
rence may be due either to recent migration in one direction or the
other, or to a long period of isolation during which the two populations
have, for some reason, failed to differentiate to the species level.

The distribution of Sphyrna zygaena is a classic example of an
antitropical distribution (Ekman, 1953, p. 250). Presumably the
now-disjunct north-south distribution of this species (map 2) was
continuous during one or more of the Pleistocene glacial periods,
when equatorial waters were cooler than at present. As the glaciers
retreated and seas became warmer, S. zygaena gradually disappeared
from the tropical regions, perhaps remaining for a time in deeper
waters where temperatures were cooler (equatorial submergence).

One may conclude from present distributions that the subgenus
Platysqualus evolved in the western Tethys Sea, a prehistoric body of
water that formerly extended from the Eastern Pacific-West Indian
region to what is now southeast Asia (Ekman, 1953, pp. 63-67).
Should it have evolved in the Indo-Pacific region, one or more species
of the subgenus would likely still be found there, since environmental
conditions in this area have remained stable over a long period of time.
The subgenus Husphyra apparently evolved in the Indo-Pacific, as
the only included species, Sphyrna blochii, is restricted to that region.
It is impossible to determine the center of origin of the subgenus
Sphyrna, since three of the species in this group have a worldwide
distribution.

Key to Species of Sphyrna
(Based primarily on specimens 1,000 mm. TI or smaller)

la. Greatest maximum width of head almost 50 percent of total body length;
outer narial groove present; nares not present near eyes, situated more
than half distance from eyes to anterior-median notch; upper precaudal
pit as in figure 21a; anterior-median pores on ventral surface of head as in

figure 22a; chondrocranium as in plate 5. . . . Sphyrna blochii (Cuvier)
Indo-Pacific region, from Persian Gulf to northern Australia and southern
Philippines.

1b. Greatest maximum width of head not more than 32 percent of total body

length; outer narial groove absent; nares present near eyes; upper pre-

caudal pit not as in figure 21a; anterior- median pores on ventral surface of

head not as in figure 22a; chondrocranium not asin plate5...... 2

2a. Anal fin and base longer, former measuring from 9.8 to 13.5 percent, and

latter from 6.5 to 9.9 percent of total body length; fifth gill slit about as

long as first gill slit, both somewhat shorter than three middle slits;

fifth gill slit situated above insertion of pectoral fin; orbit smaller, its

horizontal diameter less than shortest distance from anterior edge of
219-937—67—_2

18

3a.

4a.

PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

orbit to outer margin of narial opening (at least in smaller specimens);
teeth relatively weak, median ones often more or less needle-like in
appearance; lower precaudal pit present (may also be present in S.
lewint); posterior lobe of first dorsal fin extending beyond insertion of
pelvic fin (in S. tudes, S. corona, and S. media only); maximum size
smaller, the largest specimens probably not over 4 or 5feet TL. . . 3
Head not widely expanded, greatest width about 21 percent of total

body length (usually less than 20 percent); anterior margin of head
evenly rounded or slightly pointed in median line, never with
median notch; posterior lobe of first dorsal fin terminating anterior to
insertion of pelvic fin; origin of first dorsal fin situated well posterior
to axil of pectoral fin; eyes not situated distinctly anterior to mouth,
line drawn between posterior margins of orbits either intersecting or
passing just anterior to symphysis of lower jaw; upper precaudal pit
as in figure 21le; anterior-median pores on ventral surface of head as
in figure 22e; chondrocranium as inplates 7aA,B.........4
Head with more evenly rounded anterior margin (fig. 12b); greatest
width of head measuring slightly less as shown in table 6; fig 13
Sphyrna tiburo tiburo (Linnaeus)
Western Atlantic, from New England to southern Brazil.

4b. Head with less evenly rounded anterior margin (fig. 12a), usually

coming to a point at tip of rostrum; greatest width of head measuring
slightly more as shown in table 6; fig. 13.

Sphyrna tiburo vespertina Springer
Eastern Pacific, from southern California to Equador.

3b. Head widely expanded, the least width 22 percent of total body length;

anterior margin of head not always evenly rounded, usually with a
more or less distinct median notch (sometimes evenly rounded in S.
media); posterior lobe of first dorsal fin terminating posterior to
insertion of pelvic fin; origin of first dorsal fin situated above or slightly
posterior to axil of pectoral fin; eyes situated anterior to mouth, line
drawn between posterior margins of orbits passing well anterior to
symphysis of lower jaw; upper precaudal pit not as in figure 21e;
anterior-median pores on ventral surface of head not as in figure 22e;
chondrocranium not asin plates 7A,B... . 5M an TRS are

5a. Inner narial groove present; well-defined median note present on

anterior margin of head; anterior-median pores on ventral surface
of head as in figure 22h; chondrocranium as in plates 8a, B.

Sphyrna tudes (Valenciennes)
Gulf of Mexico south to Uruguay; also recorded from western
Mediterranean Sea.

5b. Inner narial groove absent; no well-defined median notch present on

anterior margin of head (poorly defined notch may be present in
some individuals) ; anterior-median pores on ventral surface of head
not as in figure 22h; chondrocranium not as in plates 8a, B. . . 6

6a. Distance from tip of snout to symphysis of upper jaw slightly shorter,

usually less than 40 percent of head width (table 7; fig. 16); anal
fin more deeply faleate, with a more pointed apex (figs. 14a, 15a) ;
mouth slightly broader (figs. 14b, 156); anterior-median pores on
ventral surface of head as in figure 22f; chondrocranium as in
plate 7c; maximum size possibly larger, the largest specimen

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NO. 3539

HAMMERHEAD SHARKS—GILBERT 19

examined about 34 feet long . . . . .Sphyrna media Springer
Eastern Pacific, from Panama to Gulf of California; also in
southern Caribbean and southwestern Atlantic, from Panama to
Brazil.

6b. Distance from tip of snout to symphysis of upper jaw slightly longer,
usually more than 40 percent of head width (table 8; fig. 16);
anal fin less deeply faleate, with a more rounded apex (fig. 18a) ;
mouth slightly narrower (fig. 186); anterior-median pores on
ventral surface of head as in figure 22g; chondrocranium as in
plate 7p; maximum size possibly smaller, the largest specimen
examined a little over 2 feet long. . . Sphyrna corona Springer
Eastern Pacific, from Colombia to southern Mexico.

2b. Anal fin and base shorter, former measuring from 8.2 to 10 percent, and

7a. Inner narial groove absent; pelvic fin faleate; second dorsal fin higher,

latter from 3.8 to 6.7 percent of total body length; fifth gill slit shorter
than first gill slit (fourth gill slit about as long as first gill slit); fifth gill
slit situated behind insertion of pectoral fin; orbit larger, its horizontal
diameter greater than shortest distance from anterior edge of orbit to outer
margin of narial opening; teeth broad, not needle-like in appearance; lower

precaudal pit absent; posterior lobe of first dorsal fin not extending to
above insertion of pelvic fin; maximum size larger, some individuals
reaching) 20)feet) ins. is 7

length of its anterior margin greater than length of anterior margin of
anal fin; lobe of second dorsal fin shorter, not extending nearly to
caudal pit, and, when extended vertically upward, about equal to
greatest height of fin; teeth strongly serrate, even in small individuals;
anterior-median pores on ventral surface of head as in figure 22b;
chondrocranium as in plate 64 . . . . Sphyrna mokarran (Riippell)
Circumtropical in distribution, but apparently absent from the more
isolated oceanic islands.

7b. Inner narial groove present; pelvic fin not faleate; second dorsal fin

lower, length of its anterior margin less than length of anterior margin
of anal fin; lobe of second dorsal fin longer, extending nearly to caudal
pit, and, when extended vertically upward, definitely exceeding greatest
height of fin; teeth smooth in small individuals, often weakly serrate
in large individuals; anterior-median pores on ventral surface of head
not as in figure 226; chondrocranium not asin plate 6a...... 8

8a. Median notch absent from anterior margin of head; inner narial groove

extending at least halfway from narial opening to median part of
anterior margin of head; base of anal fin shorter, about equal in
length to base of second dorsal fin and slightly less than base of
pectoral fin; rostral fenestra ordinarily absent; anterior-median
pores on ventral surface of head as in figure 22c; chondrocranium as
in plate 6c.

Sphyrna zygaena (Linnaeus)
Antitropical in distribution; occurring in cooler waters of northern
and southern hemispheres, but absent from intervening tropical
waters.

8b. Median notch present on anterior margin of head; inner narial groove

not extending halfway from narial opening to median part of anterior
margin of head; base of anal fin longer, greater in length than bases

20 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

of either second dorsal or pectoral fins; rostral fenestra present;

anterior-median pores on ventral surface of head as in figure 22d.*
Sphyrna lewini (Griffith and Smith)

Circumtropical in distribution; common in all warm seas.

*Sphyrna couardi Cadenat keys out to this point, but was not included here since a whole specimen was
notexamined. It differs from S. lewini, however, in having a deeper and more robust head (pl. 9), in minor
differences in the chondrocranium (pl. 10), in having the anteriomedian pore patch completely instead of
partially divided, and (according to the original description) in having the tips of the pectoral fins white.

Dignaosis of Family and Genus

Distinguished primarily by the shape of the head, which is expanded
laterally, and which is accompanied by a corresponding modification
of the chondrocranium. Eyes situated at tips of lateral expansions,
and nares situated at varying distances along anterior margin of head.
Olfactory capsules and orbital region widely expanded, and the three
rostal cartilages broadly expanded where joined anteriorly. Midline
of back posterior to first dorsal fin without a middorsal ridge. Second
dorsal fin always much smaller than first dorsal. Body compressed,
the depth of the trunk (measured at origin of first dorsal) from 4 to 5%
times in distance from tip of snout to upper precaudal pit. Caudal
peduncle about two-thirds as wide as deep. Upper precaudal pit
strongly developed. Lower precaudal pit present in some species,
absent in others; when present, not as prominent as upper pit and only
about half as wide.

Subgenus Eusphyra Gill

The subgenus Husphyra, which includes only Sphyrna blochii, is
distinguished by the following unique characters: Head extremely ex-
panded, the greatest width from 42.0 to 49.2 percent of total body
length; outer narial groove present; total number of vertebrae few (117
and 124 in two specimens examined); body vertebrae 51 to 54, there
being from 15 to 16 fewer body than caudal vertebrae; vertebrae over
posterior part of coelomic cavity expanded in size; upper precaudal pit
merely a narrow longitudinal groove (fig. 21a); orbit separated from
nares by a distance many times diameter of orbit; a unique anterior-
median pore patch on underside of head, the patch characterized by a
sharp, pointed angle at outer-posterior corner and a more or less
straight posterior edge (fig. 22a); and various chondrocranial charac-
ters (pl. 5).

Sphyrna (Eusphyra) blochii (Cuvier)
Ficurss 5, 2la, 22a; Puatss 1, 5
Squalus zygaena (misidentification) Bloch, 1785, vol. 1, pl. 117. (Figure.)
Zygaena blochit Cuvier, 1817, p. 127. (Original description [in footnote]:

“Ajoutez Vespéce représentée par Bl. 117, reconnaissable & ses narines
placées bien plus prés du milieu [Z. Nob. Blochii]. Sa deuxiéme dorsale est

i I ta a aia a

en i eel

ee

No. 3539 HAMMERHEAD SHARKS—GILBERT 21

aussi bien plus prés de la caudale. L’espéce & large téte, donnée, sous le
nom de pantouflier.”’ Description based on figure in Bloch, 1785; no type
specimen; no type locality.)

Zygaena latycephala van Hasselt, 1823, p. 315. (Originally written Latycephala;
type locality, Java.

Zygaena laticeps Cantor, 1837, p. 315. (Original description; based on adult
specimen of S. blochi; type locality, Calcutta.)

Zygana laticeps Swainson, 1838, p. 134. (Illustration of head; generic name mis-
spelling of Zygaena.)

Sphyrna blochii Miller and Henle, 1841, p. 54. (Locality unknown.)

Sphyrnias blochit Gray, 1851, p. 50. (India; Singapore.)

Eusphyra blochii Gill, 1861, pp. 403, 412. (Name; new generic name.)

Cestracion (Zygaena) blochii Duméril, 1865, p. 383. (Indian Ocean.)

Sphyrna zygaena (misidentification) Bridge, 1904, p. 450. (Figure.)

Zygaena blochi Volz, 1907, p. 237. (Benkulen, Sumatra.)

Sphyrna blocht Pellegrin, 1912, p. 2. (Singapore.)

Cestracion blochit Garman, 1913, pp. 156-157. (Synonymy; description; range.)

Cestracion blochi Hora, 1924, p. 464. (Tale Sap, Siam.)

Sphyrna (Eusphyra) blochit McCulloch and Whitley, 1925, p. 129. (Reference.)

SPECIMENS EXAMINED.—(Range in size: 333-880 mm. TL). Pacific
Ocean: British North Borneo: SU 13815 (1), no further data;
FMNH 21836 (1), Sandakan. Burma: SU 14495 (1), FMNH 40914
(1), Southern Moscos group, South Island. India: SU 41989 (1),
Krusadai Island, Gulf of Mannar, Pumbau District, Madras; USNM
195846 (2), ANSP 69397 (1), Bombay. Java: FMNH 15655 (1),
FMNH 15656 (1), UMMZ 177111 (6), MCZ 1389 (1 mutilated head),
Batavia. Malaya: BMNH 1902.5.28.37 (1), coast of Perak; BMNH
1860.3.19.890 (2), Singapore.

NomeEnNciature.—Although Sphyrna blochii, in common with the
other hammerheads, has been placed in several different genera,
Eusphyra is the only genus which has been erected (Gill, 1861, p. 412)
for the sole reception of this species. Gill considered that two unique
morphological features of blochii (the position of the nares, which are
nearer the tip of the rostrum than the eyes, and the extreme elongation
of the lateral extensions of the head) constituted sufficient reason for
placing the species in a distinct genus. There has been some differ-
ence of opinion among recent investigators as to whether Husphyra
should be regarded as a good genus. In some of the most important
recent works (Bigelow and Schroeder, 1948; Tortonese, 1950a) this
classification has been followed, but in an even greater number of
cases it has been considered a subgenus of Sphyrna. For reasons that
are discussed elsewhere in this paper, I follow the latter alternative.

Diaqnosis.—See diagnosis of subgenus Husphyra.

Description.—Meristic data appear in table 2. Characters men-
tioned in the diagnosis of the subgenus Eusphyra are not repeated
here. A comparison of S. blochii with other members of the genus is
presented in table 1.

ae PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Intranarial distance from one-seventh to one-sixth of TL; anterior
margin of head broadly curved medially, the lateral expansions of the
head pointing posteriorly in embryonic individuals and extending
nearly perpendicularly from body in larger specimens; distinct median
scallop present on anterior margin of head; inner narial groove present
and distinct, extending about half of distance from narial opening to

ce ks
€

Ficure 5.—Sphyrna blochii: a, female, 622 mm. TL, from Sandakan, North Borneo (FMNH
21836); b, head of adult, 0.2 X natural size (FMNH 21836); c, head of juvenile, 333 mm.
TL, from Batavia, Java, 0.6 X natural size (UMMZ 177111) d, dermal denticles (FMNH
21836); e, teeth series from left side of upper and lower jaws (FMNH 21836). (Drawings
by Dorothea B. Schultz.)

median scallop; narial depression present; outer posterior corner of
lateral expansions of head situated posterior to corner of mouth in
specimens of all sizes; narial flap broad, blunt at tip, with the outer
edge curving gradually inward; length of snout about one-eighth of
head width; greatest transverse distance between corners of mouth
about one-seventh of greatest width of head; symphysis of jaws

NO. 3539 HAMMERHEAD SHARKS—GILBERT 23

situated posterior to a transverse line drawn between posterior margin
of orbits; labial furrows absent.

Origin of first dorsal fin above or slightly anterior to axil of pectoral
fin, a perpendicular line drawn ventrally from origin usually inter-
secting axil; fifth gill slit situated distinctly behind, and fourth gill
slit above (or very slightly posterior to) insertion of pectoral fin;
posterior lobe of first dorsal fin terminating well anterior to insertion
of pelvic fin; first through fourth gill slits nearly equal in length, and
all longer than fifth gill slit; first dorsal fin relatively slender, not
perfectly erect, a line drawn perpendicularly from apex of fin inter-
secting free rear margin of fin (posterior lobe) about halfway along its
length; base of first dorsal fin about three-fifths head length and
almost twice length of pectoral fin base; anterior margin of first dorsal
fin nearly straight, slightly curved near apex; upper half of distal
margin of first dorsal fin straight, the lower half falcate; lobe of first
dorsal fin (free posterior margin) slightly less than one-half length of
base; origin of second dorsal fin situated over posterior third of anal
base; second dorsal fin moderately tall, its greatest height five-eighths
to five-sevenths as long as base, and about five-sevenths greatest
height of anal fin; second dorsal fin base about four-sevenths length of
anal base; lobe of second dorsal fin long, its length about twice greatest
height of fin, and extending almost to upper precaudal pit; anterior
margin of pectoral fin three-fourths to four-fifths length of head, and
about equal to greatest height of first dorsal; pectoral fin about three-
fifths as broad as long; pectoral fin base four-fifths to seven-eighths
length of anal fin base; anterior margin of pectoral slightly convex;
distal margin of pectoral slightly faleate; apex of pectoral pointed;
inner corner of pectoral narrowly rounded; pelvic fin about five-sixths
length of anal fin; anal fin base slightly longer than pectoral and pelvic
bases and more than 1% times as long as second dorsal fin base; height
of anal fin about two-sevenths again as much as height of second
dorsal fin; anal fin deeply faleate toward apex and nearly straight
toward tip of lobe; length of caudal fin about one-third of TL; upper
margin of caudal slightly convex; terminal part of caudal about two-
ninths of total length of fin; posterior margin of terminal part of caudal
fin shallowly falcate; terminal apex of caudal broadly pointed; lower
lobe of caudal moderately narrow and short; length of lower caudal
lobe one-third to two-fifths as long as upper lobe; lower caudal lobe
not erect, sloping posteriorly at about a 35° angle from a perpendicular
to the body axis; anterior margin of lower caudal lobe convex; posterior
margin of lower caudal lobe nearly straight; lower precaudal pit
absent.

Anterior margin of rostral node of chondrocranium straight; acces-
sory rostral cartilages absent; anterior fontanelle (viewed ventrally)

24 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

appearing extremely broad and shallowly U-shaped, with a shallow,
broad, median notch; small, pointed wings present on sides of rostral
node; anterior wing of olfactory cartilage prominent, pointed, and
projecting medially; anterior part of preorbital process long and
slender; posterior part of preorbital process terminating in an angular

knob.

15-0-16
Teeth a=
in both jaws notably oblique, with none (except the median tooth)
perfectly erect, the teeth becoming progressively more oblique toward
corners of mouth, and those nearest the corners having cusps pointing
almost straight back; cusps visible on all teeth, although those near
corner of mouth in lower jaw poorly developed; teeth 6 through 10 m
upper jaw notably larger than others; tooth 1 notably smaller than
tooth 2 in lower jaw; teeth 2 through 10 in lower jaw gradually
increasing in size.

(one specimen examined); teeth not serrated; teeth

Map 1.—Distribution of species.
A Sphyrna blochii @ Sphyrna mokarran
(Solid=specimens examined; hollow=confirmed literature references.)

Dermal denticles (examined from upper part of back beneath first
dorsal fin) not closely overlapping, the blades steeply raised; length of
denticle (measured to tip of median marginal ‘‘tooth’’) as great, or
nearly as great, as width; five strong ridges usually present, and as
many bluntly pointed teeth usually present on posterior margin of
denticle, the outer ridges and teeth sometimes rudimentary or absent,
particularly in small individuals; median tooth on denticle ordinarily
slightly longer than more distal teeth; pedicel short and broad.

Individuals gray or grayish brown dorsally, becoming more pale
below; no markings on fins.

Maximum total length not recorded, possibly not more than several
feet.

Ranee.—Sphyrna blochii is restricted to the Indo-West Pacific area
and is one of the two species of hammerheads not found in the Western
Hemisphere. It is known from the Persian Gulf (Blegvad and
L¢ppenthin, 1944, p. 41, fig. 12) eastward throughout the East Indies,

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NO. 3539 HAMMERHEAD SHARKS—GILBERT 20

and north to southern China and the southern Philippines (Herre,
1930, p. 142). The species also has been recorded from Queensland,
Australia (Ogilby, 1908, p. 4); this area apparently represents both the
eastern and southern limits of its range.

Sphyrna blochii, while perhaps not as closely restricted to continental
waters as Sphyrna tiburo, nevertheless appears to be incapable of
traversing broad stretches of ocean, as evidenced by its absence from
outlying oceanic islands (map 1).

Subgenus Sphyrna Rafinesque

The subgenus Sphyrna, which includes S. zygaena, S. lewini, S.
couardi, and S. mokarran, is distinguished by only two unique char-
acters. These are: A large orbit, the greatest horizontal diameter
greater than distance from anterior margin of orbit to narial opening;
and a large maximum size, individuals reaching a size of from 12
to 20 feet in total length.

The subgenus 1s also distinguished by: A moderately expanded head,
the greatest width from 22.9 to 30.2 percent of total body length (in
specimens less than 1,000 mm. TL); first through fourth gill slits
nearly equal in length, and all somewhat longer than fifth gill slit;
anal fin and anal fin base relatively short, the former not over 11.0
percent, and the latter not over 7.5 percent, of TL; fifth gill slit situ-
ated distinctly behind, and fourth gill slit above (or very slightly
anterior to) insertion of pectoral fin; presence of a deep narial de-
pression; absence of an outer narial groove; lobe of first dorsal fin
terminating well anterior to insertion of pelvic fin; total number of
vertebrae ranging from 174 to 206 (usually 190 or more), with those
over posterior part of coelomic cavity not expanded in size.

Sphyrna (Sphyrna) mokarran (Riippell)
Ficures 6, 7, 216, 226, Plates 2, 6a

?Squalus tiburo (misidentification) Risso, 1810, p. 35. (Identification by descrip-
tion of head and reference to Lacepede, ‘‘Squale pantouflier”’; Nice; very rare.)

Zygaena tudes Valenciennes, 1822, pp. 225-226. (In part; original description;
illustration of head; type specimens apparently represent more than one
species; types from Mediterranean Sea, Cayenne, and Coromandel; specific
name tudes subsequently restricted to a species other than one here called
Sphyrna mokarran.)

Sphyrnichthys zygaena (misidentification) Thienemann, 1828, p. 408. (General;
identification probable because teeth described as “gezidhnelte.’’)

Zygaena mokarran Riippell, 1835, pp. 66-67. (Original description; figures of
head, teeth, and body; type specimen, a male 2510 mm. TL, Senckenberg
Museum no. 3590; type locality, Massaua, Red Sea.)

Sphyrna malleus (misidentification) L. Agassiz, 1838, pp. 235, 303. (Identification
probable because teeth serrate.)

Sphyrna mokarran Miiller and Henle, 1841, p. 54. (Red Sea.)

26 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Sphyrna tudes Miiller and Henle, 1841, p. 53. (In part; identification from
reference to Valencennes, 1822.)

?Sphyrna chiereghint Nardo, 1847, p. 111. (Name only.)

Sphyrnias mokarran Gray, 1851, p. 50. (In part; identification by reference to
Valenciennes, 1822.)

Sphyrnias tudes Gray, 1851, p. 50. (In part; identification by reference to Valen-
ciennes, 1822.)

Cestracion (Zygaena) mokarran Duméril, 1865, p. 383. (Compiled.)

Cestracion (Zygaena) tudes Duméril, 1865, p. 384. (In part; description from
Valenciennes, 1822.)

Squalus zygaena (misidentification) de la Blanchere, 1868, p. 499. (Teeth de-
scribed as ‘‘dentelees.’’)

Cestracion tudes Ninni, 1872, p. 10. (Not seen.)

Cestracion mokarran Bleeker, 1873, p. 115. (Chinese drawing.)

Zygaena dissimilis Murray, 1887, p. 103. (Original description; type locality,
Kurrachee.)

Sphyrna (Platysqualus) tudes Jordan and Evermann, 1896, p. 217. (Name;
West Indies and Mediterranean; references to specimens from Gulf of
California and Indian Ocean perhaps do not pertain to this species.)

Cestracion zygaena (misidentification) Radcliffe, 1916, p. 263. (Identification
by description of teeth in illustration as serrate; specimens 12 ft., 6 in.
long; Beaufort, N.C.)

Sphyrna zygaena (misidentification in part) Coles, 1919, p. 41. (Identification
by shape of head in photo, of 13 ft., 10 in. female; Cape Lookout, N.C.)
Platysqualus tudes Howell-Rivero, 1936, p. 44. (Identification probable because

of large size, 12 to 14 ft.; Cuba.)

Sphyrna lewini (misidentification in part) Fowler, 1941, p. 215. (Figure of S.
mokarran incorrectly labelled S. lewinz.)

Sphyrna ligo Fraser-Brunner, 1950, pp. 213-219. (Original description; figure;
holotype, an embryo, BMNH 1890.9.23.231; type locality, Clarence River,
New South Wales, Australia; radiograph of head seen.)

SPECIMENS BXAMINED.—(Range in size: 435-675 mm. TL; heads
of larger specimens also examined). Atlantic Ocean: Senegal: (pho-
tograph seen), Gorée. United States (Florida): USNM 106543 (1),
USNM 108453 (1), USNM 108454 (1), FMNH 63092 (1), MCZ
35247 (1), SU 14330 (1), Englewood; FMNH 530, Captiva Pass;
USNM 186087 (2), 25°56’ N., 81°47’ W.

Pacific Ocean: Australia: USNM 40014 (1), USNM 40026 (1),
MCZ 969 (1), Richmond River, New South Wales; BMNH 1890.9.23.
231 (x-ray of head of holotype of Sphyrna ligo), Clarence River,
New South Wales. India: USNM 12622 (1), Madras. Mexico:
USNM 29645 (1), USNM 37162 (1), Mazatlan; UCLA 52-252 (1),
Bahia Almejas, Isla Santa Margarita, Baja California.

NoMENCLATURE.—Zygaena mokarran was described by Riippell
(1835, pp. 66-67, pl. 17, fig. 3) from the Red Sea, at Massaua. The
illustration (pl. 2), though inaccurate in certain details, particularly
the conformation of the fins, is readily identified by the shape of
the head. In addition, the type specimen (Senckenberg Museum
no. 3590) has been illustrated recently by Klausewitz (1960, p. 293).

NO. 3539 HAMMERHEAD SHARKS—GILBERT Di

For many years the species herein called Sphyrna mokarran was
known as Sphyrna tudes. 'Tortonese (1950b, pp. 1030-1033) was the
first to point out that the three type specimens of Zygaena tudes
actually comprise two species, one of which may be S. mokarran.
Only two of the types are apparently now extant (one from Nice,
France; the second from Cayenne, French Guiana), neither of which
is S. mokarran. However, since the missing specimen is from an
area (Coromandel, India) where the species represented by the two
existing types does not occur, and since there is a fairly close super-
ficial resemblance between the two species in question, one can assume
that the Coromandel individual was probably Sphyrna mokarran.

Ficure 6.—Sphyrna mokarran: a, juvenile male, 673 mm. TL, from Englewood, Fla.
(USNM 106543); b, head, 0.2 * (USNM 106543); c, dermal denticles, about 49
(USNM 106543). (From Bigelow and Schroeder, 1948, p. 430.)

Diacnosts.—Characters mentioned in the diagnosis of the sub-
genus Sphyrna are not repeated here.

Sphyrna mokarran is unique among members of the genus Sphyrna
in having a deeply falcate pelvic fin and strongly serrated teeth at all
sizes. S. mokarran is also characterized by a deep median indentation
(scallop) on anterior margin of head; absence of an inner narial groove,
this structure represented by a thin line; a tall second dorsal fin, the
height of the fin slightly greater than height of anal fin; a short second
dorsal lobe, the length of the lobe equal to height of fin and extending
only about one-half of distance from posterior part of fin base to upper

28 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

precaudal pit; a shallow upper precaudal pit, with an acutely pointed
posterior ridge to upper precaudal pit (fig. 216); an anterior-median
pore patch in which the outer-posterior corners are sharply angled
and the posterior edge is straight (fig. 22b); presence of a rostral
fenestra, and several other chondrocranial characters (pl. 64).

A comparison of S. mokarran with other members of the genus is
presented in table 1.

DescriptTion.—Meristic data appear in table 3. Characters men-
tioned in the diagnoses of the subgenus Sphyrna and S. mokarran are
not repeated here.

IE NIN eee ar SS Se =

Va ENN mi Ty NO 2

a Mi Ach tye |
ba eee

Ficure 7.—Sphyrna mokarran: a, head of small adult, 1,660 mm. TL, from Englewood, Fla.,
0.09 * natural size (USNM 108453); b, teeth series from left side of upper and lower
jaws (USNM 108453), about natural size; c, fifth upper tooth; d, twelfth upper tooth;
é, fifth lower tooth; f, eleventh lower tooth. (c-f, about 1.5 X.) (From Bigelow and
Schroeder, 1948, p. 429.)

Head moderately expanded, the greatest width from 22.9 to 27.3
percent of TL (in specimens less than 1,000 mm. long); shallow scal-
lops present on anterior margin of head about halfway between nares
and tip of snout; outer posterior corner of lateral expansion of head
posterior to corner of mouth in smaller specimens, anterior to corner of
mouth in larger individuals; narial flap broad, broadly pointed at tip,
with the outer edge curving abruptly inward; orbit large, the hori-
zontal diameter almost equal to length of shortest (fifth) gill slit;

NO. 3539 HAMMERHEAD SHARKS—GILBERT 29

length of snout from one-third to one-fourth of head width; greatest
transverse distance between corners of mouth about one-fourth of
sreatest width of head; symphysis of jaws situated posterior to a trans-
verse line drawn between posterior margins of orbits; labial furrows
absent.

Origin of first dorsal fin above axil of pectoral fin, a perpendicular
line drawn ventrally from origin intersecting at, or slightly behind,
this point; first dorsal fin tall, slender, and pointing strongly posteri-
orly, a line drawn perpendicularly from apex of fin passing well back
of fin lobe; base of first dorsal fin slightly less than one-half head
length and almost twice length of pectoral fin base; anterior margin
of first dorsal fin broadly curved (excessively curved in embryonic
and early post-embryonic individuals); upper two-thirds of distal
margin of first dorsal fin straight or slightly faleate toward tip, the
lower one-third distinctly falcate; lobe of first dorsal fin about one-
third length of base; origin of second dorsal fin situated slightly behind
origin of anal fin; second dorsal fin base slightly greater than four-
fifths length of anal fin base; anterior margin of pectoral fin slightly
longer than one-half length of head and a little less than greatest
vertical height of first dorsal fin; pectoral fin two-thirds as broad as
long; pectoral fin base about four-fifths length of anal fin base;
anterior margin of pectoral broadly rounded (strongly curved in
embryos and early post-embryos); distal margin of pectoral broadly
falcate; apex of pectoral broadly pointed; inner corner of pectoral
broadly rounded; length of pelvic fin almost equal to length of anal fin;
pelvic fin base slightly less than length of anal base; anterior margin
of pelvic nearly straight, except near apex, where it is broadly curved;
length of anal fin slightly greater than length of pelvic fin; anal fin
base slightly (up to one-sixth) longer than pectoral, pelvic, and second
dorsal fin bases; anal fin deeply falcate toward apex, nearly straight
toward tip of lobe; length of caudal fin nearly a third of TL; upper
margin of caudal slightly convex; terminal part of caudal about one-
fourth of total length of fin; posterior margin of terminal part of caudal
fin notably falcate; terminal apex of caudal broadly pointed; lower
lobe of caudal narrow and long, the distance from angle included by
the upper and lower caudal lobes to tip of latter going about 214
times in anterior margin of lower caudal lobe; length of lower caudal
lobe one-third to two-fifths as long as upper lobe; lower caudal lobe
not erect, sloping posteriorly at about a 45° angle from a perpendicular
to body axis; anterior margin of lower caudal lobe convex; posterior
margin of lower caudal lobe nearly straight or slightly concave.

Anterior margin of rostral node of chondrocranium straight;
accessory rostral cartilages absent; anterior fontanelle (viewed
ventrally) shallow and nearly straight on posterior margin; prominent,

30 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

broad, bluntly rounded wings present on sides of rostral node; no
anterior wing on olfactory cartilage; tip of anterior part of preorbital
process very short, blunt, and rather thick; posterior part of pre-
orbital process terminating in an evenly rounded knob.

Total number of vertebrae 205 to 206 (in two specimens examined) ;
body vertebrae 95 to 98, there being from 10 to 11 fewer body than
caudal vertebrae.

L7—2 (OGianl is :

Teeth pei oa aa teeth triangular, on expanded
bases, the first tooth erect, symmetrical, but the subsequent upper
teeth increasingly oblique toward corners of mouth; second to tenth
or eleventh teeth the largest in both upper and lower jaws; outer-
most teeth in both jaws with distinct cusps; one or two series of teeth
functional in alternating rows along sides of upper jaw and usually
two rows along sides of lower jaw.

Dermal denticles (examined from upper part of back beneath first
dorsal fin) evenly and closely spaced, but with skin often visible in
between, the blades thin and little arched; length of denticle (measured
to tip of median marginal “‘tooth’?) about same as width; small
specimens with from three to five, larger specimens with five or
even six, smooth-topped ridges, and as many short, broadly pointed
marginal teeth on dermal denticle; median tooth on denticle slightly
longer than more distal teeth; pedicel short and thick.

Individuals with gray or grayish brown dorsally, becoming more
pale below; no markings on fins.

Maximum total length reputedly about 20 feet.

VaRIaAtTIon.—Comparatively few specimens of Sphyrna mokarran
were available for study, and of these a large percentage (including
all individuals from the Indo-Pacific region) were very small embryos.
Specimens of this species are characterized by strongly curled bodies
at this stage of development, and this, together with the paucity of
specimens, tends to negate apparent differences, such as orbital
diameter and length of second dorsal fin base, between specimens
from the Atlantic and Pacific regions (table 3). In addition, radio-
graphs of the chondrocrania show no apparent differences. For
these reasons, as well as the fact that neither of the other hammer-
heads of worldwide distribution (S. zygaena and S. lewini) show any
noticeable intraspecific differentiation, it seems best to recognize
but one species, Sphyrna mokarran.

Ranee.—Sphyrna mokarran is circumtropical in distribution (map
1). However, it does not usually appear to be as common as Sphyrna
lewini, a species occupying a similar range.

S. mokarran is found in the western Atlantic, including the Gulf
of Mexico and Caribbean Sea, from North Carolina south to Brazil.

NO. 3539 HAMMERHEAD SHARKS—GILBERT 31

It is present in the eastern Atlantic, at least off the northwest coast
of Africa (Cadenat, 1951, pp. 96-97) and in the Red Sea. Despite
its presence in the last two areas, there are no substantiated records
from the intervening Mediterranean (Tortonese, 1950a, p. 36).
This is probably not due to lack of adequate collecting, as certain
other species, such as the carcharhinid shark Phizoprionodon acutus
(Victor G. Springer, pers. comm.), show a similar hiatus in
distribution.

Sphyrna mokarran is present in the eastern Pacific from the Gulf
of California south to Panama and probably northwestern South
America. It occurs in the Pacific from southern Japan south to
Australia, although it appears to be absent from New Zealand and
the more eastern parts of Oceania. S. mokarran is found through-
out the East Indies and westward, in the northern Indian Ocean, to
east Africa.

Sphyrna (Sphyrna) zygaena (Linnaeus)
Ficurss 8, 9, 21c, 22c; PuaTE 6c

Squalus zygaena Linnaeus, 1758, p. 234. (By reference to illustration by
Willughby, 1686; no type specimen; type locality, “‘Europe,’’ ‘‘America.’’)

Squalus zygena Bonnaterre, 1788, p. 9. (Atlantic and Mediterranean.)

Sphyrna zygaena Rafinesque, 1810, p. 46. (Identification by included references;
Sicily.)

?Cestrorhinus caroliniensis Blainville, 1816, p. 121. (Name only.)

?Cestrorhinus pictus Blainville, 1816, p. 121. (Name only.)

Zygaena zygaena Cuvier, 1817, p. 127. (General; identification by references.)

Zygaena malleus Valenciennes, 1822, p. 223. (‘‘French”’ coasts; Brazil.)

Zygaena vulgaris Cloquet, 1830, p. 621. (At least in part; general.)

Zygaena subarcuata Storer, 1848, p. 70. (Identified by description of 2 ft. speci-
men; type presumably in MCZ [not located]; type locality, Provincetown,
Cape Cod.)

Sphyrnias zygaena Gray, 1851, p. 48. (In part; identification by references, but
also includes lewini [misspelled ‘‘lewis72’’].)

Sphyrna malleus van der Hoeven, 1855, p. 262. (General; Mediterranean; identi-
fication by reference to Linnaeus.)

Cestracion zygaena Gill, 1861, p. 403. (Name only; identification by reference
to Linnaeus, 1758.)

Cestracion subarcuata Gill, 1862, p. 59. (Name only; evidently based on Zygaena
subarcuata Storer, 1848.)

?Cestracyon zygaena Poey, 1881, p. 348. (Refers primarily to Sphyrna lewini;
specimens from Mediterranean may apply in part to S. zygaena.)

?Zygoena malleus Sauvage, 1891, p.510. (May refer entirely or in part to Sphyrna
lewint.)

?Sphyrna (Zygaena) zygaena Imms, 1905, p. 43. (Pharyngeal denticles; name
only; may refer entirely or in part to S. lewini.)

?Sphyrna (Sphyrna) zygaena McCulloch and Whitley, 1925, p. 129. (Reference;
may refer in part to S. lewini.)

Sphyrna (Cestracion) zygaena von Bonde, 1933, p. 377. (South Africa; skull and
nerves; identification by photo of head.)

32 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Zygaena zigaena Nobre, 1935, p. 425. (Portugal; identification by description
of head.)

SPECIMENS EXAMINED.—(Range in size: 457-2,663 mm. TL). At-
lantic Ocean (including Mediterranean Sea): Azores: USNM 94502
(1), Terceira. Brazil: MCZ 451 (1), MCZ 35364-35365 (2), Rio de
Janeiro. Italy: USNM 28452 (1), Livorno; ANSP 604 (1), (no
further data). United States (Florida): UMML 5639 (head), off
Alligator Reef, Monroe Co.; UMML 10444 (1), off North Miami,
Dade Co. (in Gulf Stream). (Massachusetts): USNM 10418 (1),
USNM 83980 (1), Woods Hole; MCZ 36425 (1), USNM 33268 (1),
Buzzards Bay, Woods Hole; USNM 31527 (1), Stuart’s Pond, Buz-
zards Bay; MCZ 1159 (1), Nahant; MCZ 1416 (1), Cape Cod. (Mary-

Figure 8.—Sphyrna zygaena: a, juvenile female, 687 mm. TL, from Nahant, Mass. (MCZ
1159); b, upper and lower teeth, left side, about 1.7 3c, second upper tooth; d, ninth
upper tooth; ¢, third lower tooth; f, ninth lower tooth. (c-f, about 3.8 X.) (From
Bigelow and Schroeder, 1948, p. 437.)

land): USNM 76681-76683 (3), Ocean City. (New Jersey): USNM
93845 (3), USNM 120495 (11), Barnegat Bay, Seaside Park; ANSP
522 (1), Holly Beach. (New York): CAS 19189 (1), Graves End
Bay; USNM 116447 (5), Sandy Hook; UMMZ 115903 (1), Long
Island; MCZ 136 (2), no further data. (Virginia): USNM 119699
(1), off Virginia Beach; USNM 6090 (1), Norfolk.

Pacific Ocean: Australia: USNM 29020 (head), USNM 39992 (1),
Sydney (Port Jackson). Ecuador: USNM 127758 (head), Gulf of
Guayaquil, 81°13’ W., 4°13’ S.; MCZ 383 (1), Galapagos Islands.
Japan: USNM 23383 (stuffed skin), USNM 71779 (1), Tokyo market;

NO. 3539 HAMMERHEAD SHARKS—GILBERT 33

USNM 51289 (head), Wakanoura; USNM 51291 (head), Nagasaki;
FMNH 57423 (1), Osaka market; MCZ 1042 (1), Yenoshima; SU
24125 (1), Misaki; UMMZ 179078 (1), vicinity of Choshi; UMMZ
179079 (1), Hakata Bay, northern Kyushu; UMMZ 179080 (1),
Suruga Bay. Mexico: USNM 190592 (10), Gulf of California,
southern end of Angel de la Guarda Island; UMMZ 61032 (1), Cedros
Island. New Zealand: UMML uncataloged (2), Bay of Islands,
North Island. Panama: MCZ 515 (1), no further data. Peru:
USNM 77711 (head), I. Lobos de Tierra; MCZ 421 (1), MCZ 441
(3), Magdalena Bay; MCZ 382 (2), Payta; SIO 58-29 (1), ‘‘off Perw’’;
SU 13003 (1), no further data. United States (California): SIO
58-224, SIO 58-382 (jaws), off La Jolla, San Diego County; LACM
uncataloged (jaws and chondrocranium), off San Diego.

Draenosis.—Characters mentioned in the diagnosis of the sub-
genus Sphyrna are not repeated here.

Sphyrna zygaena is characterized (as is S. lewint, its closest relative)
by a long second dorsal fin lobe and a low second dorsal fin, the length
of the former about twice greatest height of fin. The anterior-
median pore patch on the underside of the head is unique, the patch
characterized by a broad, smoothly rounded angle at the outer-
posterior corner and a broad, pointed posterior extension in the
median-posterior area (fig. 22c).

S. zygaena is distinguished from S. lewini by lack of a median
indentation (scallop) on anterior margin of head; a longer inner
narial groove, which extends at least 50 percent of distance from

_ inner margin of nares to tip of snout; orbit closer to nares, being
separated by a distance equal to about half of greatest horizontal
_ diameter of eye; tip of second dorsal lobe not reaching as near to

upper precaudal pit, the lobe extending, at most, about two-thirds
of distance from second dorsal base to pit; a slightly more deeply
falcate anal fin, the deepest part of the notch on the distal margin
situated slightly posterior to a line drawn perpendicularly from the
posterior axil of the fin; a shorter anal base, the length of which is

slightly less than length of pectoral base; a more acutely pointed

_ posterior ridge to the upper precaudal pit (fig. 21c); absence of a

lower precaudal pit; absence of a rostral fenestra: and several other
? )

_ chondrocranial characters (pl. 6c).

A comparison of Sphyrna zygaena with other members of the genus

Sphyrna is presented in table 1.

Description.—Meristic data appear in table 4. Characters men-
tioned in the diagnoses of the subgenus Sphyrna and of S. zygaena
are not repeated here.

Head moderately expanded, the greatest width from 26.1 to 29.0
percent of TL (in specimens less than 1,000 mm. long); shallow

219-937—67——3

34 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 119

Ficure 9.—Sphyrna zygaena: a, head of specimen illustrated in figure 8 (MCZ 1159), |
ventral view, about 0.45 X natural size; b, second dorsal and anal fins; ¢, dermal denticles, |
about 45 X; d, fourth upper tooth; ¢, fifth lower tooth. (d-e, about 9X.) (From —
Bigelow and Schroeder, 1948, p. 438.) |

NO. 3539 HAMMERHEAD SHARKS—GILBERT 35

scallops present on anterior margin of head, about halfway between
nares and tip of snout; outer posterior corner of lateral expansion of
head posterior to corner of mouth, probably in specimens of all sizes;
narial flap broad, moderately pointed at tip, with the outer edge
curving evenly inward; orbit large, the horizontal diameter about
equal to length of longest (second) gill slit; length of snout from one-
fifth to one-fourth of head width; greatest transverse distance be-
tween corners of mouth from one-fifth to one-fourth of greatest width
of head; symphysis of jaws situated posterior to a transverse line
drawn between posterior margins of orbits in specimens of all sizes;
labial furrow present at corners of lower jaw only (concealed when
mouth is closed), about one-fifth as long as greatest horizontal
diameter of eye.

Origin of first dorsal fin above posterior axil of pectoral fin, a per-
pendicular line drawn ventrally from origin intersecting axil; first
dorsal fin broad, rather erect (not as erect as in large examples of
Sphyrna tudes), a line drawn perpendicularly from apex of fin passing
close behind base; base of first dorsal fin from two-fifths to nine-
twentieths of length of head, and from three-fourths again to nearly
twice length of pectoral fin base; anterior margin of first dorsal fin
evenly curved, with the part nearest the apex very strongly curved;
distal margin of first dorsal fin strongly falcate; lobe of first dorsal fin
about one-third length of base; origin of second dorsal fin situated
about halfway back above base of anal fin; second dorsal fin base from
three-fifths to four-fifths length of anal fin base; anterior margin of
pectoral fin from one-half to two-thirds length of head (higher values
usually apply to larger individuals) and about equal to or slightly
greater than greatest vertical height of first dorsal fin; pectoral fin
about two-thirds as broad as long; anterior margin of pectoral moder-
ately convex; distal margin of pectoral slightly falcate; apex and inner
corner of pectoral rounded; length of pelvic fin about four-fifths length
of anal fin; length of pelvic fin base approximately equal to length of
anal fin base; anterior margin of pelvic fin weakly convex, the distal
margin nearly straight; length of anal fin equal to or slightly greater
than length of pelvic fin; height of anal fin about a third again as much
as height of second dorsal; length of caudal fin nearly a third of TL;
upper margin of caudal fin nearly straight; terminal part of caudal
about one-fifth of total length of fin; posterior margin of terminal part
of caudal fin moderately falcate; terminal apex of caudal narrowly
rounded ; lower lobe of caudal relatively narrow and long, the distance
from angle included by upper and lower caudal lobes to tip of latter
going about twice in anterior margin of lower caudal lobe; length of
lower caudal lobe one-third to two-fifths as long as upper lobe; lower
caudal fairly erect, sloping posteriorly at about a 35° angle from a

36 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

perpendicular to the body axis; anterior margin of lower caudal lobe
weakly convex; posterior margin of lower caudal lobe nearly straight.

Anterior margin of rostral node of chondrocranium nearly straight,
or with a broad, very shallow median indentation; accessory rostral
cartilages absent; anterior fontanelle (viewed ventrally) rather deep
and evenly rounded, with no noticeable median notch; short, rounded
wings present on sides of rostral node; anterior wing of olfactory
cartilage absent, represented only by a small “bump’’; anterior part
of preorbital process long and slender, and without a sharp angle
on anterior edge; posterior part of preorbital process terminating in an
evenly rounded knob.

Total number of vertebrae 196 (in one specimen examined); body
vertebrae 101, there being six more body than caudal vertebrae.

Teeth panne Gee Oteraeto aes ae an a ao; teeth weakly serrate in large in-
dividuals; first upper tooth nearly symmetrical and erect; subsequent
upper teeth strongly oblique, their inner margins straight or evenly
convex, their outer margins deeply incised and with the outermost
teeth with very small, though well-developed cusps; lower teeth
similar to upper, though smaller; first four teeth in lower jaw rather
erect and with narrow cusps; one small symmetrical tooth at sym-
physis of lower jaw, none or one in upper jaw; one to two series of
teeth functional along sides of upper jaw, and two to three series
functional toward center; two to three series functional in lower jaw.

Dermal denticles (examined from upper part of back beneath first
dorsal fin) closely overlapping, the blades thin and moderately arched ;
length of denticle (measured to tip of median marginal ‘‘tooth’’) as
ereat, or nearly as great, as width; small specimens usually with
three, large specimens with up to seven, sharp-toothed ridges, and
from three to five pointed marginal teeth present on denticle; median
tooth on denticle slightly longer than more distal teeth; pedicel very
short.

Individuals deep olive to brownish gray dorsally, shading to white
ventrally; tip of ventral surface of pectoral fin sometimes faintly gray
black in life.

Maximum total length probably between 12 and 13 feet.

Rance.—Sphyrna zygaena has an antitropical distribution, occur- |
ring in cooler waters of both the Northern and Southern Hemispheres |

but apparently absent from the intervening warmer areas (map 2).
It is present in the western Atlantic from Nova Scotia and northern
New England south to (occasionally) southern Florida (UMML 5639
and UMML 10444), and from Argentina and Uruguay to (probably)
northern Brazil (exact northern limits of range in South American

waters uncertain). Stewart Springer (pers. comm.) states that he |

NO. 3539 HAMMERHEAD SHARKS—GILBERT on

knows of no authentic record for this species from the Gulf of Mexico
or Caribbean, all literature records applying to Sphyrna lewini, which
is common there.

In the eastern Atlantic Sphyrna zygaena is known from Scotland
south to Senegal, in northwest Africa (Cadenat, 1951, p. 46), and also
from the Mediterranean Sea. Records of S. zygaena farther south
along the coast of Africa (Irvine, 1947, pp. 89,90) and from South
Africa (Smith 1953, pp. 46, 47) may not have been based on this
species, however, since in neither case were adequate descriptions
given, and the hammerheads pictured represent species other than
zygaena. Considering that the cool waters off South Africa are suit-
able for S. zygaena, however, it is likely that the species does occur
there.

/
i=

ESI fu

!

Map 2.—Distribution of Sphyrna zygaena.
(Solid= specimens examined; hollow=confirmed literature references.)

In the eastern Pacific Sphyrna zygaena is known from western South
America (Chile, Peru, and the Galapagos Islands) and (one record)
Panama, and from southern California and Mexico (south to the
Gulf of California), but intensive collecting has failed to reveal its
presence between the above two areas. In the northwestern Pacific
S. zygaena is common around southern Japan, although the exact
northern limits of its range there are uncertain. It seems to be absent
from the Hawaiian Islands and the East Indian area, but is present
off New Zealand and southern Australia. It probably is also present
in the southern parts of the Indian Ocean as well.

Sphyrna (Sphyrna) lewini (Griffith and Smith)
Fiaures 10, 21d, 22d; PLatss 3, 6B, 9B, 10B

?Cestrorhinus caroliniensis Blainville, 1816, p. 121. (Name only.)

?Cestrorhinus pictus Blainville, 1816, p. 121. (Name only.)

Zygaena malleus Valenciennes, 1822, p. 223. (In part; description; illustration of
head; Mediterranean Sea and Atlantic Ocean; teeth described as denticulate.)

*Zygaena indica Van Hasselt, 1823, p. 315. (Type locality: Java.)

Zygaena lewini Griffith and Smith, in Cuvier, Griffith, and Smith, Animal King-
dom, vol. 10, p. 640, pl. 50, 1834. (Original description; figure; no type
specimen; type locality, south coast of New Holland.)

Sphyrnias zygaena (misidentification in part) Gray, 1851, p. 48. (In part;
malleus included in synonymy.)

38 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Sphyrna lewint Duméril, 1858, p. 261. (Possibly in part; Gorrée, West Africa.)

Cestracion leewwenii Day, 1865, p. 271. (Description; Malabar; distinguished
from C. zygaena.)

Cestracion (Zygaena) leewwenii Duméril, 1865, p. 383. (“Cotes de la terre de
Leeuwen” [Australia].)

2Cestracion (Sphyrna) zygaena (misidentification) Steindachner, 1870, p. 576.
(Possibly in part; name only; Senegambia.)

Zygaena erythraea (Ehrenberg) Klunzinger, 1871, p. 666. (Name in synonymy.)

Zygaena leuwinii Ramsay, 1881, p. 96. (Possibly in part; name only; Port
Jackson, Australia.)

Cestracyon zygaena (misidentification) Poey, 1881, p. 348. (Porto Rico, Guiana,
Mediterranean area, East Indies; refers primarily to Sphyrna lewint; speci-
mens from Mediterranean may apply in part to S. zygaena.)

?Sphyrna zygaena (possible misidentification) Studer, 1889, p. 263. (Moreton
Bay.)

?Zygoena malleus (possible misidentification) Sauvage, 1891, p. 510. (May refer
entirely or in part to Sphyrna zygaena.)

2Sphyrna (Zygaena) zygaena (possible misidentification) Imms, 1905, p. 43.
(Pharyngeal denticles; name only; may refer entirely or in part to Sphyrna
zygaena.)

?Sphyrna tudes (possible misidentification) Ogilby, 1908, p. 4. (May refer to
Sphyrna mokarran.)

Cestracion oceanica Garman, 1913, pp. 158-159. (Original description; type
locality, Society Islands.)

Cestracion lewini Ogilby, 1916, pp. 81, 94. (Moreton Bay, Queensland.)

Sphyrna (Sphyrna) lewini McCulloch and Whitley, 1925, p. 129. (Reference.)

2Sphyrna (Sphyrna) zygaena (possible misidentification) McCulloch and Whitley,
1925, p. 129. (May refer in part to Sphyrna lewini.)

Sphyrna oceanica Fowler, 1928, p. 23. (Copied.)

Cestracion zygoena (probable misidentification) Chevey, 1932, p. 6. (Cochin
China; misidentification probable because of locality.)

Sphyrna diplana Springer, 1941, pp. 46-52. (Original description; type locality,
Englewood, Fla.)

SPECIMENS EXAMINED.—(Range in size: 386-1,755 mm. TL). At-
lantic Ocean: ‘Atlantic’: MCZ 393 (1), no further data. ‘‘Europe”’:
MCZ 673 (1), no further data. ‘West Indies’: SU 11830 (1), no
further data. Bahamas: UMML 10867 (1), off west end Grand
Bahamas Island. Brazil: MCZ 462 (4), Rio de Janeiro; MCZ 314
(1), MCZ 470 (1), SU 52717-52719 (3), Rio Grande do Sul; SU 52874
(1), Florianopolis; SU 14046 (1), Santos. Cuba: USNM 111350
(chondrocranium), no further data. Italy: UP uncataloged (1),
Livorno. Guinea: USNM uncataloged (1, head and fins), 9° 34’ N.,
16° 18’ W. Jamaica: USNM 29999 (1), no further data. Liberia:
USNM 179705-179706 (2), Messurado River beach. Mexico: MCZ
39646 (1), off Ciudad Carmen, Campeche; SU 11696 (1), no further
data. Panama: USNM 79282 (1), USNM 79285 (1), ANSP 49936-
49937 (2 heads), Colon. Surinam: USNM 159196 (1), 06°23’ to
06°20’ N., 54°47’ to 54°51’ W.; USNM 159197 (1), USNM 159199 (1),

ae

NO. 3539 HAMMERHEAD SHARKS—GILBERT 39

06°27’ to 06°25’ N., 55°05’ to 55°10’ W. ‘Trinidad: FMNH 3165 (2),
no further data; SU 19193 (1), 1°29’ N., 10°39’ W. (This locality, as
listed, is not close to Trinidad.) United States (Florida): USNM
106544 (1), UMMZ 155050 (1), Englewood; USNM 108451 (holotype
of Sphyrna diplana), Englewood; USNM 133580 (1), Merritt Island,
Brevard Co.: UF 455 (1), Matanzas Inlet, St. Johns Co.; UF 4208 (1),
Fernandina, Nassau Co. (Georgia): UF 5463 (1), mouth of St. Simons
Sound, Glynn Co. (South Carolina): USNM 25180 (1), Charleston;
SNM, 32°42’ N., 79°02’ W.. (Texas): FMNH 37798 (1), Port
Aransas; USNM 158561 (1), Freeport; MCZ 35223 (1) MCZ 35826-—
35828 (3), USNM 116449 (3), USNM 118660 (1), Galveston; SU
18390 (1), no further data.

Pacific Ocean (including Indian Ocean and Red Sea): Aden:
BMNH 1925.7.20.32.37 (6), Gulf of Aden. Burma: SU 14496 (1),
South Moscos Island group. Celebes: MCZ 479 (1), no further data;
USNM 170557 (1), Makassar market. China: SU 14150 (1), Tinghai,
Chu Shan Island; USNM 130604 (1), Foochow. Costa Rica: SIO
52-94 (1), Puntarenas. Eritrea: USNM 49330 (1), Red Sea, at
Massaua. Hawaii: MCZ 471 (1), USNM 151539 (1), Honolulu;
USNM 52642 (1), USNM 52644 (1), Honolulu, 28°31’00’’ N.,
141°47’00” W.; SU 12765 (1), SU 14088 (1), MCZ 79 (1), MOZ 456
(1), no further data. India: SU 14398 (1), Calicut. Java: USNM
72476 (1), Batavia. Japan: UMMZ 177119 (1), Southern Chosen
(South Korea?). Mexico: SU 11696 (1), no further data; USNM
120376 (head), Gulf of California, off San Marcos Island. New Cale-
donia: USNM 111201 (head), USNM 111243-111244 (2 dried skins),
no further data. Panama: ANSP 565 (1), MCZ 517 (2), ‘‘Pacific
side,” no further data; USNM 29289 (1), USNM 111414 (stuffed skin),
no further data; ANSP 82186 (2), San Miguel Bay, Garachine Pt.;
MCZ 1063 (1), MCZ 1059-1061 (3), Panama Bay, San Miguel Island;
MCZ 1088-1092 (5), Panama Bay, near Panama; UCLA 53-275 (3),
Panama Bay, between Panama City and Punta Gorda. Philippines:
USNM 170554 (1), loilo market; USNM 170555-170556 (2), Manila
market; USNM 56337 (1), Cavite. Society Islands: MCZ 460 (3),
USNM 153587 (1) (syntypes of Cestracion oceanica), no further data.

Nomenciaturs.—Sphyrna lewini was originally described (as
Zygaena lewini Griffith and Smith, 1834, p. 640, pl. 50) from the
south coast of Australia, or ‘“New Holland,” as it was then called.
Although the description lists none of the important diagnostic
features, the accompanying picture (pl. 3) is identifiable, and can be
recognized by the distinct median indentation on the anterior part of
the head and the low second dorsal fin with a long lobe.

In the past Griffith has been cited as the original describer of
Sphyrna lewini, a circumstance resulting from the inclusion of his

40 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

name on the title page as author of the supplementary sections in the
English (1834) edition of Cuvier’s ‘The Animal Kingdom.” Ap-
parently overlooked, however, is the fact that there are actually two
consecutive title pages to each volume, and, on the second page of
volume 10, under the heading ‘“The Class Pisces,’ the names of
Edward Griffith and Charles Hamilton Smith appear, thus indicating
their responsibility for the new sections on the orders of fishes. The
past confusion has resulted because only Griffith’s name appears on
the first title page (entitled ‘“The Animal Kingdom’’) as author of
the additions, although the words ‘‘and others” are written in small
type underneath. It is clear from the above that Cuvier had no
part in the preparation of these supplements, thereby eliminating
him as a codescriber of the species named therein; and therefore,
Griffith and Smith should be regarded as describers of Sphyrna lewini
and other new forms appearing in this volume.

For a long time Sphyrna lewini was confused with the superficially
similar S. zygaena. Garman seems to have been the first to recognize
the distinctness of the two species, since, in his monograph of the
sharks (1913, pp. 158-159), he described a new hammerhead, Cestracion
oceanica, which he noted was “‘closely allied to C. zygaena, [and]
similar in most respects.’’ In the comparison of this species with
zygaena, reference was made to the long inner narial groove of the
latter (extending more than halfway from the nares to the middle
of the snout). The length of the inner narial groove in oceanica,
together with other characters mentioned in the original description,
would lead one to suspect that this species is identical with Sphyrna
lewini, a suspicion that is confirmed by examination of the four
syntypes (MCZ 460 [38 spec.]; USNM 153587 [1 spec.]). Although
Garman distinguished the young of the above species, there is no
evidence that he recognized the adults, a situation that is difficult |
to understand in view of the fact that the difference in length of the
inner narial groove, which he recognized in the young, remains
constant at all sizes. |

Springer (1941, pp. 46-52) was the first to demonstrate conclusively |
that S. zygaena and S. lewini are specifically distinct. At the time,
however, he was unaware that the Atlantic and Pacific populations of |
lewint are morphologically indistinguishable, and he consequently |
described the former population as a new species, S. diplana.

Tortonese (1950a, pp. 21-28), although continuing to recognize both
S. diplana and S. lewini, strongly questioned their taxonomic separa-
tion. He did not synonymize the two species, however, due to a lack |
of material from critical areas. In addition, Tortonese doubted the |
validity of Sphyrna oceanica (1950a, p. 28), which, as he correctly |
suspected, is identical with lewini.

NO. 3539 HAMMERHEAD SHARKS—GILBERT Al

Fraser-Brunner (1950, pp. 213-219) synonymized S. diplana with
S. lewini, although he continued to recognize S. oceanica as valid,
citing supposed differences in size of eye, length of snout, relative
position of first dorsal fin, size of second dorsal fin, and shape of head.
Examination of specimens from the Gulf of Aden (BMNH 1925.7.20.
32.37), which were identified by Fraser-Brunner as S. oceanica, fails
to substantiate his conclusion that this nominal species is a valid form.

Dracnosis.—Characters mentioned in the diagnosis of the subgenus
Sphyrna are not repeated here.

CS

Ficure 10.—Sphyrna lewini: a, juvenile female, 639 mm. TL, from Rio de Janeiro, Brazil
(MCZ 462); b, ventral view of head; c, upper and lower teeth, left side, about 0.55 &3
d, fourth upper tooth; e, twelfth upper tooth; f, third lower tooth; g, eleventh lower tooth;
h, dermal denticles, about 42 X. (d-g, about natural size.) (From Bigelow and Schroeder,
1948, p. 415.)

Sphyrna lewini is characterized (as is S. zygaena, its closest relative)
by a long second dorsal fin lobe and a low second dorsal fin, the length
of the former about twice greatest height of fin. The anterior-median
pore patch on the underside of the head is unique, the patch character-
ized by a rather sharp, smoothly rounded angle at the outer-posterior
corner and by a narrow triangular area, which is devoid of pores, in
the median-posterior region (fig. 22d).

S. lewini is distinguished from S. zygaena by a median indentation
(scallop) on anterior margin of head; a shorter inner narial groove,
which extends 40 to 45 percent of distance from inner margin of nares
to tip of snout; orbit farther from nares, being separated by a distance
slightly less than greatest horizontal diameter of eye; tip of second
dorsal lobe reaching closer to upper precaudal pit, the lobe extending
from three-fourths to four-fifths of distance from second dorsal base

42 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

to pit; a slightly less deeply falcate anal fin, the deepest part of the
notch on the distal margin situated slightly anterior to a line drawn
perpendicularly from the posterior axil of the fin; a longer anal base,
the length of which is about equal to length of pectoral base; a more
obtusely pointed posterior ridge to the upper precaudal pit (fig. 21d);
occasional presence of a lower precaudal pit; presence of a rostral
fenestra; and several chondrocranial characters (pl. 68).

A comparison of Sphyrna lewini with other members of the genus is
presented in table 1.

Descriprion.—Meristic data appear in table 5. Characters men-
tioned in the diagnoses of the subgenus Sphyrna and of S. lewini are
not repeated here.

Head moderately expanded, the greatest width from 24.0 to 30.2
percent of TL (in specimens less than 1,000 mm. long); shallow
scallops present on anterior margin of head, about halfway between
nares and tip of snout; outer posterior corner of lateral expansion of
head posterior to corner of mouth, probably in specimens of all sizes;
narial flap broad, moderately pointed at tip, with the outer edge
curving evenly inward; orbit large, the horizontal diameter almost
equal to length of shortest (fifth) gill slit; length of snout about
one-third of head width in smaller specimens, to about one-fifth of
head width in larger specimens; greatest transverse distance between
corners of mouth from one-fifth to one-fourth of greatest width of
head; symphysis of jaws situated posterior to a transverse line drawn
between posterior margins of orbits in specimens of all sizes; labial
furrow present at corner of lower jaw only (concealed when mouth is
closed), about one-fifth to one-fourth as long as greatest horizontal
diameter of eye.

Origin of first dorsal fin above or slightly posterior to axil of pectoral
fin, a perpendicular line drawn ventrally from origin intersecting
anterior third of pectoral fin; first dorsal fin broad, rather erect (not
as erect as in large specimens of Sphyrna tudes), a line drawn perpen-
dicularly from apex of fin passing close behind base; base of first dorsal
fin about one-half length of head and from half again to almost twice
length of pectoral fin base; anterior margin of first dorsal fin evenly
curved, with part nearest apex very strongly curved; distal margin of
first dorsal fin strongly falcate; lobe of first dorsal fin about one-third
length of base; origin of second dorsal fin situated about halfway back
above base of anal fin; second dorsal fin base from three-fifths to four-
fifths length of anal fin base; anterior margin of pectoral fin from
one-half to two-thirds length of head (higher values usually apply to
larger individuals) and about equal to greatest vertical height of first

dorsal fin; pectoral fin about two-thirds as broad as long; anterior —

margin of pectoral fin moderately convex; distal margin of pectoral |

NO. 3539 HAMMERHEAD SHARKS—GILBERT 43

straight or slightly faleate; apex and inner corner of pectoral fin
rounded; pelvic fin equal in length or slightly less (about four-fifths as
long) than anal fin; pelvic fin base approximately equal in length to
anal fin base; anterior margin of pelvic fin weakly convex, the distal
margin nearly straight; anal fin equal in length or slightly greater than
pelvic fin; height of anal fin about a third again as much as height of
second dorsal fin; caudal fin nearly a third of TL; upper margin of
caudal fin nearly straight; terminal part of caudal about one-fifth of
total length of fin; posterior margin of terminal part of caudal fin
moderately faleate; terminal apex of caudal narrowly rounded; lower
lobe of caudal relatively narrow and long, the distance from angle
included by upper and lower caudal lobes to tip of latter going from
1% to 2 times in anterior margin of lower caudal lobe; lower caudal
lobe one-third to two-fifths as long as upper lobe; lower caudal lobe
fairly erect, sloping posteriorly at about a 35° angle from a perpen-
dicular to body axis; anterior margin of lower caudal lobe weakly
convex; posterior margin of lower lobe nearly straight.

Anterior margin of rostral node of chondrocranium nearly straight,
or with a very shallow median indentation; accessory rostral cartilages
absent; anterior fontanelle (viewed ventrally) appearing broad and
shallowly U-shaped, with no noticeable median notch; prominent,
bluntly rounded wings present on sides of rostral node; anterior wing
of olfactory cartilage prominent, pointed, and projecting medially;
anterior part of preorbital process long and slender, with a sharp angle
on anterior edge; posterior part of preorbital process terminating in an
evenly rounded knob.

Total number of vertebrae 174 to 204 (in nine specimens examined;
only one individual with less than 192); body vertebrae 86 to 100, the
relative number of body to caudal vertebrae varying from six more to
twelve less.

Teeth 15 or 16-0 to 2-15 or ay

Lover 16-1 or2—15 orel6
large individuals; first tooth smaller than second, the 15th and 16th
very small; teeth 1 through 3 in upper jaw nearly symmetrical and
erect, with the subsequent teeth becoming increasingly oblique
toward corners of mouth; cusps present on all teeth; lower teeth with
somewhat narrower cusps than uppers, similarly oblique, but with
successive series tending to become more erect and their cusps relative-

teeth sometimes weakly serrate in

ly more narrow with growth.

Dermal denticles (examined from upper part of back beneath first

dorsal fin) closely overlapping, the blades thin and moderately arched;
_ length of denticle (measured to tip of median marginal ‘‘tooth’’) as

great, or nearly as great, as width; small specimens usually with three,

44 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

large specimens with four or five, sharp-toothed ridges, and as many
pointed marginal teeth present on denticle; median tooth on denticle
slightly longer than more distal teeth; pedicel very short, moderately
slender.

Individuals deep olive to brownish-gray dorsally, shading to white
ventrally; tip of ventral surface of pectoral fin faintly gray-black to
dark black in life.

Maximum total length probably between 12 and 13 feet.

VARIATION.—With the possible exception of the teeth, Sphyrna
lewini shows little noticeable geographic variation in external mor-
phological characters. This can be seen from table 5, in which mor-
phometric data have been recorded from specimens from throughout
the range. Those characters that are considered to be of basic tax-
onomic importance, such as the structure of the chondrocranium (pl.
6B), the form of the anterior-median pore patch on the underside of
the head (fig. 22d), the length and development of the intranarial
groove, the position and length of the gill slits, and the shape and
position of the fins, remain quite constant.

Although the teeth in Sphyrna lewini are ordinarily smooth, in
some cases they may be very weakly serrate. This apparently occurs
only in larger specimens, at least 5 or 6 feet in length. The serrated
condition has long been thought to be characteristic only of the
Pacific population (Sadowsky, 1965, p. 2), and, in fact, was one of the
reasons for Springer’s naming the Atlantic population as a new species,
S. diplana. However, Dr. Shelton P. Applegate has recently informed

me (in litt.) that he has seen specimens of this species from the western |
Atlantic with serrated teeth. In view of this, and because there seem |
to be no concomitant characters to suggest that more than one species |

is involved, I choose to regard S. diplana as a synonym of S. lewint.
A series of nine vertebral counts is available for specimens from
widely scattered parts of the range. Although the total number of vertebrae
ranges from 174 to 204, variation in eight of the nine individuals is
only from 192 to 204. The reduced count in the other specimen
(USNM 25180—from Charleston, S.C.) is due both to a decreased

number of body and caudal vertebrae. It is difficult to say at this |

time what the significance of this unusually low count is.

Ranee—Sphyrna lewini is circumtropical in distribution (map 8). |
In the western Atlantic it occurs from southern Brazil (and possibly ©
Uruguay) north to New Jersey (Bigelow and Schroeder, 1948, p. 419; »
Backus, 1960, p. 244). In the eastern Atlantic its range is not so
well defined. It occurs in the Mediterranean Sea (Bigelow and Schroe- |

der, 1948, p. 419) and along the African coast south at least to Liberia |
and Ghana (Bigelow and Schroeder, 1948, p. 419). The northern |

NO. 3539 HAMMERHEAD SHARKS—GILBERT 45

limits of its range in the eastern Atlantic are uncertain, although it
has never been recorded from the British Isles.

Sphyrna lewint is found in the western Pacific from southern
Australia north to southern Japan and Korea, and in the eastern
Pacific from northwestern South America to the Gulf of California.
The fact that it is the only species of hammerhead definitely recorded
from the Hawaiian Islands (Gosline and Brock, 1960, p. 91) may
indicate that it is a more pelagic species than Sphyrna mokarran,
which also logically might be expected to occur there.

Map 3.— Distribution of Sphyrna lewint.
(The most westerly record from Africa is now known to represent S. couardi; solid=speci-
mens examined; hollow=confirmed literature references.)

Sphyrna lewint apparently does not extensively penetrate cool
waters (i.e., in which the temperature does not rise above the low
70’s F.). Although its range overlaps that of S. zygaena to some extent,
there are relatively few records of the two species having been taken
together (Herald, 1961, p. 31, fig.).

Sphyrna (Sphyrna) couardi Cadenat
PLATES 9a, 104A

Sphyrna couardi Cadenat, 1951, pp. 98-99. (Original description; compared with
S. diplana and S. bigelowi; tips of pectoral fins described as white; type
locality: ‘‘off West Africa’’.)

Specimens examined.—Guinea: USNM uncataloged (1 head, 425
mm. in width), 12°06’ N, 17°22’ W.

Discussion—Sphyrna couardi was described by Cadenat (1951,
pp. 98-99) from “‘off West Africa.”” The description was very brief
and there was no illustration, although the form was said to resemble
S. diplana (=S. lewini), differing from that species (and from all other
members of the family Sphyrnidae) in having the inferior part of the
pectoral fin white and a longer and smaller head. While I have not
seen a whole specimen of S. cowardi, I have examined a large head
(USNM uncatalogued) collected by Dr. Bruce B. Collette from off
West Africa. This head, which is 425 mm. wide, has been compared
with the head of a typical specimen of S. lewini, 445 mm. wide (USNM

46 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

uncataloged), from the same geographic area. Photographs of these
appear in plate 10 and radiographs in plate 9.

The most striking difference between the two heads is the deeper,
more robust appearance of S. couardi (also see measurements in table
10). This is confusing, since Cadenat described S. couardi as having a
‘longer and smaller’ head than S. lewini; however, this apparent
discrepancy may simply be a matter of semantics. The overall
appearance of the head, in fact, seems closer to S. mokarran than S.
lewini; however, couardi and mokarran may readily be distinguished
by the presence of a distinct intranarial groove and nonserrated teeth
in the former species, and by chondrocranial structure and differences
in pore patterns on the ventral surface of the head.

The mucal pore patches on the ventral surface of the head are
basically similar in S. cowardi and S. lewini, although the anterior-
median patch in the former is completely separated medially instead
of being divided about halfway as in the latter (fig. 22d). The
chondrocrania of the two forms also are similar (pls. 9 a, B), the main
difference being in the appearance of the preorbital process. In S.
couardi this structure is shorter and has a distinct knob at the angle,
whereas in S. lewini it is longer and lacks a distinct knob. In addition,
the olfactory wings in S. couvardi are blunt and do not come to a sharp
point as in S. lewini. In this case, however, the wing tips are rather
ragged in appearance, suggesting that this possibly may be an aberrant
condition.

In conclusion, the above evidence seems to indicate that Sphyrna
couardi is a valid species of restricted range and that it is most closely
related to Sphyrna lewint.

Subgenus Platysqualus Swainson

The subgenus Platysqualus, which is here considered to include
Sphyrna tudes, S. media, S. corona, and S. tiburo, is distinguished by
the following unique characters: First and fifth gill slits nearly equal
in length and both somewhat shorter than the three middle slits; anal
fin and anal fin base longer than in any other group of sphyrnids, the
former 11.0 to 13.5 percent, and the latter 7.5 to 9.0 percent, of TL;
anal base a fourth to half again as long as pectoral base; upper pre-
caudal pit broadly semicircular or slightly pointed in shape, the ends
pointing anteriorly (figs. 21le-c); lower precaudal pit present (this
feature also sometimes present in S. lewini); fifth gill slit above in-
sertion of pectoral fin (slightly posterior in S. tiburo); lobe of first
dorsal fin extending past insertion of pelvic fin (in all but S. tiburo);
teeth weak, nonserrated, with narrow pointed cusps; accessory rostral
cartilages often present (none observed in S. tiburo).

as

NO. 3539 HAMMERHEAD SHARKS—GILBERT AT

In addition, the subgenus is characterized by: A fairly high second
dorsal fin, its greatest height about equal to greatest height of anal fin;
a shallowly falcate anal fin; eyes situated anterior to mouth, a line
drawn between the posterior margins of orbits passing anterior to sym-
physis of jaws (intersecting symphysis in larger specimens of S. tiburo) ;
a small orbit, separated from nares by a distance greater than horizon-
tal diameter of orbit; absence of an outer narial groove; a variable total
number of vertebrae, ranging in number from 142 to 202, with those
over posterior part of coelomic cavity not enlarged; a small maximum
size, probably not exceeding 4 or 5 feet. Perhaps the maximum size
in some cases is even less (e.q., S. corona).

Sphyrna (Platysqualus) tiburo (Linnaeus)

DraGcnosis.—Characters mentioned in the diagnosis of the subgenus
Platysqualus are not repeated here.

Sphyrna tiburo is unique among members of the genus Sphyrna in
having the head only slightly expanded, either spade or shovel shaped,
and measuring from 14.4 to 24.5 percent of TL in specimens up to
1,000 mm. long (higher values usually apply to smaller individuals
and/or to individuals of the subspecies vespertina). ‘The anterior-
median pore patch on the underside of the head is also unigue, the
patch characterized by a rather sharp, but smoothly rounded angle at
the posterior corner and by a long, narrow median posterior extension
(fig. 22e).

Sphyrna tiburo is also distinguished by the anterior margin of the
head being more or less evenly rounded between the eyes, with neither
a median scallop nor narial depressions present; no inner narial groove;
origin of first dorsal fin distinctly posterior to axil of pectoral fin, a per-
pendicular line drawn ventrally from origin of dorsal intersecting pos-
terior part of pectoral fin; teeth not serrated, with short, weak, rather
blunt cusps.

A comparison of Sphyrna tiburo with other members of the genus is
presented in table 1.

DerscripTion.—Meristic data appear in table 6. Characters men-
tioned in the diagnoses of the subgenus Platysqualus and of S. tiburo
are not repeated here.

Intranarial distance from one-tenth to one-sixth of TL (lower
values usually apply to larger individuals); outer posterior corners
of lateral expansions of head broadly rounded and situated posterior
to corners of mouth; narial flap broad, blunt at tip, with the outer
edge curving gradually inward; length of snout about two-fifths
of head width; greatest transverse distance between corners of mouth
from one-third to a little more than two-fifths of greatest width
of head; symphysis of jaws situated, in smaller individuals, slightly

48 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

anterior to a transverse line drawn between posterior margins of
orbits, while in larger individuals a line drawn between these points
passes through the symphysis; a weakly marked labial furrow present
on lower jaw only, the furrow extending only a short distance (con-
cealed when mouth is closed).

Fourth gill slit situated anterior to, and fifth gill slit slightly pos-
terior to, insertion of pectoral fin; posterior lobe of first dorsal fin
terminating just anterior to insertion of pelvic fin (the dorsal and
pelvic fins being in closer proximity only in S. tudes, S. corona, and
S. media); first dorsal fin relatively slender, not perfectly erect, a
line drawn perpendicularly from apex of fin barely intersecting lobe
of fin; base of first dorsal fin about two-fifths length of head and at
least half again length of pectoral fin base; anterior margin of first
dorsal fin evenly curved, with the part nearest the apex more strongly
curved; distal margin of first dorsal fin moderately falcate; lobe of
first dorsal fin from four-ninths to almost one-half length of first
dorsal base; origin of second dorsal fin situated about a third of way
back above base of anal fin; second dorsal fin rather tall, its greatest
height about two-thirds length of base, and about equal to greatest
height of anal fin; second dorsal fin base about five-eighths of anal
fin base; lobe of second dorsal fin rather long, its length about a third
again greatest height of same fin, and extending about two-thirds
of distance from posterior part of second dorsal base to upper pre-
caudal pit; anterior margin of pectoral fin from one-half to nearly
two-thirds length of head (higher values usually apply to larger
individuals); pectoral fin four-fifths as broad as long; pectoral fin
base five-eighths to three-fourths length of anal fin base; anterior
margin of pectoral moderately convex; distal margin of pectoral
nearly straight; apex and inner corner of pectoral rounded; pelvic
fin about three-fourths length of anal fin; pelvic fin base about three-
fourths length of anal fin base; anterior margin of pelvic fin weakly
convex; distal margin of pelvic fin very shallowly falcate; anal fin
about one-fourth again as long as pelvic fin; anal fin base from one-
fourth to over one-third again as long as pectoral and pelvic fin
bases, and from three-fifths again to nearly twice as long as second
dorsal fin base; height of anal fin about equal to greatest height of
second dorsal fin; anal fin weakly falcate toward apex and nearly
straight toward tip of lobe; caudal fin from one-fourth to nearly
one-third of total body length; upper margin of caudal nearly straight;
terminal part of caudal about one-fourth of total length of fin; posterior
margin of terminal part of caudal fin nearly straight or slightly
faleate; both terminal and lower apices of caudal narrowly rounded
in smaller specimens, somewhat more pointed in larger individuals;
lower caudal lobe appearing relatively broad and short, the distance

NO. 3539 HAMMERHEAD SHARKS—GILBERT 49

from the angle included by the upper and lower caudal lobes to
tip of latter going from 2 to 2% times in anterior margin of lower
caudal lobe; lower lobe of caudal one-third to two-fifths as long
as upper lobe; lower caudal lobe sloping posteriorly at about a 45°
angle from a perpendicular to the body axis; anterior margin of
lower caudal lobe weakly convex; posterior margin of lower caudal
lobe nearly straight.

Anterior margin of rostral node of chondrocranium either straight
or with a poorly defined indentation; accessory rostral cartilages
absent; rostral fenestra absent; anterior fontanelle (viewed ventrally)
appearing broad and shallowly U-shaped, with a distinct, rounded,
median notch; prominent pointed wings present on sides of rostral
node; anterior wing of olfactory cartilage prominent, pointed, and
projecting medially; anterior part of preorbital process short and
rather blunt, with a moderately sharp angle on anterior edge; pos-
terior part of preorbital process terminating in a knob, the anterior
corner of which is evenly rounded and projects forward, and the
posterior corner of which is somewhat more angulate (pls. 7 A, 8B).

Total number of vertebrae 142 to 172 (seven specimens examined) ;
body vertebrae 71 to 88, the relative number of body to caudal
vertebrae varying from 9 more to 2 less.

12 to 14-0 or 1-12 to 14
Teeth ———————————_—_ : teeth : first tooth
ee TOR enIO eon eeth not serrate; first upper too

erect and symmetrical, but subsequent upper teeth becoming in-
creasingly oblique toward corners of mouth; teeth 3 through 10 or
11 largest in both jaws; cusps evident only on teeth 1 through 9 in
specimens examined (through 11 or 12 according to Bigelow and
Schroeder, 1948, p. 423); lower teeth shorter than upper teeth, with
narrower cusps and slightly broader bases, the first through third
teeth erect and symmetrical, the fourth through seventh or eighth
slightly oblique, the eighth and subsequent teeth without cusps or
definite cutting edges; three series of teeth functional in front and
one or two series functional along side of upper jaw; four to six series
functional in front, usually two functional along side, and three
functional toward corner of mouth of lower jaw.

Dermal denticles (examined from upper part of back beneath first
dorsal fin) closely overlapping, the blades rather steeply raised;
length of denticle (measured to tip of median marginal ‘‘tooth’’)
as great, or nearly as great, as width; five strong ridges usually pres-
ent, and as many long, sharp-pointed teeth (these teeth are some-
times blunted or broken) usually present at posterior margin of
denticle, the outer ridges and teeth sometimes rudimentary or absent,
particularly in smaller individuals; median tooth on denticle ordi-
narily slightly longer than more distal teeth; pedicel short and slender.

219-937—67——4

50 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Individuals gray or grayish brown dorsally, becoming progres-
sively more pale below, and often with small dark spots on side of
body; occasional spots on caudal fin.

VARIATION.—Specimens of Sphyrna tiburo from the Pacific usually
have more pointed heads than specimens from the Atlantic (figs. 12a, 6).
This difference is also reflected in proportional measurements, the
relative width of the head and transverse distance between the nares
averaging slightly greater for specimens from the Pacific (table 6 and
fig. 13). As a result, Springer (1940a, pp. 161, 162) described the
Pacific population as a new species, Sphyrna vespertina, although
Bigelow and Schroeder (1948, p. 428) later placed it in the synonymy
of S. tiburo.

For several reasons the two populations are regarded here as having
differentiated only to the subspecies level. First, no differences com-
parable to those found for head shape were discovered for other
morphometric characters (table 6). In addition, some individuals are
intermediate in head shape (fig. 12c). As mentioned previously, such
specimens possibly may have resulted from a partial reamalgamation
of the populations following initial differentiation of the two forms.

Vertebral counts were made on four specimens of S. tiburo tiburo
and three of S. tiburo vespertina. These counts, which range from 163
to 172 for the latter form and from 142 to 170 for the former, are
interesting both because of the wide range in variation found in the
typical subspecies and also because of the close agreement in vertebral
number between two of the individuals of S. tiburo tiburo and the three
specimens of S. tiburo vespertina. The two specimens of S. tiburo
teburo with the lowest counts (142 and 143) are from the eastern Gulf
of Mexico, while those with the higher counts (159 and 170) are from
Rhode Island and Brazil. (It should be mentioned here that these
differences are not attributable solely to variation in number of caudal
vertebrae.) Although it is possible that larger series of counts will
show this apparent difference to be clinal in nature, a study of vertebral
variation in sharks, recently completed by Drs. V. G. Springer and
J. A. F. Garrick (1964) indicates the existence of a number of closely
related species differing only in vertebral counts. This suggests that
those specimens of S. tiburo tiburo from the eastern Gulf of Mexico
represent a sibling form that may be distinguished from other pop-
ulations of the same subspecies only by number of vertebrae.

RancE.—Despite statements to the contrary (Bigelow and Schroe-
der, 1948, pp. 424-425; Herre, 1953, p. 27; Briggs, 1960, p. 173; and
1961, p. 552), there appear to be no substantiated records of Sphyrna
tiburo outside of the Western Hemisphere.

Sphyrna tiburo tiburo is common in shallow, inshore waters of the
western Atlantic, where it ranges from southern Brazil northward,

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NO. 3539 HAMMERHEAD SHARKS—GILBERT dl

regularly, to southern North Carolina and, occasionally, to southern
New England. All but three records are from continental waters,
and these exceptions are from Bimini, in the western Bahamas, and
from Cuba, less than 100 miles from the mainland. Although the
species is known to be well established on Bimini, there is no evidence
of a permanent population on Cuba, and it is possible that records
from the latter area have resulted from market specimens that were
caught on the other side of the Florida Straits.

Sphyrna tiburo vespertina is common in similar habitats in the
eastern Pacific from southern California to Equador.

Locality records for Sphyrna tiburo are plotted on map 4.

Sphyrna (Platysqualus) tiburo tiburo (Linnaeus)
Fiaures 11, 120, c, 18, 2le, 22e; Puate 7a

Squalus tiburo Linnaeus, 1758, p. 234. (Original description; no type specimen;
type locality, ‘‘America’’.)

Sphyrna tiburo Rafinesque, 1810, p. 47. (New generic name for Squalus tiburo
Linnaeus, 1758; misspelled “‘liburo,’”’ obviously a typographical error.)

Cestrorhinus tiburo Blainville, 1816, p. 121. (Name only.)

Zygaena tiburo Valenciennes, 1822, pp. 226-227. (Description; illustration of
head; Brazil.)

Zygaena (Squalus) tiburo Leuckart, 1836, p. 22. (Illustration; early embryo, yolk
stalk and yolk.)

Platysqualus tiburo Swainson, 1839, p. 318. (Description; new generic name
proposed without comment.)

Sphyrnias tiburo Gray, 1851, p. 50. (In list.)

Reniceps tiburo Gill, 1861, pp. 403, 412. (Name; new generic name proposed
despite availability of Platysqualus.)

Cestracion tiburo Gill 1862, p. 59. (In North American catalogue.)

Sphyrna (Reniceps) tuburo Jordan and Evermann, 1896, p. 217. (Range.)

Cestracion zygaena (misidentification) Radcliffe, 1916, pl. 43, fig. 2. (Photo of
jaws, evidently mislabeled by accident.)

SPECIMENS EXAMINED.—(Range in size: 350-910 mm. TL). Ba-
hamas: UMMZ 174384 (1), Bimini. Brazil: MCZ 464 (3), USNM
43359 (head and tail), Bahia; MCZ 469 (1), SU 52715 (1), SU 52873
(1), USNM 104318 (2), Recife (Pernambuco); SU 52713 (1), SU
52714 (1), Ponta de Mucuripe, Fortaleza; MCZ 27 (1), MCZ 90 (2),
MCZ 153 (5), MCZ 162 (5), MCZ 394 (4), MCZ 416 (4), MCZ 439 (2),
MCZ 450 (2), MCZ 461 (5), Rio de Janeiro; SU 14047 (1), FMNH
46997 (1), Santos; SU 52872 (1), Vitoria market. Colombia: USNM
94754 (1), Caribbean Sea, near Puerto Colombia (and Barranquilla).
Cuba: USNM 2710 (1), MCZ 522 (2), no further data. Mexico:
MCZ 39645 (1), off Ciudad Carmen, Campeche. Nicaragua: USNM
78180 (1), Punta Gorda. Panama: USNM 79315 (1), ANSP 49841-
49847 (7), ANSP 49938-49940 (3 heads), Colon. Surinam: USNM
159198 (1), 06°27’ to 06°25’ N., 55°05’ to 55°10’ W.; USNM 159201
(1), 06°20’ to 06°19’ N., 54°33.5’ to 54°38.5’ W. United States

52 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

(Alabama): USNM 119827 (1), near Cedar Pt., Mississippi Sound.
(Florida): USNM 26582 (1), UF 1407 (4), UF 5649 (1), UF 7061 (1),
UF 8948 (1), Cedar Key, Levy Co.; USNM 30687 (1), Pensacola;
USNM 34733-34734 (2), St. Marys River; USNM 39882 (1), off Cape
Sable; USNM 47334 (1), USNM 184870 (1), Tampa Bay; USNM
57242 (1), Alligator Light; USNM 62773 (1), USNM 118995 (1),
FMNH 7038 (1), Key West; USNM 92594 (1), USNM 116888 (1),
MCZ 755 (1), Tortugas; USNM 108455 (1), SU 14329 (1), Englewood;
USNM 125763 (1), near West Pass, Apalachicola Bay; CAS 19306 (1),
MCZ 35847 (1), Pine Island Sound; UF 1416 (1), Daytona Beach pier,

TAIRA ; =
pen Nhe gee eee pre age

Ficure 11.—Sphyrna tiburo: a, juvenile female, 367 mm. TL, from Rio de Janeiro, Brazil
(MCZ 90); b, head of same specimen, ventral view; c, upper and lower teeth, left side, of
male, 812 mm. TL, from Tortugas, Fla. (MCZ 848), about 1.5 ; d, fourth upper tooth;
e, tenth upper tooth; f, second lower tooth; g, seventh lower tooth; h, tenth lower tooth;
i, dermal denticles, about 14 * (MCZ 90); 7, lateral and apical views of dermal denticle,
about 26 X. (d-h, about 3 X.) (From Bigelow and Schroeder, 1948, p. 421.)

Volusia Co.; UF 4208 (1), Fernandina Beach, Nassau Co.; UF 4211
(1), St. Augustine, St. Johns Co.; UF 7061 (1), Spanish Harbor Key
(west end—Atlantic side), Monroe Co. (Georgia): UF 5463 (1), off
St. Simons Island, Glynn Co. (Louisiana): USNM 127114-127115 (2),
Barataria Bay channel; USNM 127116 (1), Bayou Fifi; USNM 155648
(1), Gulf of Mexico, south of New Orleans, 28°36’ N., 91°00’ W.
(Mississippi): USNM 147798 (1), no further data; FMNH 21590 (1),
Horn Island. (New Jersey): USNM 9270 (1), Delaware Bay,
Beesleys Pt., between Great Egg and Peck Bays. (North Carolina):
USNM 51894 (3), Beaufort. (Rhode Island): ANSP 581 (1), New-
port. (South Carolina): USNM 25171 (1), MCZ 78 (skin), Charles-

NO. 3539 HAMMERHEAD SHARKS—GILBERT 53

ton. (Texas): USNM 82710 (2), Chame Point Pass; USNM 116450
(5), MCZ 35824-35825 (2), MCZ 36157 (1), MOZ 36240-36245 (9),
Galveston; USNM 127101 (1), USNM 155964 (18), Corpus Christi;
USNM 127104 (1), Harbor Island; USNM 127106 (1), Parateria
(Grand Pass) ; USNM 127109 (1), ca. 13 mi. off Bay Chalaud; FMNH
37794-37796 (3), Port Aransas, Aransas; FMNH 11195-11198 (4),
Laguna Madre, Point Isabel. (Virginia): USNM 125760 (1), Lynn-
haven Roads, just west of Cape Henry light; MCZ 138 (skin),
Hampton Roads; FMNH 1622 (1), Ocean View.

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Ficure 12.—Sphyrna tiburo: a, head of S. t. vespertina, a specimen 636 mm. TL (UCLA
58-46), from Gulf of California, showing pointed head; b, head of S. #. tiburo, a specimen
637 mm. TL (USNM 57242), from southern Florida, showing rounded head; c, head of
S. t. tiburo, a specimen 880 mm. TL (USNM 127106), from Texas, showing intermediate
condition of head. (Drawings by Dorothea B. Schultz.)

Diaenosis.—Sphyrna tiburo tiburo differs from S. tiburo vespertina
in having a slightly narrower and more evenly rounded head. These
differences are illustrated in figures 12 and 13.

Sphyrna (Platysqualus) tiburo vespertina Springer
FicurE 12a; PLATE 7B

Sphyrna vespertina Springer, 1940a, pp. 161, 164-166. (Original description; com-
parisons; range; holotype, a 769 mm. female, SU 11584; type locality, Panama
City; paratypes, a 520 mm. female, SU 11881, same locality as holotype; and
a 938 mm. male, formerly Carnegie Museum no. 5675, from Guayaquil,
Keuador.)

SPECIMENS EXAMINED.—(Range in size: 277-880 mm. TL). Ec-
uador: USNM 88677 (1), FMNH 59350 (1), Guayaquil. El Sal-
vador: USNM 21627 (1), Acajutla. Mexico: USNM 190595 (1),
Camorore Beach, ca. 3 mi. north of Mazatlan, Sinaloa; UCLA 51-4
(1), Gulf of California, Bocochibampo Bay, Sonora; UCLA 52-250
(4), Almejas Bay, at anchorage off Santa Margarita Island, Baja
California; UCLA 58-46 (1), Gulf of California, south of Bahia Top-
olobampo, off Isla San Ignacio and Isla Macapule, Sinaloa; CAS (W
55-167) (1), Gulf of California, south end of Angel de la Guarda
Island; CAS 3987 (1), Guaymas; SIO 47-53 (1), Gulf of California, 15

54 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

mi. from mouth of Rio Colorado, Baja California. Panama: USNM
190591 (2), Panama Bay, 1-3 mi. SE of Isla Taboga and 2—4 mi. from
Rio Pacora; UCLA 53-275 (3), Panama Bay, between Panama City
and Punta Gorda; UCLA 58-305 (1), Panama Bay, offshore between
Rio Chico and Punta de la Plata; FMNH 62438 (2), Punta Chame and
Punta Anton, Panama Bay; SU 11584 (holotype of Sphyrna vesper-
tina), Panama City; SU 11881 (one paratype of S. vespertina), Panama
City; MCZ 80 (1), MCZ 516 (1), no further data.

Diaenosis.—Sphyrna tiburo vespertina differs from S. tiburo tiburo
in having a slightly broader and more pointed head. ‘These differences
are illustrated in figures 12 and 13.

250
240

© SPHYRNA TIBURO VESPERTINA
220
e
se °

210
200
190
180
170
160

HEAD WIDTH IN
THOUSANDTHS OF TL

250 300 350 400 450 500 550 600 650 700 750 800 850 900 950
TL IN MM

Ficure 13.—Comparison of head width in the two subspecies of Sphyrna tiburo. (Figure by
Paul Laessle.)

Sphyrna (Platysqualus) media Springer
Ficures 14, 15, 16, 17, 21 f, g, 22f; Puats 7c

Sphyrna tudes (misidentification) Jordan, 1895, p. 383. (Mazatlan, Sinaloa.)

Cestracion tudes (misidentification) Meek and Hildebrand, 1923, p. 59. (Coldn,
Panama.)

?Sphyrna tiburo (misidentification) Kumada and Hiyama, 1937, p. 17. (Descrip-
tion could apply to either S. tiburo, S. media, or S. corona; illustration is either
of S. media or S. corona.)

Sphyrna media Springer, 1940a, pp. 161-169. (Original description; illustration of
head; holotype, a partially defleshed 900 mm. male, SU 11583; type locality,
Mazatlan, Sinaloa, Mexico; one paratype, a 745 mm. female, USNM 28160,
from Mazatlan.)

Sphyrna nana Sadowsky, 1965, pp. 1-12. (Original description; illustrations of
head and body; holotype, a 1,330 mm. female, SAo Paulo Zoological Museum
[POB] 7172; type locality, off Cananéia, Brazil [lat. 25° 8., long. 47°52’ W.].)

SPECIMENS EXAMINED.—(Range in size: 337-900 mm. TL; heads
of larger specimens also examined; estimated length of largest, ca.
1025 mm.). Atlantic Ocean (including Caribbean Sea): Brazil: SU
52716 (head only), Vitoria. Panama: MCZ 500 (1), USNM 79284
(1), Colon market.

NO. 3539 HAMMERHEAD SHARKS—GILBERT 55

Pacific Ocean: Costa Rica: USNM 196140 (6), Gulf of Nicoya.
Mexico: UCLA 58-47 (2, including one head), Gulf of California,
south of Bahia Topolobampo, off Isla San Ignacio and Isla Macapule,
Sinaloa; FMNH 63093 (2), above San Benito, Chiapas; SIO 60-88
(3), Gulf of California, 21°55’ N., 105°37’ W.; SU 11583 (holotype of
Sphyrna media—partial skin), Mazatlan, Sinaloa; USNM 28160 (1
paratype of Sphyrna media), Mazatlan, Sinaloa. Panama: USNM
190593 (4), UCLA 58-304 (4, including three heads), (all originally
UCLA 58-304), Panama Bay, offshore between Punta de Hicacal
and Rio Pasiga; USNM 50376 (1), no further data.

Initially Sphyrna media and Sphyrna corona were regarded by me as
synonymous. This conclusion was based mainly on the fact that dif-
ference in anterior contour of the head, which was the primary basis
for Springer’s original separation of the two forms, does not remain
constant, the head shape grading from the more evenly rounded con-
dition of S. media to the more lobate situation found in S. corona. In
addition, the chondrocrania of the two forms are very similar, as are

l hee = ee EEL ai Map 4.—Distribution of subspecies.
Bey Wale . © Sphyrna tiburo tiburo
Ai oe A Sphyrna tiburo ves pertina
(Solid=specimens examined; hollow=
confirmed literature references.)

the various external morphological features. Closer examination of the
chondrocrania, however, revealed several relatively small, but none-
theless constant, differences. With a firm basis for separation thus
available, it was found that other morphological differences, which
were at first thought to be due to individual variation, also could be
used to separate the two species. The various distinguishing features
are discussed in the diagnoses of the respective species.

Draanosts.—Characters mentioned in the diagnosis of the subgenus
Platysqualus are not repeated here.

Sphyrna media differs from the closely related S. tudes in the fol-
lowing characters: Anterior margin of head more rounded (partic-
ularly in large specimens), with no pronounced median notch; inner
narial groove absent; anterior-median pore patch on underside of head
with the posterior margin more broadly rounded than in S. tudes and
with a pointed median-posterior extension (fig. 22f); narial depression
very shallow; first dorsal fin of larger specimens (600 mm. or more)
less erect and less broadly triangular, a line drawn perpendicularly
from apex of fin not intersecting base of fin.

56 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Sphyrna media differs from S. corona in having a narrower postor-
bital process to the chondrocranium, with the anterior edge of this
structure posterior, rather than anterior, to the angle formed by the
juncture of the innerorbital and anterior mediorbital cartilages (pl.7c) ;
a slightly deeper anterior fontanelle; and shorter rostral cartilages.
The last difference has been objectively shown by dividing the trans-
verse distance between the tips of the preorbital cartilages into the

b

PRA MA A Atel a

Ficure 14.—Sphyrna media: a, adult male, 900 mm. TL, from Panama Bay (UCLA 58-304);
b, head of same specimen, about 0.12 X; c, dermal denticles of same specimen; d, upper

and lower teeth, left side, of same specimen, about 1.5 X. (Drawings by Dorothea B.
Schultz.)

leneth of the rostral cartilage. For S. media the range of values thus
obtained is from 18.1 to 24.0 percent, and for S. corona from 25.9 to
30.5 percent; these have been plotted and may be seen in figure 17.

Sphyrna media also differs from S. corona in having a slightly
shorter snout. A comparison of snout lengths may be seen in figures
15b, 16, and 186. Although there is some overlap in values for the
two species, when specimens of a comparable size are compared lower
values are obtained for S. media.

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NO. 3539 HAMMERHEAD SHARKS—GILBERT Dt

Sphyrna media also differs from S. corona in having: The anal fin
more deeply falcate and with a more pointed apex (figs. 14a, 15a);
anterior-median pore patch on underside of head with the posterior
margin more broadly rounded and with a pointed median-posterior
extension (fig. 22f); a more pointed upper precaudal pit in smaller
specimens (fig. 21g); and a slightly wider mouth. When the trans-
verse distance between the corners of the mouth is divided by the
head width, the values obtained are somewhat greater for S. media
Finally, Sphyrna media may be a larger species. The largest specimen
of Sphyrna media seen (a head) was calculated to be 1025 mm. TL,
while of the 17 specimens of Sphyrna corona examined, the largest
(the holotype) measured only 672 mm. More specimens of S. corona
must be seen to definitely confirm this, however.

A comparison of Sphyrna media with other members of the genus is
presented in table 1.

DescriptTion.—Meristic data appear in table 7. Characters men-
tioned in the diagnoses of the subgenus Platysqualus and of S. media
are not repeated here.

eee
pee) /)

ed

Figure 15.—Sphyrna media: a, juvenile female, 356 mm. TL, from Chiapas, Mexico (FMNH
63093); b, head of same specimen. (Drawings by Paul Laessle.)

Head moderately expanded, the greatest width measuring from
22.3 to 27.3 percent of TL; character of anterior-median margin of
head somewhat variable, evenly rounded in some individuals, with no
median indentation, and with a broad, very shallow median depres-
sion in others; outer posterior corner of lateral expansion of head
situated anterior to corner of mouth; narial flap broad, bluntly pointed
at tip, with the broadly curved outer edge curving sharply inward; a

58 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

very short labial furrow present at corner of lower jaw in larger speci-
mens, not noticeable in smaller specimens.

Origin of first dorsal fin slightly posterior to axil of pectoral fin,
a perpendicular line drawn ventrally from origin intersecting anterior
third of pectoral fin; base of first dorsal fin one-third to one-half
length of head and about twice length of pectoral fin base; anterior
margin of first dorsal fin broadly and evenly curved, particularly
in larger individuals; distal margin of first dorsal fin rather deeply
falcate, particularly on lower two-thirds of fin; lobe of first dorsal
fin from two-fifths to one-half length of first dorsal base; origin of
second dorsal fin situated about two-fifths of way back above base
of anal fin; second dorsal fin tall, its greatest height about three-
fourths length of base and from three-fourths to four-fifths of greatest
height of anal fin; second dorsal fin base from one-half to five-eighths
of anal fin base; lobe of second dorsal fin rather long, its length about
a third again greatest height of fin and extending from two-thirds
to three-fourths of distance from posterior part of second dorsal
base to upper precaudal pit; pectoral fin from two-fifths to two-thirds
length of head (higher values usually apply to larger individuals) ;
pectoral fin two-thirds as broad as long; length of pectoral fin base
from five-eighths to three-fourths length of anal fin base; inner two-
thirds of anterior margin of pectoral fin straight, outer third slightly
convex; distal margin of pectoral fin slightly faleate; apex of pectoral
broadly pointed in small specimens, slightly more acute in larger
individuals; inner corner of pectoral narrowly rounded; length of
pelvic fin from two-thirds to four-fifths length of anal fin; length
of pelvic fin base from five-ninths to three-fourths length of anal
fin base; anterior margin of pelvic fin straight or slightly convex;
distal margin of pelvic very slightly falcate; anal fin from one-tenth
to one-third again as long as pelvic fin; length of anal fin base one-
fourth to four-ninths again as long as pectoral and pelvic fin bases
and from three-fifths again to nearly twice as long as second dorsal
fin base; height of anal fin one-fifth to one-fourth greater than height
of second dorsal fin; length of caudal fin from one-fourth to nearly
one-third of total body length; upper margin of caudal fin evenly
convex; terminal part of caudal slightly less than one-fourth of total
length of fin; lower posterior margin of terminal part of caudal fin
nearly straight or slightly faleate; both terminal and lower apices
of caudal broadly pointed; lower lobe of caudal appearing relatively
narrow and long, particularly in larger individuals, the distance from
the angle included by the upper and lower caudal lobes to tip of latter
going about 1%; times in anterior margin of lower caudal lobe; lower
caudal lobe about one-third as long as upper lobe; lower caudal lobe
rather erect, sloping posteriorly at about a 25 to 30° angle from a

NO. 3539 HAMMERHEAD SHARKS—GILBERT 59

perpendicular to the body axis: anterior margin of lower caudal lobe
weakly convex toward base, more strongly convex toward apex;
posterior margin of lower caudal lobe nearly straight.

Anterior margin of rostral node of chondrocranium varying slightly,
from broadly convex to broadly and shallowly concave; rostral
fenestra absent; anterior fontanelle (viewed dorsally) deep and broadly
U-shaped, with the sides of the U slanting slightly inward; a distinct
median notch present on anterior fontanelle; fairly prominent (though
not as prominent as in S. tudes), broad, obtusely pointed wings present

130
© SPHYRNA CORONA
95 ° e SPHYRNA MEDIA

125

DISTANCE FROM SNOUT TO SYMPHYSIS
IN THOUSANDTHS OF TOTAL LENGTH

75
200 250 300 350 400 450 500 550 600 650 700 750 800 850 900 950
TLIN MM

Ficure 16.—Comparison of snout length in Sphyrna media and Sphyrna corona. (Figure
by Paul Laessle.)

on sides of rostral node; anterior wing of olfactory cartilage fairly
prominent (though not as prominent as in S. tudes), pointed, and pro-
jecting medially; tip of anterior part of preorbital process short (not
as short as in S. mokarran) and rather thick, with a sharp angle on
the anterior edge.

Total number of vertebrae 191 to 196 (10 specimens examined) ;
body vertebrae 100 to 103, there being from 7 to 14 more body than
caudal vertebrae.

13—1-12
Teeth ———_,), ———_) —____,, —__~__~ and ees

60 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

(first two counts listed by Springer, [1940a, p. 163]); upper teeth
oblique, becoming increasingly so toward corners of mouth in upper
jaw; teeth 6 to 9 or 10 longest, the outer teeth very low, but with
a definite cusp; in lower jaw, teeth 2 through 8 longest (tooth 1 not
markedly smaller however), the outermost two or three teeth very
short, round, and without a definite cusp; one or two series of teeth
functional in alternating rows along sides of upper jaw and two to
three rows along sides of lower jaw.

Dermal denticles (examined from upper part of back beneath first
dorsal fin) evenly and closely spaced, overlapping but little, the blades
thick and rather strongly arched; length of denticle (measured to tip
of median marginal ‘‘tooth’’) less than width; small specimens with
three, larger specimens usually with up to five, sharp-topped ridges,
and as many sharp-pointed marginal teeth (especially deep and
pointed in small individuals) on denticle; median tooth on denticle
slightly longer than more distal teeth; pedicel short and broad.

Individuals gray or grayish brown dorsally, becoming progressively
more pale below; no markings on fins.

Stratus or Sphyrna nana.—Sadowsky (1965) recently described a
new species of hammerhead, Sphyrna nana, from southern Brazil,
based on a single adult female, 1,330 millimeters long. This form
was said to differ from S. media in having a shorter snout, a larger
orbit, the anterior margin of the head not regularly oval and with
prenarial convexities, a shorter distance between the eye and nares,
a shorter distance between the nares, a shorter upper caudal lobe, a
greater interdorsal distance, and a longer first dorsal fin base. Un-
fortunately, no mention was made either of the morphology of the
chondrocranium or of the arrangement of the mucal pores on the
ventral surface of the head.

The following measurements were given for the holotype of S. nana:
Head width (190); snout to first gill slit (169); snout to first dorsal
origin (308); snout to second dorsal origin (637); distance between
first and second dorsal bases (233); snout to pectoral insertion (204);
snout to pelvic insertion (466); horizontal diameter of orbit (15);
length of first dorsal base (95); length of second dorsal base (43);
length of second dorsal lobe (45); height of second dorsal fin (37);
length of anal base (80); length of anal lobe (33); height of anal fin
(36); length of pectoral base (54); length of pelvic base (34); length
of caudal fin (251). If one compares these measurements with those
listed for S. media in table 7, it can be seen that, for those characters
that undergo little or no proportional change with an increase in total
body length, the values for S. nana generally fall within the range of
values for S. media. For those characters that undergo pronounced

NO. 3539 HAMMERHEAD SHARKS—GILBERT 61

allometric change with an increase in length (head width, distance
from snout to symphysis, head length, distance between first and
second dorsal bases, and snout to pectoral insertion), the values given
for S. nana are about what one would expect in individuals of media
around 1,300 millimeters long. Of the various measurements listed
for S. nana, only three (distance from snout to pelvic insertion,
horizontal diameter of orbit, and length of caudal fin) seem to fall
outside the range of variation one would normally expect to find in
a large individual of S. media.

Thus, with few exceptions the proportional measurements in S. nana
and S. media are very similar, and those in which there is slight
disagreement are not sufficiently different to warrant recognition of
a second species. The contour of the anterior margin of the head,
another character that supposedly differentiates S. nana and S. media,
falls within the normal range of variation for S. media (as discussed
elsewhere in this paper).

In conclusion, while the validity of S. nana cannot be categorically
denied until the natures of the chondrocranium and the pore patches
on the ventral surface of the head are known, I so far can find no
evidence to indicate that it is specifically distinct from S. media.

VARIATION.—Sphyrna media shows more than the usual amount of
variation in configuration of the anterior margin of the head. Some
specimens have an evenly rounded head (fig. 155), while in others
the head is faintly lobate, with a slight median indentation or scallop
(fig. 146). Those individuals with more lobate heads may be dis-
tinguished only with difficulty from specimens of S. corona. That
this condition is not strictly a function of size is shown by the fact
that one of the largest specimens of S. media examined, a 900 mm.
male (UCLA 58-304) from Panama Bay, was at first identified as
S. corona on the basis of head shape.

Variation in the median scallop on the anterior margin of the head
was found to result from a comparable variation in the tip of the
rostral cartilage. Considerable variation was found also in the degree
of development and/or presence of the accessory rostral cartilages,
a single series of specimens sometimes showing all degrees of develop-
ment.

RancE.—Sphyrna media occurs in the eastern Pacific, from the
Gulf of California south at least as far as Panama Bay, in the southern
Caribbean, and in the southwestern Atlantic (map 5).

The occurrence of Sphyrna media in the Caribbean is confirmed by
two specimens (USNM 79284, MCZ 500) from the Colon (Panama)
market, which were collected around the turn of the century. Pres-
ence of the species in this area is further substantiated by Stewart

62 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Springer (1949, pp. 17-26), who lists (under the name ‘‘scoophead
shark’’) a female, 1,125 mm. long, containing eight embryos, which
was collected in April 1949, from the Gulf of Paria, Trinidad.

There is, in the Stanford University collection, the head of a speci-
men of this species (SU 52716) from Vitoria, Brazil. Since this
locality is so far removed from Trinidad, the next closest place from
which S. media has been recorded, one might regard the record with
suspicion. However, the intervening area has not been well collected,
and further exploration will likely show the species to be more widely
distributed in the western Atlantic than presently appears.

350

O SPHYRNA CORONA
e SPHYRNA MEDIA

300

250

200

LENGTH OF ROSTRAL CARTILAGE
WIDTH OF CHONDROCRANIUM *

150
200 250 300 350 400 450 500 550 690 650 700 750 800 850 900 950

TL IN MM

Figure 17.—Comparison of rostral cartilage length (expressed as a ratio) in Sphyrna media
and Sphyrna corona. Width of chondrocranium measured from tips of preorbital cartilages.

(Figure by Paul Laessle.)

Sphyrna (Platysqualus) corona Springer
Ficures 16, 17, 18, 21h, 22g; PLATE 7p

?Sphyrna tiburo (misidentification) Kumada and Hiyama, 1937, p.17. (Descrip-
tion could apply to either S. tzburo or S. media; illustration is either of S. media
or S. corona.)

Sphyrna corona Springer, 1940a, pp. 161-169. (Original description; illustration
of head; holotype, a 672 mm. male, SU 11882; type locality, Panama Bay,
at Panama City, Panama.)

SPECIMENS EXAMINED.—(Range In size: 231-672 mm. TL). Pacific
Ocean: Colombia: UCLA 51-283 (1), Boca Candelaria, 50 mi. south
of Buenaventura. Costa Rica: UCLA 54-167 (1), Golfo de Nicoya,
Chira Flats, off Isla Chira. Mexico: FMNH 72521 (2), off Chiapas
coast, above San Benito. Panama: CAS (W 53-273) (3), Panama
Bay, 1-3 mi. southeast of Taboga, and 2—4 mi. off Rio Pacora; UCLA

NO. 3639 HAMMERHEAD SHARKS—GILBERT 63

53-275 (9), Panama Bay, between Panama City and Punta Gorda;
SU 11882 (holotype of Sphyrna corona), Panama City.

Draenosis.—Characters mentioned in the diagnosis of the subgenus
Platysqualus are not repeated here.

Sphyrna corona differs from the closely related S. tudes in much the
same way that S. media differs from that species (see diagnosis of S.
media). Unlike S. media, however, the shape of the anterior-median
pore patch on the underside of the head is very similar in S. corona
and S. tudes (figs. 22g,h).

Sphyrna corona differs from 8. media in having a broader postorbital
process to the chondrocranium, with the anterior edge of this structure
anterior, rather than posterior, to the angle formed by the juncture

Ficure 18.—Sphyrna corona: a, juvenile female, 371 mm. TL, from Chiapas, Mexico
(FMNH 72521, ex 63093); b, head of juvenile male, 402 mm. TL, from Panama Bay
(UCLA 53-275), about 0.6 X natural size. (Fig. a drawn by Paul Laessle, fig. b by
Dorothea B. Schultz.)

of the innerorbital and anterior mediorbital cartilages (pls. 7c,p); a
slightly shallower anterior fontanelle; and longer rostral cartilages.
(The last character is discussed at greater length in the diagnosis of
S. media.)

Sphyrna corona also differs from §. media in having: A slightly
longer snout (fig. 18b); a less deeply falcate anal fin with a more
rounded apex (fig. 18a); anterior-median pore patch on underside of
head with the posterior margin less broadly rounded and without a
pointed median-posterior extension (fig. 22g); a more broadly rounded
upper precaudal pit in smaller specimens (fig. 21h); and a slightly
narrower mouth. Finally, S. corona may not reach as large a max-

64 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

imum size as S. media, because the largest specimen examined (672
mm.) was little over half as large as the largest specimen of S. media
that was seen.

A comparison of Sphyrna corona with other members of the genus
is presented in table 1.

DeEscrRIPTION.—Since most of the morphological characters in
Sphyrna corona are similar to those found in S. media, the following
description has been condensed so as to avoid repetition. Only those

Map 5.— Distribution of species.
@ Sphyrna tudes A Sphyrna media
WV Sphyrna corona
* Sphyrna media and Sphyrna corona
(Solid= specimens examined; hollow=
confirmed literature references.)

characters in which actual counts or measurements are cited are
listed in the following account. For other characters see either the
diagnoses of S. corona or the subgenus Platysqualus, or the description
of S. media.

Meristic data appear in table 8.

Head moderately expanded, the greatest width measuring from

Eros
23.7 to 29.1 percent of TL; teeth a (count given for holotype
: 14-14 14-14
by Springer, 1940a, p. 163) ae and Sag

RanGce.—Sphyrna corona has one of the most restricted ranges of any
species of hammerhead, occurring only in the eastern Pacific from
southern Mexico south at least to the coast of Colombia (map 5).

Sphyrna (Platysqualus) tudes (Valenciennes)
Figures 19, 20, 212, 22h; Puarss 4, 8

Zygaena tudes Valenciennes, 1822, pp. 225-226. (Illustration of head; original
description [apparently based on two species, S. tudes and S. mokarran);
types from Nice and Cayenne are S. tudes, as here recognized, while type
from Coromandel probably is S. mokarran.)

Sphyrna tudes Miiller and Henle, 1841, p. 53. (In part; identification by reference
to Valenciennes, 1822.)

Sphyrna bigelowi Springer, 1944, pp. 274-276. (Original description; illustration;
type locality, Uruguay; holotype, a 385 mm. male [USN M 87682]; paratypes,
a 395 mm. male from Uruguay [USNM 120751], and a 900 mm. female
from Rio de Janeiro, Brazil [MCZ 463].)

SPECIMENS EXAMINED.—(Range in size: 213-900 mm. TL). West-
ern Atlantic Ocean (including Caribbean Sea and Gulf of Mexico):

NO. 3539 HAMMERHEAD SHARKS—GILBERT 65

Brazil: MCZ 463 (paratype of S. bigelowi), Rio de Janeiro. French
Guiana: MNHWN 1019 (lectoparatype of Zygaena tudes), Cayenne.
Surinam: ANSP 377 (1), ANSP 516 (1), (no further data); USNM
156721 (1), 1 mi. southeast of Paramaraibo light ship; USNM 159197
(1), USNM 159199 (1), 06°27’ to 06°25’N., 55°05’ to 55°10’W.;
USNM 159227 (2), 06°20.5’N. to 06°19.5’N., 54°54’W. to 54°49’W.;
USNM 159228 (4), USNM 159229 (4), 06°22’N., 55°04’ to 55°08’ W.
United States (Mississippi): USNM 195957 (8), Gulf of Mexico,
29°54’N., 88°06’W. (Oregon Sta. 2110). Uruguay: USNM 87682
(holotype of Sphyrna bigelowr), no further data; USNM 120751 (para-
type of S. bigelowr), no further data. Venezuela: USNM 123217
(head), Gulf of Venezuela, Anway Bay.

Mediterranean Sea: France: MNHN 1049 (lectotype of Zygaena
tudes), Nice.

NomENCLATURE.—The original description of Zygaena tudes (Va-
lenciennes, 1822, pp. 225-226, pl. 12, figs. la and 6b) was based on
specimens, in the Museum National d’Histoire Naturelle, from Nice,
France (cat. no 1049); Cayenne, French Guiana (cat. no. 1019); and
Coromandel, India (specimen lost). As has previously been shown,
these specimens comprise two species: The one currently called
Sphyrna tudes (Gilbert, 1961, p. 480), which is characterized by the
tip of the first dorsal fin extending beyond the insertion of the pelvic
fins, the first and fifth gill slits nearly equal in length, with the latter
situated above the insertion of the pectoral fin, and a small maximum
size (perhaps not over 4 or 5 feet) ; and the one presently called Sphyrna
mokarran (Gilbert, 1961), characterized by the tip of the first dorsal
fin not reaching the insertion of the pelvic fins, the first gill slit longer
than the fifth gill slit, with the latter situated posterior to the insertion
of the pectoral fin, and a large maximum size (up to 20 feet). I here
restrict the name tudes to those specimens from Nice and Cayenne,
and designate as lectotype of Zygaena tudes the 346 mm. female (no.
1049) on which the illustration in the original description was based
(pl. 4).

Diaenosis.— Characters mentioned in the diagnosis of the subgenus
Platysqualus are not repeated here.

Sphyrna tudes differs from the closely related S. corona and S. media
in the following characters: Anterior margin of head straighter (par-

ticularly in large specimens), with a pronounced median notch; inner
_narial groove present; anterior-median pore patch on underside of
head divided into two broadly rounded lobes and separated by a

deep, wide, median indentation (fig. 225); narial depression slightly

_ deeper; first dorsal fin of larger specimens (600 mm. or more) more

|

_ erect and more broadly triangular, a line drawn perpendicularly from

_ apex of fin intersecting base of fin.

219-937—67——5

66 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

A comparison of Sphyrna tudes with other members of the genus is
presented in table 1.

Descrirtion.—Meristic data appear in table 9. Characters men-
tioned in the diagnoses of the subgenus Platysqualus and of S. tudes
are not repeated here.

Head moderately expanded, the greatest width measuring from
27.5 to 31.5 percent of TL; inner narial groove extending 3.5 to 4.0
percent of distance from narial openings to median notch; outer
posterior corner of lateral expansion of head situated posterior to

SRR RR eee ice a
vi iN Ay SFR eS Eta

it ley Ain A penne Oh

A
ah ee a ee ee

Ficure 19.—Sphyrna tudes: a, subadult male, 886 mm. TL, from Rio de Janeiro, Brazil
(MCZ 463); b, upper and lower teeth, left side, of same specimen, about 1.5 X;3 c, fifth
upper tooth; d, twelfth upper tooth; ¢, fifth lower tooth; f, twelfth lower tooth; g, dermal
denticles of MCZ 463, about 17 ; h, dermal denticle, side view, about 34 X; i, dermal
denticle, apical view, about 34 X. (¢-f, about 3 X.) (From Bigelow and Schroeder,
1948, p. 410.)

corner of mouth in smaller specimens, anterior to corner of mouth in
larger individuals; narial flap rather narrow (particularly in smaller
specimens), pointed at tip, with the broadly curved outer edge curving
sharply inward; length of snout about two-fifths of head width in
smaller specimens to about one-third of head width in larger speci-
mens; greatest transverse distance between corners of mouth from
one-fifth to one-fourth of greatest width of head; a very short labial
furrow present at corner of lower jaw in larger specimens, not notice-
able in smaller specimens.

Origin of first dorsal fin slightly posterior to axil of pectoral fin,
a perpendicular line drawn ventrally from origin intersecting anterior

NO. 3539 HAMMERHEAD SHARKS—GILBERT 67

third of pectoral fin; base of first dorsal fin three-eighths to four-ninths
length of head and from a third again to not quite twice length of
pectoral fin base; anterior margin of first dorsal fin broadly and evenly
curved, particularly in larger individuals, with the part nearest the
apex a little more strongly curved; upper distal margin of first dorsal
broadly convex, lower margin broadly but distinctly falcate; lobe of
first dorsal fin a little less than one-half length of first dorsal base;
origin of second dorsal fin situated about two-fifths of way back
above base of anal fin; second dorsal fin tall, its greatest height from
two-thirds to four-fifths length of base and from two-thirds to four-

C

Figure 20.—Sphyrna tudes: a, juvenile female, 402 mm. TL, from Mississippi (USNM
195957); b, head of same specimen, about 0.3 X; c, head of small adult, from Gulf of
Venezuela (USNM 123217), about 0.18 X. (Drawings by Dorothea B. Schultz.)

fifths of greatest height of anal fin; second dorsal fin base slightly
greater than one-half of anal fin base; lobe of second dorsal fin long,
its length from a third to half again greatest height of fin, and ex-
tending from two-thirds to three-fourths of distance from posterior
part of second dorsal base to upper precaudal pit; pectoral fin from
one-half to two-thirds length of head (higher values usually apply to
larger individuals); pectoral fin two-thirds as broad as long; length of
pectoral fin base from two-thirds to three-fourths length of anal fin
base; inner two-thirds of anterior margin of pectoral fin straight,
outer third distinctly convex; distal margin of pectoral fin nearly
straight or very slightly falcate; apex of pectoral broadly pointed;

68 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

inner corner of pectoral fin narrowly rounded; length of pelvic fin
from two-thirds to nine-tenths length of anal fin; length of pelvic fin
base from five-ninths to seven-eighths length of anal fin base; anterior
and distal margins of pelvic fin nearly straight; anal fin from one-
eighth to one-third again as long as pelvic fin; length of anal fin base
from one-third to one-half again as long as pectoral and pelvic fin
bases and about twice as long as second dorsal fin base; height of

a é Z
Peres : Ye, poe 2

;

Ficure 21.—Appearance of upper precaudal pit: a, Sphyrna blochit, 615 mm. female, from
North Borneo (FMNH 21836); b, S. mokarran, 675 mm. male, from Florida (USNM
106543); c, S. zygaena, 580 mm. male, from Virginia (USNM 119699); d, S. lewini, 495
mm. female, from Texas; ¢, S. ¢. tiburo, 880 mm. female, from Texas (USNM 127106);
f, S. media, 900 mm. female, from Panama Bay (UCLA 58-304); g, S. media, 356 mm.
female, from Chiapas (FMNH 63093); h, S. corona, 371 mm. male, from Chiapas (FMNH
72521, ex. 63093); i, S. tudes, 825 mm. male, from Surinam (USNM 156721). (Figs. a-f,
i drawn by Dorothea B. Schultz; figs. g, A by Paul Laessle.)

anal fin from one-fifth to one-third greater than height of second
dorsal fin; anal fin deeply falcate toward apex, nearly straight toward
tip of lobe; length of caudal fin nearly a third of total body length;
upper margin of caudal fin slightly convex; terminal part of caudal
about one-fourth of total length of fin; lower posterior margin of
terminal part of caudal fin nearly straight or slightly falcate; both
terminal and lower apices of caudal broadly pointed; lower lobe of

|

NO. 3539 HAMMERHEAD SHARKS—GILBERT 69

caudal appearing relatively narrow and long, the distance from the
angle included by the upper and lower caudal lobes to tip of latter
going about 1% times in anterior margin of lower caudal lobe; lower
caudal lobe about one-third as long as upper lobe; lower caudal lobe
rather erect, sloping posteriorly at about a 25 to 30° angle from a
perpendicular to the body axis; anterior margin of lower caudal lobe
weakly convex; posterior margin of lower lobe nearly straight.

Anterior margin of rostral node of chondrocranium with a broad,
deep, median indentation; accessory rostral cartilages usually present;
rostral fenestra usually absent (present in one individual examined) ;
anterior fontanelle (viewed dorsally) deep and broadly V-shaped, with
an indistinct median notch on the posterior margin; prominent,
broad, obtusely pointed wings present on sides of rostral node;
anterior wing of olfactory cartilage prominent, pointed, and projecting
medially; tip of preorbital process short (though not as short as in
S. mokarran), rather thick, with a sharp angle on the anterior edge;
postorbital process terminating in an angular knob, the anterior
corner of which may be either broadly pointed or narrowly rounded,
and the posterior corner of which forms a broadly rounded angle.

Total number of vertebrae numbering from 195 to 202 (five speci-
mens examined); body vertebrae numbering from 107 to 110, there
being from 12 to 25 more body than caudal vertebrae.

15-0-15 16—0-16

Seer RICHI yoni
for lectotype) ; upper teeth oblique, becoming increasingly so toward
corners of mouth in upper jaw; teeth 4 or 5 to 11 longest, the outer
teeth very low, but with a definite cusp; in lower jaw teeth 2 to 7 or
8 longest, the outermost two very short, rounded, without a definite
cusp; one or two series of teeth functional in alternating rows along
sides of lower jaw, and two to three rows along sides of lower jaw.

Dermal denticles (examined from upper part of back beneath first
dorsal fin) evenly and closely spaced, overlapping but little, the blades
thick and rather strongly arched; length of denticle (measured to tip
of median marginal “‘tooth’’) less than width; small specimens with
three, larger specimens usually with up to five, sharp-topped ridges,
and as many sharp-pointed marginal teeth (especially deep and pointed
in small individuals) on denticle; median tooth on denticle slightly
longer than more distal teeth; pedicel long and rather slender.

Individuals gray or grayish brown dorsally, becoming progressively
more pale below; no markings on fins.

VARIATION.—Examination of the lectotype (designated elsewhere
in this paper) of Zygaena tudes, a 346 mm. female from Nice, France,
indicates that the Mediterranean and western Atlantic populations of
Sphyrna tudes are morphologically indistinguishable. The only un-

(two specimens counted; latter count

70 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

usual feature of the lectotype is the presence of a small rostral fen-
estra (pl. 88). However, the occasional presence of such a fenestra
in S. zygaena and S. blochii (in which species it is ordinarily absent)
suggests that the type specimen is merely a variant with regard to this
character.

Ere Ens

Ficure 22.—Distribution of pores on ventral side of head: a, Sphyrna blochii; b, S. mokarran;
c, S. zygaena; d, S. lewint; e, S. tiburo; f, S. media; g, S. corona; h, S. tudes. (Drawings
by Paul Laessle.)

oS
oO

Rancu.—Sphyrna tudes is restricted to the western Atlantic Ocean
and the western Mediterranean Sea. In the western Atlantic it occurs
from Uruguay (Springer, 1944, p. 274) to the northern Gulf of Mexico
(Gilbert, 1961, p. 480). In the western Mediterranean it is recorded
from Nice, France (the type locality) and Livorno, Italy (Tortonese,
1950a, pp. 31, 32; 1950b, p. 1031).

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NO. 3539 HAMMERHEAD SHARKS—GILBERT 15

PELLEGRIN, JACQUES
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76 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

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NO. 3539 HAMMERHEAD SHARKS—GILBERT 77

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NO. 3539

HAMMERHEAD SHARKS—GILBERT 79

TABLE 2.— Measurements of Sphyrna blochii (proportions expressed in thousandths
of total length)

Size range in mm.

300-399 400-599 600-799 >800
Number of specimens
4 4 2 1
Head width 337-492 (391. 2) 441-481 (463. 8) 432-456 (444. 0) 420
Internarial distance 152-165 (157. 5) 160-167 (164. 0) 153-164 (158. 5) 152
Snout to symphysis 66-78(70. 8) 63-72 (66. 5) 56-64(60. 0) 54
Head length 221-237 (229.0) 197-213(205.0) 195-197 (196.0) 195
Snout to Ist gill slit 173-176(174.5) 151-168 (159.5) 153-154(153. 5) 153
Snout to 1st dorsal origin 244-266(256.5) 217-258 (228.8) 214-219(216.5) 214
Snout to 2nd dorsal origin 665-626 (583.3) 560-575 (565.8) 577-584(580.5) 567
Distance between 1st and 2nd dorsal
bases. 191-248 (216.5) 209-242 (226.8) 243-247 (245.0) 232
Snout to pectoral insertion 187-224(204.8) 193-206 (197.5) 171-204 (187.5) 191
Snout to pelvic insertion 392-432(415.3) 390-420(408.3) 395-397 (396.0) 395
Horizontal diameter of orbit 14-21(16.5) 14-18(15.8) 13-15(14. 0) 13
Length of Ist dorsal base 105-113 (110.3) 103-114 (107.8) 111-117(114.0) 121
Length of 2nd dorsal base 35-44(39.3) 34-40(37.5) 39-40(39.5) 44
Length of pectoral base 51-56 (53.5) 50-58(53.5) 54-63 (58.5) 55
Length of pelvic base 52-59(55.3) 53-54(53.5) 59-60 (59.5) 61
Length of 2nd dorsal lobe 43-47 (44.8) 47-53 (50.0) 50-57 (53.5) 51
Height of 2nd dorsal fin 22-30 (26. 0) 22-27 (24.3) 26-27 (26.5) 28
Length of anal base 58-62(60.0) 55~-65(60.3) 69-71(70. 0) 64
Length of anal lobe 34-39 (36.3) 39-44 (40.8) 38-45 (41.5) 39
Height of anal fin 32-36 (33.8) 34-39 (37.0) 37-38 (37.5) 40
Length of caudal fin 317-329(323.5) 332-339 (335.8) 335-338 (336.5) 331

TABLE 3.—Measurements of Sphyrna mokarran (proportions expressed in thou-
sandths of total length)

N 400-599 N_ 600-799

Head width

Atlantic 3 231-244(239.3) 1 235

Pacific 4 225-230(228.3) 1) 273
Internarial distance

Atlantic 3 170-175(173.0) 1 170

Pacific 4 157-173(164.8) 1 190
Snout to symphysis

Atlantic 3 73-81(78.0) t-73

Pacific 4 66-80(74.3) 1 82
Head length

Atlantic 3 220—232(227.3) 1 226

Pacific 4 212-235(226.5) 1 250
Snout to Ist gill slit

Atlantic 3 177-187(182.3) oa

Pacific 4 166-187(179.8) 1 200
Snout to 1st dorsal origin

Atlantic 3 273-289(280.7) 1 284

Pacific 4 253-289(277.3) 1, 285

Size range in mm. (N=number of specimens)

80 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

TABLE 3.—Measurements of Sphyrna mokarran (proportions expressed in thou-
sandths of total length) —Continued

Size range in mm. (N=number of specimens)

N 400-599 N_ 600-799

Snout to 2nd dorsal origin

Atlantic 3 581-600(592.3) 1 595

Pacific 4 555-579(568.5) 1 599
Distance between Ist and 2nd

dorsal bases

Atlantic 3 193-211(202.7) 1 213

Pacific 4 189-205(196.0) 1 208
Snout to pectoral insertion

Atlantic 3 197-215(204.3) 1 207

Pacific 4 187-226(211.0) 1 244
Snout to pelvic insertion

Atlantic 3 421-436(427.3) 1 434

Pacific 4 432-449(439.5) 1 469
Horizontal diameter of orbit

Atlantic 3 20-25(22.7) Lige22

Pacific 4 29-33(81.3) 1 226
Length of 1st dorsal base

Atlantic 3 97-100(99.0) 1 98

Pacific 4 85-98(92.7) 1 106
Length of 2nd dorsal base

Atlantic 3 54-67(60.0) 1 52

Pacific 4 41-51(45.8) 1 56
Length of 2nd dorsal lobe

Atlantic 3 40-46(42.3) 1 39

Pacific 4 46-48(46.5) 1 45
Height of 2nd dorsal fin

Atlantic 3 28-39(33.3) 1 38

Pacific 4 27-41(85.3) 1 47
Length of anal base

Atlantic 3 61-67(64.0) 1 61

Pacific 4 56-73(63.5) 1 67
Length of anal lobe

Atlantic 3 33(33.0) 1 30

Pacific 4 32-39(35.3) 1 31
Height of anal fin

Atlantic 3 28-33(30.0) 1 33

Pacific 4 34-40(36.5) 1 40
Length of pectoral base

Atlantic 3 53-58(55.7) 1 54

Pacific 4 49-54(51.8) 1 58
Length of pelvic base

Atlantic 3 56-59(57.3) 1 60

Pacific 4 53-56(54.8) 1 56
Length of caudal fin

Atlantic 3 310-335(825.3) 1 309

Pacific 4 307-320(313.0) 1 289

NO. 3539 HAMMERHEAD SHARKS—GILBERT

81

TaBLE 4.—Measurements of Sphyrna zygaena (proportions expressed in thou-
sandths of total length)

Size range in mm. (N=number of specimens)

N 400-599 N 600-799 N >800
Head width
Atlantic 8 261-290(276.9) 3 262-276(268.7) 0 —
Pacific 10 261-285(273.38) 2 276-279(277.5) 1 260
Internarial distance
Atlantic 8 194-209(201.5) 3 189-198(193.3) Qo —
Pacific 10 190-207(197.38) 2 191-203(197.0) 1 186
Snout to symphysis
Atlantic 8 62-77(70.6) 3 67-70(68.3) Qo —
Pacific 10 65-73(69.6) 2 64-65(64.5) a5
Head length
Atlantic 8 219-245(230.8) 3 218-226(221.3) Oo —
Pacific 10 225-242(231.9) 2 219-227(223. 0) ee 20
Snout to 1st gill slit
Atlantic 8 170-192(178.3) 3 168-180(173.3) 0 —
Pacific 10 172-189(180.4) 2 167-171(169.0) 1 162
Snout to 1st dorsal
origin
Atlantic 8 264-291(275.6) 3 267-270(269.0) 0 —
Pacific 10 266-288(278.5) 2 263-274(268.5) i Wigs}
Snout to 2nd dorsal
origin
Atlantic 8 589-626(606.4) 3 595-602(599.0) QO —
Pacific 10 592-626(605.3) 2 602-614(608.0) 1 635
Distance between 1st
and 2nd dorsal
bases
Atlantic 8 228-245(236.1) 3 238-251(243.0) Oo —
Pacific 10 221-244(236.1) 2 241-253(247.0) 1 —
Snout to pectoral in-
sertion
Atlantic 8 207-224(214.5) 3 198-207(203.3) Oo —
Pacific 10 209-226(217.1) 2 195-202(198.5) 1 198
Snout to pelvic in-
sertion
Atlantic 8 440-468(450.3) 3 438-446(443.3) 0 —
Pacific 10 440-458(451.38) 2 439-443(441.0) 1 454
Horizontal diameter
of orbit
Atlantic 8 23-30(25.8) 3 23-25(24.0) Oo —
Pacific 10 24-31 (28.0) 2 24(24.0) ly
Length of 1st dorsal
base
Atlantic 8 90-105(95.8) 3 91-98(95.0) Oo —
Pacific 10 86-105(95.8) 2 93-100(96.5) te O2
Length of 2nd dorsal
base
Atlantic 8 28-35(381.9) 3 30-33(381.3) Oo —
Pacific 10 29-38(33.3) 2 28-38(383.7) maces

219-937—67——6

82 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

TABLE 4.—Measurements of Sphyrna zygaena (proportions expressed in thou-
sandths of total length) —Continued

Size range in mm. (N=number of specimens)
N 400-599 N 600-799 N »>s800

Length of 2nd dorsal

lobe
Atlantic 8 43-52 (47.0) 3 44-47(45.7) 0 —
Pacific 10 43-48(45.1) 2 46-49(47.5)
Height of 2nd dorsal
fin
Atlantic 8 16-22(18.9) 3 18-21(19.7) 0 —
Pacific 10 16-23(19.5) 2 21-23(22.0) 1 —
Length of anal base
Atlantic 8 438-57 (47.3) 3 42-47(45.0) 0 —
Pacific 10 44-52(46.9) 2 42-44(43.0) 1 —
Length of anal lobe
Atlantic 8 39-44(42.1) 3 42-44(43.3) 0 —
Pacific 10 38-43(41.1) 2 43-45(44.0) 1 —
Height of anal fin
Atlantic 8 24-35(27.6) 3 26-29(27.7) 0 —
Pacific 10 24-30(27.0) 2 26-27(26.5) 1 29
Length of pectoral
base
Atlantic 8 47-58(53.4) 3 50-57(54.0) 0 —
Pacific 10 49-56(52.8) 2 53-60(56.5) 1 —
Length of pelvic base
Atlantic 8 40-46(43.1) 3 39-45(42.0) 0 —
Pacific 10 41-47(44.3) 2 40-41 (40.5) 1 —
Length of caudal fin
Atlantic 8 293-311(303.5) 3 290-301 (294.7) 0 —
Pacific 10 295-312(303.1) 2 298(298.0) 1 269

TaBLE 5.—Measurements of Sphyrna lewini (proportions expressed in thousandths
of total length)

Size range in mm. (N=number of specimens)

N 300-399 N 400-599 N 600-799 N 800-900 N >1,000
Head width
Atlantic 2 280-284(282) 16 268-302(287.0) 6 258-285(274.0) 1 267 1 240
Eastern Pacific == SS 8 260-291(276.1) 4 256-277(267.5) — — _-—

Western Pacific and 2 266-285(275.5) 12 246-287(264.8) 3 261-280(269.0) — — -—-
Indo-Australia

Internarial distance

Atlantic 2 205-213(209.0) 16 190-218(203.9) 6 193-212(201.7) 1 181 1 175
Eastern Pacific — 8 187-211(200.3) 4 184-202(195.8) _ _
Western Pacific and 2 197-204(200.5) 12 180-214(193.3) 3 186-205(197.3) = ed
Indo-Australia

Snout to symphysis

Atlantic 2 81-86(83.5) 16 72-83(79.1) 6 67-77(71.3) 1 64 1 46
Eastern Pacific — — 8 73-81(77.8) 4 68-73(70.3) So as
Western Pacific and 2 83-85(84) 12 69-83(73.9) 3 66-70(67.3) — — 1 41

Indo-Australia

NO. 3539

HAMMERHEAD SHARKS—GILBERT

83

TaBLE 5.— Measurements of Sphyrna lewini (proportions expressed in thousandths
of total length) —Continued

Head length
Atlantic
Eastern Pacific
Western Pacific and
Indo-Australia
Snout to 1st gill slit
Atlantic
Eastern Pacific
Western Pacific and
Indo-Australia
Snout to 1st dorsal
origin
Atlantic
Eastern Pacific
Western Pacific and
Indo-Australia
Snout to 2nd dorsal
origin
Atlantic
Eastern Pacific
Western Pacific and
Indo-Australia
Between Ist and 2nd
dorsal bases
Atlantic
Eastern Pacific
Western Pacific and
Indo-Australia
Snout to pectoral inser-
tion
Atlantic
Eastern Pacific
Western Pacific and
Indo-Australia
Snout to pelvic inser-
tion
Atlantic
Eastern Pacific
Western Pacific and
Indo-Australia
Horizontal diameter of
orbit
Atlantic
Eastern Pacific
Western Pacific and
Indo-Australia
Length of 1st dorsal
base
Atlantic
Eastern Pacific
Western Pacific and
Indo-Australia
Length of 2nd dorsal
base
Atlantic
Eastern Pacific
Western Pacific and
Indo-Australia

N 300-399 N 400-599 N 600-799 N 800-900 N >1,000
2 253-265(259.0) 16 220-256(241.0) 6 217-238(227.7) 1 210 1 197
——— 8 221-247(233.7) 4 216-237(225.0) — — ———
2 244-273(258.5) 12 213-250(233.0) 3 203-226(215.3) — — =
2 198-200(199.0) 16 177-196(189.6) 6 166-192(177.3) 1 168 1 162
= 8 174-200(188.6) 4 160-184(173.8) — =
2 194-206(200.0) 12 169-196(181.8) 3 165-179(170.0) _ =
2 282-307(204.5) 16 261-298(285.7) 6 268-285(277.1) 1 259 1 230
a 8 279-303(291.8) 4 263-286(273.4) — — ae
2 294-301(297.5) 12 276-312(204.4) 3 254-271(263.7) — — 1 282
2 606-618(612.0) 16 587-625(603.4) 6 572-600(584.7) 1 575 1 551
— 8 591-637(612.9) 4 579-606(591.3) — — — =
2 593-622(607.5) 12 598-657(615.3) 3 591-602(595.0) — — ——
2 198-207(202.5) 16 199-224(210.0) 6 222-242(232.7) 1 233 1 227
a 8 196-222(210.4) 4 208-240(228.3) — — —_—
2 195-219(207.0) 12 194-244(213.9) 3 219-236(226.0) — — 1 281
2 230-235(232.5) 16 202-242(225.8) 6 199-221(211.8) 1 200 Atle
— 8 216-233(222.3) 4 266-232(215.3) — — —
2 225-253(239.0) 12 200-231(217.2) 3 206-215(210.3) — — 1 196
2 460-464(462.0) 16 435-472(450.1) 6 427-456(441.7) 1 422 1 390
—— 8 429-462(446.0) 4 431-459(442.0) — — — =
2 456-479(467.5) 12 404-475(444.3) 3 430-451(445.3) — — 1 464
2 26(26) 16 21-29(23.8) 6 19+22(20.8) 1 19 1 14
—— 8 21-26(23.5) 4 21-23(21.8) oo ———
2 27-28(27.5) 12 20-26(23.4) 3 21(21) — WEL
2 105-111(108.0) 16 93-113(107.7) 6 94-106(101.0) 1 95 1 94
—_ — 8 99-110(104.2) 4 92-102(98.5) — — —-—
2 97-109(163.0) 12 92-110(104.3) 3 102-110(106.3) — — 1 91
2 35-37(36.0) 16 34-42(38.6) 6 32-40(36.3) 1 35 1 35
--— 8 32-41(36.4) 4 31-38(34.8) ee ee
2 34-38(36.0) 12 28-38(34.0) 3 33-44(37.0) —_-— 1 23

Size range in mm. (N=number of specimens)

84 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

TaBLE 5.—Measurements of Sphyrna lewini (proportions expressed in thousandths
of total length) —Continued

Size range in mm. (N=number of specimens)

N_ 300-399 N = 400-599 N 600-799 N 800-900 N>1,000
Length of 2nd dorsal
lobe
Atlantic 2 49-50(49.5) 16 43-53(49.0) 6 44-51(48.2) 1 48 1 48
Eastern Pacific - — 8 45-52(48.9) 4 47-50(48.5) ==> -_--
Western Pacific and 2 45-51(48.0) 12 44-56(49.6) 3 44-52(48.7) _—- = _- —
Indo-Australia
Height of 2nd dorsal
fin
Atlantic 2 22(22.0) 16 19-24(21.8) 6 17-23(20.7) 15822 1 23
Eastern Pacific = = 8 18-23(21.0) 4 18-22(20.8) a a
Western Pacific and 2 21-23(22.0) 12 16-22(20.2) 3 20-21(20.3) _- = 1 23
Indo-Australia
Length of anal base
Atlantic 2 63-56(54.5) 16 48-60(53.8) 6 47-55(51.5) 1 47 ibe
Eastern Pacific -- 8 48-59(51.9) 4 45-51(49.0) — -
Western Pacific and 2 51-64(52.5) 12 43-54(50.2) 3 48-64(54.7) -_-- 1 40
Indo-Australia
Length of anal lobe
Atlantic 2 40(40.0) 16 33-46(38.9) 6 37-45(40.3) 1 40 1 40
Eastern Pacific -- 8 35-43(38.9) 4 37-41(38.8) _-_— _- =
Western Pacific and 2 38-47(42.5) 12 34~-44(40.2) 3 38-39(38.3) a -_- =
Indo-Australia
Height of anal fin
Atlantic 2 28-30(29.0) 16 28-34(30.5) 6 27-33(29.2) 28: 1 29
Eastern Pacific -_--— 8 26-31(28.9) 4 27-34(29.8) —_- — ==
Western Pacific and 2 26-29(27.5) 12 24-32(28.3) 3 29-82(30.0) _- 1 25
Indo-Australia
Length of pectoral base
Atlantic 2 59-61(60.0) 16 50-61(54.6) 6 48-60(53.0) 1 51 1 50
Eastern Pacific -- 8 48-57(52.1) 4 49-55(53.0) = = ==
Western Pacific and 2 61(61.0) 12 48-57(51.8) 3 51-59(55.0) a =
Indo-Australia
Length of pelvic base
Atlantic 2 53-54(53.5) 16 49-59(54.2) 6 49-54(51.5) 1 50 1 50
Eastern Pacific —_— 8 49-58(53.9) 4 47-54(51.5) ls = =
Western Pacific and 2 61-54(52.5) 12 60-60(53.9) 3 48-55(52.0) calle ed
Indo-Australia
Length of caudal fin
Atlantic 2 305-314(809.5) 16 291-332(316.4) 6 287-318(302.7) 1 295 1 284
Eastern Pacific _ 8 300-325(313.9) 4 294-327(308.0) ae =

Western Pacific and 2 308-317(812.5) 12 296-323(3813.3) 3 295-308(302.0) — a
Indo-Australia

TaBLE 6.—Measurements of subspecies of Sphyrna tiburo (proportions expressed in
thousandths of total length)

Size range in mm. (N=number of specimens, A=Atlantic, P= Pacific)
N 200-399 N 400-599 N 600-799 N >800

Head width 8

tiburo (A) 3 186-229(206.0) 14 178-204(188.6) 12 160-192(174.1) 7 143-189(168.0)
vespertina (P) 215-245(227.7) 8 194-213(202. 4) 5 185-208(193.0) 4 175-188(180. 3)
Internarial distance

tiburo (A) 8 137-156(144.9) 14 121-148(131.7) 12 114-129(119.7) 7 100-132(118. 4)
vespertina (P) 3 153-160(156.0) 8 140-155(146. 1) 6 125-144(133.2) 4 122-140(130. 8)
Snout to symphysis

tiburo (A) 8 94-99(97.0) 14 78-90(83. 9) 12 72-80(76.0) 7 73-82(75.8)

oo

vespertina (P) 93-96 (94. 5) 8 82-91(86.0) 5 75-82(79. 0) 4 73-81(77. 0)

TaBLB 6.—Measurements of subspecies of Sphyrno tiburo (proportions expressed

Head length
trburo (A)
vespertina (P)
Snout to 1st gill slit
tiburo (A)
vespertina (P)
Snout to 1st dorsal
origin
taburo (A)
vespertina (P)
Snout to 2nd dorsal
origin
tiburo (A)
vespertina (P)
Distance between 1st
and 2d dorsal bases
tiburo (A)
vespertina (P)
Snout to pectoral in-
sertion
tiburo (A)
vespertina (P)
Snout to pelvic inser-
tion
tiburo (A)
vespertina (P)
Horizontal diameter of
orbit
tiburo (A)
vespertina (P)
Length of 1st dorsal
base
tiburo (A)
vespertina (P)
Length of 2nd dorsal
base
tiburo (A)
vespertina (P)
Length of 2nd dorsal
lobe
tiburo (A)
oespertina (P)
Height of 2nd dorsal fin
tiburo (A)
vespertina (P)
Length of anal base
tiburo (A)
vespertina (P)
Length of anal lobe
tiburo (A)
vespertina (P)
Height of anal fin
tiburo (A)
vespertina (P)
Length of pectoral base
tiburo (A)
vespertina (P)
Length of pelvic base
tiburo (A)
vespertina (P)
Length of caudal fin
tiburo (A)
vespertina (P)

N

oo

© GO

Se)

ew oO

© CO

@ CO

oo

w oO

@ oO

oo

Cw

@ oo

© CO

ww oO

ew ao

2 OO

in thousandths of total length)—Continued

Size range in mm. (N=number of specimens, A=Atlantic, P= Pacific)

200-399

234-249 (242. 8)
238-246(242. 0)

186-204(195. 4)
190-196 (193. 3)

295-331 (309. 9)
297-316(307. 7)

576-611(591. 8)
587-600 (594. 3)

181-202(188. 4)
188-191(189. 7)

220-243 (233. 3)
224-240(231. 0)

421-446 (431. 3)
427-448 (436. 7)

21-23(22. 3)
21-24(22. 3)

85-97 (90. 1)
88-98(92. 0)

46-52(48. 8)
47-51 (49. 0)

38-49(44. 1)
43-47 (45. 0)

24-41 (33. 5)
26-37 (32. 0)

70-84(75. 5)
74-83(78. 3)

33-39(35. 5)
31-38(34. 7)

30-38 (34. 0)
35-38(36. 3)

49-63 (56. 3)
58-60 (59. 3)

52-63(57. 5)
52-60(55. 7)

281-307 (292. 0)
288-295 (292. 3)

N

14
8

400-599

216-244 (228. 7)
222-240 (231. 9)

173-196(183. 3)
178-194(187. 8)

287-326(303. 9)
283-312 (298. 9)

588-625 (601. 5)
556-619(598. 1)

192-219(207. 1)
189-212(202. 6)

263-230(213. 8)
200-235 (218. 2)

430-450 (438. 2)
428-470(445. 3)

19-23(20. 1)
18-23(20. 4)

90-106 (96. 1)
77-102(93. 3)

42-58(50. 8)
42-55 (48. 2)

39-51 (46. 6)
43-51 (47. 0)

31-41 (35. 8)
32-40(35. 4)

65-86(73. 4)
69-88(77. 6)

32-39(33. 6)
35-39 (37. 3)

28-35 (32. 3)
29-37 (33. 4)

53-65 (59. 6)
56-64(60. 4)

55-64(59. 4)
56-70(61. 0)

266-294 (280. 9)
278-294 (287. 5)

N

12

12

or

_
or bo

or bo

12
5

600-799

205-223(214. 9)
208-215 (214. 3)

159-185 (170. 8)
162-180(173. 0)

268-319 (295. 4)
279-333 (302. 3)

595-634 (610. 3)
585-628(608. 6)

199-233(218. 7)
204-229(212. 8)

188-222(201. 3)
195-215 (205. 1)

426-456 (444. 0)
412-446 (427. 4)

18-20(18. 8)
16-20(18. 2)

93-110(100. 4)
86-101 (92. 4)

44-57 (50. 8)
42-52(47. 2)

43-53(48. 6)
47-53(49. 8)

35~43(39. 3)
34-43(38. 2)

64-89(76. 3)
69-88 (79. 0)

27-38(34. 8)
33-39(36. 4)

30-38(33. 0)
33-38(35. 0)

51-69(59. 3)
54-60(56. 3)

58-67 (62. 3)
55-65 (58. 8)

250-291 (270. 1)
269-296 (280. 6)

N

aa

~I

3

-_

>800

201-227 (212. 3)
200-215 (208. 8)

153-174 (163. 7)
157-169 (162. 5)

277-310 (298. 6)
280-300 (290. 8)

605-634 (614. 2)
600-636 (615. 0)

214-253 (235. 0)
214-242 (230. 3)

194-214 (203. 8)
185-220(197. 3)

385-471 (442. 3)
435-455 (443. 8)

15-17(16. 0)
14-18(16, 3)

92-107 (98. 7)
98-104(166. 3)

41-57(47. 8)
45-51 (48. 6)

45-55 (50. 0)
48-56(52. 7)

35-41 (37. 5)
38-44(40. 3)

71-83 (79. 8)
73-85 (76. 8)

25-37 (32. 8)
30-37 (34. 3)

30-36 (32. 0)
26-36 (32. 3)

57-66(60. 3)
58-67 (62. 5)

60-86 (62. 3)
58-64(61. 5)

234-276 (258. 9)
249-270(255. 8)

TaBLE 7.—Measurements of Sphyrna media (proportions expressed in thousandths
of total length)

Size range in mm.

200-399 400-599 600-799
Number of specimens
7 3 2
Head width 231-333 (289.3) 223-264 (247.3) 232-236 (234.0)
Internarial distance 182-204 (194.0) 162-191 (177.0) 166-171(168.5)
Snout to symphysis 100-107 (103.7) 88-97 (91.3) 80-84 (82.0)
Mouth width 67-76(71.7) 61-71(65.7) 62-65 (63.5)
Head length 251-264 (256.7) 222-257 (242.7) 218
Snout to Ist gill slit 204-216(210.7) 183-200 (192.7) 171-179(175.0)
Snout to 1st dorsal origin 301-321(306.7) 267-322(302.0) 285-300(292.5)
Snout to 2nd dorsal origin 592-619(605.5) 575-628 (604.7) 598-618 (608.0)
Distance between Ist and 2nd 182-196 (192.0) 202-216 (207.0) 213
dorsal bases

Snout to pectoral insertion 231-251 (245.3) 206-241 (222.3) 211-226 (218.5)
Snout to pelvic Insertion 421-439(432.0) 418-446 (429.3) 421-437 (429.0)
Horizontal diameter of orbit 19-25 (20.9) 18-21(19.3) 16-17 (16.5)
Length of 1st dorsal base 97-111(103.8) 90-99 (93.7) 90-105 (97.5)
Length of 2nd dorsal base 43-52(48.0) 44-46 (45.0) 45-50(47.5)
Length of 2nd dorsal lobe 45-49 (47.1) 48-51(49.7) 51-54(102.5)
Height of 2nd dorsal fin 30-38 (34.3) 30-37 (33.0) 35-39 (37.0)
Length of anal base 72-83(80.3) 73-84(80.0) 86-90(88.0)
Length of anal lobe 36-38 (37.3) 39 31-38 (34.5)
Height of anal fin 39-47 (42.5) 38-43 (40.7) 39-46 (42.5)
Length of pectoral base 50-57 (53.8) 53-57 (54.3) 52-56 (54.0)
Length of pelvic base 54-62(59.2) 60-65 (62.3) 48-61 (54.5)
Length of caudal fin 279-297 (289.3) 290-310 (298.0) 287-297 (292.0)

>800

2

228-232 (230.0)
167-169(168.0)
76-81(78.5)

62-72(67.0)

211-238 (224.5)
176-215(195.5)
281-306 (293.5)
615-645 (630.0)
227-233 (230.0)

195-210 (202.5)
387-392 (389.5)
14-15(14.5)
106-107 (106.5)
44-49(46.5)
55-63 (59.0)

39

84-85 (84.5)
39-48 (43.5)

41

59-62 (60.5)
66-67 (66. 5)
289-306 (297.5)

TABLE 8.—Measurements of Sphyrna corona (proportions expressed in thousandths

Head width

Internarial distance

Snout to symphysis

Mouth width

Head length

Snout to 1st gill slit

Snout to 1st dorsal origin

Snout to 2nd dorsal origin

Distance between Ist and 2nd
dorsal bases

Snout to pectoral insertion

Snout to pelvic insertion

Horizontal diameter of orbit

Length of 1st dorsal base

Length of 2nd dorsal base

Length of 2nd dorsal lobe

Height of 2nd dorsal fin

Length of anal base

Length of anal lobe

Height of anal fin

Length of pectoral base

Length of pelvic base

Length of caudal fin

of total length)

200-399

Size range in mm.

400-599

Number of specimens

12

259-291 (275.4)
181-206(192.3)
110-127(117.1)
59-69(65.6)

250-281 (270.3)
209-230(218.9)
315-345 (328.6)
595-640(613.9)
174-202(188.1)

253-277 (262.3)
436-465(450.1)
17-22(18.8)
89-104(97.4)
39-47 (43.6)
40-52(45.1)
28-33(30.4)
82-92(87.5)
33-43(37.8)
32-41 (36.8)
52-61 (55.6)
48-60(53.1)
268-290(279.3)

2

261-262 (261.5)
203-210(182.0)
107—109(108.0)
60-63 (61.5)

251-261 (256.0)
203-210(206.5)
309-318(313.5)
588-617 (602.5)
185-205(195.0)

238-263(250.5)
428-447 (437.5)
17-18(17.5)

94

43-49(46.0)
46-49 (47.5)
30-34(32.0)
87-89(88.0)
40-41 (40.5)
40-43(41.5)
55-59(57.0)
53-59(56.0)
284-288(286.0)

NO. 3539

HAMMERHEAD SHARKS—GILBERT 87

TABLE 9.—Measurements of Sphyrna tudes (proportions expressed in thousandths

of total length)

Head width
Western Atlantic
Mediterranean

Internarial distance
Western Atlantic
Mediterranean

Snout to symphysis
Western Atlantic
Mediterranean

Mouth width
Western Atlantic
Mediterranean

Head length
Western Atlantic
Mediterranean

Snout to 1st gill slit
Western Atlantic
Mediterranean

Snout to 1st dorsal origin
Western Atlantic
Mediterranean

Snout to 2nd dorsal origin
Western Atlantic
Mediterranean

Distance between Ist and 2nd

dorsal bases

Western Atlantic
Mediterranean

Snout to pectoral insertion
Western Atlantic
Mediterranean

Snout to pelvic insertion
Western Atlantic
Mediterranean

Horizontal diameter of orbit
Western Atlantic
Mediterranean

Length of 1st dorsal base
Western Atlantic
Mediterranean

Length of 2nd dorsal base
Western Atlantic
Mediterranean

Length of 2nd dorsal lobe
Western Atlantic
Mediterranean

Height of 2nd dorsal fin
Western Atlantic
Mediterranean

Length of anal base
Western Atlantic
Mediterranean

Length of anal lobe
Western Atlantic
Mediterranean

Height of anal fin
Western Atlantic
Mediterranean

Size range in mm. (N=number of specimens)

N 200-399 N 400-599 N 600-799 N >800
13 299-319(310.0) 1 304 276-285(280.5) 1 280
1 321 o— — —
13 221-234(228.5) 1 219 202-209(205.5) 1 206
1 231 o— — =
13 94-117(105.1) 1 92 80(80.0) 1 72
1 104 o— — i
13 61-69(64.7) 1 68 65~69(67.0) 1 69
1 69 = — Oo —
13 229-263(242.2) 1 233 224-226 (225.0) 1 227
1 243 o— _ i
13 190-211(200.5) 1 193 186-189 (187.5) 1 184
1 197 OV _ =
13 293-334(309.8) 1 305 299-306(302.5) 1 290
1 319 0 — — 0 —
13 580-625(597.0) 1 606 601-619(610.0) 1 622
1 243 o— _ 0
13 178-209(190.7) 1 203 199-209(204.0) 1 224
1 182 o— — =
13 215-251(231.1) 1 225 227-228 (227.5) 1 212
1 246 o— — o—
13 397-427(413.5) 1 412 405-428 (416.5) 1 393
1 434 o— — 1
13 15-23(18.0) 1 15 15(15.0) 1 18
1 20 o— — 0 —
13 89-109(98.0) 1 108 103-104(103.5) 1 108
1 95 0 — — 0 —
13 41-51(44.2) 1 45 41-44 (42.5) 1 48
1 45 0 — — on
13 41-47(44.0) 1 47 53-58(55.5) 1 56
1 42 Ca = i.
13 29-33(31.0) 1 29 33-36 (34.5) 1 39
1 35 o— _ =
13 75-99(84.6) 1 83 83-93(88.0) 1 84
1 84 a — — i
13 28-33(32.1) Tesy 37-42(39.5) 1 34
1 35 o— == o—
13 36-47(40.4) 1 46 47(47.0) 1 47
1 61 o— — o—

88 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

TaBLE 9.—Measurements of Sphyrna tudes (proportions expressed in thousandths
of total length) —Continued

Size range in mm. (N=number of specimens)

N 200-399 N 400-599 WN 600-799 N >s800
Length of pectoral base
Western Atlantic 13 53-58(55.2) e167: 2 60-62(61.0) 1 59
Mediterranean 1 55 0o— 0o— 0o—
Length of pelvic base
Western Atlantic 13 50-60(55.0) 1 54 2 64-67(65.5) 1 73
Mediterranean 1 59 o— 0— o—
Length of caudal fin
Western Atlantic 13 290-313(302.4) 1 295 2 293-297(295.0) 1 286
Mediterranean 1 292 0o— o— o-—

TasLE 10.—Measurements of heads of Sphyrna lewini and S. couardi from Gulf

of Guinea }
Nares to
posterior
Snout to margin of Inner narial
Head width symphysis head 2 Mouth gape groove
S. lewint 445. 0 93. 5 93. 5 103. 5 83. 0
proportion _— . 210 . 210 . 233 . 187
S. couardt 425. 0 115. 0 eto LOSS, 89. 0
proportion — . 270 . 276 . 281 . 209

1 One specimen from each species. Under each species the first row represents the empirical lengths (in
millimeters); the second row, the proportional values (in thousandths of head width), obtained by dividing
head width into the respective measurements.

3 Shortest vertical distance.

GILBERT—PLATE 1

119

MUS. VOL.

U.S. NAT.

PROC.

“(ZIT ‘Id ‘T ‘JOA “SQ/T) Yoorg wor ‘(puavshz snjvnbs se

eel ge
: Pe poe, > Pix
° YP yfsrUnr? ) 7 4? Fe
~ — WwW
“VWNAVOAZ SrvIVaAeYy

LWT

ee *7/-
o

peyuaptsiur) 114 9074 DUIDSAT jo uoheiysnyyt [eUuIsLIO

GIEBERT—PEAGE 2

119

PROC. U.S. NAT. MUS. VOL.

GILBERT—PLATE 3

119

NAT. MUS. VOL.

PROC. U.S.

‘(os ‘14 ‘OF9 “d ‘FegT) yrTWS

pur “WIS

‘IOIAND us “YW pur YIN woz ‘wim DuevshZ JO uonrisn|! [euIsC

MmIMaT vial As

GILBERT—PLATE 4

119

PROC. U.S. NAT. MUS. VOL.

L Qe
Mg
y
i j ‘ei!
Ws taal
|

Mish il cy iit i
Adal ‘i

t

la, b).

12, figs.

—

Original illustration of Zygaena tudes, from Valenciennes (1822, p

GILBERT—PLATE 5

119

PROC. U.S. NAT. MUS. VOL.

(95981 HNINA) Parl

wolf 5

ele “WU OZLF usulioods e “114 9014 puacyds fo WUNtUeISOIpuoys fo ydvisoipey

PROG: UTS: NAT. MUS. VOE- 119 GILBERT—PLATE 6

Radiographs of chondrocrania of several species of Sphyrna: A, S. mokarran, 560 mm. TL,
from Florida (USNM 186087); B, S. lewini, 447 mm. TL, from Liberia (USNM 179706);
C, S. zygaena, 590 mm. TL, from Italy (USNM 28452).

PROC. U.S. NAT. MUS. VOL. 119 GILBERT—PLATE 7

Radiographs of chondrocrania of several species of Sphyrna: A, S. tiburo tiburo, 388 mm
TL, from Nicaragua (USNM 78180; B, S. tiburo vespertina, 545 mm. TL, from Mexico
(UCLA 51-4); C, S. media, 337 mm. TL, from Mexico (FMNH 63093); D, S. corona,
371 mm. TL, from Mexico (FMNH 72521, ex. 63093).

PROC. U.S. NAT. MUS. VOL. 119 GILBERT—PLATE 8

Radiographs of chondrocrania of Sphyrna tudes: A, a specimen 390 mm. TL (holotype of

S. bigelow1) from Uruguay (USNM 87682); B, a specimen 346 mm. TL (lectotype
S. tudes) from France (MNHN 1049).

of

— i

PROC. U.S. NAT. MUS. VOL. 119 GILBERT—PLATE 9

Radiographs of chondrocrania: A, S. couardi, head 425 mm. wide, from Guinea (USNM
uncat.); B, S. lewint, head 445 mm. wide, from Guinea (USNM uncat.).

PROC. U.S. NAT. MUS. VOL. 119 GILBERT—PLATE 10

Photographs of heads: A, S. couardi, head 425 mm. wide, from Guinea (USNM uncat.); B
S. lewini, head 445 mm. wide, from Guinea (USNM uncat.).

Proceedings of

the United States

National Museum

SMITHSONIAN INSTITUTION + WASHINGTON, D.C.

Volume 119 1966 Number 3540

NEOTROPICAL MICROLEPIDOPTERA, X!

SYSTEMATIC POSITION OF TWO TAXA
ERRONEOUSLY PLACED IN THE FAMILY STENOMIDAE
(LEPIDOPTERA)

By W. Donatp DuckwortTH
Associate Curator, Division of Lepidoptera

Revisionary studies presently being conducted on the Stenomidae
have revealed two incorrectly placed taxa. One, a monobasic genus,
is transferred to the Tortricidae; the other represents a new genus and
is transferred to the Oecophoridae.

The author wishes to acknowledge with thanks the cooperation and
aid of Mr. P. E. S. Whalley of the British Museum (Natural History)
and Dr. Fritz Kasy of the Natural History Museum, Vienna, Austria
for allowing him to study the type specimens in their charge.

The author also wishes to thank Andre Pizzini for the line draw-
ings in this paper and Jack Scott for the photographic work.

The work on this paper was aided by the National Science Foun-
dation Grant GB-1800.

1 Prepared with the aid of a National Science Foundation Grant. Previous
parts of this same series are: I and II, Clarke, 1962, Proc. U.S. Nat. Mus., vol.
113, no. 3457, pp. 373-388; III, Clarke, 1964, ibid., vol. 115, no. 3480, pp. 61-84;
IV, Duckworth, 1964, ibid., vol. 116, no. 3497, pp. 97-114; V, Obraztsov, 1964,
ibid., vol. 116, no. 3501, pp. 183-196; VI, Clarke, 1964, ibid., vol. 116, no. 3502,
pp. 197-204; VII, Obraztsov, 1966, ibid., vol. 118, no. 3727, pp. 221-232; VIII,
Duckworth, 1966, ibid., vol. 118, no. 3531, pp. 391-404; IX, Obraztsov, 1966,
ibid., vol. 118, no. 3535, pp. 577-622.

1

2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Family Oecophoridae
Mattea, new genus

Type species: Cryptolechia phoenissa Butler, 1883, Trans. Ent. Soe.
London, 1883, p. 81; pl. 11, figs. 12, 12a.

Head with appressed scales; lateral tufts spreading; antenna simple
in female, finely ciliated ventrally in male; basal segment with pecten;
labial palpus recurved, just reaching base of antenna. Forewing with
costa slightly arched, termen oblique, tornus rounded; 11 veins; 2 well
before angle; 7 and 8 coincident to termen; remainder separate.
Hindwing as broad as forewing; 8 veins; 3 and 4 connate; 6 and 7
subparallel.

Male genitalia: Uncus present, well developed. Gnathos a lightly
sclerotized, simple band. Anellus a simple plate. Harpe with
clasper. Aedeagus with inception of ductus ejaculatorius near
midpoint.

Female genitalia: Genital plate reduced to a thin, sclerotized band.
Bursa copulatrix reduced, signum absent.

Remarks: The exact position of this genus in the family Oecopho-
ridae is virtually impossible to ascertain until a revision of the Neo-
tropical species is undertaken. In addition, careful attention to the
Australian and New Zealand fauna will be necessary since the distri-
butional relationships between these areas and the west coast of
South America are poorly understood.

The genus is distinguished by veins 7 and 8 of the forewing being
coincident to the termen. Coincidence of veins 7 and 8 is not un-
common in the family, but the termination in other genera is on the
costa. The genitalia present a number of striking characters, such
as the inception of the ductus ejaculatorius being near the midpoint
of the aedeagus and the presence of a reduced bursa copulatrix in the
females. However, the genital characters of the Neotropical genera
are still poorly known, and until they are studied no definite conclu-
sions can be drawn.

Mattea phoenissa (Butler), new combination
FIGuRE 1; PLATE 1 (Fia. a)
Cryptolechia phoenissa Butler, 1883, Trans. Ent. Soc. London, 1883, p. 81, pl. 11,
figs. 12, 12a.
The original description of this species is quite accurate and reads
as follows: ‘Wings sericeous carmine-red; primaries above with a

a small triangular golden spot at outer third of dorsal margin; fringe
at apex tipped with bright ochreous; thorax purplish brown; antennae

NO. 3540 MICROLEPIDOPTERA, X—DUCKWORTH 3

carmine; palpi whitish; abdomen dark brown; wings below of a clearer
carmine-red than above; base of secondaries and pectus bronze-
brown; legs and venter whity-brown. Expanse of wings, 16 mm.

In addition to the characters given in the original description, the
details of the wing venation, head, and male and female genitalia are
illustrated in figures 1a-e.

Type: In the British Museum (Natural History).

Type locality: Corral, Chile.

Distribution: Chile: Corral; Llanquihue, Peulla (March); Centro-
Austral (Jan.—Mar.).

Remarks: This species was described originally in the family
Gelechiidae, and this association was followed by Calvert (1886) in
his catalog of Chilean Lepidoptera. Busck (1934) listed the species
as Stenoma phoenissa in his catalog of the family Stenomidae, and
this combination has persisted until now. Examination of the type
specimen, however, clearly indicates that the genus should be placed
in the Oecophoridae. The remote position of veins 6 and 7 in the
hindwing and the presence of pecten on the basal segment of the
antenna are both good oecophorid characters which are not found in
the Stenomidae.

Family Tortricidae
Subfamily Sparganothiinae
Rhynchophyllis Meyrick
Figure 2; PLATE 1 (FIG. b)

Rhynchophyllis Meyrick, 1932, Exotic Microlepidoptera, vol. 4, pp. 288-289.
Type species: Rhynchophyllis categorica Meyrick, by monotypy.
This monobasic genus was established for a new species, R. cate-

gorica, described from a single female specimen from Jaragua, “Santa
Catharina” (Santa Catarina), Brazil. Meyrick included the genus
in the Xyloryctidae (=Stenomidae) with the following comment:
“A highly peculiar insect; the hindwings are normal for the Xyloryc-
tidae, to which therefore it must be referred, but the rest of the
structure is very abnormal; possibly there is some relationship to
Faleulina, which is suggested by the superficial appearance, but
this may be delusive.’”’ Examination of the type specimen in the
Natural History Museum, Vienna, Austria, which remains the only
known example of this species, clearly indicates its proper placement
in the Tortricidae rather than in the Stenomidae. The correct
systematic position within the Tortricidae must await a revision of
the Neotropical members of the family. On the basis of the ribbon-
like signum, the genus is tentatively assigned to the subfamily Spar-
ganothiinae.

4 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

Literature Cited

Busck, A.
1934. Stenomidae. Lepidopterorum Catalogus, vol. 67, pp. 1-73.
CaLvERT, W. BARTLETT
1886. Catalogo de Los Lepidépteros de Chile. Anales Univ. Chile, vol.
69, pp. 1-44.

NO. 3540 MICROLEPIDOPTERA, X—DUCKWORTH o

Ficure 1.—Mattea phoenissa (Butler): a, wing venation; b, lateral view of head; c, male
genitalia; d, aedeagus; ¢, female genitalia.

6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Ficure 2.—Rhynchophyllis categorica Meyrick: a, female genitalia; b, lateral view of head.

U.S. GOVERNMENT PRINTING OFFICE:1966

PROC. U.S. NAT. MUS. VOL. 119 DUCKWORTH—PLATE 1

Ficure a, Mattea phoenissa (Butler). Ficure b, Rhynchophyllis

categorica Meyrick.

Proceedings of
the United States

National Museum

SMITHSONIAN INSTITUTION - WASHINGTON, D.C.

Volume 119 1966 Number 3541

BENTHIC AMPHIPODA
OF MONTEREY BAY, CALIFORNIA

By J. Laurens BARNARD

Associate Curator, Division of Crustacea

The fauna of benthic subintertidal Amphipoda in Monterey Bay,
California is compared to that of southern California south of Point
Conception. This study of northern Amphipoda complements a
study of southern Amphipoda of Baja California (Barnard, 1964a).
The two surveys have been conducted in order to determine the
affinities of various Amphipoda of southern California.

The extrinsic distributions of southern California Amphipoda are
poorly known except for those species from Baja California reported
by Shoemaker (1925, 1942), Barnard (1964a), and those from Oregon,
Washington, and British Columbia reported by Calman (1898),
Walker (1898), Thorsteinson (1941), Barnard (1954), Bousfield
(1958), and Mills (1961, 1962). Most of the cited papers have
treated only intertidal species. Although Monterey Bay has not
been considered to be a typical cold-temperate area, the present
analyses indicate the cold-temperate character of several California
Amphipoda.

I am indebted to Dr. R. E. Stevenson of Florida State University,
formerly of the Allan Hancock Foundation, who directed the Cali-
fornia State Water Pollution Control Board survey of California.

1

2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

G. F. Jones, aboard the RV/Velero IV, was instrumental in collecting
and processing the samples. Captain Fred C. Ziesenhenne identified
the ophiuroids dominating the bottom samples. Dr. Joel W. Hedg-
peth of Pacific Marine Station loaned specimens of a new species of
Protomedeia. The Beaudette Foundation and the Smithsonian In-
stitution supported the project and the National Science Foundation
provided funds for illustration (grant G—-10750).

Methods and Data

The benthos of Monterey Bay, in water depths of 15 to 116 m
(table 1) is represented by those 46 samples charted in figure1. They
are relatively evenly distributed in proportion to bathymetry. The
coastal benthos of southern California is represented by 348 samples
evenly proportioned to depths between 10 and 200 m. The two
groups of samples, although disparate, are roughly proportional in
number to their respective bottom areas. Any comparisons between
the Amphipoda of the two areas must be viewed with caution.

Samples of sediment and fauna were collected with an orange-
peel grab taking a surface area of 0.25 m?. Barnard and Jones (1960)
have considered the average mechanical efficiency of the grab to be
about 80 percent if large numbers of samples are taken on diverse
substrates. The efficiency decreases on hard-packed sand bottoms
and increases on soft silty bottoms. Calculations of frequencies of
Amphipoda in the 348 southern California samples employed the 80
percent efficiency rule. Those of Monterey Bay employed a 100
percent efficiency rule. All sedimentary samples from Monterey
Bay, with one exception, exceeded 1.5 m*, the mark of 100 percent
efficiency. Presumably differences in depth of penetration of the
grab in those samples of 100 percent areal efficiency would not affect
tallies of Amphipoda except for those listriellas inhabiting deep
burrows of polychaetes and echiuroids.

Sedimentary samples were washed through meshes of 0.7 mm
square openings and the residues preserved in a mixture of seawater
and formaldehyde. Amphipoda and those other animals considered
to be dominants in the samples were removed to alcohol in the labora-
tory. The remaining materials were represerved and stored in the
collections of the Allan Hancock Foundation. Thorson’s (1957)
methods were used to determine standing-crop dominance.

Planning and conducting of the surveys were based on USHO
charts contoured in English fathoms, but the bathymetric data have
been converted to meters herein. Hence the depth classifications of
benthic areas in southern California, quoted in meters, are not in
standard intervals of 10, 20, 30 m, etc., but are in converted intervals
of 10, 20, 30, etc., fms.

NO. 3541 BENTHIC AMPHIPODA—BARNARD 3

Geographic Positions and Environments

The region of southern California, with a median latitude of 33°
N, extends from Point Conception on the north to the international
border between California amd Mexico on the south. The coastal
length is approximately 300 km. Monterey Bay, with a latitude of
approximately 36.5° N, lies 275 km north of Point Conception. The
bay occurs near the southern end of the cold-temperate Oregonian
zoogeographic province and is within the Montereyan subprovince.
Monterey Bay is broad, semilunar, and about 30 km wide. Its
shelf is narrow and cut by the shoreward ends of the Monterey Canyon
complex occurring just north of the area shown in figure 1. The
benthic enviroment, except for intertidal areas, has been scarcely
explored, but several known environmental differences between
Monterey Bay and southern California are explained below.

Surface waters of Monterey Bay are dominated by the upwelling
of cool subsurface layers (California Cooperative Fishery Investi-
gations, 1952-1964). Sea-surface temperatures vary between 10°
and 14° C. The average sea-surface temperature of southern Cali-
fornia, at 33° N during 1957-1960, was about 17.5°, and the range
was 14° to 23° C (Jones and Barnard, 1963). Upwelling occurs from
place to place and is especially prominent in the Gulf of Santa Bar-
bara, where waters are entrained by the California current.

The shelf of the southern part of Monterey Bay lies in greater
water depths than do Santa Monica Bay (Hartman, 1956) and San
Pedro Bay (Barnard and Ziesenhenne, 1961) of southern California.
Very little of Monterey Bay is shallower than 37 m (20 fms). Those
southern bays have a large share of benthic area in the 27-46 m
(15-25 fms) depth interval. As a result, the benthic fauna of Mon-
terey Bay is dominated by deep-water ophiuroid communities, whereas
the southern bays are dominated by shallow-water ophiuroid com-
munities or nonophiuroid communities. The distribution of the
communities in Monterey Bay is shown in figure 1. Contours are
necessarily diagrammatic because of low sampling frequencies. The
principal ophiuroids are Amphioda urtica (Lyman), Amphipholis
squamata (Delle Chiaje), Amphiura arcystata H. L. Clark, and
Amphioplus strongyloplax (H. L. Clark). The southern bays are domi-
nated principally by the Amphioplus heracanthus H. L. Clark com-
munity (Barnard and Ziesenhenne, 1961), which occurs only on the
inshore margins of Monterey Bay. The deep-water Amphiodia urtica
is the common dominant of the outer shelf of southern California.
The prevalence of A. strongyloplaz, A. amphacantha, and A. arcystata
in depths of less than 100 m in Monterey Bay is a reflection of the
northern emergence of communities occurring in southern California

4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

only on coastal slopes in depths exceeding 120m. These communities
have not been sampled frequently in southern California because of
steeply sloping bottoms and diffuse sampling grids.

The headland of Monterey Peninsula has a rocky substrate but to
the northeast a long, slightly concave shoreline is composed of sand
beach. A community dominated by the polychaete Nothria species
occurs on inshore sand bottoms in depths of less than 30 m; presumably
it is similar to that described from southern California by Barnard
(1963).

The Amphipod Fauna

Because most of the Monterey shelf hes in depths exceeding 50 m,
the station grid embraces only 13 samples in depths of 10 to 50 m.
Those few samples have little comparative significance to samples
from southern California. Thus, this discussion is confined to the
33 samples from depths of 50 to 1389 m. The Amphipoda occurrng
in Monterey Bay are listed in rank in table 2, with their frequencies
of individuals and their limitations to depths of 50 to 139 m. They
are compared with rankings of species from similar depths in southern
California (tables 3, 4, 5, 6).

Half of the 20 most abundant species occurring in southern Cali-
fornian depths of 94-183 m (table 6) also occur in the first 20 of
Monterey Bay in depths of 50-139 m (table 2). Those southern
California species not found in the list of abundant Monterey species
are Ampelisca pacifica, Westwoodilla caecula, Ampelisca brevisimulata,
Orchomene decipiens, Nicippe tumida, Ampelisca pugetica, Lysianassa
holmesi, Paraphorus robustus, Pardisynopia synopiae, and Lysianassa
oculata (the latter is number 22 in Monterey Bay). Because all of
those species do occur in low frequency in Monterey Bay, this poor
comparison is probably a circumstance of low sampling frequency.
These data also indicate that shallow waters of Monterey Bay, de-
spite the occurrence of upwelling, are too warm for the occurrence
of deep-water species. |

A 50 percent correspondence in predominant species occurring |
both on the Monterey deep shelf and in southern California depths of |
75 to 92 m also occurs (table 5). Some deep-water species, Le.,
Nicippe tumida and Pardisynopia synopiae, occur dominantly in the |
southern California section but not in Monterey Bay.

Only 8 of the 20 predominant species occurring in depths of 57 to i
73 m also predominate in Monterey Bay. Similarly, neither Nicippe
tumida nor Urothoe varvarini are abundant in Monterey Bay (table 4). |
The same is true in depths of 39 to 55 m (table 3). |

These poor faunal congruencies may result from the low densities |
of the individual species in the Monterey samples. For instance, ©

NO. 3541 BENTHIC AMPHIPODA—BARNARD oD

the 20 species of Monterey Bay occurring most frequently (1-21,
less 3) tally 220 individuals per m?, whereas they tally 493 individuals
in southern California depths of 75-92 and 94-183 m. The fre-
quency of the 20th most abundant species in Monterey Bay is 2.4
individuals whereas the 20th of southern California is about 7.0 indi-
viduals per m?. Fifty-eight species occur on the deep shelf of Monterey but
more than 110 species occur in depths of 75 to 183 m in southern
California. Of the 110 southern California species, 12 have fre-
quencies of less than 0.2 individuals per m? and 39 have less than 1.0
individuals. This demonstrates the influence of sampling frequency.

All 58 species of the Monterey deep shelf occur in southern
California except for Centromedon pavor and Protomedeia penates. In
shallow waters, only Hohaustorius sencillus (of 23 species not found
in the deep zone) is not recorded from southern California. This
strong relationship is a result of the extreme intermixture of northern
and southern elements in the fauna of southern California. Perhaps
the intermixture results from southward range extensions of northern
species inhabiting areas of upwelling.

On the other hand, many of the Amphipoda of Monterey Bay
were not collected in the survey of Baja California (Barnard, 1964a).
Thirty-seven open-sea samples were collected there, an effort compa-
rable to that of Monterey Bay. A comparison of Monterey Bay and
Baja California probably suffers the effects of small numbers of
samples more than would a comparison between a large and a small
number. The sampling stations of Baja California were scattered
throughout 300 km of the coastal length and occurred in a variety
of environments. Approximately 113 species occur in the fauna of
Monterey Bay and Baja California, but only 61 species occur in
both areas. The remaining species are listed in groups of northern
and southern species in tables 7 and 8.

These data provide an estimate of the northern or southern affinities
of more than 100 of the 167 species of Amphipoda occurring on the
benthos of southern California. A large share of the species occurs
in all three areas. Records from the literature supply information
for some of the other Amphipoda, and a few species are assigned to
regions by determining the general distribution of their genera.
Facts and approximations are combined in a checklist of species from
southern California (table 9). Only those species with detectable
affinities are annotated with appropriate statements. The southern
California fauna is estimated to be comprised of 24 percent northern
or cold-water species, 13 percent southern or warm-water species,
5 percent bathyal affinity, and 58 percent interregional or poorly
known distribution. Northern species occur generally in deep water,
southern species in shallow water, and ubiquitous species in one or

6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

both. The percentages support the conclusion that southern Cali-
fornia lies at the northern end of the warm-temperate province but
that the deep-water shelf is inhabited primarily by submergent cold-
temperate species.

Presumably 16 species of table 8, marked with asterisks, represent
northeastern Pacific endemic boreal elements which may be found
throughout the Oregon province (in shallow depths in the northern
portion). Approximately 28 (85 percent) of the 81 known species of
Monterey Bay are of cold-temperate distribution. Purely tropical
influence is small. The remaining species are of presumed warm-tem-
perate affinities.

The intertidal zone of California north of Point Conception has
several cold-temperate species (Barnard, MS), whereas the middepths
of Monterey Bay and southern California have very few, if any, of
these elements. These facts indicate that amphipodan distributions
are strongly controlled by temperature and that a wide latitude of
ecothermic response is possible for mud-bottom species through sub-
mergence because of substrate similarities between shallow and deep
waters. The floral substrates of intertidal waters have a narrow
range of submergence owing to the absence of illumination in deep
waters. Thus phycophilous Amphipoda cannot submerge to the
extent of their mud-bottom congeners. The thermal tolerances of
the cold-temperate intertidal species must therefore be greater than
those of mud dwellers. The temporal evolution of these intertidal
distributions may be favorably controlled by the occurrence of a wide
variety of available niches and an extensive food supply.

Northwestern Pacific Relationships

Only 12 of the 81 species of Monterey Bay have been found in the
northwestern Pacific region embracing the Japan Sea, Okhotsk Sea,
and Bering Sea (table 10). Two of those species, Argissa hamatipes
and Nicippe tumida, may be cosmopolitan in cold water. They may
submerge to great depths in the tropics, the former possibly occurring
even in shallow tropical seas. Leucothoe spinicarpa is a eurybathic
cosmopolite occurring in sponges. The genus Anonyz is enormously
diverse in the northwestern Pacific region but declines in diversity by
southerly increments. Only 2 species of Anonyzx occur in California,
south of Monterey Bay. Corophium uenoi rarely occurs in the open
sea of California, but it is abundant in lagoons and estuaries (Newport
Bay, Morro Bay) and may have been introduced from Japan in
oyster transplants. The remaining 7 species of table 10, with the
exception of Ampelisca macrocephala and Paraphoxus obtusidens, are
scarce on the Monterey shelf. Probably they submerge or are de-
pleted to the south of southern California. Ampelisca macrocephala

NO. 3541 BENTHIC AMPHIPODA—BARNARD 7

is an extremely abundant deep-shelf species in California and is
replaced in bathyal depths by its anoculate subspecies, A. m. unsocalae.
Paraphozus obtusidens is a paneastern Pacific eurythermal species
ranging from the subarctic into the tropics. It scarcely submerges to
the south, but it may occur in warm latitudes primarily in areas of
upwelling.

TABLE 1.—Metric depths of stations in Monterey Bay shown in figure 1 (uncor-

rected for tidal levels).

Station Depth Station Depth Station Depth
6425 24 6444 17 6459 84
6426 54 6445 25 6460 87
6427 63 6446 52 6462 85
6428 61 6447 63 6463 87
6429 73 6448 55 6464 82
6430 78 6449 35 6465 93
6431 93 6450 34 6466 106
6432 98 6451 56 6469 109
6433 107 6452 53 6471 139
6435 116 6453 24 6474 115
6438 97 6454 15 6477 107
6439 67 6455 76 6478 101
6440 19 6456 77 6479 96
6442 24 6457 75 6480 103
6443 38 6458 80

TaBLE 2.—Frequency of 58 species of Amphipoda in Monterey Bay in depths of
50-139 m, with percent of restriction of each species to this depth zone (third
species, based on 33 samples, eliminated in all calculations described in text;
NSS = not statistically significant).

Percent of specimens

Individuals collected between Present in no.
Name of species per m2 50 and 139 m of samples

Paraphocus fatigans 73 76 18

(kind 1) (13) (39) (6)

(kind 2) (61) (94) (17)
Paraphozus bicuspidatus 24 100 17
Kermystheus ociosa NSS 14 100 4
Phoxocephalus homilis 13 100 6
Phottis lacia 12 100 8
Paraphozus similis 12 87 12
Photis californica 11 100 2
Ampelisca macrocephala 10 96 19
Heterophoxus oculatus 9.7 95 15
Byblis veleronis 9. 6 100 22
Ampelisca romigt 8.8 100 Z
Metaphoxus frequens a3 80 15
Paraphozxus obtusidens 4,4 94 10
Ampelisca compressa 4.1 100 10
Paraphozus variatus 3.9 60 4
Aoroides columbiae 3. 6 27 6
Ampelisca cristata 3. 2 70 7
Synchelidium sp. A 2.9 100 9

8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

TABLE 2.—Frequency of 58 species of Amphipoda in Monterey Bay in depths of
50-139 m, with percent of restriction of each species to this depth zone (third
species, based on 33 samples, eliminated in all calculation described in text;
NSS=not statistically significant )—Continued

Percent of specimens

Individuals collected between Present in no.
Name of species per m? 50 and 139m of samples
Anonyx adoxus 2nd 100 Ht
Urothoe varvarini 2. 6 100 4
Protomedeia articulata 2. 4 73 9
Lysianassa oculata 2.3 100 9
Hippomedon denticulatus 2.1 88 6
Synchelidium sp. G 2. 1 82 6
Ampelisca pacifica 2.0 100 6
Ampelisca pugetica 2. 0 21 7
Monoculodes emarginatus 2. 0 100 6
Paraphozus robustus 2.0 100 6
Ivstriella goleta lene 92 10
Orchomene pacifica 17, 100 2
Nicippe tumida 1.5 100 5
Pardisynopia synopiae 15 100 3
Dexamonica reduncans 1.4 100 3
Orchomene decipiens 1.4 100 4
Westwoodilla caecula b2 100 6
Ampelisca hancock tt 100 4
Photis brevipes a Bea 6 4
Opisa tridentata 0. 9 100 2
Melita desdichada 0.8 100 2
Ampelisca brevisimulata 0. 6 100 3
Metaphoxus fulton 0. 6 50 1
Paraphorus epistomus 0. 6 100 1
Ampelisca milleri 0.5 100 1
Acidostoma hancocki 0. 3 100 1
Lysianassa holmest 0. 3 100 2
Centromedon pavor 0. 3 100 1
Lepidepecreum gurjanovae 0.3 100 2
Maera danae 0.3 100 1
Pachynus barnardi 0.3 100 i!
Prachynella lodo 0. 3 100 2
Synchelidium shoemakeri 0.3 100 1
Anonyz carinatus 0. 2 100 1
Argissa hamatipes 0. 2 100 1
Microdeutopus schmitti 0. 2 0+ f
Photis macrotica 0. 2 100 1
Protomedeia penates 0. 2 100 1
Stenothoe frecanda 0. 2 100 1
Synchelidium sp. E 0. 2 100 1

Other species from Monterey Bay found in shallow water: Ampelisca lobata,
Amphilochus picadurus, Bathymedon roquedo, Corophium wuenoi, Eohaustorius
sencillus, Erichthonius brasiliensis, Eurystheus thompsont, Ischyrocerus litotes,
Leucothoe alata, Leucothoe spinicarpa, Liljeborgia kinahani, Listriella diffusa,
Maera simile, Meguluropus longimerus, Megamphopus mamolus, Metopella aporpis,
Monoculodes norvegicus, Monoculodes spinipes, Panoploea rickettst, Parapleustes
pugettensis, Photis bifurcata, Podocerus cristatus, Tiron biocellata.

NO. 3541 BENTHIC AMPHIPODA—BARNARD 9

TaBLE 3.—The most abundant species in depths of 39-55 m insouthern California.

Species Individuals Species Individuals
per m2 per m2
Ampelisca brevisimulata 68 Listriella goleta 14
Heterophoxus oculatus 58 Paraphozus obtusidens 14
Paraphoxus bicuspidatus 44 Ampelisca pactfica 14
Amphideutopus oculatus 41 Paraphozus epistomus 14
Paraphozus similis 36 Ampelisca pugetica 11
Metaphoxus frequens 33 Ampelisca coinpressa 11
Ampelisca cristata 32 Byblis veleronis 8.0
Photis brevipes 19 Lysianassa oculata Mee
Photis californica 19 Podocerus cristatus ete
Photis lacia iLvZs Paraphozrus stenodes G0

TaBLE 4.—The most abundant species in depths of 57-73 m in southern California.

Species Individuals Species Individuals
per m2 per m2
Paraphoxus bicuspidatus 134 Amphideutopus oculatus 12
Ampelisca brevisimulata 63 Ampelisca indentata 11
Heterophoxus oculatus 51 Photis lacia ib
Paraphozus similis 45 Ampelisca cristata 9. 8
Metaphorus frequens 35 Lysianassa oculata 9. 6
Ampelisca pacifica 31 Ampelisca macrocephala 9. 0
Ampelisca pugetica 21 Nicippe tumida 8.8
Byblis veleronis 15 Urothoe varvarini 8. 6
Photis brevipes 15 Paraphoxus robustus 8.3
Protomedeia articulata 13 Lysianassa holmesi 7. 4

Tas LE 5.—The most abundant species in depths of 75-92 m in southern California.

Individuals Individuals

Species per m2 Species per m2?
Paraphoxus bicuspidatus 125 Photis lacia 24
Metaphoxus frequens 35 Byblis veleronis 15
Ampelisca macrocephala 34 Lysianassa holmesi 11
Ampelisca pacifica 33 Photis californica 11
Ampelisca brevisimulata 31 Lysianassa oculata 11
Heterophorus oculatus 27 Paraphozus obtusidens 10
Ampelisca pugetica 26 Phoxocephalus homilis 10
Paraphozus similis 25 Ampelisca hancocki 8. 6
Paraphozus robustus 25 Pardisynopia synopiae 8. 6
Urothoe varvarini 25 Nicippe tumida 8.3

219-939—66——2

10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Tas LE 6.—The most abundant species in depths of 94-183 m in southern California.

Individuals Individuals

Species per m? Species per m?
Paraphozus bicuspidatus 98 Orchomene decipiens 12
Ampelisca macrocephala 84 Nicippe tumida 1d
Ampelisca romigi 45 Ampelisca pugetica 10
Heterophoxus oculatus 35 Protomedeia articulata 9. 6
Metaphorus frequens 33 Lysianassa holmesi 9. 2
Photis lacia 27 Paraphorus similis 8. 6
Ampelisca pactfica 21 Paraphozus robustus 8.3
Phoxocephalus homilis 19 Urothoe varvarini 8.1
Westwoodilla caecula 16 Pardisynopia synopiae vol
Ampelisca brevisimulata 13 Lysianassa oculata 6.5

TasLE 7.—Amphipoda of Baja California not recorded from Monterey Bay
(species marked with asterisks probably occur in both areas but were absent
in the collections).

Acuminodeutopus heteruropus Megamphopus martesia (intertidal N
Ampelisca indentata of Point Conception)
Ampelisca mexicana * Melphisana bola
Ampelisca shoemakert Metopa dawsonti
Ampelisca venetiensis Monoculodes hartmanae
Amphideutopus oculatus Ocosingo borlus (intertidal N of
Ampithoe ramondi Point Conception)
Cheiriphotis megacheles Orchomene magdalenensis
Chevalia aviculae Paraphozus stenodes
Elasmopus antennatus Photis viuda
Eusiroides monoculoides Platyischnopus metagracilis
Gaviota podophthalma Platyischnopus viscana
Gitana calitemplado Podocerus brasiliensis
*Listriella albina Pontogeneia quinsana
*Listriella eriopisa Pseudokoroga rima
*Tistriella melanica Rudilemboides stenopropodus
Lembos audbettius Stenopleustes monocus pis
Lysianassa dissimilis Stenothoides bicoma

Megaluropus ?agilis
Megamphopus effrenus (intertidal N
of Point Conception)

NO. 3541 BENTHIC AMPHIPODA—BARNARD iat!

TABLE 8.—Amphipoda of Monterey Bay not recorded from coastal shelf of Baja
California; some species may occur on the slope in depths exceeding 100 m
(asterisks mark possible northeastern Pacific cold-temperate species of the
coastal shelf).

Anonyx adoxus Monoculodes norvegicus
*Anonyx carinatus * Monoculodes spinipes
*Centromedon pavor *Opisa tridentata

Corophium uenor Orchomene pacifica
*Dexamonica reduncans *Paraphozus similis
* Hohaustorius sencillus *Parapleustes pugettensis
* Kermystheus ociosa * Photis lacia
*Lepidepecreum gurjanovae *Stenothoe frecanda
*Lysianassa holmest * Protomedeia penates
*Lysianassa oculata Urothoe varvarini

* Megamphopus mamolus

TasBLE 9.—Benthic Amphipoda of the coastal shelf of southern California in
depth classes of 9-18, 19-37, 38-55, 56-73, 74-92, and 93-183 m, from a survey
of 348 samples (occurrences of the species in the surveys of Monterey Bay and
Baja California are shown; other significant records are listed for various species,
and the presumed affinities where significant, are stated).

Southern
California
Baja Probable metric
Name of species Monterey California Literature records affinity depth class
Acidostoma hancocki x sp. xX Northern 18-183
Acuminodeutopus heteruropus x Southern 18-92
Allorchestes angustus Oregon Northern 18
Ampelisca brevisimulata x x 18-183
Ampelisca compressa x x 18-183
Ampelisca cristata x x 18-183
Amopelisca furcigera Okhotsk, Bering Northern 183
Ampelisca hancocki x x 18-183
Ampelisca indentata x Southern 37-92
Ampelisca lobata x x 18-92
Ampelisca macrocephala x x Cold-temperate Northern 18-183
Ampelisca milleri x x 37-183
Ampelisca pacifica x x 18-183
Ampelisca pugetica x x 18-183
Ampelisca romigi x x 37-183
Amphideutopus oculatus x Southern 18-183
Amphilochus neapolitanus x Tropicopolitan Southern 18
Amphilochus picadurus x 18-37
Ampithoe plumulosa 18
Ampithoe simulans Oregon Northern 18
Anonyz carinatus x Northern 55-183
Aoroides columbiae x x 18-183
Argissa hamatipes x 18-183
Atylus tridens British Columbia Northern 18
Batea transversa Southern 18-37
Bathymedon pumilus Bathyal 73-183
Bathymedon roquedo Bathyal 92-183
Bruzelia tuberculata Cold-temperate Northern 183
Byblis veleronis x x 37-183
Ceradocus spinicaudus Oregon Northern 18
Cerapus tubularis x 18-55
Cheiriphotis megacheles x Tropicopolitan Southern 18
Chevalia aviculae x Tropicopolitan Southern 18

Corophium acherusicum 18

12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

TaBLE 9.—Benthic Amphipoda of the coastal shelf of southern California in
depth classes of 9-18, 19-37, 38-55, 56-73, 74-92, and 93-183 m, from a survey
of 348 samples (occurrences of the species in the surveys of Monterey Bay and
Baja California are shown; other significant records are listed for various species,
and the presumed affinities where significant, are stated) Continued

Southern
California
Baja Probable metric
Name of species Monterey California Literature records affinity depth class
Corophium baconi x 18-55
Corophium uenoi x Japan Northern 18
Dexamonica reduncans x Northern 55-183
Dulichia monacantha Cold-temperate Northern 92
Elasmopus antennatus x 18-73
Eohaustorius sencillus Ke Northern 18
Evohaustorius washingtonianus Northern 18-37
Ericthonius brasiliensis x x 18-183
Ericthonius hunteri Cold-temperate Northern 73-183
Eurystheus thompsoni x x 18-183
Garosyrrhoe bigarra 55
Gaviota podophthalma x Southern 18-73
Gitana calitemplado x 18-55
Gitanopsis vilordes 18
Haploops spinosa Cold-temperate Northern 183
Harpiniopsis epistomata Bathyal 92-183
Harpiniopsis fulgens Bathyal 183
Harpiniopsis galera Bathyal 92-183
Heterophlias seclusus American tropical Southern 18
Heterophorus oculatus x X: 18-183
Hippomedon denticulatus x x Cold-temperate Northern 18-183
Hyale rubra frequens 18
Ischyrocerus litotes x x 18-183
Ischyrocerus pelagops x 18-37
Jassa falcata 18
Kermystheus ociosa x 37-183
Lembos audbettius x Southern 55-92
Lembos concavus Southern 18
Lepidepecreum garthi Northern 92-183
Lepidepecreum gurjanovae x Northern 18-183
Leucothoe alata x x 18
Leucothoe spinicarpa x x 18-183
Leucothoides pacifica Southern 18
Liljeborgia brevicornis Northern 18
Listriella albina x 18-183
Listriella diffusa x x 18-92
Listriella eriopisa x 18-183
Listriella goleta x x 18-183
Listriella melanica x 18-183
Lysianassa dissimilis x 18
Lysianassa holmesi x 18-183
Lysianassa oculata x 18-183
Maera danae x Cold-temperate Northern 183
Maera simile x x Oregon Northern 18-183
Mandibulophorus uncirostratus Indo-Pacific Southern 18
Megaluropus longimerus x x 18-37
Megamphopus mamolus x 18
Megamphopus martesia Xs Southern 18
Melita dentata Cold-temperate Northern 18
Melita desdichada x x 55, 183
Melphisana bola x 18-73
Metaceradocus occidentalis Southern 18-73
Metaphocus frequens x x 18-183
Metaphorus fultoni x x 18-183

Metopa dawsoni x 55-183

NO. 3541 BENTHIC AMPHIPODA—BARNARD 13

TaBLE 9.—Benthic Amphipoda of the coastal shelf of southern California in
depths classes of 9-18, 19-37, 38-55, 56-73, 74-92, and 93-183 m, from a survey
of 348 samples (occurrences of the species in the surveys of Monterey Bay and
Baja California are shown; other significant records are listed for various species,
and the presumed affinities where significant, are stated) —Continued

Southern
California
Baja Probable metric
Name of species Monterey California Literature records affinity depth class
Metopella aporpis x x 92-183
Microdeutopus sch mitti x x: 18-55
Monoculodes emarginatus x x 55-183
Monoculodes hartmanae x 18-183
Monoculodes murrius 18-92
Monoculodes norvegicus x Cold-temperate Northern 37-183
Najna ?consilior«um Japan Sea Northern 18
Neomegamphopus roosevelti Baja California Southern 18
Netamelita cortada 18
Nicippe tumida x x Bathyal 37-183
Ocosingo borlus x “Shallow”
Opisa tridentata x Northern 18-183
Orchomene anaquela 37-92
Orchomene decipiens x x(sp.) 18-183
Orchomene magdalenensis x Southern 18
Orchomene pacifica x Northwest Pacific Northern 55-183
Pachynus barnardi x x 18-183
Panaploea rickettsi x x 183
Parajassa angularis 18
Parametopella ninis 18-183
Paraphorus abronius 18-92
Paraphorus bicuspidatus x x 18-183
Paraphorus cognatus Pelagic
Paraphorus daboius 18-183
Paraphozus epistomus x x 18-183
Paraphorus fatigans x x 18-183
Paraphorus floridanus Florida Southern 18-55
Paraphorus heterocuspidatus 18-55
Paraphorus jonesi 18
Paraphorus lucubrans 18-92
Paraphorus obtusidens x x 18-183
Paraphozrus oculatus Cold-temperate Northern 183
Paraphorus robustus x x 18-183
Paraphorus similis x 18-183
Paraphorus spinosus 18-183
Paraphorus stenodes x 18-92
Paraphorus tridentatus 18-37
Paraphozrus variatus x x 18-92
Parapleustes oculatus Northern 183
Parapleustes pugettensis 18-183
Pardaliscella symmetrica Bathyal 92-183
Pardisynopia synopiae x x Bathyal 55-183
Photis bifurcata x x 18-92
Photis brevipes x x 18-183
Photis californica x x 18-92
Photis lacia x Northern 18-183
Photis macrotica x x 55-183
Phoxocephalus homitis x x 73-183
Platyischnopus viscana x Baja California Southern 18
Pleustes platypa Rare So. Calif. Northern 18
Podocerus brasiliensis XG Tropicopolitan Southern 18
Podocerus cristatus x x 18-183
Polycheria antarctica 18
Pontogeneia rostrata Northwest Pacific Northern 18

Prachynella lodo x x 37-183

14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

TABLE 9.—Benthic Amphipoda of the coastal shelf of southern California in
depths classes of 9-18, 19-37, 38-55, 56-73, 74-92, and 93-183 m, from a survey
of 348 samples (occurrences of the species in the surveys of Monterey Bay and
Baja California are shown; other significant records are listed for various species,
and the presumed affinities where significant, are stated) Continued

Southern
California
Baja Probable metric
Name of species Monterey California Literature records affinity depth class
Protomedeia articulata x x Northern 18-183
Pseudokoroga rima x Southern 18
Rhachotropis inflata Cold-temperate Northern 183
Rhachotropis oculata x Cold-temperate Northern 73-183
Rudilemboides stenopropodus 18-73
Schisturella cocula Bathyal 183
Socarnes illudens x Northern 183
Stenopleustes monocuspis x Southern 73-183
Stenothoe frecanda x 55-183
Stenothoides bicoma x 18-183
Stenula modosa 73-183
Sympleustes subglaber Genus northern Northern 18-183
Synchelidium rectipalmum 18-183
Synchelidium shoemakeri x 18-183
Synchelidium sp. A x 18-183
Synchelidium sp. E x 18-183
Synchelidium sp. G x 18-183
Tiron biocellata x x 18-183
Uristes entalladurus Baja California 18
Urothoe varvarini be North Pacific Northern 37-183
Westwoodilla caecula x x Cold-temperate Northern 18-183

TaBLeE 10.—Amphipoda of Monterey Bay recorded from the northwestern Pa-
cific Ocean.

Depths of 1549 m Depths of 50-139 m
Corophium uenot Ampelisca macrocephala
Leucothoe spinicarpa Argissa hamatipes
Monoculodes norvegicus Hippomedon denticulatus

Maera danae

Nicippe tumida
Orchomene pacifica
Paraphozus obtusidens
Westwoodilla caecula
Urothoe varvarini

Family Acanthonotozomatidae

Panoploea rickettsi Shoemaker
Panoploea rickettsi Shoemaker, 1931, pp. 1-5, figs. 1, 2.—Barnard, 1964a, p. 212.

Material: 6425 (1).
Distribution: Monterey Bay to San Quintin Bay, Baja California,
10-92 m.

NO. 3641 BENTHIC AMPHIPODA—BARNARD 15

Family Ampeliscidae

Ampelisca brevisimulata Barnard

Ampelisca brevisimulata Barnard, 1954a, pp. 33-35, pls. 23, 24; 1964a, p. 212.

Material: 6433 (1), 6435 (2), 6471 (1).

Distribution: Caribbean Sea and eastern Pacific Ocean from Pan-
ama to Monterey Bay, 11-172 m.

Ampelisca compressa Holmes

Ampelisca compressa.—Barnard, 1960a, pp. 31-32; 1964a, p. 213.

Material: 6427 (1), 6428 (4), 6429 (2), 6430 (3), 6435 (1), 6447 (9),
6448 (2), 6451 (1), 6452 (2), 6471 (2).

Distribution: Western Atlantic Ocean, Caribbean Sea, eastern Pa-
cific Ocean from Panama to Puget Sound, Washington, 1-266 m.

Ampelisca cristata Holmes

Ampelisca cristata.—Barnard, 1954a, pp. 26-28, pls. 17, 18; 1964a, p. 213.

Material: 6426 (3), 6427 (7), 6428 (1), 6429 (5), 6430 (2), 6446 (1),
6447 (2), 6449 (2), 6450 (1), 6453 (5).

Distribution: Caribbean Sea and eastern Pacific Ocean from Ecua-
dor to Tomales Bay, California, 6-152 m.

Ampelisca hancocki Barnard

Ampelisca hancocki Barnard, 1954a, pp. 37-38, pl. 26; 1964a, p. 213.

Material: 6433 (1), 6451 (1), 6465 (1), 6471 (2), 6474 (2).

Distribution: Eastern Pacific Ocean from Monterey Bay to Costa
Rica, 9-157 m.

Ampelisca lobata Holmes
FIGURE 2a

Ampelisca lobata.—Barnard, 1954a, pp. 11-14, pls. 5, 6; 1964a, p. 214.

Material: 6425 (24).

Distribution: Caribbean Sea, eastern Pacific Ocean from Ecuador
and the Galapagos Islands to Monterey Bay, 0-183 m.

Ampelisca macrocephala (Liljeborg)
Ampelisca macrocephala.—Bamard 1954a, pp. 41-43, pl. 29; 1960a, pp. 28-30,
fig. 7; 1964a, p. 214.

Material: 6426 (1), 6427 (2), 6428 (1), 6430 (3), 6431 (2), 6432 (4),
6433 (9), 6435 (14), 6488 (2), 6443 (2), 6446 (2), 6450 (1), 6451 (1),
6455 (2), 6460 (1), 6464 (3), 6465 (1), 6466 (8), 6470 (4), 6471 (12),
6478 (1).

Distribution: Subarctic-boreal in the North Atlantic and North
Pacific Oceans, submerging in waters of low latitudes, 5-1686 m.

16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Ampelisca milleri Barnard
Ampelisca millert Barnard, 1954a, pp. 9-11, pls. 3, 4; 1964a, p. 215.

Material: 6474 (3).

Distribution: Eastern Pacific Ocean from Ecuador and the Gala-
pagos Islands to San Francisco Bay, California, 15-187 m in the open
sea, from shallow water in San Francisco Bay.

Ampelisca pacifica Holmes
Ampelisca pacifica—Barnard 1954a, pp. 31-33, pls. 21, 22; 1964a, p. 215.

Material: 6432 (3), 6458 (4), 6466 (2), 6471 (2), 6477 (1), 6478 (1).
Distribution: Caribbean Sea, eastern Pacific Ocean from Panama
to Monterey Bay, California, 24-183 m.

Ampelisca pugetica Stimpson

Ampelisca pugetica.—Barnard 1954a, pp. 49-51, pls. 35, 36; 1960a, p. 31, fig. 9;
1964a, p. 215.

Material: 6425 (46), 6426 (1), 6427 (3), 6428 (2), 6430 (2), 6447 (3),
6452 (1), 6463 (1).

Distribution: Caribbean Sea, eastern Pacific Ocean from Peru to
Puget Sound, Washington, 9-183 m.

Ampelisca romigi Barnard

Ampelisca romigi Barnard, 1954a, pp. 18-20, pls. 10, 11; 1960a, p. 34; 1964a,
pp- 215-216.

Material: 6471 (10), 6474 (48).

Distribution: Caribbean Sea, eastern Pacific Ocean from Ecuador
to Monterey Bay, 3-504 m.

Byblis veleronis Barnard

Bypblis veleronis Barnard, 1954a, pp. 52-54, pls. 37, 38; 1964a, p. 216.

Material: 6426 (1), 6428 (3), 6429 (2), 6430 (4), 6431 (2), 6432 (8),
6433 (2), 6485 (2), 6438 (2), 6447 (1), 6451 (5), 6452 (3), 6455 (4),
6458 (2), 6462 (1), 6464 (2), 6465 (1), 6466 (6), 6470 (2), 6471 (3),
6474 (6), 6480 (1).

Distribution: Monterey Bay to the Gulf of California, 31-422 m.

Family Amphilochidae

Amphilochus picadurus Barnard
Amphilochus picadurus Barnard, 1962c, pp. 126-129, fig. 4; 1964a, p. 217.

Material: 6425 (4).
Distribution: Monterey Bay to southern California, 33° N., 4-41 m.

NO. 3541 BENTHIC AMPHIPODA—BARNARD 17

Family Aoridae

Aoroides columbiae Walker
Aoroides columbiae Barnard, 1961, p. 180; 1964a, pp. 217-218.
Material: 6425 (57), 6432 (2), 6433 (2), 6435 (2), 6440 (1), 6445 (8),
6470 (1), 6471 (6), 6474 (11).
Distribution: Puget Sound, Washington to San Quintin Bay, Baja
California, 0-180 m.

Microdeutopus schmitti Shoemaker

Microdeutopus schmitti Shoemaker, 1942, pp. 18-21, fig. 6; 1959a, pp. 32-33,
pl. 9; 1961, p. 180; 1964a, p. 218.

Material: 6425 (419), 6425 (1).
Distribution: Monterey Bay to Cape San Lucas, Baja California,
0-43 (65?) m.
Family Corophiidae
Corophium uenoi Stephensen

Corophium uenoit Stephensen, 1932, pp. 494-498, figs. 3, 4.—Barnard, 1952
pp. 28-32, pls. 8, 9; 1959a, p. 39.—Nagata, 1960, p. 178.—Barnard, 1961,
p. 183; 1964b, p. 112, chart 16.

Material: 6425 (3).

Distribution: Japan; eastern Pacific Ocean from Monterey Bay
at 24 m depth, to San Quintin Bay, Baja California, rarely occurring
in the open sea, more often in lagoons or estuaries such as Morro
Bay, Newport Bay, and San Quintin Bay, intertidal to 2 m.

Ericthonius brasiliensis (Dana)

Ericthonius brasiliensis.—Barnard 1955a, pp. 37-38; 1959a, p. 39; 1961, p. 183;
1964a, p. 219.

Material: 6425 (69), 6445 (2).
Distribution: Cosmopolitan in tropical, warm-temperate and some
boreal seas, 0-130 m.

Family Dexaminidae
Dexamonica reduncans Barnard
Dexamonica reduncans Barnard, 1958, pp. 130-132, pls. 26, 27.
Material: 6433 (2), 6471 (6), 6474 (1).
Distribution: Monterey Bay to southern California, 33° N, in

Monterey Bay from 107 to 139 m in depth, in southern California

from about 55 to 183 m.
219-939—66——_3

18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Family Eusiridae
Rhachotropis oculata (Hansen)
Figures 2b-f

Rhachotropis oculata.—Stephensen, 1944, pp. 97-98.—Gurjanova, 1951, pp. 712-
713, fig. 496.

Stebbing (1906) stated that the rostrum of this species is small.
Although the heads of the specimens at hand resemble Hansen’s
figure reproduced by Gurjanova (1951), the rostra are long and
hidden from lateral view by the first antennae. Article 3 of antenna
1 is not as short as or indistinct as that described by Stebbing or
figured by Hansen. Small specimens, difficult to separate from R.
inflata Sars, may be distinguished by the produced corner of the
second pleonal epimeron.

Material: Hancock Velero stations 1027 (2), 1149 (4), 4753 (4),
4772 (1), 4824 (1), 5615 (1), 5760 (1), 6001 (1), 6006 (1).

Records: Coastal shelf of southern California and Santa Catalina
Island, 64-274 m, caught in benthic grabs and dredges, probably
a demersal species infrequently collected in benthic closing devices.

Distribution: North Atlantic and North Pacific Oceans, Bering
Sea, Okhotsk Sea, Japan Sea, 20-274 m. These are the southern-
most records of this species in the eastern Pacific Ocean.

Family Gammaridae
Maera danae (Stimpson)

Maera danae.—Shoemaker, 1955, pp. 53-54 (with references).—Barnard, 1964b,
pp. 108-109.
Maera loveni.—J. L. Barnard, 1962b, p. 103, fig. 19 (not Bruzelius).

Material: 6462 (2), 85 m.

Additional record: 4770 (1), southern California, 33°21’ N., 117°34’
W.,)15)m.

Remarks: According to Shoemaker (1955) this species lacks an
articulated spine at the palmar defining corner of gnathopod 2, but
a spine is present on the specimens at hand, yet the second articles
of pereopods 3-5 are broad, not slender as in M. loveni, and distinct
eyes are present. Regardless of the gnathopodal spine, the specimens
are identified as M/. danae.

Distribution: North Atlantic and North Pacific Oceans, in the
North Pacific known as far south as San Quintin Bay in shallow water
but primarily a subarctic species submerging with depth toward the
tropics, rather rare in southern California and usually occurring in
depths of about 200 m.

NO. 3541 BENTHIC AMPHIPODA—BARNARD 19

Maera simile Stout

Maera simile-—Barnard, 1959a, pp. 24-25, pl. 4; 1964a, p. 222.

Material: 6425 (1).

Distribution: Coos Bay, Oregon to San Quintin Bay, Baja Cali-
fornia, 0-43 m.

Megaluropus longimerus Schellenberg

Megaluropus longimerus.—Barnard, 1962b, p. 103, figs. 20, 21; 1964a, p. 224.

Material: 6440 (8), 6442 (3).

Distribution: Lagos, Nigeria; in the eastern Pacific Ocean from
Monterey Bay to San Ramon Bay, Baja California, 10-108 m.

Melita desdichada Barnard

Melita desdichada Barnard, 1962b, p. 110, fig. 22; 1964a, p. 224.

Material: 6455 (4), 6460 (1).

Distribution: Monterey Bay to San Ramon Bay, Baja California,
10-108 m.

Family Haustoriidae
Eohaustorius sencillus Barnard

Eohaustorius sencillus Barnard, 1962f, pp. 249-252, figs. 1, 2.

Material: 6444 (19), 6445 (1), 6454 (28).

Distribution: Monterey Bay to Point Conception, California,
14-25 m.

Urothoe varvarini Gurjanova

Urothoe varvarini Gurjanova, 1953, pp. 219-221, figs. 3, 4.—Barnard, 1957,
pp. 82-84.—Gurjanova, 1962, pp. 426-428, fig. 142.

Material: 6433 (1), 6438 (1), 6471 (6), 6474 (9).

Distribution: Northwestern Pacific Ocean, Japan Sea, Okhotsk
Sea, 5-13 m; California from Monterey Bay to southern California
at 33° N, 40-200-+ m.

Family Isaeidae (=Photidae)
Eurystheus thompsoni (Walker)

Eurystheus thompsoni.—Barnard, 1959a, p. 36; 1961, p. 182; 1964a, p. 237.

Material: 6425 (44), 6445 (1).

Distribution: Puget Sound, Washington to Magdalena Bay, Baja
California, 0-135 m.

Kermystheus ociosa Barnard

Kermystheus ociosa Barnard, 1962a, p. 23, fig. 8.

Material: 6432 (2), 6435 (2), 6471 (7), 6474 (S1).

Distribution: Monterey Bay to the coastal shelf of southern
California between Point Conception and San Diego, 27-165 m.

20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Megamphopus mamolus Barnard
Megamphopus mamolus Barnard, 1962a, pp. 23-26, fig. 9.

Material: 6425 (68), 6445 (9).

Distribution: Monterey Bay to Point Conception, California, at
the latter in a red algal Diopatra bed, 16 m; at Monterey in depths
of 24-25 m.

Photis bifurcata Barnard
Photis bifurcata Barnard, 1962a, pp. 30-31, fig. 10; 1964a, p. 240.

Material: 6425 (3832), 6445 (15).
Distribution: Monterey Bay to San Cristobal Bay, Baja Cali-
fornia, 11-93 m.
Photis brevipes Shoemaker

Photis brevipes Shoemaker, 1942, pp. 25-27, fig. 9—Barnard, 1962a, pp. 31-33,
fig. 11; 1964a, pp. 240-241.
Material: 6425 (58), 6445 (7), 6471 (2), 6477 (5).
Distribution: Coos Bay, Oregon to Magdalena Bay, Baja Cali-
fornia, 0-135 m.
Photis californica Stout

Photis californica.—Barnard, 1962a, pp. 33-36, figs. 12, 13; 1964a, p. 241.

Material: 6471 (12), 6474 (58).

Distribution: Monterey Bay to San Cristobal Bay, Baja Cali-
fornia, 10-98 m, in southern California; maximum depth in Monterey
Bay 139 m.

Photis lacia Barnard

Photis lacia Barnard, 1962a, pp. 42-44, fig. 18.

Material: 6425 (4), 6430 (?2), 6432 (2), 6433 (9), 6465 (1), 6469
(3), 6471 (34), 6474 (24), 6477 (2).

Distribution: Monterey Bay to southern California, 32° N, 9-146
m, abundant on the shelf in depths of 73-92 m (88 individuals per m?).

Photis macrotica Barnard
Photis macrotica Barnard, 1962a, p. 44, fig. 19; 1964a, p. 241.

Material: 6474 (1).

Distribution: Monterey Bay to San Cristobal Bay, Baja Cali-
fornia, 55-157 m.

Protomedeia articulata Barnard

Protomedeia articulata Barnard, 1962a, pp. 48-50, fig. 21; 1964a, p. 242.

Material: 6426 (1), 6429 (1), 6438 (1), 6445 (6), 6455 (6), 6456 (1),
6457 (2), 6458 (2), 6466 (1), 6471 (71).

Distribution: Monterey Bay to San Quintin Bay, Baja California,
18-200+ m.

NO. 3641 BENTHIC AMPHIPODA—BARNARD Dt

Protomedeia penates, new species

FIGURE 3

Diagnosis of male: Article 2 of gnathopod 1 lacking posterodistal
swelling; article 7 of gnathopod 1 overlapping palm by more than
75 percent of its length, palm with triangular process and defined by
one stout spine; palm of gnathopod 2 with defining tooth exceeding
transverse palm and bearing small basal tooth anterior to it; inner
ramus of uropod 8 slightly shorter than outer ramus.

Female: Gnathopods with oblique palms defined by large spines.

Notes: Article 2 of pereopod 5 has a small dorsoposterior tooth;
pleonal epimeron 3 bears lateral setae, variable in extent and often
absent, especially in males.

Holotype: Allan Hancock Foundation no. 5923, male 6.0 mm.

Type locality: White Gulch, Tomales Bay, California, sta. 1-59-10,
June 29, 1959, depth of 48 feet, on dark sand and mud, associated
with Chone and Pectinaria, collected by Dr. Joel W. Hedgpeth and
Dr. Ralph G. Johnson, 8 specimens.

Material: 6455 (4).

Relationship: Morphologically, this species is most closely related
to Protomedeia fasciatoides Bulycheva (1952) but differs in having a
longer palmar tooth of male gnathopod 2 and oblique palms on the
female gnathopods. The absence of a posterodistal prominence on
article 2 of gnathopod 1 distinguishes P. penates from P. fasciata
Kr¢gyer (Sars, 1895, pl. 196), a species having a distinctive aspect as
illustrated by Gurjanova (1951). The largest male, 7.0 mm, from
station 6455, is obviously fully mature and lacks the gnathopodal
prominence. Protomedeia popovi Gurjanova (1951) is also similar to
P. penates, but the male has a more oblique gnathopodal palm lack-
ing the subsidiary palmar tooth.

Distribution: Tomales Bay to Monterey Bay, California, 15-76 m.

Family Ischyroceridae
Ischyrocerus litotes (Barnard)
Ischyrocerus litotes—Barnard, 1962a, pp. 53-56, figs. 23, 24; 1964a, pp. 226-227.

Material: 6425 (1).

Remarks: I may have been in error in removing this species from
its original position in Microjassa because of the slight difference in
size of coxae 5 and 6. As in Microjassa, coxa 1 of this species is
small in contrast to members of Ischyrocerus. Both genera prob-
ably should be emended to permit J. litotes to be included with

32 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Microjassa. Ischyrocerus litotes at least forms a bridge between the
two concepts. Its small size, very shiny, slick body, and poor pig-
ment in alcohol make it highly distinctive from other species of
Ischyrocerus.

Distribution: Monterey Bay to San Cristobal Bay, Baja Cali-
fornia, 1-157 m.

Family Leucothidae

Leucothoe alata Barnard

Leucothoe alata Barnard, 1959a, pp. 19-20, pl. 1; 1962c, p. 132, figs. 7, D, E, F;
1964a, p. 227.

Material: 6425 (5).
Distribution: Monterey Bay to San Ramon Bay, Baja Cali-
fornia, 0-24 m.
Leucothoe spinicarpa (Abildgaard)
Leucothoe spinicarpa—Sars, 1895, p. 100, pl. 101, fig. 1—Barnard, 1962c, p. 132,
figs. 7, A, B, C; 1964a, p. 227.

Material: 6425 (2).
Distribution: Cosmopolitan, 0-1505 m.

Family Liljeborgiidae
Liljeborgia kinahani (Bate)

Liljeborgia kinahani.—Sars, 1895, pp. 532-533, pl. 188, fig. 1—Chevreux and
Fage, 1925, p. 157, fig. 157.— Barnard, 1964a, p. 228.
Material: 6425 (5).
Distribution: Possibly bipolar, rare in southern California, in the
eastern Pacific Ocean known as far south as off San Quintin Bay,
Baja California, 24-41 m.

Listriella diffusa Barnard
Listriella diffusa Barnard, 1959b, pp. 18-20, figs. 3-5; 1964a, p. 228.
Material: 6440 (1).
Distribution: Monterey Bay to San Cristobal Bay, Baja Cali-
fornia, 12-172 m.
Listriella goleta Barnard
Listriella goleta Barnard, 1959b, pp. 20-22, figs. 5-7; 1964a, p. 229.
Material: 6427 (1), 6428 (2), 6429 (1), 6431 (1), 6446 (1), 6448
(1), 6450 (1), 6451 (2), 6466 (1), 6477 (1), 6480 (1).
Distribution: Monterey Bay to San Cristobal Bay, Baja Cali-
fornia, 12-200+ m.

NO. 3541 BENTHIC AMPHIPODA—BARNARD 23

Family Lysianassidae

Acidostoma hancocki Hurley
Acidostoma hancocki Hurley, 1963, pp. 37-40, figs. 9, 10.

Material: 6429 (2).
Distribution: Monterey Bay to southern California, 33° N, 22-
73 m.
Anonyx adoxus Hurley
FIGurE 4
Anonyx adoxus Hurley, 1963, pp. 108-112, figs. 35, 36.

Diagnosis: Eyes dark, rather small for genus, forming an elongated
oval of even dimensions, not expanded at either end; epistome and
upper lip not differentially produced; coxa 1 rounded ventrally;
gnathopod 1 with article 6 concave posteriorly, narrowing distally,
palm short, coarsely serrate, dactyl strongly overlapping palm; palm
of gnathopod 2 short, slightly produced, dactyl short, fitting palm
perfectly; pereopods 1 and 2 with hooked, unstriated spine at base
of article 7; all pereopods with large spines on sixth articles; inner
ramus of uropod 2 not abnormally shortened, bearing small con-
striction on dorsal margin; rami of uropod 3 subequal in length,
article 2 of outer ramus very small (see figure) ; first pleonal epimeron
differing on the two sides of the adult specimen, on the left side with
the anterior edge straight, on the right side produced anteriorly
(see figures); second pleonal epimeron with rounded anteroventral
corner, posterior corner with small, blunt tooth; third pleonal epimeron
rounded at anteroventral corner, posterior corner with moderately
slender, medium-sized tooth; body lacking sculpture or minute
ornamentation.

Material: 6432, female, 10.0 mm, and 17 hatched juveniles.

Remarks: This adult specimen has the special characteristics noted
by Hurley in his key (1963, p. 103), but it differs slightly in the con-
figuration of its epistome-labrum complex, the accessory flagellum is
shorter, and marginal spines are absent from the telsonic lobes. The
weakly hooked, blunt distal spines of the sixth articles on pereopods
1-2 are small, as shown by Hurley. However, the minute ornamen-
tation of the spines has the appearance of the large spines of the A.
liljeborgi group of the genus.

Anonyz adoxus most closely resembles Anonyzx nugax (Phipps) and
its varieties as arranged by Gurjanova (1962), but the eyes of A.
adoxus are not differentially widened. The following species, refer-
ences to which may be found in Gurjanova (1962), differ from the
specimen at hand in the morphology of the parts and appendages
listed: A. liljeborgi, wpper lip and uropod 2, third pleonal epimeron;
A. affinis, A. minimus, uropods 2 and 38; A. validus, third pleonal

24 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

epimeron; A. kurilicus, A. magnus, uropod 2; A. ampulloides, A.

laticoxae, uropod 2 and eyes; A. ochoticus, eyes and epistome; A.

compactus and A. oculatus, eyes and spines of pereopods 1 and 2.
Distribution: Monterey Bay (type locality), 18-98 m.

Anonyx carinatus (Holmes)

Lakota carinata Holmes, 1908, pp. 498-500, fig. 9.—Gurjanova, 1962, pp.
302-303, fig. 100.

Anonyz carinatus.—Hurley, 1963, pp. 103-108.

Material: 6477 (1).

Distribution: Monterey Bay to southern California, 33° N.,
69-200 m.

Centromedon pavor, new species
FIGure 5

Diagnosis: Lateral cephalic lobes short, rounded terminally, sep-
arated from large rostral area by deep concavity; antenna 1 very stout,
articles 2 and 3 short, telescoped into article 1, article 1 of flagellum
heavily armed with aesthetes; coxa 5 with well-defined posterior lobe;
urosomite 1 bulbous dorsally.

Holotype: Allan Hancock Foundation no. 5922, female, 2.9 mm.

Type locality: Station 6459, Monterey Bay, California, 84 m,
September 1959.

Material: Twospecimens from the type locality and 6462 (1), 6477 (1).

Relationship: The genus Centromedon became monotypic when
Barnard (1962d) removed all but its type species to the genus Uristes.
Centromedon pumilus (Liljeborg) differs from C. pavor in having very
acute, projecting lateral cephalic lobes, slender first antennae, a sym-
metrically lobed fifth coxa, and a poorly ornamented first urosomite.
As the specimens at hand and that figured by Sars are females, these
differences cannot be attributed to sexual dimorphism. ‘The mouth-
parts of C. pavor correspond to those figured for C. pumilus by Sars
(1895, pl. 34, fig. 2).

Hippomedon denticulatus (Bate)

Hippomedon denticulatus—Hurley, 1963, pp. 137-140, fig. 45.—Barnard, 1964a,
p. 230; 1964b, pp. 80-82.
Material: 6426 (1), 6430 (1), 6445 (2), 6447 (1), 6471 (4), 6474 (2),
6477 (5).
Distribution: Subarctic-boreal of North Atlantic and North Pacific
Oceans, 0-924 m; in the eastern Pacific its southern record is off San
Cristobal Bay, Baja California.

Lepidepecreum gurjanovae Hurley
Lepidepecreum gurjanovae Hurley, 1963, pp. 49-53, figs. 13, 14.
Material: 6469 (1), 6474 (1).

NO. 3541 BENTHIC AMPHIPODA—BARNARD 295

Distribution: In the northeastern Pacific Ocean from about 33° N
to about 49° N, 109-1740 m.

Lysianassa holmesi (Barnard), new combination

Aruga holmesi Barnard, 1955b, p. 100, pls. 27, 28.—Gurjanova, 1962, pp.
299-301, figs. 98, 99.
Lysianopsis holmesi.—Hurley, 1963, pp. 74-75, fig. 21b.

Material: 6431 (1), 6474 (1).

Distribution: Monterey Bay to Ecuador, 1-183 m.

Remarks: I am carrying Hurley’s synthesis of various lysianassid
genera one more step by fusing Lysianopsis Holmes, along with Aruga
Holmes, Shoemakerella Pirlot, Arugella Pirlot, and Pronannonyz
Schellenberg to Lysianassa. This procedure is discussed in another
paper (Barnard, MS).

Lysianassa oculata (Holmes), new combination
Aruga oculata Holmes, 1908, pp. 505-507, figs. 14, 15.—Barnard, 1955b, p. 98,
pl. 29, figs. a-f,h,j—Gurjanova, 1962, pp. 294-296, fig. 96.
Lysianopsis oculata.—Hurley, 1963, p. 74, fig. 21c.

Material: 6426 (1), 6428 (1), 6429 (1), 6431 (6), 6438 (1), 6439 (1),
6448 (1), 6451 (2), 6455 (1).

Distribution: Monterey Bay to southern California, 32° N, 18 to
?300 m, generally shallower than 100 m.

Opisa tridentata Hurley
Opisa tridentata Hurley, 1963, pp. 26-30, figs. 4, 5.

Material: 6435 (1), 6471 (8), 6474 (2).
Distribution: Monterey Bay to southern California, 33° N, 44-
183 m.
Orchomene decipiens Hurley, new combination
Orchomenella decipiens Hurley, 1963, pp. 127-130, figs. 43, 44.
Orchomene species.—Barnard, 1964a, p. 231.
Material: 6435 (1), 6471 (5), 6474 (2), 6477 (1).
Distribution: Monterey Bay to San Cristobal Bay, Baja California,
35-180 m.
Remarks: Barnard (1964b) synonymized Orchomenella Sars with
Orchomene Boeck.
Orchomene pacifica (Gurjanova)
Orchomenella pacifica Gurjanova, 1938, pp. 252-254, fig. 3; 1951, p. 287, fig. 155;
1962, pp. 174-177, figs. 52, 53.
Orchomene pacifica—Barnard, 1964b, pp. 92-93, fig. 13.
Material: 6471 (7), 6474 (4).
Distribution: Northwestern Pacific Ocean, Japan Sea, Okhotsk
Sea, 29-129 m; California from Monterey Bay to southern California,
33° N, 46-183 m.

26 PROCEEDINGS OF THE NATIONAL MUSEUM Vou. 110

Pachynus barnardi Hurley

Pachynus barnardi Hurley, 1963, pp. 31-35, figs. 6, 7.
Pachynus species, Barnard, 1964a, p. 232.
Material: 6458 (1), 6469 (1).
Distribution: Monterey Bay to San Ramon Bay, Baja California,
12-183 m.
Prachynella lodo Barnard

Prachynella lodo Barnard, 1964a, p. 233, fig. 7.

Material: 6426 (1), 6471 (1).
Distribution: Monterey Bay to San Cristobal Bay, Baja California,
10-157 m.
Family Oedicerotidae

Monoculodes emarginatus Barnard
Monoculodes emarginatus Barnard, 1962e, pp. 361-363, fig. 4; 1964a, p. 234.

Material: 6432 (2), 6435 (2), 6438 (1), 6466 (3), 6469 (1), 6471 (4).

Distribution: Monterey Bay to San Quintin Bay, Baja California,
55-200-+ m.

Monoculodes norvegicus (Boeck)
Monoculodes norvegicus.—Sars, 1895, pp. 301-302, pl. 107, fig. 1.—Barnard, 1962e;
p. 367 (with references).

Material: 6425 (1).

Distribution: Subarctic-boreal in the North Atlantic and North
Pacific Oceans, occurring in southern California as about 1.3 individ-
uals per m? on the coastal shelf, in depths of 20-146 m.

Monoculodes spinipes Mills

Monoculodes spinipes Mills, 1962, pp. 12-14, fig. 3—Barnard, 1962e, pp. 368-369,
fig. 10.
Material: 6454 (1).
Distribution: British Columbia to Point Conception, California,
intertidal to 20 m.
Synchelidium species A

A new species with vestigial rostrum, manuscript in preparation.

Material: 6427 (1), 6428 (1), 6429 (7), 6430 (2), 6431 (3), 6438 (2),
6465 (1), 6466 (1), 6474 (1).

Distribution: Monterey Bay to southern California, 32° N, 10-
100+ m.

Synchelidium species E

A new species, manuscript in preparation.

Material: 6435 (1), 116 meters.

Distribution: Monterey Bay to southern Califronia, 32° N, about
40-183-+- m.

NO. 3541 BENTHIC AMPHIPODA—BARNARD D7

Synchelidium species G

A new species, manuscript in preparation.

Material: 6435 (2), 6439 (4); varietal type: 6426 (2), 6427 (3),
6433 (2) 6440 (3), 6469 (1).

Distribution: Monterey Bay to southern California, 32° N, 10 to
slightly more than 120 m.

Synchelidium shoemakeri Mills
Synchelidium shoemakeri Mills, 1962, pp. 15-17, figs. 4, 6A.

Material: 6432 (2), 98 m.

Distribution: British Columbia south to southern California, inter-
tidal in northern end of its range, subtidal in southern California, to
depths slightly exceeding 100 m but very rare below 40 m.

Westwoodiila caecula (Bate)
Westwoodilla caecula.— Mills, 1962, pp. 509, figs. 1, 6A.—Barnard, 1962e, p. 370;
1964a, p. 235.

Material: 6427 (1), 6431 (1), 6435 (3), 6438 (1), 6469 (1), 6474 (1).

Distribution: A cold-temperate species of the North Atlantic and
North Pacific Oceans, in the eastern Pacific known as far south as
Todos Santos Bay, Baja California; intertidal in British Columbia but
occurring in southern California in 12-200 m.

Family Pardaliscidae

Nicippe tumida Bruzelius
Nicippe tumida.—Barnard, 1959c, pp. 39-40, figs. 1, 2; 1964a, p. 235.

Material: 6431 (2), 6433 (1), 6457 (1), 6465 (2), 6466 (4).
Distribution: Apparently cosmopolitan, submerging in the tropics,
34-1367 m.

Pardisynopia synopice Barnard
Pardisynopia synopiae Barnard, 1962b, pp. 77-79, figs 3, 4; 1964a, pp. 235-236.
Material: 6431 (2), 6435 (2), 64388 (3), 6465 (1), 6469 (1), 6471 (1).
Distribution: Monterey Bay to San Quintin Bay, Baja California,
53-200+ m.
Family Phoxocephalidae

Heterophoxus oculatus (Holmes)
Heterophoxus oculatus—Barnard, 1960b, pp. 320-324, pls. 59-61; 1964a, p. 242.
Material: 6425 (3), 6431 (2), 6432 (8), 6483 (2), 6455 (5), 6456 (1),
6457 (4), 6458 (5), 6459 (1), 6462 (1), 6464 (8), 6465 (8), 6471 (1),
6474 (8), 6477 (13), 6480 (1).
Distribution: Puget Sound, Washington to Panama, 13-1785 m;
in San Quintin Bay, Baja California, 2 m.

28 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Metaphoxus frequens Barnard
Metaphoxus frequens Barnard, 1960b, pp. 304-306, pls, 51, 52; 1964a, p. 242.

Material: 6425 (12), 6426 (1), 6428 (4), 6429 (2), 6430 (4), 6431
(1), 6432 (5), 6433 (12), 6435 (6), 6438 (3), 6458 (1), 6466 (2), 6470
(2), 6471 (3), 6477 (1), 6480 (1).

Distribution: Monterey Bay to Isabel Island, Mexico, 13-458 m.

Metaphoxus fultoni (Scott)

Metaphoxus fultoni—Chevreux and Fage, 1925, pp. 106-107, figs. 96, 97.—
Barnard, 1964a, pp. 242-243.
Material: 6425 (4), 6474 (4).
Distribution: Eastern Atlantic Ocean and Mediterranean Sea,
from England to Tunisia; in the eastern Pacific Ocean from Monterey
Bay to San Cristobal Bay, Baja California, 0-170 m.

Paraphoxus bicuspidatus Barnard

Paraphozus bicuspidatus Barnard, 1960b, pp. 218-221, pls. 15, 16; 1964a, p. 243,
fig. 12.

Material: 64380 (8), 6481 (21), 6482 (22), 6483 (23), 6485 (18),
6438 (13), 6447 (2), 6448 (6), 6451 (13), 6452 (12), 6455 (4), 6456 (2),
6457 (1), 6458 (5), 6466 (7), 6469 (4), 6471 (5).

Distribution: Monterey Bay to Santa Maria Bay, Baja California,
8-210 m.

Paraphoxus epistomus (Shoemaker)
Paraphozrus epistomus.—Barnard, 1960b, pp. 205-209, pls. 6-8; 1964a, p. 243.
Material: 6429 (4).
Distribution: Mendocino County in northern California to Panama,
0-182 m; northwestern Atlantic Ocean from New Hampshire to
South Carolina.

Paraphoxus fatigans Barnard

FIGurREs 6, 7
Paraphozxus fatigans Barnard, 1960b, pp. 209-210, pl. 9; Barnard, 1964a, p. 244.

Material: Kind 1: 6426 (48), 6427 (14), 6428 (1), 6429 (3), 6440
(5), 6442 (17), 6443 (18), 6444 (6), 6445 (32), 6446 (18), 6448 (1),
6449 (7), 6450 (6), 6453 (2), 6454 (40). Kind 2: 6427 (88), 6428
(33), 6429 (54), 6430 (60), 6431 (30), 6432 (14), 6433 (23), 6435
(11), 6438 (24), 6445 (2), 6446 (8), 6447 (26), 6448 (41), 6449 (4),
6451 (6), 6452 (4), 6453 (18), 6466 (25), 6469 (1), 6471 (1).

Remarks: The most common members of Paraphorus in Monterey
Bay are the most difficult to identify. These specimens have several
characters which are intermediate between those of P. fatigans and
P. daboius (both Barnard, 1960b). Two kinds of fifth pereopods

NO. 3541 BENTHIC AMPHIPODA—BARNARD 29

occur which resemble the figures of P. fatugans published by Barnard.
The first gnathopods of P. fatigans are slender, those of P. daboius
strongly expanded and those of the specimens at hand are intermediate
between the two species. The two species and the specimens at hand
have small female eyes. The epistomal cusp is usually longer than
it is in either P. fatigans or P. daboius.

The two configurations of pereopod 5 are: Article 2 has more than
3 very small teeth crowded together on the posterior margin, the
oblique ventral edge being slightly convex rather than truncate as in
P. fatigans from southern California (figs. 6d,k,l); article 2 has only
2 or 3 slightly enlarged, less crowded teeth, the oblique ventral margin
being nearly truncate (fig. 6c). A third kind of rare occurrence, is
shown in figures 6a,¢,i,7; the posterior teeth of article 2 are very large
and resemble those of P. variatus Barnard (1960b).

Although the gnathopods of the specimens at hand are stouter
than those of P. variatus, the enlarged teeth of pereopod 5 on some
individuals demonstrate the close relationship of the P. fatigans
complex with P. variatus. Barnard (1960b) has already considered
the possibility that P. fatigans is a phenotype of P. variatus, but a
clarification of the problem is complicated by the additional dif-
ferences displayed by the specimens from Monterey Bay.

The Monterey fatigans complex resembles Paraphoxus epistomus,
of which only a few specimens have been found in the present samples.
Paraphozus epistomus may be distinguished by the horizontal ventral
edge (either truncate or slightly convex) of article 2 on pereopod 5
and by the slightly stouter gnathopods. A few specimens assigned
to P. fatigans show a relationship to P. epistomus because of the
peduncular setosity of uropod 2. Most of these specimens have the
elongated epistomal process.

Distribution: Monterey Bay to Todos Santos Bay, Baja California,
12-162 m.

Paraphoxus obtusidens (Alderman)

Paraphoxus obtusidens.—Barnard, 1960b, pp. 249-259, pls. 33-37; 1964a, p. 244.
Material: 6429 (1), 6430 (1), 6431 (6), 6432 (8), 6433 (5), 6435

(1), 6438 (1), 6439 (1), 6444 (1), 6453 (1), 6469 (3), 6471 (5).
Distribution: Kurile Islands to Columbia, South America, 0-180 m.

Paraphoxus robustus Holmes

Paraphoxus robustus—Barnard, 1960b, pp. 235-236, pl. 25; 1964a, p. 244.

Material: 6431 (1), 6432 (2), 6433 (4), 6466 (1), 6471 (1), 6474 (4).

Distribution: Puget Sound, Washington to San Quintin Bay, Baja
California, 4-183 m.

30 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Paraphoxus similis Barnard
Paraphozus similis Barnard, 1960b, pp. 230-233, pls. 22, 23.

Material: 6431 (2), 6432 (6), 6438 (3), 6444 (11), 6456 (1), 6458
(11), 6459 (1), 6465 (10), 6466 (6), 6474 (17), 6477 (13), 6478 (4),
6480 (2).

Distribution: Puget Sound, Washington, to southern California,
32° N, 31-324 m, abundant on the shelf of southern California in
depths of 55-110 m.

Paraphoxus variatus Barnard

Paraphocus variatus Barnard, 1960b, pp. 198-202, pls. 3, 4; 1964a, p. 245.

Material: 6426 (14), 6428 (2), 6446 (2), 6447 (8), 6450 (17).

Distribution: Monterey Bay to San Ramon Bay, Baja California,
5-93 m.

Phoxocephalus homilis Barnard

Phozocephalus homilis Barnard, 1960b, p. 301, pls. 49, 50; 1964a, p. 245.

Material: 6433 (1), 6456 (10), 6471 (12), 6474 (51), 6477 (9),
6480 (5).

Distribution: Monterey Bay to San Cristobal Bay, Baja Cali-
fornia, 62-200-+ m.

Family Pleustidae

Parapleustes pugettensis (Dana)

Parapleustes pugettensis——Barnard and Given, 1960, pp. 43-45, fig. 4 (with
synonymy).—Shoemaker, 1964, pp. 410-413, fig. 10.

Material: 6445 (1), 25 m.

Distribution: West coast of Alaska, 62° N to southern California,
32° N, where it is moderately abundant in the intertidal zone and
on shallow water epifloras.

Family Podoceridae

Pedocerus cristatus (Thomson)
Podocerus cristatus—Barnard, 1962a, pp. 67-69, figs. 31, 32; 1964a, p. 246.

Material: 6425 (11).

Distribution: Probably cosmopolitan in the Indo-Pacific tropical
and warm-temperate regions, southwest Africa, New Zealand, Hawaii,
Australia, in the eastern Pacific Ocean from Monterey Bay to Turtle
Bay, Baja California, 0-171 m.

Family Stenothoidae
Metopella aporpis Barnard
Metopella aporpis Barnard, 1962c, pp. 142-145, figs. 12, 13; 1964a, p. 246.
Material: 6425 (1), 24 m.

NO. 3541 BENTHIC AMPHIPODA—BARNARD 31

Distribution: Monterey Bay to San Cristobal Bay, Baja Cali-
fornia, 24-140 m, south of Point Conception not shallower than 84 m.

Stenothoe frecanda Barnard
Stenothoe frecanda Barnard, 1962c, p. 151, fig. 18.

Material: 6471 (1)
Distribution: Monterey Bay to southern California shelf, 64-92 m.

Family Synopiidae (=Tironidae)
Tiron biocellata Barnard
Tiron biocellata Barnard, 1962b, p. 75, fig. 2; 1964a, p. 247.

Material: 6440 (1).
Distribution: Monterey Bay to San Cristobal Bay, Baja California,
11-180 m.
Literature Cited

ALDERMAN, A. L.

1936. Some new and little known amphipods of California. Univ. California

Publ. Zo6l., vol. 41, pp. 53-74, figs. 1-51.
BaRNARD, J. L.

1952. Some Amphipoda from central California. Wasmann Journ. Biol.,
vol. 10, pp. 9-36, pls. 1-9.

1954a. Amphipoda of the family Ampeliscidae collected in the eastern
Pacific Ocean by the Velero III and Velero IV. Allan Hancock
Pacific Exped., vol. 18, no. 1, 137 pp., pls. 1-38.

1954b. Marine Amphipoda of Oregon. Oregon State Monogr. Stud. Zool.,
no. 8, pp. 1-103, pls. 1-33, fig. 1.

1955a. Gammaridean Amphipoda (Crustacea) in the collections of Bishop
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1955b. Notes on the amphipod genus Aruga with the description of a new
species. Bull. Southern California Acad. Sci., vol. 54, pp. 97-103,
pls. 27-29.

1957. A new genus of haustoriid amphipod from the northeastern Pacific
Ocean and the southern distribution of Urothoe varvarini Gurjanova.
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1958. A new genus of dexaminid amphipod (marine Crustacea) from
California. Bull. Southern California Acad. Sci., vol. 56, pp.
130-1382, pls. 26, 27.

1959a. Estuarine Amphipoda. Jn Ecology of Amphipoda and Polychaeta of
Newport Bay, California. Allan Hancock Found. Publ. Occas.
Pap. 21, pp. 13-69, pls. 1-14.

1959b. Liljeborgiid amphipods of southern California coastal bottoms, with
a revision of the family. Pacific Nat., vol. 1, no. 4, pp. 12-28, figs.
1-12.

1959c. The common pardaliscid Amphipoda of southern California, with a
revision of the family. Pacific Nat., vol. 1, no. 12, pp. 36-43, figs.
1-4,

1960a. New bathyal and sublittoral ampeliscid amphipods from California,
with an illustrated key to Ampelisca. Pacific Nat., vol. 1, no. 16,
pp. 1-36, figs. 1-11.

32 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

1960b. The amphipod family Phoxocephalidae in the eastern Pacific Ocean,
with analyses of other species and notes for a revision of the family.
Allan Hancock Pacific Exped., vol. 18, no. 3, pp. 175-368, pls. 1-75.

1961. Relationship of California amphipod faunas in Newport Bay and in
the open sea. Pacific Nat., vol. 2, no. 4, pp. 166-186, figs. 1-2.

1962a. Benthic marine Amphipoda of southern California: Families Aoridae,
Photidae, Ischyroceridae, Corophiidae, Podoceridae. Pacific Nat.
vol. 3, no. 1, pp. 1-72, figs. 1-32.

1962b. Benthic marine Amphipoda of southern California: Families Tironidae
to Gammaridae. Pacific Nat., vol. 3, no. 2, pp. 73-115, figs. 1-23.

1962c. Benthic marine Amphipoda of southern California: Families Amphi-
lochidae, Leucothoidae, Stenothoidae, Argissidae, Hyalidae. Pa-
cific Nat., vol. 3, no. 3, pp. 116-163, figs. 1-23.

1962d. South Atlantic abyssal amphipods collected by R.V. Vema. In
Abyssal Crustacea, Vema Res. Ser., no. 1, pp. 1-78, figs. 1-79.

1962e. Benthic marine Amphipoda of southern California: Family Oedicero-
tidae. Pacific Nat., vol. 3, no. 12, pp. 349-371, figs. 1-10.

1962f. A new species of sand-burrowing marine Amphipoda from California.
Bull. Southern California Acad. Sci., vol. 61, pp. 249-252, figs. 1-2.

1963. Relationship of benthic Amphipoda to invertebrate communities of
inshore sublittoral sands of southern California. Pacific Nat.,
vol. 3, no. 15, pp. 437-467, figs. 1-7.

1964a. Los anfipodos bentonicos marinos de la costa occidental de Baja
California. Rev. Soc. Mexicana Hist. Nat., vol. 24, pp. 205-274,
figs. 1-11.

1964b. Marine Amphipoda of Bahia de San Quintin, Baja California. Pa-
cific Nat., vol. 4, no. 3, pp. 55-139, figs. 1-21.

In press. Intertidal Amphipoda of California: Monterey to La Jolla.

BaRNARD, J. L., and Givmen, R. R.

1960. Common pleustid amphipods of southern California, with a projected
revision of the family. Pacific Nat., vol. 1, no. 17, pp. 37-48,
figs. 1-6.

BARNARD, J. L., and JONES, G. F.

1960. Techniques in a large scale survey of marine benthic biology. In

Pearson, Waste disposal in the marine environment, pp. 413-447.
BARNARD, J. L., and ZIESENHENNE, F. C.

1961. Ophiuroid communities of southern Californian coastal bottoms.

Pacific Nat., vol. 2, no. 2, pp. 131-152, figs. 1-8.
BovusFIELp, E. L.

1958. Notes on the amphipod genus Orchestoidea on the Pacific coast of
North America. Bull. Southern California Acad. Sci., vol. 56,
pp. 119-129, pls. 24, 25.

ButycuHeva, A. I.

1952. Novye vidy bokoplavov (Amphipoda, Gammaridea) iz Japonskogo
morja. Trudy Zool. Inst. Akad. Nauk SSSR, vol. 12, pp. 195-250,
figs. 1-39. [In Russian.]

CALIFORNIA COOPERATIVE FISHERIES INVESTIGATIONS
1952-64. California Cooperative Oceanic Fisheries Investigations Reports
[California Marine Research Committee Progress Reports],
vols. 9+-.
Catman, W. T.

1898. On a collection of Crustacea from Puget Sound. Ann. New York

Acad. Sci., vol. 11, pp. 259-292, pls. 31-34.

NO. 3541 BENTHIC AMPHIPODA—BARNARD 33

CHEVREUX, E., and Faas, L.

1925. Amphipodes. Faune de France, vol. 9, pp. 1-488, figs. 1-438.
EKMAN, S.

1953. Zoogeography of the sea, pp. i-xiv + 1-417, figs. 1-121.
GartTH, J.S.

1955. The case for a warm-temperate marine fauna on the west coast of
North America: Essays in the natural sciences in honor of Captain
Allan Hancock, on the occasion of his birthday, July 26, 1955,
pp. 19-27.

Gursanova, E. F.

1938. Amphipoda, Gammaroidea [sic] of Siaukhu Bay and Sudzukhe Bay
(Japan Sea). Jn Reports of the Japan Sea Hydrobiological Ex-
pedition of the Zoological Institute of the Academy of Sciences of
the USSR in 1934, pt. 1, pp. 241-404, figs. 1-59. [In Russian
with English summary.]

1951. Bokoplavy morej SSSR isopredel’nykh vod (Amphipoda-Gammaridea) .
Opredel. Faune SSSR Akad. Nauk SSSR, vol. 41, pp. 1-1029,
figs. 1-705. [In Russian.]

1953. Novye dopolnenija k dal’nevostochnoi faune morskik bokoplavov.
Trudy Zool. Inst. Akad. Nauk SSSR, vol. 18, pp. 216-241, figs.
1-19. [In Russian.]

1962. Bokoplavy severnoi chasti Tixogo Okeana (Amphipoda-Gammaridea)
chast’ 1. Opredel. Faune SSSR Akad. Nauk SSSR, vol. 74, pp.
1-440, figs. 1-143. [In Russian.]

Hartman, O.

1956. Contributions to a biological survey of Santa Monica Bay, California,
161 pp. [Mimeographed report to Hyperion Engineers Inc. by
Geology Dept., Univ. Southern California, Los Angeles.]

Homes, 8. J.

1908. The Amphipoda collected by the U.S. Bureau of Fisheries Steamer,
Albatross, off the west coast of North America, in 1903 and 1904,
with descriptions of a new family and several new genera and
species. Proc. U.S. Nat. Mus., vol. 35, pp. 489-543, figs. 1-46.

Hurtey, D. E.

1963. Amphipoda of the family Lysianassidae from the west coast of North
and Central America. Allan Hancock Found. Publ., Occas. Pap.
no. 25, pp. 1-165, figs. 1-49.

Jones, G. F. and Barnarp, J. L.

1963. The distribution and abundance of the inarticulate brachiopod
Glottidia albida (Hinds) on the mainland shelf of southern Cali-
fornia. Pacific Nat., vol. 4, pp. 27-52, figs. 1-14.

Mitts, E. L.

1961. Amphipod crustaceans of the Pacific coast of Canada, 1: Family
Atylidae. Nat. Mus. Canada Bull. 172, pp. 13-33, figs. 1-6.

1962. Amphipod crustaceans of the Pacific coast of Canada, 2: Family
Oedicerotidae. Nat. Hist. Paps. Nat. Mus. Canada, vol. 15, pp.
1-21, figs. 1-6.

Nagata, K.

1960. Preliminary notes on benthic gammaridean Amphipoda from the
Zostera region of Mihara Bay, Seto Inland Sea, Japan. Publ. Seto
Mar. Biol. Lab., vol. 8, pp. 163-182, figs. 1-2, pls. 13-17.

34 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Sars, G. O.
1895. Amphipoda. Vol. 1 of An account of the Crustacea of Norway with

short descriptions and figures of all the species, viii + 711 pp., 240
pls., 8 suppl. pls.
SHOEMAKER, C. R.
1925. The Amphipoda collected by the United States Fisheries Steamer
Albatross in 1911, chiefly in the Gulf of California. Bull. American
Mus. Nat. Hist., vol. 52, pp. 21-61, figs. 1-26.
1931. Anew species of amphipod crustacean (Acanthonotozomatidae) from
California, and notes on Eurystheus tenuicornis. Proc. U.S. Nat.
Mus., vol. 78, pp. 1-8, figs. 1-4.
1942. Amphipod crustaceans collected on the presidential cruise of 1938.
Smithsonian Misc. Coll., vol. 101, no. 11, pp. 1-52, figs. 1-17.
1955. Amphipoda collected at the arctic laboratory, Office of Naval Research,
Point Barrow, Alaska, by G. E. Macginitie. Smithsonian Misc.
Coll., vol. 128, no. 1, pp. 1-78, figs. 1-20.
1964. Seven new amphipods from the west coast of North America with
notes on some unusual species. Proc. U.S. Nat. Mus., vol. 115,
pp. 391-4380, figs. 1-15.
STEBBING, T. R. R.
1906. Amphipoda, 1: Gammaridea. Pt. 21 in Das Tierreich, pp. 1-806,
figs. 1-127.
STEPHENSEN, K.
1932. Some new amphipods from Japan. Annot. Zool. Japonensis, vol. 13,
pp. 487-501, figs. 1-5.
THORSTEINSON, E. D.
1941. New or noteworthy amphipods from the North Pacific Coast. Univ.
Washington Publ. Oceanogr., vol. 4, pp. 50-96, pls. 1-8.
THORSON, G.
1957. Bottom communities. Ch. 17 in vol. 1 of Hedgpeth et al., Treatise on
marine ecology and paleoecology.
WALKER, A. O.
1898. Crustacea collected by W. A. Herdman, F.R.S., in Puget Sound,
Pacific coast of North America, September 1897. Trans. Liverpool
Biol. Soc., vol. 12, pp. 268-287, pls. 15, 16.

NO. 3541 BENTHIC AMPHIPODA—BARNARD 35

*6480 +6479 lleac2 - 6460

ico. «= 6474 «#6477 «+6478 fh G464 +6459

6471 °*6470 ¢6466 *6465 #6463 ° 6458

[

6435 —« 6433+ 6432—* 6438—-- 6431» 6430—« 6447

©6436 © 6437 4 6439/ ©6429 «6427
50
40

e 6428’ «6426

wo

[=| srarion|
= [Beam Fas

36°35 'N

MONTEREY

12|°60 W
121°55' Ww

36°35'N

Ficure 1.—Upper, station locations in Monterey Bay; lower, distribution of community
dominants.

36 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Ficure 2.—Ampelisca lobata Holmes, female, 4.0 mm, sta. 6425: a, pereopod 5. Rhacho-
tropis oculata (Hansen), male, 8.0 mm, sta. 1149 (southern California): b, dorsally toothed
segments, left to right, pereonal 7, pleonal 1-4; c, peduncle of antenna 1; d, head; e, cuticu-
lar scales of chitin; f, pereopod 5.

NO. 3541 BENTHIC AMPHIPODA—BARNARD 37

8

Fo
1 rr]
Si 1 T (oS

———

V2

Ficure 3.—Protomedeia penates, new species, White Gulch, Tomales Bay, female, 7.0 mm:
a, lateral view; b, antenna 1 to show accessory flagellum; c, article 5 of pereopod 4; d-f,
uropods 1, 2, 3; g, telson; h, pleonal segment 2; 7, base of peduncle of antenna 234, articles
2-3 of pereopod 5; k,/, gnathopods 1, 2; m, head; n,o, articles 6-7 of gnathopods 1, 2.

Male, 6.0 mm: 9,9, gnathopods 2, 1; r,s, palms of gnathopods ee

38 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

{

Ficure 4.—Anonyx adoxus Hurley, female, 10.0 mm, sta. 6432: a, head; 6, antenna 1;
c,d, gnathopod 1; e,f, gnathopod 2; g—k, pereopods 1, 2, 3, 4, 5; 1,m, uropods 1, 2; n,o, uropod
3 with and without setae; p, telson; g, pleonal epimera 1-3, left to right, right side of
pleonal epimeron 1 offset.

|
|
|
|

Figure 5.—Centromedon pavor, new species, holotype, female, 2.9 mm, sta. 6459: a, lateral
view; b,c, gnathopods 1, 2; d, antenna 1; e-g, uropods 1, 2, 3; h, telson; 71, pereopod 1.

40 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

Ficure 6.—Paraphoxus fatigans Barnard, male, 2.5 mm, sta. 6448, variant: a, pereopod 5;
b, epistome. Female, 2.8 mm, sta. 6493, kind 2: ¢, pereopod 5. Male, 3.1 mm, sta. 6426,
kind 1: d, pereopod 5. Male, 3.5 mm, sta. 6448, variant: ¢, pereopod 5; f, head; g, epi-
stome. Female, 3.5 mm, sta. 6448, variant: h, head; 7, pereopod 5; 7, epistome. Female,
3.8 mm, sta. 6426, kind 1:k, pereopod 5. Female, 3.0 mm, sta. 6454, kind 1:2, pereopod 5.

EXPLANATION OF FicurE 7
Ficure 7.—Paraphoxus fatigans Barnard, female, 3.0 mm, sta. 6454, kind 1: 4,b, gnathopod 1.
Female, 3.5 mm, sta. 6448, variant: c,d, gnathopod 1. Male, 2.5 mm, variant: e,f,
gnathopod 1. Male, 3.1 mm, sta 6426, kind 1: g,h, gnathopod 1. Male 3.5 mm, sta.
6448, variant: 1,7, gnathopods 1, 2. Female, 2.8 mm, sta. 6493, kind 2: k, gnathopod 1.

NO. 3541

BENTHIC AMPHIPODA—BARNARD

4]

Figure 7.—Explanation on facing page.

U.S. GOVERNMENT PRINTING OFFICE: 1966

mrocee din ss ot

the United States

National Museum

SMITHSONIAN INSTITUTION + WASHINGTON, D.C.

Volume 119 1966 Number 3542

A NEW GENUS AND SIX NEW SPECIES
OF ENTOCYTHERID OSTRACODS
(OSTRACODA, ENTOCYTHERIDAE)

By Horton H. Hosss, Jr., and MARGARET WALTON *

The conservatism that marked the supraspecific treatment of this
group of epizootic ostracods for 58 years was broken with the estab-
lishment of the family Entocytheridae by Howe (1961) and the ap-
pearance of a familial revision in the following year by Hart (1962).
Even with the elevation of Hoff’s (1942) subfamily Entocytherinae
in 1961, only two genera were recognized. In his generic revision,
however, Hart proposed the recognition of two subfamilies containing
11 genera. We have followed his recommendations and are adding
an additional genus to the subfamily Entocytherinae.

At the time of the preparation of this manuscript, descriptions of
at least three more genera of entocytherids are in press. Therefore,
it seems somewhat futile to offer a key for the separation of established
genera that will be obsolete prior to its publication; furthermore,
there are undescribed species available to those currently studying
these animals which will require the erection of additional genera.

The assemblage of species described here was collected from various
areas in the United States, and all of them were associated with cray-
fishes. The ostracod genera represented are Dactylocythere, Don-

1 Hobbs: Department of Zoology, Smithsonian Institution; Walton: 1116 Rich-

mond Blvd., Danville, Va.

1
219-940—66

2 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

naldsoncythere, Entocythere, the new genus Thermastrocythere, and
Unicnocythere.

We wish to thank the following persons who have made available
to us the collections on which this paper is based: Messrs. J. W. Berry,
C. W. Hart, Jr., J. M. Stubbs, Drs. Fulton Fite, P. C. Holt, L. R.
McManus, G. C. Smart, P. J. Spangler, and W. 8. Woolcott.

Genus Dactylocythere Hart, 1962
Dactylocythere brachystrix, new species
Figures la-d

Maur.—Eye present. Shell (fig. 16) with highest portion posterior
to midlength and with dorsal and ventral margins more gently
tapered anteriorly than posteriorly; ventral margin of shell entire;
submarginal setae anteriorly, ventrally, and posteriorly somewhat
evenly spaced, absent dorsally. Range of shell size of 10 specimens
recorded in table 1.

Copulatory complex (figs. la, d) with posterior margin of peniferum
expanded into three humps; both anteroventral and posteroventral
extremities directed anteroventrally. Penis small, its length approxi-
mately one-third the anterior-posterior dimension of distal portion
of peniferum, and situated at level of base of ventral two-fifths of
peniferum. Accessory groove extending dorsally almost to ventral
extremity of spermatic duct. Dorsal finger slender with its terminal
seta extending to ventral margin of finger guard; ventral finger gently
curved throughout its length. Finger guard prominent with its
anterior margin straight and its ventral margin emarginate with
three small prominences. Clasping apparatus with vertical and
horizontal rami somewhat distinctly delimited; vertical ramus with
a bend of about 45° just proximal to midlength and joining horizontal
ramus (external borders) at an angle of about 100°; internal border
or horizontal ramus with three evenly spaced teeth and apex with
three denticles; external borders of both rami entire.

FrmMa.e.—Shell of triunguis female (fig. 1c) distinctly larger than
that of male, with its greatest height slightly posterior to midlength
and distinctly lower anteriorly than posteriorly; ventral margin entire;
marginal setae as in male. Prominent J-shaped rod and ruffled
amiculum present. Range of shell size of 8 specimens recorded in
table 1.

TYPE-LOCALITY AND RANGE.—Otter Creek, Cumberland County,
Tenn., approximately 3.5 miles upstream from the Obed River on
the Cumberland Plateau. This species is known only from the type
locality where it was found associated with Dactylocythere pachy-
sphyrata, new species.

NO. 3542 OSTRACODS—-HOBBS AND WALTON 3

Typres.—The holotypic male, the allotypic female, and a dissected
paratypic male are deposited in the U.S. National Museum (nos.
111251, 111252). Paratypes are in the collections of C. W. Hart, Jr.,
and the joint collection of the authors.

Hosr.—An undescribed species of the genus Cambarus.

RELATIONSHIPS.—Dactylocythere brachystriz is closely related to
D. daphnioides (Hobbs, 1955), D. runki (Hobbs, 1955), D. chalaza
(Hobbs and Walton, 1962), and D. pachysphrata, new species, but
differs from all of them in possessing an accessory groove on the pen-
iferum which extends dorsally only to the ventral extremity of the
loop of the spermatic duct. In the other species, the groove reaches
the dorsal extremity of the loop.

Remarks.—The name brachystriz is derived from the Greek Bpaxus,
meaning short, and from orpé, meaning groove, referring to the short
accessory groove of the peniferum of the male.

Dactylocythere pachysphyrata, new species

FIGureEs le-g

Ma.e.—Eye present. Shell (fig. 1g) highest posterior to midlength
and with dorsal and ventral margins more gently tapered anteriorly
than posteriorly; submarginal setae limited largely to anterior,
anterodorsal, posterior, and posterodorsal areas, nowhere abundant,
rare ventrally, and absent dorsally. Range of shell size of eight
specimens recorded in table I.

Copulatory complex (figs. 1e, f) with posterior margin of peniferum
entire; anteroventral extremity directed ventrally and posteroventral
extremity directed anteriorly. Penis small, its length about one-
fourth of anterior-posterior dimension of distal portion of peniferum,
and situated at level of base of ventral two-fifths of peniferum.
Accessory groove extending dorsally to level of dorsalmost extremity
of loop of spermatic duct. Dorsal finger moderately stout with its
terminal seta extending to ventral margin of finger guard; ventral
finger slender, curved throughout its length, somewhat strongly so
at base of proximal two-fifths. Finger guard prominent with its
anterior margin straight and its ventral margin slightly emarginate
but without distinct prominences. Clasping apparatus with vertical
and horizontal rami indistinctly delimited by thickened ‘“ankle’’;
vertical ramus with a bend of about 30° at distal end of proximal two-
fifths; internal border of vertical ramus inclined anteroventrally at
approximately a 45° angle to main axis; internal border of horizontal
ramus with three teeth; apex of clasping apparatus with three denti-
cles; external borders of both rami entire, their posteroventral ex-
tensions forming a right angle.

Fremate.—Unknown.

4 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

TYPE-LOCALITY AND RANGE.—Otter Creek, Cumberland County,
Tenn., approximately 3.5 miles upstream from the Obed River on
the Cumberland Plateau. This species is known only from the type
locality where it was found associated with Dactylocythere brachystriz.

Typrs.—The holotypic male and a dissected male paratype are
deposited in the U.S. National Museum (nos. 111253 and 111254).
Paratypes are in the collections of C. W. Hart, Jr., and the joint col-
lection of the authors.

Host.—An undescribed species of the genus Cambarus.

RELATIONSHIPS.—Dactylocythere pachysphyrata is related to the same
group of species as is D. brachystriz. While they share a number of
features in common, the most conspicuous is the thickened “ankle”
of the clasping apparatus. It is less closely allied to D. steevesi (Hart
and Hobbs, 1961) in which the vertical ramus of the clasping apparatus
is more uniformly thickened. It may be distinguished from daph-
nioides by lacking a posteroventral extension of the shell; from
chalaza by possessing more than one tooth on the internal border of
the clasping apparatus; and from runki by possessing a much more
prominently thickened “ankle.”

Remarxs.—The name pachysphyrata is derived from the Greek
maxus, Meaning thick, and cdvpov, meaning ankle, referring to the
thickened junction of the vertical and horizontal rami of the clasping
apparatus of the male.

Genus Donnaldsoncythere Rioja, 1942
Donnaldsoncythere cayugaensis, new species
Figures 1lh-k

Maue.—KHye present. Shell (fig. 17) with highest portion posterior
to midlength; dorsal margin tapering more gently anteriorly than
posteriorly, and ventral margin entire; submarginal setae somewhat
evenly, if widely, spaced around entire perimeter of shell. Range of
shell size of 10 specimens recorded in table 1.

Copulatory complex (figs. 17, #) with peniferum terminating distally
in a bilobed rounded prominence; anteroventral margin of posterior
lobe thickened. Dorsal and ventral fingers slender and almost straight
with terminal seta of dorsal finger reaching level of penis guides on
peniferum. Clasping apparatus with distal portion directed at about
a 45° angle to basal portion and not clearly divisible into vertical and
horizontal rami; margins entire except distal portion of internal border
with two teeth and distal extremity with four denticles.

FEmMALE.—Shell of triunguis female (fig. 14) much higher than that
of male and higher posteriorly than anteriorly; ventral margin with a

NO. 3542 OSTRACODS—-HOBBS AND WALTON 5

b e

osm
LS Sculsinton

HGS

a 2mm
een}
Scale2 ii

pen
Scales
/
I 2mm 1
Scaie4
0s mm_,
Scales 5
n

Ficure 1.—Dactylocythere brachystrix: a, copulatory complex of male; b, right valve of male;
c, right valve of triunguis female; d, clasping apparatus of male. Dactylocythere pachy-
sphyrata: e, clasping apparatus of male; f, copulatory complex of male; g, right valve of male.
Donnaldsoncythere cayugaensis: h, right valve of triunguis female; 7, right valve of male;
j, clasping apparatus of male; k, copulatory complex of male. Entocythere kanawhaensis:
1, right valve of male; m, right valve of triunguis female; n, clasping apparatus of male;
0, copulatory complex of male. (Scale 1: figs. a, d, ¢, f, k, n, 0; scale 2: figs. b, c, g; scale 3:
figs. h, 1; scale 4: figs. 1, m; scale 5: fig. 7.)

219-940—66——2

6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

distinct concavity just anterior to midlength. Length of allotype
.360 mm.; height .240 mm.

TyYPE-LOCALITY AND RANGE.—Tributary to Fall Creek, 1 mile east
of Summer Hill Village, Cayuga County, N.Y., on Route 90 (elevation
1440 ft.). Fall Creek is a tributary to Cayuga Lake. Here it was
found on Cambarus robustus Girard in association with a species near
Donnaldsoncythere scalis Hobbs and Walton (1963).

Two additional localities on Salmon Creek and Fall Creek in
Tompkins County, N.Y., are in the Cayuga Lake drainage. A fourth
locality is the Salmon River, 8 miles east of Pulaski, Oswego County,
N.Y., on Route 13. The Salmon River drains directly into Lake
Ontario.

Tyres.—The holotypic male, the allotypic female, and a dissected
paratypic male are deposited in the U.S. National Museum (nos.
111255 and 111256). Paratypes are in the collections of C. W. Hart,
Jr., and the joint collection of the authors.

Host.—In all the localities cited above, this ostracod was found
on Cambarus robustus Girard.

RELATIONSHIes.—Donnaldsoncythere cayugaensis is most closely
allied to D. ileata Hobbs and Walton (1963) and differs from it chiefly
in the bilobed ventral extention of the peniferum and in the slightly
more bent clasping apparatus. On the average, it is slightly larger
than ileata, although some specimens of the latter are longer and
higher than the smallest specimen of cayugaensis.

REeMARKS.—The name cayugaensis refers to the presence of the
species in the Lake Cayuga drainage system.

Genus Entocythere Marshall (1903)

Entocythere kanawhaensis, new species
Fiaures Il-o

Ma.e.—Eye present. Shell (fig. 1/) distinctly elongate, highest
just posterior to midlength, and with dorsal margin evenly contoured,
tapering gently both anteriorly and posteriorly; ventral margin
entire; submarginal setae present except dorsally between level of
eye and anus; setae mostly regularly spaced but somewhat closer
together posteriorly. Length of holotype .615 mm., height .300 mm.;
length of paratype male .600 mm.; height .300 mm.

Copulatory complex (figs. 1n, 0) with posterior and ventral margins
of peniferum entire, the latter rounded; entire peniferum of almost
uniform thickness and without ornamentation. Penis small, its
length approximately one-fourth the anterior-posterior dimension of
distal portion of peniferum, and situated very dorsal to ventral ex-
tremity of peniferum. Dorsal finger slender with its terminal seta

NO. 3542 OSTRACODS—HOBBS AND WALTON 7

reaching level of penis; ventral finger, also slender, and sinuous.
Clasping apparatus with distinct vertical and horizontal rami meeting
at an angle of approximately 100°. Vertical ramus slightly convex
anteriorly with both margins entire; horizontal ramus slightly convex
ventrally with external border entire, internal border with three
distinct teeth and six apical denticles.

Frmaute.—shell of triunguis female (fig. 1m) proportionately higher
than that of male and larger; posterior declivity of dorsal margin
more sudden than in that of male; ventral margin entire. Sub-
marginal setae as in male. Length of allotype .698 mm., height .398
mm.

TYPE-LOCALITY AND RANGE.—Rapids in New River at Pembroke,
Giles County, Va., where it was associated with Donnaldsoncythere
ileata Hobbs and Walton (1963).

Two additional specimens were taken from the Cascades on Little
Stony Creek, also in Giles County. Here it was associated with
D. ileata, D. scalis Hobbs and Walton (1963), and Phymocythere phyma
(Hobbs and Walton, 1962).

Typrs.—The holotypic male and the allotypic female are deposited
in the U.S. National Museum (no. 111257). A paratype male is in
the joint collection of the authors.

Hosts.—At the type locality, the specimens were taken from
Cambarus sciotensis Rhoades. At the Cascades the specimen was
found in a collection of crayfishes including C. sciotensis and C. 6.
bartonii (Fabricius).

RELATIONSHIPS.—Entocythere kanawhaensis seems to have its
closest affinities with EF. dorsorotunda Hoff (1944) and E. elliptica
Hoff (1944) but differs from both species by possessing only three
teeth along the internal border of the horizontal ramus of the clasping
apparatus of the male.

ReMARKS.—The name kanawhaensis refers to the fact that the
species is an inhabitant of the Kanawha River drainage system.

Thermastrocythere, new genus

Dracnosis.—Terminal tooth of mandible with cusps. Copulatory
complex of male without finger guard; ventral portion of peniferum
appearing to be deeply incised (actually apparent rami at least
partially ensheathed by delicate membrane), its posterior ‘‘ramus’’
slender and curved with apex directed anteriorly, its apparent anterior
“ramus” also curved with tip directed ventrally—the two rami ap-
pearing to oppose one another; anterior ‘‘ramus’’ consisting of the
prominent penis; penis large, its length subequal to anterior-posterior
dimension of distal portion of peniferum. Clasping apparatus ex-
tending ventrally beyond peniferum with distinct slender vertical

8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

and horizontal rami, the former one-third longer than latter; extensions
of rami forming an angle of approximately 80° to 85°; internal and
external borders of vertical ramus and external border of horizontal
ramus entire; internal border of horizontal ramus with two teeth, one
just proximal to midlength and the other subapical; apex with four
denticles.

Typn-species.— Thermastrocythere harti, new species.

Remarxs.—The name Thermasirocythere is derived from the Greek
fepuaorps, Meaning tongs or pincers, referring to the forcipate
appearance of the ventral portion of the peniferum.

Thermastrecythere haréi, new species
Figures 2e-g

Maue.—Eye present. Shell (fig. 2e) with highest portion slightly
posterior to midlength and with dorsal margin more gently tapering
posteriorly than anteriorly; ventral margin of shell entire; submarginal
setae present anteriorly, posteriorly, and ventrally but absent dorsally,
anterior and anteroventral setae closer together than posterior and
posteroventral ones. Range of shell size of 10 specimens recorded
in table 1.

Copulatory complex (figs. 2f, g) as described in the diagnosis. In
addition, dorsal finger slender with tip of apical seta reaching level of
base of penis; ventral finger with moderate curve near base and a less
prominent one a short distance proximal to base of terminal seta.

FemaLe.—Female unknown.

TYPH-LOCALITY, RANGE, AND HoSTs.—Cache Creek at White Wolf
Crossing, Fort Sill, Comanche County, Okla., on Orconectes nais
(Faxon).

Additional localities include: Six Mile Creek near Waunakee, Dane
County, Wis., on Orconectes p. propinguus (Girard) and 0. virilis
(Hagen): McSpadin Falls, 10 miles northeast of Talequah on Route
10, Cherokee County, Okla., on Orconectes meeki brevis Williams, 0.
nana Williams, and QO. neglectus neglectus (Faxon); small stream, 13
miles west of Madison, Dane County, Wis., on O. p. propinquus and
O. virilis; Yahara River near Sun Prairie where it crosses Route 19,
Dane County, Wis., on O. p. propinquus and in association with
Entocythere cambaria Marshall; White Pines State Forest, Ogle County,
Ill., on O. virilis; and 5.9 miles west of Eyota on Route 14, Olmsted
County, Minn., on Cambarus d. diogenes Girard and O. virilis, in
association with Rhadinocythere serrata (Hoff).

Types.—The holotypic male and two paratypic males are deposited
in the U.S. National Museum (nos. 111258 and 111259). Paratypes

NO. 3542 OSTRACODS—-HOBBS AND WALTON 9

are in the collections of C. W. Hart, Jr., and the joint collection of
the authors.

RELATIONSHIPS.— Thermastrocythere harti has its closest affinities
with members of the genus Uncinocythere (Hart, 1962) and is perhaps
more closely allied to U. simondsi (Hobbs and Walton, 1960) than to
any other species; similarities exist in the armature of the clasping
apparatus and in the distal portion of the peniferum; however, the
penis in harti reaches the anteroventral extremity of the peniferum—
an arrangement that does not exist in any known species of the genus
Uncinocythere.

Remarks.—We are pleased to name this species in honor of our
friend and colleague, C. W. Hart, Jr., who has contributed much to
our knowledge of the entocytherids.

Genus Uncinocythere Hart, 1962

Uncinocythere stubbsi, new species

Figures 2a-d

Maue.—Bye present. Shell (fig. 2b) highest near midlength and
with dorsal margin evenly contoured anteriorly and posteriorly;
ventral margin convex without an emargination; submarginal setae
evenly but widely spaced anteriorly, ventrally, and posteriorly, absent
dorsally. Range of shell size of 10 specimens recorded in table 1.

Copulatory complex (figs. 2a, d) with posterior margin of peniferum
entire, its ventral margin excised with anteroventral extremity directed
ventrally and posteroventral extremity directed anteroventrally;
anteroventral portion with a heavy, acute, dorsally directed penis
guide. Penis of moderate size, its length approximately one-half
anterior-posterior dimension of distal portion of peniferum and situated
far distal to base of clasping apparatus within ventral one-third of
area between dorsal margin of spermatic loop and ventral extremity
of peniferum. Dorsal and ventral fingers slender, latter with strong
caudal bend at distal end of proximal one-third. Clasping apparatus
with vertical and horizontal rami of subequal lengths, and extensions
of their rounded junction forming angle of less than 90°. Vertical
ramus entire but with distal one-third curved posteriorly. External
border of horizontal ramus gently rounded and entire; internal border
with three teeth—largest immediately proximal to midlength and
two smaller ones near distal extremity; terminal extremity with
three teeth.

FrEMALE.—Shell of triunguis female (fig. 2c) distinctly larger than
that of male with its greatest height slightly posterior to midlength
and distinctly lower anteriorly than posteriorly; ventral margin with

10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

emargination near midlength; submarginal setae as in male. Range
of shell size of 10 specimens recorded in table 1.

TYPE-LOCALITY AND RANGE.—A tributary of Big Turnbull Creek
(to Harpeth and Cumberland Rivers) on the property of Bethany
Hills Church Camp in the southern portion of Cheatham County,
Tenn.

TaRLE 1.— Measurements (in mm.) of shells

Sex Length Height Length/Height
Dac. brachystrix
Mean a . 473+. 009 . 273+. 008 1. 73+. 031
Minimum d . 465 . 263 Gu
Maximum J . 488 . 293 leeaie
Mean Q . 483+. 022 . 296+. 012 1. 63+. 030
Minimum 9 . 442 . 210 1. 59
Maximum 2 . 510 . O29 1. 70
Dac. pachysphyrata
Mean J . 486+. 004 . 2538+. 015 1. 73+. 039
Minimum a . 428 . 240 1. 68
Maximum J . 443 200 1S
Don. cayugaensis
Mean J . 450+. 008 . 248+. 007 1. 82+. 062
Minimum oS . 435 . 240 ea
Maximum J . 460 . 260 1. 92
T. harti
Mean J . 358+. 013 . 201+. O11 1. 78+. 093
Minimum J . 338 . 188 1. 65
Maximum J . 390 . 218 1. 92
U. stubbsi
Mean a . 281+. 008 . 161+. 003 1. 75+. 067
Minimum J . 270 S158 1. 64
Maximum J . 293 . 165 1. 85
Mean Q . 324+. 009 . 189+. 005 1. 74+. 113
Minimum Q oly . 180 1. 62
Maximum 2 . 845 . 195 1. 83

This species is known only from the type locality.

Typxs.—The holotypic male, the allotypic female, and a dissected
paratypic male are deposited in the U.S. National Museum (nos.
111260 and 111261). Paratypes are in the collections of C. W. Hart,
Jr., and the joint collection of the authors.

Host.—Orconectes rusticus subsp. (?)

Revationsuies.—Uneinocythere stubbsi has its closest affinities
with those species of the genus having a distinct bifid ventral (or
posteroventral) margin of the peniferum: U. ericksoni (Kozloff, 1955),
U. caudata (Kozloff, 1955), U. neglecta (Westervelt and Kozloff, 1959),

!

NO. 3542 OSTRACODS—HOBBS AND WALTON E1

U. simondsi (Hobbs and Walton, 1960), and U. pholetera (Hart and
Hobbs, 1961). It differs from U. caudata in lacking a posteroventral
acute projection of the shell; from U. ericksoni and U. neglecta in
possessing only three teeth at the extremity of the ventral ramus of
the clasping apparatus; from U. pholetera in having the vertical and
horizontal rami of the clasping apparatus subequal in length and with
the distal two teeth of the horizontal ramus closer together, the distal

2m
[= scaleea
03 mm_,
Scaler, 7

ae 2m
arscateue
k -O2mm {
Scale9Q

d

Ficure 2.—Uncinocythere stubbsi: a, copulatory complex of male; }, right valve of male;
c, right valve of female; d, clasping apparatus of male. Thermastrocythere harti: e, right
valve of male; f, copulatory complex of male; g, clasping apparatus of male.

one more distally situated than in pholetera; and from U. simondsi
in having the penis situated in the distal third of the area between
dorsal margin of the spermatic loop and the ventral margin of the
peniferum. It differs from all of its relatives in the arrangement of
the penis guide.

Remarxs.—It is a pleasure to name this species in honor of
John M. Stubbs, of the Tennessee Game and Fish Commission, who
has so generously aided us in our studies of crayfishes and their
epizootic ostracods in Tennessee.

12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 119

Literature Cited
Hart, C. W., JR.

1962. A revision of the ostracods of the family Entocytheridae. Proc.

Acad. Nat. Sci. Philadelphia, vol. 114, no. 3, pp. 121-147, 18 figs.
Hart, C. W., Jz., and Honrs, Horton H., Jr.

1961. Hight new troglobitic ostracods of the genus Entocythere (Crustacea,
Ostracoda) from the eastern United States. Proc. Acad. Nat. Sci.
Philadelphia, vol. 113, no. 8, pp. 173-185, 32 figs.

Hosgss, Horton H., Jr.

1955. Ostracods of the genus Entocythere from the New River system in
North Carolina, Virginia, and West Virginia. Trans. Amer.
Micros. Soc., vol. 74, no. 4, pp. 325-333, 10 figs.

Hospss, Horton H., Jr., and Watton, MARGARET

1960. Three new ostracods of the genus Entocythere from the Hiwassee
drainage system in Georgia and Tennessee. Journ. Tennessee
Acad. Sci., vol. 35, no. 1, pp. 17-23, 20 figs.

1962. New ostracods of the genus Entocythere from the Mountain Lake
region, Virginia (Ostracoda, Entocytheridae). Virginia Journ.
Sci., vol. 12, no. 2, pp. 42-48, 13 figs.

1963. Four new species of the genus Donnaldsoncythere (Ostracoda, Ento-
cytheridae) from Virginia with a key to the species of the genus.
Trans. Amer. Micros. Soc., vol. 82, no. 4, pp. 363-370, 26 figs.

Horr, C. CLAYTON

1942. The subfamily Entocytherinae, a new subfamily of freshwater
cytherid ostracods, with descriptions of two new species of the
genus Entocythere. Amer. Midl. Nat., vol. 27, no. 1, pp. 63-73,
13 figs.

1944. New American species of the ostracod genus Entocythere. Amer.
Midl. Nat., vol. 32, no. 2, pp. 327-357, 33 figs.

Howe, Henry V.

1961. Entocytheridae. P. 300 zm Part Q: Anthropoda, 3: Crustacea,
Ostracoda zn Moore and Pitrat, Treatise on invertebrate paleon-
tology, xiiit442 pp. University of Kansas Press.

Koz.orr, EUGENE N.

1955. Two new species of Entocythere (Ostracoda: Cytheridae) commensal
on Pacifastacus gambelii (Girard). Amer. Midl. Nat., vol. 53,
no. 1, pp. 156-161, 24 figs.

MarsHA.L., W.S.

1903. Entocythere cambaria (nov. gen. et nov. sp.), a parasitic ostracod.
Trans. Wisconsin Acad. Sci. Arts and Letters, vol. 14, no. 1, pp.
117-144, 30 figs.

Rroya, ENRIQUE

1942. Estudios carcinologicos, 13: Consideraciones y datos acerca del
genero Entocythere (Crust. Ostracodos) y algunas de sus especies,
con descripcion de una nueva. Anal. Inst. Biol. Mexico, vol. 13,
no. 2, pp. 685, 697, 21 figs.

WESTERVELT, CLINTON A., JR., and Kozuorr, EuGENE N.

1959. Entocythere neglecta sp. nov., a cytherid ostracod commensal on
Pacifastacus nigrescens (Stimpson). Amer. Midl. Nat., vol. 61,
no. 1, pp. 239-244, 14 figs.

U.S. GOVERNMENT PRINTING OFFICE:1966

Proceedings of

the United States

National Museum

SMITHSONIAN INSTITUTION «© WASHINGTON, D.C.

Volume 119 1966 Number 3543

NEOTROPICAL MICROLEPIDOPTERA, XI!

REVISION OF GENUS IDOLATTERIA
(LEPIDOPTERA: TORTRICIDAE)

By Nicwotas S. Osraztsov ?

The genus Jdolatteria Walsingham is endemic to the Neotropical
region, and includes, as far as known, few species. They are poorly
represented in the collections, and the material available for the
present study amounted to only 15 specimens consisting of eight
species, three of which were new. No ecological information is
known concerning this genus; thus, it is not known whether the moths

1 Prepared with the aid of a National Science Foundation Grant. Previous
parts of this same series are: I and II, Clarke, 1962, Proc. U.S. Nat. Mus., vol.
113, no. 3457, pp. 373-388; III, Clarke, 1964, ibid., vol. 115, no. 3480, pp. 61-84;
IV, Duckworth, 1964, ibid., vol. 116, no. 3497, pp. 97-114; V, Obraztsov, 1964,
ibid., vol. 116, no. 3501, pp. 183-196; VI, Clarke, 1964, ibid., vol. 116, no. 3502,
pp. 197-204; VII, Obraztsov, 1966, ibid., vol. 118, no. 3527, pp. 221-232; VIII,
Duckworth, 1966, ibid., vol. 118, no. 3531, pp. 391-404; IX, Obraztsov, 1966,
ibid., vol. 118, no. 3535, pp. 577-622; X, Duckworth, 1966, ibid., vol. 119, no.
3540, pp. 1-6.

2 Deceased May 6, 1966.

2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

are actually rare in nature, or if their apparent rarity is due merely
to the fact that their habitats have not been discovered by collectors.

In their appearance the Jdolatteria species are strikingly colored
moths, superficially very similar to those of the genera Pseudatteria
Walsingham and Aiteria Walker. Especially impressive is the fact
that the resemblance is displayed even in a parallelism of the various
types of the wing pattern observed in these three genera. For
instance, Jdolatteria zanthocapna (Meyrick) might easily be confused
with Pseudatteria heliocausta (Dognin), and the two new Jdolatteria
species, J. fasciata and I. cantharopisca, could be mistaken for dwarfs
of P. cantharopa (Meyrick). Only careful examination of structural
characters can provide a definitive generic separation of the specimens
belonging to the three named genera.

The most important generic distinction is present in the genitalia,
and according to this character all three genera must be referred to
three separate tribes of the subfamily Tortricinae of the family
Tortricidae. Pseudatteria belongs to the tribe Polyorthini, Atteria
to Anacrusiini, and Jdolatteria to Archipini. Actually the wing
venation alone is satisfactory for separating the three genera. Jdolat-
teria and Pseudatteria differ from Atteria in having the veins R, and
R; of the forewing separate; in the latter genus they are stalked. In
Idolatteria the vein R; of the forewing runs to the termen, in Pseudat-
teria to the wing apex.

The author acknowledges with thanks the kind cooperation of
Mr. J. D. Bradley of the British Museum (Natural History) [BM],
Dr. J. F. Gates Clarke of the U.S. National Museum [USNM],
Dr. H. J. Hannemann of the Zoological Institut and Museum of the
Humboldt University in Berlin [ZMB], and Dr. F. H. Rindge of the
American Museum of Natural History [AMNH], who supplied the
materials for the present paper.

Genus Idolatteria Walsingham, 1913
Figures 1-3; Puates 1-8

Atteria (not Walker).—Druce, 1901, Ann. Mag. Nat. Hist., ser. 7, vol. 7, p. 440.

Pseudatteria (in part).—Meyrick, 1912, in Wagner, Lepidopterorum catalogus,
pt. 10, p. 16; 1913, zn Wytsman, Genera insectorum, fasc. 149, p. 22; 1930,
Exotic Microlepidoptera, vol. 3, p. 607.—Clarke, 1955, Catalogue of the type
specimens of Microlepidoptera in the British Museum described by Edward
Meyrick, vol. 1, p. 227, 326; 1958, ibid., vol. 3, pp. 199, 200.

Idolatieria Walsingham, 1913, in Godman and Salvin, Biologia Centrali-Ameri-
cana, Lepidoptera Heterocera, vol. 4, p. 214; 1914, ibid., vol. 4, p. 270.—
Obraztsov, 1966, Proc. U.S. Nat. Mus., vol. 118, no. 3535, p. 619.

Type species: /dolatteria simulatrix Walsingham, 1913; by monotypy
and original designation.
Head smooth, sides of vertex with cristae of longer and raised

NO. 3543 MICROLEPIDOPTERA, X1—-OBRAZTSOV 3

scales. Antenna dentate, in female with teeth shorter, in both sexes
biciliate, with setae shorter than width of antennal shaft; scapus
cylindrical. Labial palpus about three times as long as width of
eye, obliquely ascending, with apex rather porrect; basal segment
pronounced; second segment longest of all segments, smoothly scaled,
slightly dilated apicad; terminal segment about half as long as second,
tapering distad, and ending rather acutely. Proboscis moderate.
Thorax smooth.

Forewing smoothly scaled, elongate subrectangular; costa strongly
arched in basal third, then gently arcuate to almost straight; apex
rotundate; termen convex, slightly oblique; tornus broadly rotundate;
dorsum very gently arched to almost straight or flatly undulate.
No costal fold in male. Twelve veins, all separate: S gently sinuate;
R, from about middle of discal cell; Rz less than twice as close to R;
as to R,; R3 twice as remote from Ry, as latter from R,;; Ry running to
costa, R; to termen, never in apex, although sometimes rather close
to it; M, nearer to R; than to M2; Mz, M3, and Cu, almost equidistant,
latter originating from lower angle of discal cell; Cu, from about
two-thirds of discal cell; A, indiscernible basally, distinct tornally;
basal fork of Az;3 about one-fourth as long as entire vein. Hindwing
subtrapezoidal, as broad as forewing or narrower; costa gently arched
or slightly sinuate; apex rotundate; termen gently convex to almost
flat; tornus broadly and flatly rotundate; dorsum straight or slightly
concave in external portion, strongly curved basally. Eight veins:
S straight or slightly sinuate; R and M, closely parallel to about their
half, then diverging; M, gently descending basad, and at basis about
twice as close to Mz; as at termen; M; and Cu; connate or stalked,
originating at lower angle of discal cell; Cu, from two-thirds of discal
cell; A, well developed or vestigial; A, with basal fork, twice as close
to A; as to A;. No cubital pecten.

Mate GeniTatiA.—EKighth abdominal segment with a subtri-
angular mensis ventralis having a slightly stronger sclerotized proximal
angle. Uncus hooklike, moderately long, strongly sclerotized;
enathos with a dilated middle process; socius absent or indicated by
few hairs on internolateral fold of tegumen. Tegumen moderately
broad with shoulders oblique; pedunculi moderately broad, narrowing
ventrad; saccus broad, not deep, laterally dilated. Valva moderately
sclerotized, roundly and broadly dilated in basal two-thirds, and
ending with a much narrower cucullus turned upward; valvula with
ventral portion longitudinally folded and reaching into cucullus, and
with basal portion bearing a haired pulvinus; sacculus stronger
sclerotized, rather broad, without a free tip, and with ventral edge
subangular. Fultura superior complete, narrowed at middle; fultura
inferior flatly subcordate, with weak, haired dorsal papillae; caulis

4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

moderately long. Aedeagus moderately long, tubular, narrowed and
pointed ventroapically, with a long, obliquely located orificium
dorsoapically; coecum penis curved downward, slightly tapering to
bottom, ending with a narrow process, or without such a process;
cornuti flat, rather long, narrowed at base and apically, and lying
over one other.

FEMALE GENITALIA—Papillae anales pelmiform, oblong, soft, and
haired. Sinus vaginalis infundibular or somewhat tureen shaped,
cephalically rounded or flat. Lamella antevaginalis narrow, en-
circling sinus vaginalis cephalically; lamella postvaginalis composed
of two triangular pieces located caudolaterad of ostium bursa and
connected to dorsal, membranous wall of sinus vaginalis. Antrum
short, tubular, membranous or slightly sclerotized, bearing two nar-
row, lateral collicule; ductus bursa rather short; bursa copulatrix
membranous; cervix bursae variously long and broad, but generally
moderate; corpus bursa rotundate or slightly elongate, smooth or
somewhat rugose; signum strongly sclerotized, with a basal plate
elevated over external surface of corpus bursa, narrowly extended
caudad and cephalad, and having a variously pronounced capitulum;
internal process of signum dagger shaped, straight or curved.

Remarks.—In spite of its very distinct appearance, Jdolatteria is
morphologically very close to the genus Argyrotaenia Stephens, having
similar wing venation and very similar genitalia. The genitalic re-
semblance of these two genera is so complete, even in details, that it
is impossible to separate one genus from the other, using only these
characters. The external distinction consists of very long labial palpi
of Idolatteria and the bright coloring of the wings and the body of the
moths of this genus. Also the pattern of the wings is completely
distinct in both Jdolatteria and Argyrotaenia. The labial palpi of
Idolatteria further differ from those of Argyrotaenia in having a taper-
ing, subacute terminal segment which in the latter genus is more
cylindrical and blunt.

Key to Idolatteria Species

1. Forewing with not less than nine costal streaks; at least two subtermina
spots basad of terminal spots; bands in discal area (if any present)
DATTOM, ANG irregulati, <loerd jort- ysrah dys ee Hmevand aes ae BEE

Forewing with four costal spots, all of them in external third of costa;
a very large, round or somewhat piriform spot in subterminal area;
two broad bands crossing forewing from costa to (or almost to)

LG TM UATE ok LITER RRS eal Ree SORTING LES oat OAT REISS SINS MEA eres ee eee
2. Forewing without spots on dorsum .....- ++ 2 +s 2 ee ses 3
Forewing with spots on dorsum... . . ery oA

3. Termen of forewing with three smaller and two larger spots; hindwing darker
than forewiNe:< so sos. ceused +. >) sche epee, ws ae Re OCaDnE Meyrick

SS

NO. 3543 MICROLEPIDOPTERA, X1—OBRAZTSOV 5

Termen of forewing with two large spots; hindwing concolorous with fore-

WN Phe os oy ce bac . . » . Mmydros, new species
4. Discal spots of forewitig os oe faced me freemen rows . . maon Druce
Discal spots of forewing not fused into transverse rows. . . . ooo

5. Forewing with few spots in discal cell and between its end and terminal spots;
few spots on dorsum and none in supradorsal area.

simulatrix Walsingham
Forewing with discal spots numerous, also in supradorsal area; many spots on
dorsum . . . Meas Wptic, LO
6. Hindwing with cone oad pabberanall avers ree shen foe discal spots.
orgias Meyrick
Hindwing with terminal and subterminal i smaller than many of discal
spots .. . . pyropis Walsingham

7. Cilia of both mie see. cuenee pearicnts on “ake of hindwing.
cantharopisca, new species
Cilia of forewing black; that of pairs black or orange checked with black
in front of terminal dots... . One . . fasciata, new species

Idolatteria xanthocapna (Meyrick)

PLATE 1

Pseudatteria xanthocapna Meyrick, 1930, Exotic Microlepidoptera, vol. 3, p.
607.—Clarke, 1955, Catalogue of the type specimens of Microlepidoptera
in the British Museum described by Edward Meyrick, vol. 1, p. 326; 1958,
ibid., vol. 3, p. 200, pl. 100, figs. 4—4b.

Idolatteria xanthocapna.—Obraztsov, 1966, Proc. U.S. Nat. Mus., vol. 118, no.
3535, pp. 619.

MALE GENITALIA.—Uncus moderately long, curved, tapering apicad,
with tip obtuse. Valva roundly dilated in basal portion; cucullus
rather narrow, far not reaching upper level of dilated portion of valva.
Aedeagus with a very short apical tip.

Typr.—Holotype, male (genitalia on slide 4428, JFGC), Manizales,
Caldas, Colombia (A. M. Patino); BM.

ReEMARKS.—Known as a single male specimen. Superficially this
species reminds one of Pseudatteria heliocausta (Dognin) but differs
from the latter both structurally and in the wing markings.

Idolatteria simulatrix Walsingham

Ficure 1; Puate 2

Idolatteria simulatrix Walsingham, 1913, in Godman and Salvin, Biologia Cen-
trali-Americana, Lepidoptera Heterocera, vol. 4, p. 214; 1914, ibid., vol. 4,
p. 270, pl. 8, fig. 8.

FEMALE GENITALIA.—Sinus vaginalis infundibular, abruptly nar-
rowed toward ostium bursae; lamella antevaginalis narrow, with
lateral portions inclined mediad and incurved, and medial portion
forming a short arch bent cephalad. Antrum coincident with short,
cylindrical portion of ductus bursa bearing two lateral colliculi.

219-941-662

6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Signum narrow, dagger shaped, very insignificantly curved; caudal
and cephalic extensions of its basis narrow and almost equally long;
capitulum somewhat flat.

Type.—Holotype, female (genitalia on slide 2770), Geronimo,
Vera Paz, Guatemala (Champion; Godman and Salvin Collection;
66270); BM.

RemMarkK.—The holotype is the only known specimen of this
species.

Idolatteria mydros, new species

PLATE 3

Fema.e.—Antenna [only basal third present] black. Labial palpus
dark brown, basal segment and inner surface of second segment
yellow. Head dark brown; face and marginal cristae yellow. Thorax
dark brown with posterior margin orange yellow; most of tegula
yellow. Abdomen dark brown. Forewing reddish orange with
markings dark brown, iridescent bluish separated by cream-white
interspaces, and arranged as follows: nine variously broad streaks
on costa, last two of them occasionally connected at tips; a circular,
moderately sized apical spot closely touching a larger terminal spot,
located below and slightly incised at termen and interiorly; a second
terminal, slightly elongate spot, located closer to tornus, and also
incised at termen and interiorly; an elongate spot at tornus; a more or
less round spot dorsad of last two costal streaks and basad of upper
terminal spot, slightly larger than latter; a small spot (on right
forewing accompanied by a very minute dot) located above tornal
spot; some little, faint dots along external borders of orange area;
cilia cream white, in front of terminal spots and around tornus black.
Length of forewing 11 mm. Hindwing red orange with black spots:
a preapical spot on costa, an apical spot and six marginal spots along
termen and tornus; an obscure spot almost midway between dis-
cocellulars and termen; cilia grayish black.

Mauze.—Unknown.

FEMALE GENITALIA.—Sinus vaginalis flatly infundibular, narrowed
at ostium bursae; lamella antevaginalis narrow, with lateral portions
inclined mediad and medial portion forming a short arch, bent cepha-
lad. Antrum short, joined to a narrower, almost cylindrical portion
of ductus bursae bearing two lateral colliculi. Signum curved,
dagger shaped; caudal and cephalic extensions of its basis strongly
thickened, almost equally long and broad; capitulum not separated,
forming highest point of basis of signum.

Typr.—Holotype, female (genitalia on slide 1-Obr., Jan. 22, 1961),
Loja vicinity, Ecuador, 1887; USNM 67735.

ReMARKs.—Somewhat similar to simulatrix Walsingham, but with

PROC. U.S. NAT. MUS. VOL. 119 OBRAZTSOV—PLATE 1

- ‘

Figures 1-3.—Idolatteria xanthocapna (Meyrick), holotype, male: 1, left wings; 2, caudal

aspect of genitalia with valvae spread and aedeagus removed; 3, lateral aspect of aedeagus.
(From Clarke, 1958.)

PROC. U.S. NAT. MUS. VOL. 119 OBRAZTSOV—PLATE 2

3 4

Ficures 1-4.—/dolatteria simulatrix Walsingham, holotype, female: 1, left wings; 2, ventral
aspect of genitalia; 3, detail of sinus vaginalis; 4, detail of signum.

PROC. U.S. NAT. MUS. VOL. 119 OBRAZTSOV—PLATE 3

Ficures 1-4.—Idolatteria mydros, new species, holotype, female: 1, left wings; 2, ventral
aspect of genitalia; 3, detail of sinus vaginalis; 4, detail of signum.

PROC. U.S. NAT. MUS. VOL. 119 OBRAZTSOV—PLATE 4

Ficures 1-4.—Jdolatteria pyropis Walsingham, females: 1, holotype, left wings; specimen
from Monteverde, Costa Rica: 2, ventral aspect of genitalia; 3, detail of sinus vaginalis;
4, detail of signum.

PROC. U.S. NAT. MUS. VOL. 119 OBRAZTSOV—PLATE 5

|
|

Figures 1-4+.—/dolatteria maon (Druce), holotype, female: 1, right wings (image reversed)
2, ventral aspect of genitalia; 3, detail of sinus vaginalis; 4, detail of signum.

PROC. U.S. NAT. MUS. VOL. 119

OBRAZTSOV—PLATE 6

3

holotype, female: 1, left wings; 2, ventral
(From Clarke, 1958.)

Figures 1—4.—Idolatteria orgias (Meyrick),
aspect of genitalia; 3, detail of sinus vaginalis; 4, detail of signum.

PROC. U.S. NAT. MUS. VOL. 119 OBRAZTSOV—PLATE 7

4 | 7

Figures 1-7.—Idolatteria fasciata, new species, holotype, male: 1, left wings, 2, caudal
aspect of genitalia with valvae spread and aedeagus removed; 3, lateral aspect of aedeagus;
allotype, female: 4, right wings (image reversed); 5, ventral aspect of genitalia; 6, detail
of sinus vaginalis; 7, detail of signum.

PROC. U.S. NAT. MUS. VOL. 119 OBRAZTSOV—PLATE 8

Ficures 1|-4.—/dolatteria cantharopisca, new species, holotype, male: 1, left wings; 2, caudal
aspect of genitalia with valvae spread and aedeagus removed; 3, lateral aspect of aedeagus;
4, ventral aspect of mensis ventralis.

|

NO. 3543 MICROLEPIDOPTERA, X1—OBRAZTSOV c

only two terminal spots and without any dorsal spot on the forewing.
The submarginal spots of the forewing are dissimilar in shape, and
on the hindwing the submarginal row of spots is represented by only
one indistinct spot. The female genitalia resemble those of simula-
tri, but the signum is of a quite different shape. The specific name
is derived from the Greek word pvédpos, meaning ‘“‘a red-hot mass.”

Idolatteria pyropis Walsingham
Figure 2; Puatse 4

Idolatteria pyropis Walsingham, 1914, in Godman and Salvin, Biologia Centrali-
Americana, Lepidoptera Heterocera, vol. 4, p. 270, pl. 8, fig. 9.

FEMALE GENITALIA.—Sinus vaginalis tureen shaped; lamella
antevaginalis narrow, straight cephalically. Antrum membranous,
broader than adjacent, cylindrical portion of ductus bursae bearing
two lateral colliculi. Signum dagger shaped, slightly curved apically;
caudal extension of its basis longer than cephalic extension; capitulum
semirotundate.

Type.—Holotype, female (abdomen missing), Volcan de Irazu,
Costa Rica, 6000-7000 ft. (H. Rogers; Godman and Salvin Collection;
66225); BM.

OTHER SPECIMEN EXAMINED.—One female (genitalia on slide
757-Obr.), Monteverde, Puntarenas, Costa Rica, 4600 ft., Feb. 28,
1962 (C. W. Palmer); AMNH.

Remarxs.—No male of this species is known. There are some
color differences between the holotype, which appears somewhat
faded, and the other specimen examined, due probably to different
collection times. The antennae, labial palpi, head, thorax, and the
wing markings, described by Walsingham as being ‘‘purplish fuscous”’
or ‘‘dark purple,” are found by the present author as having these
colors in the holotype. In the newly collected specimen they are
almost black with a slight bluish hue. The head would best be
described as being black with a large, cream-white spot on the face
and with concolorous cristae externad of the eyes. The patagia are
black. The thorax is also black with two anterior, mediolateral
yellow streaks becoming pale orange distally; the posterior margin of
the thorax is orange; the external cristae of the tegulae are cream
white. The abdomen is bluish black with narrow postsegmental
bands orange dorsally, whitish laterally and ventrally, and also
whitish on the dorsal surface of the tip. The arrangement of the wing
spots is approximately the same in both of the examined specimens,
although the size and shape of separate spots are slightly distinct.
In spite of all the mentioned differences, there are no grounds to treat
the above specimens as belonging to two separate species.

8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 119

Idolatteria maon (Druce)

PLATE 5

Atteria maon Druce, 1901, Ann. Mag. Nat. Hist., ser. 7, vol. 7, p. 440.

Pseudatteria maon.—Meyrick, 1912, im Wagner, Lepidopterorum catalogus,
pt. 10, p. 16; 1913, in Wytsman, Genera insectorum, fasc. 149, p. 22.

Idolatteria maon.—Durrant, 1914, in Godman and Salvin, Biologia Centrali-
Americana, Lepidoptera Heterocera, vol. 4, p. 270.

FEMALE GENITALIA.—Sinus vaginalis tureen shaped; lamella ante-
vaginalis narrow, straight cephalically. Antrum membranous,
broader than adjacent, cylindrical portion of ductus bursae bearing
two lateral colliculi touching each other medially. Signum strong,
dagger shaped, insignificantly curved; caudal extension of its base
a little longer than cephalic extension; capitulum broadly rounded,
but moderately prominent.

Typr.—Holotype, female (genitalia on slide 5762), Chiguinda,
Ecuador (C. Buckley; 6610); BM.

ReMARK.—The holotype is the only specimen known.

Idolatteria orgias (Meyrick)
PLATE 6

Pseudatteria orgias Meyrick, 1930, Exotic Microlepidoptera, vol. 3, p. 607.—
Clarke, 1955, Catalogue of the type specimens of Microlepidoptera in the
British Museum (Natural History) described by Edward Meyrick, vol. 1,
p. 227; 1958, ibid., vol. 3, p. 199, pl. 99, figs. 3-3c.

Idolatteria orgias.—Obraztsov, 1966, Proc. U.S. Nat. Mus., vol. 118, no. 3535,
pp. 619.

FEMALE GENITALIA.—Sinus vaginalis tureen shaped; lamella
antevaginalis narrow, straight cephalically. Antrum membranous,
infundibular, much broader than adjacent portion of ductus bursa
bearing two lateral colliculi. Signum dagger shaped, broad, curved;
caudal extension of its basis narrow and long; cephalic extension
much broader and slightly shorter, and with scalloped margins;
capitulum with tip slightly narrowed.

Typr.—Holotype, female (genitalia on slide 4440, JFGC), Pacho,
East Cordilleras, Colombia, 7250 ft. (Paravicini Collection); BM.

OTHER SPECIMENS EXAMINED.—One female (one wing and abdomen
missing), Bogota, Cundinamarca, Colombia (Felder Collection;
400159); BM. One female (genitalia on slide 4441, JFGC), same
locality; USNM.

REMARKS.—The male is unknown. All three female specimens are
very similar, varying slightly in the shape and size of separate wing
spots.

NO. 3543 MICROLEPIDOPTERA, X1—OBRAZTSOV 9

Idolatteria fasciata, new species

PLATE 7

Mauz.—Antenna blackish brown with black annulation; tip white
scaled. Labial palpus black with inner surface of basal and second
segments cream yellow. Head black; face encircled by cream yellow.
Thorax black with prismatic blue hue; its middle, posterior margin,
and tips of tegula orange. Abdomen brown black with orange yellow,
on ventral surface paler, postsegmental rings. Forewing orange with
markings prismatic blue or violet, arranged as follows: A narrow band
at wing basis, oblique externad; a broad, almost vertical band before
middle of forewing and crossing it from costa to dorsum; a third band,
parallel to and almost as broad as former, and located just beyond
middle of forewing; a large, round or slightly piriform spot in external
third of forewing, separated from or connected to a much smaller
tornal spot; a triangular costal streak in interspace of first and second
bands; two or three more or less rotundate costal dots distad of third
transverse band, and a similar dot on forewing apex; five more or less
separate dots on termen; cilia black. Length of forewing 9.5-10.0
mm. Hindwing orange with brownish-black spots: an irregularly
shaped spot at wing basis, separate or fused with a dorsal spot; a more
or less rotundate spot at middle of disc, separate or fused with a dorsal
spot, other than already mentioned; sometimes these two dorsal spots
fused together; a rotundate or slightly piriform spot in external
portion of wing, lying free or occasionally reaching to termen; a small
preapical dot on costa; a more or less rotundate apical dot; two or
three terminal dots; cilia orange yellow, slightly checked with black
in front of terminal dots, or entirely black.

FrmaLe.—Similar to male, but with antenna not white at tip; third
band of forewing not reaching dorsum, or connected with it by means
of a narrow streak; interspace of second and third bands of forewing
with spots, a larger one on costa, a smaller on dorsum; spots in basal
half of hindwing fused together. Length of forewing 11 mm.

MALE GENITALIA.—Uncus moderately long, tapering apicad, bas-
ally distinctly broader than apically, with tip somewhat obtuse.
Valva strongly rotundate dilated in basal portion; cucullus moderately
broad; sacculus before middle with an arcuate elevation directed
dorsad. Aedeagus with a moderately long tip; coecum penis elongate,
acutely narrowed at bottom.

FEMALE GENITALIA.—Sinus vaginalis tureen shaped, rounded at
bottom; lamella antevaginalis narrow, equally arcuate. Antrum
cylindrical, fused with adjacent portion of ductus bursa bearing two
lateral colliculi. Signum straight, dagger shaped; caudal and cephalic

10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

extensions of its basis almost equally long, but former with a weakly
sclerotized prologation reaching into cervix bursae; capitulum rotun-
date, slightly prominent.

Typres.—Holotype, male (genitalia on slide 8667), Bolivia, 1903
(“Staudinger”); BM. Allotype, female (genitalia on slide 6603), Rio
Zongo, Bolivia, 750 m. (A. H. Fassal); BM. Paratypes: One male
(genitalia on slide 6604), Andes, Bolivia, 1920; one female, Bolivia;
BM. One male, Rio Zongo, Yungas, Bolivia, 1200 m., 1895-1896
(Garlepp); ZMB.

Remarks.—Very similar to the following species, cantharopisca,
with which it is compared. As a manuscript name, fasciata has been
used by Walsingham on the labels of some specimens, now becoming
the types of this species.

Idolatteria cantharopisca, new species
FicurE 3; PLATE 8

Maur.—Antenna entirely black with a prismatic blue hue. Labial
palpus black, at basis and on inner surface of basal and second seg-
ments orange. Head black with greenish hue; face encircled by
orange. Thorax black with greenish hue, at posterior margin and on
tips of tegulae orange. Abdomen entirely greenish black. Forewing
orange with prismatic green markings narrowly outlined with black
and arranged as follows: A narrow, transverse basal arcuate out-
ward; a very broad, transverse band in basal half of forewing, ex-
ternad of former band; a slightly narrower, transverse band crossing
forewing just externad of its middle; a large, round spot in external
third of forewing; three costal dots in same area; an apical dot, five
terminal dots, and a slightly larger tornal dot; cilia entirely orange.
Length of forewing 10 mm. Hindwing orange with black markings,
arranged as follows: A wide area consisting of at least three large spots
fused together, occupying more than basal half of hindwing, and in-
cluding some orange dots; a piriform, black spot externad of this area
and reaching termen; a black costal spot connected with basal area;
a preapical dot on costa; an apical dot, accompanied by a smaller dot
on termen; a larger spot on tornus; cilia orange, slightly grayish at
tips.

Frma.te.—Unknown.

MALE GENITALIA.—Uncus rather long, narrow, tapering apicad, and |
ending acutely. Valva with a rather narrow cucullus; sacculus be- |
fore middle with a triangular tooth directed dorsad. Aedeagus mod- —
erately long, with a narrow, long tip; coecum penis elongate, rounded |
at bottom.

NO. 3543 MICROLEPIDOPTERA, X1—OBRAZTSOV 11

Typr.—Holotype, male (genitalia on slide B.26), ‘‘Loeotal,’
Bolivia, 2600 m., 1891 (Garlepp); ZMB.

RemMARKS.—Very similar to Jdolatteria fasciata, new species, but
differs from it in some details. The antenna is not white at the tip,
and is not annulated. The labial palpus has the basis and the inner
surface of the basal and second segments deep orange. On the fore-
wing the markings are green. Most of the discal spots of the hind-
wing are joined into a common, black area. The cilia of both wings
are entirely orange, only those of the hindwing turn slightly grayish
toward the tips. The uncus of cantharopisca is somewhat longer and
narrower than in fasciata, and has a more acute tip. The tooth in
the basal half of the sacculus is triangular, not obtuse as in fasciata.
The bottom of the coecum penis is rotundate, and has no process.
Superficially the new species reminds one of a little specimen of
Pseudatteria cantharopa (Meyrick). The specific name cantharopisca
is derived from the name of this Pseudatteria species and the Greek
diminutive suffix tcxn.

12 PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 119

Ficures 1—3,—Idolatteria Walsingham, morphological characters: 1, wing venation of
I. simulatrix Walsingham, holotype, female; 2, head of J. pyropis Walsingham, female,
Monteverde, Costa Rica; 3, cornuti of J. cantharopisca, new species, holotype.

U.S. GOVERNMENT PRINTING OFFICE:1966

Proceedings of

the United States

National Museum

SMITHSONIAN INSTITUTION + WASHINGTON, D.C.

Volume 119 1966 Number 3544

RANGE AND VARIATIONS OF SUBSPECIES
OF CAMBARUS LONGULUS
(DECAPODA, ASTACIDAE)

By Hueco A. James!

As a result of the loss of the type specimens and confusion arising
from an erroneous restriction of the type locality of the typical sub-
species, the crayfish Cambarus longulus Girard, 1852, has been poorly
understood; furthermore, the ranges of its two previously described
subspecies have never been clearly delineated.

This report is based on a restudy of much of the material seen by
previous authors and of representatives of all the collections of
C. longulus in the U.S. National Museum (including those previously
a part of the collection of H. H. Hobbs, Jr.). In addition, personal
field studies within the ranges of all three subspecies herein recognized
have supplemented the study of preserved specimens. A total of
approximately 1300 specimens in 265 collections from 219 localities
have been included.

_ The maps have been prepared from drainage, contour, and base
maps obtained from the U.S. Geological Survey, Washington, D.C.

The present paper is a portion of a master’s thesis presented to the
Graduate Faculty of the University of Virginia. Completion of the
study was supported, in part, by grants from the National Science

1 University of Bridgeport, Bridgeport, Connecticut.

2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Foundation and the University of Bridgeport Faculty Research Fund.
The author also is deeply grateful to Dr. Horton H. Hobbs, Jr., for
his advice and assistance.

A History of the Species

1852. Girard, having examined specimens of crayfish of uncertain
locality (somewhere ‘‘within the middle States of the Union’’), con-
cluded that they were sufficiently different from the then-known forms
of C. diogenes and C. bartonii to warrant the designation Cambarus
longulus (p. 90). In this work, he first employed Cambarus as a
subgeneric name.

1870. The genus Cambarus was divided by Hagen into four groups
(p. 31), C. bartonii being made the type for his Group III into which
all similar forms were placed. Having examined Girard’s type of
C. longulus, Hagen indicated that it was probably an abnormal
C. bartonii (pp. 78, 79) and he erroneously placed it in his Group IT]
as ‘‘Cambarus Bartoni.”

1885. Faxon, having examined Hagen’s description of Girard’s
type of C. longulus, as well as several similar specimens in the USNM
collections, intimated that Girard’s C. longulus was valid (1885a,
p. 66), yet he did not consider his own total number of specimens
sufficiently adequate to warrant reestablishing longulus as a species.
Instead, on the basis of specimens taken from eastern Tennessee,
West Virginia, and Cumberland Gap, Va., he described and named
longirostris as a variety of the species C. barton (1885a, pp. 65, 66)
and stated: ‘‘The specimens described above under the name of
C. bartonii var. longirostris, perhaps are the same form as C. longulus

. in accord with Hagen’s description of Girard’s type” (1885a,
p. 66).

1890. Faxon“. . . after examining the large number of specimens |
(over one hundred, including females and both forms of the male)...” |
restored longulus ‘to the full rank of a species.”’ The variety longi- |
rostris, however, was retained as a varietal form of C. bartonzi on the
basis of an inadequate number of specimens (pp. 623, 624). In this |
paper, Faxon erred in designating C. longirostris as C. spinirostris |
but corrected his own error: ‘“(lapsu calami pro ‘longirostris’)” (Faxon |
1914, p. 424). .

1898. In his ‘Observation on the Astacidae,’’ Faxon listed several |
new localities for specimens of both “Cambarus longulus Girard” and |
“Cambarus bartonii longirostris” deposited in the USNM. It should |
be noted that he still retained the specific name of longulus (pp. 649- |
650). |
1899. Hay, in “Synopsis of North American Invertebrates,”

NO. 3544 SUBSPECIES OF CAMBARUS LONGULUS—JAMES a

placed longirostris with C. longulus as a subspecies of the latter (p.
966) and characterized both subspecies in a clear, concise key to the
North American Astacidae.

1903. Harris, in a compilation of data (1903, p. 58), referred to
C. b. longulus and C. 6. longirestris. JI have presumed this to be in
error owing to Harris’ interpretation of Faxon (1885a).

1905-1912. Having erected four subgenera of Cambarus, Ortmann
established the subgenus Bartonius (1905, p. 120), which was declared
a synonym of Cambarus for reason of priority by Fowler, who listed,
in the place of Bartonius, the subgenus Cambarus (1912, pp. 340-341).

1914. For some unexplained reason, Faxon reduced longulus to
subspecific rank calling it ‘“Cambarus bartoni longulus” and retained
longirostris as ‘“C. barton longirostris.’ He completely omitted
Hay’s designation and wrote: ‘“The character of the suborbital margin
of the carapace seems to be very constant within the limits of a good
subspecies, and it may prove to be the really diagnostic feature for
separating C. b. longulus and C. b. longirostris” (pp. 389, 424). Fifteen
years before, Hay (1899, p. 966) had utilized this identical diagnostic
feature in his key.

1931. Ortmann, following Fowler, listed both ‘‘Cambarus
(Cambarus) longulus longulus Girard (1852)” and ‘‘Cambarus (Cam-
barus) longulus longirostris (Faxon) (1855)” (pp. 118, 121). Ortmann
considered the type locality for longirostris to be ‘The first exact
locality given by Faxon (’85b) ... Doe River, Carter Co.,
Tennessee’”’ (p. 121).

1942. Ten years later, in a revision of the genus Cambarus, Hobbs,
primarily adopting Ortmann’s ideas of subgeneric relationships,
elevated the previously recognized subgenera to the rank of genus
within the new subfamily Cambarinae.

1959-1961. Hobbs (1959, p. 896) indicated that there are two
subspecies of Cambarus longulus and subsequently indicated to the
present writer that a third (form) probably should be recognized.

Cambarus longulus Girard

The species Cambarus longulus Girard (1852) belongs to the Bartoni
Section (Ortmann, 1931, p. 105) of Cambarus Erichso.1 (1846) (as
redefined by Hobbs 1942a, p. 354) and, as herein recognized, consists
of three subspecies: Cambarus longulus longulus, C. longulus longi-
rostris Faxon, and C. longulus chasmodactylus, new subspecies.

GEOGRAPHICAL LimITs—The limits of the range of the species C.
longulus, although more specifically designated within the geographical
discussions of the respective subspecies are: North: represented
by C. l. longulus, in the upper James drainage in Greene County
(on the eastern side of the Blue Ridge) and Highland County (to the

4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

west of the Blue Ridge) in Virginia, and by C. l. chasmodactylus in
Greenbrier drainage in Pocahontas County, W. Va. South: rep-
resented by OC. l. longulus in the upper Yadkin drainage of Wilkes
County (east of the Blue Ridge) and by ©. l. longirostris in Will’s
Creek (Coosa River drainage), DeKalb County, Ala. East: in the
upper piedmont province by C. J. longulus from the Rivanna River
(James drainage), Fluvanna County, Va., southward to the Yadkin
River in North Carolina. West: by C. l. chasmodactylus from the
Greenbrier River in West Virginia and C. 1. longirostris from a direct
tributary to the Tennessee River in Lawrence County, Tenn. (Map 1).

Key to Subspecies of Cambarus longulus

1. Suborbital angle well defined and acute. . - - longirostris Faxon (1885)
Suborbital angle markedly reduced or absent, never acute. 276.5) ais 26 te 2
2. Length of dactyl of chela less than twice the length of the inner margin of
Rhe paley 2. bese wp te eer Ae ae eee longulus Girard (1852)
Length of dactyl of chela twice (or more) the length of the inner margin of
GhospAlNA Yo ee dp pets Sree Sd Enel) pa chasmodactylus, new subspecies

Cambarus longulus longulus Girard

Cambarus longulus Girard, 1852, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, p. 90.

Cambarus longulus Faxon, 1890 (part) pp. 623, 624.—Ortmann, 1902, p. 277.—
Harris, 1903 (part) p. 107.—Fowler, 1912, p. 344.—Ortmann, 1913 (part),
pp. 335, 337, 339, 352, 353, 375, 376.—Brimley, 1938, p. 503.

Cambarus Bartonii Hagen, 1870 (part), pp. 7, 9, 75, 78, 79.—Faxon, 1884 (part),
p. 145; 1885a, pp. 11, 64, 66.

Cambarus bartonii longulus.—Faxon, 1885a, p. 66 (by implication).—Harris, 1903
(part), pp. 58, 107, 138, 142, 154-155, 159.—Faxon, 1914, pp. 390, 424.
Cambarus longulus longulus.—Hay, 1899 (by implication) (part), pp. 959, 966.—

Ortmann, 1913 (part), pp. 336, 337, 375.—Hobbs 1950, p. 349; 1959 (part),
896.—Johnson, R. M., 1957, pp. 178, 182; 1959, pp. 181, 183.—Johnson,
T. L., 1960, p. 229.
Cambarus (Bartonius) longulus—Ortmann, 1905 (part), pp. 120, 122, 128, 129.
Cambarus (Cambarus) longulus longulus.—Ortmann, 1931 (part), pp. 106, 107,
108, 118-121, 123, 124, 128, 134.

Dracnosis.—Concolorous, or speckled, in shades of blue to orange.
Rostrum with swollen margins, lacking marginal spmes or tubercles,
tapering somewhat abruptly to a short acumen, length .40-1.1 (aver-
age .95) times postorbital width. Suborbital angle and branchio-
stegal spines strongly reduced or absent; lateral spines or tubercles
on carapace usually present, but absent in certain populations;
postorbital spines and ridges weak. Areola from 29 to 42 (average
37) percent of length of carapace: 2.3-5.9 (average 3.5) times longer
than broad; and with 4-6 punctations across narrowest part. An-
tennal scale 1.7-3.2 (average 2.5) times longer than broad (pl. 1a).
Chela (pl. 1j-m) almost devoid of tubercles; inner margin of palm

NO. 3544 SUBSPECIES OF CAMBARUS LONGULUS—JAMES 5

with an indistinct row of weak serrations; all surfaces with scattered,
deep punctations. Palm broad, with widely gaping (in adults),
subcylindrical fingers which meet only at their tips; inner base of
immovable finger with tuft of plumose setae in immature animals, and,
except in upper James River, to some degree in older forms. Proxi-
momesial angle of inner margin of palm strongly hooked proximally,
forming, with carpus, a deep-curved acute angle (pl. 1mm: C); length
of inner margin of palm (pl. 1mm: A) at least twice that of distance
between spine of carpus and proximal extremity of inner margin of
palm when the chela is fully extended (pl. 3mm: B) (in well over 75
percent of examined specimens). Width of palm .53-1.0 (average
.80) times dactyl length; length of outer margin of chela 1.5-3.6
(average 2.7) times that of inner margin of palm and 1.0-2.6 (average
1.7) times greater than the length of dactyl. First pleopod of first-
form male reaches coxa of third pereiopod when abdomen is flexed.
(For detailed description of first pleopod of first-form male, see Hart,
1952, p. 47; Parish, 1948, figs. 2, 4.)

Remarks.—Faxon (1898, p. 650) (1914, pp. 389, 424) lists as C. 1.
longulus several collections which should be referred to C. 1. chas-
modactylus or C. longulus longirostris; Newcombe (1929, maps 268, 278,
286) and Fleming (1938) repeat the errors of Faxon. All collections
erroneously recorded as C. l. longulus are listed by me with their
proper designations in list I (p. 7).

Of the three subspecies, C. l. longulus appears to be the most
variable. Although most populations possess a tuft of plumose
setae at the base of the immovable finger of the chelae, those of the
upper James drainage typically lack such a tuft. Ortmann (1931,
pp. 118-124) notes the absence of lateral spines on the carapace of
nine of his specimens of C. 1. longulus. He was correct only with
respect to a minority of populations; most have spines.

Some taxonomic characters, previously used by other investigators,
must be disregarded or used only in part. Size, although of interest,
gives little indication of subspecific variation; most ©. 1. longulus
are, on the average, smaller than C. longulus longirostris or C. 1.
chasmodactylus; the carapace length of the largest first-form male
C. 1. longulus examined is 34 mm. and its hand length 29 mm. This
male is larger than some first-form males of C. longulus longirostris
and ©. 1. chasmodactylus: hence, size (sexually mature adults) as
a taxonomic criterion must be used in combination with other char-
acters. Color and color pattern of living specimens are sometimes
useful in separating populations of C. l. longulus (speckled green
to orange versus concolorous blue greens and browns). Color pat-
tern is of slight intersubspecific value in that neither of the other

6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

subspecies shows the speckled condition; however, neither do the
majority of the population of C. l. longulus.

SPECIMENS EXAMINED.—I have examined approximately 100 col-
lections from 78 localities in Virginia, West Virginia, and North
Carolina, including adults of both sexes, both forms of male, and
juveniles, representing a minimum of 500 specimens. I have no
authenticated records of this subspecies having been taken from
Tennessee (see list III, p. 7).

GEOGRAPHIC DISTRIBUTION.—It would appear that C. J. longulus
is limited geographically to the James, Roanoke, and Yadkin River
systems of the Atlantic drainage (maps1,2). Thesouthernmost record
is from the Yadkin River drainage, Wilkes County, N.C. (map 2,
no. 92). The northern limit seems to be in the headwaters of the
James in Highland County, Va., west of the Blue Ridge (map 2, nos.
51-53) and in Greene County, east of it (map 2, nos. 43-50). It
is found as far east as the Rivanna River (James drainage) and its
tributaries (map 2, no. 36) but does not, from my data, extend west
into or beyond the New River drainage or north into the Shenandoah
as some records erroneously indicate (see lists I, II).

A list of all localities and drainage systems in which C. 1. longulus
has been found has been deposited with the U.S. National Museum.?
Many collections incorrectly labeled as C. l. longulus must herewith
be removed and reassigned to other taxa.

Because the labels for his type specimens had been lost, Girard
(1852, p. 90) cites the locality only as ‘Middle States.”” Ortmann
(1931, p. 118), listing the first of Faxon’s reported localities (1890,
p. 623), designates ‘‘the first exact locality . . . as a supplementary
type-locality . . . South River, Waynesboro, Augusta Co., Virginia.
(Faxon) (to Shenandoah and Potomac).’”’ This choice of type locality
seems inadmissible, for no member of the species C. l. longulus has
been found since in the Potomac drainage. Neither Girard (loc. cit.)
nor Hagen (1870, p. 78) mentions the presence of a lateral spine on
the type specimen, but Hagen does note that between the fingers there
‘Gs a large bunch of hairs.”” From this combination of characteristics,
it would seem more likely that Girard’s type came from the James
drainage, east of the Blue Ridge.

SYNONYMICAL REASSIGNMENTS.—On the basis of available data,
many collections (listed below), previously recorded as C. l. longulus,
now must be reassigned either to another species or to the subspecies
C. longulus longirostris or C. l. chasmodactylus. Inappropriate names
and/or localities are so indicated in the synonomy.

2 Copies may be obtained by writing to the author or to the U.S. National
Museum, where these data are on file (no 254736) in the office of the Registrar.

NO. 3544 SUBSPECIES OF CAMBARUS LONGULUS—JAMES iG

List I
(Forms recorded as C. 1. longulus that must be relegated to other species)

Faxon, 1890. 1. Waynesboro, Potomac River—no C. longulus are known
from this drainage.

Harris, 1903. 2. A compilation of data, repeating the errors after Faxon
1884-1890.

Ortmann, 1913. 3. “Shenandoah River (Faxon),’’ Potomac drainage—see 1
above.

Ortmann, 1931. 4. As in Ortmann 1913—see 3 above.
Fleming, 1988. 5. Errors as in Faxon to 1914.

List II
(Forms recorded as C. J. longulus that must be relegated to C. l.
chasmodactylus)
Faxon, 1890. 1. Wytheville, Va. (New River).
2. Reed Creek, Wytheville, Va. (New River).
Hay, 1899. 3. Hay derived his key on the basis of known collec-
tions that include C. l. chasmodactylus.
Harris, 1903. 4. See list I, no. 2.
Ortmann, 1905. 5. Errors as in Faxon through 1898.
Ortmann, 1913. 6. Upper Kanawha River (presumed to be New River
drainage; if not, this collection then belongs in
list I).
7. Greenbrier River (New River).
8. New River.
Faxon, 1914. 9. Greenbrier River (New River).
10. Bluestone River (New River).
Newcombe, 1929. 11. A compilation, errors as in Faxon, 1914 (list IT,
nos. 9, 10).
Ortmann, 1931. 12. A listing of known localities, errors as in Faxon,
1890 (list I, no. 1) and Ortmann, 1913 (list II,
nos. 6, 7, 8).
Fleming, 1938. 13. Errors as in Faxon to 1914.
Hobbs, 1959. 14. According to collections on which Hobbs key was
based, part of C. l. longulus belongs here.
List IIT
(Forms recorded as C. 1. longulus that must be relegated to C. longulus
longirostris)
Faxon, 1890. 1. South Fork of Holston River (Tennessee River)—

C. l. longulus is entirely absent from Tennessee and
the Tennessee River system.

Spring Creek to French Broad River.

Watauga River (Holston River).

. Knoxville, Tenn.

. Eastern Tennessee.

. Holston River drainage system.

Don mp wb

8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Faxon, 1898. 7. Tennessee, Cumberland Gap.
8. Tennessee, Tazewell.
9. Tennessee, Greeneville.
10. Tennessee, Knoxville.
Harris, 1903. 11. See list I, no. 2.
Ortmann, 1905. 12. Errors as in Faxon to 1898.
Ortmann, 1913. 13. Holston and Clinch River systems—corrected by
Ortmann, 1931.
Brimley, 1938. 14. Lists both C.l. longulus and C. longulus longirosilis—
both are C. longulus longirostris from French Broad

drainage.
Fleming, 1938. 15. Errors as in Faxon to 1914.
Hobbs, 1959. 16. These forms of C. l. longulus from Tennessee and the

Tennessee River system in southwestern Virginia
are C. longulus longirostris.

List IV

(Other synonymical records not fitting into lists I, II, IIT)

Hagen, 1870 1. ‘“‘A female type of C. longulus . . . differs from C. Bartonii in
having its hands smooth .. . I think it is C. Bartonii’’—
This specimen more logically belongs to C. l. longulus.

Faxon, 1914 2. “I have seen an interesting lot of specimens (from above
Kanawha Falls) that combine the characters of C. 6b.
montanus and C. b. longulus. ... These specimens are in the
U.S. National Museum, No. 23990, and in the Museum of
Comparative Zoology, No. 7401.’”—Having examined the
former collection, the present writer, although uncertain of
its specific status, excludes the above from the C. longulus

group.

CoLORATION AND COLOR PATTERN.—Most C. 1. longulus are con-
colorous, usually a brown or green with shades of ivory to tan beneath.
A mottled pattern is known from two widely separated localities
(fig. 2c). One, the Swift Run, a tributary of the North Fork of the
Rivanna River (James drainage) in Greene County, Va. (map 2, no.
50) is entirely of the brown phase (orange to dark brown). The other
mottled pattern is found in some tributaries of the Smith and South
Mayo Rivers (Dan River to Roanoke drainage) in Patrick County
(map 2, nos. 65-70) ; these are variously colored within individual popu-
lations. Colors range from yellow orange through shades of green
and brown and, like the James River specimens, have their underparts
tinted ivory to tan. It would appear that the background color of
these mottled ‘“Roanoke longulus’’ is in shades of tan and that only
the mottling seems to vary. These two very similarly patterned and
colored populations of crayfish occur in streams almost 125 miles apart.

NO. 3544 SUBSPECIES OF CAMBARUS LONGULUS—JAMES 9

Cambarus longulus longirostris Faxon

Cambarus Bartonii var. longirostris—Faxon, 1885a, Mem. Mus. Comp. Zool.,
vol. 3, p. 64; 1885a (part), p. 64; 1885b, p. 358; (as C. bartonii longirostris) ;
1890, pp. 623, 624; 1898 (part), p. 649.—Harris, 1903 (part), pp. 58, 75, 154.—
Ortmann, 1905 (part), pp. 128, 129, 135.—Faxon, 1914, pp. 389, 424.—New-
combe, 1929, p. 286.

Cambarus bartonii spinirostris.— Faxon, 1890, pp. 623, 624.

Cambarus longulus.—Faxon, 1890 (part), pp. 623, 624; 1898 (part), p. 650.—
Harris, 1903 (part), pp. 58, 107, 138, 154, 155, 159.—Ortmann, 1913 (part),
pp. 335, 337, 362, 375.—Fleming, 1938, pp. 299, 300, 301.—Brimley, 1938,
p. 502.—Hobbs, 1959 (part), p. 898;

Cambarus longulus longirostris—Hay, 1899, pp. 959, 966; Johnson, 1957, pp. 178,
182; 1959, pp. 181, 183.—Hobbs, 1959 (part), p. 898.

Cambarus (Cambarus) longulus longirostris—Ortmann, 1931 (part), pp. 121-124.

Cambarus longerosilis Brimley, 1938, p. 503.

Dracnosis.—Concolorous in shades of blue green through orange,
or bicolorous with two dark dorsal saddles of varying widths; one on
posterior part of carapace and the other immediately cephalic to cer-
vical groove (figs. 24,6). Rostrum with strongly swollen margins,
without spines or tubercles, abruptly tapering to moderately short
acumen, length .78-1.1 (average 1.0) times postorbital width. Sub-
orbital angle and postorbital spines and ridges strong (pl. 17). Bran-
chiostegal spines and lateral spines or tubercles on carapace absent.
Length of areola 2.3-5.9 (average 4.0) times width, 30-39 (average
36) percent of length of a carapace, and with 3-10 punctations across
narrowest part (average 7-8). Antennal scale 1.7-3.0 (average 2.3)
times longer than broad. Chela mostly as in C. 1. longulus (pl.
1n,o), length of outer margin of chela 1.5-1.8 (average 1.7) times
length of dactyl and 2.6-4.4 (average 3.1) times length of inner
margin of palm, length of dactyl of chela 1.5-3.0 (average 1.9) times
length of inner margin of palm, and width of palm .52-.88 (average
.72) times length of dactyl. First pleopod of first-form male as
pictured (pl. 1f,9).

Remarks.—Faxon (1885a, p. 64) indicates that his collection of
three specimens“‘from Cumberland Gap (Claiborne Co., Tenn.) have
well-marked lateral spines on the carapace.’ Since the chela of this
Powell River form differs from that of the typical C. longulus, it is,
presumably (awaiting further data), other than the species C. longulus.
Some specimens of C. longulus longirostris do have a minute tubercle
in place of the lateral spine.

Little difference exists between C. longulus longirostris and the
other two subspecies. Variations in all characteristics thus far ob-
served, exclusive of the suborbital angle, overlap to some degree
those of the other subspecies: average ratios calculated for C. longulus
longirostris usually lie midway between those calculated for C. l.

219-942 662

10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

longulus and C. l. chasmodactylus. 'The setaceous tuft at the base of
the immovable finger of the chela may be present or absent but, as
in C. 1. longulus, it seems to be consistent within a population; older,
late intermolt animals do not seem to lose the tuft of setae as readily
as do those of C. l. longulus or C. l. chasmodactylus. Most C. longulus
longirostris are larger than C. l. longulus and smaller than C. 1. chas-
modactylus. The largest, a first-form male, has a carapace that
measures 43 mm. in length and a chela, 56 mm. long. Both color
and color pattern vary in different parts of the range; figures 2a,b
illustrate two of the pattern variations. Neither C. J. longulus nor
C. 1. chasmodactylus possesses the vivid saddle pattern seen in so many
C. longulus longirostris populations, particularly those orange-colored
animals from Lawrence County and the less colorful individuals from
the Hiwassee drainage, Tennessee. Most C. longulus longirostris are
a blue green or brown not unlike the concolorous C. l. longulus.

There are so many variations in this crayfish that, beyond the
presence of the suborbital angle, no characteristic has been observed
that will serve to distinguish C. longulus longirostris from the other
subspecies.

SPECIMENS EXAMINED.—1 have examined approximately 500 speci-
mens from 113 collections taken from 97 localities in Alabama,
Georgia, North Carolina, Tennessee, and Virginia. Many of these
collections contain both sexes and both forms of the male.

GEOGRAPHIC DISTRIBUTION.—Cambarus longulus longirostris is
confined to tributaries of the Tennessee and Coosa Rivers. Reports
from the Clinch River, ‘‘West Virginia” are erroneous; the Clinch
River, Tennessee drainage, does not extend into West Virginia; one
need only note this location to know that Faxon erred and probably
meant Clinch River in western Virginia. The northern limit appears
to be in the upper Clinch (map 2, no. 202), Tazewell County, Va.;
its southernmost boundary is Will’s Creek (Coosa drainage) (map 1,
no. 96), DeKalb County, Ala. Having recorded five collections
from Will’s Creek in DeKalb County, I believe it is highly probable
that this is the locality meant by Faxon (1898, p. 649), not ‘Will’s
Creek, Pollard, Escambia Co. [italics mine], Alabama.”’ I have no
knowledge of a Will’s Creek in Escambia County. Ortmann (1931,
p. 123), assuming that the county listed by Faxon was correct, notes
this record as being ‘‘extremely doubtful” giving instead as the south-
ernmost locality, Catoosa County, Ga. (Tennessee River drainage).
Two collections from Lawrence County, (map 1, nos. 209, 210) and
one from Lauderdale County, Ala. (map 1, no. 98) mark the western-
most limit of the range. Although many collections have been made
both east and west of the southwestern localities, no C. longulus
longirostris has been found closer than those collected from Marion

NO. 3544 SUBSPECIES OF CAMBARUS LONGULUS—JAMES 11

County, Tenn. (map. 1, nos. 143-145). It would seem that here the
ecologically restricted C. longulus longirostris has been forced into
its isolated strongholds by the invasion of members of the genus
Orconectes. It is probable that these western records, especially
those of Lawrence County, Tenn., are relict populations. No eco-
logical data are available for the Lauderdale, Ala. specimens, although
I suspect they would conform to those of Lawrence County. The
northeastern limit is marked by the North Fork of the Holston River,
Bland County, Va. (map. 2, no. 186).

That the three subspecies of C. longulus are allopatric is clearly
evidenced where the headwaters of separate drainage systems inter-
digitate. For example, White Top Creek (map 2, near no. 195)
northeast of Konnarock, in Smyth County, Va. (Holston drainage),
in which C. longulus longirostris is found, is less than a mile from
Lewis Creek (map 2, no. 248) near Troutdale, Grayson County (new
drainage) where C. l. chasmodactylus occurs.

In many localities C. longulus longirostris is particularly abundant;
in one such locality (map 2, no. 195), Big Laurel Creek, Smyth
County, Va., Hobbs (personal communication) collected 99 specimens
in a single seine haul over approximately 100 square feet.

Ortmann (1931, p. 123) wrote that C. longulus longirostris had “not
been found in the Tennessee River below Knoxville, and the mouth
of the Clinch, nor in its eastern tributaries (Little River, Little
Tennessee, and Hiawassee [sic] Rivers).’’? I have recorded specimens
from Louden County (below Knoxville), Roane County (below the
mouth of the Clinch); Monroe County (Little Tennessee), McMinn,
Bradley, and Polk Counties (Hiwassee); also previously unrecorded
are localities in Lawrence County, Tenn., Lauderdale County, Ala.,
and Armuchee Creek, Floyd County, Ga. (Coosa drainage).

Faxon’s type locality (1885b, p. 358) is ‘Doe River, Elizabethon,
Carter County, Tenn.,”’ from which he had three females. Ortmann
(1931, p. 121), Hobbs, and Holt subsequently have collected specimens
from this locality (map 2, no. 112).

The only incorrect listing of C. longulus longirostris of which I am
aware is “Cumberland Gap” (Faxon, 1885a), as previously discussed.
This locality has been repeated in Faxon (1898), Harris (1903), and
again in Ortmann (1931). The error is logical; except for the differ-
ence in chelae and the presence of lateral spines on the carapace, this
form is superficially much like C. longulus longirostris.

Those locality records listed by previous authors for C. l. longulus
that actually apply to C. longulus longirostris are included in list ITI,

Det.

PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

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Cambarus longulus chasmodactylus, new subspecies

Cambarus Bartoniit Hagen, 1870 (part), p. 76.—Faxon, 1885a (part), pp. 60, 61.

Cambarus longulus Faxon, 1890 (part), pp. 623, 624.—Ortmann, 1913 (part),
pp. 335, 337, 375.

Cambarus longulus longulus—Hay, 1899 (part), pp. 959, 966.—Hobbs, 1959
(part), p. 898.

Cambarus bartonii longulus.—Harris, 1903 (part), pp. 58, 107, 138, 146, 148, 154,
155, 159.—F axon, 1914, p. 389, 390.—Newcombe, 1929, map, pp. 268, 278,
280, 286.

Cambarus (Bartonius) longulus—Ortmann, 1905 (part implied), pp. 120, 122,
128, 129.

Cambarus (Cambarus) longulus longulus—Ortmann, 1931 (part), pp. 106, 107,
108, 118-121, 123, 128, 134-136.

Cambarus subspecies.—Johnson, 1957, pp. 178, 182; 1959, pp. 181, 183.

Draenosis.—Concolorous, blue green tinged with shades of cream
to rust. Rostrum with strongly swollen margins, without marginal
spines or tubercles, acumen tapering abruptly to upturned apex, with
a punctate depression posteromedially; length of rostrum 1.5-2.2
(average 1.9) times longer than wide; .69-1.1 (average 1.0) times
postorbital width and .17—.28 (average .23) times carapace length.
Suborbital angle and branchiostegal spines absent, lateral spines
absent except in specimens from one locality in Carroll County, Va.
(map 2, no. 240); postorbital ridges and spines strong, cheeks with
scattered low tubercles. Areola 3.5-6.0 (average 4.4) times longer
than broad, length 34-37 (average 27) percent of length of carapace
and with 4-10 (average 6-7) punctations across narrowest part.
Antennal scale 1.6-2.7 (average 2.2 times longer than broad), spine
strong. Inner margin of palm of chela with a row of squamous tuber-
cles, remainder smooth; all surfaces with scattered, deep punctations,
some on dorsal surfaces form linear rows. Palm broad, fingers of
adults intensely gaping, meeting only at tips; immovable finger, sub-
triangular in cross section, with basal tuft of plumose setae, frequently
reduced or absent in adult forms (pl. 1y,z) but present in all juveniles
(pl. 1w,x); dactyl ovate to subcylindrical in cross section. Both
fingers with single row of low tubercles on opposable surfaces, single
tubercle near distal end of immovable finger often enlarged. ‘Hook’
of proximomesial angle in inner margin of palm weak or obsolete,
forming with carpus a deep weakly curved obtuse angle (pl. 1zz: C);
length of inner margin of palm (pl. 122: A) of fully extended chela less
than twice that of distance between spine of carpus and proximal
extremity of inner margin of palm (pl. 1zz: B). Width of palm
.48-.74 (average .61) times length of dactyl, length of outer margin of
chela 2.6-4.7 (average 3.6) times length of inner margin of palm and
1,2-1.8 (average 1.6) times greater than length of dactyl. First
pleopod of first- and second-form males and annulus ventralis of fe-

NO. 3544 SUBSPECIES OF CAMBARUS LONGULUS—JAMES 15

Ficure 1.—Cambarus longulus chasmodactylus, new subspecies: a, lateral view of first
pleopod of first-form male; b, lateral view of carapace; c, mesial view of first pleopod of
first-form male; d, lateral view of first pleopod of second-form male; ¢, mesial view of
first pleopod of second-form male; f, epistoma; g, ventral view of annulus ventralis;
h, dorsal view of carapace; i, upper surface of right antennal scale; j, basipodite and
ischiopodite of third right pereiopod of first-form male; k, upper surface of right chela and
carpus of first-form male.

16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

male imperceptibility different from C. 1. longulus (pl. 1d,e,u; figs.
1a,c—e,g).

HOLOTYPE MALE, FORM I.—Body about 1.5 times broader than deep.
Greatest width of carapace at level of midlength of areola (21.4 mm.).
Abdomen shorter than carapace (34.0 and 39.7mm.) and narrower
(16.0 and 21.4). Areola about 4.1 times longer than broad with six
punctations across narrowest part. Cephalic section of carapace
about 1.6 times length of areola. Areola about 38 percent of entire
length of carapace (figs. 16,h).

Rostrum with thickened margins, without marginal spines or
tubercles, gently tapering, dorsally concave with deep punctations
scattered in caudally situated oval depression; rostrum with thick-
ened, cephalically converging margins forming an indistinct acumen
with upturned apex, a row of setaceous punctations along mesial base
of thickened margins. Subrostral ridges weak but visible along entire
length of rostrum in dorsal aspect.

Postorbital ridges conspicuous, terminating cephalically in a small
corneous spine; grooves of postorbital ridges well developed and
bearing fine setae. Suborbital angle obsolete, replaced by a slight
rounded projection just above level of base of antenna. Branchioste-
gal spine reduced to a small angular prominence. Carapace without
lateral spines. Upper surface of carapace and lateral portion of
branchiostegites punctate. Few granulations on lateral surface im-
mediately caudal to cervical groove, but many on cephalolateral
regions.

Cephalic section of telson with two spines in each caudolateral
corner. Epistome with a slight cephalomedian projection (fig. 1/).

Antennules of usual form with a strong spine present on ventral
side of basal segment. Antennal scale broadest just proximal of
midlength, with outer distal margin terminating in a long, acute,
corneous spine.

Right chela (fig. 1k) conspicuously punctate above and below,
flattened, and about twice wider than deep (20-10 mm.). Inner
margin of palm with single median row of nine squamous tubercles
scarcely rising above contour of margin; longitudinal row of deep
punctations present above and lateral to row of squamous tubercles.
Upper surface of palm with two large tubercular swellings at base of
dactyl. Immovable finger strongly convex laterally and without
well-defined grooves or ridges; proximal half subtriangular in cross
section, distal half ovate; basal portion almost as deep as wide;
opposable margin with a tuft of setae proximally and with a more
distal row of 16 rounded, corneous tubercles; larger tubercle present
just below tubercular row and slightly distal to midlength; a row of
minute denticles on distal third of opposable margin just ventral to

NO. 3544 SUBSPECIES OF CAMBARUS LONGULUS—JAMES 17:

row of tubercles. Dactyl convex mesially along proximal two-thirds
and without ridges or grooves. Concave opposable margin with row
of 18 rounded corneous tubercles and distal third with a row of minute
denticles ventral to tubercular row.

Carpus of right pereiopod longer than broad (14.0-10.5 mm.), with
a deep longitudinal furrow above. Dorsal and lateral surfaces
punctate; mesial surface with one large spinous tubercle near mid-
length and a much smaller one proximally; ventral surface with two
tubercles on distal margin.

Merus of first right pereiopod punctate laterally and mesially. A
small tubercle near upper distal margin. Lower surface with a row of
eight tubercles mesially (the two most distal enlarged and spinous),
and two laterally.

Hooks on ischiopodites of third pereiopods only (fig. 17); hooks
heavy and projecting proximally a little beyond distal end of
basipodite. Coxae of fourth pereiopods with caudomesial swollen
prominences.

Ficure 2.—a, Saddle pattern of “‘orange phase” of Cambarus longulus longirostris from
Lawrence County, Tenn.; 5, saddle pattern of the less colorful phase of C. longulus longiros-
tris from Hiwassee drainage system in Polk County, Tenn.;c, a mottled phase of C. /. long-
ulus from Dan River headwater (map 2, no. 70).

First pleopod reaching coxopodite of third pereiopod when abdomen
is flexed. Tip terminating in two parts (figs. 2a, c); mesial process
bulbous, noncorneous, bent caudolaterally and bearing a minute spine
at its tip. Slight shoulder on lateral surface arising gradually from
base of central projection.

18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

MorpHOTYPE MALE, FoRM 11.—Differs from the holotype as follows:
Cephalic section of telson with three spines in right and two in left
caudolateral corners. Minute spinelike tubercle on left side of cara-
pace immediately caudal to cervical groove. Areola with seven
punctations across narrowest part. Inner margin of palm of chela,
with a row of six squamous tubercles; opposable margin of immovable
finger with 17 tubercles and that of dactyl with 19. Upper distal
portion of merus with two tubercles and lower surface with a mesial
row of ten tubercles. Hooks on ischiopodite of third pereiopods much
reduced in size. First pleopod without corneous elements (figs. 1d, e) ;
swollen prominence of fourth pereiopods smaller.

ALLOTYPE FEMALE.—Differs from the holotype as follows: Areola
with five punctations across narrowest part. Opposable margin of
immovable finger of right chela with 13 rounded tubercles, and that
of dactyl with 15. Upper distal portion of left merus with two
tubercles and lower surface with mesial row of seven. Rhomboid
annulus ventralis with a high V-shaped ridge caudally and a medial
longitudinal trough, deepest at apex of bend in sinus; sinus originates
at caudal end of trough, makes a V-shaped arc dextrally and then
bends caudally cutting midcaudal margin of annulus (pl. 1u; fig. 1g).

Measurements (in millimeters) of C. 1. chasmodactylus are as follows:

rostrum holotype allotype morphotype

length 9.1 8.4 8.3

width 4.5 4.3 4.2
postorbital ridges (distance between) 9.1 8.5 8.9
carapace

maximum depth 14.3 14.0 15.0

maximum width 21.4 20.0 21.6

length 39.7 36.5 38.8

cephalic length 24.2 22.0 24.0
areola

length 15.2 14.1 14.9

width 3.7 2.6 3.4
abdomen

length (including telson) 34.0 34.5 35.0

width (at widest part) 16.0 Wied 16.6
antennal scale

length 5.1 5.1 5.6

width 2.5 2.5 2.6
right chela

palm width 20.0 14.0 17.5

palm depth 10.0 7.5 8.8

inner margin of palm length 13.6 9.1 7
right chela hand (outer margin)

length 47.2 32.0 42.9

length of dactyl 29.0 20.6 27.8
right carpus

length 14.0 9.0 12.0

width 10.5 8.0 9.0

NO. 3544 SUBSPECIES OF CAMBARUS LONGULUS—JAMES 19

Ratios of measured characters of C. 1. chasmodactylus are as follows:

holotype] _allotype mor photy pe

carapace W/L 1.50 1.40 1.44
L of cephalic part of carapace; areola L 1.59 1.56 1.61
Rostrum L/carapace L .23 .23 sak
Rostrum L/W .20 .20 .20
Rostrum L/postorbital width 1.00 .99 .94
areola L/W 4.10 5.40 4.40
areola L/carapace L 38 .39 38
antennal scale L/W 2.40 2.00 2.20
palm of chela W/L of dactyl .69 .68 .63
hand of chela L/L inner margin of palm 3.50 3.50 3.70
hand of chela L/L of dactyl 1.60 1.60 1.50
dactyl of chela L/L inner margin palm 2.10 2.30 2.40

Typr-LocaLity.—‘‘East Fork of the Greenbrier River 9.7 mi. W.
of Virginia State line on U.S. Hwy. 250, Pocahontas County, West
Virginia [map 2, no. 263]. Here the river is between 50 and 100 feet
in width and up to 3 feet in depth. Its slightly clouded water
flows sluggishly to moderately over tilted bed rock strata on which
are strewn many stones and a considerable amount of silt. The
crayfish are abundant and lie concealed beneath the loose rocks”
(Hobbs, pers. comm.).

DisposiTIon oF tTypE.—The holotype male (USNM_ 115513),
allotype female (USNM 115514), and the morphotype male (USNM
115516) are deposited in the U.S. National Museum. Of the remaining
161 paratypes (USNM 115515), 17 males, Form I, 5 males, Form IT,
16 females, 3 juvenile males, and 2 juvenile females are also in the
U.S. National Museum; 7 males, Form I, 5 males, Form II, and 1
female are in: (a) Museum of Comparative Zoology, Harvard; (b)
the collection of G. H. Penn, Tulane University; (c) the collection of
Alejandro Villalobos, Institute de Biologia, Mexico, D.F.; and (d) 2
males, Form J, 2 males, Form IT, 2 females, and 1 juvenile male are
in the collection of the author at the University of Bridgeport, Bridge-
port, Conn.

Remarks.—Ortmann (1931, p. 120) was aware of the differences
between C. 1. longulus and C. l. chasmodactylus and wrote that in those
“specimens from the New River drainage . . . the moveable finger
may be over twice as long as the inner margin of the palm.” This
has been a completely reliable characteristic for distinguishing all the
normal (chelae unregenerated) specimens I have seen.

On the basis of Faxon’s work (1885a, pp. 60, 61; 1890, p. 623), I
suspect that Hagen’s ‘Greenbrier River’ record (1870, p. 76) is C. l.
chasmodactylus, but I do not have sufficient information to synony-
mize it. Further investigation may identify the above as well as
Faxon’s records of “Reed Creek, West of Wytheville, Wythe Co.,

20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

(Virginia)”’; and those “near White Sulphur Springs, Greenbrier Co.,
West Va.” (1885a, pp. 60, 61) as C. l. chasmodactylus.

While C. 1. longulus is the most variable of the subspecies, C. J.
chasmodactylus appears to be the most stable. Little or no differences
occur between even widely separated populations. One exception is
the collection of nine specimens from Carroll County, Va., in which all
juveniles and adults (as well as males and females) possess strong
lateral spines on the carapace. This collection (map 2, no. 240) was
made in a tributary of Crooked Creek (New River drainage). All
other specimens including those from Reed Island Creek, an adjacent
tributary to the New River, lack these lateral spines. Additional
collecting in this area will be necessary before further comment may
be made.

The largest available specimen of C. 1. chasmodactylus is a female
with a carapace length of 55 mm.; the largest chela I have seen
measures 78 mm. in length by 26.5 mm. in width; only the hand of
this animal is available. Size, however, must be disregarded as a
diagnostic feature since many first-form males have a carapace length
as little as 30 mm. and, as noted, the largest first-form male of C. l.
longulus (the smallest of the subspecies) is 34 mm.

SPECIMENS EXAMINED.—I have examined approximately 275 speci-
mens in 56 collections from 44 localities; all are confined to the New
River drainage system of North Carolina, Virginia, and West Virginia.

Entirely confined to the New River system, the known range of
C. 1. chasmodactylus extends northward to the upper reaches of the
Greenbrier River in West Virginia (map 2, nos. 261-268); the most
southern record is from Watauga County, N.C., in the headwaters
of the South Fork of the New River (map 2, no. 229). Both eastern
and western boundaries are formed by those Appalachian Mountains
delimiting the New Valley.

There appears to be complete geographic isolation between this
and the other two subspecies. In Alleghany County, N.C. (map 2,
no. 215), C. l. chasmodactylus was collected from a tributary of the
New River only a few miles from where C. 1. longulus was collected
in the Yadkin drainage in Wilkes County (map 2, no. 105), but
nowhere in the entire range is there evidence of sympatry.

Ecological Distribution

The general habitat of the species Cambarus longulus has been
noted by various investigators (Ortmann, 1913, pp. 375, 376; 1931,
pp. 119, 123; Hobbs, 1950, p. 349; Reid, 1961, p. 249). Members of
C. longulus are highly restricted stream or river inhabitants that
live under or between rocks, away from the shore, in moderate to
swiftly flowing, cool to cold water. The species does not, from all

NO. 3544 SUBSPECIES OF CAMBARUS LONGULUS—JAMES 21

accounts, frequent quiet pools, springs, or mountain ‘feeder brooks.”
Stream beds are typically of rock-strewn sand or gravel and relatively
free of silt deposits.

Most specimens of C. l. longulus were taken from riffle areas in the
lower mountain or upper piedmont streams. The water, clear or
comparatively so, ranges from 2°-24° C. The current, over riffles,
has been recorded as moderate to rapid. In streams with both pool
and riffle areas, C. 1. longulus was found only in the riffles. Stream
beds of sand or gravel and strewn with rocks are characteristic of
the habitats in which this crayfish usually is found. The size of the
stream seems to have little, if any, influence on whether or not C. J.
longulus frequents it.

The subspecies C. 1. chasmodactylus varies from the typical sub-
species in seeming to prefer larger, often turbulent streams. It is
found abundantly in Reed Creek, Wythe County, Va.; typically, this
animal, as described by Ortmann (1931, p. 119) lives “in the usual
way under stones in flowing water [italics mine].”’

Of the three subspecies, C. longulus longirostris is by far the least
ecologically restricted. A cursory examination of the wide distri-
bution (maps 1, 2) should suggest the apparent adaptability of this
crayfish to the larger streams and rivers as well as to the smaller
tributaries. In McMinn County, Tenn., it was found to be numerous
in beds of Nasturtium sp. in a spring run; nearby, occasional animals
were found in debris littering a stream bed of silt and sand, where there
were but few rocks.

Serological Affinities

Johnson, in 1957 and again in 1959, using agar diffusion and tube
precipitation techniques, serologically compared, among others, the
three subspecies of Cambarus longulus (Cambarus sp. = C. l. chasmo-
dactylus). Antigen (crayfish serum) and antibody (rabbit anti-
crayfish serum) reactions, when compared (1957, p. 182; 1959, p. 183),
indicate the close affinities of the three.

Summaries of Ratios

The tabulation below summarizes the collective quantitative data
(figures of each column represent ratios of measurements of selected
characters; upper row, minimum ratio; middle, maximum; lower,
average):

Characteristic Ratio longulus longirostris chasmodactylus
0. 10 0. 19 0. 17
rostrum L/carapace L 0. 26 0. 28 0. 28
0. 20 0: 22 0: 25
itispl ic: 1.5

22 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Characteristic Ratio longulous longirostris chasmodactylus

rostrum L/W Dae, aS 22,
1.6 1.8 19

0. 40 0. 78 0. 69
rostrum L/post orbital 11 pt ey
Ww* 0. 95 1.0 1.0
2.3 2.3 3. 5
areola L/W 5. 9 5. 9 6. 0
3) 4.0 4.4

0. 20 0. 30 0. 34

areola L/carapace L 0. 42 0. 39 0. 47

0. 37 0. 36 0. 37
Lise 1% iG
antennal scale L/W Se 350 2.4
2. 5 3.3 2.2

0. 53 0. 52 0. 48

palm W/dactyl L 120 0. 88 0. 74

0. 80 0. 72 0. 61
1.5 2.6 2. 6
hand of chela L/inner 3. 6 4.4 4.7
palm margin L on 3.1 3. 6
1.0 145 E2
hand of chela L/dactyl 2. 6 1.8 1.8
L 1 ae Aa a)
1.3 1.5 2510
dactyl of chela L/inner 2.0 3. 0 3. 4
palm margin L 1.6 19 2.4

*Distance between anterolateral margins of postorbital spines or ridges.
Summary

As a result of the present study, three allopatric subspecies of
Cambarus longulus Girard have been recognized. Cambarus longulus
longulus is found in the lower mountain and piedmont streams of
the Atlantic drainage, from the James south to the Yadkin Rivers.
Cambarus longulus chasmodactylus, herein described, is restricted to
the New River system from North Carolina to western West Virginia.
Cambarus longulus longirostris frequents the Tennessee drainage
system above Wilson reservoir, and the Coosa River drainage in
northwestern Georgia and eastern Alabama.

Although all three subspecies are associated with riffle areas of
lotic habitats, C. longulus longirostris appears to have a broader
ecological tolerance than do the two other subspecies.

While C. l. chasmodactylus exhibits few variations throughout its
range, O. 1. longulus, perhaps because of its low vagility, has within
its range several recognizable local populations.

It is concluded that these three crayfishes represent offshoots
of a common stock which, while believed to be not too far removed
from the primitive members of the genus, have become, for the most
part, ecologically, highly specialized for life in restricted areas of
“young” but not “infant” streams.

NO. 3544 SUBSPECIES OF CAMBARUS LONGULUS—JAMES 23

Literature Cited

BRIMLEY, C. 5S.
1938. The insects of North Carolina, 560 pp.

ERICHSON, W. F,

1846. Ubersicht der Arten der Gattung Astacus. Arch. fir Naturg. vol. 12,

no. 1, pp. 86-103.
Faxon, WALTER

1884. Descriptions of new species of Cambarus: To which is added a
synonymical list of the known species of Cambarus and Astacus.
Proc. Amer. Acad. Arts Sci., vol. 20, pp. 107-156.

1885a. A revision of the Astacidae. Mem. Mus. Comp. Zool., vol. 10, no. 4,
pp. 1-186, 10 pls.

1885b. A list of the Astacidae in the United States National Museum. Proc.
U.S. Nat. Mus., vol. 8, no. 23, pp. 356-361.

1890. Notes on North American crayfishes, family Astacidae. Proc. U.S.
Nat. Mus., vol. 12, no. 785, pp. 619-634.

1898. Observation on the Astacidae in the United States National Museum
and in the Museum of Comparative Zoology, with descriptions of
new species. Proc. U.S. Nat. Mus., vol. 20, no. 1136, pp. 643-694,
pls. 62-70.

1914. Notes on the ecrayfishes in the United States National Museum and in
the Museum of Comparative Zoology, with descriptions of new
species and subspecies to which is appended a catalogue of the
known species and subspecies. Mem. Mus. Comp. Zool., vol. 40,
no. 8, pp. 347-427, pls. 1-11.

FLEmine, R. 8.

1938-1939. The larger Crustacea of the Nashville region. Journ. Tennessee
Acad. Sci., vol. 13, no. 4, p. 296-324; vol. 14, no. 2, p. 261-264;
vol. 14, no. 3, pp. 299-324.

Fow ter, H. W.

1912. Crustacea of New Jersey. Rep. New Jersey State Mus., 1911, pt. 2,

p- 31-650, pls. 1-150.
GIRARD, C.

1852. A revision of the North American Astaci, with observations on their
habits and geographical distribution. Proc. Acad. Nat. Sci.
Philadelphia, vol. 6, pp. 87-91.

Hagen, H. A.

1870. Monograph of the North American Astacidae, 3. Cat. Mus. Comp.

Zool., no. 3, pp. 1-109, pls. 1-11.
Harris, J. A.

1903. An ecological catalogue of the crayfishes belonging to the genus

Cambarus. Kansas Univ. Sci. Bull., vol. 2, no. 3, pp. 51-187.
Hart, C. W.

1952. The exoskeleton and musculature of the appendages of the first three
abdominal segments of C. longulus longulus Girard (Decapoda,
Astacidae). Virginia Journ. Sci., vol. 3, pp. 39-48.

Hay, W. P.

1899. Synopses of North American invertebrates, 6: The Astacidae of North

America, Amer. Nat., vol. 33, pp. 957-966.

24 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

fel@rpss} 18l, JEL, die
1942. A generic revision of the crayfishes of the subfamily Cambarinae
(Decapoda, Astacidae) with the description of a new genus and
species. Amer. Mid. Nat., vol. 28, no. 2, pp. 334-357.
1950. Observations on the ecological distribution of three Virginia cray-
fishes. Virginia Journ. Sci., vol. 1, n. s., no. 4, p. 349.
1959. In W. T. Edmondson, Fresh water biology, xix+1248 pp., figs.
3.1-46.1.
JOHNSON, R. M.
1957. The Agar diffusion technique as applied to the study of serological
relationships among crayfishes. Virginia Journ. Sci., vol. 3, pp.
177-184.
1959. Serology of crayfishes. Virginia Journ. Sci., vol. 10, n. s., no. 8,
pp. 181-185.
JoHNSON, T. L.
1960. Histological observations on the sperm duct of Cambarus longulus
Girard. Trans. Amer. Micros. Soc., vol. 79, no. 2, pp. 229-238.
NEwcomesgE, C. L.
1929. The crayfishes of West Virginia. Ohio Journ. Sci., vol. 29, pp.
267-288.
OrTMaANN, A. E.
1902. The geographical distribution of freshwater Decapods and its bearing
upon ancient geography. Amer. Philos. Soc., vol. 41, no. 171,
pp. 217-400, figs. 1-8.
1905. The mutual affinities of the species of the genus Cambarus, and their
dispersal over the United States. Proc. Amer. Philos. Soc., vol.
44, no. 180, pp. 91-136, 1 map.
1913. The Alleghenian Divide and its influence upon the freshwater fauna.
Proc. Amer. Philos. Soc., vol. 52, no. 210, pp. 287-390, 3 pls.
1931. Crawfishes of the Southern Appalachians and the Cumberland
Plateau. Ann. Carnegie Mus., vol. 20, no. 2, pp. 61-160.
ParisH, C.
1948. A contribution toward a knowledge of the crayfish Cambarus longulus
dasydactylus, with special reference to the annulus ventralis.
Unpublished master’s thesis, University of Virginia.
Rep, G. K.
1961. Ecology of inland waters and estuaries. xvi+375 pp.
Snopaerass, R. E.
1952. A textbook of arthropod anatomy, viii+363 pp.

U.S. GOVERNMENT PRINTING OFFICE:1966

PROC. U.S. NAT. MUS. VOL. 119 JAMES—PEATE 1

Cambarus longulus longulus: a, dorsal view of antennal scale; 7, /, lower surface of chelae

showing extreme variation in setaceous condition; k, m, upper surface of same; h, mesial
view of first pleopod of first-form male; 7, lateral view of same; 9, lateral view of cephalic
region of carapace; t, dorsal view of first-form male. C. longulus longirostris: c, dorsal
view of antennal scale; f, g, first pleopod of first-form male; , 0, chelae; 7, lateral view of
cephalic region of carapace; s, dorsal view of first-form male. C. longulus chasmodactylus,
new subspecies: b, antennal scale; d, ¢, first pleopod of first-form male; g, lateral view of
cephalic region of carapace; u, annulus ventralis (typical of all C. longulus); v, dorsal
view of holotype; w, x, lower and upper surface of chela of juvenile; y, z, lower and upper
surface of chela of adult; zz, upper surface of chela showing key characteristics (compare
with C. 1. longulus, fig. mm.). (Scale line equals 30 mm. where given; other photographs
not to scale.)

Proceedings of

the United States

National Museum

SMITHSONIAN INSTITUTION + WASHINGTON, D.C.

Volume 119 1967 Number 3545

THE LIZARDS OF ECUADOR,
A CHECK LIST AND KEY

By JAmes A. PETERS
Curator, Division of Reptiles

This paper constitutes the second of a series devoted to the establishment
of a framework upon which additional studies on the ecology and zoogeog-
raphy of the Ecuadorian herpetofauna can be based. The first paper dealt
with the snakes (J. Peters, 1960), and later studies will be concerned with
the amphibians. The principle established in the earlier list has been
abrogated, at least in part, since I have been forced to revise the genus
Ameiva strictly on the basis of the Ecuadorian political unit. But I repeat
my earlier opinion that such analyses are dangerous and can easily result
in perpetuation of difficulties.

Methods

The method of organization in this check list is the same as that of the
list of Ecuadorian snakes (J. Peters, 1960). The genera are presented
alphabetically, and the species are alphabetical within their genus. It is,
I think, true that the average user of the list will be interested in ease and
speed of use, not in my contribution to the intricacies of the higher categories

I

2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

of lizard classification (the basic phylogenetic position of the genera con-
cerned is presented on p. 3). For each taxon I have presented a very brief
synonymy, beginning with a citation to the original description of that
taxon and its type locality, plus the holotype and its location in paren-
theses. Similar citations and information are included for all taxa
described from Ecuador that have been synonymized with other species.
I have attempted to include all erroneous records in the literature based
upon Ecuadorian specimens, listing them within the synonymy of the species
to which they properly belong under the name used by the author, and
separated from the citation by a period and dash. Obviously, this is im-
possible where the specimens have not yet been reexamined, either by
myself or by someone discussing them in print, and without question some
erroneous identifications have served as the basis for inclusion of some taxa
within this list.

An asterisk following either a generic or specific name indicates that that
taxon has not yet been collected in Ecuador. In all cases there is reason
to anticipate the occurrence of these taxa in Ecuador, but it should never
be assumed that all likely members of the fauna have been anticipated.
The failings attendant upon all keys exist here as well. Keys must be
written on the basis of the known or expected variation, and it is im-
possible to include all of the individual divergencies from type of which the
genome is capable. Merely to reach a name does not put the final stamp
of adequate identification on the specimen, but only provides the basis for
a thorough check against the available descriptions and comparative
material. Two species may share all of the characters utilized in the
dichotomies and still be strikingly different from one another.

The catalog number assigned to the type specimen or series is given in
parentheses following the citation. The abbreviations used refer to the
following museums:

AMNH_ American Museum of Natural History, New York.

ANSP Academy of Natural Sciences, Philadelphia.

BerM Zoologisches Museum, Berlin, Germany.

BM British Museum (Natural History), London.

GottM Géttingen Museum, Géttingen, Germany.

HM Zoologisches Museum, Hamburg, Germany.

IRB Institut Royale d’Histoire Naturelle de Belge, Brussels.

LeyM Museum, Leyden, Netherlands.

MCZ Museum of Comparative Zoology, Harvard University, Cambridge, Mass.
MunM_ Zoologische Sammlung des Bayerischen Staates, Munich, Germany.
PM Muséum National d’Histoire Naturelle, Paris, France.

RMS Royal Museum, Stockholm, Sweden.

TurM ‘Turin Museum, Italy.

UMMZ University of Michigan Museum of Zoology, Ann Arbor.

USNM _ United States National Museum, Washington, D.C.

VM Naturhistorisches Museum, Vienna, Austria.

NO. 3545 LIZARDS—PETERS 3

Classification

Since the check list is arranged entirely on an alphabetical basis to
facilitate quick and effective use, it tells nothing about the relationships
and phylogenetic position of the genera and species. The list below will
fill this gap, placing the genera in their familial positions. It will be noted
that I have not indicated position of the families within the higher categories.
This is a consequence of the dilemma posed by the two most recent reviews
of lizard classification, published by Romer (1956) and by Underwood
(1957). In an analysis of the Ecuadorian lizards alone I find that these
authors differ either in the level of category or in the name used (or both)
in 28 of 32 instances. To accept either would indicate a completely un-
justified negation of the other, unless done on the basis of adequate and
thorough review, which I have not undertaken. I follow Underwood
(1954) in recognizing the Sphaerodactylidae as a family distinct from the
Gekkonidae.

IGUANIDAE
Anolis, Basiliscus, Enyalioides, Enyalius, Iguana, Morunasaurus, Ophryoessoides,
Plica, Polychrus, Proctotretus, Stenocercus, Tropidurus, Uracentron
GEKKONIDAE: GEKKONINAE
Phyllodactylus, Thecadactylus
SPHAERODACTYLIDAE
Gonatodes, Lepidoblepharis, Sphaerodactylus
‘TEIIDAE
Alopoglossus, Ameiva, Anadia, Arthrosaura, Callopistes, Dicrodon, Echinosaura,
Ecpleopus, Euspondylus, Iphisa, Kentropyx, Leposoma, Macropholidus,
Monoplocus, Neusticurus, Ophiognomon, Pholidobolus, Prionodactylus,
Proctoporus, Ptychoglossus, Tupinambis
SCINCIDAE
Ablepharus, Mabuya
ANGUIDAE: DIPLOGLOSSINAE
Diploglossus
AMPHISBAENIDAE
Amphisbaena

Omissions

There are many species that have been recorded as members of the
Ecuadorian fauna on the basis of erroneous identification of individual
specimens. Where possible, these errors have been placed in their proper
species in this check list and can be found in the index. This is based
either upon a reidentification appearing in the literature, or upon my own
reexamination of the specimens. In addition, however, I have presumed
to omit several things, even though a recheck has not been possible. Thus,
I have omitted LEcpleopus gaudichaudii Duméril and Bibron, which
was recorded from Ecuador by F. Miiller (1882, p. 157), since it is quite

4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

unlikely, although not impossible, that it actually occurs in Ecuador.
The same statement applies to Anolis pulchellus Duméril and Bibron,
which Cornalia (1849, p. 308), recorded from Guayaquil, based with
little question upon a misidentification. A discussion on the occurrence
of Polychrus liogaster Boulenger has been published earlier (J. Peters, 1959).

Acknowledgments

Again I must record my debt to friends, colleagues, and institutions whose
assistance facilitates my efforts. It becomes increasingly obvious to me
that completion of a work of this type is totally subject to the continued
good will and cooperation of many people, and I am pleased that I continue
to work without incurring their disfavor.

Dr. Gustavo Orcés-Villagomez, of the Escuela Polytecnica Nacional,
in Quito, continues to provide facilities, materials, specimens, knowledge,
and friendship. His constant tolerance of my faults, both while I shared
his laboratory during my visits to Ecuador and while I bombard him with
impossible requests during my stays at home, has never ceased to amaze
me. The !ate Robert Copping, of the British Embassy staff in Quito, was
more than congenial as a fellow collector, host, and intermediary in prob-
lems, political and otherwise. Robert Mullen and Peter Spoecker, students
at Valley State College, and Manuel Olalla, an Ecuadorian citizen, spent
the summer of 1962 in the field with me, enduring mule kicks, fungus in-
fections, cold mountain passes, and occasional beer shortages, all in my
behalf. Spoecker and Stephen Austin have spent many hours checking
the key against Ecuadorian specimens.

M. Boesman, Doris Cochran, J. A. Cochrane, J. Eiselt, Norman Hartweg,
Werner Ladiges, Edmond V. Malnate, George Myers, Thomas Uzzell,
Charles Walker, and Ernest E. Williams have loaned specimens, provided
information, or made work space available at their respective institutions.

I have now made three trips to Ecuador for collecting purposes; the
first, in 1954, supported by the Penrose Fund ofthe American Philosophical
Society, the second, in 1958-59, under tenureship of a Fulbright Professor-
ship, and the third, in 1962, under the sponsorship of the National Science
Foundation, Grant No. G-21010. To the authorities in charge of each,
my sincere thanks.

Key to Genera of Lizards Known or Expected in Ecuador

(Asterisk indicates genus or species has not yet been collected in Ecuador)

i) Feet)\(at least the forefect) presentsjeye delinite 7) =) i ee ee 2
Without feet; eye reduced to an indefinite point covered by the skin.
Amphisbaena

046 LIZARDS—PETERS 5

Eyelids absent, dorsal scales small and numerous, usually granular and not

NDT A LS prema ata AL roto ake ee ed al canoe RN rahe NPT pts Rm a Maas Te gles “Pc
Eyelids present (or, when lacking, the scales of the body are arranged in 13-15

rows/and are broad, smooth, and imbrieate) . .. . 20%) tee odes, 99
No ,adhesive.;mads on LOes, <7 <)))2, ticcirinda ta Mies ane eae eee
Hoes with adhesive pads, below “. 5 di, valwl-Olane fae bate oe tiles yl
INailscovercanby.a/sheathiol scalesy & cauyraris, vanirualesnt-l evisiwes frets Es. 7O
INailseuncovercdss witout arsheathy paula anne nnn en Gonatodes,
Sheath of nail symmetrical, as seen from above (fig. 1b) ........ 6
Sheath of nail asymmetrical, as seen from above (fig. la). . . Sphaerodactylus
Five scales in nail sheath (fig. 1b)... .... =... . . Pseudogonatodes*
Six scales in nail sheath (fig. Ic) . . ...... 2... . . Lepidoblepharis

Ficure 1.—Nail sheaths: a, Sphaerodactylus, dorsal view; b, Pseudogonatodes, five scales;
c, Lepidoblepharts, six scales. Tail with rings of spiny scales: d, Morunasaurus
annularis.

a:

10.

MW.

Digits with two rows of transverse lamellae below throughout length . . . 8
Digits with single row of lamellae or tubercles below, two large plates under

EXPAMOCatPIOUGISTE NL AM «we uy ckissh oa sce CORA . Phyllodactylus
Digits only partially dilated, ultimate phalanx compressed, arising from

dorsum: or dilated part of digit... 2. 2 = & . Hemidactylus*'
Digits totally dilated, ultimate phalanx not compressed but part of dilation.
Thecadactylus

Top of head with granular or irregular flat scales, the largest usually arranged
in semicircles over each eye; lacking a median frontal between the eyes;
tonsue usually fleshy:and not, extensible.) eqs ai4em . soe WO
Top of head with regular flat plates; a median frontal between the eyes;
tongue not fleshy although extensible and bifurcate . . . ..... . 30
ailyprovided|-with spines arranged injrings, =. siege, Ge So. We eee)» lil
AcaileWithOltispiny TAS MEN id cee pice di germ, ug Ate ieey Khaw at eal a PLL
Tail cylindrical or compressed vertically, rings of spiny scales separated
ito SIG) ease tec Near oat Ae a ae PRM (a Nya 06 A! ot ellen ah meatal 2
Tail flattened in a horizontal plane, all caudal scales spiny . . . . Uracentron

1 Known from both Colombia and Peru.

12:

13:

14.

15.

16.

Ficure 2.—Toe of hindfoot: a, Basiliscus; b, Corythophanes.

Vi
18.
19;
20.
21.
22.

23.

PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119
A strong transverse gular fold. . he OY 13
No transverse gular fold . . Stenocercus
Dorsal crest present . Enyalioides
Dorsal crest absent. aya . Morunasaurus
Scales below ear subequal in size . . ee
A large, round, flat scale below ear . PASE Se ARO eS Iguana
Head produced posteriorly forming a vertical fin (in female, especially young
ones, sometimes very inconspicuous) . . B
Head not produced posteriorly. .......-.-.. » ales
Toes of hindfoot with fringe of flat scales (fig. 2a). . . Basiliscus
Toes of hindfoot without fringe (fig. 2b) . . . Corythophanes*
Be
UN
AD
WAU
A
WAU
WY
UI
Wu
AU
WAU
LN
UAT
WAY
UI
NJ
UN
ONL ah
UXT

species; d, Tropidurus holotropts.

Toes not dilated but quite cylindrical or compressed. .
Toes dilated and flattened. .

A raised dorsal crest or slightly enlarged row of vertebral scales .

Dorsal crest absent .

No femoral pores.

Femoral pores present w= alien ent esa bo ee) GAGs
A transverse gular fold extending completely across throat .
No transverse gular fold extending completely across throat
Infradigital lamellae distinctly keeled . a5
Infradigital lamellae smooth or vaguely keeled . .

Anterior maxillary teeth longest .

Maxillary teeth subequal .

@ccipital’small (Gig w26)hs 4. ea) eae

Occipital greatly enlarged (fig. 2d). . .

Occipital: c, Leiocephalus

ibe lS
. Anolis
rl9
. 26
5820

. Enyalioides

ed
123

ee

. Enyalius
Plica?

. Tropidurus

ioe

. Tropidurus

2 When this key is used for generic identification of non-Ecuadorian forms, Uranoscodon
will key to Plica, since they are identical in external characters used. Uranoscodon lacks
a sternal fontanelle, Plica has one. It is unlikely that Uranoscodon occurs in Ecuador.

NO. 3545 LIZARDS—PETERS Z

PPro dil rontiOnshOulder es” oye.) ale snk Wh ie ar Maw tet ooh eee | IDB
Notoldinuronttor shoulder.) 0.5: . a. = “yer bet inn Oplryoessotdes
ZoemUipper head scales -keeledi. ru 6s goal Sus cp nye icd sei rey 4 ey E roctotnetas
Wiper aead sealesysmiooth, 2. 4. Sb eye t) espe pan as 4 a WO EEROCETCUS,

Poa Hemoralpporesra ascntte seer sie alae. Mab: tyr sb fot) dys peSetre lage fh eB
Femoral pores present. . . . . aha cleeceh cmesnol|

27. Ventrals large, square, in transverse end jengitudinati rows aa distinct from
venvesmall. pranular corals.) 6 5.9) 2, -) 6), ~) 6, cle, esi ist AMAVE
Ventrals not as above... . ‘ pl Beaitesdens Bays Paiyehras

28. A transverse gular fold, earapletetar across divans eee Suicere trie at 2329
No complete gular fold, although a fold in front of ahoulden may extend part
Wayaomtosthnoate gyal Ages is je a lod oad whist is iaeeals) OS

29>, Dorsals keeled and timbricateys <4). )<)) scar ny -A ys Gpagidans (torquatus ealy)
Dorsals smooth and granular. . . . . ts. (obese Gallopestes:

30. Body scales not semicircular, not very eabricate (Githonen sometimes they are
arranged in 16 or fewer rows); lack bony plates underlying the scales . . 31

Body scales semicircular, usually smooth or slightly keeled and striated (fig.

3a), and very imbricate, with underlying bony plates (fig. 3) . . . . 60

Bony Ptare

Posrerion Enp

a b

Figure 3.—Body scales of Diploglossus monotropis: a, dorsal view; }, cross section.

31. Dorsal scales heterogeneous; large keeled scales mixed in with smaller granular

onesé 4)... Pe eat eden Seley Ud 1 i ENE Re ot ee A Lem Oe
Dorsal scales hetueeeneous: ee ee : ; HA erat Ome

32. Males with 10 or fewer femoral pores; inner eer at least aigualy
Keele Soy. 2 ats s ie urs ‘ Fc. Ras Oo

Males with 12 or more fempral pores; inner ventvals smipot . . Neusticurus

33. Nostril in suture between two nasals; ventrals small, elongate, subquadrangular;
doublecollar foldi 220". %) Jian de eae nae Aeneas
Nostril in single nasal; ventrals ae eee nace ence fold . Echinosaura

34. Dorsal scales usually granular, always much smaller than ventrals. . . . 35
Dorsal scales as nee as, or only slightly smaller than, ventrals, not gran-
ibaa ee ee Seer cen feet eh ark SOME Pm, SOARS tr ea

35. More than 20 rows af ventral rates! Sie REL ADRS D BOS NSIC) SPOKE: TPN PR TO)
Sine Como LOwaromventraliscaless yy sae. Flys vat ie ie tay fle a. alee NOT

20.) Pall strongly bicarinate’dorsalty 3 toe). 2 Ae. ek S@rocodelurus*®

Tail rounded, no raised ridges dorsally ...... .. .. . .Tupinambis

Sil.

38.

hb

40,

41,

42.

43.

44,

PROCEEDINGS OF THE NATIONAL MUSEUM vou. 119

Ventral scales‘ keeled \. <. 40,5) en SE Ot Ret ae ee
Ventral scales'smooth... < 0x. Ws, 2 ep era a) ee Pe eee
Femoral. pores present). 502 ce a gees. @ Dn a entropy
Femoral pores absent. . .. . oe oe he Monoplocus:
Ventral surfaces of limbs and tail pale, contrasmae strongly with the dark
belly ie. acdc . 40
Ventral surfaces of ibs and oil dark, not contrastine atronely witht venter of
bodys). ot5 5... elt US Ameina

Patch of enlarged seals eae on eo inoat beeween ‘he eaples BE; jaw (fig. 4a);
all supraoculars in contact with central head shields; males with heavy
spines on sides of anus. . . . . . . Ameiva

No enlarged patch of scales racaially on renroae beaween enales of jaw (fe. 4b);
last two supraoculars separated from frontal and frontoparietal by row

of granules; males with no heavy anal spines. . . . . . . . . Ducrodon
Five toes’on forefoot $F)" ) £5 REL SE Ce Oe eae
Fewer than five toes, if any, on forefeor OPIS? FOE IDA ee 2
No greatly enlarged dorsal and ventral scale rows. . . . 4a
Dorsum and venter with two rows of extremely broad scales, ucparated Aeterauy

by four.rows ot scales: «fy Aca, te 2, a ee es
Avclawion everysinger 0) 5..x, 2 Be A ced: 1 ed ee eee
innermost. tmeenlacks:arclaw. 4 ~ os 2a. = -1 os = alu }Galusemcopass
INotpretrontals (ie. !4c)i (oA ps ee Peo Wot a EY eh Se ee
Prefrontals present\(iies 4d)... «Peete es ce le ee ee

c

Ficure 4.—Scales medially on throat between angle of jaws: a, Ameiva edracanitha; b,
Dicrodon guttulatum. Dorsal view of head: c, Pholidobolus montium; d, Neusticurus

strangulatus.

45, Dorsalsshéxazonalorssubhexagonal..- 565 2.0 sais ols aces ee ee
Dorsals rectangular... . 2 st ape suas acy dy crackoberus,
46. No clear separation between vena aad doris. .. . . . « Macropholidus

Dorsals and ventrals separated by two or more rows of granular scales
Pholidobolus
47. Dorsal and ventral scales similar, aaa dorsals in transverse and
longitudinal rows. . . . etl kcal Phe) tude een cee

Dorsal scales not arranged in oeeidieal TOWSic 2 ty, pac) ae ee ean mee

NO. 3545 LIZARDS—PETERS 9

48.

eos

Dorsalscales/strongly keeled /)-ai4n-) yi) foe (0a 2 an) a Cencoxaura®
Dorsal scales smooth or weakly keeled . 2 2) 10: 5%: . 0... 5 sv Anadia
Dorsal scales considerably narrower than ventrals; in transverse rows (fig.

Eid) PPP ML eerik Fe mci ros is, tai eeu eta ths hos ly a Pelvic TNL Artal . 50
Dorsal scales in oblique rows, (fig. 54) notably imbricate and keeled . . 56

50.

51.

a2.

53.

OAs

oo.

56.

57.

58.

D9.

60.

Ficure 5.—Dorsal scales: a, Alopoglossus festae; b, A. copit.

Dorsal scales keeled or with the center of each scale distinctly raised along

TAGES ee ee. ee ee OY ee ee, Parnes Meee to COM
Dorsaliscales'smooth or weakly keeled: "> 295592 32 5 Gc = 4 se. 98
Lateral and dorsal scales subequal in size; laterals hexagonal, flat. . . . 52

Lateral scales much smaller than dorsals, tubercular or granular. Prionodactylus
Ventral plates rounded posteriorly, overlapping next row; dorsal caudal

scales sharply pointed, overlapping... ....... . . . Arthrosaura
Ventral plates square or truncate posteriorly, not overlapping next row;
caudal scales truncate, not overlapping, not pointed. . . . . Ptychoglossus
Bateral scales:much smaller than/dorsals}) 7.09 ).9. 059. 8? ne. OF
Lateral scales and dorsal scales subequal, ventrals and dorsals not separated
byzzonelolsmaller/scaleg 5) 2.) nyo eek Gp eis nae
Strong collar fold; male: with femoral'pores . . ..........-. =. 99
Slight or weak collar fold; male without femoral pores. . . . . . Ecpleopus
Head only slightly larger than neck; 5-7 posterior pre-anals; few large flat
temporals Ape hog eee nein Cay he . Euspondylus
Head clearly larger than neck; less than 5 posterior pre-anals; many small
granular temporals..). :. . = < : Shao aoe ta ee NELESTICUTUES,

Head scales with many longitudinal striations; interparietal longer than
PARLCEAIS VSO SieEy Vane aee. vin that) yy MSGi meee Mhunead Peman . Leposoma
Head scales smooth, may have some striae or keels on the interparietal and

parietals; interparietal equal in length to parietals . . . . . . Alopoglossus
Dorsum: with moreithan two rows of scales. : =. . 2... - «...- =» .:08
Dorsum with two greatly enlarged rows of scales . ...... . . -Dphisa
Rear tinb.radimentary or absents\i5 =) 40s Vo 5 ee ke Sa fens ey ee
Rear limb well developed, pentadactyl . . . . .. . . . Gymnophthalmus*
An unpaired frontonasal separates nasals. . . ..... =... +. . Bachia*®
Nasals forming a suture on dorsal midline of head. . . . . . . Ophtognomon
Atleastitwounterasals: a. ei wre Bs SPS ee Ee OL
Initermasal/ single 4) Sees 265 Ge oy ee) ee Gymnophihalmust

219-943—67——_2

10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

61. One pair of internasals present; body usually striped. ........ . 62
Two pairs of internasals present; body large and not striped. . . Diploglossus

62. Frontoparietals and interparietal fused into a single shield . . . . Ablepharus
Frontoparietals and interparietal separate, distinct (although latter may fuse
with parietals))is Ais 29 t. SS Oe eH ins atitoe. eee eeanaia

63. Scales on tail arranged in concentric. eel FIMGS 2. 5 2 a = a Sbenocercus

Scales on tail not arranged in vertical rings but in diagonal series . Proctotretus

Genus ABLEPHARUS Fitzinger

Ablepharus boutonii poecilopleurus Wiegmann

Ablepharus poecilopleurus Wiegmann, 1835, Nov. Act. Acad. Caes.
Leop. Carol., ‘vol. 17, pt. 1, p. 202.— Pisacomats;. Perus(type

unknown).
Rance: Islands off west coast of Peru and Ecuador; Sandwich Is.,
Savage Is.
Genus ALOPOGLOSSUS Boulenger
1. Gulars not arranged in two longitudinal rows; four pre-anal scales. . . . 2
Gulars transversely enlarged and arranged in two aa rows; usually
3 pre-analscales, 42) ys. . festae
2. Scales on the side of the neck ines ge Aad Geel: wees on the ene half
ofthe. dorsum in longitudinal mows. yey) 91.04. 4) yoneue> eel alie os er CODES
Scales not as above... . 540g <6 eo ea ee Cea
3. Scales on the side of the neck pean bleacst sranulens > ky ween hyde BDC Lee

Scales on the side of the neck keeled, imbricate, not granular . .carinicaudatus
Alopoglossus buckleyi (O’Shaughnessy)

Leposoma buckleyt O’Shaughnessy, 1881, Proc. Zool. Soc. London, p.
233, pl. 22, fig. 2. Canelos, Ecuador (BM 1946.8.31.66, male).

RANGE: Amazonian Ecuador and Peru; has also been recorded from
Pacific slope of Ecuador (Babahoyo, by Werner, 1910, Mitt. Nat.
Mus. Hamburg, vol. 27, p. 30).

Alopoglossus carinicaudatus (Cope)

Leposoma carinicaudatum Cope, 1876, Journ. Acad. Nat. Sci. Philadelphia,
ser. 2, vol. 8, p. 160. Valley of Rio Marafion, Peru (type unknown).

RanGE: Periphery of Amazon Basin, in British Guiana, Ecuador, and
Peru.

Alopoglossus copii Boulenger

Leposoma carinicaudatum.—O’Shaughnessy, Proc. Zool. Soc. London,
1S31-sp. 233.
Alopoglossus copii Boulenger, 1885, Cat. Lizards British Mus., vol. 2,

NO. 3545 LIZARDS—PETERS hat

p. 383, pl. 20, fig. 1. Pallatanga and Canelos, Ecuador (BM 1946.8.31.-
58-59; 1946.9.8.17-18).
Rance: Amazonian slopes of Ecuador.
Alopoglossus festae Peracca
Alopoglossus festae Peracca, 1904, Boll. Mus. Zool. Univ. Torino, vol.
19, no. 465, p. 7. Vinces, Ecuador (TurM 2875).
RANGE: Pacific slope of Ecuador.

Genus AMEIVA Meyer

1. Frontal plate divided into smaller scales, usually both longitudinally and

transversely; prefrontals absent, or if present, notin contact. ......2
Frontal plateventiressprefrontallscales;in/contact. )9.) 2 wi 5 244

2. Mesoptychium with enlarged scales; humerus with row of very large, flat,
smooth scales on anterior aspect... . cyan an SORE REL
Mesoptychium with subequal scales; humerus wt 1 =F rows of sliently enlarged,
strongly keeled scales... . . A) ee es Be nord pests

3. Eight rows of ventrals on much of ven ee ancl aol peas prefrontals;
dorsal head scales not ridged and pitted .... . sh keuinisy os SON CESS,

Six rows of ventrals on all of venter; several small eal ee prefron-
tals; dorsal head scales heavily ridged and usually pitted . . . septemlineata

4, Eight rows of ventral plates; males with group of spines on each side of pre-anal
FEPTION «=. ; : sh es) pe cedracantha

Ten to twelve rows of renal ie no spines in ane mreanal REGION 45)

5. Last two or three supraoculars bordered entirely by granules, not in contact
with frontal and frontoparietals . ... . . . . . . btfrontata divisa*

No supraoculars except tiny fourth bordered ey by granules, in contact
with frontal and frontoparietals .........2. 2... =. . ameiva petersi

Ameiva ameiva petersi Cope
Ameiva petersi Cope, 1868, Proc. Acad. Nat. Sci. Philadelphia, vol. 30,
p. 99. Napo or Marafion, Ecuador (USNM 6639, now lost).
Rance: Upper Amazonian Basin.
Ameiva bridgesii (Cope)
Holcosus bridgestt Cope, 1869, Proc. Acad. Nat. Sci. Philadelphia, vol.
20, 1868 (1869), p. 306. Ecuador? (ANSP 9651).
Rance: Northwestern coastal areas of Ecuador; Chocé of Colombia,
Gorgona Island.

Ameiva edracantha Bocourt

Ameiwwa edracantha Bocourt, 1874b, Mission sci. Mexique, p. 263.
Mexico (PM 4202).

Rance: All records of the species, with the exception of the type, are
from the coastal area of Ecuador and Peru, from Playas south-
ward.

12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Ameiva orcesi J. Peters
Ameiva orcesi J. Peters, 1964, Bull. So. California Acad. Sci., vol. 63,
p. 123. %km. NE of Abdon Calderon, Azuay Prov., Ecuador

(USNM 149655, male).
RANGE: Valley of the Rio Jubones, Azuay Prov., Ecuador.

Ameiva septemlineata A. Duméril

Ameiva septemlineata A. Duméril, 1851, Cat. Méth. Coll. Reptiles, p.
114. ‘‘Amerique meridionale” (PM 4198).

Ameiva sexscutata Giinther, 1859b, Proc. Zool. Soc. London, p. 402.
Andes of Western Ecuador (BM 1946.8.28.35).

Rance: Moister areas of coastal area of Ecuador, from Guayaquil
northward.

Genus AMPHISBAENA Linnaeus

1. A single annulus at midbody contains less than 60 segments; median ventral

Sepinents not longer tam’ broad srs. 9.) ae ee es 9 a 2
A single annulus at midbody contains more than 60 segments; median ventral
Scoments longer’ thanproads ~ cots nee noe eee ee en areal ee alba

2. Abdomen as densely colored as dorsum; 190-205 body annuli; 23-27 tail annuli.
fuliginosa varia

Abdomen usually nearly immaculate, much lighter than dorsum; 204-215
bedy- annulis’’ 26-30) taalanmulie.. 3, 2 ee ie ee fuliginosa basslert

Amphisbaena alba Linnaeus

Amphisbaena alba Linnaeus, 1758, Syst. Nat., 10th ed., vol. 1, p. 229.
‘*America” (RMS).

RancE: Tropical South America. (No good Ecuadorian records are
known.)

Amphisbaena fuliginosa bassleri Vanzolini

Amphisbaena fuliginosa bassleri Vanzolini, 1951, Bull. Mus. Comp.
Zool., vol. 106, no. 1, p. 61. Roaboya, Loreto, Peru (AMNH 56606,
male).

Rance: Upper Amazonian basin of Peru and Ecuador.

Amphisbaena fuliginosa varia Laurenti

Amphisbaena varia Laurenti, 1768, Synopsin Reptilium, p. 66. Barro
Colorado Is., Panama (MCZ 22070, neotype, Vanzolini, 1951,
Bull. Mus. Comp. Zool., vol. 106, no. 1, p. 61).

RANGE: Pacific Coast of Ecuador and Colombia; Panama to Villavi-
cencio, Colombia, in Amazonian South America; east to near
Trinidad in Venezuela.

NO. 3645 LIZARDS—PETERS iS
Genus ANADIA Gray

1. Series of black, blue-centered ocelli on sides; 52-58 scales from occipital to tail;

30-32 scales about body .... net Locate
No black, blue-centered ocelli on dee 48 peace fon Beene to base of tail;
33 scales about body... . . Bids ‘gakloap lect loe.” 53 siaglinin, compere Maint rhombifera

Anadia ocellata Gray

Anadia ocellata Gray, 1845, Cat. Lizards British Mus., p. 74. ‘Tropical
America” (BM 1946.8.2.2, male).
RancE: Known from Jerico, Colombia, and Loja, Ecuador.
Anadia rhombifera (Giinther)

Cercosaura rhombifera Giinther, 1859b, Proc. Zool. Soc. London, p. 405,
pl. 20, fig. A. Western Ecuador (BM 60.6.16.11, female).
RANGE: Pacific slope of Ecuador.

Genus ANOLIS Daudin?

1, End of snout raised, elongated, or with prominent bulge. . . ..... 2
End of snout normal, not swollen or elongated... .. . ae Late ES

2. End of snout produced into an elongate, leaflike structure BEOe as long as
ECAC ACO eeeretse cata te sents Gewese A hs oe eh ay de ee oy BSA es proboscis

End of snout swollen, eae aon Rencreeel a iene distance beyond tip of
TARA ATV a ae i Bo RN Gok 5 Gul yes on to | oh xp a, Spunotatus, boulengers

MV CNIET ALIS AICSISIMOOUM Moc eViyarre comet ce lay” is ois ce ap ah ne oad ee A
Wentral scales keeled 5 5 3s: wy RGA LE alps eh

4. At least two rows of middorsal ents dearly argc an other dorsal and
lateral scales... t05,. Sarah Rn Oe Sate c/s erate Ae

All dorsal scales subequal, Sonate Shes Site geo cy ce gage ick ae emma aoc

3 vemlarsed dorsal scales'smooth, a a) (5s ac se ee ce eo, ee
Enlarged dorsal scales keeled. . . . . . Sle eee eG

6. Enlarged dorsal scales comparatively eae ina oe rows orinch rapidly grade
initombeteralsy ot 6 ch we) Jes pkey tog ce ee ton NR OS eo
Enlarged dorsal scales spate freee fae eee in Sane 12 regular
GOW SOM terse Me) ca eects ie ERMAN Lars Ser 8 cop Jal Sesrel IReleo aa Bae bee Ceara 2h lionotus*

7. 5-6 enlarged papenenare ordered by granules, consisting of one hexagonal
smooth plate surrounded by five similar or smallerones . . .. . bocourti*
11-12 small anne which are only slightly larger than scales on the
MUZZICS ee. 3 Sey oA erie ee Ai Contes ea Nes eM eST CE

8. Head scales enlarged . ates ea rr th om att VS, a a A
iEfead pscalesteranilat.) Soe! se ce a) 3 ae ee ey Sy a PQSENE

Os, Elead scales keeled o-50 7) 5).2) Sieh Uewcn Kiettacy utah. NO ae oe eee a)
Head scales smooth. . . . . ee on aon So, Scored 7, Su ay 1 ge ein Ce GUGLONIGIES.

3 Two new taxa, A. biporcatus parvauritus Williams and A. nigrolineatus Williams, have
been added to the list of species but are not included in the key to the genus Anolis.

14
10.

I

L3i

Ls

ld:

18.

20.

21.

22.

23

2.
2a.
26.

27.

PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

No see einaes larger thanithe seales‘omsnout.4 95 se 2 eo
Some or many supraoculars enlarged, larger than snout selec Se hed Socata
6=8 upperdabials tocenter of eye (to) Giese See 1h cee
10-12 upper labials to center ofeye. . . . - [ha .lQbejrons
White beneath, dotted and vermiculated a black: Mout rows of scales
between supraorbital semicircles. . . . . .. . ventrimaculatus*
Uniform white beneath, two rows of scales benween supracrnieal semicircles.
eulaemus

Supraorbital semicircles separated by one or two rows of scales. . . . . 14

Supraorbital semicircles separated by two to four rows of scales . _fuscoauratus
Interparietal (occipital) larger than ear ee and in contact with the
supraorbital semicircles . . . . . . « macrolepis*
Interparietal (occipital) equal to or mnaller hein the ear opening, and separated
from the supraorbital semicircles by three or four rows of scales . peraccae

Head 'scalesigmooth: +4 2.455.500 ) aes ieee ral tee bens, fe ee ee
Fleadscales not'smooth (2.5). «ies agioes, Eesti
Supraocular scales large, about sevenin number . . . . ete Gay,
Supraocular scales small, about size of scales on rest of hewt very numerous.
chloris

Occipital larger thamiear Opening. 6. cf. ii) oe chine, es ee ee ROOD
Occipital not larger than ear opening. . . . ..... . . . transversalis
lead ‘scales mlcose is S40 Ae en elm fe eae oy ee alee aoe ey
Pieadtseales keeled: Gi! cis cs eos ot, hes yen aioe eagle au eee es ee
Supraocularscalessmooth. 6c sta Bos Gi eee coo ue heen Madaawen
Supraocular scales carinate .. . . . gemmosus
Eight labials to below center of eye; eile emcee nee recone and third
phalanges of fourth toe. . . . . . « maculiventris
10-12 labials to below center dee eye; + 20-22 iamicltae under second and third
phalanges of fourth toe. . . . Sh ies: vicky hy hse baat ay ete ROTI ES
At least two rows of enlarged Banal acute ab wha od do gay plea al Us ee eel age ee
All dorsal scales granular . . . . Snes Pug tee
Legs long, hindlimb reaches end ars snout; eee feebly ied 3. Dal a Seen
Legs short, hindlimb reaches ear opening; digital expansions strongly developed.
Srasert

Dorsum uniform brown or olive; may have few small dark spots, distinct dark
lateral band from eye to side of body .... . . . . . granuliceps

Dorsum with series of large angular spots or ay often confluent into
zigzag band; females may have broad, light, dark-edged vertebral band.

scypheus
Enlarged supraccular scales kecled 5 boys to teh on os ay fe, eS
Enlarged supraocular scales smooth. ............ . . binotatus
Dorsal scales small, rhomboidal, subimbricate. . . . . 3 ike cA
Dorsal scales large, hexagonal, juxtaposed, in fonerecieal SERIES. myc 4p ea
Supraorbital semicircles separated by two rows of scales . . . . tropidogaster
Supraorbital semicircles in contact or a by one row of scales gracilipes
Ventral scales larger than dorsals. . . . . zl ie OU eames ate
Dorsal scales very large, largest twice as eee as penal . . . . notopholis*
Five labials to point below centeriof eyes. 4 4-2 ia 2 ee) amrales
Eight or more labials to belowcenter ofeye ......... . . bitectus

NO. 3545 LIZARDS—PETERS 15

Anolis aequatorialis Werner

Anolis aequatorialis Werner, 1894b, Zool. Anz., vol. 17, p. 157. ‘‘Ecua-
dor” (VM 16233).
Rance: Middle altitudes of western slopes in Ecuador.

Anolis auratus Daudin

Anolis auratus Daudin, 1802, Hist. Nat. Reptiles, vol. 4, p. 89. Un-
known (type also unknown).
RANGE: Northern South America into Central America.

Anolis binotatus W. Peters

Anolis binotatus W. Peters, 1863, Monatsb. Akad. Berlin, p. 140. Gua-
yaquil, Ecuador (BM 4685).
RANGE: Pacific Ecuador and Colombia; southern Central America.

Anolis biporcatus parvauritus Williams

Anolis biporcatus parvauritus Williams, 1966, Breviora, Mus. Comp. Zool.,
no. 239, p. 7, illustr. Gorgona Is., Cauca, Colombia, 5-45 m.
(MCZ 78935).

RANGE: Lowlands west of the Andes in Colombia and Ecuador.

Anolis bitectus Cope

Anolis bitectus Cope, 1864, Proc. Acad. Nat. Sci. Philadelphia, vol. 16,
p. 171. West Ecuador (BM 60.6.16.33).
RANGE: Pacific lowlands of Ecuador.

Anolis chloris Boulenger

Anolis chloris Boulenger, 1898, Proc. Zool. Soc. London, p. 110, pl. 10,
fig. 3. Paramba, Ecuador (BM 98.4.28.10).
RANGE: Pacific lowlands of Ecuador; Darien of Panama.

Anolis eulaemus Boulenger

Anolis eulaemus Boulenger, 1908, Ann. Mag. Nat. Hist., ser. 8, vol. 2,
p. 516, fig. 1. Pavas, Colombia (BM 1909.4.30.61).

RANGE: Southwestern Colombia; also reported from Ecuador on basis
of specimens that differed in several respects from Boulenger’s type
description, by Despax, 1911, Bull. Mus. Nat. Hist. Paris, vol. 17,
Os, Spe, 0:

Anolis festae Peracca

Anolis festae Peracca, 1904, Boll. Mus. Zool. Univ. Torino, vol. 19,
no. 465, p. 4. Balzar, Ecuador (TurM 2872).
RancE: Lowlands of western Ecuador.

16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Anolis fraseri Giinther

Anolis cristatellus —Ginther, 1859a, Proc. Zool. Soc. London, p. 89.

Anolis fraseri Giinther, 1859b, Proc. Zool. Soc. London, p. 407. Andes
of Western Ecuador (BM 1946.8.8.47, lectotype, per Williams, 1966,
p12):

Anolis devillei Boulenger, 1880, Bull. Soc. Zool. France, p. 42. Andes of
Ecuador (IRB 2006).

Rance: Higher western slopes of the Andes in Ecuador and Colombia.

Anolis fuscoauratus fuscoauratus Duméril and Bibron

Anolis fusco-auratus Duméril and Bibron, 1837, Erp. Gén., vol. 4, p. 110.
““Chile,”’ corrected to Rio Mamore, between Loreto and “‘le confluent
du Rio Sara,” Prov. of Moxas, Bolivia, by Bocourt, 1870, Bull.
Nouv. Arch. Mus. Paris, vol. 6, p. 15 (PM 798).

Anolis viridiaeneus W. Peters, 1863, Monatsb. Akad. Berlin, p. 147.
Quito, Ecuador (BerM 3889).

Anolis apollinaris—Burt and Burt, 1930, Proc. U.S. Nat. Mus., vol. 78,
art:,6,0p:,.6-

Rance: Amazonian slopes of Andes from Ecuador to Bolivia.

Anolis gemmosus O’Shaughnessy

Anolis gemmosus O’Shaughnessy, 1875, Ann. Mag. Nat. Hist., ser. 4,
vol. 15, p. 280. Type locality unknown (BM 71.4.16.27).

Anolis squamulatus—Boulenger, 1882, Ann. Mag. Nat. Hist., ser. 5,
vol. 9, p. 458.

Anolis andianus Boulenger, 1885, Cat. Lizards British Mus., vol. 2,
p. 60. Milligalli, Ecuador, 6200 feet (BM 82.7.26.7).

Anolis fasciatus Boulenger, 1885, Cat. Lizards British Mus., vol. 2,
p. 59, pl. 3, fig. 1. Guayaquil, Ecuador (BM 60.6.16.35).

Anolis elegans Boulenger, 1898, Proc. Zool. Soc. London, p. 109, pl. 10,
fig. 2. Chimbo, Ecuador (BM 98.4.28.9).

RANGE: Pacific lowlands of Ecuador.

Anolis gracilipes Boulenger
Anolis gracilipes Boulenger, 1898, Proc. Zool. Soc. London, p. 112, pl.
11, fig. 3. Paramba, Ecuador (BM 98.4.28.22-25).
RANGE: Known only from type locality.

Anolis granuliceps Boulenger

Anolis granuliceps Boulenger, 1898, Proc. Zool. Soc. London, p. 111, pl.
11, fig. 2. Paramba, Ecuador (BM 98.4.28.15-20; TurM 235i)
UMMZ 59002).

Anolis breviceps Boulenger, 1913, Proc. Zool. Soc. London, p. 1031,
pl. 107, fig. 1. Pefia Lisa, Condoto, Colombia (BM 1913.11.12.12-
14).

Rance: Pacific lowlands of Colombia and Ecuador, in Choco region.

|
|

a a

NO. 3545 LIZARDS—PETERS 7

Anolis latifrons Berthold

Anolis latifrons Berthold, 1846, Nachr. Univ. und KGnigl. Gesell. Wiss.
Gdttingen, nos. 8-10, p. 11. Popayan, Colombia (GottM).

Rance: Northwestern Ecuador through the Chocé of Colombia to
southern Central America.

Anolis maculiventris Boulenger
Anolis maculiventris Boulenger, 1898, Proc. Zool. Soc. London, p. 111,
pl. 11, fig. 1. Paramba, Ecuador (BM 98.4.28.11—12).
RANGE: Lowlands of northwestern Ecuador.

Anolis nigrolineatus Williams

Anolis nigrolineatus Williams, 1965, Breviora, Mus. Comp. Zool.,
no. 233, p. 4, illustr. Machala, El Oro Prov., Ecuador (MCZ 38940).

RANGE: Known only from Machala and Guayaquil, in Ecuador.
There is reason to doubt the validity of both localities, according to
Williams, loc. cit.

Anolis ortoni Cope

Anolis ortont Cope, 1868, Proc. Acad. Nat. Sci. Philadelphia, vol. 30,
p. 97. Napo or Upper Marafion, Ecuador-Peru (present location of
type unknown).

Anolis bouviert.—O’Shaughnessy, 1881, Proc. Zool. Soc. London, p. 243.

Rance: Amazonian Basin.

Anolis peraccae Boulenger
Anolis peraccae Boulenger, 1898, Proc. Zool. Soc. London, p. 108, pl.
10, fig. 1. Chimbo, Ecuador and Rio Peripa, Ecuador (BM 98.4.-
28.4-8; TurM 2358).
Anolis irregularis Werner, 1901, Verh. Zool.—Bot. Gesell. Wien, vol.
51, p. 594. Ecuador (BerM 16592).
Rance: Northwestern Ecuador.

Anolis princeps Boulenger
Anolis princeps Boulenger, 1902, Ann. Mag. Nat. Hist., ser. 7, vol. 9,
p. 54. Carondolet (BM 1901.6.27.2), Rfo Lita (BM 1901.3.29.83);
Paramba (BM 1901.3.29.95-96). Specimens from St. Javier and
Salidero are not marked as types in the BM, although these places
were mentioned in the type description.
RANGE: Lowlands of Northwestern Ecuador.

Anolis proboscis J. Peters and Orcés-V.

Anolis proboscis J. Peters and Orcés-V., 1956, Breviora, Mus. Comp.
Zool., no. 62, p. 2, illustr. Cunuco, 5 km. northwest of Mindo,
1200 m., Pichincha Prov., Ecuador (MCZ 54300).

Rance: Northwestern Ecuador.

219-943-673

18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Anolis punctatus boulengeri O’Shaughnessy

Anolis nasicus.—O’ Shaughnessy, 1880, Proc. Zool. Soc. London, p. 491.

Anolis boulengert O’Shaughnessy, 1881, Proc. Zool. Soc. London, p. 242,
pl. 24. Canelos, Ecuador (BM 80.12.8.43).

RaNncE: Amazonian Ecuador.

Anolis scypheus Cope

Anolis scypheus Cope, 1864, Proc. Acad. Nat. Sci. Philadelphia, vol. 16,
p. 172. Caracas, Venezuela, according to Barbour (BM 1946.-
8.8.55, also XXII.5.3.F).

Anolis chrysolepis—O’ Shaughnessy, 1880, Proc. Zool. Soc. London, p.
491.

Rance: Amazonian drainage of Venezuela, Peru, Ecuador.
Anolis transversalis A. Duméril

Anolis transversalis A. Duméril, 1851, Gat. Méth. Coll. Reptiles, p. 57.
South America (actually Sarayacu, Peru, according to E. E. Williams)
(PM).

Anolis buckleyi O’Shaughnessy, 1880, Proc. Zool. Soc. London, p. 492,
pl. 49. Canelos, Ecuador (BM 80.12.8.45-46).

Rance: Amazonian Ecuador and Peru.

Anolis tropidogaster Hallowell

Anolis tropidogaster Hallowell, 1857, Proc. Acad. Nat. Sci. Philadelphia,
vol. 8, 1856 (publ. 1857), p. 224. Colombia (ANSP 7618).

Anolis stigmosus Bocourt, 1869, Bull. Nouv. Arch. Mus. Paris, vol. 5,
p. 43. Magdalena River, Colombia (PM 2427).

Anolis chrysolepis. —Boulenger, 1882, Ann. Mag. Nat. Hist., ser. 5, vol. 9,
p. 458.

Anolis lemniscatus Boulenger, 1898, Proc. Zool. Soc. London, p. 113,
pl. 10, fig. 4. Chimbo, Ecuador (BM 98.4.28.27-31; TurM one
syntype; MCZ 16783).

RancE: Western slopes of Colombia and Ecuador.

Genus ARTHROSAURA Boulenger

Arthrosaura reticulata reticulata (O’Shaughnessy)

Cercosaura (Pantodactylus) reticulata O’Shaughnessy, 1881, Proc. Zool.
Soc. London, p. 230, pl. 22, fig. 1. Canelos, Ecuador (BM 1946.-
9.1.5).

RANGE: Amazonian Ecuador.

Genus BASILISCUS Laurenti

Ll. Mentral’sealées:smoothe 20 soe) css catia ac ee a aera 2
Ventral scales keeled

NO. 3545 LIZARDS—PETERS 19

2. A high dorsal crest and a high caudal crest, both crests with the upper edges
serrate but not scalloped, both covered with thin, somewhat enlarged scales.
basiliscus

No high dorsal crest but a serrate series of compressed trihedral tubercles,
touching or separated by smaller scales; a low, serrate ridge on tail. galeritus

Basiliscus basiliscus (Linnaeus)

Lacerta basiliscus Linnaeus, 1758, Syst. Nat., 10th ed., vol. 1, p. 206.
“America australi” (type unknown).

Rance: Northwestern Ecuador and Colombia; southern Central
America.

Basiliscus galeritus A. Duméril

Basiliscus galeritus A. Duméril, 1851, Cat. Méth. Coll. Reptiles, p. 61.
**N.-Grenade,”’ which is Colombia (PM 2130-31).

Ptenosaura seemani Gray, 1852, Ann. Mag. Nat. Hist., ser. 2, vol. 10,
p. 438. ‘“‘Quibo, on West coast of America’? (BM).

RANGE: Pacific slopes of Colombia and Ecuador to Panama and Costa
Rica.

Basiliscus vittatus Wiegmann

Basiliscus vittatus Wiegmann, 1828, Isis von Oken, vol. 21, p. 373.
Mexico, restricted by Smith and Taylor, 1950, Bull. U.S. Nat.
Mus., no. 199, p. 72, to Veracruz, Veracruz (BerM 549-551).

Rance: Mexico through Central America on both coasts as far as
Colombia. Recorded from Ecuador by Boulenger, 1885, Cat.
Lizards British Mus., vol. 2, p. 109.

Genus CALLOPISTES Gravenhorst

Callopistes flavipunctatus (Duméril and Bibron)

Aporomera flavipunctata Duméril and Bibron, 1839, Erp. Gén., vol. 5,
p. 72. South America (PM 8298, 2 specimens).
RancE: Inter-Andean valleys of Peru and southern Ecuador.

Genus DICRODON Dumeéril and Bibron
Dicrodon guttulatum Duméril and Bibron

Dicrodon guttulatum Duméril and Bibron, 1839, Erp. Gén., vol. 5, p.
138. Peru (PM 1116).

Cnemidophorus lentiginosus Garman, 1892b, Bull. Essex Inst., vol. 24, p.
92. San Francisco de Posorja, Ecuador (MCZ 10775-76).

Ameiva leucostigma Boulenger, 1899a, Proc. Zool. Soc. London, p. 517.
Guayaquil, Ecuador (BM 1946.8.8.69-70).

Rance: Dry coastal Ecuador, north to southern limits of Esmeraldas
Prov. ; coastal Peru.

20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Genus DIPLOGLOSSUS Wiegmann

Diploglossus monotropis (Kuhl)

Scincus monotropis Kuhl, 1820, Beitr. Zool. und Vergl. Anat., p. 128.
“Jamaica,” probably erroneous (type unknown).

Tiliqua jamaicensis Gray, 1839, Ann. Nat. Hist., vol. 2, p. 293. Jamaica
(Mus. Chatham).

RANGE: Pacific coast of Ecuador and presumably Colombia; Central
America to Costa Rica.

Genus ECHINOSAURA Boulenger

Echinosaura horrida horrida Boulenger

Echinosaura horrida Boulenger, 1890, Proc. Zool. Soc. London, p. 82,
pl. 8, fig. 1. Ecuador (BM 1946.8.31.60-61).
RancE: Pacific lowlands of Ecuador.

Genus ECPLEOPUS Duméril and Bibron

Ecpleopus affinis W. Peters

Ecpleopus affinis W. Peters, 1862b, Abh. Akad. Wiss. Berlin, p. 199,
pl. 3, fig. 1. Unknown (MunM).

Cercosaura gaudichaudi— Giinther, 1859a, Proc. Zool. Soc. London, p. 89.

Rance: Higher Pacific slopes and inter-Andean valleys, from Ambato
south, in Ecuador.

Genus ENYALIOIDES Boulenger

1: Dorsal crest distinct §\:; 2 G22 bl A eh ee
Dorsal crest indistinct... . Sntelng ee wabteteniak

2. Spines on nuchal crest not poled fr om spines on domed GESE 52 hanLete, Seats
Spines of nuchal crest prominent and completely isolated from spines on dorsal
ELeSt | ee Uae agen ee Rares to oe ore COLES

3. One or two Pearal pores on Rada nde rath RMN rh eb Reh nde! baa onl Sh
‘Three or four femoral pores onveach side 5 Aeea ete, | 1 epee

4. Ventrals keeled 37> =" 2 = PNT ee Ae. SONA (uc cs ptt Ree
Ventrals smooth or fecisunetly eclede fee . . . praestabilis

5. Dorsal granules very fine, more than sixteen Benseen ae a ceatedianen and
dorsalcrest =) S.-75- . . . macrolepis
Dorsal granules larger, eves ‘han sixteen pestides Piet aepeeainean and
dorsalterest cy, rate toe ae Ue oe Se Oe pS LEDER NT

6... Dorsal scales heteropencous’msize? 0%. ss ee es eee
Dorsal scales homogeneous . . . RAAMSPY, 58

7. Larger scales forming two irregular iene udiial’ series on abs me of back and
irregular vertical series on flanks. .... . ip yet be CO elerolepes

A single series of enlarged scales on each side of daraemn Siena) Meermicrolepes

NO. 3545 LIZARDS—-PETERS al

8. Ventral scales smooth or indistinctly keeled... ....-.- ; laticeps laticeps
Ventral scales usually strongly keeled... --- +--+: > laticeps festae
Enyalioides heterolepis (Bocourt)

Enyalius heterolepis Bocourt, 1874, Ann. Sci. Nat., ser. 5, vol. 19, art. 4,
p. 1. Veragua, Panama (PM 4067).

Enyalioides mocquardi Despax, 1911, Bull. Mus. Nat. Hist., vol. 17, no.
1, p. 10. “Ecuador” (PM 06-226 to 06-228).

Rance: Northwestern Ecuador into Panama.

Enyalioides laticeps laticeps (Guichenot)

Enyalius laticeps Guichenot, 1855, in Castelnau, Exp. Amér. Mérid.,
Reptiles, p. 20. Fonteboa, Upper Amazon, Brazil (PM 6821-22).

Rance: Upper Amazon?

Enyalioides laticeps festae Peracca

Enyalioides festae Peracca, 1897, Bo. Mus. Zool. Univ. Torino, vol.
12, no. 300, p. 3. Valley of the Rio Santiago, Ecuador (TurM 2169,
2 syntypes).

Rance: Amazonian Colombia and Ecuador.

Enyalioides microlepis (O’Shaughnessy)

Enyalius microlepis O’Shaughnessy, 1881, Proc. Zool. Soc. London, p.
938, pl. 24, fig. 2. Sarayacu, Ecuador (BM 1946.8.5.70).

RancE: Pacific lowlands of Ecuador.

Enyalioides oshaughnessyi (Boulenger)

Enyalius oshaughnessyi Boulenger, 1881, Proc. Zool. Soc. London, p.
246, pl. 26. Ecuador (IRB 2009).

Rance: Amazonian Ecuador and Colombia.

Enyalioides praestabilis (O’Shaughnessy)

Enyalius praestabilis O’Shaughnessy, 1881, Proc. Zool. Soc. London, p.
240, pl. 25, fig. 1. Pallatanga and Canelos, Ecuador (BM 1946.-
8.9.15).

Rance: Amazonian Ecuador.

Genus ENYALIUS Wagler

Enyalius zonatus Wettstein
Enyalius zonatus Wettstein, 1926, Anz. Akad. Wiss. Wien, vol: 63, p. I.
“Ecuador” (VM 17188-89).
RANGE: Known only from types.

Genus EUSPONDYLUS Tschudi

1. Largest infraorbital about equal in size to smaller labials. . ... - guenthert
Largest infraorbital much smaller than any Fatale ce ahs: orton: maculatus

22 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119
Euspondylus guentheri (O’Shaughnessy)

Ecpleopus (Euspondylus) guentheri O’Shaughnessy, 1881, Proc. Zool. Soc.

London, p. 235, pl. 23, fig. 1. Sarayacu, Ecuador (BM 1946.8.8.99).

RANGE: Amazonian Ecuador.

Euspondylus maculatus Tschudi
Euspondylus maculatus Tschudi, 1845, Archiv fiir Naturg., vol. 11, p.
161. Vicinity of Moyabamba, Peru (type unknown, not in VM).
Ecpleopus frasert O’Shaughnessy, 1879, Ann. Mag. Nat. Hist., ser. 5,
vol. 4, p. 296. Guayaquil, Ecuador (BM 58.7.25.14).
RANGE: Coastal areas of northern Peru and southern Ecuador.

Genus GONATODES Fitzinger
caudiscutatus
. concinnatus

1. Basal phalanges of digits cylindrical (fig. 6a) .
Basal phalanges of digits slightly but distinctly depressed (fig. 64).

b ¢
Ficure 6.—Basal phalanges of digits: a, Gonatodes caudiscutatus; b, G. concinnatus. Scales
in occipital region: c, Ophryoessoides iridescens; d, O. guenthert.

Gonatodes caudiscutatus caudiscutatus (Giinther)
Gymnodactylus caudiscutatus Giinther, 1859b, Proc. Zool. Soc. London,

p. 410. W. Ecuador (BM 1946.9.7.6-9).
Gonatodes collaris Garman, 1892, Bull. Essex Inst., vol. 24, p. 83.

Wreck Bay, Chatham Is., Galapagos (MCZ 9432).
RANGE: Chocé of Colombia and Ecuador; Galapagos Is.

Gonatodes concinnatus O’Shaughnessy
Goniodactylus ferrugineus.—Cope, 1868, Proc. Acad. Nat. Sci. Philadelphia,

vol. dO; (p.coi.

NO. 3545 LIZARDS—PETERS 23

Goniodactylus concinnatus O’Shaughnessy, 1881, Proc. Zool. Soc. London,
p. 237, pl. 23, fig. 2. Canelos, Ecuador (BM 1946.9.7.10-12).

Goniodactylus buckleyt O’Shaughnessy, 1881, Proc. Zool. Soc. London,
p. 238, pl. 23, fig. 3. Pallatanga and Canelos, Ecuador (BM 1946.-
9.7.13-15).

RANGE: Lower Amazonian slopes of Ecuador.

Genus GYMNOPHTHALMUS Merrem*

Gymnophthalmus speciosus speciosus (Hallowell) *

Blepharactisis speciosus Hallowell, 1861, Proc. Acad. Nat. Sci. Phila-
delphia, 1860 (publ. 1861), p. 484. Nicaragua (type lost).

RancE: Uncertain. Presumably Central America and part of northern
South America, with a questionable record from Chile. Existence
in Ecuador not verified by any specimens known to me.

Genus IGUANA Laurenti

Iguana iguana iguana (Linnaeus)

Lacerta iguana Linnaeus, 1758, Syst. Nat., 10th ed., vol. 1, p. 206.
“Indiis”’ (type unknown).

Iguana tuberculata Laurenti, 1768, Syn. Reptilium, p. 49 (type and
type locality unknown to me).

Rance: Northern South America to Costa Rica; on both Pacific
and Amazonian slopes in Ecuador.

Genus IPHISA Gray

Iphisa elegans Gray

Iphisa elegans Gray, 1851, Proc. Zool. Soc. London, vol. 19, p. 39.
Para, northern Brazil (BM 1946.9.1.1).

RANGE: Known from type locality, which may be erroneous, the
Guianas, and from Santiago-Zamora Prov., Ecuador.

Genus KENTROPYX Spix

ts Dorsal seales, keeled, at leastiin: parts. 4") sew a) sya iie or 8 hag ete 2
iDorsalkscalesksmooth weet oe ee eee aes ee ee altamazonicus

2. Femoral scales which touch the row of femoral pores anteriorly hardly as large
asjthe micdian: oularsines ta Fis? oan ts ioe Ga us tinh ce oda eh Amat calcaratus

Same |scales)larger than) the median gulars)2\)..)..)s)<¢.5)).0 |. fae pelviceps

24 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Kentropyx altamazonicus Cope
Centropyx altamazonicus Cope, 1876, Journ. Acad. Nat. Sci. Philadelphia,
ser. 2, vol. 8, pt. 2, p. 162. Moyobamba, Peru (ANSP 13105).
Rance: Amazonian Ecuador and Peru.

Kentropyx calcaratus Spix
Kentropyx calcaratus Spix, 1825, Spec. Nov. Lacert. Brasil, p. 21, pl. 22,
fig. 2. Itapicurt, in Provincia Maranhao, Brazil (Leipzig?).
RANGE: Northern Amazonian Basin of South America.

Kentropyx pelviceps Cope

Centropyx pelviceps Cope, 1868, Proc. Acad. Nat. Sci. Philadelphia,
vol. 30, p. 98. Napo or Upper Amazon of Ecuador (formerly
USNM 6638, now ANSP 9556).

Centropyx dorsalis——O’Shaughnessy, 1881, Proc. Zool. Soc. London,
ps 226.

RANGE: Amazonian lowlands of Ecuador.

Genus LEPIDOBLEPHARIS Peracca

i. Dorsal-scalesshomogenecous: . i. 2 <tpu cu acne ee iene ee eee 2
Dorsal scales granular with enlarged scales scattered irregularly over back.

ruthvent

2. ‘scales on snout larger than those on'restief- head Means nen el ee 3

No enlarsediscalesvonisnoute! 1G iss ai. Gee teas festae

2: (Noienlarged anterior gulars:,. . afore elton Pinca oe 4

Scales immediately behind mental larger than rest of gulars . . . . buchwaldi

4: Mental: without clefts <2..." hy ae hese ere. ol cieon eee oxycephalus

wo cleits;in rear:part of mental .7\.-0 a + sh co ee eee intermedius

Lepidoblepharis buchwaldi Werner
Lepidoblepharis buchwaldi Werner, 1910, Mitt. Nat. Mus. Hamburg,
vol. 27, p. 8. Hacienda Clementina, Babahoyo, Ecuador (HM 808).

RANGE: Known only from types.

Lepidoblepharis festae Peracca

Lepidoblepharis festae Peracca, 1897, Bol. Mus. Zool. Univ. Torino,
vol. 12, no. 300, p. 2 with fig. San José de Cuchipamba, Ecuador
(TurM 2163).

RANGE: Amazonian slopes of Ecuador and Rio Jurua region of Brazil.

Lepidoblepharis intermedius Boulenger

Lepidoblepharis intermedius Boulenger, 1914, Proc. Zool. Soc. London, p.
814, pl. 1, fig. 2. Pefia Lisa, Condoto, Colombia (BM 1914.5.21.5).
Rance: Chocé of Colombia and Ecuador.

NO. 3545 LIZARDS—PETERS Di

Lepidoblepharis oxycephalus (Werner)

Gonatodes oxycephalus Werner, 1894a, Zool. Anz., vol. 17, p. 413.
Ecuador (type unknown; not in VM).
RancE: Unknown.

Lepidoblepharis ruthveni Parker

Lepidoblepharis ruthvent Parker, 1926, Ann. Mag. Nat. Hist., ser. 9,
vol. 17, p. 295. Chimbo, Ecuador (BM 98.4.28.3, female; TurM
2645).

RancE: Pacific slope of Ecuador.

Genus LEPOSOMA Spix

Leposoma parietale (Cope)

Mionyx parietalis Cope, 1885, Proc. American Philos. Soc., vol. 23,
p. 96. Pebas, Peru (probably ANSP).
RanGcE: Amazonian Colombia, Ecuador, and Peru.

Genus MABUYA Fitzinger

Mabuya mabouya (Lacépéde)

Lacerta mabouya Lacépéde, 1788, Hist. Nat. Quad. Ovip. Serpents,
vol. 1, p. 378, pl. 24. Type and type locality unknown; restricted
to the Antilles by Latreille, 1802 (fide Dunn); further restricted to
the Lesser Antilles by Dunn, 1936, vol. 87, p. 544; still further
restricted to St. Vincent Is. by Smith and Taylor, 1950, p. 156.

Mabuia cepedei—Cope, 1868, Proc. Acad. Nat. Sci. Philadelphia, vol. 30,
p. 100.

Mabuia aenea.—O’ Shaughnessy, 1881, Proc. Zool. Soc. London, p. 236.

Mabuya agilis—Peracca, 1897, Bol. Mus. Zool. Univ. Torino, vol. 12,
no. 300, p. 13; Boulenger, 1887, Cat. Lizards British Mus., vol. 3, p.
190; Parker, 1934, Ann Mag. Nat. Hist., ser. 10, vol. 14, p. 271.

Rance: Low and moderate elevations from central Mexico to Brazil.

Remarks: The proper name for the species of Mabuya from Ecuador
is very much in doubt. I follow Stuart (1963, Misc. Publ. Mus.
Zool. Univ. Mich., no. 122, p. 76), in using this combination, although
Taylor (1956, U. Kans. Sci. Bull., vol. 38, pt. 1, p. 297) said that
‘Sit must be regarded as doubtful whether the species Mabuya mabouya
is represented on the mainland of Central America.’ A thorough
review, both biological and taxonomic, is very much in order for
this species in order to permit accurate definition of subspecies and,
perhaps, species masquerading under this name.

219-943—67—_—4

26 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119
Genus MACROPHOLIDUS Noble

Macropholidus annectens Parker

Macropholidus annectens Parker, 1930, Ann. Mag. Nat. Hist., ser. 10,
vol. 5, p. 569. Loja City, Ecuador (BM 1930.1.30.2, female).
Rance: Upper drainage of Rio Zamora, Ecuador.

Genus MONOPLOCUS Giinther

Monoplocus dorsalis Giinther

Monoplocus dorsalis Giinther, 1859b, Proc. Zool. Soc. London, p. 404.
““Andes of Western Ecuador”’ (originally BM, now lost).

RancE: Uncertain; this has not been taken since the type was col-
lected. The type is no longer extant.

Genus MORUNASAURUS Dunn

Morunasaurus annularis (O’Shaughnessy)

Hoplocercus annularis O’Shaughnessy, 1881, Proc. Zool. Soc. London,
p. 244, pl. 25, fig. 2. Canelos, Ecuador (BM 1946.8.10.35).
RANGE: Amazonian Ecuador.

Genus NEUSTICURUS Duméril and Bibron *

Pronounced. tuberclesion limbs <p c) Ts ch slats nO cin ac et ee Soe 2
Na tubercleson limps. ie hl gol nye: cymes eon eee strangulatus strangulatus

2. Granular scales of sides interspersed with fairly regular rows of enlarged, keeled
SGALES Ty Ber cst Se Rees ae oh GA Pe tee) Ran et nor cen en area nn cae ecpleopus
Granular scales of sides uniform in size; no keeled, enlarged scales. . cochranae

Neusticurus cochranae Burt and Burt

Neusticurus ecpleopus cochranae Burt and Burt, 1931, Bull. American
Mus. Nat. Hist., vol. 61, p. 350. San José de Sumaco, Ecuador
(AMNH 28891, male).

RanGE: Amazonian lowlands of Northern Ecuador.

Neusticurus ecpleopus Cope

Neusticurus ecpleopus Cope, 1876, Journ. Acad. Nat. Sci. Philadelphia,
ser. 2, vol. 8, pt. 2, p. 161. Peru (probably ANSP).

Neusticurus tuberculatus Shreve, 1935, Occ. Pap. Boston Soc. Nat. Hist.,
vol. 8, p. 209. Sarayacu, Ecuador (MCZ 37711, male).

Custa bicarinata—Giinther, 1859b, Proc. Zool. Soc. London, p. 404.

Rance: Amazonian lowland in Ecuador and northern Peru.

4 Arrangement following Uzzell, Bull. American Mus. Nat. Hist., in press.

NO. 3545 LIZARDS—PETERS 24

Neusticurus strangulatus strangulatus (Cope)

Euspondylus strangulatus Cope, 1868, Proc. Acad. Nat. Sci. Philadel-
phia, vol. 30, p. 99. Ecuador (ANSP 7538).

Euspondilus festae Peracca, 1897, Boll. Mus. Zool. Univ. Torino, vol. 12,
no. 300, p. 10. Valleys of the Rio Zamora (TurM 2157) and the
Rio Santiago (TurM 2156), Ecuador.

RancE: Eastern slopes of the Ecuadorian Andes, and a dubious
record from Pallatanga, on the western slopes of Ecuador.

Genus OPHIOGNOMON Cope

1. Hindlimb half as long as pre-anal plates; 20 scales around the body . ¢risanale
Hindlimb reduced to tiny stub; 26-28 scales around the body . . . abendrothii

Ophiognomon abendrothii (W. Peters)

Chalcides (Hapalolepis) Abendrothii WW. Peters, 1871, Monatsb. Akad.
Berlin, p. 399. Sarayacu, Peru (BerM 7132-7134 and BM 73.-
4.30.9).

RancE: Amazonian slopes in Ecuador and Peru.
Ophiognomon trisanale Cope

Ophiognomon trisanale Cope, 1868, Proc. Acad. Nat. Sci. Philadelphia,
vol. 30, p. 100. ‘‘Napo or Upper Marajion, Ecuador” (formerly
USNM 6637, now ANSP 9637).

RancE: Amazonian Ecuador.

Genus OPHRYOESSOIDES Duméril

1. Scales in occipital region large (fig. 6¢,p. 22)... ... Resi? Ait ee 24
Scales in occipital region small (fig.6d) ........ potd Aa hee on eS

2. Head scales quite rugose; pair of dorsolateral crests present. . . . . aculeatus
Head scales not rugose; no dorsolateral crests. . . . . ... = . . indescens

Ae SUPLAUCHI ALS ECE AISVEISClyiGU ALC fy is og tin ge eel eh ek eee ou
iINollarve: dilated: supraoculars” 050 Nth aT NAONS UN aih tee al a

A. Four broadly dilated’supraoculars: . 9.7) 2 eS. formosus
Sim broadly dilated/supraceulars 2. a ee 2 Se haensehe

5: Weessithan\ 55) scales around: body? <0) ) 2 ishkerso eo) Baan eae ere
More than 56 scales around body. ......... . . . guenthert

6. Sides of belly black; small postfemoral and postaxillary eel pouches; 272) 7
Sides of belly red or pink; large postaxillary and postfemoral dermal pouches.
rhodomelas

J: (47 Gr more,scalessaroundyubogy, Ven eM) Pe) PR WO festae

tessithan: 47iscalestaround body?) iy) Asustek borle SW es . . ornatus

28 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Ophryoessoides aculeatus (O’Shaughnessy)
Leiocephalus aculeatus O’Shaughnessy, 1879, Ann. Mag. Nat. Hist.,
ser. 5, vol. 4, p. 303. Moyobamba, Peru (BM 1946.8.12.33-36).
Letocephalus angulifer Werner, 1901, Verh. Zool.-Bot. Gesell. Wien,
vol. 51, p. 595. Ecuador (BerM 16594).
Rance: Amazonian slopes of Ecuador and Peru.

Ophryoessoides festae (Peracca)
Lewocephalus festae Peracca, 1897, Bol. Mus. Zool. Univ. Torino, vol. 12,

no. 300, p. 6. Cuenca, Ecuador (TurM 2619).
RANGE: Inter-Andean plateau in Cuenca Hoya, Ecuador.

Ophryoessoides formosus (Boulenger)
Liocephalus formosus Boulenger, 1880, Bull. Soc. Zool. France, p. 43.

Andes of Ecuador (IRB 2007).
RANGE: Known only from the type.

Ophryoessoides guentheri (Boulenger)

Liocephalus ornatus.—Ginther, 1859b, Proc. Zool. Soc. London, p. 408.

Liocephalus trachycephalus.—Boulenger, 1882, Ann. Mag. Nat. Hist., ser. 5,
vol. 9, p. 458.

Liocephalus guenthert Boulenger, 1885, Cat. Lizards British Mus., vol. 2,
p. 169, pl. 13. Guayaquil and Sarayacu, Ecuador and a questioned
report from Colombia (BM. 58.7.25.16-18; 59.9.26.6; 60.6.16.18—-21;
Ale 27 -LOs 7 46532 G051228-93)).

RANGE: Apparently confined to the Inter-Andean plateau in Ecuador.

Ophryoessoides haenschi (Werner)

Liocephalus haenschi Werner, 1901, Verh. Zool.—Bot. Gesell. Wien,
vol. 51, p. 595. Balzapamba, Ecuador (BerM 16595).
RANGE: Known only from the type.
Ophryoessoides iridescens (Giinther)
Liocephalus iridescens Giinther, 1859b, Proc. Zool. Soc. London, p. 409,
pl. 20, fig. B. Andes of Ecuador (BM 60.6.12.2—7).
RANGE: From Guayaquil to El Oro Prov., in drier coastal areas.
Ophryoessoides ornatus ornatus (Gray)
Lewocephalus ornatus Gray, 1845, Cat. Lizards British Mus., p. 219.
Guayaquil, Ecuador (BM 1946.8.29.72).
Rance: West Coast of Ecuador.
Ophryoessoides rhodomelas (Boulenger)
Liocephalus rhodomelas Boulenger, 1899b, Ann. Mag. Nat. Hist., ser. 7,
vol. 4, p. 455. Ofia, Ecuador (BM 1946.8.29.77—80).
RancE: Inter-Andean plateaus of southern Ecuador.

NO. 3545 LIZARDS—PETERS 29
Genus PHOLIDOBOLUS Peters

Pholidobolus montium W. Peters

Ecpleopus (Pholidobolus) montium W. Peters, 1862b, Abh. Akad. Wiss.
Berlin, p. 196, pl. 2, fig. 3. Quito, Ecuador (BerM 900, LeyM
3401).

Cercosaura gaudichaudi.—Boulenger, 1882, Ann. Mag. Nat. Hist., ser.
DyFvolt9:M9..459:

RANGE: Inter-Andean Ecuador.

Genus PHYLLODACTYLUS Gray

Phyllodactylus reissii W. Peters

Phyllodactylus reissii W. Peters, 1862a, Monatsb. Berlin Akad., p. 626.
Guayaquil, Ecuador (BerM 3734).

Phyllodactylus guayaquilensis Werner, 1910, Mitt. Nat. Mus. Hamburg,
vol. 27, p. 4. Guayaquil, Ecuador (HM 989).

Phyllodactylus abruptisertiatus Werner, 1913, Mitt. Nat. Mus. Hamburg,
vol. 30, p. 4. ‘‘wahrscheinlich Brasilien’? (formerly HM, now
destroyed).

RANGE: Coastal Ecuador up to 1250 meters, perhaps also coastal Peru.

REMARKS: These three species are regarded as synonymous by James
R. Dixon (1962, personal communication), and his scheme is
being followed in anticipation of publication of his results. P.
tuberculosus is not Ecuadorian, and references to it in the literature
are probably this species.

Genus PLICA Gray

1. Side of neck with tufts of small, erect, spinelike scales
Side of neck without spines

Plica plica (Linnaeus)
Lacerta plica Linnaeus, 1758, Syst. Nat., 10th ed., vol. 1, p. 208.
‘Indiis” (type not located).
Hypsibates agamoides—Cope, 1868, Proc. Acad. Nat. Sci. Philadelphia,
volw3O) p89 7
Rance: Northern South America to Bolivia, east of the Andes.

Plica umbra (Linnaeus)

Lacerta umbra Linnaeus, 1758, Syst. Nat., 10th ed., vol. 1, p. 207.
**Meridionalibus” (type unknown).
Rance: Northern South America, east of the Andes.

30 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119
Genus POLYCHRUS Cuvier

1. Scales of sides same size or smaller than dorsals and not separated by tiny

pranmulés.iis. 3. Jpsse weds. Veeteye ic uae Soe ERE Cae Ae ee 2
Scales of sides slightly larger than dorsals and separated from each other by
tiny, granular scales. 176) we eek Perea wh, fy eater on ieee Nee Femoralts

2. Canthus rostralis at least somewhat rounded; scales on pectoral region smooth
or very Weakly keeled (sg van, peones UN hs, “Sante ena eee 3
Canthus rostralis distinctly angular; scales on pectoral region strongly keeled,
may be bi- or tricarinate, usually unicarinate. .... . gutturosus gutturosus

3. Low series of raised scales forming midventral crest from mental to gular
APDENGARE 2 ia mal pola mise denen al ee ere code rence marmoratus

No low series of raised scales enme aarcaitel crest on chin.
gutturosus spurrellit

Polychrus femoralis Werner

Polychrus femoralis Werner, 1910, Mitt. Nat. Mus. Hamburg, vol. 27,
p. 21. Guayaquil, Ecuador (formerly HM, apparently destroyed).
RANGE: Lowlands in southwestern Ecuador.

Polychrus gutturosus gutturosus Berthold

Polychrus gutturosus Berthold, 1846, Nachr. Univ. und Kénigl. Gesell
Wiss. Gottingen, nos. 8-10, p. 11. Popayan, Colombia (GottM).
Rance: Higher western Andean slopes of Ecuador and Colombia and
northward to Costa Rica and Nicaragua.
Polychrus gutturosus spurrellii Boulenger

Polychrus spurrellit Boulenger, 1914, Proc. Zool. Soc. London, p. 814.
Pefia Lisa, Condoto, Colombia (BM 1946.8.8.33-34).
Rance: Chocé of northwestern Ecuador and Colombia.
Polychrus marmoratus (Linnaeus)

Lacerta marmorata Linnaeus, 1758, Syst. Nat., 10th ed., vol. 1, p. 208.
“Hispania” (type unknown).
Rance: Amazonian basin of South America.

Genus PRIONODACTYLUS O’Shaughnessy

1. Fewer than 25 transverse rows of ventral plates from the edge of the collar to

pte-amalshivelclsta 5 4 PEE YEG ee PSE oe PGBS Aen ER 9 eee 2

25 or more transverse ventral rows . . . Bape & Pete geaiet Wey PeeOCCEnt nen

2. 35 or more scales from occiput to base of a Jee dolce hte et et ees Re ee 3
Fewer than 35 scales from occiput to base of tail. . . . . . . . . vertebralis

3. Fewer than 40 scales about middle of body ....... . . oshaughnessyt

40 or more’scales about middle of body) | 4./...) GA. :snaee pees manicatus

NO. 3545 LIZARDS—PETERS ub

Prionodactylus manicatus (O’Shaughnessy)

Cercosaura (Prionodactylus) manicata O’Shaughnessy, 1881, Proc. Zool.
Soc. London, p. 231, pl. 22, fig. 3. Canelos and Pallatanga, Ecuador
(BM 1946.8.2.1).

Rance: Amazonian Ecuador.

Prionodactylus ocellifer Werner

Prionodactylus ocellifer Werner, 1901, Verh. Zool.-Bot. Gesell. Wien,
vol. 51, p. 596. Ecuador (BerM 16593).
RANGE: Known only from type, which lacks specific data.

Prionodactylus oshaughnessyi Boulenger

Cercosaura (Pantodactylus) argulus——O’Shaughnessy, 1881, Proc. Zool.
Soc. London, p. 229.

Prionodactylus oshaughnessyi Boulenger, 1885, Cat. Lizards British Mus.,
vol. 2, p. 392, pl. 21, fig. 1. Canelos and Pallatanga, Ecuador
(BM 1946.8.31.18-20).

Rance: Amazonian Ecuador.

Prionodactylus vertebralis (O’Shaughnessy)

Cercosaura (Pantodactylus) vertebralis O’Shaughnessy, 1879, Ann. Mag.
Nat. Hist., ser. 5, vol. 4, p. 298. Intac, Ecuador (BM 1946.8.31.35).

Rance: Higher Pacific slopes of Ecuador; also reported from Zamora,
Ecuador, by Parker, 1934, Ann. Mag. Nat. Hist., ser. 10, vol. 14,
p. 270. Darien, Panama.

Genus PROCTOPORUS Tschudi

1. Pores present in pre-anal area; at least one supraocular in contact with upper

palpebrals; adpressed limbs do not touch. . . . . . SP yells NE

No pores in pre-anal area; superciliary series usually complet except in
columbianus: adpressea: limbs usually meet: © EE i ee 5

2. Supraoculars four; two, three, and four usually in contact ae upper palpebrals;
no sexual dimorphism in femoral porenumber.......... meleagris
Supraoculars three; usually only two in contact with palpebrals; sexual di-
morphismiin femoral pore number .).5..) ei) + + + oe entcolor

3. Fewer than 50 scales from occiput to base of ea Ly GND eee OUP Ai Be a
More than 50 scales from occiput to base of tail; disc in lower eyelid not divided

by vertical grooves into two or three sections. . . . . “iy Js ow Sah ACIEIENIESS

4. Superciliary series incomplete, second or second and hid aipraceulers in
contact with upper palpebrals; adpressed limbs separated or barely touching.
columbianus

Superciliary series usually complete (occasionally not in striatus); limbs usually
broadly overlapping when adpressed. ..-) £0. © J). sh Beg os eS

oo PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

5: Dorsal scales'smooth. >.) Ate Re oe ee ees ee stmoterus
Dorsal scales striatedion keeled Foe els ate cun tn ue eA eed EN 6

6. Males with ocelli in pattern; a dorsolateral light stripe present; dorsal scales
keeled sta pi ae wh oe Pay es wie toni Rin, cee ait oe Oe ene ca oculatus
Males without ocelli in pattern; dorsals keeled or striate ........ 7

7. Dorsal pattern of longitudinal dark brown stripes on light brown, or uniform
Light brown; ‘scales striate’... 2h)... Dee ed Senter aly. aaah ant ane striatus
Dorsal pattern not linear, consisting of small black spots on a dark brown
background; dorsals feebly but distinctly keeled... . . .... . hypostictus

Proctoporus columbianus Andersson
Proctoporus columbianus Andersson, 1914, Arkiv fiir Zool., vol. 9, no. 3,
p. 3, fig. 1. Colombia (RMS).

Proctoporus oculatus (part).—Burt and Burt, 1931, Bull. American Mus.
Nat. Hist., vol. 61, p. 369.

RANGE: Eastern slopes of Andes in Colombia and Ecuador.

Remarks: The record from Abitagua, by Burt and Burt (ibid.), probably
belongs to an undescribed subspecies of this species, according to
Uzzell, 1958, Occ. Pap. Mus. Zool. U. Mich., no. 597, p. 7.

Proctoporus hypostictus Boulenger

Proctoporus hypostictus Boulenger, 1902, Ann. Mag. Nat. Hist., ser. 7,
vol. 9, p. 55. Paramba, Ecuador (BM 1901.3.29.105).

Rance: Higher western slopes of Andes in Ecuador.

Proctoporus meleagris Boulenger
Proctoporus meleagris Boulenger, 1885, Cat. Lizards British Mus., vol. 2,
p. 415, pl. 22, fig. 2. Western Ecuador (BM 60.6.16.18).

RancE: Higher western slopes of Andes in Ecuador, up to 3000
meters.

Proctoporus oculatus (O’Shaughnessy)

Ecpleopus oculatus O’Shaughnessy, 1879, Ann. Mag. Nat. Hist., ser. 5,
vol. 4, p. 297. Intac, Ecuador (BM 78.1.25.5).
Rance: Higher western slopes of Andes in Ecuador.

Proctoporus simoterus (O’Shaughnessy)

Emphrassotis simoterus O’Shaughnessy, 1879, Ann. Mag. Nat. Hist.,
ser. 5, vol. 4, p. 296. Intac, Ecuador (BM 1946.9.1.6).
RaNcE: Western slopes of Andes in Ecuador.

Proctoporus striatus (W. Peters)
Ecpleopus (Oreosaurus) striatus W. Peters, 1862b, Abh. Akad. Wiss.
Berlin, p. 201. Santa Fe de Bogoté, Colombia (BerM).

Proctoporus oculatus (part).—Burt and Burt, 1931, Bull. American Mus.
Nat. Hist., vol. 61, p. 369.

Rance: Upper eastern slopes of the Andes in Colombia; El Chiral,
El Oro Prov., Ecuador(?).

NO. 3545 LIZARDS—PETERS ao

Remarks: Uzzell, 1958, Occ. Pap. Mus. Zool. U. Mich., no. 597,
p. 7, indicates that the specimen Burt and Burt listed as Proctoporus
oculatus from El Chiral may belong to P. striatus Peters, but that it
is not clearly so, showing several distinct differences. The record
certainly makes no zoogeographic sense at all, and it is likely that
the El Chiral specimen represents a new species.

Proctoporus unicolor (Gray)

Riama unicolor Gray, 1858, Proc. Zool. Soc. London, p. 446, pl. 15,
fig. 2. Western Ecuador (BM 53.7.25.44).

Proctoporus pachyurus—Ginther, 1859a, Proc. Zool. Soc. London, p. 89,
and 1859b, ibid., p. 407.

Proctoporus lividus Thominot, 1889, Bull. Soc. Philom. Paris, ser. 8,
vol. 1, p. 25. Ecuador (PM 5812, 2 specimens).

Rance: Western slopes and inter-Andean valleys of northern Ecuador.

Genus PROCTOTRETUS Duméril and Bibron

Proctotretus ornatissimus (Girard)

Saccodeira ornatissima Girard, 1858, Proc. Acad. Nat. Sci. Philadelphia,
1857 (1858), p. 198. Obrajillo and Yanga, Peru (USNM 5655).
RancE: Interior highlands of Ecuador and Peru; southern Ecuado-

rian hoyas only.

Genus PTYCHOGLOSSUS Boulenger

1. Eight rows of ventral plates; three pairs of chinshields, two in contact on midline;

frontal not longer than broad; four supraoculars . ..... . brevifrontalis
Ten rows of ventral plates; four pairs of chinshields, one pair in contact on
midline; frontal longer than broad; three supraoculars.. . ... . picticeps

Ptychoglossus brevifrontalis Boulenger

Ptychoglossus brevifrontalis Boulenger, 1912, Ann. Mag. Nat. Hist., ser. 8,
vol. 10, p. 421. El Topo, Rio Pastaza, Ecuador (BM 1946.8.31.63).

Rance: Amazonian slopes of Ecuador.

Ptychoglossus picticeps (Cope)

Leposoma picticeps Cope, 1885, Proc. American Philos. Soc., vol. 23,
p. 99. Pebas, Peru (type not located).

Ptychoglossus bilineatus Boulenger, 1890, Proc. Zool. Soc. London, p. 84,
pl. 10, fig. 2. Ecuador (BM 1946.8.2.38).

Rance: Amazonian Ecuador and Peru.

34 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119
Genus SPHAERODACTYLUS Wagler

1. No median row of enlarged subcaudals; no black lines on dorsum of head; a

darkband. across the scapularwerion. 4 —oiu.0b, . aa ile eae scapularis
A median row of enlarged subcaudals; five to seven dim longitudinal dark lines
on head; no dark band across the scapular region. . . . . . . . lineolatus

Sphaerodactylus lineolatus Lichtenstein

Sphaerodactylus lineolatus Lichtenstein, 1856, Nomencl. Mus. Zool. Berlin,
p. 6. Veragoa, Panama, which is Veragua (BerM 417).

RancE: Central America from British Honduras south to Panama;
Colombia; reported in Ecuador on the basis of USNM 65451, from
Macas.

Sphaerodactylus scapularis Boulenger

Sphaerodactylus scapularis Boulenger, 1902, Ann. Mag. Nat. Hist., ser. 7,
vol. 9, p. 54. St. Javier, Ecuador (BM 1946.8.30.70).
Rance: Northwestern Ecuador.

Genus STENOCERCUS Duméril and Bibron

1: (G@audal seales without spies icon 2O/ whee Ue! 2 psig ey see I Sagte Shame men
@audal scales strongly Spmose, + iy. %) egaus yell) say phere ee J et ame

2. Vertebral scales raised and pointed, forming a denticulate ridge. . . .. . 3
Nowaised and pomted vertebraliscales.. 3-02 42-22 = eve Pee he OGRE:

3. Ventral scales approximately same size as largest dorsals. . . . ..... 4
Ventral scales considerably larger than largest dorsals. . . . . . . Aumeralis

4. Dorsal scales mucronate; caudal scales mucronate. ..... . nigromaculatus
Dorsal scales not mucronate; caudal scales not or but very shortly mucronate.
boettgert

5. Caudal whorls subequal... . . BESS Tsay lt. Be .. Simonsit
Caudal whorls alternately larger and eae. pees Sa el ate ee carriont

Stenocercus boettgeri Boulenger

Stenocercus boettgeri Boulenger, 1911, Ann. Mag. Nat. Hist., ser. 8, vol. 7,
p. 22. Huancabamba, Peru (BM 1946.8.11.92-99).
Rance: Highland areas of Peru and Ecuador.

Stenocercus carrioni Parker
Stenocercus carrioni Parker, 1934, Ann. Mag. Nat. Hist., ser. 10, vol. 14,
p. 264. Zamora, Ecuador (BM 1933.6.24.75, male).
Rance: Known only from type locality.

Stenocercus humeralis (Giinther)

Microphractus humeralis Giinther, 1859a, Proc. Zool. Soc. London, p. 90.
Andes of Western Ecuador (BM 1946.8.11.76-77).

NO. 3545 LIZARDS—PETERS 35

RANGE: Interandean Plateau of Ecuador, from Cuenca Valley south-
ward.

Remarks: Boulenger, 1885, Cat. Lizards British Mus., vol. 2, p. 134,
notes two specimens from Guayaquil and indicates that they are the
types. However, all records of the species have come from moun-
tainous areas.

Stenocercus nigromaculatus Noble
Stenocercus nigromaculatus Noble, 1924, Occ. Pap. Boston Soc. Nat.
Hist., vol. 5, p. 112. Huancabamba, Peru (MCZ 17975).
RANGE: Highlands of Northern Peru.

Stenocercus simonsii Boulenger

Stenocercus simonsii Boulenger, 1899b, Ann. Mag. Nat. Hist., ser. 7,
vol. 4, p. 454. Ona, Ecuador (BM 1946.8.11.73-74).
RancGE: Highlands of Ecuador.

Stenocercus varius Boulenger
Stenocercus varius Boulenger, 1885, Cat. Lizards British Mus., vol. 2,
p. 134, pl. 8, fig. 3. Unknown (BM 71.4.16.53).
RancE: Highlands of Ecuador.

Genus THECADACTYLUS Oken

Thecadactylus rapicaudus (Houttuyn)

Gekko rapicaudus Houttuyn, 1782, Verh. Zeeuw. Genootsch. Wet.
Vlissingen, vol. 9, p. 323, pl. 3, fig. 1. “American Islands,” restricted
to Chichen Itza, Yucatan, Mexico, by Smith and Taylor, 1950,
U.S. Nat. Mus. Bull., no. 199, p. 49 (type unknown).

Rance: Lesser Antilles, Mexico, Central America, northwestern
South America. Known from lowlands on both sides of the Andes
in Ecuador.

Genus TROPIDURUS Wied

eat leastsa few, dorsal, scales Keele <a ious verges ars os Nocgy ome eR 2
Worsal scalesjcmOotay tps yct ue wee S) bts Nee nek or) ide See peruvianus
Pe EntralscalessmMootie te tees oo ty cl cee CRT Vice. Deck at ty gach oer. eats 3
Woentraliscalesikeeledin, Sehtre nt ce (rene sh ie een ree PILLS Lakh AMS holotropis
ge 2~ dorsalydenticulation or ierest,4).-) SPO UI se occipitalis occipitalts
No dorsalidenticulationsorierest’ (Sind) eR) eo. ei ae torquatus*

Tropidurus holotropis Boulenger

Tropidurus holotropis Boulenger, 1912, Ann. Mag. Nat. Hist., ser. 8,
vol. 10, p. 420. Alpayacu, Rfo Pastaza, Ecuador (BM 1946.8.29.64).
RaNncGE: Amazonian slopes in Ecuador and Peru.

36 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Tropidurus occipitalis occipitalis W. Peters

Tropidurus (Laemopristus) occipitalis WW. Peters, 1871b, Monatsb. Akad.
Wiss. Berlin, p. 645. Peru (BerM 6446).

Tropidurus continentalis Miller, 1924, Mitt. Zool. Mus. Berlin, vol. 11,
no. 1, p. 82. Machalilla, Ecuador (BerM 26397).

RANGE: Coastal areas of southwestern Ecuador and _ northwestern
Peru.

Tropidurus peruvianus peruvianus (Lesson)

Stellio peruvianus Lesson, 1826, in Duperrey, Voy. “Coquille,” Reptiles,
vol. 5, pl. 2, fig. 2; vol. 2, pt. 1, 1830, p. 40. Callao and Payta,
Peru (PM 6873).

RANGE: Coastal southwestern Ecuador and northwestern Peru.

Genus TUPINAMBIS Daudin

1. About 30 ventral scales across middle of belly; oneloreal . . . . nigropunctatus
36 to 40 ventral scales across middle of belly; two loreals. . . . . . teguixin*

Tupinambis nigropunctatus Spix

Tupinambis nigropunctatus Spix, 1825, Spec. Nov. Lacert. Brasiliam,
p. 18, pl. 20. Brazil (Leipzig?).

RANGE: Amazon Basin.

Remarks: Tupinambis teguizin (Linnaeus) was recorded from “‘Napo or
the Upper Marafion” by Cope, 1868, Proc. Acad. Nat. Sci. Phila-
delphia, vol. 30, p. 99. It is very likely that the species is Ecuadorian,
but I know of no other records.

Genus URACENTRON Kaup

Uracentron flaviceps (Guichenot)

Doryphorus flaviceps Guichenot, 1855, in Castelnau, Exp. Amér. Mérid.,
Reptiles, p. 26, pl. 3, fig. 2. Sarayacu, Peru (PM 6882).

Rance: Upper Amazonian areas of Ecuador and Peru.

Remarks: Guibé, 1954, Cat. Types Lézards Paris Mus., p. 41, indicates
that flaviceps is a synonym of Uracentron azureum (Linnaeus). I have
seen no documentation of this synonymy and retain flaviceps until
such documentation is published. Uracentron castor (Cope), 1870, was
described from Pebas, ‘‘Ecuador,” but this locality is actually in
Peru. No Ecuadorian specimens are known to me, but the species
is to be expected in Ecuador, if it is validly distinct from /laviceps.

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40 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

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NO. 3545 LIZARDS—PETERS 41

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Wiss. Wien, vol. 63, pp. 1-3.

NO. 3545 LIZARDS—PETERS 43

WIEGMANN, A. F.
1828. Beitrage zur Amphibienkunde. Isis von Oken, vol. 21, pp. 364-383.
1835. Beitrage sur Zoologie, gesammelt auf einer reise um die Erde, con
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WILuiAMs, ERNEsT E.
1965. South American Anolis (Sauria, Iguanidae): Two new species of the
punctatus group. Breviora, Mus. Comp. Zool., no. 233, pp. 1-15, 3 figs.
1966. South American Anoles: Anolis biporcatus and Anolis fraseri (Sauria,

Iguanidae) compared. Breviora, Mus.Comp. Zool., no. 239, pp. 1-14,
5 figs.

abendrothii, Chalcides (Hapalolepis), 27 | Anolis, 3, 13

Ophiognomon, 27
Ablepharus, 3, 10

boutonii poecilopleurus, 10

poecilopleurus, 10
abruptiseriatus, Phyllodactylus, 29
aculeatus, Leiocephalus, 28

Ophryoessoides, 28
aenea, Mabuia, 25
aequatorialis, Anolis, 15
affinis, Eepleopus, 20
agamoides, Hypsibates, 29
agilis, Mabuya, 25
alba, Amphisbaena, 12
Alopoglossus, 3, 10

buckleyi, 10

carinicaudatus, 10

copii, 10

festae, 11
altamazonicus, Centropyx, 24

Kentropyx, 24
Ameiva, 3, 11

ameiva petersi, 11

bifrontata divisa, 11

bridgesii, 11

edracantha, 11

leucostigma, 19

orcesi, 12

petersi, 11

septemlineata, 12

sexscutata, 12
Amphisbaena, 3, 12

alba, 12

fuliginosa bassleri, 12

fuliginosa varia, 12

varia, 12
Amphisbaenidae, 3
Anadia, 3, 13

ocellata, 13

rhombifera, 13
andianus, Anolis, 16
Anguidae, 3
angulifer, Leiocephalus, 28
annectens, Macropholidus, 26
annularis, Hoplocercus, 26

Morunasaurus, 26

44

aequatorialis, 15
andianus, 16
apollinaris, 16
auratus, 15
binotatus, 15
biporecatus parvauritus, 15
bitectus, 15
bocourti, 13
boulengeri, 18
bouvieri, 17
breviceps, 16
buckleyi, 18
chloris, 15
chrysolepis, 18
cristatellus, 16
devillei, 16
elegans, 16
eulaemus, 15
fasciatus, 16
festae, 15
fraseri, 16
fusco-auratus, 16
fuscoauratus fuscoauratus, 16
gemmosus, 16
gracilipes, 16
granuliceps, 16
irregularis, 17
latifrons, 17
lemniscatus, 18
lionotus, 13
macrolepis, 14
maculiventris, 17
nasicus, 18
nigrolineatus, 17
notopholis, 14
ortoni, 17
peraccae, 17
princeps, 17
proboscis, 17
pulchellus, 4
punctatus boulengeri, 18
scypheus, 18
squamulatus, 16
stigmosus, 18
transversalis, 18

NO. 3545

Anolis—Continued
tropidogaster, 18
ventrimaculatus, 14
viridiaeneus, 16

apollinaris, Anolis, 16

Aporomera flavipunctata, 19

argulus, Cercosaura (Pantodactylus), 31

Arthrosaura, 3, 18
reticulata reticulata, 18

auratus, Anolis, 15

azureum, Uracentron, 36

Bachia, 9
Basiliscus, 3, 18
basiliscus, 19
galeritus, 19
vittatus, 19
basiliscus, Basiliscus, 19
Lacerta, 19
bassleri, Amphisbaena fuliginosa, 12
bicarinata, Custa, 26
bilineatus, Ptychoglossus, 33
binotatus, Anolis, 15
bitectus, Anolis, 15
Blepharactisis speciosus, 23
bocourti, Anolis, 13
boettgeri, Stenocercus, 34
boulengeri, Anolis, 18
Anolis punctatus, 18
bouvieri, Anolis, 17
breviceps, Anolis, 16
brevifrontalis, Ptychoglossus, 33
bridgesii, Ameiva, 11
Holecosus, 11
buchwaldi, Lepidoblepharis, 24
buckleyi, Alopoglossus, 10
Anolis, 18
Goniodactylus, 23
Leposoma, 10

calearatus, Kentropyx, 24
Calliscincopus, 8
Callopistes, 3, 19
flavipunctatus, 19
carinicaudatum, Leposoma, 10
carinicaudatus, Alopoglossus, 10
carrioni, Stenocercus, 34
castor, Uracentron, 36
caudiscutatus, Gonatodes caudiscuta-
tus, 22
Gymnodactylus, 22
Centropyx altamazonicus, 24
dorsalis, 24
pelviceps, 24

LIZARDS—-PETERS

45

cepedei, Mabuia, 25

Cercosaura, 9
gaudichaudi, 20, 29
(Pantodactylus) argulus, 31

reticulata, 18
vertebralis, 31

(Prionodactylus) manicata, 31
rhombifera, 13

Chalcides (Hapalolepis) Abendrothii, 27

chloris, Anolis, 15

chrysolepis, Anolis, 18

Cnemidophorus lentiginosus, 19

cochranae, Neusticurus, 26
Neusticurus eepleopus, 26

collaris, Gonatodes, 22

columbianus, Proctoporus, 32

concinnatus, Goniodactylus, 23
Gonatodes, 22

continentalis, Tropidurus, 36

copii, Alopoglossus, 10

Corythophanes, 6

eristatellus, Anolis, 16

Crocodilurus, 7

Custa bicarinata, 26

devillei, Anolis, 16
Dicrodon, 3, 19

guttulatum, 19
Diploglossinae, 3
Diploglossus, 3, 20

monotropis, 20
divisa, Ameiva bifrontata, 11
dorsalis, Centropyx, 24

Monoplocus, 26
Doryphorus flaviceps, 36
Dracaena, 7

Echinosaura, 3, 20
horrida, 20
horrida horrida, 20
Ecpleopus, 3, 20
affinis, 20
(Euspondylus) guentheri, 22
fraseri, 22
gaudichaudii, 3
oculatus, 32
(Oreosaurus) striatus, 32
(Pholidobolus) montium, 29
ecpleopus, Neusticurus, 26
edracantha, Ameiva, 11
elegans, Anolis, 16
Iphisa, 23
Emphrassotis simoterus, 32

46 PROCEEDINGS OF THE NATIONAL MUSEUM

Enyalioides, 3, 20
festae, 21
heterolepis, 21
laticeps festae, 21
laticeps laticeps, 21
leechi, 20
microlepis, 21
mocquardi, 21
oshaughnessyi, 21
palpebralis, 20
praestabilis, 21

Enyalius, 3, 21
heterolepis, 21
laticeps, 21
microlepis, 21
oshaughnessyi, 21
praestabilis, 21
zonatus, 21

eulaemus, Anolis, 15

Euspondilus festae, 27

Euspondylus, 3, 21
guentheri, 22
maculatus, 22
strangulatus, 27

Euspondylus, subg., 22

fasciatus, Anolis, 16
femoralis, Polychrus, 30
ferrugineus, Goniodactylus, 22
festae, Alopoglossus, 11
Anolis, 15
Enyalioides, 21
Enyalioides laticeps, 21
Euspondilus, 27
Leiocephalus, 28
Lepidoblepharis, 24
Ophryoessoides, 28
flaviceps, Doryphorus, 36
Uracentron, 36
flavipunctata, Aporomera, 19
flavipunctatus, Callopistes, 19
formosus, Liocephalus, 28
Ophryoessoides, 28
fraseri, Anolis, 16
Eepleopus, 22
fusco-auratus, Anolis, 16
fuscoauratus, Anolis fuscoauratus, 16

galeritus, Basiliscus, 19
gaudichaudi, Cercosaura, 20, 29
gaudichaudii, Ecpleopus, 3
Gekko rapicaudus, 35
Gekkonidae, 3

VOL. 119

Gekkoninae, 3
gemmosus, Anolis, 16
Gonatodes, 3, 22
caudiscutatus caudiscutatus, 22
collaris, 22
concinnatus, 22
oxycephalus, 25
Goniodactylus buckleyi, 23
concinnatus, 23
ferrugineus, 22
gracilipes, Anolis, 16
granuliceps, Anolis, 16
guayaquilensis, Phyllodactylus, 29
guentheri, Ecpleopus (Euspondylus), 22
Euspondylus, 22
Liocephalus, 28
Ophryoessoides, 28
guttulatum, Dicrodon, 19
gutturosus, Polychrus, 30
Polychrus gutturosus, 30
Gymnodactylus caudiscutatus, 22
Gymnophthalmus, 9, 23
speciosus speciosus, 23

haenschi, Liocephalus, 28
Ophryoessoides, 28
Hapalolepis, subg., 27
Hemidactylus, 5
heterolepis, Enyalioides, 21
Enyalius, 21
Holcosus bridgesii, 11
holotropis, Tropidurus, 35
Hoplocercus annularis, 26
horrida, Echinosaura, 20
Echinosaura horrida, 20
humeralis, Microphractus, 34
Stenocercus, 34
hypostictus, Proctoporus, 32
Hypsibates agamoides, 29

Iguana, 3, 23
iguana iguana, 23
tuberculata, 23

iguana, Iguana iguana, 23
Lacerta, 23

Iguanidae, 3

intermedius, Lepidoblepharis, 24

Iphisa, 3, 23
elegans, 23

iridescens, Ophryoessoides, 28
Liocephalus, 28

irregularis, Anolis, 17

jamaicensis, Tiliqua, 20

NO. 3545 LIZARDS—PETERS 47

Kentropyx, 3, 23
altamazonicus, 24
calearatus, 24
pelviceps, 24

Lacerta basiliscus, 19
iguana, 23
mabouya, 25
marmorata, 30
plica, 29
umbra, 29
Laemopristus, subg., 36
laticeps, Enyalioides laticeps, 21
Enyalius, 21
latifrons, Anolis, 17
leechi, Enyalioides, 20
Leiocephalus aculeatus, 28
angulifer, 28
festae, 28
ornatus, 28
lemniscatus, Anolis, 18
lentiginosus, Cnemidophorus, 19
Lepidoblepharis, 3, 24
buchwaldi, 24
festae, 24
intermedius, 24
oxycephalus, 25
ruthveni, 25
Leposoma, 3, 25
buckleyi, 10
carinicaudatum, 10
parietale, 25
picticeps, 33
leucostigma, Ameiva, 19
lineolatus, Sphaerodactylus, 34
Liocephalus formosus, 28
guentheri, 28
haenschi, 28
iridescens, 28
ornatus, 28
rhodomelas, 28
trachycephalus, 28
liogaster, Polychrus, 4
lionotus, Anolis, 13
lividus, Proctoporus, 33

mabouya, Lacerta, 25
Mabuya, 25
Mabuia aenea, 25
cepedei, 25
Mabuya, 3, 25
agilis, 25
mabouya, 25
macrolepis, Anolis, 14

Macropholidus, 3, 26
annectens, 26

maculatus, Euspondylus, 22

maculiventris, Anolis, 17

manicata, Cercosaura (Prionodactylus),

ol

manicatus, Prionodactylus, 31

marmorata, Lacerta, 30

marmoratus, Polychrus, 30

meleagris, Proctoporus, 32

microlepis, Enyalioides, 21
Enyalius, 21

Microphractus humeralis, 34

Mionyx parietalis, 25

mocquardi, Enyalioides, 21

Monoplocus, 3, 26
dorsalis, 26

monotropis, Diploglossus, 20
Scincus, 20

montium, Ecpleopus (Pholidobolus), 29
Pholidobolus, 29

Morunasaurus, 3, 26
annularis, 26

nasicus, Anolis, 18
Neusticurus, 3, 26
cochranae, 26
ecpleopus, 26
ecpleopus cochranae, 26
strangulatus strangulatus, 27
tuberculatus, 26
nigrolineatus, Anolis, 17
nigromaculatus, Stenocercus, 35
nigropunctatus, Tupinambis, 36
notopholis, Anolis, 14

occipitalis, Tropidurus (Laemopristus),
36

Tropidurus occipitalis, 36
ocellata, Anadia, 13
ocellifer, Prionodactylus, 31
oculatus, Ecpleopus, 32

Proctoporus, 32
Ophiognomon, 3, 27

abendrothii, 27

trisanale, 27
Ophryoessoides, 3, 27

aculeatus, 28

festae, 28

formosus, 28

guentheri, 28

haenschi, 28

iridescens, 28

ornatus ornatus, 28

48 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Ophryoessoides—Continued
rhodomelas, 28
orcesi, Ameiva, 12
Oreosaurus, subg., 32
ornatissima, Saccodeira, 33
ornatissimus, Proctotretus, 33
ornatus, Leiocephalus, 28
Liocephalus, 28
Ophryoessoides ornatus, 28
ortoni, Anolis,17
oshaughnessyi, Enyalioides, 21
Enyalius, 21
Prionodactylus, 31
oxycephalus, Gonatodes, 25
Lepidoblepharis, 25

pachyurus, Proctoporus, 31, 33
palpebralis, Enyalioides, 20
Pantodactylus, subg., 18, 31
parietale, Leposoma, 25
parietalis, Mionyx, 25
parvauritus, Anolis biporcatus, 15
pelviceps, Centropyx, 24
Kentropyx, 24
peraccae, Anolis, 17
peruvianus, Stellio, 36
Tropidurus peruvianus, 36
petersi, Ameiva, 11
Ameiva ameiva, 11
Pholidobolus, 3, 29
montium, 29
Pholidobolus, subg., 29
Phyllodactylus, 3, 29
abruptiseriatus, 29
guayaquilensis, 29
reissii, 29
picticeps, Leposoma, 33
Ptychoglossus, 33
Plica, 3, 29
plica, 29
umbra, 29
plica, Lacerta, 29
Plica, 29
poecilopleurus, Ablepharus, 10
Ablepharus boutonii, 10
Polychrus, 3, 30
femoralis, 30
gutturosus, 30
gutturosus gutturosus, 30
gutturosus spurrellii, 30
liogaster, 4
marmoratus, 30
spurrellii, 30

praestabilis, Enyalioides, 21
Enyalius, 21
princeps, Anolis, 17
Prionodactylus, 3, 30
manicatus, 31
ocellifer, 31
oshaughnessyi, 31
vertebralis, 31
Prionodactylus, subg., 31
proboscis, Anolis, 17
Proctoporus, 3, 31
columbianus, 32
hypostictus, 32
lividus, 33
meleagris, 32
oculatus, 32
pachyurus, 31, 33
simoterus, 32
striatus, 32
unicolor, 35
Proctotretus, 3, 33
ornatissimus, 33
Pseudogonatodes, 5
Ptenosaura seemani, 19
Ptychoglossus, 3, 33
bilineatus, 33
brevifrontalis, 33
picticeps, 33
pulchellus, Anolis, 4

rapicaudus, Gekko, 35
Thecadactylus, 35

reissii, Phyllodactylus, 29

reticulata, Arthrosaura reticulata, 18
Cerecosaura (Pantodactylus), 18

rhodomelas, Liocephalus, 28
Ophryoessoides, 28

rhombifera, Anadia, 13
Cercosaura, 13

Riama unicolor, 33

ruthveni, Lepidoblepharis, 25

Saccodeira ornatissima, 33
scapularis, Sphaerodactylus, 34
Scincidae, 3
Scincus monotropis, 20
secypheus, Anolis, 18
seemani, Ptenosaura, 19
septemlineata, Ameiva, 12
sexscutata, Ameiva, 12
simonsii, Stenocercus, 35
simoterus, Emphrassotis, 32
Proctoporus, 32

NO. 3545

speciosus, Blepharactisis, 23
Gymnophthalmus speciosus, 23
Sphaerodactylidae, 3
Sphaerodactylus, 3, 34
lineolatus, 34
scapularis, 34
spurrellii, Polychrus, 30
Polychrus gutturosus, 30
squamulatus, Anolis, 16
Stellio peruvianus, 36
Stenocercus, 3, 34
boettgeri, 34
carrioni, 34
humeralis, 34
nigromaculatus, 35
simonsii, 35
varius, 35
stigmosus, Anolis, 18
strangulatus, Euspondylus, 27
Neusticurus strangulatus, 27
striatus, Ecpleopus (Oreosaurus), 32
Proctoporus, 32

teguixin, Tupinambis, 36

Teiidae, 3

Thecadactylus, 3, 35
rapicaudus, 35

Tiliqua jamaicensis, 20

torquatus, Tropidurus, 35

trachycephalus, Liocephalus, 28

transversalis, Anolis, 18

trisanale, Ophiognomon, 27

tropidogaster, Anolis, 18

LIZARDS—PETERS 49

Tropidurus, 3, 35
continentalis, 36
holotropis, 35
(Laemopristus) occipitalis, 36
occipitalis occipitalis, 36
peruvianus peruvianus, 36
torquatus, 35

tuberculata, Iguana, 23

tuberculatus, Neusticurus, 26

Tupinambis, 3, 36
nigropunctatus, 36
teguixin, 36

umbra, Lacerta, 29
Plica, 29
unicolor, Proctoporus, 33
Riama, 33
Uracentron, 3, 36
azureum, 36
castor, 36
flaviceps, 36
Uranoscodon, 6

varia, Amphisbaena, 12
Amphisbaena fuliginosa, 12
varius, Stenocercus, 35
ventrimaculatus, Anolis, 14
vertebralis, Cercosaura (Pantodacty-
lus), 31
Prionodactylus, 31
viridiaeneus, Anolis, 16
vittatus, Basiliscus, 19

zonatus, Enyalius, 21

U.S. GOVERNMENT PRINTING OFFICE: 1967

Proceedings of

the United States

National Museum

SMITHSONIAN INSTITUTION +« WASHINGTON, D.C.

Volume 119 1966 Number 3546

NOCTUID MOTHS
OF THE
AMERICAN GENUS EUSCEPTIS HUBNER

By E. L. Topp?

This paper, a revision of the genus Husceptis Hiibner, is the first
of a proposed series of revisionary papers on the American species of
the noctuid subfamily Acontiinae. The purpose of this paper is to
provide keys, illustrations, and descriptions that will facilitate
identifications, and to assemble the accumulated knowledge pertaining
to the distribution and biology of the species of the genus. The
revision is the result of a study of the literature and the specimens in
the collection of the United States National Museum, supplemented by
specimens and information received from the following individuals
and institutions: D. 8. Fletcher, British Museum (Natural History),
London, England; W. Forster, Zoologische Sammlung des Bayerischen
Staates, Munich, Germany; H. K. Clench, Carnegie Museum, Pitts-
burgh, Pa.; J. G. Franclemont, Cornell University, Ithaca, N.Y.;
F. H. Rindge, American Museum of Natural History, New York,
N.Y.; F. F. Yépez, Universidad Central de Venezuela, Maracay,
Venezuela; P. Kohler, Buenos Aires, Argentina; C. M. Biezanko,
Pelotas, Rio Grande do Sul, Brazil; and C. V. Covell, Jr., Louisville,
Ky. Their assistance is gratefully acknowledged.

1 Entomology Research Division, Agriculture Research Service, U.S. Depart-
ment of Agriculture, Washington, D.C.

1

2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

The line drawings of genitalia were prepared by Arthur Cushman,
Scientific Illustrator, Entomology Research Division. The photo-
graphs of adult moths were made by staff photographers of the
Photographic Services, U.S. Department of Agriculture.

Genus Eusceptis Hiibner

Eusceptis Hiibner, 1823, Zutrige zur Sammlung exotischer Schmettlinge [sic],
Zweites Hundert, p. 21.—Hampson, 1910, Catalogue of the Lepidoptera
Phalaenae in the British Museum, vol. 10, p. 793.—Sherborn, 1922, Index
animalium 1801-1850, p. 2250.—Schulze, et al, 1929, Nomenclator animalium
generum et subgenerum, vol. 2, p. 1232.—Hemming, 1937, “Hubner,” vol. 2,
p. 198.—Neave, 1939, Nomenclator zoologicus, vol. 2, p. 370. (Type species:
Eusceptis irretita Hiibner, monobasic.)

Eugraphia Guénée, 1852, Histoire naturelle des insectes: Species général des
lépidoptéres, vol. 6 (Noctuélites, II), p. 208.—Walker, 1857, List of the
specimens of lepidopterous insects in the collection of the British Museum,
pt. 12, p. 776.—Scudder, 1882, Bull. U.S. Nat. Mus., no. 19, pt. 2, p. 119.—
Druce, 1889, in Godman and Salvin, Biologia Centrali-Americana, Insecta,
Lepidoptera, Heterocera, vol. 1, p. 304; 1896, Ann. Mag. Nat. Hist., ser. 6,
vol. 18, p. 42.—Hampson, 1910, Catalogue of the Lepidoptera Phalaenae in
the British Museum, vol. 10, p. 792.—Strand, 1913, Arch. Naturg., vol. 79
(A), nos. 7-9, p. 62; 1916, Arch. Naturg., vol. 82 (A), nos. 1-13, p. 39.—
Schulze, et al, 1929, Nomenclator animalium generum et subgenerum, vol. 2,
p. 1274.—Draudt, 1939, in Seitz, Gross-Schmetterlinge der Erde, vol. 7,
p. 397.—Neave, 1939, Nomenclator zoologicus, vol. 2, p. 326.—Biezanko,
Ruffinelli and Carbonell, 1957, Rev. Fac. Agron. Univ. Rep. Uruguay, no. 46,
p. 53. (Type species: Eusceptis irretita Hiibner, monobasic.) [New synon-
ymy.]

The generic names Husceptis Hiibner and EHugraphia Guénée are
isogenotypic. Guénée, 1852, accepted Hiibner’s specific name, but
not his generic name. Instead, he proposed Hugraphia as the generic
name for irretita Hiibner. The Hiibnerian generic name has also
been ignored in all subsequent taxonomic treatments and in all
catalogs.

Head with proboscis well developed; labial palpi small, nearly
porrect, very slightly curved dorsad, third segment very short, second
seoment three or four times as long, vestiture mainly of appressed
scales except longer and looser along ventral margin especially of two
basal segments; frons slightly bulbous, smooth or roughened medially,
clothed mostly with appressed, incurved scales; eyes large, hemi-
spherical, naked; ocellus small, adnate to upper margin of eye im-
mediately caudad of base of antenna; antenna usually more or less
filiform, oval or cuneate in cross section, pubescent ventrally and
shortly spiculate, males of E. effusa (Druce) and E. atriora, new
species, with longer spicules. Vestiture of thorax a mixture of scales
and hair, no obvious crests present, metathoracic scales occasionally
as slight decumbent crest; tympanum shielded dorsally by a large

NO. 3546 MOTHS OF GENUS EUSCEPTIS—TODD 3

alular fan of very large scales. Abdomen clothed with hair and scales,
no dorsal tufts present, terminal tufts about genitalia in males of
effusa and atriora. Legs rather stout, tarsus longer than tibia; tibia
of foreleg about half as long as femur, bearing a well-developed
epiphysis, tibia of middle and hindlegs longer; inner spurs of pairs
on middle and hindlegs about twice as long as outer spur; tibia of
middle leg rough scaled with median and apical tufts. Forewing
traingular, apex rounded, termen rounded except slightly excavate
before tornus, inner margin slightly sinuous, excavate distally; hind-
wing more or less rounded, very slightly angled at Cu,, inner margin
expanded over abdomen in effusa and atriora, a hair pencil on ventral
surface of expansion; venation of general noctuid type, forewing
with R; from R, adnate with R, to form small elliptical accessory
cell, R; stalked with R, for one-fourth their length from apex of
accessory cell, stalk of R3;+ R, connate with R; at apex of accessory
cell, M, from mdc slightly below accessory cell, M2 from just above
lower angle of discal cell, M; from lower angle of discal cell, Cu, from
just basad of lower angle of discal cell, Mz and Cu, approximately
equidistant from one another, R, and Cu, from the outer third of the
two sides of the discal cell; hindwing with Rs and M, connate from
upper angle of discal cell, M2 from below middle of discocellulars,
very weak or partially obsolescent, parallel to M;, M; and Cu, connate
or shortly stalked from lower angle of discal cell; length of forewing
9 to 17 mm.

Male genitalia with moderate, slightly curved uncus, uncus with a
minute hooked spine at apex, a row of basally directed spines along
either ventrolateral margin of uncus; scaphium of two straplike
sclerotizations, each terminating in a dense hair tuft; juxta irregularly
diamond shaped with an elongate, dorsal, straplike sclerotization;
valves moderately large, widest just before apex, interfacial processes
when present usually asymmetrical, sacculus well developed, usually
bearing a large apical or costal spine, a large clasper usually present
on one or both valves, absent in E. irretita Hiibner; when present
apex usually hooked, length of clasper usually bearing a row of
fascicles of hair and frequently a row of sclerotized triangular denta-
tions; corona short, consisting of about 10 spines, absent in E. irretita
and E. koehleri, new species, coronal spines nearly as long as width
of corona; aedeagus moderate, slightly shorter than valve, slightly
sinuous, largest at apex, vesica with scobinate areas but no conspicuous
cornuti. Female genitalia with valve of ovipositor roundly pointed,
moderately sclerotized bearing rows or clumps of large setae basally
and clumps of smaller setae distally, posterior apophyses slightly
longer than anterior pair; posterior margin of seventh abdominal
sternite usually emarginate medially, the lateral lobes thus formed

4 PROCEEDINGS OF THE NATIONAL MUSEUM vou, 119

usually slightly different in size and shape; ductus bursae usually
with a sclerotization near ostium, absent in F. obscura (Schaus),
remainder broad, twisted, and irregularly furrowed; bursa copulatrix
membranous, lacking signa, internally uniformly spiculate, the
spicules extremely minute except in E. obscura.

The immature stages and food plants of the species of Husceptis
Hiibner are completely unknown.

The genus Eusceptis Hiibner belongs to the tribe Acontiini and
apparently is most closely related to the genus Acontia Ochsenheimer
(=Tarache Hiibner). The two were separated in the “Key to the
Genera” (Hampson, 1910, Catalogue of the Lepidoptera Phalaenae
in the British Museum, vol. 10, pp. 7 and 8) by choices ‘‘d?’ and
‘““e?,”’ the difference being whether the frons possessed a rounded
prominence or no prominence. The species of Acontia usually do
possess a more bulbous frons than the species of Husceptis, but it is
a very slight difference of degree and scarcely sufficiently distinctive
to be considered diagnostic for these genera. In the generic diagnoses
in the same work, Hampson (op. cit., pp. 723 and 792), indicates that
Acontia (as Tarache) species have a decumbent metathoracic crest
which is lacking in Husceptis (as Hugraphia) species. In some species
of Acontia the decumbent crest is no more developed than in the
species of Husceptis. The two genera are very closely related, but
may be separated by the nature of the uncus of the male genitalia.
In Eusceptis the uncus has a row of basally directed spines along
either ventrolateral margin. The uncus is simple in Acontia.

The species of the genus are essentially found in the neotropics,
but some species approach the temperate regions of North and South
America.

Key to Adults of Eusceptis

(Based on wing maculation)

13 Hindwing black)... —) 2.4 Seoul Rika re obscurai(ps6)
Hindwing mostly white, crane or recliony Sh Fis Beweaen rien yi kal coe a earn!

2. Forewing with some yellow or orange in the terminal area between vein M,
andthe tornus ... ay lia Jay eg weed Gust eset A SG ia a ae
Terminal area of foe dnl bh eNotes et lls ort Oar Sa oy eee

3. Dark marginal band of hindwing of male nearly anifer m in ree same band
of female usually terminating at about Cu... ... . . . .effusa (p.7)

Dark marginal band of hindwing of male tapering toward anal angle, width

at Rs+M, about twice width at Cu,; same band of female reaching anal

Bm Ce cj Son an, Le io nn Poe bas ae aes Ona ake Beane eee al COLE (tS)

4. Outer, dark costal mark of forewing about equal in width to median mark. 5
Outer, dark costal mark of forewing about twice as wide as median mark . 6

NO. 3546 MOTHS OF GENUS EUSCEPTIS—TODD o

5. Outer, dark costal mark of forewing looped basad immediately above lower
angle of cell; orange terminal line of forewing curved around apex, nearly
reaching costa ... . . .. . . Koehleri (p. 11)
Outer, dark costal mark oe foromnie Gaus ot looped basad above lower
angle of cell; orange terminal line of forewing not curved around apex.
irretita (p. 10)

6. Dark basal part of ocellate spots of postmedian area of forewing separate;

fringe usually paler than oblique bar, especially at outer edge... . 7
Dark basal part of ocellate spots of forewing united into a sinuous line; fringe
uniformly dark 23 =) -)- 5 ds 29

7. Hindwing of male mostly te ait Beane near weutcn margin only; sub-
terminal, oblique, gray bar of forewing of female wider at basal end than at
apex tanean: . . . extensa (p. 17)

Hindwing of ee mosey, pemaee: perineal! Gphane: gray bar of forewing
of female usually widest near apex; distal yellow area level with apex of
Cu, as wide as or wider than base of oblique, gray bar. ....... 8

8. Fringe of forewing more or less concolorous with gray terminal line; terminal

line extending from dark apex almost to tornus (South America).
splendens (p. 13)
Fringe of forewing usually paler than gray terminal line; terminal line not
reaching apex or crossing anal vein (Central America) . . lelae (p. 16)

9. Hindwing of male orange; oblique, subterminal, gray bar of forewing divided

or paler at middle; hindwing of female lacking a fuscous spot at apex.
robertae (p. 18)
Hindwing of male with basal half whitish; oblique, subterminal, gray bar not
divided or paler at middle; female with fuscous spot present at apex of
ing wintmepe- ey lewe stun eee ne va paraguayensis: (ps 19)

Key to Males of Eusceptis

(Based on genitalia)

1. Valves lacking a corona (figs. 19 and 20) .... NL eae ee neg
Valves with a conspicuous short corona (example, fy. 22) oe ap er pee ase

2. Sacculus of right valve greatly enlarged, right valve with clasper present
distad of sacculus (fig. 20). .... . . . . . Koehleri (p. 11)
Sacculus of right valve not enlarged, pope i sacculus of left valve, valves
Bimiple; ‘clasper absent, (fig) 19) 2 So, oe 2s se irretita (p:. 10)

3. Left valve lacking a clasper (fig. 22). ........ .. . . obscura (p. 6)
ett valve with’aclasper (example, fig. 23). 055 2 SO ai es

4. Sacculus of left valve more than half as long as valve, apex of sacculus pro-
duced into a large dorsally directed spine (figs. 23 and 24) ...... 5

Sacculus of left valve short, usually decidedly less than half as long as valve,
spine of sacculus small or arising from near middle of costal margin (exam-
De OEA 7) ete rts FG eo te em OR Cle URC oo sib is Ut mies nO

5. Costal spine of sacculus of right valve longer than distance from its base to
apex of sacculus; sacculus of left valve only slightly wider than sacculus of
right valve (fig. 24) . RAD te othe eens eitasar (pend)

Costal spine of sacculus of right wale ahrovter thax muatanied from its base to
apex of sacculus; sacculus of left valve distinctly wider than sacculus of
right valve (fig. 23) ney adie oy bea sitcuuc, po een aula atyhoh eA ORA (Dine)

6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

6. Clasper of left valve short, apex scarcely reaching apical third of valve. . 7
Clasper of left valve long, reaching beyond apex of valve. .. . 9
7. Sacculus of right valve with a spine poe from the dorsal oes ee
29) pays & acs arte . .. . extensa (p. 17)
Sacculus of right oalve pane a spine on ane. Aomal margin (figs. 25 and
Da Te es Shae RS
8. Clasper of left ae ee eeu, ree ehenels, alubbed (Giz. 25),
lelae (p. 16)
Clasper of left valve hooked, apex usually pointed (fig. 27) . splendens (p. 13)
9. Clasper of right valve short, reaching about to middle of valve (fig. 31).
robertae (p. 18)
Clasper of right valve long, reaching slightly beyond apex of valve (fig. 34).
paraguayensis (p. 19)

Eusceptis obscura (Schaus), new combination
Figures 7, 8, 22

Acontia obscura Schaus, 1898, Journ. New York Ent. Soc., vol. 6, no. 2, p. 117.

Eugraphia obscura (Schaus).—Hampson, 1910, Catalogue of the Lepidoptera
Phalaenae in the British Museum, vol. 10, p. 795, pl. 174, fig. 3—Draudt,
1939, in Seitz, Gross-Schmetterlinge der Erde, vol. 7, p. 397.

Acontia trilinea Schaus, 1898, Journ. New York Ent. Soc., vol. 6, no. 2, p. 117.
[New synonymy.]

Eugraphia trilinea (Schaus).—Hampson, 1910, Catalogue of the Lepidoptera
Phalaenae in the British Museum, vol. 10, p. 795, pl. 174, fig. 2.—Draudt,
1939, in Seitz, Gross-“Schmetterlinge der Erde, vol. 7, p. 397.

The uniformly black hindwings of this species separate it from all
the other known species of the genus. Length of forewing: male,
16 mm.; female, 17mm.

Types: The type of obscura (USNM 10658), a female from Guada-
lajara, Mexico, and that of trilinea (USNM 10659), a male from
Jalapa, Mexico, are in the collection of the U.S. National Museum.

Distribution: Known only from Guadalajara and Jalapa, Mexico.
Only four specimens are known to have been collected.

Remarks: Schaus indicated in the original descriptions that
trilinea (fig. 7) and obscura (fig. 8) might represent the two sexes of
a single species. The two names apply to the same species, but it
is possible that the specimen named ¢rilinea may represent a distinct
form. The two known male specimens have the same genitalia
(fig. 22), but they differ in maculation of the forewing, one specimen
being dark like the type of obscura. Because only two males are
available for study, it is not known whether specimens exist that are
intermediate in maculation of the forewing between the dark type of
obscura and the orange-streaked type found in ¢rilinea. The solution
of this problem must of necessity await the collection of more speci-
mens of this species.

Method of determination: By examination of the type.

NO. 3546 MOTHS OF GENUS EUSCEPTIS—TODD iG

Eusceptis effusa (Druce), new combination
Fiaurss 4, 5, 24

Eugraphia effusa Druce, 1889, in Godman and Salvin, Biologia Centrali-Ameri-
cana, Insecta, Lepidoptera, Heterocera, vol. 1, p. 304, table 28, fig. 12; 1898,
ibid, vol. 2, p. 492.—Hampson, 1910, Catalogue of the Lepidoptera Phalaenae
in the British Museum, vol. 10, p. 794.—Strand, 1913, Arch. Naturg., vol.
79 (A), nos. 7-9, p. 62.—Draudt, 1939, in Seitz, Gross-Schmetterlinge der
Erde, vol. 7, p. 397.

This species and the following one, EF. atriora, new species, differ
from the other species of the genus in which the ground color of the
hindwing is orange, yellow, or white (figs. 1-3, 9-18) in that the
forewing distad of the postmedial band (excluding the costal pale
spot) is dark (figs. 4-6), lacking the yellow or orange line that extends
from tornus toward apex in the terminal area of the forewing of the
other species. They also have a medial line of metallic scales in the
basal and distal black costal marks of the forewing. The costal
marks are uniformly dark in the other species. In addition, the
hindwings of males of effusa and atriora are modified. The inner
margin of the hindwing is expanded, upturned and curved over the
abdomen. ‘This modified area of the hindwing forms a concavity on
the ventral surface and contains a tuft of long hairs. E. effusa
(Druce) and FE. atriora, new species, are extremely closely related
entities; however, there are a number of differences in maculation
and in the male genitalia. One of the more obvious differences is
the degree of development of the dark marginal band of the hind-
wing. In effusa males the marginal band of the hindwing is nearly
uniform in width and extends from the apex to about Cu,. In
males of atriora the marginal band at Rs+M, is about twice as wide
as at Cue, the band being distinctly tapered and extending from apex
to beyond Cup. In the females of effusa the marginal band is rather
similar to that of the males of atriora, but usually it is broader and more
truncate at about Cue. In females of atriora the marginal band is
much like that of effusa to Cus, but continues to taper on to the anal
angle. One female specimen from Cayuga, Guatemala does have the
marginal band terminating in a tapered point, but the band does not
extend to the anal angle. There are other differences in maculation
between the two species. Those differences and the differences in the
male and female genitalia will be discussed in the description of atriora.
Length of forewing: male, 14 mm.; female, 14 to 15 mm.

Type: The type, a female from Volcan de Chiriqui, Panama, is in
the collection of the Kénigliche Zoologische Museum, Berlin, Germany.

Distribution: Southern Texas to Panama. Specimens from the fol-
lowing localities have been examined. Texas: Brownsville. Mexico:
Misantla, Tamazunchale, Poza Rica. Guatemala: Cayuga, Quirigua.

8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 119

A female from Santa Clara Valley, Costa Rica has been referred to
this species by Druce and by Hampson. It will be necessary to
examine the specimen to determine whether it is correctly placed.
It is possible that it should be referred to the following species, atriora.

Remarks: Because I have this species only from localities in or
to the north of Guatemala, and because it was not found in the large
series of noctuids collected by Zetek and others on Barro Colorado
Island, Canal Zone, Panama, the type locality (Panama) cited by
Druce might be incorrect. Unless Druce misread the locality label
on the type, it does not seem likely that it could be proved that an
error might have occurred. On the other hand, the existence of
specimens from Costa Rica and/or Panama or the subsequent collec-
tion of the species from those countries would certainly eliminate any
doubt as to the correctness of the type locality.

Method of determination: This species is easily recognized by the
excellent illustration of the female type accompanying the original
description.

Eusceptis atriora, new species
FIGurEs 6, 23

Head with proboscis well developed; labial palpi small, very slightly
curved dorsad, apical segment slightly exceeding ventral margin of
frons, third segment very short (0.3 mm.), second segment four times
as long, vestiture mainly of appressed, pale yellow scales except longer
and looser along ventral margin, especially first segment and base of
second segment, dorsum and dorsolateral area of second segment dark
brown or black; frons smooth, scarcely exceeding anterior margin of
eye, vestiture of appressed, down and incurved pale yellow scales; eyes,
large, hemispherical, naked, about equal to frons in width; ocellus
small (0.1 mm.), adnate to upper margin of eye immediately caudad of
base of antenna; antenna black, filiform, spiculate laterally and
ventrally, the spicules of the male about twice as long (0.1 mm.) as
those of the female and more numerous. Vestiture of patagia,
tegulae, and thorax a mixture of pale yellow hair and scales; a slight
decumbent, metathoracic crest present. Abdomen orange dorsally
and laterally, pale yellow or white ventrally with terminal black scales
on segments 3 to 7 in the females, the males with much less black
scaling except on segment 7; dorsal tufts absent. Pectus clothed with
large, pale yellow or white scales and long, sparse, white hair; tympa-
num shielded dorsally by a very large alular fan of broad pale yellow
scales. Legs banded with dark brown and pale yellow or white scales,
the dark brown scales highly iridescent in certain lights; foreleg with
tibia and tarsus clothed with dark brown scales except some pure white
scales at base and apex of first tarsomere and at base of second tar-

Beaugenze MOTHS OF GENUS EUSCEPTIS—TODD 9

somere, scales of distal three-fifths of femur dark brown, scales of basal
part white, trochanter clothed with longer, less appressed, pale yellow
scales; middle leg similar to foreleg except tibia with a broad median
ring of loose pale yellow scales, the dark scales at apex long, loose,
down curved, forming a broad tuft, white scales at base of femur
extending along dorsum to apex, tibial spurs dark brown except
extreme apex of long, inner spur white; hindleg similar to middle leg
except scales of tibia appressed, pale median part white and extending
to base along dorsum, basal tibial spurs white except apical half of
short outer spur. Venation of wings as for genus, accessory cell very
small and narrow as in effusa. Wing shape as in effusa (figs. 4-6),
male with inner margin of hindwing modified, upcurved over abdomen
and bearing a tuft of long hairs in the pocket formed on the ventral
surface. Pattern of maculation of upper surface of wings as illus-
trated (fig. 6, female), male resembling female except dark marginal
band of hindwing less extensive, reaching only slightly beyond Cup.
Ground color of basal part of forewing pale yellow; dark part of dark
brown scales some iridescent brassy green in certain lights, some pale
blue white scales usually present above tornus in cells Cu, and Cup;
Y-shaped transverse line of metallic gray scales heavily bordered by
dark brown scales. Hindwing yellow orange except dark brown mar-
ginal band, fringe dark with some white at outer edge; ventral surfaces
of wings similar to dorsal surfaces except metallic gray Y-shaped
transverse line of forewing absent, apical orange spot of hindwing
larger and fringe of hindwing mostly white, basal costal band of dorsal
surface of forewing absent on ventral surface of that wing in males.
Length of forewing: male, 13 to 14 mm.; female, 14 to 15 mm.

Male genitalia as illustrated (fig. 23), very similar to that of effusa
(fig. 24) except sacculus of left valve distinctly wider than sacculus
of right valve, dorsally directed process of sacculus of left valve ex-
tending beyond costal margin of valve at a point well below corona,
costal spine of sacculus of right valve shorter than distance from its
base to apex of sacculus, and costal angle of apex of right sacculus
forming a short, triangular projection. Female genitalia very similar
to those of effusa except right lobe of seventh abdominal sternite
weakly sclerotized along median emargination, left lobe of seventh
abdominal sternite with a short, blunt projection medially at base of
lobe, dorsal sclerotization of vaginal chamber about twice as long as
wide.

Type: Type female, Sixaola River, Costa Rica, April 1907, Wm.
Schaus (USNM 64639); 1 female paratype, same place, September;
and 1 female paratype, Guapiles, Costa Rica, 850 ft. alt., May 1907,
Wm. Schaus, in the U.S. National Museum, Washington, D.C. Two
male paratypes, Sixaola River, Costa Rica, September, in the Carnegie

219-944 662

10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Museum, Pittsburgh, Pa. One female paratype, Costa Rica, Never-
mann in the Zoologische Sammlung des Bayerischen Staates, Munich,
Germany.

Distribution: Known only from Costa Rica.

Remarks: At the beginning of this study I thought the three fe-
male examples in the U.S. National Museum probably represented
a dark form of effusa in which there was a greater development of
the dark marginal band of the hindwing. This seemed to be a reason-
able conclusion because other species of the genus, especially those
of the splendens complex, do show considerable variation in the de-
gree of development of the dark marginal band of the hindwings.
However, when males became available for study, constant differences,
even if slight, were found to exist in the genitalic characters and other
differences in maculation were recognized. The broad sacculus of
the left valve, the differently directed apical process of the left sac-
culus, the shorter costal spine of the right sacculus of the male
genitalia, the differences in the sclerotized parts of the female geni-
talia, the heavy dark markings of the forewing, the presence of blue
white patches of scales in cells Cu, and Cuz of the forewing, and the
dark fringe of the hindwing (yellow in effusa) convinced me that
this entity should be considered to be a species distinct from effusa.

The known localities for the species are both from the Caribbean
side of Costa Rica. It is possible, therefore, that this species is
geographically isolated from effusa, which may occur on the Pacific
side of Costa Rica. Proof of the correctness of this supposition must
of necessity depend on further collections of material from that
country.

Eusceptis irretita Hiibner
Figures 1, 19

Eusceptis irretita Hiibner, 1823, Zutrage zur Sammlung exotischer Schmettlinge
[sic], Zweites Hundert, p. 21, figs. 305, 306.

Eugraphia irretita (Hiibner).—Guénée, 1852, Histoire naturelle des insectes:
Species général des lépidoptéres, vol. 6 (Noctuélites, II), p. 208.—Walker,
1857, List of the specimens of lepidopterous insects in the collection of the
British Museum, pt. 12, p. 776.—Druce, 1889, in Godman and Salvin,
Biologia Centrali-Americana, Insecta, Lepidoptera, Heterocera, vol. 1, p.
304.—Hampson, 1910, Catalogue of the Lepidoptera Phalaenae in the
British Museum, vol. 10, p. 793, fig. 211.—Strand, 1913, Arch. Naturg.,
vol. 79 (A), nos. 7-9, p. 62.—Hemming, 1937, ‘‘Hiibner,’’ vol. 2, p. 198.—
Draudt, 1939, in Seitz, Gross-Schmetterlinge der Erde, vol. 7, p. 397.

Noctua melanogramma Perty, 1833, Delectus animalium articulatorum quae in
itinere per Brasiliam . . ., p. 163, pl. 32, fig. 10.—Walker, 1857, List of the
specimens of lepidopterous insects in the collection of the British Museum,
pt. 12, p. 776 [as a synonym of zrretita Hiibner].

|
|
|
|

NO, 3546 MOTHS OF GENUS EUSCEPTIS—TODD 11

This species, the type of the genus, is also the smallest species of
the genus. It agrees with Z. koehleri, new species, and differs from
the other similarly marked species in that the outer dark costal mark
of the forewing is about equal in width to the medial costal mark.
E. irretita Hiibner differs from koehleri by its slightly smaller size,
by the absence of a loop in the outer costal mark of the forewing
immediately above the lower angle of the cell, by the orange terminal
line of the forewing not being curved around the apex of the wing,
and by the characteristic male genitalia (fig. 19). The valvae of the
male genitalia are simple, without spines or processes developed from
or immediately beyond the sacculus. Length of the forewing: male,
9 to 10 mm.

Types: The present location of the type of irretita is unknown.
It may be in the Naturhistorisches Museum, Vienna, Austria.
Hiibner’s type was from Brazil. The type of melanogramma was
from “Provincia Bahiensi,’’ Brazil. It may be in the collection of
the Zoologische Sammlung des Bayerischen Staates, Munich,
Germany. At least part of Perty’s types were received by that
institution.

Distribution: . irretita apparently is restricted to Brazil, possibly
to the northeastern part of that country. Only four males have been
examined. One is unlabeled, the other three are from Bahia and
Pernambuco, Brazil.

Remarks: The illustration of Perty’s melanogramma appears to
be of afemale. At least the marginal band of the hindwing is exten-
sive and similar to that of the female of the related koehlerc.

Method of determination: The species was identified from the
excellent illustrations accompanying the original description. Perty’s
melanogramma was recognized as a synonym by means of the good
illustration in his work.

Eusceptis koehleri, new species
Figures 2, 3, 20, 21

Head with proboscis well developed; labial palpi small, very
slightly upcurved, apical (third) segment slightly exceeding ventral
margin of frons, third segment very short, second segment about four
times as long, vestiture mainly of closely appressed pale yellow scales,
except longer and looser at venter of first segment and second segment
black dorsally and dorsolaterally; frons slightly bulbous, exceeding
anterior margin of eye about one-fourth length of eye, rather rough
medially, depressed before slightly porrect ventral margin, vestiture
of appressed, down and incurved pale yellow scales; eyes large, about
equal to frons in width, naked, hemispherical; antennae black, filiform,

12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

densely pubescent ventrally, with minute spicules present laterally
and ventrally. Vestiture of patagia, tegulae, and thorax a mixture of
pale yellow hair and scales; a slight decumbent, metathoracic crest
present. Abdomen orange dorsally and laterally, pale yellow ven-
trally with lateral and/or terminal black scaling on segments 3 through
7; dorsal tufts absent. Pectus clothed with large, pale yellow or
white scales and long, sparse, white hair; tympanum nearly covered
by very large alular fan of broad yellow scales. Legs mainly pale
basally, dark distally; foreleg with tibia and tarsus dark brown except
small white spot at middle of tibia and white rings at base and apex
of first tarsomere and apex of second; middle leg pale yellow except
for dark brown knee spot, broad apical brown band on tibia (includ-
ing spurs) and dark tarsus, tarsomeres 1 and 2 marked with white
like those of foreleg; hindleg colored like preceding leg except dark
knee spot absent. Venation of wings as for genus. Maculation of
male similar to that of irretita (see figs. 1 and 2), except that outer,
dark costal mark of forewing is distinctly looped basad immediately
above lower angle of cell and terminal orange line extends around
apex of wing. Basal half of upper surface of forewing lemon yellow,
transverse lines black, subterminal oblique mark and fringe slate gray.
Upper surface of hindwing orange around margins, white medially,
afuscous spot at apex. Ventral surface of forewing with ground color
orange, median and apical costal marks, subterminal oblique mark
and fringe dark fuscous or black; hindwing ventrally like upper surface
except orange of costa brighter and small postmedial and antemedial
spots present. Female colored as in male except upper surface of
hindwing orange with broad marginal fuscous band, the latter in-
curved slightly at Cuz; ventrally postmedial fuscous mark extending
nearly to anal angle, broadest between M, and caudal end, antemedial
spot variable in size, absent in one specimen. Length of forewing:
male, 12 mm.; female, 12 to 13 mm.

Male genitalia of type specimen partially destroyed by dermestids;
however, the valves are preserved and are distinctive (fig. 21). Right
valve enlarged, much broader than left valve, sacculus broad, pro-
duced and curved toward costa at apex, a longer, curved process
developed immediately distad of sacculus from near ventral margin of
the valve. Female genitalia as illustrated (fig. 20), ductus bursae
broad, rather long, only slightly sclerotized; caudal margin of seventh
abdominal sternite not asymmetrical as in the females of the splendens
complex.

Types: Type male and two female paratypes, Alta Gracia, La
Granja, Sierras de Cordoba, Argentina, C. Bruch, in the private collec-
tion of P. Kéhler, Buenos Aires, Argentina. One female paratype,
same data, in the U.S. National Museum.

PR eemn! 5 6

Ficures 1-6.—Dorsal view of adults of Eusceptis species: 1, irretita, male, Bahia, Brazil;
2, koehleri, male, type, Alta Gracia, La Granja, Sierra de Cordoba, Argentina; 3, koehlert,
female, paratype, same place; 4, effusa, male, ‘‘Central America”; 5, effusa, female,
Misantla, Mexico; 6, atriora, female, type, Sixaola River, Costa Rica.

gg

uh

!
‘

oe 12

Freures 7—-12.—Dorsal view of adults of Eusceptis species: 7, obscura, male, type of trilinea
Schaus, Jalapa, Mexico; 8, obscura, female, type, Guadalajara, Mexico; 9, robertae,
male, type, Tucuman, Argentina; 10, robertae, female, paratype, same place; 11, para-
guayensis, male, Misiones, Argentina; 12, paraguayensis, female, Sapucay, Paraguay.

\ a ‘ , ¢
m s
3 % = a
* \ ’ a -
:

+ ? oe =
ie a

ff @ ©&

ae me 18

Frcoures 13-18.—Dorsal view of adults of Eusceptis species: 13, lelae, male, type, Avangarez,
Costa Rica; 14, lelae, female, paratype, Mexico; 15, splendens, male, Zaruma, El Oro,
Ecuador; 16, splendens, female, Rancho Grande, Aragua, Venezuela; 17, extensa, male,
Vicosa, Minas Geraes, Brazil; 18, extensa, female, Puerto Bertoni, Paraguay.

Figures 19-24.—Male and female genitalia of Eusceptis species (aedeagus of male removed
and figured separately): 19, irretita, male, Bahia, Brazil; 20, koehlerit, male, type, Alta
Gracia, La Granja, Sierra de Cordoba, Argentina (valves only); 21, koehleri, female,
paratype, same place; 22, obscura, male, Guadalajara, Mexico; 23, atriora, male, para-
type, Sixaola River, Costa Rica; 24, effusa, male, Cayuga, Guatemala.

Ficures 25-29.—Male and female genitalia of Eusceptis species (aedeagus of male removed
and figured separately): 25, lelae, male, type, Avangarez, Costa Rica; 26, lelae, female,
paratype, Mexico; 27, splendens, male, Zaruma, El] Oro, Ecuador; 28, splendens, female,
Aroa, Venezuela; 29, extensa, male, Misiones, Argentina.

Ficures 30-34.—Male and female genitalia of Eusceptis species (aedeagus of male removed
and figured separately): 30, extensa, female, Puerto Bertoni, Paraguay; 31, robertae,
male, paratype, Tucuman, Argentina; 32, robertae, female, paratype, same place; 33,
paraguayensis, female, Sapucay, Paraguay (ductus bursae); 34, paraguayensis, male,
Misiones, Argentina.

NO. 3546 MOTHS OF GENUS EUSCEPTIS—TODD 13

Distribution: Known only from the type locality.

Remarks: This species agrees with irretita and differs from the
similarly marked species in that the outer, dark costal mark of the
forewing is about equal in width to the median costal mark. The
species is slightly larger than zrretita, has the outer, dark costal
mark of the forewing distinctly looped immediately above lower
angle of the cell, and has the terminal orange line extending around
the apex of the wing. In addition, the male genitalia are specifically
distinct.

Eusceptis splendens (Druce), new combination

Figures 15, 16, 27, 28

Eugraphia splendens Druce, 1896, Ann. Mag. Nat. Hist., ser. 6, vol. 18,
p. 42.—Hampson, 1910, Catalogue of the Lepidoptera Phalaenae in the British
Museum, vol. 10, p. 793, pl. 174, fig. 1 [in part].—Strand, 1913, Arch. Naturg.,
vol. 79 (A), nos. 7-9, p. 62.— Kaye and Lamont, 1927, Mem. Dept. Agric.
Trinidad and Tobago, no. 3, p. 66 [in part]—Draudt, 1939, in Seitz, Gross-
Schmetterlinge der Erde, vol. 7, p. 397 [in part].

Eugraphia irretita (Hiibner)—Weymer and Maasen not Hiibner, 1890, Lepi-
dopteren gesammelt auf einer Reise durch Colombia, Ecuador, Peru,
Brasilien, Argentinien und Bolivien in den Jahren 1868-1877 von Alphons
Stubel, p. 67.

The splendens complex is composed of at least five species, some
of which have been misidentified as splendens or treated as forms or
aberrations of that species. The species of this complex (figs. 9-18)
may be separated from the other species of the genus by the following
combination of maculational characters: hindwing with some orange,
yellow, or white, terminal area of forewing with some yellow between
vein M, and tornus, and outer, dark costal mark of forewing about
twice as wide a median costal mark.

The species of the splendens complex form three segregates on
genitalic and maculational characters. . splendens (Druce) and
E. lelae, new species, extremely closely related species, form one segre-
gate. Two other species, EL. paraguayensis (Draudt) and E. robertae,
new species, belong to a second distinct segregate. The fifth species,
EH. extensa (Strand), is intermediate on maculation and genitalic
characters and is considered to be a separate segregate. The species
and the segregates of the splendens complex can be identified by
means of the characters utilized in the keys.

The known geographic distribution of the four species of the
splendens complex in South America is illustrated on map 1. The
fifth species, E. lelae, new species, is known to occur in Mexico,
Guatemala, and Costa Rica.

E. splendens (Druce) is redescribed in this paper because other
species of the complex have been confused with it in the past and

14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

because a detailed description of splendens will simplify the descriptions
of new species of the complex.

Head with proboscis well developed; labial palpi small, very
slightly curved dorsad, apical segment slightly exceeding ventral
margin of frons, apical segment very short (0.3 mm.), second segment
about three times as long, vestiture mainly of appressed, pale yellow
scales except longer and looser along ventral margin of first and
second segments, dorsum and dorsolateral area of second segment
light brown; frons smooth, scarcely exceeding anterior margin of eye,
vestiture of appressed, pale yellow scales; eyes large, hemispherical,
naked, about equal to frons in width; ocelli present, moderate, adnate
to upper margin of eye immediately caudad of base of antenna;
antenna dark reddish brown, filiform, ventral pubescent area wedge-
shaped in the male, round in the female, weakly spiculate. Vestiture
of patagia, tegulae, and thorax a mixture of pale yellow hair and
scales; a slight, decumbent, metathoracic crest present. Abdomen
mostly orange dorsally and laterally, the two basal segments pale
yellow, the remainder orange, the terminal segments darkest; venter
pale yellow except caudolateral angles and terminal margins of
sternites light to dark brown, the amount of brown variable, last
three sternites of females usually brown; dorsal tufts absent. Pectus
clothed with large white, appressed scales and long pale yellow hair
and scales; tympanum shielded dorsally by a very large alular fan
of very large pale yellow scales. Legs banded with various shades
of brown and pale yellow or white scales, foreleg darkest; tarsus of
foreleg dark brown except base and apex of first tarsomere and apex
of second tarsomere ringed with white, tibia dark brown distally,
light yellowish brown basally, a small median white point between
the two shades of brown, femur with basal third white, remainder
yellowish brown; tarsus of middle leg similar to that of foreleg, white
bands of tarsomeres slightly broader, tibia with a broad band of loose
pale yellow scales, base yellowish brown, apex darker forming a
terminal brush, base and dorsum of femur white, remainder yellowish
brown, tibial spurs usually white except basal half of short outer
spur yellow brown; hindleg similar to middle leg except white bands
of tarsomeres broader, scales of tibia more appressed, pale band
white, dark terminal part with shorter scales and forming a shorter
less prominent tuft, tibial spurs usually white except basal part of
short outer spur of terminal pair yellowish brown. Venation as for
genus. Wing shape and pattern of maculation of dorsal surface of
wings as illustrated (figs. 15 and 16). Ground color of dorsal surface
of forewing pale lemon yellow; transverse lines, oblique subterminal
bar and fringe gray brown, subterminal bar darkest toward apex,
paler and slightly narrowing toward base, fringe and terminal line

Bee MOTHS OF GENUS EUSCEPTIS—TODD 15

more or less unicolorous, the dark scaling usually extending to tornus;
some dull reddish brown shading basad of base of oblique subterminal
bar and beyond upper ocellate spot of postmedial band; ocellate spots
of postmedial band composed of basal brown crescent, median blue
eray line and an outer dark mark, outer dark mark of lower ocellate
spot nearly black, usually darker than similar mark of other ocellate
spot. Hindwing yellow orange, paler in male than in female, especially
toward base; a dark brown apical spot usually present, larger in females
than in males, in some females a dark subterminal spot present on
costa. Ventral surface of wings mostly yellowish orange, paler,
nearly white before apical dark spots; dark brown marks of forewing
consisting of oblique subterminal bar, two outer costal marks and
a dark outer mark of lower ocellate spot, oblique subterminal bar
with apical third much darker than remainder of bar, basal part
vague or absent in males. Length of forewing: male, 12 to 14 mm.;
female, 12 to 15 mm.

Male and female genitalia as illustrated (figs. 27, 28). Clasper
of left valve of male genitalia hooked or bent dorsad, the apex of the
clasper variable in shape, pointed or weakly clavate; left sacculus
only with a costal spine. Ductus bursae of female genitalia with a
sclerotized groove to the right; ductus seminalis arising from a lobe
on the right side of bursa copulatrix.

Type: A female from Guayaquil, Ecuador in the British Museum
(Natural History), London, England via the Druce collection. In
the original description Druce does not indicate the number of speci-
mens, but he indicates the specimen described was a male. The
statement in the original description (1896, p. 42) ‘. . . secondaries
bright chrome-yellow, with three black spots close to the apex... .”
proves that the specimen was a female and that Druce was in error
as to the sex. Males either lack an apical black spot on the hindwing
or have only a single spot, females may have one, two, or three spots.
Examples appearing to have three spots actually have the apical spot
divided into two spots by yellow orange scales along vein M,;. In the
British Museum there are four other specimens from Guayaquil,
Ecuador (Dolby-Tyler collection); but they were in that collection
before the type (see Hampson, 1910, p. 793).

Distribution: This species occurs in northern South America.
Thirty specimens from the following localities have been examined.
Ecuador: El Oro, Zaruma. Colombia: Popayan. Venezuela: Aroa;
Las Quiguas, Carabobo; Rancho Grande, Aragua; Alto de Yuma,
near Giigiie, Carabobo; El Limén, near Maracay, Aragua; Maracay,
Aragua; and Santa Lucia, Miranda. Trinidad: Caparo. In addition
to the five specimens from Guayaquil, Ecuador, there is one female
from Trinidad in the British Museum (Natural History).

16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Remarks: This species is very similar to EF. lelae, new species,
from which it may be separated by slight differences in maculation,
genitalia, and by geographical distribution as explained in the dis-
cussion of lelae. It may be separated from the other species of the
splendens complex by the characters indicated in the keys.

Method of determination: The species was identified from the
illustration in Hampson, 1910, pl. 174, fig. 1. A photograph of a
specimen so identified was then compared with the type for me by
Mr. D.S. Fletcher at the British Museum.

Eusceptis lelae, new species

Figures 13, 14, 25, 26

This species is very similar to splendens except the gray brown
terminal line usually does not cross the anal vein or reach the apex
of the forewing, the fringe usually paler; the dull reddish-brown
shading basad of base of oblique subterminal bar and distad of upper
ocellate spot of forewing paler; outer dark mark of upper ocellate spot
nearly as dark as similar mark of lower ocellate spot; clasper of left
valve straight or nearly so, clubbed apically (fig. 25). Length of
forewing: male, 13 to 14 mm.; female, 14 to 15 mm.

Type: Type male, Avangarez, Costa Rica, July-August, William
Schaus (USNM 64640); 1 male paratype, same place, July, Schaus
and Barnes collectors; 1 male paratype, Chejel, Guatemala, June,
Schaus and Barnes collectors; 1 male paratype, Cayuga, Guatemala,
April, Schaus and Barnes collectors; 1 female paratype, Mexico, no
date or collector, and 1 female paratype, Mexico City, Mexico, no
date or collector, in the collection of the U.S. National Museum.
One male paratype, Quirigua, Guatemala, September, Schaus and
Barnes collectors, in the Carnegie Museum, Pittsburgh, Pa.

Distribution: This species is known to occur in Mexico, Guatemala,
and Costa Rica.

Remarks: J. lelae, new species, and EF. splendens (Druce) differ in
maculation from EF. extensa (Strand) in that the gray brown sub-
terminal oblique bar of the female is not widest at base and in that
the hindwing of the male is mostly yellow orange. They differ from
the other two species of the complex in that the basal parts of the
ocellate spots of the forewing do not form a sinuous line and in that
the fringe is not uniformly dark. In the male and female genitalia
lelae and splendens agree and differ from the other species of the
splendens complex because the right valve lacks a costal spine of the
sacculus and the sclerotized part of the ductus bursae forms a groove
to the left side.

NO. 3546 MOTHS OF GENUS EUSCEPTIS—TODD 17

This species is named after my mother in recognition of her en-
couragement of my boyhood entomological interests.

Eusceptis extensa (Strand), new combination
Figures 17, 18, 29, 30

Eugraphia extensa Strand, 1913, Arch. Naturg., vol. 79 (A), nos. 7-9, p. 62.
Eugraphia splendens f. extensa Strand.—Draudt, 1939, 1m Seitz, Gross-Schmetter-
linge der Erde, vol. 7, p. 397.

The pattern of maculation of the dorsal surface of the forewing of
this species is like that of splendens except the gray brown oblique
subterminal bar is widest toward the base. The pattern of macula-
tion of the dorsal surface of the hindwing is more like that of E.
paraguayensis (Draudt). The males have the hindwing mostly white
with a narrow marginal band of yellow and a dark brown apical spot.
The hindwing of the female is yellowish orange usually with a heavy
dark brown marginal band, the latter being widest at costa, bent basad
at Ms, usually broken at Cuz and followed by a short recurved spur of
brown scales. The ventral surfaces of the wings of the male are like
those of splendens except the hindwing is mostly white. The markings
of the ventral surface of the forewing of the female are very dark and
usually fused together into a large median and subterminal spot con-
nected to the dark costal marks. The amount of dark scaling present
on both surfaces of the hindwing of females is variable, but more so
on the ventral surface, varying from two moderate spots, one at apex
and one on costa, to a repetition of the broad marginal band usually
occurring on the dorsal surface. Length of forewing: male, 14 mm.;
female, 14 to 15 mm.

Male and female genitalia as illustrated (figs. 29 and 30), inter-
mediate to those of the other two segregates of the splendens complex.
Left clasper of male genitalia decidedly smaller than clasper of right
valve as in splendens and lelae, but costal margin of sacculus of right
valve with a well-developed spine as in robertae and paraguayensis.
Sclerotized groove in ductus bursae of female genitalia short, extend-
ing along ventral surface, not directed to either the left or right side.

Type: A female from Huayabamba, Peru, in the collection of the
K6nigliche Zoologische Museum, Berlin, Germany.

Distribution: Ten examples of this species from the following
localities have been examined. Brazil: Vigosa, Minas Geraes, and
Chapada near Cuyabé, Matto Grosso. Argentina: Remsers, Missiones
and ‘‘Misiones.” Paraguay: Puerto Bertoni. Bolivia: Prov. del
Sara. There are two females of this species from Salto Grande,
Parané and Campinas, Sao Paulo, Brazil in the British Museum
(Natural History). This species appears to be intermediate in dis-

18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

tribution, occupying a range between those of the splendens-lelae and
the paraguayensis-robertae segregates.

Remarks: There are several reasons for questioning the determina-
tion that the species treated is actually extensa. First, the type is
officially unavailable to me and no illustration of it exists. Second,
I have not seen any examples of this genus from Peru and I do not
know of the existence of other museum specimens. Peruvian ento-
mologists were unable to supply examples. Third, there is a question
about the type locality. There are at least three populated places
named Huayabamba in Peru. Two are in the northern part of San
Martin and Cajamarca. The other is in the southern part of Cusco.
In the original description the abbreviation ‘or.’ follows the type lo-
cality. I presume it refers to the Spanish word, oriente, meaning east.
All three Peruvian localities named Huayabamba are on the eastern
side of the Andes, but the one in Cusco is at least 5° east of the other two
and it is much nearer to the known range of the species I am identifying
as extensa in this paper. An examination of the genitalia of the type
would verify whether the name eztensa is correctly applied.

Method of determination: This species has been determined as
extensa by comparison with the original description, especially the re-
marks on the maculation of the hindwing. It is true that the macula-
tion of that wing does vary to some extent in the other species of the
complex and therefore, the character alone is not completely reliable;
but considering our total knowledge of the complex, I believe the name
is correctly applied.

Eusceptis robertae, new species
Figures 9, 10, 31, 32

Structure, vestiture, and coloration of head, thorax, and abdomen as
in splendens. Coloration of legs also similar except base of tibia of
foreleg as dark as apical part, other two pairs of legs with dark areas
reduced, femora nearly completely white, tibia of middle leg with
smaller tufts. Pattern of maculation of forewing differing from that
of splendens in that the basal parts of the two ocellate spots form a
sinuous line, the two basal transverse lines are darker than the broad
costal spot and the oblique subterminal bar, the oblique subterminal
bar usually obsolescent in the middle, and the fringe uniformly dark.
Ventral surface of forewing of male mostly yellowish orange with two
black costal spots, basal part only of oblique subterminal bar usually
present, entirely absent in some specimens. Ventral surface of fore-
wing of female with more dark markings than male, oblique subter-
minal bar divided into two spots or with apex and basal parts darker
than median area. Hindwings of male yellowish orange, lacking dark
marks on both surfaces. Position and shape of dark marks on dorsal

NO. 3546 MOTHS OF GENUS EUSCEPTIS—TODD 19

surface of hindwing of female variable, located between Rs and
anal veins (fig. 10) on two examples, but extending from apex to Cu;
on another example. Ventral surface of hindwing of female yellowish
orange with a dark median costal mark. Length of forewing: male,
13 to 14 mm.; female, 14 to 15 mm.

Male genitalia distinctive (fig. 31), clasper of left valve long and
slender, exceeding apex of valve; clasper of right valve reduced, less
than one-third length of valve. Female genitalia (fig. 32) with the
sclerotized fold and groove on the right side of ductus bursae; ductus
seminalis from right side of bursa copulatrix; right side of posterior
margin of seventh abdominal sternite forming a short, broad lobe,
the shape and size somewhat variable.

Type: Type male, Tucumdn, Argentina, R. Schreiter (USNM
64641); 5 males and 1 female paratypes, same place and collector; 1
male and 1 female paratypes, same place, P. Girard in the U.S.
National Museum, Washington, D.C. Two males and 1 female
paratypes, Tucumdan, Argentina, P. Kohler, in the personal collection
of Mr. Kohler, Buenos Aires, Argentina.

Distribution: Known only from the type locality.

Remarks: The sinuous line formed by the basal parts of the
ocellate spots of the forewing, the long clasper of the left valve of the
male genitalia, and the sclerotized groove of the right side of the
ductus bursae of the female genitalia will enable the entomologist to
separate this species and EL. paraguayensis (Draudt) from the other
species of the splendens complex. The obsolescence of the median
part of the oblique subterminal bar on either or both surfaces of the
forewing, the yellowish-orange hindwing of the male, the absence of
dark apical marks on the ventral surface of the hindwing of the female,
and the reduced clasper of the right valve of the male genitalia dis-
tinguish examples of robertae from paraguayensis. I name this species
with affection after my wife, Roberta.

Eusceptis paraguayensis (Draudt), new combination and new status
Fieures 11, 12, 33, 34

Eugraphia splendens ab. 1, Hampson, 1910, Catalogue of the Lepidoptera Pha-
laenae in the British Museum, vol. 10, p. 793. ([Infrasubspecific, not
available. ]

Eugraphia splendens ab. paraguayensis Strand, 1916, Arch. Naturg., vol. 82 (A),
nos. 1-13, p. 39. [Infrasubspecific, not available.]

Eugraphia splendens f. paraguayensis Strand (= ab. 1 Hampson), Draudt, 1939,
in Seitz, Gross-Schmetterlinge der Erde, vol. 7, p. 397.

Eugraphia splendens paraguayensis Strand.—Biezanko, Ruffinelli, and Carbonell,
1957, Rev. Fac. Agron. Univ. Republ. Uruguay, no. 46, p. 53.

20 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

This species agrees with robertae and differs from the other species
of the splendens complex in that the basal part of the ocellate spots of
the forewing form a sinuous line; in that the fringe of the forewing is
uniformly dark; in that the clasper of the left valve of the male
genitalia is long, exceeding the apex of the valve; and in that the
sclerotized groove of the ductus bursae and the origin of the ductus
seminalis of the female genitalia are located on the right side. £.
paraguayensis (Draudt) differs from robertae as follows: hindwing of
male white with a yellowish orange marginal band; oblique subtermi-
nal bar of forewing not obsolescent in median area on either surface of
the wing; and clasper of right valve (fig. 34) not reduced, exceeding
apex of valve. Length of forewing: male, 12.5 mm.; female, 13 to
15 mm.

Type: A type specimen has not been designated for this species.
The treatments by Strand and Draudt were based on the description
of specimens studied and referred to ‘Ab. 1” of Hugraphia splendens
by Hampson. Hampson did not state how many specimens were
examined, but the descriptive paragraph indicates both sexes and at
least two females were present. ‘Three specimens, one male and two
females, from Sapucay, Paraguay are in the British Museum (Natural
History). These are, without doubt, the series studied by Hampson.
I hereby select the male of that series as the lectotype of the species.

Distribution: This species is recorded from Paraguay southeast
through Misiones, Argentina into southern Brazil and northwestern
Uruguay. I have examined only four specimens from the following
localities—Paraguay: Sapucay. Argentina: Misiones. Brazil: Gua-
rani, Rio Grande do Sul. Biezanko, Ruffinelli, and Carbonell record
the species from Artigas, Uruguay.

Remarks: Hampson recognized that examples of this species
were different from those of splendens, but he treated them as an
unnamed aberration of that species. Subsequently Strand proposed a
name, but continued to consider it to be an aberration of splendens.
According to Articles 1 and 45 (c) of the International Code of
Zoological Nomenclature, 1961, this infrasubspecific usage is to be
excluded from the species-group names and is not available from that
date. Draudt then treated the name as a form of splendens. On
this point, Article 45 (e) (i) states, “Before 1961, the use of either of
the terms ‘‘variety”’ or “form” is not to be interpreted as an express
statement of either subspecific or infrasubspecific rank.” I accept
Draudt’s usage of ‘form’’ in this genus to be equivalent of subspecific
rank and in accordance with Article 10 (b) cite him as author of the
name and consider it available as of that date.

Method of determination: A photograph of a male specimen that
agrees with the original description was compared with the specimen

NO. 3546 MOTHS OF GENUS EUSCEPTIS—TODD Pall

selected as lectotype. Mr. D. 8. Fletcher who made the comparison
for me indicates they are the same.

Name of Undetermined Application

Eugraphia splendens f. seriata

Eugraphia extensa ab. seriata Strand, 1913, Arch. Naturg., vol. 79 (A), nos. 7-9,
p. 63. [Infrasubspecific, not available.]

Eugraphia splendens f. seriata Strand.—Draudt, 1939, in Seitz, Gross-Schmetter-
linge der Erde, vol. 7, p. 397.

This name, like paraguayensis, must be credited to Draudt and date
from that work. The specimen studied by Strand, a female from
Brazil, is in the Kénigliche Zoologische Museum, Berlin, Germany.
The description of the dark markings of the ventral surface of the
hindwing indicates a pattern unlike any example of the splendens
group known to me. It is, however, most likely that the type is
either an aberrant specimen of eztensa or paraguayensis. The
description of the dorsal surface of the hindwing agrees with the pattern
of maculation of those species. An examination of the nature of the
basal part of the ocellate spots of the forewing would indicate to which
seriata is related, but would not necessarily indicate it was either
particular species. There is, of course, the possibility that it repre-
sents a distinct species. Because I have not seen examples that
agree with the description, because large areas of Brazil remain
practically unknown entomologically, and because the type is officially
unavailable to me, I feel obligated to consider seriata as a name of
undetermined application.

Species Transferred to Other Genera
Tarachidia bruchi (Breyer), new combination
Eugraphia bruchi Breyer, 1931, Rev. Soc. Ent. Argentina, vol. 3, p. 309.

This species, the type of which I have examined through the courtesy
of Mr. Pablo Kohler, Buenos Aires, Argentina, is closely related to,
but specifically distinct from, Tarachidia viridans Schaus. It is
not at all closely related to the species of Lusceptis Hiibner. Recent
studies of mine at the British Museum (Natural History) revealed
to me that EL. bruchi Breyer is the same species described by Hampson
as Tarachidia albisecta (1910, Catalogue of the Lepidoptera Phalaenae
in the British Museum, vol. 10, p. 689, pl. 169, fig. 22). The Hamp-
son name, Zarachidia albisecta, is the correct name for the species.

22, PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

/

*% SPLENDENS

A EXTENSA

@ PARAGUAYENSIS
© ROBERTAE

Map 1.—Localities of known occurrence of the South American species of the Eusceptis
splendens complex (Goode Base Map, courtesy Dept. of Geography, University of Chicago).

Proceedings of

the United States

National Museum

SMITHSONIAN INSTITUTION + WASHINGTON, D.C.

Volume 119 1966 Number 3547

REVISION OF NEARCTIC GELECHITDAE, I

THE LITA GROUP
(LEPIDOPTERA: GELECHIOIDEA)

By Ronatp W. Hopces *

Introduction

Busck’s (1903) revision of the North American Gelechiidae is the
only comprehensive treatment of the family for a major part of the
Nearctic Region. Its usefulness is greatly limited by the large
number of species described since 1903, the number of species described
before that date but not correctly determined by Busck, the number of
undescribed species, the inadequacy of descriptions based on macula-
tion and venation for specific determination, and the recognition of
the male and female genitalia as character systems with taxonomic
value.

Subsequent to 1903 numerous gelechiids have been described by
Busck, Braun, Walsingham, Kearfott, Meyrick, Forbes, Keifer, Free-
man, Clarke, and Powell. Of these authors, only the last five have
figured the genitalia of new species. In 1939 Busck published an ex-
tremely important work, ‘Restriction of the genus Gelechia ... ,”’

1 Entomology Research Division, Agriculture Research Service, U.S. Depart-
ment of Agriculture, Washington, D.C.

1

2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

in which he defined several genera and associated species (formerly in
Gelechia) with each genus. Only one or two representatives of each
genus were figured; thus, most of the names, even though correctly
associated generically, are not recognizable as species in that work.
The result is that most of the described Nearctic gelechiids cannot be
determined from the existing literature.

Failure to associate species with earlier names has led nearly all
workers to describe and redescribe several species while leaving
closely related, valid species unnamed. To avoid this situation I
have examined nearly all of the existing type specimens and before
completion of this series of papers plan to study all of them. Some
types, particularly those of Chambers, are no longer extant. An
attempt will be made to associate his names with species whenever
possible.

This paper on the Jita group is the first of a series, the purpose of
which is to make known the described and undescribed Nearctic
gelechiids, to illustrate each species, and to present keys for identifi-
cation of the genera and species. The most useful part of the work
will appear last: the summary of major groups and keys to genera.
However, until the fauna is moderately well known, generic definitions
will continue to change.

Because of their small size, Microlepidoptera, in general, have
been poorly collected; thus, distributional information usually is
inadequate to give a complete picture of the geographic range of any
species. One serious ramification of this point is that clinally variable
character systems may remain unrecognized, and thus two or more
populations may be given names when they represent discontinuous
samples of a species rather than samples of two or more species.
Variation appears to be the rule in the Lepidoptera and may occur in
maculation, coloration, vestiture, size and shape of abdominal sclerites,
width of antennal segments, or characters of the genitalia. However,
suspected or anticipated variation cannot be confirmed on the basis of
few specimens from widely scattered localities. Until much more
diligent collecting is done for nearly all major Nearctic locality types,
works of this nature are necessarily sketchy and subject to further
revision.

As would be anticipated for a relatively large family, the larval
habits are moderately diverse; but on the whole, living plant tissues
are attacked. Some species are leaf or needle miners for all or part
of the larval stage; many are leaf rollers or tiers; some are stem borers
or gall formers; some feed in developing seed heads of composites;
some are external feeders; and some feed on dried seeds. Normally,
some shelter is present, whether it be a silk webbing, a gall, or a
leaf roll. A few species, such as the pink bollworm [Pectinophora

NO. 3547 NEARCTIC GELECHIIDAE—HODGES 3

gossypiella (Saunders)], the Angoumois grain moth [Sttotroga cerealella
(Olivier)], the potato tuberworm [Phthorimaea operculella (Zeller),
and the lodgepole needle miner [Coleotechnites milleri (Busck)], are
economically important. Crasimorpha infuscata Hodges is being
studied for potential use as a biological control agent of a weed in
Hawaii. The food plants and life histories of most species are un-
known, and their study offers a challenge to anyone interested in
rearing insects.

To insure relatively uniform handling and observation of specimens
for study purposes, I have included the following comments: (1)
Adult moths should be mounted and spread, and care should be taken
so that the maculation is preserved and all parts are present. (2)
Genitalic preparations should be made from the best specimens avail-
able so that later maculational comparisons can easily be made.
(3) Age of the specimen must be considered when colors are observed.
Several colors change over a period of time; thus, specimens that
appear dark black or brown when relatively fresh become rather
uniform yellow brown upon aging. Many gelechiids have a light
dusting of blue green when they are alive; this color usually disappears
within afew days after death. (4) The source of light available when
the specimens are observed and the angle of light incidence on the
scale surfaces will affect the appearance of colors. In this study the
colors were seen under incandescent illumination (GE transparent
bulb in an AO illuminator). (5) Maculation of most species is
variable to some degree, and comparisons with illustrations and
descriptions must be made with this fact in mind. (6) Both the
male and the female genitalia can vary; so, several genitalic prepara-
tions may be necessary to learn the range of variation. (7) In many
genera (e.g., Recurvaria Haworth, Coleotechnites Chambers, Lita
Treitschke) the tegumen and vinculum should be opened so that they
are approximately in the same plane rather than being in parallel
planes. In most illustrations a ventral view of the male genitalia is
shown to present characters of the juxta, gnathos, and uncus; however,
a ventrolateral or lateral view of some species is also necessary to show
the shape of the valvae. (8) Species of some genera, particularly
Dichomeris Hiibner, have the female frenulum fused so that the
position of the retinaculum (medial rather than subcostal) or presence
of the ovipositor must be observed to verify the sex. (9) Terminology
of male genitalic structures, particularly of the valva, juxta, and
vinculum, is apt to be somewhat misapplied because of fused parts.
In some instances the valvae and vinculum appear to flow into one
another rather than being separate. Reference to the illustrations
should clarify any questions concerning the name used for a particular
part.

4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

In the systematic part of this paper I describe the maculation and
color pattern of an individual specimen which usually is the one, or
one of those, figured. Label data are presented verbatim with the
exception that dates have been converted to a standard form.

The following individuals have been extremely helpful in the loan
or donation of material, rearing of specimens, and sharing of informa-
tion: J. D. Bradley, Annette F. Braun, J. F. G. Clarke, H. Clench,
P. J. Darlington, D. R. Davis, E. A. Dickason, W. D. Duckworth,
H. E. Evans, D. C. Ferguson, O. S. Flint, Jr., J. G. Franclemont,
T. N. Freeman, M. O. Glenn, H. Grant, R. C. Hall, H. J. Hannemann,
Lars Hedstrém, C. P. Kimball, A. B. Klots, C. D. MacNeill, L. M.
Martin, J. A. Powell, F. H. Rindge, K. Sattler, P. J. Spangler, G. A.
Struble, E. L. Todd, W. G. Tremewan, P. Viette, and A. K. Wyatt.

Abbreviations used for location of specimens are as follows:

ANSP Academy of Natural Sciences, Philadelphia, Pennsylvania.

AMNH American Museum of Natural History, New York, New York.

AFB Collection of Annette F. Braun, Cincinnati, Ohio.

BMNH British Museum (Natural History), London, England.

CAS California Academy of Sciences, San Francisco, California.

CNC Canadian National Collection, Ottawa, Canada.

CM Carnegie Museum, Pittsburgh, Pennsylvania.

CPK — Collection of Charles P. Kimball, West Barnstable, Massa-
chusetts.

CNHM Chicago Natural History Museum, Chicago, Illinois.

CU Cornell University, Ithaca, New York

JGF Collection of John G. Franclemont, Ithaca, New York.

LACM Los Angeles County Museum, Los Angeles, California.

MOG _ Collection of Murray O. Glenn, Henry, Illinois.

MCZ Museum of Comparative Zoology, Cambridge, Massa-
chusetts.

NSMS Nova Scotia Museum of Science, Halifax, Nova Scotia.

USNM United States National Museum, Washington, D.C.

UCB University of California, Berkeley, California.

Mr. J. Scott, staff photographer, Smithsonian Institution, made the
photographs used in the illustrations.

Work on this revision was started while I was a postdoctoral fellow
with the National Science Foundation.

Taxonomic TREATMENT.—Suprageneric taxa are present in the
Gelechiidae; however, their definition and relationships are in part
connected with comparable units in the Gelechioidea. On the basis
of material seen to date the Nearctic genera can be segregated into
five groups; and for convenience the well-known genera, Jsophrictis
Meyrick, Recurvaria, Gelechia Hiibner, Dichomeris, and Anacampsis

NO. 3547 NEARCTIC GELECHIIDAE—HODGES 5

Curtis, are used to exemplify them. I am not certain what status
should be assigned to these groups. For example, the Jsophrictis
group is as distinct from the other four as the timyrids are from the
five; in other words, the two are equal in rank. Thus, I am inclined
to treat each as a subfamily or tribe of the Gelechiidae. This problem
becomes even more complex when other units are studied, e.g., the
Xylorictidae, Copromorphidae, Oecophoridae, Momphidae, Sym-
mocidae. After studying the venational and genitalic illustrations in
Clarke’s (1955, 1963, and 1965) work on the Meyrick types, the
interrelationships among these so-called families become apparent— or
put in another manner—the means of distinguishing one from another
appear to be lacking. Several genera of xylorictids have oecophorid-
type genitalia. The male genitalia of Jsophrictis are more closely
related to those of oecophorids than to the Gelechia type, but the wing
shape and venation ally them with the gelechiids. In the same manner
“Hypatuma”’ zesticopa Meyrick (Gelechiidae) is close to Meleonoma
stomata (Meyrick) (Oecophoridae). The question then must be asked:
What criteria are valid for establishing relationships? Obviously, our
present system does not truly reflect a natural system; but at this
time I am in no position to rally adequate evidence to form the basis
of a sound system. I do feel that we should be very cautious about
proposing new taxa of family rank because this practice eventually
would lead to a system wherein many small groups would be recognized
but no interrelationships shown.

The Inta group, a member of the larger taxon exemplified by
Gelechia, is closely related to Gelechia and Gnorimoschema Busck, and
is arbitrarily separated from them by the presence of a terminal row
(or rows) of caudally directed, modified setae on the uncus. In Lita,
Arla Clarke, Neodactylota Busck, and Eudactylota Walsingham, these
setae are scalariform; in Friseria Busck, Sriferia, new genus, Rifseria,
new genus, Schizovalva Janse, Parapsectris Meyrick, Araeovalva Janse,
and Leuronoma Meyrick, these setae are stout and usually long.
Neofriseria Sattler is closely related to the Lita group, particularly in
the structure of the valvae; but because the uncus lacks the terminal
setae, it is excluded. A feature, apparently common to all of these
genera and to Gelechia and Gnorimoschema, is the culcitula (new term
proposed for the membranous pillowlike base of the gnathos); how-
ever, because I know the South African genera only through Janse’s
(1949-1964) diagnoses and illustrations, I cannot be certain whether
this structure is always present.

The geographic distribution of the Lita group may be summarized
as follows: Lita, 1 Holarctic, 1 Palearctic, and 20 Nearctic species;
Arla, Neodactylota, Eudactylota, Sriferia, Rifseria, and Friseria are
Nearctic;and Parapsectris, Araeovalva, Leuronoma, and Schizovalva are

6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Ethiopian (southern Africa). The South African genera are closely
allied with Friserra. Sattler (1960) has shown that none of the
Palearctic species of the Gelechia complex are referable to the Lita
group with the exception of two species of Lita. Also, no Indo-
Australian or Neotropical gelechiid examined to date is a member of
the Lita group.

Key to the Genera of the Lita Group

1. Valva consisting of asimple lobe... . . gies, 0n a eee
Valva consisting of a compound lobe or two or more Miobes Se eee
2. Valva very broad at base, apex narrow. ... . . . . Araeovalva Janse
Valvalinearyshichtlytexpanded!apically, “29a... Sasi eal ee ae ee
3. Lobes of juxta separate (figs. 100, 103) . . . . . Eudactylota Walsingham

Lobes of juxta connected (at least by membrane) nearly to apex (fig. 66).
Lita Treitschke

4. Aedeagus linear, length more than 10 times maximum width .. Arla Clarke
Aedesgus'‘stout'or with expanded base 4 (2-55 )' RRP eg) a a

5. Caudal setae of uncus sealariform ..... .. . . Neodactylota Busck
Caudal setae of uncus not flattened . . 4 36

6. Valva consisting of two pag lobes, Baer ieee a Somewhat pape
distallyaessers ee he Ne hat ee 7,
Valva consisting of aa or more lobes epee comptes or oben lobe very
broad basally ... . s AUASIAIE EAS

7. Aedeagus broad basally, very eleaden on distil fireestanmtne: costal lobe of
valva shorter than medial lobe... . srl cey es estseniat new genus
Aedeagus stout, narrowest beyond rain apex Some expanded; costal
lobe of valva longer than medial lobe . ... . . . Sriferia, new genus

8. Costal lobe of valva slender, simple; saccal lobe very broad basally, becoming
narrow, then relatively slender to apex . . . .. . . Schizovalva Janse
Costal lobe of valva with medial or basal projections .........49

9. Uncus emarginate medially ........ .. .. . . Leuronoma Meyrick
Uncus even or produced medially .. . . sky LATS. lO

10. Hook of gnathos curved before one half, ener str pene pad eeedaelly tapering
to apex... . . . Parapsectris Meyrick

Hook of gnathos pared ae or ibenoee Paiduile: abruptly turned just before
apextieis.o cote clitds.. Wm. te! PATE) 2 Serre By. aw riseriaybusck

Lita Treitschke
Lita Treitschke, 1833, Die Schmetterlinge Europas, vol. 9 (pt. 2), p. 76.

Type-species: Tinea wirgella Thunberg, 1794 (=Inta zebrella
Treitschke, 1833), designated by Walsingham, 1915, 7 Godman and
Salvin, Biologia Centrali-Americana, vol. 42 (Lepidoptera—Heterocera,
vol. 4), p. 410.

Head: smooth scaled; frontovertical processes present or absent;
tongue moderate, scaled on basal one-third to one-half; labial palpus
recurved, second and third segments slender, subequal in length, a
short tuft on anterior surface of second segment; antenna two-thirds

NO. 3547 NEARCTIC GELECHIIDAE—HODGES a

to four-fifths length of forewing, simple. Forewing: broadly lanceo-
late; 12 veins present; 2 from cell at three-fourths length of cell; 3 and 4
distant to short stalked; 7 and 8 stalked. Hindwing: subquadrate,
outer margin gradually moving to apex; 8 veins present; 3 and 4
connate or stalked; 5 closer to 4 than to 6 basally, curved upward
from origin; 6 and 7 separate. Male genitalia: vinculum moderately
broad; saccus developed; juxta fused basally with vinculum, lobes
setate apically; valva linear, simple, slightly expanded apically;
aedeagus moderately slender to stout, a linear, heavily sclerotized
flange often present; manica with numerous short spicules near
juncture with aedeagus; tegumen long; uncus broad, short with an
apical row of broad scales; gnathos present, a strong hook; culcitula
present. Female genitalia: corpus bursae membranous with numerous
spinules; two signa present; an incomplete, sclerotized ring between
ostium bursae and inception of ductus seminalis; ostium bursae a
simple opening at base of eighth sternum, no associated sclerotized
plates present; apophyses anteriores shorter than apophyses
posteriores.

Lita species form an extremely compact group based upon the sum
of all examined characters. Two basic wing patterns occur: one with a
series of lines and/or dashes paralleling the veins, the other with
spots, sometimes forming bands, at the middle and apex of the cell and
occasionally with an oblique fascia at the base of the forewing. One
species has both types within its range of variation. Frontal modifica-
tions in the form of raised prominences occur in eight species. When
fully developed, three elements are present, a dorsal protuberance
rising from the vertex, a medial protuberance rising from the fronto-
clypeus, and a ventral, low-margined, elliptical depression. The
dorsal and medial protuberances become partially or wholly fused in
some species. The male genitalia are nearly homogeneous among the
species: aedeagal differences serve to distinguish groups of species.
The female genitalia are also relatively uniform; however, the relative
degree of sclerotization and curvature of the apophyses anteriores and
the shape of the signa show minor differences. In variabilis (Busck)
variation in the relative length of the ductus and corpus bursae occurs;
thus, variation may be anticipated in other species. For males the
sternum and tergum of the eighth abdominal segment differ in the
shape of the anterior and posterior margins. Specific variation occurs,
but many species can be defined by these characters. The relative
width of antennal segments, particularly in males, and the width
and shape of the sensory areas are diagnostic characters for some
species. In others both characters seem to vary.

The group of species with a striate wing pattern forms, in part, a
most confusing complex for satisfactory morphological analysis.

8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Unfortunately, the food plant is known for only three species of the
complex, and in most instances few representatives of any one species
are present in collections. Moreover, most specimens are from
scattered localities; moderate series are available for only two species;
and, in each of these two series variation seems to be relatively limited.

The species involved are jubata, new species, nefrens, new species,
princeps, recens, new species, rectistrigella, sironae, new species, thaliae,
new species, and variabilis. Within the series of variabilis in the
USNM collection are specimens with the same maculation as rectistri-
gella, but which were not so identified. Also some of the specimens
that were in the determined series of rectistrigella have been referred
elsewhere. ‘This is not to say that some species of the variabilis group
do not have constant maculational differences. L. princeps, recens,
and sironae usually can be identified on the basis of maculation alone.

Representatives of some populations, here designated as species,
may prove to be races of variable species; however, the morphology
of a limited number of specimens leads me to segregate several entities.
Busck (1939) resurrected Treitschke’s genus Lita and defined it using
characters of the male and female genitalia. He included ten species
of which diversella (Busck) and prorepta (Meyrick) are referable to
Arla and Sriferia, new genus, respectively. Lita crocipunctella
Walsingham is clearly a species of Chionodes and so is transferred.
Thirteen new species are described in this paper, and princeps (Busck)
is transferred from Gnorimoschema Busck; thus, the known world
fauna consists of 22 species. Lita solutella (Zeller) is Palearctic,
virgella (Thunberg) is Holarctic, and the remaining species are Ne-
arctic, mainly western.

Key to Species of Lita

1. Frons and vertex produced ..... ASS TTS 2 og sees Sarees
Frons and vertex smooth, not produced <Scyreie i: tO

2. Distal portion of dorsal frontal protuberance with nacre elliptical cross
section (fig. 64)... 2). . . . . .jubata, new species
Distal portion of dorsal font Droriberance with broadly elliptical or
circular cross section ... . oA eee

3. Forewing pattern a series of jones nelencal ines (Ge 33, 34).
nefrens, new species
Forewing pattern with transverse or oblique elements (figs. 8,15) ... 4
4. Forewing with a dorsal, transverse band (or two spots anise to form
band) at one-third (figs. 15,17) . . 2. . Lecter Rc tae EO
Forewing lacking a dorsal, transverse band at one- fade SS eee ee
5. Dorsal element of frontal protuberance tapering to apex; ventral element
raised and partially fused with dorsal one (fig. 65) . . puertella (Busck)
Dorsal element of frontal protuberance slightly expanded apically; ventral
element low (fig. 59)... . . . . . . . deoia, new species
6. Margin of dorsal element of frontal er Ornternes heavily sclerotized (figs.
GL, C2) NS eS SER eeree | eee eee me cerments)

NO.

N

10.

bts

12.

13.

14.

15.

16.

Le

18.

i)

3547 NEARCTIC GELECHIIDAE—HODGES 9

Margin of dorsal element of frontal protuberance not sharply defined (fig.
58); forewing with a faint oblique band running from costa at one-sixth
to dorsum at one-fourth (fig. 9) ... . . . imcicur, new species

Forewing with costa pale buff, strongly Corice with dorsum (fig. 35);
dorsal element of frontal protuberance broad to apex (fig. 59).

invariabilis (Kearfott)

Forewing with costa generally concolorous with dorsum (fig. 14); dorsal
element of frontal protuberance tapering to apex (fig. 62) ..... . 8

Dorsal element of frontal protuberance longest on dorsal margin, ventral
portion open to ventral element (fig. 62) . .. . . geniata, new species

Dorsal element of frontal protuberance longest on ventral margin, ventral
margin present (fig. 61). .... . . . . . barnesiella (Busck)

Forewing pattern a series of lines ariel ic veins) (fig. 24) 0 ons. oer 4 olS

Forewing pattern a series of dots and/or transverse fasciae (figs. 1, 20). . 10

A transverse bar (formed by coalescing of two spots) in cell at one-third
length of ae (fig. 18), remainder of wing without prominent mark-

AIRY. \) 31-7 . . . . . dialis, new species
Forewing eee a bar a eaetaed if en other dark marks promi-
Wveiva Woe apc tea cgustaree LUE
Forewing with an ohlique Bi running ero eect near hare to dorsum at
one-fourth (fig. 1) ... . . . . Virgella (Thunberg)
Forewing lacking such a complete ancl Geensionnliy. with a band from
middle of wing to dorsum .. . shy as es ete GND,
Forewing with a faint dark patch baadliy: iter third then te veins dark
(fig. 19) . : . . . . pagella, new species
Forewing Eo basal fare patch, eee third with scales on veins con-
colorous with those on Membrane. ......... esha et LO
PRICAECHICISUGCICS ef en hace tes Cuheee as wend oe ol aa polatelis (Zeller)
Nearctic species. . . . : Nei arene
Forewing with a faint eNiinae ine fon ete rane near DOLE) at base to
middle of wing at one-fifth (fig. 26) . . . . . thaliae, new species, in part
Forewing lacking basal oblique line . . . sale Mai tele

Forewing usually with three well-defined dark spate in cell (fig. 28).
texanella (Chambers)

Forewing lacking distinct spots in cell (fig. 22) ........... 16
Antenna unicolorous cream to cream white basally ......... 17
Antenna brown and white basally. ... .. . . maenadis, new species

Inner surface of second segment of labial palpus gray buff with at least a
few brown or orange-brown scales; apex of valva globose.
obnubila, new species
Inner surface of second eran of labial palpus white; apex of valva ellip-
BOG Acme oy owns : : ; . .. . . Veledae, new species
Forewing with black ane in sail ponfivent i ak strongly contrasting
with remainder of wing (fig.31).... . . . . princeps (Busck)
Forewing with two separate black dashes in ceul or none present ... 19
Male antenna with sensory areas extending to dorsal surface of basal seg-
ments; female antenna with ventral sensory area bordered anteriorly with
dark brown (darker than scales separating sensory areas).
recens, new species
Male antenna with sensory areas not extending to dorsal surface, rows of
scales separating sensory areas ocherous; female antenna with unicolorous
BRtCriORDOFGGTS = e.atey tele iste pes. ol Wine Ava peltsie-.ipeetths 40. ¢20

10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

20. Vein 3 of forewing from before end of cell, 3 and 4 moderately distant; dorsal
margin tapering gradually to apex, dorsal and costal margins slightly
convex before apex; flange of aedeagus without a well-defined apical area.

21

Vein 3 of forewing usually from end of cell, 3 and 4 connate or short stalked;
dorsal margin slightly angled at end of 2, apex relatively acute; free por-
tion of sclerotized flange of aedeagus with relatively parallel edges, apex
pluntiae. es eee . . . Variabilis (Busck)

21. Smaller species Goceamne fence ores 7.9 mm, range 7.0-8.5 mm);
forewing with a pale yellow to brownish-orange subcostal streak from near
base to one-third, a strong black dash on dorsal margin of cell from near
base of two-fifths and another on costal margin of cell from two-fifths to
three-fifths (fig. 25). .... . . . Sironae, new species

Larger species (forewing length: peene 9.4 mm, range 8.2-11.5 mm);
forewing usually lacking pale subcostal streak, many specimens lack the
black dashes bordering the cell . . . : tis) oon oe

22. Forewing red brown with dark brown to a binch creas (fig. Die

thaliae, new species, in part

Forewing gray brown, buff brown, or brown, usually not strong red brown;
anterior margin of eighth tergum more or less smoothly curved (fig. 24).
rectistrigella (Barnes and Busck)

Lita virgella (Thunberg)

Figures 1, 66, 136, 169, 170

Tinea virgella Thunberg, 1794, Dissertatio entomologica sistens insecta Suecica,
p. 92.

Lita virgella—Walsingham, 1915, in Godman and Salvin, Biologia Centrali-
Americana, vol. 42 (Lepidoptera-Heterocera, vol. 4), p. 410.

Anacampsis longicornis Curtis, 1827, British Ent., vol. 4, p. 189.

Tinea histrionella Geyer, 1832, Samml. Europiischer Schmett., Tineae, pl. 70.

Lita zebrella Treitschke, 1833, Die Schmett. Europa, vol. 9, pt. 2, p. 82.

Gelechia alternella Kearfott, 1908, Journ. New York Ent. Soc., vol. 16, p. 185.

Gelechia petulans Braun, 1925, Canadian Ent., vol. 57, p. 125.

Maculation: as in figure 1. Head: anterior surface of tongue covered
with white and pale brown scales from base to one-third length;
maxillary palpus white, sometimes with dusting of brown, porrect or
curved over base of tongue; first segment of labial palpus white with
ocherous stripe on outer surface, second segment white with dusting of
brown scales, becoming more dense distally, third segment brown with
dusting of white scales; scape of antenna brown heavily dusted with
white, dorsal surface of shaft with segments brown on basal half,
brown mixed with buff on distal half, ventral surface with more buff
scales basally, brown distally; frons with a series of whitish scales from
margin of eye to base of tongue, scales light brown and pale buff
medially; vertex and occiput with scales light to medium brown
apically, paler basally. Thorax: uneven brown. Forewing: dark
brown with varying amounts of white defining pattern, distal third of
wing often suffused with ocherous scales, cilia fuscous, Hindwing:

NO. 3547 NEARCTIC GELECHIIDAE—HODGES 11

membrane and cilia fuscous. Prothoracic leg: coxa brown mixed with
white on mesal surface, white on outer surface; femur white mixed
with brown, mesal surface darker than lateral; tibia brown with white
scales basally and apically; tarsus brown, white on base and apex of
first segment and apex of second. Mesothoracic leg: similar to
prothoracic leg, tibia with more white scales. Metathoracic leg: coxa
and femur white with some brown scales; tibia brown laterally with
dusting of white basally, a white fascia at one-half, white apically,
dorsal tuft white to pale buff; tarsus white with apices of first through
third segments white, base of first segment white. Abdomen: brown
and buff dorsally, mainly white ventrally. Alar expanse: 12.5-17.0
mm. Male genitalia: as in figure 66 (RWH slide 2563). Female
genitalia: as in figure 136 (RWH slide 2555).

Food plant: Calluna, Erica, Vaccinium (Ericaceae). These are
European records. JL. virgella has not been reared in the Nearctic
Region.

Types: vwirgella, lectotype, present designation, male, bearing
following label: lectotype, male, Tinea virgella Thunberg, by R. W.
Hodges, in Thunberg Collection, Uppsala, Sweden; longicornis, lecto-
type, present designation, female, bearing following labels: female
genitalia slide 3586 R. W. Hodges, lectotype, female, Anacampsis
longicornis Curtis, by R. W. Hodges 1965, National Museum of
Victoria, Melbourne, Australia; histrionella, lost; zebrella, Hungarian
National Museum, Budapest, Hungary; alternella, female, USNM;
petulans, male, CNC.

Specimens examined:

UNITED STATES: Atasxa: White Horse, 2 #7, 2 9, May 30, 31, 1916,
USNM. Catirornia: Deer Park Springs, Lake Tahoe, 1 ? , June 24-30, LACM;
Mineral King, Tulare Co., 19, July 8-15, CAS; Mt. Shasta, Siskiyou Co., 6 0,
29, Aug. 2 through Sept. 1, 1871, Walsingham, BMNH; Mt. Shasta, 7000 ft.,
19, July 7-24, LACM; Mt. Shasta, 19, July 16-23, USNM; Panther Meadow,
Mt. Shasta, 37, July 14, 1962, J. Powell, UCB; Ski Bowl, 14 road mi. E Shasta
City, 7800 ft., Siskiyou Co., 2¢, 39, July 23, 1962, Rentz and MacNeill, CAS.
Coxtorapo: Chimney Gulch, Golden, 4 o, Apr. 14, 1908, Oslar, USNM; Denver,
S. Park, 19, USNM; Dream Lake, Rocky Mt. N. P., 1c’, Aug. 1, 1929, A. F.
Braun, AFB; Hall Valley, Arctic Alpine zone, 11500-12500 ft., Park Co., 107,
July 13-15, 1935, A. B. Klots, AMNH; Loveland, 17, July 1891, Smith, BMNH.
Montana: Glacier Nat. Park, 19, July 28, 1928, A. F. Braun, AFB. OREGON:
Red Top Mountain, 6000 ft., Klamath Co., 1¢, 19, July 7, 1955, J. F. G.
Clarke, USNM; Skyline Ridge, Mt. Baker District, 1¢, Aug. 27, 1932, J. F.
Clarke, CU; Strawberry Lake, Malheur Nat. Forest, 7000 ft., 19, July 13, 1955,
J. F. G. Clarke, USNM. Uvran: Mirror Lake, Uintah Mts., Duchesne Co.,
1d, July 12, 1936, Klots, AMNH. Wasuineton: Marten Lake, Whatcom
Co., 19, Aug. 15, 1931, J. F. G. Clarke, USNM; Skyline Ridge, Mt. Baker
Dist., 5c’, 19, Aug. 25, 27, 1932, J. F. G. Clarke, USNM;; Slate Peak, 7200 ft.,
Okanogan Co., 5<, July 28, 1962, J. F. G. Clarke, USNM; Table Mt., Whatcom
Co., 2¢, Aug. 28, 1927, J. F. G. Clarke, USNM. Wyomine: University of

12 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

Wyoming Campus, Snowy Range, 10000-10500 ft., Albany Co., 97, July 17-23,
1935, A. B. Klots, AMNH.

CANADA: AusBerra: Laggan, 40’, 49, July 4, 10, 1925, O. Bryant, MCZ,
USNM; Moraine Lake, 1c, Aug. 6, 1923, J. McDunnough, AFB; Nordegg,
34, 59, June 8-19, 1921, J. McDunnough, CNC; Rocky Mountain House,
29, June 7, 1921, J. McDunnough, CNC; Shovel Pass, Jasper Park, 19, July 1,
1915, CU; Waterton Lakes, 19, June 20, 1923, J. McDunnough, CNC; same
locality, 10,19, May 29, 1922, C.H.Young, CNC. Britisn Cotumszia: Clinton,
19, June 14, 1938, J. K. Jacob, CNC; Duncans, Vancouver Is., 30, 59, May 18
through June, Henham, CNC, LACM, USNM;; Hedley, 107, 19, July 19, 1923,
C. B. Garrett, CNC; Kathleen Mt., Peachland, 2’, May 23, 1936, A. N. Gartrell,
CNC; Mt. Lolo, Kamloops, 2%, May 31 and July 2, 1938, G. S. Walley, CNC;
Paradise, 47, July 14, 1923, W. B. Anderson, CNC; Quamicham Lake, Vancouver
Is., 1c", May 10, 1908, CNC; Royal Oak, 19, May 20, 1917, R. C. Treherne,
CNC; Sahtlam, 5’, May 18, 1925, E. H. Blackmoore, CNC; Wellington, 1,
G. W. Taylor, USNM. Lasrapor: Goose Bay, 19, June 17, 1948, W. W. Judd,
CNC. Manrrosa: Aweme, 1c’, 69, May 15 through 28, N. Criddle, CNC,
USNM; Riding Mt. Pk., 50, 39, June 2-5, 1938, J. MeDunnough, CNC.
New Brunswick: Waweig, 20’, June 6, 1938, T. N. Freeman, CNC. Norrts-
west TerriroriEs: Aklavik, 1, June 27, 1931, Bryant, USNM; Saw Mill Bay,
200, 59, June 12, 1948, D. F. Hardwick, CNC, USNM. Nova Scotia: Mt.
Uniacke, 16, June 8, 1950, D. C. Ferguson (McDunnough slide, Gel. 2s), NSMS;
Bog, Prospect Road, Halifax, 19, June 4, 1954, D. C. Ferguson, NSMS; West
Dover, Halifax, 1, June 16, 1954, D. C. Ferguson, NSMS. Onrario: Gerald-
ton, 19, June 6, 1956, J. C. E. Riotte, AMNH; Smokey Falls, Mattagami River,
1¢, June 13, 1934, G. S. Walley, CNC. Quersec: Mare du Sault, Laurentides
Park, 2550 ft., 19, July 10, 1954, Klots and Rindge, AMNH; Thunder River,
19, June 18, 1930, W. J. Brown, CNC. White River, 1c, June 24, 1907, Knab,
USNM. Yuxon Territory: Rampart House, 1’, May 31, 1951, C. C. Loan,
CNC.

Discussion: The maculation of virgella is highly variable, with the
amount of white and intensity of the dark brown appearing somewhat
different in each specimen. However, as a variable species, the
maculation is still distinct from the other species of Lita. The
pattern of dark brown bars separated by lighter areas is unique in
the genus.

Lita solutella (Zeller)

Fieures 2, 67, 143, 171, 172

Gelechia solutella Zeller, 1839, Isis von Oken, p. 199.

Lita solutella—Busck, 1929, Proc. U.S. Nat. Mus., vol. 86, p. 573.

Gelechia nigrobipunctatella Lucas, 1932, Bull. Soc. Ent. France, vol. 37, p. 168.
Food plant: Sarothamnus (Spartium) scoparius L. and Genista spp.
Types: solutella, BMNH; nigrobipunctatella, male, Muséum

National d’Histoire Naturelle, Paris.

Distribution: Austria, Bulgaria, England, France, Greece, Hungary,

Italy, Poland, Spain, Syria, and Turkey.

Discussion: L. solutella is included in this treatment of the Nearctic
fauna to complete the picture of the genus inasmuch as it is the only

NO. 3647 NEARCTIC GELECHIIDAE—HODGES 13

extralimital species. It is an extremely variable species in maculation ;
some specimens are immaculate; others are well marked as is shown in
figure 2.

Lita variabilis (Busck)
Figures 3-7, 68-70, 146, 151, 173, 174

Gelechia variabilis Busck, 19038, Proc. U.S. Nat. Mus., vol. 25, p. 871.
Lita variabilis——Busck, 1939, Proc. U.S. Nat. Mus., vol. 86, p. 573.

Maculation: as in figures 3-7. Head: base of tongue white medially,
brown laterally and distally; maxillary palpus white to buff white,
folded over base of tongue; labial palpus with first segment pale
brown, second segment white with scattered pale brown scales, third
segment slightly darker than second with more brown scales; scape
of antenna ocherous brown, shaft ocherous brown with more yellow
basally than distally; frons, vertex, and occiput pale buff brown,
apices of scales slightly darker buff brown. Thorax and forewing:
yellow brown, scales with dark medial areas, dark areas of wing black
or dark brown, some red brown streaks present; cilia paler than wing,
first scale row black tipped, outer scale rows pale brown tipped.
Hindwing: fuscous, veins lightly outlined in pale reddish orange,
cilia buff. Prothoracic and mesothoracic legs: coxa and femur
whitish laterally, brown anteromedially, tibia and tarsus brown,
apices slightly pale. Metathoracic leg: coxa and femur whitish with
some brown scales; tibia buff gray, apices of scales pale brown, dorsal
tuft pale buff; tarsus brown, apices of first, second, and third segments
off white. Abdomen: terga ocherous, apices of terga pale, terga 3-8
darker than one and two; first two sterna white, caudal ones gray
brown. Alar expanse: 14.5-22.0 mm. Male genitalia: as in figures
68-70 (RWH slides 2572, 2588, and 2614). Aedeagus relatively
linear; heavily sclerotized flange with broad apical extension at right
angle with aedeagus. Female genitalia: as in figures 146 and 151
(RWH slides 2735 and 2744).

Food plant: Unknown.

Type: Male, USNM.

Specimens examined:

UNITED STATES: Arizona: Fort Valley, 7350 ft., 74% mi. NW Flagstaff,
Coconino Co., 24o¢, 29, Aug. 10-Sept. 6, 1961, R. W. Hodges (RWH slide
2588), CU, USNM; Hart Prairie, 8500 ft., 10 mi. NNW Flagstaff, Coconino
Co., 796, 189, Aug. 11-Sept. 6, 1961, R. W. Hodges (RWH slides 2259, 2584-
7, 2589, 2744, and 2745), CU, USNM; Paradise, Cochise Co., 2o, Oct. 1-7,
USNM;; Prescott, 1, Sept. 24-30, USNM; Redington, 14, USNM; White
Mountains near McNary, Apache Co., 1, Sept. 15-30, 1925, O. C. Poling,
USNM. Catirornia: Azusa, Los Angeles Co., 17, 29, Oct. 12, 1939, Lloyd
Martin, LACM; Barton Flats, San Bernardino Co., 1, Oct. 1, 1945, Sperry,
AMNH; Cedarville, 37, Sept. 14, 1933, Jones (AB slide Oct. 10, 1933), USNM;

14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Ibanpah Mts., San Bernardino Co., 30’, 19, Oct. 5, 1940, Henne and Comstock,
LACM; Jacumba, 407, Sept. 28—-Oct. 1, 1924, Piazza (RWH slide 2917), LACM;
USNM; La Puerta Valley, 8, Sept. (RWH slide 2921), USNM; Lone Min.,
San Francisco Co., 307, Oct. 3-10, 1909, F. X. Williams, CAS; Monachee Mea-
dows, Tulare Co., 8000 ft., 15¢%, 39, Aug. 8-23 (RWH slides 3151, 3157, and
3158), USNM; near Mono Pass, NW Inyo Co., 12000 ft., 1o@, Aug. 7, 1959,
C. D. MacNeill, CAS; Mt. Shasta, Siskiyou Co., 1¢, Aug. 1871, Walsingham,
BMNH;; Mt. Shasta, 19, Aug. 19, 1939, E. C. Johnston, CPK; Mt Shasta City,
Nevada Co., 167, September (RWH slide 2615), USNM;; Siskiyou Co., 1c, Aug.
22, 1958, J. Powell, UCB; Placer Co., 3o°, 89, August-September (RWH slides
2628 and 2920; AB slide Apr. 3, 1931; RWH wing slide 53), AMNH, LACM
MCZ, USNM;; Rock Creek, 1 mi. SW Tom’s Place, Mono Co., 27, Aug. 9, 1961
and Sept. 6, 1960, MacNeill, Rentz, and Lundgren, CAS; Ruby Lake, NW Inyo
Co., 11500 ft., 29, Aug. 13, 1957, MacNeill and Powell, CAS, UCB; Mouth
San Gabriel Canyon, Los Angeles Co., 2c’, Nov. 10, 1945, C. Henne, USNM;
Upper Santa Ana River, San Bernardino Co., 36’, 19, Sept. 7-14, Sperry and
Melander (RWH slide 2922), AMNH, USNM;; Silver Lake, Amador Co., 207,
Aug. 21, 1936, E. C. Johnston (RWH slides 2572, and 3150), CNC, CPK; Twin
Lakes, Alpine Co., 3’, Aug. 21, 1938, E. C. Johnston (RWH slide 2614), CNC,
CPK, USNM. Cotorapo: no locality, 7 @ (RWH slides 3430 and 3431; AB
slide Oct. 16, 1933), LACM, USNM;; no locality, 7000 ft., 1c (RWH slide 3442),
BMNH;; no locality, 7000 ft., 37, August 1891, BMNH; no locality, 5000 ft.,
49, July and August 1891, BMNH; Bear Creek, Morrison, 1<, Oslar, USNM;
Boulder, 1, Cockerell, USNM; Capitol City, Hinsdale Co., 30’, July 25, 26,
1936, Klots, AMNH; Cripple Creek, 167, Sept. 3, 1899, USNM; Denver, 627,
19, Oslar (RWH slide 2924; AB slide Oct. 16, 1933), LACM, USNM; Durango,
14, Sept. 26, 1939, E. C. Johnston (RWH slide 2581), CNC; Estes Park, 10,
Aug. 12, Mrs. Dyar, USNM; Glenwood Springs, 97, 39, August-September
(RWH slide 2923), MCZ, USNM; Morrison, 1<¢’, August 1891, BMNH; Rocky
Mtn. Natl. Park, 107, Aug. 15, 1987, Klots, AMNH; Silverton, 27, Aug. 8-15,
ex larva, USNM; South Park, 2°, Aug. 19, 21, 1905, USNM. Connecticut:
Putnam, Windham Co., 19, Sept. 5-7, 1961, A. B. Klots, AMNH. Massa-
CHUSETTS: Barnstable, 6°, Sept. 5-12, C. P. Kimball (JFGC slide 10167).
Montana: Boulder, 1, 1892, T. Ulke, USNM; Bozeman, 1.7, Aug. 20, 1928,
J. MeDunnough, CNC; Richel Lodge, 1¢, Aug. 20, 1939, Sperry, USNM.
Nevapa: Mt. Magruder, Esmeralda Co., 19, Sept. 19, 1939, G. Willett, LACM.
New Mexico: Hell Canyon, 1<, Sept. 16, 1916, C. Heinrich, USNM; Indian
Spring, 19, Sept. 9, 1916, C. Heinrich (RWH slide 2735), USNM; Mescalero,
7000 ft., 14, Oct. 27, BMNH; Therma, 3¢7, Aug. 12, 1932, AMNH. Uran:
Buckboard Flap Camp, 7 mi. W Monticello, 8800 ft., San Juan Co., 1c, July
28, 1960, Rindge (RWH slide 2882), AMNH. Dividend, 2.7, Sept. 26, 28,
BMNH; Eureka, 87, Aug. 20-27, 1911, Tom Spalding (RWH slide 2918),
LACM, USNM; Stockton, 14, September 1908, BMNH. WasuinetTon:
Kushi Canyon, Yakima Co., 130, 19, Aug. 20-Sept. 17, E. C. Johnston (RWH
slide 2736), CNC; Satus Creek, Yakima Co., 60’, 19, Aug. 19 and Sept. 19,
1949, E. C. Johnston (RWH slides 2632 and 2633), CNC, USNM. Wromine:
Green River L., Wind River Range, 1, July 24-Aug. 7, 1935, A. B. Klots,
AMNH;; Sacajawea Camp, 24 mi. W. Big Piney, 8400 ft., Sublette Co., 20,
Aug. 2, 1959, Rindge, AMNH; Sheepeater Cliffs, Yellowstone N. P., 1%, Aug.
17, 1962, Spangler (RWH slide 2640), USNM.

CANADA: British CotumBia: Kamloops, 1c’, Sept. 1, 1918, BMNH;
Peachland, 2¢’, Aug. 21, 22, 1909, J. B. Wallis, USNM. Manitospa: Aweme,

No. 3547 NEARCTIC GELECHIIDAE—HODGES 15

9%, 29, Aug. 14-29, N. Criddle (RWH slides 2748, 2749, 3434-3436; RWH
wing slide 56), CNC, USNM;; Beulah, 4.7, Aug. 15, (RWH slides 2625 and 2626),
USNM;; Westbourne, 1”, Aug. 26, 1908, J. B. Wallis, USNM. SasxarcHEewan:
Attons Lake, 2c‘, Aug. 20, 1940, A. R. Brooks (RWH slides 2746 and 2747),
CNC, USNM; Earl Grey, 10’, Aug. 2, 1924, J. D. Ritchie (RWH slide 3443),
CNC, Harlan, 2¢’, Aug. 10, 11, 1940, A. R. Brooks (RWH slide 3437), CNC;
Saskatoon, 1o7, Aug. 14, 1940, A. R. Brooks, CNC.

Discussion: The moths illustrated indicate the range of macula-
tional variation to be anticipated in this species. I have seen a
moderate series of specimens from one area; and in it, although
maculational variation is great, the extremes in size and color pattern
are not present. As a guess, it would seem as though the species is
extremely variable but that any given population will have a number
of phenotypes smaller than the total of the species. Variability is not
confined to maculation, color, and size. It is also expressed in the
shape of the apical portion of the heavily sclerotized flange of the
aedeagus in the male genitalia and in the relative length of the
ductus bursae and corpus bursae in the female genitalia.

As might be expected, several species have been confused under
the name variabilis. Most of them are easily distinguished by the
frontal processes, antennal characters, and the shape of the aedeagus.
L. variabilis is the only known species of Lita with the aedeagus
shape as illustrated in figures 68-70.

Lita barnesiella (Busck)

Ficurss 8, 61, 85, 139, 175, 176

Gelechia barnesiella Busck, 1903, Proc. U. S. Nat. Mus., vol. 25, p. 875.
Lita barnesiella.—Busck, 1939, Proc. U. S. Nat. Mus., vol. 86, p. 573.
Maculation: as in figure 8. Head: tongue pale gray brown
basally, becoming darker brown distally; maxillary palpus ascending,
nearly attaining ventral margin of frontal depression, pale gray with
scattered light brown scales; labial palpus pale gray with a few brown
scales on posterior and lateral surfaces of second and third segments,
anterior surface brown with a few pale gray scales, third segment
brown with dusting of pale gray scales; scape of antenna brown buff,
shaft piceous dorsally, ocherous ventrally; frons, vertex, and occiput
pale buff, frontovertical processes as in figure 61. Thorax, forewing,
and abdomen, pale buff overlaid with dark brown and varying shades
of red brown, cilia of forewing pale buff. Hindwing: pale fuscous,
veins slightly darker fuscous, cilia and tuft of scales on vein 1 pale
buff. Prothoracic leg: coxa brown overlaid with pale buff, apex pale
buff; femur and tibia dark brown with scattered pale buff scales,
buff scales more abundant on outer surfaces; tarsus dark brown,
apices of first, second, and third segments pale buff, ventral surface

16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

of first segment pale buff. Mesothoracic leg: coxa off white on outer
surface; femur and tibia dark brown with some pale buff flecks, a
pale buff fascia just beyond middle of tibia, apex of tibia pale buff;
tarsus dark brown dorsally, apices of first, second, and third segments
pale buff, apex of fourth segment pale buff laterally. Metathoracic
leg: coxa off-white; femur brown, heavily overlaid with pale buff;
tibia pale buff with a few brown scales near base, a broad brown
fascia just before middle and apex, dorsal scale tuft pale buff; tarsus
dark brown externally, base of first segment and apices of all segments
pale buff. Alar expanse: 15-25 mm. Male genitalia: as in figure 85
(RWH slide 2618). Heavily sclerotized flange of aedeagus with
moderately acute apex, ductus ejaculatorius simplex one-and-one-half
times length of aedeagus. Female genitalia: as in figure 139 (RWH
slide 3144).

Food plant: Unknown.

Type: Male, USNM.

Specimens examined:

Arizona: Maricopa, Pinal Co., 10%, Oct. 17, 1927, J. A. Kusche, CAS;
Kingman, 1.7, Oct. 1-7, USNM; Mohave Co., 1<7, Oct. 8-15, USNM; same
locality, 14, BMNH; Paradise, Cochise Co., Oct. 1-7, USNM; 4 mi. ESE Pine,
5400 ft., Gila Co., 27, Sept. 5, 1961, R. W. Hodges, USNM;; Prescott, 8, 89,
Oct. 1-30 (AB slide Aug. 16, 1933), USNM. Catirornia: Loma Linda, 19,
Noy. 6, G. R. Pilate, AFB; Southern Calif., 19 (RWH slide 2619), USNM;
Willow Creek, Siskiyou Co., 14<7, Sept. 10-11, 1871, Walsingham (RWH « slide
2879), BMNH, USNM. Cotorapo: Denver, 607, Sept. 15, 1910, Oslar,
USNM;; same locality, 14, 49 (AB slide Aug. 11, 1933), USNM; Durango, 12,
19, Oslar, CAS, USNM; Glenwood Springs, 57, 19, July 16-September (AB
slides Aug. 4, 1933 and Apr. 30, 1937), USNM; same locality, 3, Aug. 2, 22,
1892, W. Barnes, USNM; same locality, 2°, September 1906, BMNH, LACM;
Lamar, 407, 19, Sept. 24, 1945, E. C. Johnston (RWH slide 2573), CNC, USNM;
Larimer Co., 20’, 19, June, 407, July, 2007, 29, August 1891, BMNH; Mesa-
verde, 167, Sept. 27, 1945, E. C. Johnston (RWH slide 2618), CNC; Rock Creek
Canyon, 307, Sept. 26-28, 1957, Margot May, CPK; no further locality, 110,
BMNH, LACM, USNM. New Mexico: Bent, 207, 19, 10.27, BMNH; Gallup,
1, Sept. 11, 1961, R. W. Hodges, USNM; Jemez Springs, 167, Oct. 1-7, USNM;
Mescalero, 167, 10.27, BMNH. Texas: Bosque Co., 1c’, Oct. 6, 1876, Belfrage,
BMNH. Uraun: Eureka, 207, Sept. 6, 9, 1910, T. Spalding, USNM; Stockton,
33,19, Aug. 31-Sept. 10, T. Spalding (AB slides Aug. 8, 1933 and Apr. 29, 1937),
USNM. Wyomrne: Torrington, 1<, Sept. 2, 1948, R. E. Pfadt (JFGC slide
9896), USNM.

Discussion: The maculation of barnesiella is relatively constant.
The major variation is in the amount of dark brown and red brown on
the forewing and thorax. Some specimens are very pale with the
basal and antemedial triangular dark markings as the prominent
features; others are moderately dark and have a general suffusion of
dark brown on the forewings.

The frontal processes (fig. 61) are the major diagnostic feature of

NO. 3547 NEARCTIC GELECHIIDAE—HODGES 17

barnesiella. Of the four species that might be confused with it by
maculation, tezanella, pagella, incicur, and geniata, the first two have
a plain frons. JL. incicur (fig. 58) has the interior of the dorsal pro-
jection spinose, that of barnesiella is smooth. The dorsal and medial
frontal processes of geniata (fig. 62) lack separating margins; those of
barnesiella are separated by the strong ventral margin of the dorsal
process.
Lita incicur, new species

Figures 9-13, 58, 78, 137, 140, 177, 178

Maculation: as in figures 9-13. Head: tongue white to pale buff
basally, becoming brown subbasally; maxillary palpus off white,
folded over base of tongue; labial palpus, first segment with yellow
brown and pale brown scales, second segment white with scattered
yellow and pale brown scales, third segment with a mixture of white
and brown scales; scape of antenna buff with scattered brown scales
on dorsal surface, shaft greasy brown, some scales of distal half of
basal segments greasy buff; frontovertical processes as in figure 58;
frons buff white; vertex and occiput buff. Thorax; varying shades of
buff and buff brown. Forewing: pale buff to orange; dark markings
brown; cilia pale buff, apices of scales darker. Hindwing; shining
fuscous, veins with orange brown, cilia buff. Prothoracic leg: white
on posterolateral surface, pale brown with whitish scales on fronto-
medial surface, apex pale buff; femur brown with whitish scales;
tibia brown with a white fascia slightly beyond middle and another
at apex; tarsus brown, apices of first three and base of first segment
white. Mesothoracic leg: nearly as for prothoracic leg. Meta-
thoracic leg: tibia white with an antemedial and preapical patch of
brown scales. Abdomen: terga pale yellow basally, gray brown
distally, apices of sterna buff white. Alarexpanse: 17-27mm. Male
genitalia: as in figure 78 (RWH slide 2570). Apex of flange of
aedeagus sickle shaped. Female genitalia: as in figures 137 and 140
(RWH slides 2616 and 2751).

Food plant: Unknown.

Type: Male, Smokey Valley, 6300 ft., Tulare Co., California,
Sept. 28, 1946, C. Henne (RWH slide 2610), USNM type 67643.

Paratypes:

CALIFORNIA: Mammoth Lake, Inyo Co., 16, Aug. 15, 1921, O. C. Poling,
USNM; Mono Pass, Inyo Co., 40, Aug. 13, 1957, J. Powell (RWH slides 2611
and 2612), UCB, USNM; Monachee Meadows, 8000 ft., Tulare Co., 207, Aug.
16-23, 1936, USNM;; Mt. Shasta, Siskiyou Co., 127, 13 9, August 1871, Walsing-
ham, BMNH, USNM;; same locality, 70, 59, Aug. 19, 1939, E. C. Johnston
(RWH slide 2571), CNC, USNM; Nevada Co., 27, September, CAS, USNM;
same locality as type, 2c’, CU, USNM; Tuolumne Meadows, Tuolumne Co.,
1o, Aug. 27, 1960, W. E. Ferguson, UCB; Twin Lakes, Alpine Co., 407, Aug. 21,

219-945—66——2

18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

1936, E. C. Johnston (RWH slide 2570), CNC, USNM; White Mt. Peak, 12500
ft., 5 mi. S, Mono Co., 16’, 19, Aug. 15, 1960, S. F. Cook (RWH slides 2559
and 2560), UCB, USNM; Willow Creek, Siskiyou Co., 8, 39, Sept. 10-11,
1871, Walsingham (RWH slides 2877, 2878, 2880, and 3019), BMNH, USNM.
Cotorapo: Glenwood Springs, 407, Aug. 16-18, 1892, W. Barnes (AB slide Aug.
12, 1933; RWH slide 2609), USNM. Orgaon: Crater Lake Park, South Rim,
7100 ft., 1o’, Sept. 13, 1930, H. A. Scullen, CU. Wyomine: Green River L.,
Wind River Range, 10, July 24-Aug. 7, 1935, A. B. Klots, AMNH; Sacajawea
Camp, Middle Piney Creek, 8400 ft., Sublette Co., 1, Aug. 14, 1953, Rindge
(RWH slide 2883), AMNH; Yellowstone Natl. Park, Madison Jet., 1d, 29;
Aug. 19, 1962, Spangler (RWH slides 2606 and 2616), USNM.

Other specimens examined:

Ca.LiFrorNiA: Mono Pass, Inyo Co., 1¢, Aug. 13, 1957, J. Powell, UCB, same
locality, 12000 ft., 10c°, 49, MacNeill, and MacNeill, Rentz, and Landgren,
CAS; Rock Creek, 1 mi. W. Tom’s Plane, Mono Co., 1%, 39, Sept. 6, 1960,
MacNeill, CAS.

Discussion: LZ. incicur is similar to texanella and obnubila but
differs from them in having frontovertical processes.

Lita geniata, new species
Figures 14, 62, 81, 179, 180

Maculation: as in figure 14. Head: tongue and maxillary palpus
pale buff white, latter folded over base of tongue; labial palpus nearly
white, outer surface of first segment and base of second segment with
a few yellow-brown scales, scattered pale brown scales apically on
second segment and generally on third segment; scape of antenna
buff, shaft greasy brown (male, no females seen); scales of frons,
vertex, and occiput buff; frontovertical processes as in figure 62.
Thorax: varying shades of brown. Forewing: varying shades of
brown; darker basally, becoming buff white apically, spots on dise
dark brown, almost black; apices of cilial scales darker than remainder
of scale. Hindwing: fuscous buff, veins with more yellow orange;
cilia pale buff. Legs: buff to off white basally, becoming brown
distally; apices of basal tarsal segments off white; metathoracic tibia
with pale buff dorsal tuft and three brown scale patches on outer
surface, one subbasal, one antemedial, and one preapical. Abdomen:
first three terga yellow, remaining terga yellow brown with pale
apices; sterna greasy buff white, bases darker than apices. Alar
expanse: 15.0-19.5 mm. Male genitalia: as in figure 81 (RWH slide
2604). Apex of heavily sclerotized flange of aedeagus narrow, sickle
shaped. Female genitalia: no specimens available.

Food plant: Unknown.

Type: Male, La Puerta Valley, So. Calif., USNM type 67644.

Paratypes:

CALIFORNIA: same data as for type, 167 (RWH slides 2604 and 2605), BMNH,

NO. 3547 NEARCTIC GHLECHIIDAE—HODGES 19

CNC, CU, UCB, USNM; La Puerta, 307, Stephens, USNM; La Puerta Val.,
16, Oct. 16-23, USNM; Loma Linda, 1.7, Oct. 22, G. R. Pilate, AFB; So. Calif.,
16’, USNM.

Discussion: ZL. geniata may be separated from barnesiella as indi-
cated under the latter. The maculation of geniata is relatively
constant with the exception of the costal area which may be brown to
buff. Inasmuch as the known specimens are from one locality (the
“So. Calif.” specimen may represent the same population), the
amount of variation is likely to be small. When other populations
are discovered, more phenotypes may be observed.

Lita puertella (Busck)

Figures 15, 16, 65, 80, 149, 181, 182

Gelechia puertella Busck, 1916, Proc. Ent. Soc. Washington, vol. 18, p. 148.
Lita puertella——Busck, 1939, Proc. U.S. Nat. Mus., vol. 86, p. 573.

Maculation: as in figures 15 and 16. Pale form: head, thorax, and
forewing pale yellowish white; labial palpus with a few pale ocherous
scales scattered on lateral surface; maxillary palpus folded over base
of tongue; shaft of antenna ocherous basally becoming darker distally;
markings on forewing brown; hindwing pale fuscous yellow; cilia pale
buff; abdomen pale ocher dorsally, pale yellowish white ventrally;
legs pale yellowish white, becoming brown distally, prothoracic leg
darker than metathoracic leg, apices of tarsal segments yellowish
white except last two of prothoracic leg and ultimate segment of
mesothoracic leg. Dark form: much as given for pale form but with
overlay of brown and red brown. Lower portion of frontal process
(fig. 65) with strongly carinate margin, inner surface pitted; dorsal por-
tion with a small, circular cross section, margin even and strongly
carinate. Alar expanse: 16-20 mm. Male genitalia: as in figure 80
(RWH slide 2598). Heavily sclerotized flange of aedeagus generally
uniform in width, tapering to acute apex. Female genitalia: as in
figure 149 (RWH slide 3145).

Food plant: Unknown.

Type: Male, USNM.

Specimens examined:

Arizona: Maricopa, Pinal Co., 16, Oct. 17, 1927, J. A. Kusche, CAS; Phoenix,
16, Oct. 29, Cockerell, USNM. Catirornia: Desert Springs, San Bernardino
Co., 47, Oct. 17, 1960, Hurd and Powell (RWH slides 2567 and 2568), UCB,
USNM;; near Essex, San Bernardino Co., 167, 39, Oct. 30, 1939, J. A. Comstock
(RWH slide 3145), LACM, USNM; Ivanpah Mts., San Bernardino Co., 11¢,
Oct. 5, 1940, J. A. Comstock (RWH slide 2597), LACM, USNM; same locality,
4, Oct. 8, 1940, C. Henne, LACM; La Puerta Valley, So. Calif., 740, Sep-
tember through Oct. 23 (AB slides), Sept. 10, 1930, Mar. 2, 1931, and Apr. 28,
1937 (RWH slides 2566, 2598, 2599, and 2600), LACM, USNM; Providence
Mts., 3c’, 22, Oct. 8, 1936, Sperry, AMNH, CNC; Rock Cr., 1 mi. W Tom’s

20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Place, Mono Co., 37, Aug. 9, 1959, C. D. MacNeill, CAS; West Riverside, 10,
Oct. 28, 1905 (AB slide Aug. 1, 1933), USNM.

Discussion: JL. puertella is a highly variable species in regard to
maculation, ranging from pale yellowish-white specimens with two
prominent dark brown spots on the forewings to brown specimens
with the dark brown spots barely discernible. I have seen numerous
intermediate stages between the two extremes.

L. deota and dialis superficially resemble puertella. L. dialis is
easily separated because it lacks a frontal process. The lower section
of the frontal process of puertella has a strong, high ventral margin;
that of deoia has a rounded, poorly defined ventral margin.

Lita deoia, new species
FiaurE 17, 82, 183, 184

Maculation: as in figure 17. Head: buff white, maxillary palpus
folded over base of tongue; outer surface of first segment of labial
palpus pale brown, apex of third segment brown; scape of antenna
with a dorsomedial brown patch, shaft greasy brown; frontovertical
protuberance much as for invariabilis (fig. 59), dorsal protuberance
more nearly circular, ventral margin slightly flattened. Thorax:
buff white. Forewing: buff white, tinged with pale yellow and brown,
dark markings brown. Hindwing: shining pale fuscous, veins
slightly darker, cilia buff. Prothoracic leg: coxa and femur white on
outer surface, inner surface of coxa white with scattered brown scales,
inner surface of femur brown with scattered white scales; tibia brown
with a postmedial and apical white band; base and apex of basitarsus
and apex of second tarsal segment white. Mesothoracic leg: as for
prothoracic leg but with more white scales on inner surface of femur,
inner surface of tibia white. Alar expanse: 20.6mm. Male genitalia:
as in figure 82 (RWH slide 2556). Apex of flange of aedeagus broad
and blunt, much as for variabilis. Female genitalia: no specimens
available.

Food plant: Unknown.

Type: Male, Smokey Valley, 6000 ft., Tulare Co., Calif., Sept. 28,
1946, C. Henne (RWH slide 2556), USNM type 67645.

Discussion: Superficially, deova looks like the light form of puertella
or dialis. It differs from dialis in having frontovertical processes.
It may be distinguished from puertella (fig. 65) by the ventral margin
of the lower frontal processes being poorly defined; in puertella this
margin is raised and heavily sclerotized. The frontal process of
deoia is broad, that of pwertella is narrow; the medial process of
puertella is raised more than its apical width, that of deova is raised
less than one-third its apical width.

L. deoia and puertella are very close, and it may be that deova

NO. 3547 NEARCTIC GELECHIIDAE—HODGES il

represents a population of puertella; however, evidence to support
this hypothesis is lacking.

Lita dialis, new species
Figures 18, 83, 132, 185, 186

Maculation: as in figure 18. Head: tongue buff; maxillary palpus
pale buff, folded over base of tongue; labial palpus white, apex of
third segment tinged with yellow brown; scape and base of antenna
buff, shaft greasy buff brown apically; frons, vertex, and occiput pale
buff. Thorax: buff white. Forewing: buff white to buff, dark
spot brown, cilia buff white. Hindwing: shining pale fuscous, veins
darker and tinged with yellow gray, cilia buff. Legs brown with
scale bases pale brown to white; a white fascia at middle and apex of
each tibia, dorsal tuft on metathoracic tibia off white, apices of first
three or four tarsal segments and base of basitarsi white. Abdomen:
terga greasy ocher, paler on basal terga, apices of terga pale; sterna
greasy brown, apices of segments greasy buff, many scales with buff
lines. Alar expanse: 17.5-20.0 mm. Male genitalia: as in figure 83
(J.F.G.C. slide 10208). Apex of flange of aedeagus tapering to blunt
tip. Female genitalia: as in figure 132 (RWH slide 2603).

Food plant: Unknown.

Type: Male, Paradise, Cochise Co., Ariz., March (RWH slide
2602), USNM type 67646.

Paratypes:

Arizona: Southern Arizona, 16’, USNM; Wickenburg, 19, May 20, 1957,
Sperry, USNM. New Mexico: Carlsbad, Eddy Co., 2°, May 17, 1950, E. C.
Johnston (RWH slide 2603), CNC. Trxas: Alpine, 5000-8000 ft., 407, April-
July, AFB; Brewster Co., 5000 ft., 70%, 29, 4.26, BMNH, USNM; no fur-

ther locality, 1, May 19, 1953, O. J. Robertson (JFGC slide 10208), USNM.
Mexico: Durango, La Resolana, 1<, Feb. 21, 1953, E. I. Schlinger, UCB.

Discussion: ZL. dialis is similar in appearance to deoia and puertella
but differs from them in lacking frontovertical processes. Individual
specimens differ in the amount of brown scaling so that the moths
appear pale buff to gray brown.

Lita pagella, new species
FicurE 19, 20, 84, 145, 187, 188

Maculation: as in figures 19 and 20. Head: tongue buff brown
to brown laterally, some pale buff scales medially; maxillary palpus
pale buff, folded over base of tongue; first segment of labial palpus
white on inner surface, outer surface white basally, brown from
middle to apex, pale buff on ventrodistal portion, second segment
buff white to white with scattered brown scales, third segment mainly

22 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

brown on anterior surface, white on posterior surface; scape of antenna
orange brown dorsally, buff white ventrally, shaft greasy brown; frons
buff white; vertex and occiput buff. Thorax: scales brown, buff
brown to pale buff, most scales paler basally than apically. Fore-
wing: orange brown and varying shades brown, cilia orange buff
basally, pale buff distally. Hindwing: fuscous, veins dark, cilia pale
buff. Prothoracic leg: coxa with anterior surface brown, basal row
of scales buff white, posterolateral surface white; femur brown an-
teriorly, mottled brown and white posteriorly; tibia brown, a buff
fascia beyond middle, apex white; tarsus brown, base and apex of
first segment buff white. Mesothoracic leg: as for prothoracic leg,
outer surface of coxa white, apex of tibia buff. Metathoracic leg:
coxa white, outer surface of femur white with scattered brown scales;
tibia greasy brown black, an oblique buff white subbasal fascia,
another at three-fifths, apex white, dorsal tuft off white; tarsus
brown black, base of first segment and apices of all segments pale.
Abdomen: first three terga greasy pale yellow, remaining terga pale
buff; sterna white to buff white. Alar expanse: 17-19 mm. Male
genitalia: as in figure 84 (RWH slide 2608). Apex of flange of
aedeagus acute, free portion nearly at right angle with aedeagus.
Female genitalia: as in figure 145 (RWH slide 2574).

Food plant: Unknown.

Type: Female, Fort Valley, 7350 ft., 74 mi. NW Flagstaff,
Coconino Co., Ariz., Aug. 27, 1961, Ronald W. Hodges (RWH slide
2574), USNM type 67647.

Paratypes:

Arizona: Mohave Co., 19, Sept. 8-16, USNM; Paradise, Cochise Co., 8?
Aug. 24-Oct. 7, USNM; White Mts., 7200-11500 ft., 1, Aug. 10-30, 1925,
O. C. Poling, USNM. Catirornia: Jacumba, 3c, Sept. 27-Oct. 2, 1924,
Piazza (RWH slide 2608), LACM, USNM. Cotorapo: Colorado Springs,
Fountain Valley School, 16, Aug. 20-31, 1932 (ABK slide 18 Jan. 23), AMNH;
Rock Creek Canyon, 1, Sept. 10, 1947, Margot May (RWH slide 2876), CPK;
Salida, 19, Aug. 27, 1938, Sperry (RWH slide 2884), AMNH. New Mexico:
Bent, 3o7%, October 1927 (RWH slide 3020), BMNH, USNM; Mountain Park,
19 August 1927, BMNH.

Discussion: L. pagella may be separated from barnesiella and
incicur by the smooth frons and from thaliae by the entire basal area
of the forewing being dark or with dark spots. Some specimens are
much paler than the one described.

Lita obnubila, new species
FiaursEs 21, 87, 135, 189, 190

Maculation: as in figure 21. Head: tongue pale buff, some scales
brown tipped; maxillary palpus pale buff, folded over base of tongue;

NO. 3547 NEARCTIC GELECHIIDAE—HODGES 23

first segment of labial palpus white on inner surface, buff brown on
outer surface, second and third segments buff white with scattered
brown scales; scape of antenna buff white lightly dusted with brown,
shaft greasy buff basally becoming buff brown distally; frons buff
white; scales of vertex and occiput buff white with pale brown apices.
Thorax: gray buff, scales streaked with gray, many scales with narrow
brown border at apex; last row of scales of tegula buff white. Fore-
wing: white, buff white, pale yellow, red brown, and brown; many
scales of cilia tipped with red brown. Hindwing: fuscous, veins
broadly covered with scales that appear shining orange at some angles
of light incidence, cilia buff. Legs: mottled buff white and brown,
each succeeding pair paler than the preceding one, apices of tarsal
segments and sometimes dorsal surfaces off white. Abdomen:
anterior two terga pale yellow, succeeding ones buff white; sterna
buff white with scattered brown scales, apices unicolorous. Alar
expanse: 19-23 mm. Male genitalia: as in figure 87 (RWH slide
3459). Apex of valva globose. Female genitalia: as in figure 135
(RWH slide 3460).

Food plant: Unknown.

Type: Male, Fort Davis, Tex., 5000 ft., 5.28 (RWH slide 3459),
BMNH.

Paratypes:

Same data as for type, 3c’, 19 (RWH slides 2881 and 3460), BMNH, USNM.

Discussion: Superficially, obnubila appears like some specimens of
texanella; however, the forewings of obnubila lack distinct spots in the
cell; the apices of the valvae are globose, ellipsoidal in texanella; the
two projections on the anterior margin of the eighth sternum (fig. 189)
are uniformly broad to the apex, in teranella these projections taper
somewhat to the apex; the ratio of width of the segment bearing the
apophyses anteriores to the total length from the apex of the papillae
anales to the anterior margin of the bursa copulatrix is 1:5 in obnubila,
1:7 in tezanella. L. obnubila may be separated from veledae by the
apices of the valvae being globose and the inner surface of the second
segment of the labial palpus being mottled.

Lita maenadis, new species
FIGURES 22, 133
Maculation: as in figure 22. Head: tongue white basally becoming
pale brown medially; maxillary palpus white, folded over base of
tongue; inner surface of first segment of labial palpus white, scales of
outer surface pale buff basally, brown distally, second segment white
with scattered brown scales, third segment brownish, bases of scales
nearly white; dorsal surface of scape pale buff with brown blotches,

24 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

ventral surface buff, ventral surface of shaft buff, individual scales of
dorsal surface pale buff basally, dark brown distally, several half
segments pale buff basally; frons nearly white; vertex and occiput
buff white, brown tuft of scales above eye. Thorax: buff white, many
scales with darker streaks. Forewing: buff white to white, several
brown or red-brown tipped scales; first row of scales of cilia
brown tipped, others unicolorous. Hindwing: pale fuscous, veins
darker, cilia pale buff. Prothoracic leg: coxa white with several brown
tipped scales on anterior surface, posterolateral surface white; femur
and tibia brown, bases of scales pale buff, apex of tibia white; tarsus
brown, base of first segment and apices of first and second segments
white. Mesothoracic leg: much as for prothoracic leg but somewhat
lighter, outer surface of coxa white, a white streak just beyond middle
of tibia. Metathoracic leg: as for mesothoracic leg but lighter, dorsal
tuft on tibia buff white. Alar expanse: 18-20 mm. Male genitalia:
no specimens available. Female genitalia: as in figure 133 (RWH
slide 2583).

Food plant: Senecio species.

Type: Female, Blanco’s Corral, White Mts., Mono Co., Calif.,
10150 ft., Aug. 25, 1960, P. D. Hurd, Senecio (RWH slide 2583),
CAS.

Paratype:

Placer Co., Calif., 19, August, Koebele, CAS.

Discussion: L. maenadis is nearest to obnubila but may be separated
by the antenna being multicolored basally; in obnubila the antenna is
unicolorous off white basally.

Lita veledae, new species
Fiagure#s 23, 88, 191, 192

Maculation: as in figure 23. Head: tongue pale buff to buff white;
maxillary palpus buff white, somewhat curved over base of tongue;
first segment of labial palpus white on inner surface, white with a few
orange scales on outer surface; second segment white on inner surface,
white with scattered orange scales on outer surface, third segment
mainly white but with numerous orange and pale brown scales anteri-
orly; scape of antenna buff white on ventral surface, brownish on
dorsal surface, shaft pale buff basally (antennae not complete on
unique specimen); frons buff white; vertex and occiput buff white
tinged with yellow. Thorax: buff white with three brown streaks
(possibly grease marks), tegulae tinged with yellow. Forewing: buff
white to white, suffused with pale orange brown on dorsal third, a
row of yellow scales on outer margin. Hindwing: shining gray white,
cilia buff white. Prothoracic leg: coxa white on posterolateral surface,

No. 3547 NEARCTIC GELECHIIDAE—HODGES 25

anterior surface buff white; femur white on outer surface, pale brown
medially, gray white on edges; tibia pale brown with a gray white
fascia at three-fifths, apex gray white; tarsus brown, base and apex
of basitarsus and apex of second segment white. Mesothoracic leg:
absent on type. Metathoracic leg: coxa and femur buff to buff white,
many scales streaked with gray, trochanter nearly white; tibia white
with a faint streak of orange-brown scales at one-fourth and another
at four-fifths; tarsus white to gray brown depending upon angle of
light incidence, ventral surface orange brown basally, brown distally,
apices of segments pale. Abdomen: not observed before dissection
was made. Alar expanse: 17 mm. Male genitalia: as in figure 88
(RWH slide 2885). Female genitalia: no specimens available.

Food plant: Unknown.

Type: Male, Dixieland, Imperial Co., Calif., Mar. 15-30, 1922,
O. C. Poling (RWH slide 2885). USNM type 67648.

Discussion: L. veledae may be distinguished from obnubila by having
the inner surface of the second segment of the labial palpus pure
white and by having the apices of the valvae ellipsoidal. In many
superficial respects veledae is similar to maenadis but differs in having
the base of the shaft of the antenna unicolorous, whereas that of
maenadis is brown and white.

All of the characters examined and used to separate veledae and
maenadis may be indicative of nothing more than the two sexes of a
species or of two semiseparate populations of a species; however,
without further knowledge based on more specimens (and preferably
reared series) of either, I am recognizing two entities.

Lita rectistrigella (Barnes and Busck)

Figures 24, 72, 144, 193, 194

Gelechia rectistrigella Barnes and Busck, 1920, Contr. Nat. Hist. Lep. N. America,
vol. 4, p. 229.

Lita rectistrigella.—Busck, 1939, Proc. U.S. Nat. Mus., vol. 86, p. 573.
Maculation: as in figure 24. Head: tongue gray white basally:
maxillary palpus pale buff, folded over base of tongue; labial palpus
eray white, liberally dusted with brown; scape of antenna dark brown,
shaft ocherous basally becoming brown distally in male, gray white
interspersed with brown basally becoming brown distally in female,
sensory areas of male usually covering one-half to nearly all of
ventral surface of one-half of each segment basally; frons gray white;
vertex and occiput buff. Thorax: gray white overlaid with brown.
Forewing: streaks of gray white between brown on veins, outer mar-
gin with dark brown at ends of veins; scales of cilia gray white, apex
of each scale pale brown. Hindwing: fuscous, cilia buff. Legs:

26 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

brown with buff to gray white at apices of tarsal segments. Abdo-
men: ocherous basally, paler distally, posterior margin of segments
pale buff; ventral surface yellow white with scattered brown scales.
(This description was taken mainly from the specimen used for the
photograph. Others differ in the amount of brown present on
various surfaces.) Alar expanse: 17.0-20.5 mm. Male genitalia:
as in figure 72 (RWH slide 2642). Female genitalia: as in figure
144 (RWH slide 2629).

Food plant: Unknown.

Type: Female, USNM.

Specimens examined:

UNITED STATES: Arizona: Douglas, 19, Sept. 16, 1923, USNM;
Mohave Co., 19, Sept. 16, 1923, USNM; Paradise, Cochise Co., 19, Oct. 1-7
(RWH slide 2591), USNM; White Mts., 7200 ft., 1, Aug. 15, 1925, O. C. Poling
(RWH slide 2641), USNM. Catirornia: La Puerta Valley, 40°, 29, Septem-
ber-October 23 (RWH slides 2629 and 2642), USNM; 7 mi. N Temecula, River-
side Co., 19, Oct. 19, 1960, J. Powell, UCB; Upper Santa Ana River, San
Bernardino Co., 1c’, 19, Sept. 15-19, Sperry, USNM; West Riverside, 207,
19, Oct. 26, 27, 1925 (AB slide Mar. 21, 1933, RWH slide 2590), USNM.
Coxuorapo: Bear Creek, Morrison, 3’, Aug. 23, 1904 (RWH slide 34382), USNM;
Denver, 7.7, Sept. 5, 9, Oslar (RWH slides 3424-3429), USNM; Durango, 1?
Sept. 26, 1945, E. C. Johnston (RWH slide 2637), CNC; Lamar, 3.7, Sept. 24,
1945, E. C. Johnston (RWH slides 2734, 3433, and 3441), CNC; Rock Creek
Canyon, 3, Sept. 10-29, 1957, Margot May (RWH slides 3438-3440; RWH
wing slide 55), CPK, USNM; no locality, 5000 ft., 2%, August 1891, BMNH.
Montana: Boulder, 16%, Titus Ulke (RWH slide 2750), USNM; Butte, 40’,
Sept. 12, 1945, E. C. Johnston (RWH slide 2635), CNC, USNM. New Mexico:
Ft. Wingate, 19 , Sept. 1-7, USNM;; Hell Canyon, 1, Sept. 15, 1916, C. Heinrich
(RWH slide 2624), USNM; Mesilla, 1, C. N. Ainslie, USNM. Utan: Deer
Creek, Provo Canyon, 1<, Sept. 16, 1918, T. Spalding, USNM; Eureka, 10,
Aug. 27, 1911, T. Spalding (JFGC slide 9906), USNM; Lehi, 307, Sept. 29, 30,
1939, H. F. Thornley, CNC; Salt Lake City, 19, 5-90, Ainslie (RWH slide
2643), USNM; Spanish River, 1, Sept. 4-8, CNC. Wasuineton: Satus Creek,
Yakima Co., 1’, Sept. 16, 1949, E. C. Johnston (RWH slide 2752), CNC.

CANADA: AxsBerta: Dorothy, 17, 1921, W. G. Hodgson, BMNH.

Discussion: L. rectistrigella, as defined above, includes a relatively
homogeneous appearing group of moths. The major criteria for
recognition of the males are the width of the sensory areas of the
antennae, the aedeagus of the male genitalia, and the smooth frons.
A series of specimens from Colorado and Montana have the sensory
areas of the male antennae about one-third to one-half the width of a
segment, and two well-defined bars are present on the forewings.
They would seem to be variabilis from these characters; however, the
male genitalia place them with rectistrigella. Three possible interpre-
tations exist for this series: (1) The width of the male antennal sensory
areas is variable and thus has no taxonomic significance in this
species. (2) The width of the male antennal sensory areas is not

No. 3647 NEARCTIC GELECHIIDAE—HODGES 20

variable and thus is taxonomically significant. (3) The width of the
male antennal sensory areas has taxonomic significance at the sub-
specific level. Of the three possibilities, I have tentatively concluded
that the first is correct; however, until the species has been reared
and the progeny of known females studied, I cannot be certain.
Because no females have been correlated with the series of males under
discussion, I have discarded the second interpretation. However,
females of several species of Lita are uncommon to rare in collections,
and it could be that none have been collected. That the series might
represent a subspecies of rectistrigella seems unlikely inasmuch as a
‘normal’ male is known from Denver, Colorado; six of the second
type of male are from Denver. Thus, no geographic separation is
apparent.

L. rectistrigella may be separated from the other striate species of
Lita as follows: jubata and nefrens have frontal processes; the aedeagus
of variabilis has the apex of the flange blunt, that of rectistrigella is
poorly defined; princeps has a well-defined bar on the forewing but
no other lines, rectistrigella has series of lines; sironae is a smaller
species (alar expanse 14-17 mm.) that consistently has two bars on
the forewing, rectistrigella is a larger species (alar expanse: 17.0—20.5
mm.) that usually lacks dark bars on the forewing; the antennal seg-
ments of recens are bicolored (buff white and pale brown) and have
the male sensory areas extending to the dorsal surface, the antennal
segments of rectistrigella are not bicolored, and the sensory areas are
confined to the ventral surface; the anterior margin of the eighth
tergum of thaliae is roughly triangular (fig. 197), that of rectistrigella
is more evenly rounded (fig. 193).

Lita sironae, new species
Fieures 25, 73, 142, 195, 196

Maculation: as in figure 25. Head: tongue with buff white scales
basally, brown medially; maxillary palpus with buff white scales, some
scales streaked with pale brown; first segment of labial palpus buff
brown on outer surface, white on inner surface, second and third
segments white with scattered brown scales; ventral surface of scape
of antenna greasy buff, some scales brown tipped, dorsal surface pale
buff with numerous brown scales, shaft orange becoming darker dis-
tally (co), individual shaft segments white on basal half, pale orange
on distal half basally, becoming darker distally (2); frons, vertex, and
occiput pale buff with pale brown to brown apices. Thorax: scales
brown streaked with pale buff, scales of distal half of tegula buff white
streaked with pale brown. Forewing: streaked with white, buff
white, pale yellow, brown, and black; many scales streaked; scales of
cilia buff white, apices of basal row tipped with brown, apices of

28 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

medial and distal rows tipped with pale brown. Hindwing: fuscous
with numerous orange scales, cilia buff, apices of scales slightly darker.
Prothoracic leg: coxa, femur, and tibia dark brown, scales pale basally,
outer surface of coxa gray brown; tarsus yellow brown, apices of first
and second and base of first segment white. Mesothoracic leg: coxa
buff white; femur, tibia, and tarsus brown, a dorsal white streak at
middle of tibia, apex of tibia white, base of basitarsus white, apices
of tarsal segments slightly paler than remainder of segments. Meta-
thoracic leg: coxa and femur buff white, many scales of femur streaked
with brown; tibia buff white, apices of many scales pale brown to
brown, dorsal tuft pale buff, apex white; tarsus pale brown, base of
first segment white, apices of segments buff white. Alar expanse:
14-17 mm. Male genitalia: as in figure 73 (RWH slide 2630).
Female genitalia: as in figure 142 (RWH slide 2631).

Food plant: Unknown.

Type: Male, San Diego, Calif. (RWH slide 2630), USNM type
67649.

Paratypes:

CaiFoRNia: same locality as type, 41¢, 119, May 1-7 and Oct. 8-30 (RWH
slide 2631, JFGC slides 9900 and 9901), BMNH, CNC, CU, and LACM; same
locality as type, 57, Oct. 1-16, 1920, Karl Coolidge, LACM, USNM; same
locality as type, 367, 29, Sept. 21-Nov. 20, 1923, Piazza, LACM, USNM;; Desert
Springs, San Bernardino Co., 19, October 17, 1960, Hurd and Powell, UCB;

Los Angeles Co., 19 , September, CAS; Mouth, San Gabriel Canyon, Los Angeles
Co., 16°, Nov. 11, 1945, C. Henne, USNM.

Other specimens examined:
CauirorNia: San Diego, 10, 22, Oct. 8-15, CAS.

Discussion: L. sironae is nearest to rectistrigella, differing in having
a pale yellow to orange-brown subcostal streak on the base of the
forewing and in the smaller average size (7.9 mm. for sironae, 9.4 mm.
for rectistrigella). The signa of sironae are from 0.28-0.38 mm. long,
and the distance from the outer surface to the apex of the inwardly
projecting termen is 0.16—-0.22 mm. The same dimensions for rectis-
trigella are 0.72—0.78 mm. and 0.28-0.29 mm. In sironae the inwardly
projecting termen of the signum is more nearly vertical with the outer
margin than is that of rectistrigella. In the latter this projection is not
abruptly angled from the outer margin.

Lita thaliae, new species

Figures 26, 27, 74, 150, 197, 198

Maculation: as in figures 26 and 27. Head: tongue white basally,
becoming pale brown medially; maxillary palpus white, folded over
base of tongue; first segment of labial palpus pale brown with a
dorsobasal white patch; second and third segments white with scat-

NO. 3547 NEARCTIC GELECHIIDAE—HODGES 29

tered pale brown scales; scape of antenna white to buff white with
pale brown dusting dorsally; shaft greasy pale orange basally becoming
orange brown distally (o’), shaft blotchy greasy brown dorsally, buff
ventrally (@ ); frons off white; vertex and occiput pale buff. Thorax;
mainly buff, some scales streaked with brown and a few brown scales
streaked with buff. Forewing: off white, pale buff, pale yellow, and
brown; ridges of many scales white or off white; cilia pale to dark
buff, scale apices of basal row brown. Hindwing: pale fuscous, veins
and costal margin shining pale orange; cilia buff. Prothoracic leg:
outer surface white, anteromedial surface a mixture of gray brown
and white; femur and tibia pale brown mixed with white, apex and
epiphysis of tibia white; tarsus pale brown, base of first segment and
apices of first and second segments white. Mesothoracic leg: coxa
white; femur appearing gray on outer surface; tibia pale brown, a
white streak at middle, apex white; tarsus gray brown, apices of seg-
ments and base of first segment pale gray to white. Metathoracic
leg: coxa, femur, and tibia white, tibia with a few brown scales;
tarsus off white with scattered brown scales. Alar expanse: 19.0—
20.5mm. Male genitalia: as in figure 74 (RWH slide 2578). Female
genitalia: as in figure 150 (RWH slide 2582).

Food plant: Chrysothamnus nauseosus (Pallas) Britten.

Type: Male, Eureka, Utah, August 16, 1911, Tom Spalding, USNM
type 67650.

Paratypes:

Catirornia: Nevada Co., 19, September, CAS; near Sonora Peak, Tuolumne
Co., 11000 ft., 16, Aug. 6, 1959, C. D. MacNeill, CAS; 1 mi. W Tom’s Place,
Mono Co., 19, Aug. 13, 1957, ex Chrysothamnus nauseosus consimilis, J. Powell
(RWH slide 2582), UCB. Cotorapo: Glenwood Springs, 27, August 1892,
W. Barnes (AB slide Aug. 14, 1933; RWH slide 3249); same locality, 107,
August 1889, MCZ; no further locality, 1%, AMNH. Uvrau: same locality as

type, 20c7, 119, Aug. 16-Sept. 9, 1911 (RWH slides 2577-2580 and 2594), CAS,
USNM;; Stockton, 1.7, Sept. 6, 1907, USNM.

Discussion: The faint oblique line from the costa at the base of
the forewing to the middle of the forewing at one-fifth will separate
nonstriate specimens of thaliae from pagella, barnesiella, and texanella.
Superficially, striate specimens of thaliae are similar to some specimens
of weicur, variabilis, and rectistrigella. By lacking frontovertical
processes thaliae differs from incicur, and in the shape of the aedeagus
(rectistrigella type) it differs from variabilis. L. thaliae differs from
most known males of rectistrigella by having narrow sensory areas on
the basal segments of the antenna. The basal extension of the
aedeagus differs from that of rectistrigella by being more heavily
sclerotized. Females of thaliae have the heavy setae at the base of
the papillae anales numerous and in several rows; females of recti-

30 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

strigella have a single row of heavy setae in this position. The series
of specimens from Utah tends to be relatively uniform in hue, whereas
the specimens from Colorado and California are generally darker,
often having white areas replaced by pale buff ones.

Lita texanella (Chambers)
Ficurss 28, 29, 76, 148, 199, 200

Anesychia texanella Chambers, 1880, Journ. Cincinnati Soc. Nat. Hist., vol. 2,
. 179.
Lita rie ean tr 1939, Proc. U.S. Nat. Mus., vol. 86, p. 573.

Maculation: as in figures 28 and 29. Head: tongue with a few
white scales at base, scales elsewhere pale basally becoming dark
brown medially; maxillary palpus white (some scales brown tipped),
folded over base of tongue; labial palpus, scales of first segment gray
brown basally, dark brown apically, second and third segments white
with heavy dusting of dark brown scales; scales of scape of antenna
buff with brown tips, shaft yellow orange (<7), buff white with scat-
tered brown scales basally, becoming brown distally (@); scales of
frons pale gray white with pale brown apices, scales of vertex and
occiput buff white with brown apices. Scales of thorax and forewing
buff white with brown apices and medial streaks, dark marks on
forewing formed by scales with broad brown borders, scales of cilia
lack medial dark markings. Hindwing: fuscous, cilia buff. Pro-
thoracic leg: coxa white with a few dark scales laterally, anteromedially
dark brown, apex buff white; scales of femur, tibia, and tarsus dark
brown apically, paler basally, apices of first and second tarsal segments
pale. Mesothoracic leg: much as for prothoracic one. Metathoracic
leg: paler than prothoracic leg, dorsal tuft on tibia pale buff. Abdomen:
ocherous dorsally, segments pale apically, ventral surface pale brown,
segments lighter apically. Alar expanse: 21-27 mm. Male genitalia:
as in figure 76 (RWH slide 2569). Apex of heavily sclerotized flange
of aedeagus blunt. Female genitalia: as in figure 148 (RWH slide
3148).

Food plant: Unknown.

Type: Lectotype, present designation, male, bearing following labels:
Type, 1424; 12/4; Chambers, Keatueky; Anesychia texanella, 'Texas-
Cham.; Lectotype by R. W. Hodges. MCZ.

Specimens examined:

UNITED STATES, Txxas: Fort Davis, 37, 3.28, BMNH; Kerrville, 207, 29
(AB slide Sept. 25, 1934), USNM; same locality, 1c, March 10, H. Lacey
(RWH slide 2569), USNM; no further locality, 1h, MCZ.

MEXICO, Nuevo Leon: 3 mi. E Galeana, 5000 ft., 667, 82, Aug. 7-9, 1963,
Duckworth and Davis (RWH slides 3146-3149), CNC, CU, USNM.

Discussion: Some specimens of incicur are much like texanella on

NO. 3547 NEARCTIC GELECHIIDAE—-HODGES 31

the basis of maculation; however, teranella lacks a frontovertical
process, incicur has one.

Some specimens, particularly females, have more yellow in the basic
color scheme than occurs in the specimen described for the species.

Lita recens, new species
Fiaurss 30, 75, 147, 201, 202

Maculation: as in figure 30. Head: buff white medially, brown
laterally; maxillary palpus buff white, folded over base of tongue;
first segment of lavial palpus white on inner surface, pale brown on
outer surface, second segment brown heavily dusted with white
basally, anterior surface lacking white scales, third segment brown
with some white scales, anterior surface off white; dorsal surface of
scape of antenna brown, ventral surface buff, a faint anterior white
line at base, shaft white and red brown, sensory areas on most of
antenna extending to anterior surface (o), shaft brown with few
white scales, sensory areas narrow, obliquely placed on segments,
bordered with brown anteriorly (2); frons, vertex, and occiput
mottled buff brown. Thorax: buff brown. Forewing: scales white,
buff white, pale yellow, brown, and dark brown, many scales with
apices, many scales streaked with a second color; cilia gray white,
apices of scales brown. Hindwing: fuscous, scales shining orange
at certain angles of light incidence, veins somewhat darker, cilia
buff. Prothoracic leg: anterior surface of coxa mottled buff brown,
apex pale, posterolateral surface with brown and white scales; femur
and tibia brown with few pale scales, posterior surfaces whitish,
apex of tibia white; tarsus brown, base and apex of basitarsus and
apices of second and third segments white. Mesothoracic leg: as for
prothoracic leg but with anterior surface of coxa white. Metathoracic
leg: coxa and femur white mixed with gray brown; tibia mottled white,
gray brown, and brown on outer surface, dorsal tuft buff white;
tarsal segments pale brown to brown, apices pale. Alar expanse:
17-22 mm. Male genitalia: as in figure 75 (RWH slide 2595).
Female genitalia: as in figure 147 (RWH slide 2627).

Food plant: Stenatopsis linearifolius (de Candolle) Rydberg, leaf
tier. Ericameria cuneata (Gray) McClatchie.

Type: Male, Mt. Shasta, Calif., Aug. 19, 1939, E. C. Johnston
USNM type 67651.

Paratypes:

UNITED STATES: Cauirornra: same data as for type, 1767, 49, CNC,
CPK, USNM; 8 mi NW Chester, Plumas Co., 16, Aug. 18, 1958, J. Powell,
UCB; Desert Springs, San Bernadino Co., 16, Oct. 17, 1960, Hurd and Powell,
UCB; Hungry Valley, 5 mi. S Gorman, Ventura Co., 19, Oct. 1, 1959, J. Powell,
UCB; Jacumba, 107, Sept. 28, 1924, USNM; La Puerta Valley, 27, Oct. 16,

32 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

1923 (JFGC slide 9909), MCZ, USNM; same locality, 167, Stephens, USNM;
Lovejoy Butte, Los Angeles Co., 10, iss. Sept. 18, 1942, C. Henne, ex larva on
Ericameria cuneata spathulata, USNM; Palmdale, Los Angeles Co., 7c, 29,
iss. Sept. 8-25, 1939, J. A. Comstock, larva leaf tyer on Stenatopsis linearifolius
(JFGC slide 9911), LACM, USNM; Sheep Rock, Siskiyou Co., 10, Sept. 3,
1871, Walsingham, BMNH; Truckee, 1, Sept. 8-15, USNM. Co.orapo:
Glenwood Springs, 1?, Aug. 22, 1892, Barnes (RWH slide 2627), USN M;; Salida,
13, Aug. 27, 1938, Sperry, USNM. Montana: Butte, 26’, Sept. 12, 1945,
E. C. Johnston, CNC. New Mexico: Therma, 1o', Aug. 12, 1932, Klots,
AMNH. Orercon: Baker, 1c’, Sept. 24, 1946, J. H. Baker, USNM; Biggs,
23, Oct. 2, 1945, E. C. Johnston, CNC; Crooked R., nr. Ft. Klamath, Jackson
Co., 19, Sept. 21-23, 1871, Walsingham, BNMH; Lake Co., T 23 S, R 14 B,
S21, 307, Aug. 20, 1958, G. R. Kraft (RWH slide 2595), USNM. Uvtau: Eureka,
13, 19, Aug. 19, 27, 1911, T. Spalding (JFGC slide 9910), USNM; Stockton,
13, Sept. 1, 1904, T. Spalding, USNM. Wasuineron: Wenatchee, 20, Aug.
23 and Sept. 12, 1929, J. F. Clarke (JFGC slide 9908), USNM.

CANADA: British CotumBia: Nicola, 1c’, Sept. 6, 1923, E. R. Buckell,
CNC; Oliver, 267, Sept. 5, 16, 1953, D. F. Hardwick, J. E. H. Martin, CNC.

Other specimens examined:

Cauirornia: nr. Mono Pass, NW, 12,000 ft., Inyo Co., 1c’, Aug. 9, 1959,
C. D. MacNeill, CAS; Victorville, San Bernardino Co., 192, Oct. 12, 1936, C.
Dammers (RWH slide 2753), USNM. Mownrana: Boulder, 19, T. Ulke
(RWH slide 2751), USNM.

Discussion: L. recens is much like some specimens of variabilis,
sironae, and rectistrigella. 'The most obvious differences are those of
the antennae (particularly in the male) in which the sensory areas
extend to and slightly on the dorsal surface. In the females the
anterior dark brown border of the sensory areas will distinguish this
species from the others.

Lita princeps (Busck)
Ficures 31, 86, 153, 203, 204

Gnorimoschema princeps Busck, 1910, Proc. Ent. Soc. Washington, vol. 11, p. 175.
Lita princeps.—Busck, 1939, Proc. U.S. Nat. Mus., vol. 86, p. 573.
Maculation: as in figure 31. Head: tongue with white scales
basally; maxillary palpus white with a few pale gray brown scales,
folded over base of tongue; first segment of labial palpus white
mesally, brown laterally, second and third segments white with
scattered brown scales; scape of antenna white with dusting of brown
scales, dorsal surface of shaft with segments brown basally, buff
white distally, ventral surface buff except for piceous sensory areas;
frons white with some pale brown tipped scales; vertex and occiput
buff with pale brown tipped scales. Thorax: tegula white with
brown tipped scales basally, pale yellow distally; disk pale yellow.
Forewing: scales on costal half at base and costal margin buff white
with brown tips, scales on area costad and distad of medial brown

NO. 3547 NEARCTIC GELECHIIDAE—HODGES 30

streak white to buff white with pale brown apices, scales dorsad of
brown streak pale yellow, a few buff tipped scales on outer margin,
scales of cilia white to buff white with pale brown tips. Hindwing:
fuscous, veins slightly darker than membrane, cilia pale buff brown.
Prothoracic leg: coxa white on posterior surface, anterior surface
brown (scales white tipped with brown); femur, tibia, and tarsus
brown with scattered white scales, solid brown on third through fifth
tarsal segments, apices of first and second and base of first tarsal
segment white. Mesothoracic leg: as for prothoracic leg but with
lateral surface of femur and tibia white and a few white scales at apex
of third tarsal segment. Metathoracic leg: coxa and femur white
with scattered pale brown scales; tibia white ventrally, buff white
with some pale brown tipped scales dorsally, dorsal tuft buff white;
first tarsal segment white with some brown scales, segments two
through five brownish, apices whitish. Abdomen: dorsal surface
ocherous basally, mixed pale brown, white, and buff distally and on
ventral surface. Alar expanse: 16.5-20.0 mm. Male genitalia: as
in figure 86 (RWH slide 2565). Aedeagus stout, maximum width
one-fifth maximum length. Female genitalia: as in figure 153 (RWH
slide 2564).

Food plant: Unknown.

Type: Female, USNM.

Specimens examined:

UNITED STATES: Catirornia: Barton Flats, San Bernardino Co., 16, Oct.
13, 1945, Sperry, AMNH; Jacumba, 1 9, Oct. 1, 1924, (RWH slide 2596), USNM;
Rock Creek, 1 mi. W of Tom’s Place, Mono Co., 3o Sept. 6, 1960, C. D. Mac-
Neill, CAS; Upper Santa Ana River, San Bernardino Co., 16, 129, Sept. 14-20,
1947 and 48, Sperry (RWH slide 2564) CPK, USNM; no further locality, 19,
BMNH. Cotorapo: Durango, 19, Sept. 26, 1945, E. C. Johnston, CNC;
Mesaverde, 27, Sept. 27, 1945, E. C. Johnston (RWH slide 2565), CNC, USNM.
Ipano: Hagerman, Blue Gulch, 192, Sept. 23, 1932, USNM. Monrana: Butte,
19, Sept. 12, 1945, E. C. Johnston, CNC. New Mexico: Frijoles Canyon,
1, Sept. 8, 1941, Sperry, USNM. Oregon: Crooked River, nr. Ft. Kiamath,
Jackson Co., 6c7, 169, Sept. 21-23, 1871, Walsingham, BMNH; 15 mi. NW of
Vale, 167, Sept. 9, 1962, K. Goeden, USNM. Uran: Dividend, 5o7, 9.28, BMNH;
Eureka, 167, 19, Aug. 27, 1911, T. Spalding, LACM, USNM;; Stockton, 167,19,
Aug. 29 and Sept. 12, T. Spalding, USNM. Wasuinerton: Dry Falls, 107, 39,
Sept. 11, 1945, E. C. Johnston, CNC; Pullman, 1.7, Sept. 23, 1925, J. F. Clarke
(JF GC slide 3237), USNM; Satus Creek, Yakima Co., 206’, 19, Sept. 16, 19,
1949, E. C. Johnston, CNC; Wenatchee, 57, Aug. 21—-Sept. 5, 1929, J. F. Clarke
(AB slide Apr. 4, 1931), USNM.

CANADA: British Cotumstia: Oliver, 1000 ft. 3%, Aug. 28-Sept. 5, 1953,
D. F. Hardwick, CNC; same locality, 207, 19, Sept. 8-15, 1953, J. E. H. Martin,
CNC.

Discussion: Lita princeps differs from the other species of Lita by
having the two black dashes in the cell usually joined and broad. In
219-945—66——3

34 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

most specimens the remainder of the forewing is pale and thus contrasts
sharply with the black dash.

Color variation occurs in princeps to the extent of a general darken-
ing of all areas. The color description given above was made from a
relatively light colored specimen. In some specimens the longitudinal
streak on the forewing is divided into its two component parts;
however, I have seen no specimens in which the faint longitudinal
streaking, typical of variabilis, is present.

Lita jubata, new species
FIGurRES 32, 63, 64, 77, 141, 205, 206

Maculation: as in figure 32. Head: tongue whitish basally; maxil-
lary palpus buff white, folded over base of tongue; labial palpus
whitish, first segment with pale brown scales basally, second segment
with brown scales on anterior, medial, and lateral surfaces, third
segment solid white on anterior edge, with brown scales elsewhere;
antenna cinereous brown, scape and apical half of shaft darker than
base of shaft; lower portion of frons pale buff; vertex buff brown;
occiput paler than vertex; frontovertical processes as in figures 63
and 64. Thorax and forewing pale buff; darker markings of forewing
brown and black; cilia pale, scale apices slightly darker. Hindwing:
fuscous, cilia fuscous yellow. Legs: buff white overlaid with brown
basally, darker distally, apices of tibiae and first through third or
fourth tarsal segments pale. Abdomen: first three terga cinereous
yellow, caudal ones greasy pale brown, sterna greasy brown with
pale buff apices. Alar expanse: 16-20 mm. Male genitalia: as in
figure 77 (JFGC slide 9903). Female genitalia: as in figure 141
(JFGC slide 9905).

Food plant: Chrysothamnus viscidiflorus Nuttall and C. nauseosus
(Pallas) Britten. Specimens were reared from the flowers.

Type: Male, Satus Creek, Yakima Co., Washington, Sept. 19,
1949, E. C. Johnston, CNC.

Paratypes:

CALIFORNIA: 9 mi. S Bridgeport, Mono Co., 39, Sept. 10, 1957, ex flowers
Chrysothamnus, J. A. Chemsak, UCB, USNM; 1 mi. S Hobart Mills, 19, Sept. 1,
1957, Chrysothamnus viscidiflorus typicus, J. M. Linsley, UCB; Nevada Co., 29,
September (JFGC slide 9905), USNM; Sagehen near Hobart Mills, 12, Sept. 5,
1957, B. J. Adelson, UCB; Sheep Rock, Siskiyou Co., 1, Sept. 3, 1871, Wal-
singham, BMNH;3 mi. N Termo, Lassen Co., 12, Sept. 7, 1957, Chrysothamnus
nauseosus speciosus, J. A. Chemsak, UCB; Twin Lakes, Alpine Co., 1c, Aug. 21,
1936, E. C. Johnston (JFGC slide 4079), USNM. Cotorapo: Denver, 19,
Oslar, LACM; Glenwood Springs, 6", August and September (JF GC slide 9902),

MCZ, USNM; Rock Creek vic. Colorado Springs, 1¢7, Aug. 25, 1935, Klots,
AMNSH; Salida, 307, Aug. 26, 27, 1938, Sperry, AMNH, USNM;; South Park,

NO. 3547 NEARCTIC GELECHIIDAE—HODGES 35

10, Aug. 21, 1905, LACM. Ivano: Hagerman-Blue Gulch, 27, Sept. 23, 1932,
USNM;; Idaho Falls, Bonneville Co., 19, Sept. 9, 1950, Rindge, AMNH. Mon-
TANA: Boulder, 1c’, 19, USNM; Bozeman, 1’, Aug. 20, 1928, J. McDunnough,
CNC. Uta: Stockton, 4.7, Sept. 1-7, 1904, T. Spalding (JFGC slide 9903),
USNM. WasuinaTon: Pullman, 10, Sept. 18, 1930, J. F. Clarke, CU; Satus
Creek, Yakima Co., 21¢', 192, Sept. 16, 19, 1949, E. C. Johnston (RWH slide
2634), CNC, USNM; Wenatchee, 207, Sept. 5, 1929 (JF GC slide 9904), USNM,
Discussion: L. jubata is a striate winged Lita of the rectistrigella
genitalic group. It may be distinguished from the other known
species by the frontovertical processes (figs. 63, 64). In jubata the
vertical process has a linear cross section; other striate winged species
have a circular cross sectioned vertical process or lack processes.

Lita nefrens, new species
FiauRss 33, 34, 57, 60, 71, 138, 207, 208

Maculation: as in figures 33 and 34. Head: scales of tongue
brown, pale basally, a medial row of white scales basally; maxillary
palpus buff white, folded over base of tongue; first segment of labial
palpus white mesally, brown on lateral surface, second segment mainly
white with a few brown scales on mesal and posterior surfaces, a
small dorsobasal patch of brown scales on lateral surface, and a narrow
row of brown scales on anterior surface, third segment mainly brown
with scattered buff white scale segments, anterior surface white;
scape of antenna brown with scattered buff scale segments, shaft
ocherous becoming darker apically (), shaft brown dorsally, ocherous
ventrally, brown apically (2); frons buff white; vertex and occiput
buff brown, scales pale basally, darker apically; frontovertical
processes as in figures 57 and 60. Thorax: brown, many scales with
buff brown streaks, tegulae brown basally, buff brown apically.
Forewing: brown, many scales with buff streaks, ight streaks white,
buff white or buff; cilia buff white, scale apices pale brown. Hindwing:
fuscous with pale ocher reflections, cilia buff. Prothoracic leg:
brown; posterolateral surface of coxa white; posterior surface of femur
and tibia with many white scales; apices of coxa, femur, tibia, and
first two tarsal segments white or off white. Mesothoracic leg:
much as for prothoracic leg but with more white scales, tibia with a
medial white fascia, base of basitarsus white. Metathoracic leg:
with more white than first two, dorsal tibial tuft buff white, dorsal
surface of tarsal segments buff white apically. Abdomen: greasy
cinereous, dorsum lighter than venter, apices of segments pale.
Alar expanse: 15-22 mm. Male genitalia: as in figure 71 (RWH
slide 2638). Aedeagus of rectistrigella type. Female genitalia:
as in figure 138 (RWH slide 2639).

219-945—66——4

36 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Food plant: Chrysothamnus nauseosus (Pallas) Britten.

Type: Male, Sonora Pass, Tuolumne Co., Calif., Aug. 21, 1959,
J. Powell, CAS.

Paratypes:

CALIFORNIA: 9mi.S Bridgeport, Mono Co., 49, Sept. 10, 1959, J. A. Chemsak,
UCB; Monachee Meadows, 8000 ft., Tulare Co., 19, Aug. 8-15, USNM; Nevada
Co., 19, Sept., G. Willett, LACM; Ravendale, Lassen Co., 27,3 2, Sept. 7, 1957,
Chrysothamnus nauseosus speciosus, B. J. Adelson, UCB; same data as type,
4,19 (RWH slide 2638), UCB, USNM; same locality as type, 19, Aug. 15,
1959, G. I. Stage (RWH slide 2639), USNM; 1 mi. W Tom’s Place, Mono Co.,
1%, Aug. 13, 1957, J. Powell, Chrysothamnus nauseosus consimilis, UCB; Upper
Santa Ana R., San Bernardino Co., 16, Sept. 22, 1947, Sperry, USNM; Westgard
Pass, Inyo Co., 10, 49, Sept. 14, 15, 1938, G. Willett, LACM. Cotorapo:
Durango, 3c Sept. 26, 1945, E. C. Johnston, CNC; Glenwood Springs, 1¢,
August 1899, MCZ; Lamar, 7 <7, Sept. 24, 1945, E. C. Johnston, CNC, USNM.
New Mexico: Embido, 1, Sept. 26, Cockerell, USNM. Nuvapa: Montgomery
Pass, Mineral Co., 37, 32, Sept. 16, 1939, Geo. Willett, LACM; Mt. Magruder,
Esmeralda Co., 39, Sept. 19, 1939, Geo. Willett, LACM, USNM. Utau:
Stockton, 16, Sept. 5, 1904, T. Spalding, CU. Wasxineton: Dry Falls, 10,
Sept. 11, 1945, E. C. Johnston, CNC.

Other specimens examined:

Ca.irorNiA: Rock Cr., 1 mi. W Tom’s Place, Mono Co., 292, Sept. 6, 1960,
C. D. MacNeill, CAS; Sonora Pass, Tuolumne Co., 14%, Aug. 21, 1959, C. W.
O’Brien, CAS.

Discussion: L. nefrens is a striate-winged species with frontovertical
processes. It may be separated from jubata by the circular cross
section of the dorsal process, that of 7wbata has a linear cross section.

The color description of nefrons was taken from the type. Some
specimens are lighter in hue.

Lita invariabilis (Kearfott)
Fieurss 35, 59, 79, 134, 209, 210

Gelechia invariabilis Kearfott, 1908, Journ. New York Ent. Soc., vol. 16, p. 184.
Lita invariabilis.—Busck, 1939, Proc. U.S. Nat. Mus., vol. 86, p. 573.
Maculation: as in figure 35. Head: tongue pale gray basally,
slightly darker distally; maxillary palpus pale gray, folded over base
of tongue; first segment of labial palpus nearly white, outer surface
tinged with yellow, second segment pale gray dusted with brown, third
segment brown dusted with pale gray; scape of antenna pale gray
basally becoming light piceous distally, shaft piceous on dorsal surface,
ocherous on ventral surface of basal half, piceous on distal half; frons
pale buff; vertex and occiput with buff brown-tipped scales medially.
Thorax: tawny with scattered pale buff and brownscales. Forewing: costal
margin brown intermixed with buff from base to two-thirds length of
wing; a broad pale buff band from near base to three-fourths length

NO. 3547 NEARCTIC GELECHIIDAE—-HODGES Sid.

of wing, margined dorsally by off white; sinuous marking dark brown;
area dorsad of this line fulvous; an off-white fascia at three-fourths
followed by a mixture of dark brown, fulvous, and gray brown scales;
cilia medium gray. Hindwing: pale fuscous, veins slightly darker;
cilia and tuft of scales on vein 1 pale buff. Prothoracic leg: coxa,
femur, and tibia dark brown overlaid with off white scales; tarsal seg-
ments dark brown on anterior and lateral surfaces, base of first and
apices of first through third segments off white. Mesothoracic leg:
coxa white tinged with pale buff; femur off white, heavily dusted with
dark brown; tibia brown with a fascia at one-half and another at apex;
tarsus dark brown with base of first and apices of first through third
segments off white. Metathoracic leg: coxa and femur off white with
scattered brown scales; tibia off white, dorsal scale tuft pale buff, a
brown scale patch at two-fifths and another at four-fifths; tarsus
brown, first segment with off white basally and apically, second through
fifth segments off white apically. Abdomen: buff dorsally, off white
to buff ventrally. Alar expanse: 18-22 mm. Male genitalia: as in
figure 79 (RWH slide 2592). Heavily sclerotized flange of aedeagus
sickle shaped, apex sharply acute, ductus ejaculatorius simplex one-
and-one-third to one-and-one-half length of aedeagus. Female geni-
talia: as in figure 134 (RWH slide 2593).

Food plant: Unknown.

Type: Male, USNM.

Specimens examined:

Arizona: Mohave Co., 10’, Sept. 24-30, USNM. Catirornia: Inyo Co.,
5-6000 ft., 407, 12, Oct. 15-30, 1922, O. C. Poling (RWH slides 2592 and 2593),
USNM; Round Valley, Inyo Co., 407, 19, Oct. 15-30, 1921, O. C. Poling (AB
slides Apr. 20, 1937 and Apr. 21, 1987), CAS, USNM; Willow Creek, Siskiyou
Co., 19, Sept. 10-11, 1871, Walsingham, BMNH. Cotorapo: Boulder, 19,
Sept. 5, Cockerell, USNM; Salida, 27, Aug. 26, 1938, Sperry, USNM. New
Mexico: Gallinas Canyon, 19, BMNH; Gallup, 1<7, Sept. 11, 1961, R. W.
Hodges, USNM. Oregon: Biggs, 19, Oct. 2, 1945, E. C. Johnston, CNC.
Uran: Dividend, 2?, 9.28, BMNH; Eureka, 17, Sept. 2, 1911, T. Spalding,
USNM;; Stockton, 3c’, 39, Sept. 5-8, 1904, T. Spalding (AB slides Sept. 26,
1934 and Apr. 28, 1937), BMNH, USNM.

Discussion: No other known species of Zita has the strongly marked,
angular line on the forewing. JL. invariabilis has, as the name implies,
a stable color pattern. Observed variation occurs on the forewings
with the dorsal half basally and the apical third ranging from pale to
dark red brown.

Arla Clarke
Arla Clarke, 1942, Proc. U.S. Nat. Mus., vol. 92, p. 269.

Type-species: Arla tenuicornis Clarke, 1942, original designation.
Head: smooth scaled; tongue extending beyond metathoracic

38 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

coxa, scaled to one-third; maxillary palpus folded over base of tongue;
labial palpus recurved, second segment slightly longer than third, all
segments slender, apex acute; antenna simple, two-thirds to three-
fourths length of forewing, length of basal segments two-thirds width
in male, three-halves width in female, pecten absent. Forewing:
broadly lanceolate, apex broadly acute; 12 veins present; 6 closer to
7 and 8 than to 5 basally; 7 and 8 stalked, 7 to costa. Hindwing:
trapezoidal, outer margin moving gradually to termen, termen slightly
produced; 8 veins present; 3 and 4 approximate basally; 5 slightly
arched costally near base; 6 separate, straight; 7 slightly sinuous;
R, running into Se at one-sixth. Male genitalia: valva bilobate,
each lobe extending to base, costal lobe longer than ventral one;
saccus developed, tapering to apex; apex of juxta nearly attaining
apex of valva; aedeagus slender, a heavily sclerotized flange on caudal
half; manica heavily spined or unarmed; tegumen long; uncus shaped
like capital letter omega, wide scales on caudal margin; gnathos hook-
shaped; culcitula present. Female genitalia: signum absent; ductus
bursae heavily sclerotized posteriorly, membranous anteriorly;
eighth abdominal segment almost membranous; ostium bursae near
anterior margin of eighth sternum; apophyses posteriores more than
twice length of apophyses anteriores.

Arla is closely related to Lita but may be readily separated by
the bilobed valvae and the lack of a signum. The broad basal
antennal segments would appear to be a further development of some
Inta species that have thickened antennal segments.

Only two species of Arla are known, and both of them are from the
western tier of states.

Key to Species of Arla

BASED ON MACULATION

1. Frons gray white; forewing usually red brown; a brown spot above fold at
one-third, one on fold at one-half, and one above end of fold, one at two-
thirds length of cell, one at end of cell, and one on costa opposite one at
end of fold; alar expanse 14.5-16.5 mm... . . . . diversella (Busck)

Frons streaked gray white and gray brown; forewing gray brown; three faint
brown spots on forewing, one at middle of fold, one in cell slightly beyond
one on fold, and one at end of cell; alar expanse 18.0-22.5 mm.

tenuicornis Clarke

BASED ON MALE GENITALIA

1. Manica with scalelike spines; apex of flange of aedeagus acute; lateral margins
of eighth tergum excavated anteriorly (fig. 90). . .tenuicornis Clarke
Manica without scalelike spines; apex of flange of aedeagus relatively blunt;
lateral margins of eighth tergum straight (fig.91). . . diversella (Busck)

NO. 3647 NEARCTIC GELECHIIDAE—HODGES 39

BASED ON FEMALE GENITALIA

1. Sclerotized portion of ductus bursae seven times width of ostium bursae; same
region longer than apophyses anteriores . . . . . . diversella (Busck)
Sclerotized portion of ductus bursae one and two-thirds width of ostium

bursae; same region shorter than apophyses anteriores.
tenuicornis Clarke

Arla tenuicornis Clarke

Figures 36, 89, 90, 152, 211, 212
Arla tenuicornis Clarke, 1942, Proc. U.S. Nat. Mus., vol. 92, p. 269.

Maculation: as in figure 36. Head: tongue mixed gray white and
brown; maxillary palpus gray white, apex brown; outer surface of
first segment of labial palpus pale brown, scale bases gray white,
inner surface off white, second and third segments mixed brown and
gray white; dorsal surface of antenna greasy brown, ventral surface
ocher in male, alternating bands of greasy brown and ocher in female;
frons, vertex, and occiput mixed gray white and pale brown. Thorax:
light brown, scale bases pale. Forewing: most scales brown tipped,
gray white basally, some scales buff tipped. Hindwing: fuscous,
cilia gray buff. Prothoracic leg: coxa mixed gray white and brown;
femur, tibia, and tarsus brown (scale bases gray white) a few gray
white scales on dorsal surface of tibia at base of epiphysis, apex of
tibia, apices of tarsal segments, and base of basitarsus nearly white.
Mesothoracic leg: coxa off white; femur, tibia, and tarsus mixed
brown and gray white, apices of tibia and tarsal segments and base
of basitarsus off white. Metathoracic leg: coxa off white, several
scales gray white; trochanter, femur, tibia, and tarsus mixed gray
white and pale brown; dorsal tibial tuft off white; base and apex of
basitarsus and apices of segments two through four off white. Abdo-
men: dorsal surface mottled gray white and pale brown. Alar
expanse: 18.0-22.5 mm. Male genitalia: as in figures 89 and 90
(RWH slide 3226). Female genitalia: as in figure 152 (RWH slide
3154).

Food plant: Unknown.

Type: Male, USNM.

Specimens examined:

CaLirornia: The Geysers, Sonoma Co., 5c? 82, June 3, 21, 1938, E. C. Johns-
ton (RWH wing slide 58), CNC, USNM;; Lytle Creek, 19, May 18, 1936, Sperry
(RWH slide 3152), USNM; Scott’s Valley, Lake Co., 22, June 17-19, 1871,
Walsingham, BMNH; Shasta Retreat, Siskiyou Co., 3c, 49, June 16-23
(AB slides Feb. 14, 1933 and Sept. 4, 1936; JF GC slide 3768; RWH slide 2613),
USNM; Yosemite Valley, 1¢, July 1925, E. H. Nast, CAS. WasHineTon:

Brooks Mem. Park, Klickitat Co., 2, May 30, 1949, E. C. Johnston, CNC;
Kusshi Canyon, Yakima Co., 20’, May 28, 1949, E. C. Johnston, CNC; Satus

40 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Creek, Yakima Co., 27, May 28, 1949, E. C. Johnston (RWH slide 3226), CNC.
USNM;; Stinson Creek, Mason Co., 4¢°, June 18, 1949, E. C. Johnston, CNC;
Warwick, Klickitat Co., 1¢, June 9, 1931, T. C. Clarke (JFGC slide 3767),
USNM.

Discussion: A. tenuicornis may be separated from dwersella by
the relatively uniformly colored forewing, the presence of scalari-
form spines of the manica, and the broad opening of the ductus
bursae. Females of tenuicornis approach some specimens of L. vari-
abilis in appearance. The lack of a signum will quickly distinguish
them from any species of Lita.

Arla diversella (Busck), new combination

Fiaures 37, 91, 92, 154, 213, 214
Gelechia diversella Busck, 1916, Proc. Ent. Soc. Washington, vol. 18, p. 149.

Maculation: as in figure 37. Head: tongue gray white for short
distance basally, then brown; maxillary palpus gray white; outer
surface of first segment of labial palpus brown, inner surface gray
white, second segment gray white dusted with brown, third segment
as for second but with more brown on inner surface; dorsal surface of
antenna greasy brown, ventral surface ocher (<o”); dorsal surface of
scape and base of shaft mixed brown and gray white, scales of dorsal
surface of shaft brown apically, pale basally, ventral surface alter-
nating buff and brown (9); frons buff white; vertex and occiput pale
eray, scales tipped with pale brown. ‘Thorax: red brown, basal row
of scales brown; scales on metathorax gray white. Forewing: red
brown, brown, and pale gray, cilia pale gray. Hindwing: fuscous,
cilia gray buff. Prothoracic leg: coxa and femur brown mixed with
eray white; tibia and tarsus brown, scale bases gray white, apex of
tibia, base and apex of basitarsus, and apices of second and third
tarsal segments gray white. Mesothoracic leg: coxa gray and white;
femur, tibia, and tarsus as for prothoracic leg but with more gray
white, apex of fourth tarsal segment gray white. Metathoracic leg:
much as for mesothoracic leg but paler, dorsal tibial tuft off white.
Abdomen: dorsal surface shining pale gray; ventral surface gray
brown, apices of sterna pale. Alar expanse: 14.5-16.5 mm. Male
genitalia: as in figures 91 and 92 (RWH slide 2557). Female geni-
talia: as in figure 154 (RWH slide 2558).

Food plant: Lotus species.

Type: Female, USNM.

Specimens examined:

CALIFORNIA: Deer Creek, El Dorado Co., 16, 19, iss. May 31 and June 6,

1935, reared from Lotus, H. H. Keifer, USNM; La Mesa, San Diego Co., 10,
Apr. 15, 1950, E. C. Johnston, CNC; Moraga Ridge, Alameda Co., 1c’, June 11,

NO. 3647 NEARCTIC GELECHIIDAE—HODGES 41

1936, H. H. Keifer, CAS; Morro Rock, San Louis Obisbo Co., 19, May 4, 1962,
J. Powell, UCB; San Diego Co., 286’, 49, Apr. 16-June 13, W. S. Wright,
Ricksecker (AB slide Mar. 5, 1931; RWH slides 2557 and 2558), LACM, MCZ,
USNM.

Discussion: A. diversella may be separated from tenuicornis as
indicated in the keys. The two valval lobes, slender aedeagus,
sclerotized ductus bursae, and lack of signum associate diversella with
Arla rather than Lata.

The collection record from El Dorado county indicates that diver-
sella at least has a wider distribution than coastal Southern California
and that it may be found in Oregon and Washington.

Neodactylota Busck
Neodactylota Busck, 1903, Proc. U.S. Nat. Mus., vol. 25, p. 835.

Type-species: Dactylota snellenella Walsingham, 1888, original
designation.

Head: smooth scaled; tongue moderate, scaled for short distance
basally; maxillary palpus slightly folded over base of tongue to
drooping; labial palpus recurved, slender, second and third segments
subequal in length; antenna with pecten absent, two-thirds to four-
fifths length of forewing, simple. Forewing: broadly lanceolate; 12
veins present; 2 from cell at two-thirds length of cell; 2, 3, 4, and 5
subparallel; 6 closer to 7 than to 5; 7 and 8 stalked. Hindwing:
dimorphic o’, outer margin deeply emarginated; 6 veins present, 4 and
5 absent; 6 and 7 united to three-fifths, 6 to outer margin, 7 to costa;
9, wing quadrate, termen produced; 8 veins present; 3 and 4 separate
to short stalked; 5 arising closer to 4 than to 6. Male genitalia: valva
with two costal lobes; juxta fused with vinculum basally; aedeagus
relatively linear. Female genitalia: ostium bursae on anterior margin
to middle of eighth sternum; eighth sternum unadorned to slightly so;
apophyses posteriores longer than apophyses anteriores.

The species included in Neodactylota, as defined above, can be
separated from those in Eudactylota by the presence of well or moder-
ately well defined spots on the disk of the forewing; by the hindwing
shape being dimorphic between the sexes (the outer margin is deeply
emarginated in the male with a concomitant loss of veins 4 and 5,
whereas the outer margin of the female is normal); and by the valva
having two costal lobes.

With the addition of three new species the total known for the
genus is four. However, further collecting in Mexico and adjacent
areas of the Southwest may reveal that others exist. Unfortunately,
very little is known of the distribution of any of the species.

42 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Key to Species of Neodactylota

BASED ON MACULATION

1. Head with metallic lead-colored scales on vertex and occiput.
Sea new species

Head without metallic colored scales... . ‘ sIMeESOS 62
2. Dark spots of forewing well defined, siréounded ee narrow margin of pale
colored scales... . Stan euelo

Dark spots of forewing non weil deanede ae seecinedl be Pele calered scales.
egena, new species

3. Second and third segments of labial palpus dark gray with scattered gray
white scales, an anterior gray white line on third segment.

snellenella (Walsingham)

Second and third segments of labial eee pale yellow, third segment be-

coming brown apically. .... . . . . .liguritrix, new species

PARTIAL KEY BASED ON MALE GENITALIA

1. Medial invagination of juxta extending to base of juxta.
snellenella (Walsingham)

Medial invagination of juxta one-third length ofjuxta. .........2
2. Distal costal lobe of valva curved and slightly twisted, very slender on distal
Dali pn 2 Fe . . .liguritrix, new species

Distal costal lobe enn Sentry eee to apex.
basilica, new species

PARTIAL KEY BASED ON FEMALE GENITALIA

1. Highth tergum produced medially (fig. 157). . . . basilica, new species
Kighth tergum not produced medially... . . Sate. eee
Eighth sternum heavily sclerotized, caudal margin ienbaineee medially, a
semicircular sclerotized band parallel with caudal margin (fig. 156).
egena, new species
HKighth sternum lightly sclerotized, caudal margin not defined, no special
sclerotized bands (fig. 155)... . .. =. . .snellenella (Walsingham)

no

Neodactylota snellenella (Walsingham)

Fieures 38, 938, 95, 155, 215, 216

Dactylota snellenella Walsingham, 1888, Insect life, vol. 1, p. 84.
Neodactylota snellenella.—Busck, 1903, Proc. U.S. Nat. Mus., vol. 25, p. 836.
Maculation: as in figure 38. Head: tongue mottled gray white
and brown basally; maxillary palpus gray white with a few brown
scales, folded over base of tongue; first segment of labial palpus
mainly white with some brown and brown-tipped scales; second seg-
ment brown with several white scales, white scales more numerous
basally, third segment mainly brown, anterior surface white; scape of
antenna brown, scale bases pale, with an anterior white line and a
posteroapical white scale row, shaft pale yellow on anterior surface,
dorsal surface brown basally, segments brown and pale yellow short

No. 3547 NEARCTIC GELECHIIDAE—HODGES 43

distance beyond base; lower portion of frons gray white, many scales
brown tipped medially; scales of vertex and occiput buff brown with
brown apices, scales of medial area of vertex gray with metallic
reflections. Thorax: scales gray, tipped and streaked with brown;
apex of tegula pale buff. Forewing: scales yellow, white, gray, buff,
red brown, and brown, often streaked with a light or dark hue; most
scales tipped with brown. Hindwing: fuscous, margins of scales
darker than remainder of scale, scales shining orange at some angles
of light incidence; cilia pale buff. Prothoracic leg: anterior surface
of coxa gray white, many scales tipped with brown; femur and tibia
brown with several gray white scales, outer surface of femur mainly
white, tibia with a white scale row just beyond middle and at apex;
tarsus dark brown, scales lighter basally, base and apex of basitarsus
and apex of second segment white, apex of third segment with a few
white scales. Mesothoracic leg: lighter than prothoracic leg; tibia
with an oblique white fascia at one-fourth and another at one-half,
apex white; base and apex of basitarsus and apices of second and
third segments white. Metathoracic leg: coxa and femur mainly
white with scattered brown scales; tibia with a broad white streak at
one-fourth and another at three-fifths, apex white, scales of dorsal tuft
pale buff basally becoming white distally; tarsal segments gray
brown basally, broadly marked with white or gray white apically.
Abdomen: terga greasy ocher basally, gray brown distally, apices of
terga pale; sterna buff white, scales becoming greasy shortly after
death of specimen. Alar expanse: 15.5-18.0 mm. Male genitalia:
as in figures 93 and 95 (RWH slide 2150). Distal costal lobe of
valva about one-fourth length of basal costal lobe; lobes of juxta
nearly separated. Female genitalia: as in figure 155 (RWH slide
allo):

Food plant: Unknown.

Type: Lectotype, present designation; male, bearing following labels:
(1) Type. (2) Arizona, Morrison, 1883, 35345. (3) Walsingham
Collection, 1910-427. (4) Dactylota snellenella, Wlsm., U.S. Dept
Agr. Div Ent. Ins. Life. I-84 (1888); Type o, descr. (5) BM(NH) &
genitalia slide no. 9209. In BMNH.

Specimens examined:

Arizona: Huachuca Mts., 16’, Sept. 17, 1903, Oslar, BMNH; Santa Rita
Mts., Madera Canyon, 4880 ft., Santa Cruz Co., 27, 19, May 3-June 7, 1963,
J. G. Franclemont (RWH wing slide 80), JGF; no further locality, 1, 1881,
Morrison (RWH slide 2150), USNM; no further locality, 19, 1883, Morrison
(RWH slide 3155), BMNH. Texas: Ft. Davis, 5000 ft., 1¢, 5. 28, BMNH.

Discussion: The alar expanse of snellenelia is at least 3 mm. more
than that of basilica, liguritriz, or egena. The lack of metallic colored

44 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

scales on the vertex and the gray white labial palpi will separate
snellenella from basilica and liguritriz, respectively.

In Madera Canyon snellenella appears to be a spring species inas-
much as no specimens were taken during the 1959 season from June
29-November 11. On the other hand the specimen from the Huachuca
Mts. (less than 50 miles distant) was collected in September; thus,
the species may emerge sporadically during the year.

Neodactylota liguritrix, new species
Ficures 39, 102, 106, 217, 218

Maculation: as in figure 39. Head: tongue with mixed buff white
and brown scales basally, becoming brown; maxillary palpus buff
white, somewhat drooping and curved over base of tongue; first
segment of labial palpus nearly white on inner surface, outer surface
with brown, second segment pale yellow with some pale brown scales
on outer surface at base, third segment yellow, becoming brown
apically on inner and anterior surfaces; scape of antenna buff, shaft
buff basally, becoming greasy brown distally; frons pale buff; vertex
and occiput pale buff, medial scales with dark margins. Thorax:
base of tegula and medial area of mesothorax gray brown; apex of
tegula and lateral margins of mesothorax pale buff; a lateral dark
brown spot on mesothorax near apex of tegula and one at middle of
caudal margin; metathorax gray brown to lead colored. Forewing:
most scales pale buff, many tipped with gray brown; costal margin
gray brown; spots on disk dark brown. Hindwing: fuscous, cilia
dark fuscous yellow. Prothoracic leg: gray brown, scale bases pale;
some off white scales at base and apex of basitarsus and at apex of
second tarsal segment. Mesothoracic leg: coxa with numerous buff
white scales on outer surface; femur gray brown; tibia mottled buff
and gray brown, mainly buff; dorsal surface of basitarsus and apices
of second and third segments off white, remainder gray brown.
Metathoracic leg: outer surface of coxa mainly buff, medial scales
streaked with brown; trochanter gray brown; femur buff on dorso-
lateral margin, gray brown on lateral surface; tibia gray brown
laterally, a buff fascia at one-fifth and one at base of first pair of
tibial spurs, dorsal tuft buff; apex white; tarsus gray brown on outer
surface, dorsal surface of first two and base of third segment pale
buff, apices of first four segments pale buff. Abdomen: greasy gray
brown dorsally, first two terga with ocher cast; scales of sterna gray
brown, pale basally. Alar expanse: 10.5-12.0 mm. Male genitalia:
as In figures 102 and 106 (RWH slide 3153). Distal costal lobe slender,
somewhat sinuous; apex of basal costal lobe twisted; apex of juxta
emarginated. Female genitalia: no specimens available.

Food plant: Unknown.

NO. 3547 NEARCTIC GELECHIIDAE—HODGES 45

Type: Male, Corpus Christi, Tex., Mar. 28, 1943, at light W. M.
Gordon, CU type 4219.

Paratypes:

Same locality as type, 40’, Mar. 28 and May 22, 1943, W. M. Gordon (RWH
slide 3153, RWH wing slide 75), CU, USNM.

Discussion: In appearance, liguritriz is nearest to basilica; it differs
in lacking metallic lead-colored scales on the vertex and occiput. WN.
liguritriz has the dark spots on the disk much more distinct than does
egenda.

Neodactylota basilica, new species
Figures 40, 94, 98, 157, 219, 220

Maculation: as in figure 40. Head: tongue buff white basally;
maxillary palpus buff white, somewhat drooping; first segment of
labial palpus buff white with some gray-brown scales before middle,
second segment pale yellow, third segment pale yellow becoming
brown black from middle to apex on anterior and inner surfaces;
scape of antenna buff white dorsally, gray brown ventrally, shaft
ocher basally becoming greasy brown distally, several segments bi-
colored; frons buff white; scales of middle of vertex and occiput
shining lead colored, lateral scales yellow. Thorax: irregularly
mottled with buff white, gray brown, and dark brown; apex of tegula
buff white; a lateral dark brown spot on thorax at apex of tegula;
caudal margin of mesothorax with a small dark brown spot medially;
metathorax with lead gray-colored scales. Forewing: scales buff white
with gray-brown apices, spots dark brown; costal margin, especially
basally, gray brown. MHindwing: fuscous, cilia slightly paler than
membrane. Prothoracic leg: gray brown; a few buff white scales at
apex of coxa and at two-thirds and apex of tibia, apices of first two
tarsal sezments white. Mesothoracic leg: coxa with buff-white scales
on outer surface; tibia buff white on dorsal surface; tarsus with buff
white on dorsal surface and apex of first and second segments. Meta-
thoracic leg: gray brown; dorsal tuft of tibia buff white; dorsal surface
of first two tarsal segments off-white, apex of third and fourth seg-
ments buff. Abdomen: lead gray, basal terga with dark ocher re-
flections; sterna gray brown, first segment buff. Alar expanse 10.5-
12.0 mm. Male genitalia: as in figures 94 and 98 (RWH slide 721).
Distal costal lobe of valva about one-half length of basal costal lobe;
apex of juxta emarginated. Female genitalia: as in figure 157 (RWH
slide 1245).

Food plant: Unknown.

Type: Male, Sycamore Canyon, 3800 ft., Santa Cruz Co., Ariz.,
Sept. 25, 1959, R. W. Hodges, CU type 4220.

46 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Paratypes:

Arizona: Pefia Blanca Canyon, Santa Cruz Co., 2c, 19, Aug. 7-Sept. 4,
1959, R. W. Hodges (RWH slides 721, 941, and 1245; RWH wing slides 76 and
79), CU, USNM.

Discussion: The metallic lead-colored scales on the vertex and
occiput will separate basilica from snellenella, liguritriz, and egena.

Neodactylota egena, new species
Figures 41, 156

Maculation: as in figure 41. Head: tongue pale buff; maxillary
palpus pale buff, porrect to drooping along base of tongue; labial
palpus pale buff to pale yellow, third segment with brown scales; scape
of antenna buff with overlay of brown tipped scales, segments of shaft
bicolored basally, pale buff and brown, mainly brown distally; frons
pale buff; vertex and occiput gray buff with purple reflections medially,
a tuft of orange scales above eye. Thorax: gray brown, scale bases
pale, caudolateral margins buff. Forewing: nearly uniform gray
brown, scale bases pale; marks on disk brown, not strongly contrasting
with gray-brown background; cilia buff. Hindwing: fuscous. Pro-
thoracic leg: coxa pale buff; femur, tibia, and tarsus gray brown; femur
with numerous pale buff scales; tibia with apex and some scattered
scale rows pale buff; base and apex of basitarsus and apex of second
tarsal segment pale buff. Mesothoracic leg: much as for prothoracic
leg but with more pale scales. Metathoracic leg: coxa buff white;
femur buff white with some gray scales; tibia pale brown ventro-
laterally, dorsal tuft pale buff, a fascia at base of first pair of spurs and
apex pale buff; tarsus pale brown basally becoming darker distally,
dorsal surface of basitarsus and part of second segment pale buff,
apices of segments pale. Abdomen: not observed before dissection
was made. Alar expanse: 10.5 mm. Male genitalia: no specimens
available. Female genitalia: as in figure 156 (RWH slide 1532).

Food plant: Unknown.

Type: Female, Madera Canyon, 4880 ft., Santa Rita Mtns., Ariz.
Oct. 19, 1959, R. W. Hodges (RWH slide 1532), CU type 4221.

Discussion: The small size immediately separates egena from
snellenella; the relative indistinctness of the dark spots of the forewings
separates it from basilica and liguritria.

Eudactylota Walsingham

Eudactylota Walsingham, 1911, in Godman and Salvin, Biologia Centrali-
Americana, vol. 42 (Lepidoptera-Heterocera, vol. 4), p. 54.

Type-species: Neodactylota barberella Busck, 1903, original desig-
nation.

NO. 3547 NEARCTIC GELECHIIDAE—HODGES 47

Head: smooth scaled; tongue moderate, scaled to one-third or
one-half length; maxillary palpus porrect to drooping, somewhat folded
over base of tongue; labial palpus recurved, slender, second and third
segments subequal in length; antenna with pecten absent, two-thirds
to three-fourths length of forewing, simple. Forewing: broadly
lanceolate; 12 veins present; 2, 3, 4, and 5 equidistant, subparallel;
7 and 8 stalked, 7 to costa. Hindwing: subquadrate, apex strongly
produced, outer margin indented between 5 and 6; 6 and 7 stalked,
6 to outer margin; R; showing as cross vein between Scand Rs. Male
genitalia: valva with one costal lobe; juxta fused with vinculum
basally, lobes separate, stout setae present near inner margin on distal
half; saccus moderate; aedeagus somewhat globose basally; tegumen
broad; spined culcitula present; gnathos a strong hook. Female
genitalia: ductus bursae long; corpus bursae with a single signum;
ostium bursae on eighth sternum, eighth sternum usually with medial
folds with numerous fine spines; apophyses posteriores much longer
than apophyses anteriores.

Walsingham proposed Fudectylota for barberella (Busck). Later
authors have treated the genus as a synonym of Neodactylota; however,
the two are distinct on characters of facies, wing shape, venation, and
male genitalia as is indicated under Neodactylota.

Stomopteryx iobapta Meyrick is transferred to Hudactylota, and two
new species are described bringing the number of known species to
four. Nothing is known of the life history of any species, and very
little is known of their distribution.

Key to Species of Eudactylota

BASED ON MACULATION

bo

1. Anterior surface of third segment of labial palpus with a white line .
Anterior surface of third segment of labial palpus without a white line.
barberella (Busck)
2. Second segment of labial palpus with interior ventral margin white, ventral

marcin blacks.) sles . . . . . diadota, new species
Ventral margin of second senthaut of eel papa with mixed black and white
scales yaaa ‘ a0

3. Outer, apical retaghoracie tibial spur aphite, Gasallvae gray Bisee rail
(usually); pinkish white fascia on forewing relatively narrow, confined
apically, buff streaks along costal and outer margins absent or faint.

iobapta (Meyrick)

Outer, apical metathoracic tibial spur white basally and distally, gray black
medially; pinkish white fascia on forewing relatively broad, gradually
darkening to apex; buff streaks along costal and outer margins prominent.
abstemia, new species

48 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

BASED ON MALE GENITALIA

1. Lobes of juxta distant basally, width of opening greater than basal width of
lobe) 5 seas tebe) 22a te Ree ee
Lobes of juxta spends! Basal. Seopa is ip eh RO

2. An invaginated sclerotized band between lobes of te oe 100).
diadota, new species
No sclerotized band between lobes of juxta . . . . . . barberella (Busck)
3. Lobes of juxta separated to base, inner margin uniformly sclerotized (fig. 96).
iobapta (Meyrick)
Lobes of juxta separated to one-third, inner margin where lobes join heavily
sclerotized “(figs 105).° 7. “iei.) Sey Set VS") i abstemia, new species

BASED ON FEMALE GENITALIA

1. Anterior margin of eighth sternum emarginated medially (fig. 160) . . . .2
Anterior margin of eighth sternum not emarginated medially (fig. 158) . . .3
2. Lateral sclerites (folds) of eighth sternum touching medially, caudal margins
relatively abruptly angled (70-80 degrees) beyond ostium bursae, no heavily
sclerotized ridges caudad of ostium bursae (fig. 160).
abstemia, new species
Lateral sclerites (folds) of eighth sternum not touching medially, caudal
margins diverging after point of closest proximity, later diverging with
approximately a 45 degree angle; heavily sclerotized ridges caudad of
ostium bursae present (fig. 159) . ..... . . . .diadota, new species
3. Lateral sclerites of eighth sternum nearest anteriorly, gradually diverging
posteriorly (fig: 158)) 2. 2... . . .. .lobapta (Meyrick)
Lateral sclerites of eighth peed ences panies becoming indistinct
beyond point of proximity (fig. 161)... . . . . . .barberella (Busck)

Eudactylota barberella (Busck)
FicureEs 42, 101, 103, 161, 221, 222
Neodactylota barberella Busck, 1903, Proc. U.S. Nat. Mus., vol. 25, p. 836.
Eudactylota barberella—Walsingham, 1911, ix Godman and Salvin, Biologia
Centrali-Americana, vol. 42 (Lepidoptera-Heterocera, vol. 4), p. 55.

Maculation: as in figure 42. Head: tongue mottled with uni-
colorous brown and white scales; maxillary palpus white with a few
brown scales, curved over base of tongue; first segment of labial
palpus white, a dorsolateral row of brown scales, second and third
segments brown with scattered white scales, white scales more
numerous on basal two-thirds of second segment; dorsal surface of
scape of antenna brown, an anterior white line, apex mainly white,
ventral surface white, shaft brown with an anterior broken white line,
many segments with white scales; frons, vertex, and occiput buff
brown, scales with dark apices. Thorax: anterior margin of tegula
brown, remainder red brown; mesothorax dark gray brown medially,
red brown laterally; metathorax pale buff. Forewing: brown, red
brown, gray brown, gray white, and white; many scales with purple
reflections. Hindwing: pale fuscous, shining orange at some angles
of light incidence, cilia pale fuscous. Prothoracic leg: coxa brown,

NO. 3547 NEARCTIC GELECHIIDAE—HODGES 49

heavily overlaid with white; femur brown, moderately overlaid with
white; tibia and tarsus brown, scale bases pale, especially on tibia, a
white fascia at middle and another at apex of tibia, apices of first three
tarsal segments white or with white scales. Mesothoracic leg: as
for prothoracic leg. Metathoracic leg: coxa white basally, gray
brown with purple reflections distally; trochanter gray brown with a
few partially white scales medially, pale buff apically; femur brown
heavily overlaid with white; tibia brown, scale bases pale, an oblique
white fascia at base of first pair of spurs, apex white, dorsal tuft
streaked buff and fuscous; tarsus brown, apices of first four segments
white. Abdomen: greasy gray brown dorsally, first two terga some-
what ocherous; ventral surface gray brown, first sternum pale yellow,
caudal margin of second sternum pale yellow. Alar expanse: 11-14
mm. Male genitalia: as in figures 101 and 103 (RWH slides 2151
and 2152). Female genitalia: as in figure 161 (RWH slide 2153).

Food plant: Unknown.

Type: Female, USNM.

Specimens examined:

UNITED STATES: Arizona: Dewey, 19, June 16-23, USNM; Fort Valley,
7% mi. NW Flagstaff, 7350 ft., Coconino Co., 100%, 39, June 21—Aug. 21, 1961,
R.W. Hodges (RWH slides 2152-2154; RWH wing slide 67), BMNH, CU,
USNM;; Hart Prairie, 10 mi. NNW Flagstaff, 8500 ft., Coconino Co., 367, June 23
and July 14, 1961, R.W. Hodges (RWH slide 2151), CU, USNM; Williams,
19, May 27, H.S. Barber (RWH slide 2149), USNM. Co.xorapo: Boulder,
19, USNM; Boulder, University of Colorado Campus, 19, July 16, T.D.A.
Cockerell, USNM. New Mexico: Las Vegas, 19, Aug. 11, H.S. Barber (RWH
wing slide 68).

MEXICO: Guerrero: Amula, 6000 ft., 19, Aug. 18, H.H. Smith, USNM.

Discussion: E. barberella may be quickly separated from the other
species of EHudactylota by the lack of an anterior white line on the
third segment of the labial palpus. The male genitalia are nearest
those of tobapta but differ by having the lobes of the juxta slightly
swollen before the apex; those of iobapta taper gradually to the apex.

Eudactylota iobapta (Meyrick), new combination

Ficures 43, 96, 97, 158, 223, 224
Stomopteryx tobapta Meyrick, 1927, Exot. Micr., vol. 3, p. 352.

Maculation: as in figure 43. Head: tongue with white and gray
brown scales basally; maxillary palpus white, somewhat drooping at
side of tongue; first segment of labial palpus white with a brown
saddle near apex, second segment brown heavily dusted with white,
third segment brown with an anterior white line and an interrupted
white line on inner surface from near base to three-fifths, posterior
surface mainly white; antenna brown, scape with an anterior white line

50 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

continued on shaft as a broken line, a posterior broken white line on
shaft from base to beyond one-half; ventral margin of frons white
followed by a series of brown and white scales running from anterior
margin of eye to middle; vertex and occiput buff brown, scales with
dark apices, all with metallic reflections. Thorax: tegula and meso-
thorax shining orange brown, mesothorax gray brown mesally; meta-
thorax buff. Forewing: mainly shining orange brown, costal margin
mixed brown and white, fascia off white, a buff streak on costal
margin starting at fascia, cilia buff. Hindwing: fuscous, shining
orange brown at certain angles of light incidence, cilia pale fuscous.
Prothoracic leg: coxa and femur brown moderately dusted with
white; tibia brown with a few scattered white scales, a white streak
above base of epiphysis, apex white; tarsus brown, apices of first
three segments white. Mesothoracic leg: coxa mainly white, some
gray brown scales medially; femur speckled greasy white and brown;
tibia brown, scale bases pale, a white fascia at one-half, apex white;
tarsus brown, apices of first four segments white. Metathoracic
leg: coxa white, gray medially; trochanter mixed gray brown and
white; femur speckled brown and white; tibia mainly brown on
outer surface, scale bases pale, a white fascia at base of first pair of
spurs, apex white, tuft buff to white; tarsus brown, scale bases pale on
first segment, apices of first four segments white. Abdomen: greasy
gray brown dorsally, first two terga somewhat dark ocher; ventral
surface gray brown, scales at apices of sterna with buff white streaks,
first sternum pale yellow. Alar expanse: 9-13 mm. Male genitalia:
as in figures 96 and 97 (RWH slide 2164). Female genitalia: as
in figure 158 (RWH slide 2161).

Food plant: Unknown.

Type: Male, BMNH.

Specimens examined:

UNITED STATES: Arizona: Chiricahua Mts., nr. Portal, 39, July 4 and
8, 1939, A.F. Braun, AFB; Madera Canyon, 4880 ft., Santa Rita Mtns., Santa
Cruz Co., 18¢%°, 139, June 29-Aug. 30, 1959, R.W. Hodges (RWH slides 944,
2160, 2164; RWH wing slide 69), CU, USNM; Palmerlee, 14°, BMNH; Pefia
Blanca Canyon, Santa Cruz Co., 1c, 159, Aug. 7-26, 1959, R.W. Hodges
(RWH slides 2161 and 2166; RWH wing slide 70), CU, USNM; same locality,
19, June 7, 1963, J.G. Franclemont, JGF; 4 mi. ESE Pine, 5400 ft., Gila
Co., 29, Sept. 1 and 5, 1961, R.W. Hodges (RWH slide 2163), USNM; West
Fork, 6500 ft., 16 mi. SW Flagstaff, Coconino Co., 16, July 13, 1961, R.W.
Hodges (RWH slide 2162), USNM. Texas: Alpine, Brewster Co., 1c’, 29,
May 22, 1950, E.C. Johnston, CNC; Ft. Davis, Jeff Davis Co., 29, May 20,
1950, E.C. Johnston, CNC; Limpia Canyon, Jeff Davis Co., 40, 19, May 20-
June 4, 1950, E.C. Johnston, CNC; Brewster Co., 5000 and 7000 ft., 467, 59,
3.28-4.26, AFB, BMNH.

MEXICO: Mexico: ruins of Teotihuacdn, 167,19, July 25, 1963, Duckworth
and Davis (RWH slides 3462 and 3463), USNM. Nugvo Leon: 3 mi. E

No. 3547 NEARCTIC GELECHIIDAE—HODGES Sit

Galeana, 5000 ft., 2o7, 89, Aug. 7-9, 1963, Duckworth and Davis (RWH
slides 3333-3335), USNM.

Discussion: Some specimens of tobapta are very near abstemia in
maculation, and for these the genitalia should be examined for place-
ment. The uniformly sclerotized inner margin of the lobes of the
juxta will separate iobapta from abstemia and diadota; the lobes of
the juxta tapering to the apex will separate iobapta from barberella.

Specimens vary in the coloration of the forewings: Some are golden
brown, others dark brown.

Eudactylota diadota, new species
Figures 44, 99, 100, 159, 225, 226

Maculation: as in figure 44. Coloration: as for cobapta except for
the following: second segment of labial palpus with a black ventral
line, third segment with a white line on anterior and posterior sur-
faces; forewing with pink scales on costal margin from fascia to apex;
metathoracic tibia with an oblique white fascia starting at one-fifth,
another starting just before first pair of spurs; abdomen gray buff
dorsally, apices of segments buff white, pale gray buff ventrally,
apices of segments pale. Alar expanse: 9.5-12.0 mm. Male geni-
talia: as in figures 99 and 100 (RWH slide 2157). Female genitalia:
as In figure 159 (RWH slide 946).

Food plant: Unknown.

Type: Male, Madera Canyon, 4880 ft., Santa Rita Mtns., Ariz.,
June 29, 1959, R. W. Hodges (RWH slide 2156), CU type 4222.

Paratypes:

Same locality as type, 67, 139, June 29-Aug. 27, 1959, R.W. Hodges (RWH
slides 946, 2157, and 2158; RWH wing slides 71 and 72), BMNH, CU, USNM.

Discussion: Superficially, diadota is similar to iobapta and abstemia
but differs in having a black line on the anterior surface of the labial
palpus. The pink on the outer costal margin will separate diadota
from the other species of Hudactylota.

Eudactylota abstemia, new species
Fiaures 45, 104, 105, 160, 227, 228

Maculation: as in figure 45. Coloration: as for iobapta; forewing
with more pinkish-white scales. Alar expanse: 8-11 mm. Male
genitalia: as in figures 104 and 105 (RWH slide 942). Female
genitalia: as in figure 160 (RWH slide 2155).

Food plant: Unknown.

Type: Male, Madera Canyon, 4880 ft., Santa Rita Mtns., Ariz.,
Aug. 12, 1959, R.W. Hodges (RWH slide 942), CU type 4223.

Paratypes:

Same locality as type, 56’, 219, July 14-Aug. 28, 1959, R.W. Hodges (RWH

219-945—66——_5

52 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

slides 943, 945, and 2155; RWH wing slides 73 and 74), AMNH, BMNH, CAS,
CNC, CU, and USNM.

Discussion: FE. abstemia may be separated from barberella by the
presence of an anterior white line on the third segment of the labial
palpus; from diadota by the ventral margin of the second segment of
the labial palpus having white scales; and from iobapta by the juxtal
lobes being heavily sclerotized on the inner basal one-third. The
females of abstemia and iobapta may be separated by the ostium bursae
being near the caudal margin of the eighth sternum in abstemia, at
one-third in iobapta.

Friseria Busck
Friseria Busck, 1939, Proc. U.S. Nat. Mus., vol. 86, p. 573.

Type-species: (Gelechia lindenella Busck, 1903) =Gelechia cockerella
Busck, 1908, original designation.

Head: smooth scaled; tongue moderate, scaled to one-third or one-
half; maxillary palpus folded over base of tongue; labial palpus
recurved, second and third segments subequal in length, second seg-
ment slightly thicker than third, apex acute; antenna two-thirds length
of forewing, ciliate in male, simple in female, pecten absent. Fore-
wing: broadly lanceolate, apex broadly acute; 12 veins present; 7 and
8 stalked, 7 to costa. Hindwing: trapezoidal, outer margin gradually
moving to termen, apex scarcely produced; 8 veins present; 3 and 4
approximate to connate; 5 arching costally beyond base; 6 and 7
approximate to stalked, usually separate; 7 sinuous; R; running into
Se at one-fifth. Male genitalia: valva with costal lobe free, two other
lobes from ventral margin of costal lobe (variously developed) ;
vinculum narrow laterally, saccus broadly rounded; aedeagus with
base expanded; tegumen broad, gnathos hook shaped; culcitula pres-
ent; uncus short, caudal margin with long, stout setae. Female
genitalia: signum a plate with an inwardly projecting flange from each
end (flange very small to large and heavily sclerotized) ; eighth sternum
with lateral folds; ostium bursae near caudal margin of eighth ster-
num; apophyses posteriores longer than apophyses anteriores.

Friseria is nearest to Rifseria and Sriferia. The genera may be
separated as indicated under Rifseria.

Some species of Friseria have maculation very similar to that of
Parastega Meyrick species, and the venation of neither is diagnostic;
however, the male genitalia of Parastega are characterized by having
asymmetrical, single lobed valvae and a long uncus which lacks
enlarged setae on the caudal margin. The signum of Parastega is as
for Gelechia.

The known species of Friseria occur in the Southwest and Mexico;
thus, the genus may have evolved in the Mexican highlands.

NO. 3647 NEARCTIC GELECHIIDAE—HODGES 53

Busck (1939) proposed Friseria for cockerelli, lindenella, malindella,
sarcochlora, repentina, and fuscotaeniaella. Of these lindenella, malin-
della, and sarcochlora are synonyms of cockerelli, and fuscotaeniaella is
removed to Rifseria, new genus; F. infracta (Walsingham) and
lacticaput (Walsingham) are transferred from Gelechia, and acaciella
(Busck) is transferred from “Telphusa.’’ 'Two new species, nona and
caveta, are described, bringing the number of recognized species to
seven.

Key to Species of Friseria
BASED ON MACULATION

1. Forewing white overlaid with black; two patches of raised scales on fold, one

dorsad of fold, and one at end of cell. . . . . . .. caieta, new Sue
Forewing brown, no patches of raised scales. . . .2

2. Forewing dark purplish brown basally, followed aaa an vopnace sitar or put
fascia running from costa to half way between fold anddorsum. ... .5
Forewing pale brown basally, if dark jared not followed by pale colored
Tasclaee ie FA ey es ae ee ey eh ee ae

3. Forewing with rmddie third Pamala places SEA ed 2tROE REG!
Forewing varying shades of brown, never with a precel piace panel at mid-

Glee esa eas . . .. cockerelli (Busck)

4. Forewing with an Wenn date Brown ee ag one-third, bordered basally
by orange scales; outer third of costal margin uniformly dark with black
tipped, gray scales... . . . . repentina (Walsingham)

Forewing usually lacking bomapieee! dank prow fascia at one-third; costal
margin pale orange brown with dark Bee; ie scales usually tipped with

white, preceded by black... . . . Nona, new species

5. Head and basal half of thorax eaitecale pat : _ lacticaput (Walsingham)
Head and thorax mottled brown. . . Rei ies rds fee te

6. Apex of white fascia attenuated (fig. 52) . Sst ic, laws Taeaciella (Busck)
Apex of white fascia not attenuated (fig. 54) . . . . infracta (Walsingham)

BASED ON MALE GENITALIA

ihmCuiciiulayspinedey {pics times Au. Rotke coors Gaed aot 2
Culcitula bare... . Maa Scockerelli (Basco

2. Culcitula large, ene fepeeaity pecan peel of gnathos, culcitular setae
more than twice medial height of culcitula (fig.112)......... .38

Culcitula small, not extending laterally beyond base of gnathos; culcitular
setae not extending laterally beyond base of gnathos; culcitular setae less
than one-fourth medial height of culcitula (fig.118)... . sll ee

3. Medial lobe of valva with distal margin convex, apex iEuneate (fig. 116).
nona, new species

Medial lobe of valva with distal margin sinuate, apex acute (fig. 122).

repentina (Walsingham)
4. Costal lobe of valva with a lobe from ventral surface at one-half its length,

valva appearing quadrilobate (fig. 117)... . . . . acaciella (Busck)
Costal lobe of valva lacking such a lobe, valva appearing trilobate (fig. 118) . 5
5. Costal lobe of valva with a ventral flange on distal half. . .. . “uhO

Costal lobe of valva without a ventral flange on distal half (fig. 120).
lacticaput (Walsingham)

54 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

6. Lobes from ventral margin of valva more than one-half length of valva

figs UTS) ak ei BN RO aed ees nae caieta, new species
Lobes from ventral margin of valva less than one-fourth length of valva
(Gig EED) Sc Ne ers cteeewcr tema. tatia ta! rah rsy ae wee fan gare infracta (Walsingham)

Friseria cockerelli (Busck)

Ficurss 46, 47, 107-110, 162, 229, 230

Gelechia cockerelli Busck, 1903, Proc. U.S. Nat. Mus., vol. 25, p. 871.
Friseria cockerelli.—Busck, 1939, Proc. U.S. Nat. Mus., vol. 86, p. 573.
Gelechia lindenella Busck, 1903, Proc. U.S. Nat. Mus., vol. 25, p. 876. [New
synonymy.]
Gelechia malindella Busck, 1910, Proc. Ent. Soc. Washington, vol. 11, p. 179.
[New synonymy.]

Gelechia sarcochlora Meyrick, 1929, Exot. Micr., vol. 3, p. 491. [Newsynonymy.]

Maculation: as in figures 46 and 47. Head: tongue mixed gray
white and pale brown; maxillary palpus creamy white; outer surface
of first segment of labial palpus gray brown (scales pale basally),
inner surface white, outer surface of second segment gray brown on
basal half, mixed white and gray brown distally, inner surface white
with some gray-brown scales near apex, third segment mixed gray
brown and pale gray, posterior surface nearly white; dorsal surface
of scape of antenna dark brown, ventral surface buff, individual scales
of dorsal surface of shaft ocher basally, brown distally, ventral surface
pale ocher; frons and lower vertex creamy white, a few dark brown
scales before anterior margin of eye; scales of upper vertex and
occiput pale buff, apices pale gray brown. Thorax: mixed brown and
buff. Forewing: buff, brown, gray brown, and red brown. Hind-
wing: fuscous, cilia pale buff. Prothoracic leg: coxa mixed creamy
white and brown, apex creamy white; femur mixed gray white and
brown with a purple cast at certain angles of light incidence; tibia
brown with a buff fascia at base of epiphysis and at apex, scale bases
pale; tarsus brown, base and apex of basitarsus and apex of second
segment pale buff. Mesothoracic leg: coxa and trochanter creamy
white; femur mixed gray white and brown; tibia brown, scale bases
pale, a scale tuft at middle, apex of scale tuft and of tibia pale buff;
tarsus brown, base and apex of basitarsus and apex of second segment
pale buff. Metathoracic leg: coxa creamy white basally, gray
distally; trochanter creamy white; femur mixed white and brown;
tibia mixed creamy white, brown, and pale brown, tuft pale buff,
outer tibial spurs brown; tarsus brown and creamy white, apices of
first four or all segments pale. Abdomen: mottled gray brown, ventral
surface with several gray-white scales. Alar expanse: 12-17 mm.
Male genitalia: as in figures 107-110 (RWH slides 1706 and 3224).
Female genitalia: as in figure 162 (RWH slide 3225).

Food plant: Prosopis spp. Leaf feeder.

NO. 3547 NEARCTIC GELECHIIDAE—HODGES 55

Types: cockerelli, female, USNM; lindenella, male, USNM; mal-
indella, male, USNM;; and sarcochlora, male, BMNH.
Specimens examined:

UNITED STATES: Arizona: Ajo, 5c’, 19, Mar. 1-30 and May 1-15, O.C.
Poling, MCZ, USNM;; Baboquivari Mts., Pima Co., 107, 119, Apr. 23-Sept. 15,
Poling, Sperry, Snow (AB slide June 16, 1937), AMNH, MCZ, USNM;; Chiri-
cahua Mts., 19, Sept. 22, 1927, J. Kusche, CAS; Fairbanks, San Pedro River,
19, Sept. 6, 1927, J. Kusche, CAS; Fish Creek, Tonto Nat. Forest, 1, May 9-10,
1918, J.C. Bradley, CU; Ft. Grant, Pinaleno Mts., 3¢7, 19, July 15-19, 1917,
J.C. Bradley, CU; Hot Springs, 1¢, 19, June 26, H.S. Barber, USNM;; Kensley,
Arivaca Rd., 167, May 6, 1958, ex mesquite leaves, Stephenson and Alexander
(JFGC slide 1111), USNM; Madera Canyon, Santa Rita Mtns., 4000-5600 ft.,
136, 379, July 1—-Oct. 26, 1959, R.W. Hodges (RWH slides 1710, 1717, and
3223-3225), CU, USNM; Mohave Co., 267, 19, May 1-15, USNM;; Organ Pipe
Cactus Nat. Monument, 19, Aug. 31, 1961, D. Rentz, CAS; Paradise, Cochise
Co., 2c’, March and July, USNM; Pefia Blanca Canyon, Santa Cruz Co., 2607,
489, Aug. 7-Sept. 1, 1959, R.W. Hodges (RWH slides 1706-1709), CU,
USNM;; Phoenix, 207, 29, Kunze, USNM; 30 mi. E Quijotoa, Pima Co., 107,
Aug. 28-29, 1927, J.C. Bradley, CU; Ramsey Canyon, Huachuca Mts., 19,
Sept. 1-2, 1927, J.C. Bradley, CU; San Carlos, 2‘, May 12-13, 1918, J.C.
Bradley, CU; Sells P.O., Indian Oasis, Pima Co., 20¢, 189, Apr. 1-30, 1923,
O.C. Poling (AB slides Aug. 11, 1934, Sept. 28, 1936, and June 11, 1937), MCZ,
USNM;; Wellton, Yuma Co., 4<7, May 5-6, 1918, J.C. Bradley, CU; no further
locality, 4c, 29, Morrison, BMNH. Cattrornia: Chino Canyon, Palm
Springs, 16’, Apr. 19, 1950, E.C. Johnston, CNC; Cronise Valley, San Bernardino
Co., 1c", Apr. 29, 1956, J. Powell, UCB; Dixieland, Imperial Co., 27, Apr. 1-30,
1922, O.C. Poling, USNM;; Indio, 22, May 1, 1918, J.C. Bradley, CU; La Puerta,
1¢, 19, July 1911, Wright and Field, USNM; Mason Valley, 1<, Aug. 10,
1938, C. M. Dammers, USNM; Mirage Lake, San Bernardino Co., 207, 19,
Apr. 18, 1935, C. Dammers, MCZ, USNM; Needles, 19, Apr. 1-6, 1918, J.C.
Bradley, CU; Palm Springs, 3<7, 1-6, 1921, K.R. Coolidge, USNM; Surprise
Canyon, Panamint Mts., Inyo Co., 1c’, 19, Apr. 24, 1957, J. Powell, UCB;
Thermal, 100 ft. below tide, 1, Aug. 17-18, 1927, CU; 29 Palms, San Bernardino
Co., 29, Apr. 20-21, 1950, E.C. Johnston, CNC. Nevapa: Charleston Mts.,
Kyle Canyon, Clarke Co., 19, Apr. 26, 1950, E.C. Johnston, CNC; Clarke Co.,
20, Apr. 24-30, USNM. New Mexico: Bent, 19, 7.27, USNM; Carlsbad,
Otero Co., 14%, 39, May 17, 1950, E.C. Johnston, CNC, USNM; 10 mi. E
Deming, 26’, July 12, 1917, J.C. Bradley, CU; Mesilla, 2¢, 89, C.N. Ainslie
(AB slides Aug. 12, 1934 and Sept. 24, 1934), USNM; Mesilla Park (mesquite
near), 50’, 39, July 12, 1917, J.C. Bradley, CU; Otero Co., 2¢°, 39, May 10,
1950, E.C. Johnston, CNC, USNM; White City, Eddy Co., 2,89, May 14-17,
1950, E.C. Johnston, CNC, USNM. Texas: Alpine, Brewster Co., 19, May 22,
1950, E.C. Johnston, CNC; Big Bend Nat. Park, Brewster Co., 1c, 39,
June 20, 1950, E.C. Johnston, CNC; Bosque Co., 1, May 27, 1876, BMNH;
Brownsville, 3o’, 59, Feb. 27 through June, Barber, Freeman, and Townsend,
CNC, MCZ, USNM; Burnett Co., 1, September, F.G. Sharpp, USNM;
Corpus Christi, 1@, June 17, 1943, W.M. Gordon, CU; Cotulla, 1¢, May 12,
1906, Crawford and Pratt, USNM; Fedora, 5’, Mar. 31 and Apr. 1, MCZ;
Fever, 19, Apr. 1, 1897, USNM; Ft. Davis, Jeff Davis Co., 10, 19, May 20,
1950, E.C. Johnston, CNC; Kerrville, 14, F.C. Pratt, USNM; Limpia Canyon,
Jeff Davis Co., 1¢,19, May 20, 1950, E.C. Johnston, CNC; Marathon, Brewster
Co., 1o@, 39, May 23, 1950, E.C. Johnston, CNC; Odessa, Ector Co., 1c,

56 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

May 30, 1950, E.C. Johnston, CNC; Pecos, Reeves Co., 2°, 179, May 18 and
June 2, 1950, E.C. Johnston, CNC, USNM;; Sabinal, 37, Feb. 26 and Mar. 14,
1910, F.C. Pratt (AB slide Sept. 30, 1937), USNM; San Antonio, 14, May
20-22, 1918, J.C. Bradley, CU; San Benito, 1c’, 29, August and Sept. 8-15,
USNM.

MEXICO: Basa Cauirornta: Chapala Dry Lake, 19, Sept. 25, 1941,
Ross and Bohart, CAS; El Marmol, 19, Sept. 24, 1941, Ross, and Bohart, CAS.
Hipauco: 3 mi. E Zimapan, 6400 ft., 2oc°7, 49, July 31-Aug. 1, 1963, Duck-
worth and Davis, USNM. Nvegvo Leon: 3 mi. E Galeana, 5000 ft., 1¢,
19, Aug. 7-9, 1963, Duckworth and Davis, USNM. Tamauuipas: 4 mi.
SW Ciudad Victoria, 1100 ft., 19, July 10, 1963, Duckworth and Davis, USNM;
6 mi. S Ciudad Victoria, 1050 ft., 1c’, 19, Aug. 6, 1963, Duckworth and Davis,
USNM.

Discussion: F. cockerelli may be readily separated from the other
species of Friseria by the buff to dark brown color of the forewings and
by the lack of fasciae. . caveta appears blue white and black, the
other species have a complete or partial oblique fascia at one-third
or one-fourth on the forewing.

The various synonyms of cockerelli reflect the species variability
in maculation. Meyrick’s sarcochlora is a pale form; the three Busck
names cover pale, medium, and dark forms. In a given locality all
gradations between nearly immaculate buff to dark brown specimens
are likely to occur. Neither the male nor female genitalia reflect the
maculational differences.

Friseria caieta, new species
Ficures 48, 111, 113, 124, 163, 231, 232

Maculation: as in figure 48. Head: tongue white with a few
brown flecks; maxillary palpus white, a few apical scales tipped with
brown; first segment of labial palpus white with a dorsal brown
saddle, second segment white with an incomplete brown band near
base and a brown band on outer and anterior surfaces before apex,
third segment white with a brown ring at one-fourth and a brown
ring before apex; dorsal surface of scape of antenna brown with white
scales subbasally and at apex, ventral surface white; shaft mixed
brown and white or gray basally, becoming mainly brown distally;
frons white; vertex and occiput white, some scales with brown apices.
Thorax: white, several scales on tegula and middle of mesothorax
dark brown tipped. Forewing: white, dark brown, and orange
brown, many brown scales with gray bases. Hindwing: fuscous,
cilia pale buff. Prothoracic leg: anterior surface of coxa mixed
white and brown, femur and tibia brown with some white scale
bases, apex of tibia and a fascia at base of epiphysis white; tarsus
brown, apices of first three segments white. Mesothoracic leg:
coxa and femur mainly white with some brown scales; tibia mainly

NO. 3547 NEARCTIC GELECHIIDAE—HODGES 57

brown, scale bases white, apex and medial tuft white; tarsus brown,
apices of first three segments white. Metathoracic leg: coxa and
trochanter white with some gray brown scales; femur white with
brown scales; tibia brown (scale bases pale), a subbasal white fascia,
another at base of first pair of spurs, apex white; base and apex of
basitarsus and apices of second and third segments white. Abdomen:
mottled gray brown, buff, and gray, ventral surface paler than dor-
sal surface. Alar expanse: 12-14mm. Male genitalia: as in figures
111, 113, and 124 (RWH slide 3219). Female genitalia: as in figure
163 (RWH slide 1238).

Food plant: Unknown.

Type: Male, Madera Canyon, 4880 ft., Santa Rita Mtns., Aug. 23,
1959, R.W. Hodges, CU type 4224.

Paratypes:

ARIZONA: same locality as type, 890’, 869, July 11-Sept. 3, 1959, R.W.
Hodges (RWH slides 1235-1238 and 3219; RWH wing slides 61 and 62), AMNH,
BMNH, CAS, CNC, CU, UCB, USNM;; same locality except for elevation, 4000

ft.. 2c, 59, Aug. 9, 1959, R.W. Hodges, CU, USNM; Pefia Blanca Canyon,
Santa Cruz Co., 907, 159, Aug. 7-Sept. 1, 1959, R.W. Hodges, CU, USNM,

Other specimens examined:

Arizona: Baboquivari Mts., Pima Co., 1c, 4?, June 1-Oct. 30, O.C. Poling,
USNM; Madera Canyon, 4400 ft., Santa Rita Mts., Pima Co., 14, 19, June 8
and 9, 1963, J.G. Franclemont, JGF; Paradise, Cochise Co., 1 ? , June 1-7, USNM.
Texas: Alpine, Brewster Co., 19, May 22, 1950, E. C. Johnston, CNC; Brewster
Co., 7000 ft., 19, 4.26, BMNH; Ft. Davis, Jeff Davis Co., 29, May 20, 1950,
E. C. Johnston, CNC.

Discussion: F’. caieta may be quickly separated from the other
species of Friseria by the presence of white scales over most of the
forewings and head.

Friseria repentina (Walsingham)
Figures 49, 119, 122, 123, 167, 233, 234

Gelechia repentina Walsingham, 1911, in Godman and Salvin, Biologia Centrali-
Americana, vol. 42 (Lepidoptera-Heterocera, vol. 4) p. 65.
Friseria repentina.—Busck, 1939, Proc. U.S. Nat. Mus., vol. 86, p. 573.
Maculation: as in figure 49. Head: tongue pale buff and pale
brown; maxillary palpus orange; outer surface of first segment of
labial palpus orange with a few gray-brown scales, inner surface pale
buff, second segment orange with a brown ring (incomplete posteriorly)
at one-third and another at three-fourths, third segment orange with
a brown ring at one-third and another at two-thirds; dorsal surface of
scape of antenna brown, an orange band at one-fourth, apex orange;
frons pale gray; vertex and occiput orange, most scales with orange
brown apices. Thorax: mottled orange brown, brown, and orange;

58 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

apex of tegula mainly orange; disk mainly brown. Forewing: orange,
red brown, brown, gray brown, and buff; gray-brown scales with purple
reflections. Hindwing: fuscous, cilia buff. Prothoracic leg: coxa and
femur pale orange heavily dusted with brown; tibia brown with a
faint subbasal orange band, another at base of epiphysis, apex orange;
tarsus brown, base and apex of basitarsus and apices of second and
fifth segments buff. Mesothoracic leg: coxa mainly pale buff, a row
of brown-tipped scales before apex; femur buff mottled with brown;
tibia brown with subbasal, medial, and apical pale buff fasciae; tarsus
brown, base and apex of basitarsus and apices of second and fifth
segments pale buff. Metathoracic leg: coxa buff and gray white with
some gray-brown splotches; femur buff flecked with brown, brown
fasciae at one-fourth and two-thirds; tibia pale brown (scale bases pale)
with pale buff to buff white fasciae at one-fifth, middle, and apex,
dorsal tuft pale buff, outer spurs pale basally, brown distally; tarsus
brown, base and apex of basitarsus and apices of remaining segments
pale buff. Abdomen: not available. Alar expanse: 13-15 mm.
Male genitalia: as in figures 119, 122, and 123 (AB slide June 10, 1937).
Female genitalia: as in figure 167 (RWH slide 3477).

Food plant: Unknown.

Type: Female, BMNH.

Specimens examined:

MEXICO: Guerrero: Amula, 6000 ft., 1d, 29, Sept. 18, H. H. Smith
(AB slide June 10, 1937; RWH slide 3477; RWH wing slide 84), BMNH, USNM.

Discussion: F’. repentina is close to nona but may be separated on
maculation as indicated in the key. Unobserved variation may occur
in the maculation; however, only three specimens are available for
examination. The male genitalic differences as pointed out in the
key and shown in the figures are diagnostic.

All of the specimens with the facies of repentina and nona from
Arizona have been nona. Further collecting in northern and central
Mexico has not turned up additional specimens of either, but this may
be a result of each being a fall-emerging species.

Friseria nona, new species
FicureEs 50, 112, 116, 121, 166, 235, 236

Maculation: as in figure 50. Coloration much as for repentina
with the exception that the orange of repentina is generally replaced
with buff. Abdomen: mottled buff and gray brown, becoming greasy.
Alar expanse: 10.0-13.5 mm. Male genitalia: as in figures 112,
116, and 121 (RWH slide 3220). Female genitalia: as in figure 166
(RWH slide 3222).

Food plant: Unknown.

NO. 3547 NEARCTIC GELECHIIDAE—HODGES 59

Type: Male, Madera Canyon, 4880 ft., Santa Rita Mtns., Ariz.,
Oct. 25, 1959, R. W. Hodges, CU type 4225.

Paratypes:

ARIZONA: same locality as type, 4400-5600 ft., 9567, 479, June 26—Oct. 26,
1959, R. W. Hodges (RWH slides 712, 713, 3220-3222, and 3237-3244; RWH
wing slides 63 and 64), AMNH, BMNH, CAS, CNC, CU, USNM;; Pefia Blanca
Canyon, Santa Cruz Co., 50°, 59, Aug. 7-Sept. 1, 1959, R. W. Hodges (RWH
slides 3245 and 3246), CU, USNM.

Other specimens examined:

Arizona: Aravaipa Canyon, Pinal Co., 19, May 21, 1953, Dietrich, CU;
Baboquivari Mts., 6c°7, 992, Apr. 15-July 15, 1924, O. C. Poling, USNM;; Chiri-
cahua Mts., near Portal, 19, July 5, 1939, A. F. Braun, AFB; Patagonia, 19,
May 26, 1953, Dietrich, CU; Pinal Mts., 2°, July and August, 1900, Kunze,
USNM;; Santa Catalina Mts., Pima Co., 19, July 24-31, USNM.

Discussion: See the comments under repentina for means of
separating the two species.

Friseria lacticaput (Walsingham), new combination

Figures 51, 120, 126, 164, 237, 238

Gelechia lacticaput Walsingham, 1911, in Godman and Salvin, Biologia Centrali-
Americana, vol. 42 (Lepidoptera-Heterocera, vol. 4), p. 61.

Maculation: as in figure 51. Head: tongue buff with a few pale
brown-tipped scales basally; maxillary palpus pale buff basally,
darker distally; first segment of labial palpus mottled buff and pale
brown, inner surface pale buff, second segment pale buff and orange
with incomplete pale brown rings at two-fifths and three-fourths,
posterior surface nearly white, third segment pale buff with a brown
ring at two-fifths and another at four-fifths; dorsal surface of scape
orange with a pale brown medial band, ventral surface pale buff,
shaft buff on first eight or nine segments then mainly brown; frons,
vertex, and occiput buff, many scales tipped with orange. Thorax:
orange on basal half, apex of tegula and a line across mesothorax pale
buff; apex of mesothorax dark brown; metathorax pale buff. Fore-
wing: dark brown, base orange and pale buff, fascia buff white, tornal
area creamy white. Hindwing: fuscous, cilia pale fuscous and buff.
Prothoracic leg: coxa and femur buff white, a few brown scales at
middle of coxa, a brown fascia just beyond middle of femur; tibia
brown and black dorsally; pale buff ventrally, base, part of medial
tuft and apex of apical tuft pale buff; tarsus buff with brown-black
dorsal saddles on first four segments. Mesothoracic leg: much as for
prothoracic leg. Metathoracic leg: coxa, trochanter, and femur buff
white, a few post medial brown scales on femur; tibia buff white, a
small subbasal patch of brown scales, a pale brown streak at one-
fourth, middle of outer tibial spurs dark brown; tarsus mainly buff.

60 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Abdomen: buff white mottled with gray scales on caudal segments.
Alar expanse: 10-16 mm. Male genitalia: as in figures 120 and 126
(AB slide Oct. 14, 1936). Female genitalia: as in figure 164 (RWH
slide 3476).

Food plant: Unknown.

Type: Female, BMNH.

Specimens examined:

MEXICO: GuerrrRERo: Amula, 6000 ft., Sept. 18, H.H. Smith (AB slide
Oct. 14, 19836; RWH wing slide 84), USNM. Yucatan: Mérida, 16, 19,
July 29, 1964, P.J. Spangler (RWH slides 3475 and 3476), USNM.

Discussion: F. lacticaput may be separated from cockerelli, caieta,
repentina, and nona by the presence of the buff white to white costal
fascia at one-third; and from acaciella and infracta by the head and
anterior half of the thorax being buff.

Forewings of the fresh specimens are nearly black with a slight
luster as contrasted with the dark brown of the specimens collected
for the ‘Biologia.’

Friseria acaciella (Busck), new combination

Ficures 52, 53, 117, 118, 125, 165, 239, 240
Telphusa acaciella Busck, 1906, Proc. U.S. Nat. Mus., vol. 30, p. 722.

Maculation: as in figures 52 and 53. Head: tongue mottled buff
and brown, dark basally; maxillary palpus pale brown, flecked with
buff; outer surface of first segment of labial palpus gray brown, inner
and dorsal surfaces buff white, second segment gray brown, apex
dark orange, third segment dark brown (scale bases pale), base and
middle with orange scales, apex buff; dorsal surface of scape of an-
tenna brown, apex pale, ventral surface buff white, first two scale
rows brown, dorsal surface of shaft brown (scale bases pale), ventral
surface brown with each half segment buff; scales of frons buff with
eray brown apices; vertex and occiput with gray brown to brown-
tipped scales, a fascicle of orange scales from dorsal margin of eye.
Thorax: mottled dark brown, brown black, and red brown. Fore-
wing: black, dark gray brown, brown, buff white, and orange. Hind-
wing: fuscous, cilia gray brown tinted with buff. Prothoracic leg:
coxa and femur buff flecked with brown, a transverse fascia on femur
at three-fourths; tibia brown with orange at base of epiphysis, apex,
and at apex of epiphysis; tarsus brown, base and apex of basitarsus,
apex of second and entire fifth segment pale buff. Mesothoracic leg:
much as for prothoracic leg; medial tibial tuft orange basally, dark
brown before apex, gray apically; apex of fourth tarsal segment pale
buff. Metathoracic leg: coxa, trochanter, and femur buff white, a
transverse brown band near base of trochanter, near base of femur,

NO. 3547 NEARCTIC GELECHIIDAE—HODGES 61

and at two-thirds of femur; tibia mainly pale buff, gray brown between
tibial spurs, outer tibial spurs dark gray brown medially, pale buff
basally and apically; apices of all tarsal segments pale buff. Abdo-
men: mottled pale buff and pale brown. Alar expanse: 11.0-12.5 mm.
Male genitalia: as in figures 117, 118, and 125 (RWH slides 3328 and
3330). Female genitalia: as in figure 165 (RWH slide 3331).

Food plant: Acacia farnesiana Willdenow and Mimosa species.

Type: Female, USNM.

Specimens examined:

UNITED STATES: Lovistana: New Orleans, 4c, 19, Oct. 17, 1904,
E.8.G. Titus (RWH slides 3328 and 3332), USNM. Trxas: Brownsville,
7, 129, Jan. 14-July 4, Barber, Piazza (AB slide Sept. 22, 1936; RWH wing
slides 77 and 78), CAS, USNM; Corpus Christi, 1, iss. May 17, 1923, ex Acacia
farnesiana USNM; Los Borregos, Brownsville, 1, June 5, 1904, H.S. Barber,
USNM;; Mercedes, 167, 19, Jan. 2, 1956, P.T. Riherd, H.L. Schamlzied, CPK,
USNM;; Nueces River, 5 mi. SW Mathis, 167, 39, Aug. 12, 1963, Duckworth and
Davis, USNM; Richmond, Brazos River, 6°, 59, June 22, 1917, J.C. Bradley,
CU, San Benito, 6c’, 99, April-Sept. 15 (AB slide Aug. 1, 1933), USNM;
San Diego, 16, May 23, 1895, E.A. Schwarz, USNM;; Victoria, 67, 109, June,
Caudell, Schwarz, Mitchell, Hinds, Bradley, AMNH, CU, USNM.

MEXICO: Coanuita: 10 mi. N Monclova, 1500 ft., 1¢, July 7, 1963, Duck-
worth and Davis, USNM. Hrpaueo: 3 mi. E Zimapan, 6400 ft., 19, July 31-
Aug. 1, 1963, Duckworth and Davis, USNM. Nuevo Leon: Anegade Arroya,
1250 ft., 16 mi. S Linares, 567, 39, July 19, 1963, Duckworth and Davis, USNM;
3 mi. E Galeana, 5000 ft., 19, Aug. 7-9, 1968, Duckworth and Davis (RWH
slide 3329), USNM. Tamauv.ipas: 4 mi. SW Ciudad Victoria, 1200 ft.,1¢7, 59,
Duckworth and Davis (RWH slide 3331), USNM; 6 mi. S Ciudad Victoria,
1050 ft., Aug. 6, 1968, Duckworth and Davis, USNM; El Salto Falls, 2000 ft.,
26 mi. W Antiguo Morelos, 2? , July 11-14, 1963, Duckworth and Davis, USNM.

Discussion: F’. acaciella may be separated from cockerelli, caveta,
repentina, and nona by the fascia from the costa at one-third, from
lacticaput by the dark thorax; and from infracta by the outer margin
of the fascia being irregular, pointing to the tornus, whereas that of
infracta is rounded.

In some specimens the fascia is buff brown; however, no other
characters are correlated with this feature, and gradients between this
condition and that given in the present description exist.

Friseria infracta (Walsingham), new combination

Figures 54, 114, 115, 128, 241, 242

Gelechia infracta Walsingham, 1911, 7n Godman and Salvin, Biologia Centrali-
Americana, vol. 42 (Lepidoptera-Heterocera, vol. 4), p. 61.

Maculation: as in figure 54. Head: tongue missing on examined
specimen; maxillary palpus greasy brown, pale basally; first segment
of labial palpus brown on outer surface, dorsal and inner surfaces
buff white, second segment brown with pale orange to buff white

62 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

bands at one-third, two-thirds, and apex, third segment orange with
a brown band at one-third and another at two-thirds; dorsal surface
of scape of antenna brown with some orange basally and apically,
ventral surface buff, dorsal surface of shaft greasy brown, ventral
surface buff and brown; scales of frons, vertex, and occiput gray brown
apically, pale basally, a fascicle of orange scales above eye. Thorax:
dark gray brown, metathorax buff. Forewing: pale buff, white,
dark gray brown, and black. Hindwing: fuscous, cilia fuscous
tinged with buff. Prothoracic leg: coxa and femur streaked buff
and brown; tibia brown, apex and an incomplete band at base of
epiphysis orange to buff white; tarsus brown, base and apex of
basitarsus and apices of second and fifth tarsal segments buff.
Mesothoracic leg: coxa buff, many scales streaked with gray, tro-
chanter pale buff streaked with brown; femur buff and brown; tibia
brown, apex yellow and white; tarsus brown, base and apex of
basitarsus and apices of second, fourth, and fifth segments pale buff.
Metathoracic leg: coxa and trochanter with some pale brown scales;
femur pale buff streaked with brown; tibia pale brown, a pale subbasal
streak, another at base of first pair of tibial spurs, and apex nearly
white, dorsal tuft streaked buff and buff white; tarsus brown, base of
basitarsus, apices of segments two through four, and all of fifth
segment buff white. Abdomen: not observed before dissection was
made. Alarexpanse: 15mm. Male genitalia: as in figures 114, 115,
and 128 (RWH slide 3461). Female genitalia: no specimens
available.

Food plant: Unknown.

Type: Male, BMNH.

Specimen examined:

MEXICO: Guerrero: Amula, 6000 ft., 17, Sept. 18, H. H. Smith (RWH
slide 3461; RWH wing slide 83), USNM.

Discussion: F. infracta may be separated from cockerelli, caieta,
repentina, and nona by the presence of the costal, white fascia on the
forewing; from lacticaput by the dark head and thorax; and from
acaciella by the rounded apical portion of the fascia. The valvae of
the male genitalia of wnfracta have an extremely heavily sclerotized
costal lobe; the two lobes from the ventral surface are short; whereas
the costal lobe of acaciella is less heavily sclerotized, has an additional
ventrodistal lobe, and one of the lobes from the ventral margin is
more than one-half the length of the costal lobe.

Rifseria, new genus
Type-species: Gelechia fuscotaeniaella Chambers, 1878.
Head: smooth scaled; tongue moderate, scaled to one-half; max-
illary palpus short, folded at base of tongue; labial palpus recurved,

No. 3547 NEARCTIC GELECHIIDAE—HODGES 63

second and third segments subequal in length, second segment slightly
thickened with scales ventrally, third segment slender, apex acute;
antenna two-thirds length of forewing, simple. Forewing: broadly
lanceolate, apex acute; 12 veins present; 6, 7, and 8 stalked, 6 to
dorsum, 7 and 8 to costa. Hindwing: trapezoidal, apex produced,
acute; 8 veins present, 3 and 4 short stalked; 5 from 3+4 at a right
angle; 6 and 7 stalked, 6 to termen, 7 to costa, 7 sinuous. Abdomen:
second and third terga with lateral patches of stout setae. Male
genitalia: valva with two lobes separate to juncture of tezumen and
vinculum, dorsal lobe slender; vinculum broad, quadrate medially;
aedeagus with expanded base; tegumen long; uncus hood shaped, with
an apical row of short, stout setae; gnathos present, hook shaped;
culcitula present. Female genitalia: signum lightly sclerotized,
elliptical, two short ridges near each end; ostium bursae at caudal
margin of eighth sternum; eighth segment heavily sclerotized, anterior
margin a thickened sclerotized ridge; apophyses anteriores short,
apophyses posteriores moderate.

Rifserra is near Friseria and Sriferia but differs from both by
having 6, 7, and 8 of the forewing stalked, terga two and three with
permanent setae, and the apical setae of the uncus short and stout.
It differs from Friseria in that the costal lobes of the valvae are
separate to the base in the males and that the eighth abdominal seg-
ment of the female is heavily sclerotized.

Rifseria fuscotaeniaella (Chambers), new combination

Figures 55, 129, 130, 168, 243, 244

Gelechia fuscotaeniaella Chambers, 1878, Bull. U.S. Geol. Geogr. Surv. Terr.,
vol. 4, p. 89.

Maculation: as in figure 55. Head: tongue and maxillary palpus
brown; first segment of labial palpus brown on outer surface, white
on inner surface, outer surface of second segment brown (scales pale
basally) on basal third or half, white elsewhere, third segment white
except for brown apex and sometimes a brown band at two-thirds;
dorsal surface of antenna brown, ventral surface of scape off white,
ventral surface of shaft greasy pale brown basally; frons with a row
of brown scales from anterior margin of eye to base of tongue, re-
mainder of frons, vertex, and occiput white; a row of brown scales
behind eye. Thorax: white; base of tegula brown, apex sometimes
red brown; apex of mesothorax dark brown; metathorax gray white.
Forewing: white, dark marks brown, some red brown, buff, and pale
gray present; cilia white below apex, buff at tornus. Hindwing:
fuscous, cilia buff. Prothoracic leg: anteromesal surface brown (scales
pale basally), apices of femur, tibia, and tarsal segments buff white.
Mesothoracic leg: coxa mainly white, some scales gray; femur, tibia,

64 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

and tarsus brown (scales pale basally), apices of tibia and tarsal
segments buff white. Metathoracic leg: coxa white basally, gray
white distally; trochanter pale gray brown, scales with light streaks;
femur white dorsally becoming brown ventrally; tibial scales mixed
buff white and pale brown, tuft buff white; apices of first four tarsal
segments off white, outer surface with a brown dorsal saddle. Ab-
domen: buff white dorsally, permanent setae orange; ventral surface
pale buff, segments two through six with a lateral gray-brown patch.
Alar expanse: 10.5-16.0 mm. Male genitalia: as in figures 129 and
130 (RWH slide 1249). Female genitalia: as in figure 168 (RWH
slide 1251).

Food plant: Unknown.

Type: Male, MCZ.

Specimens examined:

UNITED STATES: Arizona: Ajo, Pima Co., 1 9, Mar. 1-15, 1923, O.C.
Poling, USNM; Baboquivari Mts., Pima Co., 1 ?, Oct. 15-30, 1924, O.C. Poling,
USNM;; Fort Valley, 712 mi. NW Flagstaff, 7350 ft., Coconino Co., 1 #, July 17,
1961, R.W. Hodges, USNM;; Hart Prairie, 10 mi. NNW. Flagstaff, 8500 ft., Coco-
nino Co., 1 @, July 12, 1961, R.W. Hodges, USNM; Huachuca, 1 9, Aug. 3, 1899,
MCZ; Madera Canyon, 4880 and 5600 ft., Santa Rita Mtns., 367, 59, Sept. 13-
Oct. 27, 1959, R.W. Hodges (RWH slides 1249-1252), CU, USNM; same locality,
2,19, May 11-16, 1963, J. G. Franclemont, JGF; Molino Basin, Santa Cata-
lina Mts., Pima Co., 19, June 10, 1959, MacNeill, CAS; Palmerlee, 19, USNM;
Paradise, Cochise Co., 2%, June 1-7, USNM; Ramsay Canyon, Huachuca Mts.,
16, July 10-15, 1941, A.B. Klots, AMNH; Redington, 19, USNM; Vail Lake
Road, 9% mi. SE Flagstaff, 6500 ft., Coconino Co., 1¢, 19, July 11, 18, 1961,
R.W. Hodges, USNM. Catirornia: Atascadero, 19, July 26, 1935, E.C.
Johnston, USNM; Avalon, Santa Catalina Is., 1, May 17, 1932, MCZ; 12 mi.
SE Ivanpah, San Bernardino Co., 20°, 19, May 1, 1956, J. Powell, UCB; Los
Angeles, 19, Oct. 26, 1920, K.R. Coolidge (AB slide Sept. 21, 1934), USNM;
Lytle Creek, San Bernardino Co., 19, June 29, 1946, C. Henne (RWH slide
3248), USNM; Mt. Palomar, San Diego Co., 1<, July 18, 1963, J. Powell, UCB;
San Diego, 1<’, Oct. 20, 1920, K.R. Coolidge (AB slide Sept. 21, 1934), USNM;
Walnut Cr., foot Shell Ridge, Contra Costa Co., 1 9, Apr. 22, 1962, J. Powell,
UCB. Cotorapo: Boulder, 19, Cockerell (AB slide June 11, 1937), USNM;
Chimney Gulch, Golden, 19, June 7, 1907, Oslar, USNM; Denver, 19, MCZ;
Glenwood Springs, 26‘, August 1899, MCZ, USNM. Nevapa: Clarke Co.,
19, May 16-23, USNM. Utau: Eureka, 3c’, 19, Aug. 13-22, 1911, Tom
Spalding, USNM; Warner Ranger Sta., 9000 ft., La Sal Mts., 19, July 1933,
A.B. Klots, AMNH.

CANADA: BritisH Cotumsia: Fraser Mills, 1 &, June 22, 1921, L. E. Mar-
mont, USNM. Manrrosa: Aweme, 5 o’, 2 9, Sept. 5-25, N. Criddle (AB slide
June 11, 19837; RWH wing slide 60), CNC, USNM.

Discussion: R. fuscotaeniaella is moderately variable in maculation
with gradations between nearly all white and dark specimens known.

The head, thorax, and left wings remain of the type specimen. Itisa
nearly white representative of the species.

NO. 3647 NEARCTIC GELECHIIDAE—HODGES 65

Sriferia, new genus

Type-species: Gelechia prorepta Meyrick, 1923.

Head: smooth scaled; tongue moderate, scaled to one-half; maxillary
palpus porrect; labial palpus recurved, sickle shaped, second and
third segments subequal in length, second segment thicker than third;
antenna one-half length of forewing, simple. Forewing: broadly
lanceolate, apex almost rounded; 12 veins present; 7 and 8 stalked,
7 to costa. Hindwing: trapezoidal, outer margin gradually moving
to apex, apex slightly produced, broadly acute; 8 veins present;
3 and 4 short stalked; 5 at right angle from 3 and 4; 6 and 7 separate;
7 sinuous; R; merging with Se near base. Male genitalia: saccus
developed, slightly expanded anteriorly; vinculum narrow; valva with
two linear costal lobes, separate to juncture between tegumen and
vinculum; aedeagus stout, basal half expanded; tegumen long, uncus
moderate with apical row of long, stout setae; gnathos hook shaped;
culcitula present. Female genitalia: unknown.

Sriferia may be separated from Friseria by the lobes of the valvae
being separate to the base and by the quadrate saccus and from
Rifseria by the long apical setae on the uncus, lack of permanent
setae on the abdominal terga, and by vein 6 of the forewing being
separate from 7 and 8.

Sriferia prorepta (Meyrick), new combination
Figures 56, 127, 131, 245, 246

Gelechia prorepta Meyrick, 1923, Exot. Micr., vol. 3, p. 19.
Gelechia fulmenella Busck, 1910, Proc. Ent. Soc. Washington, vol. 11, p. 178.
[Preoccupied.]

Maculation: as in figure 56. Head: tongue and maxillary palpus
buff white to white; first segment of labial palpus cream white to
white, second segment cream white, becoming brownish on anterior
surface, third segment brown, posterior surface gray white to cream
white basally; dorsal surface of antenna brown, ventral surface of
scape and base of shaft buff white; frons white; vertex and occiput
gray brown. Thorax: gray brown. Forewing: dark scales gray to
purple gray, tipped with brown; fascia white; cilia fuscous, Hind-
wing: fuscous, cilia gray brown. Prothoracic leg; coxa white basally
and apically, gray brown medially; femur, tibia, and tarsus brown,
apices of femur, tibia, and first two tarsal segments white. Meso-
thoracic leg: coxa and femur white, femur gray apically; tibia and
tarsus gray brown, apices of tibia and first two tarsal segments white
or pale. Metathoracic leg: coxa, trochanter, and femur white with
some pale gray scales; tibia and tarsus dark gray to gray brown, a
white fascia at base of first pair of tibial spurs, extending into dorsal

66 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

tuft, gray; apices of first two tarsal segments white or pale. Abdomen:
eray, apices of segments pale, first, second, and third terga with
orange scales. Alar expanse: 11.0-14.5 mm. Male genitalia: as in
figures 127 and 131 (RWH_ slide 2624). Female genitalia: no speci-
mens available.

Food plant: Unknown.

Type: Male, USNM.

Specimens examined:

Arizona: Mohave Co., 2c’, May 1-7 (RWH wing slide 59), USNM.
CaLiFoRNIA: Essex, San Bernardino Co., 26’, Apr. 10, 1936, C. Dammers (RWH
slide 2621), USNM; Split Rock Tank, Mohave Desert, 1h, May 20, 1938,
Sperry, AMNH.

Discussion: No females of prorepta are known; thus, the genus
cannot be diagnosed for this sex.

References

Buscx, AuGust
1903. A revision of the American moths of the family Gelechiidae, with
descriptions of new species. Proc. U.S. Nat. Mus., vol. 25, pp.
767-938.
1939. Restriction of the genus Gelechia (Lepidoptera: Gelechiidae), with
descriptions of new genera. Proc. U.S. Nat. Mus., vol. 86, pp.
563-591.
CiaARKE, J. F. GATES
1955, 1963, 1965. Catalogue of the type specimens of Microlepidoptera in
the British Museum (Natural History) described by Edward
Meyrick, vols. 2, 4, and 5.
JANSE VAG Jee
1949-1964. The moths of South Africa, vols. 5 and 6.
SATTLER, KLAus
1960. Generische Gruppierung der europdischen Arten der Sammelgattung
Gelechia (Lepidoptera, Gelechiidae). Deutsche Ent. Zeitschr., new
series, vol. 7, p. 10-118.

U.S. GOVERNMENT PRINTING OFFICE: 1966

PLATES

(Figures 1-246)

PROC. U.S. NAT. MUS. VOL. 119 HODGES—PLATE 1

Figures 1-8.—Left wings: 1, Lita virgella, Golden, Colorado; 2, L. solutella, Europe; 3,
L. variabilis, Ft. Valley, Flagstaff, Ariz.; 4, L. variabilis, Satus Creek, Yakima Co.,
Wash.; 5, L. variabilis, Ft. Valley, Flagstaff, Ariz.; 6, L. variabilis, Monachee Meadows,
Tulare Co., Calif., 7, L. variabilis, Hart Prairie, Flagstaff, Ariz., 8, L. barnesiella, Pine,
Gila Co., Ariz.

PROG. U.S. NAT. MUS. VOL. 119 HODGES—PLATE 2

i

Ficures 9-16.—Left wings: 9, Lita incicur, Yellowstone Nat. Park, Madison Junction,
Wyo.; 10, L. incicur, Smokey Valley, Tulare Co., Calif.; 11, Z. inctcur, near White Mt.
Peak, Mono Co., Calif.; 12, L. incicur, Yellowstone Nat. Park, Madison Junction, Wyo.; |
13, L. incicur, Willow Creek, Siskiyou Co., Calif.; 14, L. geniata, La Puerta Valley,
Calif.; 15, L. puertella, La Puerta Valley, Calif.; 16, LZ. puertella, La Puerta Valley, Calif. |

PROC. U.S. NAT. MUS. VOL. 119 HODGES—PLATE 3

24 \

Ficures 17-24.—Left wings: 17, Lita deoia, Smokey Valley, Tulare Co., Calif.; 18, £. dialis,
Brewster Co., Tex.; 19, L. pagella, Jacumba, Calif.; 20, L. pagella, Ft. Valley, Flagstaff,
Ariz.; 21, L. obnubila, Ft. Davis, Tex.; 22, L. maenadis, Placer Co., Calif.; 23, L. veledae,
Dixieland, Imperial Co., Calif.; 24, L. rectistrigella, Durango, Colo.

PROC. U.S. NAT. MUS. VOL. 119 HODGES—PLATE 4

32

Figures 25—32.—Left wings: 25, Lita sironae, San Diego, Calif.; 26, L. thaliae, Glenwood
Springs, Colo.; 27, L. thaliae, Eureka, Utah; 28, 29, L. texanella, Galeana, Nuevo Leon,
Mexico; 30, L. recens, Mt. Shasta, Calif.; 31, LZ. princeps, Jacumba, Calif.; 32, L. jubata,
Satus Creek, Yakima Co., Wash

PROC. U.S. NAT. MUS. VOL. 119 HODGES—PLATE 5

Ficures 33-40.—Left wings: 33, Lita nefrens, Sonora Pass, Tuolumne Co., Calif.; 34,
L. nefrens, Lamar, Colo.; 35, L. invariabilis, Gallup, New Mex.; 36, Arla tenuicornis,
Brooks Memorial Park, Klickitat Co., Wash.; 37, 4. diversella, La Mesa, San Diego Co.,
Calif.; 38, NV. snellenella, Huachuca Mts., Ariz.; 39, Neodactylota liguritrix, Corpus Christi,
Tex.; 40, NV. basilica, Sycamore Canyon, Santz Cruz Co., Ariz.

PROC. U.S. NAT. MUS. VOL. 119 HODGES—PLATE 6

Ficures 41-48.—Left wings: 41, Neodactylota egena, Madera Canyon, Santa Rita Mts,
Ariz.; 42, Eudactylota barberella, Ft. Valley, Flagstaff, Ariz.; 43, E. tobapta, Pena Blanca
Canyon, Santa Cruz Co., Ariz.; 44, E. diadota, Madera Canyon, Santa Rita Mts., Ariz.;
45, E. abstemia, Madera Canyon, Santa Rita Mts., Ariz.; 46, Friseria cockerelli, Pena
Blanca Canyon, Santa Cruz Co., Ariz.; 47, F. cockerelli, Madera Canyon, Santa Rita
Mts., Ariz.; 48, F. caieta, Pena Blanca Canyon, Santa Cruz Co., Ariz.

PROC. U.S NAT. MUS. VOL. 119

HODGES—PLATE 7

Ficures 49-56.—Left wings: 49, Friseria repentina, Amula, Guerrero, Mexico; 50, F.
nona, Madera Canyon, Santa Rita Mts., Ariz.; 51, F. lacticaput, Amula, Guerrero, Mexico;
52, F. acaciella, Corpus Christi, Tex.; 53, F. acaciella, New Orleans, La.; 54, F. infracta,
Amula, Guerrero, Mexico; 55, Rifseria fuscotaentaella, Madera Canyon, Santa Rita
Mts., Ariz.; 56, Sriferia prorepta, Split Rock Tank, Mohave Desert, Calif.

‘ppjapeand “T “Sg <vyoqnt “7 ‘49 <vyoqnt “7 ‘¢9 ‘vpoiued “7 ‘79
Spyjaisaudvg “TJ ‘19 fsuadfau "J ‘Og ‘sijiqviavaur “7 “66 f4no19ut “7 ‘g¢ fsuadfau DT ‘L$ :speay7—'S9-/§ SAUNOIY

HODGES—PLATE 8

PROC. U.S. NAT. MUS. VOL. 119

PROC. U.S. NAT. MUS. VOL. 119 HODGES—PLATE 9

= seme

Bees a
aI r
wits Ee
asks Sein

--

ty
igh

Ficures 66-72.—Male genitalia: 66, Lita virgella; 67, L. solutella, aedeagus; 68-70, L*°

variabilis, aedeagi; 71, L. nefrens, aedeagus; 72, L. rectistrigella, aedeagus.

PROC. UeSe NAT. MUS. VOL. 119

HODGES—PLATE 10

Arian —
2

Figures 73-79.—Aedeagi: 73, Lita sironae; 74, L. thaliae; 75, L. recens; 76, L. texanella;
77, L. jubata; 78, L. incicur; 79, L. invariabilis.

PROC. U.S. NAT. MUS. VOL. 119 HODGES—PLATE 11

Ficures 80-86.—Aedeagi: 80, Lita puertella; 81, L. geniata; 82, L. deoia; 83, L. dialis; 84,
L. pagella; 85, L. barnesiella; 86, L. princeps.

PROC. U.S. NAT. MUS. VOL. 119 HODGES—PLATE 12

Figures 87—92.—Male genitalia: 87, Lita obnubila, aedeagus; 88, L. veledae, aedeagus;
89, Arla tenuicornis, ventral view; 90, 4. tenuicornis, aedeagus; 91, 4. diversella, aedeagus;
92, A. diversella, ventral view.

PROC. U.S. NAT. MUS. VOL. 119 HODGES—PLATE 13

\ py
NU
\ ra

Se

if

fort . ahs

Ficures 93-98.—Male genitalia: 93, Neodactylota snellenella, ventral view; 94, N. basilica,
ventral view; 95, N. snellenella, aedeagus; 96, Eudactylota iobapta, ventral view; 97, E.
tobapta, aedeagus; 98 N. basilica, aedeagus.

PROC. U.S. NAT. MUS. VOL. 119 HODGES—PLATE 14

Ficures 99-103.—Male genitalia: 99, Eudactylota diadota, aedeagus; 100, £. diadota,
ventral view; 101, E£. barberella, aedeagus; 102, Neodactylota liguritrix, ventral view;
103, £. barberella, ventral view.

PROC, U.S. NAT. MUS. VOL. 119 HODGES—PLATE 15

Ficures 104-110.—Male genitalia: 104, Eudactylota abstemia, aedeagus; 105, E. abstemia,
ventral view; 106, Neodactylota liguritrix, ventral view; 107, Friseria cockerelli, aedeagus;
108, F. cockerelli, ventral view of tegumen; 109, F. cockerelli, ventral view of vinculum
and valvae; 110, F. cockerelli, lateral view of valva.

PROC. U.S. NAT. MUS. VOL. 119 HODGES—PLATE 16

Ficures 111-116.—Male genitalia: 111, Friseria caieta, ventral view of tegumen; 112,
F. nona, ventral view of tegumen; 113, F. caieta, ventral view of vinculum and valvae;
114, F. infracta, ventral view of tegumen; 115, F. infracta, ventral view of vinculum and
valvae; 116, F. nona, ventral view of vinculum and valvae.

PROC. U.S. NAT. MUS. VOL. 119 HODGES—PLATE 17

Figures 117—124.—Male genitalia: 117, Friseria acaciella, ventral view of vinculum and
valvae; 118, F. acaciella, ventral view of tegumen; 119, F. repentina, ventral view of
tegumen; 120, F. lacticaput, lateral view of valva; 121, F. nona, aedeagus; 122, F. repentina,
lateral view of valva; 123, F. repentina, aedeagus; 124, F. caieta, aedeagus.

PROC. U.S. NAT. MUS. VOL. 119

HODGES—PLATE 18

Ficures 125-131.—Male genitalia: 125, Friseria acaciella, aedeagus; 126, F. lacticaput,
aedeagus; 127, Sriferia prorepta, aedeagus; 128, F. infracta, aedeagus; 129, Rifseria

fuscotaeniaella, ventral view; 130, R. fuscotaeniaella, aedeagus; 131, S. prorepta, ventral
view.

HODGES—PLATE 19

Ficures 132-136.—Ventral view of female genitalia: 132, Lita dialis; 133, L. maenadis;
134, L. invariabilis; 135, L. obnubila; 136, L. virgella.

PROC. U.S. NAT. MUS. VOL. 119

HODGES—PLATE 20

Ficures 137-140.—Ventral view of female genitalia: 137, Lita incicur; 138, L. nefrens; 139,
L. barnesiella; 140, L. incicur.

—

PROC. U.S. NAT. MUS. VOL. 119 HODGES—PLATE 21

Ficures 141-145.—Ventral view of female genitalia: 141, Lita jubata; 142, L. sironae; 143,
L. solutella; 144, L. rectistrigella; 145, L. pagella.

PROC. U.S. NAT. MUS. VOL. 119 HODGES—PLATE 22

Ficures 146-149.—Ventral view of female genitalia: 146, Lita variabilis; 147, L. recens;
148, L. texanella;: 149, L. puertella.

PROC. U.S. NAT. MUS. VOL. 119 HODGES—PLATE 23

Ficures 150-154.—Ventral view of female genitalia: 150, Lita thaliae; 151, L. variabilis;
152, Arla tenuicornis; 153, L. princeps; 154, A. diversella.

PROC. U.S. NAT. MUS. VOL. 119 HODGES—PLATE 24

Ficures 155-157.—Ventral view of female genitalia: 155, Neodactylota snellenella; 156,
N. egena; 157, N. basilica.

PROC. U.S. NAT. MUS. VOL. 119 HODGES—PLATE 25

158

Ficures 158-159.—Ventral view of female genitalia: 158, Eudactylota iobapta; 159, £.
diadota.

PROC. U.S. NAT. MUS. VOL. 119

HODGES—PLATE 26

Freures 160-162.—Ventral view of female genitalia: 160, Eudactylota abstemia; 161, E.
barberella; 162, Friseria cockerelli.

PROC. U.S. NAT. MUS. VOL. 119 HODGES—PLATE 27

Ficures 163-165.—Ventral view of female genitalia: 163, Friseria caieta; 164, F. lacticaput;
165, F. acaciella,

PROC. U.S. NAT. MUS. VOL. 119 HODGES—PLATE 28

Ficures 166-168.—Ventral view of female genitalia: 166, Friseria nona; 167, F. repentina;
168, Rifseria fuscotaeniaella.

PROC. U.S. NAT. MUS. VOL. 119 HODGES—PLATE 29

/ — Yi
169 ion 170 \ Sct I72
eee eee |
| —— a — =
aT ee 7 _
/ - —-

173 [ 174 eee cor ( I76
ee, = ——— | \ ae ee
ENS | ——~ | aie |

\ .
| ~ J V
| a a LE
. 3 / \ i . . LE ee
/ / x ,
/ \ / \ Vi \
177 Wie = Sites ( 180 \
|
J
ee i NC
Si NG i

_

178
. \ V/A

4

/ oN

A xe Z BX n= on

iB! ( 182 \ 183 ( Is4

mann iw ann

\
y pom e pe i
Va < / \ :
a 7 186 - 187 | ([ 188 _
ON aN ee 2 )
/ =, \ ie ~ . . ae : ,

er ie aes —— rea

\89 190 \ jeniola i ee
yaaa 5 sa — y a

| YN <— : es =

2 ee |
193 4 194 | [ 195 \ / 196

Ny / LET — . = =
v ) SS ae 7 Se! :

197 I98 199 200
\ gS y a
nx [ =e Cea

Figures 169-200.—Eighth abdominal segment (odd numbers=terga, even numbers =
sterna): 169, 170, Lita virgella; 171, 172, L. solutella; 173, 174, L. variabilis; 175, 176,
L. barnesiellas 177, 178, L. incicur; 179, 180, L. geniata; 181, 182, L. puertella; 183, 184,
L. deoia; 185, 186, L. dialis; 187, 188, L. pagella; 189, 190, L. obnubila; 191, 192, L. veledae;
193, 194, L. rectistrigella; 195, 196, L. sironae; 197, 198, L. thaliae; 199, 200, L. texanella.

PROC. U.S. NAT. MUS. VOL. 119 HODGES—PLATE 30

y 203 204
aN Ul a \ Sy SSS)
v \ Ne ee ae a oy)
sae pe me wae SS
| 205 206 eer 208
|
ee as cans J Ree
209 210 2 212
———— =——— ie ‘ /
— is Sf oan }
\ ay
213 214 \ { 215 / 216
; a ges | ~
hONi¢ f 218 219 220 \
6 . ee ed Se
ee aa ae
22! 222 ee —-
a Ne ee
i.
Oe aN \
225 € 3296 » JeeGy a
: i aes
229 2508) esi) Sese \ 233 234
ie — ; | ; Ser oe j / ee

Ficures 201-234.—EKighth abdominal segment (odd numbers=terga, even numbers=
sterna): 201, 202, Lita recens; 203, 204, L. princeps; 205, 206, L. jubata; 207, 208, L.
nefrens; 209, 210, L. invariabilis; 211, 212, Arla tenuicornis; 213, 214, A. diversella; 215,
216, Neodactylota snellenella; 217, 218, N. liguritrix; 219, 220,..N. basilica; 221,, 222.
peat oe barberella; 223, 224, E. iobapta; 225, 226, E. diadota; 227, 228, E. abstemia;

229, 230, Friserta eee 231, 232, F. caieta; 233, 234, F. repentina.

PROC. U.S. NAT. MUS. VOL. 119 HODGES—PLATE 31

( gues ™ ;
235 \ [| 236 [oar / 238 \
—s NSS ( ) | \
Ca Ss) ONS, | \
Vo pe
/ | J i NN le ss. =
\ ih \ \
239 240 ) 241 / 242
oe
ete \ - Ley aa : K vorere

Ficures 235-246.—Eighth abdominal segment (odd numbers=terga, even numbers=
sterna): 235, 236, Friseria nona; 237, 238, F. lacticaput; 239, 240, F. infracta; 241, 242,
Rifseria fuscotaentaella.

U.S. GOVERNMENT PRINTING OFFICE:1966

Proceedings of

the United States

National Museum
SMITHSONIAN INSTITUTION + WASHINGTON, D.C.

Volume 119 1966 Number 3548

NOTES ON ARADIDAE IN THE U.S. NATIONAL MUSEUM, IV
(HEMIPTERA: HETEROPTERA) !

By Nicwoxias A. KorMiLey ”

For the privilege of studying the unidentified Aradidae in the
collections of the U.S. National Museum, Smithsonian Institution,
I wish to express my sincere appreciation to Dr. J. F. Gates Clarke,
former Chairman and presently Senior Scientist of the Department
of Entomology, to Dr. Richard C. Froeschner, Associate Curator in
Charge, Division of Hemiptera, and to the late Dr. Carl J. Drake,
Honorary Research Associate. The deposition of types of new
species is stated with the descriptions.

In the descriptions, 25 units equal 1 mm. The order of the figures
in the ratios follows the descriptive text leading to the ratios.

Subfamily Aradinae Amyot and Serville, 1843

Except for a few strong flying species of the lugubris group which
occur in the Southern Hemisphere, the subfamily Aradinae is holarctic
in distribution. The subfamily in North America was relatively

1 Previous parts of this same series are: I, Kormilev, 1958, Proc. U.S. Nat.
Mus., vol. 109, no. 3413, pp. 209-222; II, Kormilev, 1960, Journ. New York
Ent. Soc., vol. 68, pp. 36-47; III, Kormilev, 1964, Proc. U.S. Nat. Mus., vol.
115, no. 3483, pp. 245-258.

2 365 Lincoln Place, Brooklyn, New York 11238.

2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

well worked out in the first half of the century by various American
entomologists, particularly by Van Duzee, Parshley, and Usinger.
Parshley’s (1921) key for the species of the genus Aradus Fabricius,
though now more than forty years old, is still the basic aid for identi-
fication of American species. Nevertheless, even today new species
are being found. Three of them are described herewith.

Genus Aradus Fabricius
Aradus Fabricius, 1803, Systema rhyngotorum, p. 116.

Aradus saileri, new species
FicurE 1

Maus.—Elongate ovate, slightly widening backward; body finely
oranulate.

Head shorter than width across eyes (22:26); distance between
eyes shorter than length of the head (17:22). Anterior process
moderately long, slightly tapering anteriorly, tip rounded. Anten-
niferous tubercles short, robust, acute, parallel exteriorly, reaching
middle of antennal segment I. Eyes subpedunculate, kidney-shaped,
almost touching foreborder of pronotum. Lateral tooth of anten-
niferous tubercles absent. Preocular tubercles small, acute; post-
ocular small, slightly blurred. Depressions of vertex deep, curved,
connected posteriorly. Antennae robust, less than twice as long as
length of head (40:22): antennal segment I cylindrical; II clavate;
III cylindrical, as thick as tip of II; IV ovate, thinner than III;
proportions of segments 6:15:11:8. Rostrum reaching foreborder
of mesosternum.

Pronotum less than half as long as width across humeri (20:44).
Anterior border slightly sinuate; anterolateral angles rounded; lateral
borders very finely crenulate, strongly convex, rounded at humeri,
very finely granulate, slightly sinuate anteriorly. Inner carinae
strong, parallel, running from the foreborder to hindborder; middle
carinae much thinner and lower, restricted to posterior two-thirds
of disc, reaching hindborder; outer carinae blurred. Disc densely,
finely granulate.

Scutellum much longer than width at base (27:19); lateral borders
shghtly convex, rounded, moderately raised. Apex narrowly rounded.
Prebasal elevation of disc small. Disc finely granulate.

Hemelytra complete, distinctly wider than pronotum (50:44), apex
of membrane reaching expanded border of genital capsule. Corium
long, reaching hindborder of connexival segment V, its basolateral
border expanded and reflexed; apical border convex exteriorly, sinuate
interiorly; veins raised, finely granulate.

NO. 3548 ARADIDAE—KORMILEV 3

Abdomen longer than width across segment VI; connexivum wide,
finely granulate; postero-exterior angles II to V not protruding, VI
slightly protruding, VII produced backward, rounded apically.

Cotor.—Black; posterior borders of connexiva, inner borders of
expanded genital lobes, and tips of femora, white or whitish; remainder
of femora and the tibiae dark brown.

MasurEMEnNts.—Total length 5.0 mm.; width of pronotum 1.76
mm.; width of abdomen 2.24 mm.

Hotorypr.—Male, Riverside Tanana R., Alaska, Lindroth collec-
tor, June 29, 1958. Bears label: ‘“‘Avadus n.sp. nr duzeei det. R.I.
Sailer’ (USNM type 67580).

It is a pleasure to dedicate this species to Dr. Reece I. Sailer who
first identified it as a new species.

Aradus saileri, new species, is related to A. montanus Bergroth but
differs from it by the following characteristics: head is much shorter
than its width across eyes; antenniferous tubercles without lateral
tooth; connexivum bicolored; and size much smaller.

Aradus ovatus, new species
FIGuREs 2, 3

Frmatze.—Brachypterous; ovate, regularly tapering forward, less so
backward; densely and finely granulate.

Head as long as width across eyes (25:25.5); distance between eyes
longer than length of antennal segment II (17:12). Anterior process
long, parallel-sided, reaching basal fourth of antennal segment IT.
Antenniferous tubercles conical, acute, directed a little downward;
outer borders parallel, without tooth. Preocular tubercles rather
obsolete, postocular absent; depressions of vertex shallow, wide, not
connected posteriorly. Eyes globose, very prominent, but not
stalked. Antennae short, less than one and a half times as long as
head, (32:25), slightly narrower than fore-femora (4:5); antennal seg-
ment I ovate, II clavate, III tapering toward base, IV obovate;
proportions of segments 5:12:8:7. Rostrum short, reaching front
coxae.

Pronotum one-third as long as width across humeri (14:41). Ante-
rior border deeply sinuate, finely crenulate; anterior angles rounded;
lateral borders finely crenulate, slightly reflexed, straight and con-
vergent anteriorly, arcuate posteriorly; hindborder deeply sinuate.
Disc rather flat, densely granulate; inner carinae thin, finely granulate,
slightly convergent backward; middle and outer carinae obsolete.

Scutellum triangular, slightly longer than width at base (20:18);
lateral borders very slightly reflexed at apical half; apex narrowly
rounded; disc uneven, densely granulate, slightly depressed at two-
fifths its length and before tip.

4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Hemelytra abbreviated, wider than pronotum; membrane reduced
to a narrow rim at tip of corium interiorly; clavus not discernible;
exocorium strongly developed, outer border arcuate, finely crenulate;
posterolateral borders of main corium simple, not crenulate.

Abdomen ovate, slightly longer than width across segment V
(70:67). Lateral borders finely crenulate, widely rounded; postero-
exterior angles IT to V not protruding, VI slightly protruding, VII
almost forming a right angle. Connexivum wide, densely granulate,
its limit with tergum rather blurred. Tergum flat, slightly convex
medially; limit between terga VII and VIII clearly defined medially,
almost indistinguishable laterally. Lobes of VIII large, pincer-
shaped, surrounding small tergum IX, but not contiguous at tips.
Tergum IX reaches middle of lobes.

Cotor.—Testaceous; occiput, and postero-exterior angles of con-
nexiva II to VII, pale reddish brown, almost whitish; outer borders of
connexival segments in front of postero-exterior angles brown; head
and pronotum ferruginous.

MEASUREMENTS.—Total length 5.2 mm.; width of pronotum 1.64
mm.; width of abdomen 2.68 mm.

Houotypr.—Female, Onaga, Kans., Crevecoeur collector (USNM
type 67581).

Aradus ovatus, new species, is related to A. montanus Bergroth but
may be separated by the following characteristics: antennal segment
III two-thirds as long as IJ; middle and outer carinae of the pronotum
obsolete; pronotum widest far behind middle; size much smaller;
and by color.

Aradus barberi, new species
FIGURE 4

Ma.zr.—Elongate ovate, macropterous.

Head slightly longer than width across eyes (25:23); distance
between the eyes less than length of antennal segment II (15:22.5).
Anterior process long, slightly enlarged medially, produced beyond
tip of antennal segment J. Antenniferous tubercles dentiform,
acute, divergent, lateral tooth distinct. Eyes very prominent,
kidney shaped. Preocular tubercles small, acute; postocular tuber-
cles prominent, blunt. Depressions of vertex deep, slightly convergent
backward, connected posteriorly. Antennae long, slender, less than
twice as long as head (47:25). Proportions of antennal segments
5.5:22.5:9:10. Rostrum reaching to foreborder of mesosternum.

Pronotum less than half as long as width across humeri (19:43).
Anterior border straight, anterolateral angles produced forward,
subangular. Lateral borders sinuate and reflexed anteriorly, strongly

NO. 3548 ARADIDAE—KORMILEV 5

convex, rounded posteriorly. Hindborder sinuate. Interlobal de-
pression shallow, distinct; inner carinae thin, parallel between them-
selves; middle carinae convergent on forelobe, parallel on the hindlobe;
outer carinae short, robust, parallel.

Scutellum triangular, longer than basal width (23:18). Lateral
borders slightly arcuate, rimmed, reflexed; tip narrowly rounded;
disc slightly raised medially, densely granulate anteriorly and around
elevation.

Hemelytra reaching lobes of genital capsule; exterior border of
corium slightly arcuate, almost straight; hemelytra as wide as pro-
notum; clavus longer than scutellum; corium reaching hindborder of
connexival segment IV; membrane large.

Abdomen elongate ovate, longer than width across segment IV
(63:46); lateral borders evenly, widely rounded; postero-exterior
angles not protruding; VII forms an acute angle. Connexivum nar-
row anteriorly, widening posteriorly; lobes of genital capsule large,
contiguous at their tips.

Cotor.—Greyish brown to sepia; corium and membrane variegated
with whitish spots. Each lobe of genital capsule with a large, whitish
spot. Antennae sepia, tips of segments II and III whitish.

MeEasurEMENTsS.—Total length 5.32 mm.; width of pronotum 1.72
mm.; width of abdomen 1.84 mm.

Hototryre.—Male, Evergreen, Colo., collector not known. Col-
lected on Pinus ponderosa. Bears a label: ‘“Aradus n.sp. near
marginatus Uhl. det H. G. Barber’? (USNM type 67582).

ParaTYPEs.—Five males, deposited in the same collection and in
the collection of the author.

This species is dedicated to the memory of the late Mr. H. G.
Barber, an eminent American hemipterologist, who first identified it
as a new species.

Aradus barberi, new species, runs in Parshley’s key to A. marginatus
Uhler, as Barber has indicated. It differs by having antennal seg-
ment II one and one-half times as long as the distance between eyes,
while in A. marginatus these two measurements are equal.

Subfamily Calisiinae Stal, 1873
Genus Calisiopsis Champion

Calisiopsis Champion, 1898, in Godman and Salvin, Biologia Centrali-Americana,
Rhynchota: Heteroptera, vol. 2, p. 67.

The genus Calisiopsis Champion has had only three species: two
from Central America, and one from southeast Brazil. Among speci-
mens examined was one from Finca Florida, Valle Tablones, Colombia,
1300 m. that belongs to Calisiopsis. Judging by the shape of the

6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

head and position of the eyes, it belongs to a new species; but because
both of the taxonomically important antennae are missing, I prefer
not to describe it.

Subfamily Aneurinae Douglas and Scott, 1865

Aneurinae contains only two genera: Aneurus Curtis, 1825, with a
worldwide distribution, and Aneuraptera Usinger and Matsuda, 1959,
with a single micropterous species from New Zealand. From the
Western Hemisphere twenty species have been recorded. ‘Two more
are described herewith: one from Texas, the other from Guadeloupe.

Genus Aneurus Curtis

Aneurus Curtis, 1825, British Ent., vol. 2, pl. 86.

Aneurus pygmaeus, new species

Ficures 5-7

Mate.—Elongate ovate, slightly widening backward. Head flat,
almost as long as width across eyes (o7,11:12; 9,12:13). Anterior
process conical, rounded anteriorly, slightly produced over tip of
antennal segment I. Antenniferous tubercles small, acute. Eyes
moderately convex. Postocular tubercles blunt, slightly produced
over outer border of eyes. Vertex with two (1+1) small, ovate
callosities; surface finely, transversely rugose behind them. Antennae
slender; first two segments obovate, last two cylindrical; proportions
of segments o’, 4:3:4:8; 9, 4:3.5:4.5:9.

Pronotum trapezoidal, shorter than width across humeri (4,
10:22.5; 2, 10:25). Collar very fine, slightly sinuate in front; antero-
lateral angles rounded, slightly produced forward beyond collar;
lateral borders of the forelobe straight, convergent; lateral borders of
the hindlobe convex, rounded; both very finely crenulate; hindborder
widely sinuate medially, convex at hindangles. Foredisc with two
(1+1) round, callosities; hinddisc finely striated.

Scutellum semicircular, shorter than width at the base (07, 9.5:14.5;
9, 10:15); base slightly raised; disc concentrically striated.

Hemelytra long, reaching foreborder of tergum VII (<”), or almost
to hindborder of tergum VII (9). Corium reaching two-fifths of
length of scutellum; membrane large, finely wrinkled. Hindwings
reduced, reaching only to hindborder of tergum III.

Abdomen ovate, much longer than width across segment IV (<7,
43:32.5; 9, 48:33). Exterior borders slightly convex, moreso
posteriorly; postero-exterior angles not protruding. Connexiva very
finely granulate at exterior borders. Each connexival segment from
III to VI with two ovate, calloused spots; connexival segment VII

NO. 3548 ARADIDAE—KORMILEV a

with a single spot. Exterior borders of abdomen finely crenulate.
Hypopygium small, ovate, slightly depressed, finely granulate;
paratergites relatively large, flat, reaching tip of hypopygium. In
females, tergum VII deeply sinuate posteriorly for reception of tergum
VIII; segment IX very short and wide (1:5), straight posteriorly;
paratergites small, rounded posteriorly, reaching to tip of segment
IX. Spiracles II lateral; III to V ventral, not visible from above; VI
and VII lateral, visible from above; VIII terminal.

Cotor.— Dark ferruginous, partially blackish; tergum and venter
yellow brown; membrane white, transparent, brownish at the base.

MeEASUREMENTS.—T otal length: @, 3.0 mm.; 9, 3.28 mm.; width
of pronotum; o’, 0.9 mm.; 9, 1.0 mm.; width of abdomen: &, 1.3
mm.; 9, 1.32 mm.

Hototyre.—Male, Aransas Co., Tex., Mar. 31, 1954, D. J. and
J. N. Knull collectors, ex J. L. Lutz collection, USNM collection
(USNM type 67583).

ALLoTryPeE.—Female, collected with holotype; in the same collection.

Paratyre.—One female, Mississippi, Jan. 9, 1946; deposited in
the collection of author.

Aneurus pygmaeus, new species, is related to A. minutus Bergroth,
from which it differs by the following: scutellum regularly semi-
circular, in A. minutus more narrowed at the tip, almost subtriangular ;
paratergites of male larger, and widened toward the tip, in A. minutus
narrower, almost subcylindrical; whitish semitransparent membrane,
in A. minutus brown and not transparent; dark ferruginous body,
in A. minutus yellow brown (I hold the former color as distinctive) ;
in the female, abdomen narrow and long, in A. minutus more ovate;
paratergites are also different.

Aneurus nasutus, new species
FicuRgss 8, 9

Ma.z.—Elongate ovate, shiny; lateral borders of pronotum and
abdomen with semiobliterated granulation.

Head as long as width across eyes (13:13.5); anterior process stout,
tapering forward, rounded apically, reaching tip of antennal segment
I. Antenniferous tubercles small, blunt, convex exteriorly. Eyes
semiglobose, moderately protruding. Postocular tubercles small,
blunt, not reaching outer border of the eyes. Mesad of the latter
two (1+1) large, ovate, glabrous spots. Vertex roughly, transversely
rugose. Antennae slender, less than twice as long as head (24:13);
proportions of antennal segments 5:4:5:10.

Pronotum half as long as width across humeri (14:30). Collar
thin, sinuate anteriorly. Anterolateral angles rounded, neither pro-
duced forward nor sideways; lateral borders of forelobe straight,

8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

strongly convergent; lateral borders of hindlobe parallel, rounded and
convergent anteriorly. Hindborder sinuate medially. Foredise with
four (2+2) large, flat callosities. Hind-disc finely, transversely
rugose; with two (1+1) large, transverse, elongate, glabrous spots
mesad of humeri.

Scutellum short, wide at base (12:21), widely rounded apically;
disc concentrically rugose.

Hemelytra almost reaching hindborder of tergum VII. Corium
very short, reaching middle of scutellum. Membrane large, finely
wrinkled.

Abdomen ovate, longer than width across segment IV (52:39).
Outer borders of connexiva with semiobliterated granulation; hind-
borders of connexival segments glabrous, shiny. Spiracles II lateral
and visible from above; III to V sublateral, but not visible from above;
VI and VII lateral and visible; VIII terminal. Paratergites small,
strongly divergent, not reaching tip of hypopygium. Latter small,
lateral borders convergent; tip rounded; disc moderately convex.

Femora moderately inflated, fusiform.

Cotor.—Dark brown; hindhalf of head, middle of pronotum, and
base of scutellum, yellow brown; membrane black.

MEASUREMENTS.—Total length 3.84 mm.; width of pronotum 1.2
mm.; width of adbomen 1.56 mm.

Ho.totryrr.—Male, Guadeloupe; deposited in Drake collection
(USNM type 67584).

Aneurus nasutus, new species, is not particularly related to any of
Central American species; it appears to be most like A. subdipterous
Burmeister from southeastern Brazil, but is smaller. Antennae are
similar to those of A. tenuis Champion but the shape of the head is
quite different: antenniferous and postocular tubercles small, and
blunt; scutellum of different shape; lateral borders of the pronotum
and abdomen are smoother, not finely crenulate; hypopygium is
larger; and color is different.

Subfamily Carventinae Usinger, 1950

Usinger (1950, p. 176) divided the Mezirinae into two tribes,
Mezirini and Carventini. Later, Usinger and Matsuda (1959a, p. 56)
elevated the latter to a subfamilial rank. All members of the Car-
ventinae are tropical or subtropical, none occur in the holarctic
region. Many of them are apterous or micropterous. The ivory-like
incrustation on the body of some species, as in the genus Prozius,
may accumulate on tufts of hair and produce projections which give
the insect a bizarre appearance.

No. 3548 ARADIDAE—KORMILEV 9

Genus Proxius Stal

Prozius Stal, 1873, Kongl. Svenska Vet.-Akad. Handl., vol. 11, p. 141.

Proxius has had six species which Usinger and Matsuda (1959,
pp. 113-114) placed into three subgenera: two of these, Prozxius
Stal sensu stricto, with a single species and Neoprorius Usinger and
Matsuda, with four species, are American; the third one, Nesoproxius
Usinger and Matsuda, with a single species, is Sumatran. One new
species, belonging to the subgenus Neoprowius, is described herewith.

Proxius (Neoproxius) peruvianus, new species

Ficures 10, 11

FrmaLe.—Elongate ovate; partially incrustated with an ivory-
liked incrustation.

Head longer than width across eyes (20:16), as long as width across
postocular projections (20:20). Anterior process long, parallel-
sided, deeply cleft anteriorly; genae much longer than clypeus,
cylindrical and truncate anteriorly, surpassing the tip of antennal
segment I by one-fifth of their length. Antenniferous tubercles
dentiform, slender, parallel-sided, reaching a little beyond middle
of antennal segment I. Eyes small, semiglobose, moderately pro-
truding. Postocular parts of head produced into large, partially
punctured, triangular processes reaching far beyond outer border
of eyes. Vertex with a punctured elliptical, flattened elevation
running from clypeus to hindborder of head; head produced back-
ward, forming a long “neck.’’ Antennae, clypeus, and two (1+1)
ovate, glabrous spots laterad of the elevated portion of vertex, rostral
atrium, and rostrum, not incrustated; all other parts of head covered
with thick, ivory-like, incrustations. Antennae long and slender,
one and a half times as long as length of the head (32:20). Pro-
portions of antennal segments 6:5:13.5:7.5. Rostrum short, reach-
ing hindborder of narrow, deep, and short rostral groove; latter
closed posteriorly. Rostral atrium closed.

Pronotum subrectangular, shorter than width across humeri (20:35).
Forelobe almost as wide as hindlobe (32:35). Collar small, convex,
foreborder straight. Anterior angles produced forward far beyond
collar as two (1+1) lobes, these rounded interiorly and sinuate
exteriorly. Anterolateral angles subrectangular; lateral borders of
forelobe slightly sinuate, provided with a row of short, erect bristles
projecting through incrustation. Interlobal notch deep and narrow.
Lateral borders of hindlobe short, convex, deeply notched medially.
Hindborder slightly convex medially, posterior angles angularly pro-

219-946—66——2

10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

duced backward. Foredisc flat medially, with a small, relatively high,
median ridge on posterior half of disc; strongly inflated laterally in a
shape of an S with eight (4+4) dots on inner sides, deeply excavated
laterad of these inflations. Collar, and two (1+1) round dots laterad
of the median ridge, without incrustation; other parts of forelobe
heavily incrusted. Hind-disc with two (1+1), L-shaped, high, trans-
verse carinae medially, turned with their bases to each other, and
with four (2+2) short, oblique ridges laterally. Latter produced over
actual lateral border forming lateral teeth of the hindlobe. Hind-dise
without incrustation, except along carinae.

Secutellum subtriangular, shorter than width at base (11:19),
rounded posteriorly. Disc flat, without incrustation except on the
two (1+1) lateral, high, hatchet-shaped ridges with their blades
sideways.

Hemelytra reaching apical third of tergum VII. Corium very
short, outer border carinate, incrusted; membrane transparent, finely
corrugate.

Abdomen ovate, longer than width across segment IV (59:45).
Lateral borders slightly, evenly convex. Connexival segments IJ and
III fused, provided with two longitudinal ridges converging anteriorly.
Other connexival segments rather flat, heavily incrusted; their outer
borders slightly convex from II to IV, angularly produced on hind-
halves from V to VII. Tergum incrusted laterally, posteriorly on
tergum VII; outer margin of incrustation obliquely raised. Parater-
gites small, triangular, incrusted, reaching tip of segment IX. Latter
subtruncate posteriorly. Spiracles very small, from II to IV ventral
but progressively nearing margin; lateral and visible from above
from V to VIII. Venter incrusted only laterally. Sternum incrusted
along borders. Propleura incrusted.

Legs relatively long, slender; femora fusiform; tibiae cylindrical,
slender. Tarsi with arolia.

Cotor.—Under incrustation and on free parts, yellow brown to
brown; incrustation ivory.

MEASUREMENTS.—Total length 4.44 mm.; width of pronotum 1.4
mm.; width of abdomen 2.36 mm.

Ho.otyrr.—Female, Satipo, Peru, P. Paprzycki collector, Apr. 26,
1941 (USNM type 67585).

Proxius peruvianus, new species, is related to P. palliatus Champion
and may be separated from the latter by the following: genae produced
beyond the tip of antennal segment I; absence of excavation on postoc-
ular lobes; median ridge of the pronotum reduced to a very short, thin
carina on the hindhalf of forelobe; different shape of carinae on the
hindlobe of pronotum and scutellum.

NO. 3548 ARADIDAE—-KORMILEV Lit

Genus Kolpodaptera Usinger and Matsuda

Kolpodaptera Usinger and Matsuda, 1959, Classification of the Aradidae, p. 144.
Kolpedaptera minuta, new species

Maue.—Apterous; subovate; from metanotum to abdominal seg-
ment VI lateral borders parallel. Entire body covered with thick
layer of greyish incrustation.

Head shorter than width across eyes (11:12.5). Anterior process
short, robust, sides parallel; genae slightly produced beyond the tip of
clypeus, not contiguous, forming a shallow notch. Antennal segment
I produced by two-thirds its length over tip of anterior process.
Antenniferous tubercles robust, subacute apically, outer borders sub-
parallel. Eyes moderately protruding. Postocular borders conver-
gent backward. Vertex with a thin, triple, longitudinal carina,
laterally with two (1+1) oblique rugae. Antennae strong, almost
twice as long as head (19.5:11); proportions of the antennal segments
7:3:4.5:5. Rostrum reaching hindborder of rostral groove; latter
wide, shallow.

Pronotum much shorter than width across humeri (8:21). Collar
slender. Anterolateral angles together with lateral borders form
a regularly rounded are. Disc with a median carina occupying
posterior three-fifths of disc and running backward across meso- and
metanotum and tergum J, progressively widening, becoming a double
carina on metanotum and tergum I. Pronotum irregularly rugose
laterad of median carina. Mesonotum wider than pronotum (24:21).
Metanotum wider than mesonotum (25:24). Mesonotum laterad of
median carina with four (2+2) larger, and two (1+1) smaller, flat-
tened elevations separated from each other by fine sulci. Mesonotum
rugose along lateral borders. Median carina of mesonotum fused
posteriorly with median portion of metanotum; latter in turn fused
posteriorly with tergum I; all together forming a longitudinally rugose
triangular plate. Laterad of triangular plate metanotum has two
(1+1) rugose elevations.

Abdomen subrectangular, slightly longer than width across seg-
ment II (27.5:26). Tergum I fused with metanotum anteriorly and
with tergum II posteriorly; latter placed at a slightly lower level.
Central dorsal plate consist of terga III to VI, limited by fine sulci.
Median portion of central dorsal plate elevated, forming a median
ridge, laterad of it with pattern of rugae and round callous spots.
Tergum VII raised posteromedially for reception of hypopygium.
Connexivum broad, segments II and III fused, others separated from
each other by fine sulci; disc of each segment bearing round calloused

2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

spots with rugae around them; postero-exterior angles II to V not
protruding; VI makes a step with VII; the latter produced backward
as a triangular process, reaching to tip of hypopygium. Paratergites
cylindrical, directed obliquely up and backward. Hypopygium dorso-
caudal, flattened on disc, with an ovate median elevation on lower
half. Spiracles II to VII lateral, visible from above; those of VIII
terminal.

Cotor.— Yellow brown, partially darker.

MEASUREMENTS.—Total length 2.36 mm.; width of pronotum
0.84 mm.; width of abdomen 1.04 mm.

Hoxuotyee.—Male, Livingston, Guatemala; deposited in Drake
collection (USNM type 67586).

Kolpodaptera minuta, new species, is related to K. panamensis
Usinger and Matsuda from which it may be separated by head shorter
than width across the eyes; eyes less prominent; antenniferous
tubercles relatively shorter; postero-exterior angle of connexival
segment VII also relatively shorter; proportions of antennal segment
14:6:9:10, whereas in K. panamensis they are 15:9:10:14.

Kolpodaptera rugosa, new species

Frema.Le.—Apterous; ovate; thickly covered with greyish incrusta-
tion and accumulated dirt; under incrustation glabrous and shiny.

Head shorter than width across eyes (14:15.5). Anterior process
short and robust, cleft anteriorly, genae longer than clypeus and
pointed, reaching basal one-third of antennal segment I. Anten-
niferous tubercles short, robust, broad at base, subacute apically.
Eyes moderately prominent, with convex facets. Postocular borders
strongly convergent. Vertex with a short longitudinal ridge flanked
by two (1+1) thin carinae, behind them with two (1+1) small, but
prominent tubercles. Antennae strong, more than twice as long as
head (29.5:14); proportions of antennal segments 10:5:7:7.5. Ros-
trum short, reaching hindborder of shallow, wide rostral groove.

Pronotum much shorter than width across humeri (8:28); mesono-
tum wider than pronotum (34:28); metanotum wider than mesonotum
(39:34). Anterolateral angles of the pronotum rounded; lateral
borders divergent backward. Collar thin, smooth. Median carina
starts at middle of disc and fuses poteriorly with triangular plate
formed by median portions of meso-, metanotum, and tergum I.
Laterally pro-, meso-, and metanotum separated from each other by
deep, transverse furrows. Median, triangular plate on meso-, and
metanotum with a double median carina, roughly, longitudinally
rugose laterally. Laterad of the median triangular plate, pro-, meso-,
and metanotum roughly, irregularly rugose. Along lateral borders
pro-, meso-, and metanotum roughly granulate.

NO. 3048 ARADIDAE—-KORMILEV is

Abdomen as long as width across segment IV (42:42.5). Tergum I
completely fused with metanotum (median, triangular plate), and
with tergum IT; latter can be recognized only because it is at a slightly
lower level. Central dorsal plate of the abdomen consists of terga
III to VI, with a slightly elevated median line narrowed on tergum
III, much wider on terga IV and V, and with a double longitudinal
carina on tergum VI; laterad of the median elevation disc has a pattern
of irregular rugae and round callous spots. Tergum VII convex
medially, sloping laterally; VIII very short and wide. Connexivum
wide; discs of connexival segments with a pattern of rough rugae and
eranules; exterior borders of each segment slightly convex, entire
lateral border weakly festooned; postero-exterior angles from IT to
IV not protruding, V slightly protruding, VI forming an obtuse angle,
and VII forming a right angle. Paratergites rounded, produced
backward beyond tip of very short segment IX. Spiracles II to
VIII lateral, visible from above.

Cotor.—Brown; lateral borders and central dorsal plate of the
abdomen partially mottled with black.

MEASUREMENTS.—T otal length 3.24 mm.; width of pronotum 1.12
mm.; width of abdomen 1.56 mm.

Hoxuotyre.—Female, Cayamas, Cuba; deposited in the Drake
collection (USNM type 67587).

Kolpodaptera rugosa, new species, is related to K. prominens Usinger
and Matsuda and may be separated from it mainly by head shorter
than width across the eyes; antennae relatively longer, more than
twice as long as the head (less than twice in K. prominens); and
different proportions of the antennal segments which are 20:10:14:15,
whereas in K. prominens they are 25:11:16:17.

Subfamily Merizinae Oshanin, 19038

Mezirinae is the largest subfamily of the Aradidae. Previously, it
was considered as a family under the name of Dysodiidae Reuter, but,
as Usinger and Matsuda pointed out (1959, p. 54), that would distort
the actual relationship between subfamilies so far included in Dyso-
diidae; therefore, they accepted Aradidae as a single family, consisting
of eight subfamilies, including Mezirinae, and leaving only Termita-
phididae as a second family in the superfamily Aradoidea.

Mezirinae has an almost worldwide distribution. It is absent from
the frigid areas and develops the highest number of genera and species
in the tropical and subtropical zones. Strange as it may seem, from
the entire European and Siberian part of the Palaearctic region there
is recorded only a single species of Mezirinae, Mezira tremulae
(Germar), 1822. The Manchurian subregion of the Palaearctic region
contains several genera and numerous species.

219-946—66——3

14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Genus Miorrhynchus Champion

Miorrhynchus Champion, 1898, in Godman and Salvin, Biologia Centrali-Ameri-
cana, Rhynchota: Heteroptera, vol. 2, p. 75.

The genus Miorrhynchus Champion has contained ten species which
may be separated into two groups: those with spiracles from VI to
VIII lateral and visible from above; and those with only spiracles
VII to VIII lateral and visible from above. Two new species belong-
ing to the first group are described herewith.

Miorrhynchus angulatus, new species

FiGuREsS 16, 17

Ma.zr.—Elongate, widening backward, subtruncate posteriorly,
partially covered with relatively long, incrusted, curled hairs.

Head longer than width across eyes (25:22). Anterior process long
and moderately stout, covered with incrusted hairs which make it
look stouter; reaching basal fourth of antennal segment I. Anten-
niferous tubercles parallel, directed forward. Eyes moderately large,
semiglobose, protruding. Postocular borders straight; postocular
tubercles placed far from eyes, dentiform, directed sideways. Vertex
with two (1+1) rows of parallel, setigerous tubercles, running from
base of clypeus to hindborder of head, two (1+1) pyriform callosities
laterad of each row. Antennae very long, more than three times as
long as head (81:25); first two segments covered with incrusted hairs,
those on first segment longest; apical two segments except for a brush
of erect bristles on tip of IV, naked. Proportions of the antennal
segments: 29:12:30:10. Rostral groove long, deep, closed posteriorly ;
rostrum reaching hindborder of latter.

Pronotum much shorter than width across the humeri (32:50);
forelobe much narrower than hindlobe (36:50); interlobal depression
deep. Collar wide, with strongly sinuate foreborder. Antero-
lateral angles rounded, with a fringe of long, incrusted hairs; lateral
borders slightly convex on the forelobe, more convex on the hindlobe;
hindborder almost straight. Foredisc with two (1+1) round callo-
sities; laterad of them with two (1-+1) ridges covered with incrusted
hairs; hinddise rather flat, covered with remote granulations and
short, incrusted hairs.

Scutellum shorter than width at base (20:28); carinate at all
borders; lateral borders straight; tip pointed; disc rugose, with a low,
median ridge, covered with long, curled, incrusted hairs.

Hemelytra reaching middle of tergum VII. Basolateral borders
of corium reflexed; apical border convex exteriorly, sinuate interiorly;
apical angle rounded, reaching hindborder of connexivum II. Mem-
brane large, veins anastomosed.

NO. 3548 ARADIDAE—KORMILEV 15

Abdomen longer than width across segment VI (90:72.5); lateral
borders feebly convex from II to V; angularly produced on VI and
VII. Exterior borders of connexival segments II to V slightly con-
vex, making lateral borders look festooned. Connexival segment VI
trapezoidal, with postero-exterior angle rounded at tip; VII angu-
larly produced backward as far as the tip of hypopygium. Parater-
gites flat, fusiform, reaching to tip of hypopygium. Latter subcordate,
shorter than maximal width (15:19), divided by a _ transverse
furrow in upper and lower lobes. Upper lobe subtriangular, with a
relatively narrow median ridge; lower lobe semiglobose; both lobes
covered with incrusted, curled hairs. Spiracles from II to V ventral,
placed far from borders; VI to VIII lateral and visible from above.

Cotor.—Ferruginous; base of antennal segment I, antennal
segment III, with exception of tip, and apical half of IV, yellow
brown. Round calloused spots on connexival segments III to VII
yellow.

MrasurEMENTS.—Total length 6.8 mm.; width of pronotum 2.0
mm.; width of abdomen 2.9 mm.

Hototyre.—Male, Callanga, Peru, 1300 m., F. Woytkowski col-
lector, Feb. 13, 1953 (USNM type 67588).

Miorrhynchus angulatus, new species, is related to M. longipes
Champion and may be separated from the latter by the angularly
produced connexivum VJ, which is rounded in M. longipes.

Miorrhynchus undulatus, new species

FiaureEs 18, 19

Maur.—Elongate ovate; partially covered with curled, incrusted
hairs.

Head longer than width across eyes (25:21). Anterior process
moderately stout, covered laterally with incrusted hairs; reaching
basal fourth of antennal segment I. Antenniferous tubercles short,
blunt, slightly divergent. Eyes semiglobose, protruding, but less
than in preceding species. Postocular borders convex; postocular
tubercles spiniform, placed far behind eyes, directed sideways and a
little backward. Vertex with V-shaped row of setigerous tubercles,
and laterad of them with two (1+1) ovate callous spots. Antennae
long, more than three times as long as head (80:25); proportions of
antennal segments 28:12.5:28:11.5. Rostrum reaching hindborder
of rostral groove, latter closed posteriorly.

Pronotum shorter than width across humeri (35:53). Forelobe
much narrower than hindlobe (36:53). Collar wide, slightly sinuate
anteriorly. Anterolateral angles rounded; lateral borders convex on
the forelobe, strongly convex on the hindlobe. Hindborder slightly

16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

convex in the middle. Foredise with two (1+1) ovate calloused
spots, laterad of them with two (1+1) longitudinal ridges, the latter
covered with incrusted, curled hairs. Hind-dise partially rugose
with half obliterated setigerous granules. Humeri raised.

Scutellum shorter than width at base (20:26). All borders
carinate; lateral borders straight, tip pointed. Disc transversely
rugose; median carina tapering backward, covered with dense,
curled, incrustate hairs.

Hemelytra reaching apical third of tergum VII. Basolateral
borders of corium reflected; apical border straight exteriorly, sinuate
interiorly. Apical angle of corium pointed, reaching slightly over
the hindborder of connexival segment II. Membrane large, with
anastomosed veins, thickly covered with whitish incrustation.

Abdomen longer than width across segment IV (90:66); lateral
borders moderately convex. Exterior borders of connexival segments
II to V slightly convex; postero-exterior angles protruding, and
rounded on connexival segments II to IV; subangular but not pro-
truding on VI and VII; VI forms a slightly obtuse angle, with rounded
tip; VII forms a slightly acute angle, directed backward and slightly
sideways, tip rounded reaching to middle of paratergites. Parater-
gites small, clavate, reaching to middle of hypopygium; latter large,
subcordate, with a moderately wide median ridge. Spiracles ventral
on II to V; lateral and visible from above on VI to VIII.

Cotor.—Ferruginous to dark ferruginous; tibiae with a wide
yellow-brown subbasal ring. Connexival segments III to VI each
with two, connexival segment VII with one, round, yellow-brown,
calloused spots.

MzASUREMENTS.—Total length 6.84 mm.; width of pronotum 2.14
mm.; width of abdomen 2.64 mm.

Ho.totyre.—Male, Callanga, Peru, 1300 m., F. Woytkowski col-
lector, Feb. 13, 1953 (USNM type 67589).

Miorrhynchus undulatus, new species, is closely related to M.
champion Kormilev but differs from it by the undulate lateral borders
of abdomen which are not undulate in M. championi.

Genus Placogenis Usinger and Matsuda

Placogenis Usinger and Matsuda, 1959, Classification of the Aradidae, p. 352.
Diphyllonotus Kormilev, 1959, Proc. Ent. Soc. Washington, vol. 61, p. 61.

In a manuscript submitted for publication in 1955, the author
originally proposed the name Diphyllonotus with the single included
species D. explanatus. Unfortunately, the manuscript was lost by
the printer. Thus the second species of the genus, D. brachypterus,
was published in 1956 before the genus was validated. It was not

NO. 3548 ARADIDAE—KORMILEV 7

until 1958 that the author learned the first manuscript was definitely
lost; his redescription was printed in a different magazine in March
1959. Meanwhile, in January 1959, the genus was described by
Usinger and Matsuda under the name of Placogenis; because the lat-
ter has priority, Diphyllonotus went into its synonymy.

The species of Placogenis are generally macropterous, but P.
brachyptera (Kormilev) was recorded as both macropterous and
brachypterous. ‘The species may be best separated by the relative
lengths of antennal segments.

Key to Species of Placogenis

1. Antennal segment I long, produced beyond apex of genae by at least two-fifths

of its length; head much longer than width across the eyes... ... 2

Antennal segment I shorter, produced beyond apex of genae by at most one-

fourth of its length; head as long as width across the eyes. ...... 3

2. Antennal segment I longer than IV (11: 9.2), and much shorter than III

GIST) en Ne roy oh area Ae cockerelli Usinger and Matsuda, 1959
Antennal segment I as long as IV (11:11), and shorter than III (11:13).

explanata (Kormilev), 1959

3. Antennal segment I as long as (10:10, brachypterous form), or slightly longer

than IV (11:10, macropterous form), and shorter than III (10:13).

brachyptera (Kormilev), 1956

Antennal segment I shorter than IV (10:11), and much shorter than III

CLOSES iese ny se icue adie eke cee yoeiel le ey tea ws clarkei, new species

Placogenis clarkei, new species

Figures 12, 13

FEMALE.—Ovate; macropterous.

Head as long as width across eyes (21.5:22). Anterior process
long, parallel-sided, anteriorly cleft; genae much longer than clypeus,
reaching apical fourth of antennal segment I. Antenniferous tubercles
acute, parallel-sided reaching basal fourth of antennal segment I.
Eyes small, semiglobose, protruding. Postocular tubercles small,
acute, reaching outer border of eyes; postocular borders oblique,
converging backward. Vertex with V-shaped rows of small tubercles.
Antennae long, strong, twice as long as head (44:21.5); proportions
of antennal segments 10:8:15:11. Rostrum short, reaching hind-
border of rostral groove, latter open posteriorly.

Pronotum trapezoidal, half as long as width across humeri (22:44).
Collar thin, well separated from the disc. Anterolateral angles
rounded, expanded, slightly reflexed; produced forward far beyond
the collar; lateral borders parallel at hindlobe, convergent anteriorly.
Lateral notch almost imperceptible. Hindborder feebly and widely
sinuate. Foredisc with four (2+2) calloused spots, surrounded by
a row of granules. Hind-disc roughly granulate.

18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Scutellum shorter than width at base (15:22). Lateral borders
carinate, almost straight; tip rounded. Disc inflated, transversely
rugose, provided with a stout median carina, tapering backward.

Hemelytra complete, reaching apical fourth of tergum VII.
Corium short, not reaching hindborder of connexival segment II;
basolateral border straight and reflexed, then flat and convex;
apical border evenly sinuate; apical angle rounded.

Abdomen longer than width across segment IV (72:63). Lateral
borders evenly convex; postero-exterior angles not protruding. Suture
between connexival segments II and III not discernible. Tergum
VII with a transverse carina along the hindborder, and a transverse
sulcus in front of it. Paratergites large, triangular, reaching middle
of segment IX; latter notched posteriorly. Spriacles II to V ventral,
placed far from border; VI to VIII lateral and visible from above.
Female with second valvula longer than second valvifer.

Cotor.—Ochraceous; lateral borders and apex of scutellum, antero-
exterior angles of connexiva II to VII, spots on terga near joints of
connexival segments III and IV, IV and V, V and VI, a spot in middle
of hindborder of tergum VII, and the base of tergum VIII, fuscous
to piceous.

MEASUREMENTS.—Total length 5.28mm.; width of pronotum 1.76
mm.; width of abdomen 2.52 mm.

Ho.iotyre.—Female, Ciudad Universitaria, Tucuman, Argentina,
J.F.G. Clarke collector, Feb. 19, 1959 (USNM type 67590).

It is a pleasure to dedicate this species to the collector, Dr. J. F.
Gates Clarke, Senior Scientist in the Department of Entomology in
the U.S. National Museum, Washington, D.C.

Placogenis clarkei, new species, is related to P. brachyptera (Kormi-
lev) from which it may be separated at once by different proportions
of antennal segments, particularly by the relatively longer segment
es

Genus Notapictinus Usinger and Matsuda

Notapictinus Usinger and Matsuda, 1959, Classification of the Aradidae, p. 361.

The genus Notapictinus was established by Usinger and Matsuda
for reception of the American species lacking the stridulatory appara-
tus and previously referred to the genus Pictinus Stal, 1873.

Usinger and Matsuda originally referred six species to Notapictinus
but the genus is much larger. In my key for separation of the species
of the genus Notapictinus (1964) are included 25 species, three more
are described herewith, and we may expect that further species will
be described before long.

No. 3548 ARADIDAE—-KORMILEV 19

Notapictinus uruguayensis, new species

Figures 14, 15

Fremate.—Elongate ovate, slightly widening backward; macrop-
terous.

Head shorter than width across eyes (16:17.5). Anterior process
moderately stout, reaching almost to apex of antennal segment Ti;
sides slightly convex, tip deeply cleft; genae much longer than clypeus,
with blunt tips. Antenniferous tubercles short, dentiform, divergent.
Eyes large, protruding, their longitudinal diameter longer than anten-
niferous tubercles (5:4). Postocular tubercles slender, dentiform,
reaching as far as, or a little beyond, the outer border of the eyes.
Infraocular carinae low, finely granulate, converging from foreborder
of eyes to union at middle of posterior border of head. Vertex with
M-shaped row of granules, laterad of this with two (1+1) ovate
callosities. Antennae moderately strong, more than one-and-a-half
times as long as head (26:16); proportions of antennal segments, I to
IV: 6:5:7.5:7.5. Rostrum reaches to hindborder of rostral groove,
latter closed posteriorly.

Pronotum trapezoidal, much shorter than width across humeri
(19:36). Forelobe narrower than hindlobe (27:36). Collar tiny,
with a straight foreborder. Anterolateral angles slightly expanded,
rounded, and reflexed, produced beyond the foreborder of collar.
Lateral borders finely crenulate, straight and parallel at humeri,
strongly converging anteriorly. Hindborder slightly convex medially
and slightly sinuate mesad of hindangles. Foredisc with two (1+1)
stout, granulate ridges, convergent anteriorly, laterad of them with
two (1-+1) lower, granulate inflations. Hind-dise roughly granulate.

Scutellum shorter than width at base (12:18). All sides carinate.
Median carina T-shaped, moderately stout, granulate; disc also
granulate.

Hemelytra complete, reaching apical fourth of tergum VII. Corium
reaching middle of connexival segment III, basolateral border reflexed,
forming a low carina, then slightly sinuate, receding, and not re-
flexed. Apical border bisinuate, shallowly exteriorly, and deeper
interiorly. Membrane large and corrugate.

Abdomen longer than width across segment IV (60:45). Lateral
borders evenly convex, rounded; postero-exterior angles barely pro-
truding: VII forming a rounded lobe, reaching to the middle of
tergum VIII. Posterior border of tergum VII carinate. Paratergites
large, triangular, reaching apical fourth of segment 1X. Latter
tricuspidate; median portion acute, lateral (valves) rounded. Spir-

20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

acles from II to VI ventral, placed far from border, VII and VIII
lateral and visible from above.

Cotor.—Testaceous; head behind eyes, posterolateral angles of
pronotum, lateral and basal carinae of scutellum, outer borders of
connexiva, hindborder of tergum VII, entire terga VIII and IX, and
two (1+1) large, dark spots on sterna IIT to V, infuscated. Hind-
borders of connexival segments IT to VI yellow.

MASUREMENTS.—Total length 4.40 mm.; width of pronotum 1.44
mm.; width of abdomen 1.80 mm.

Houotyre.—Female, Atlantide (Maldonado), Uruguay, H.L.
Parker collector, Dec. 24, 1942; deposited in the Drake collection
(USNM type 67591).

Notapictinus uruguayensis, new species, is related to N. incaicus
Kormilev (1964, p. 473), from which it may be separated by the longer
anterior process of the head; much shorter antennae; more produced
anterolateral angles of the pronotum; granulate and not transversely
rugose scutellum; and yellow hindborders of connexival segments.

Notapictinus platyceps, new species
Figures 20, 21

Mauz.—Elongate, with subparallel sides; macropterous.

Head shorter than width across eyes (15:17). Anterior process
short, tapering forward, subtruncate anteriorly, reaching basal third
of antennal segment I; genae as long as clypeus. Antenniferous
tubercles short, dentiform, slightly divergent. Eyes relatively large,
moderately prominent. Postocular borders rounded, without visible
tubercle or tooth, not reaching outer border of eyes. Vertex with
semiobliterated granulation. Entire upper surface of the head flat-
tened. Antennae strong and long, more than twice as long as head
(35:15). Proportions of the antennal segments 9:7:11:8. Rostrum
reaching hindborder of rostral groove; latter wide, closed posteriorly.

Pronotum much shorter than width across humeri (20:37). Fore-
lobe narrower than hindlobe (27:37). Collar sinuate anteriorly,
rather indistinctly separated from the disc. Anterior angles produced
forward far beyond the collar as rounded lobes. Lateral borders
slightly convex, rounded on the forelobe; strongly convex on the
hindlobe. Hindborder slightly convex in the middle. Interlobal
sulcus thin but distinctly visible. Forelobe with two (1+1) almost
obliterated callosities. Hindlobe granulate.

Scutellum shorter than width at base (12:18). All borders cari-
nate; disc roughly, transversely rugose, with a moderately stout
median carina.

Hemelytra reaching apical fourth of tergum VII. Basolateral
border of corium reflexed forming a low carina. Apical angle

NO. 3548 ARADIDAE—KORMILEV Zi

produced into a long point, reaching to middle of connexival segment
III. Apical border of corium deeply and widely sinuate.

Abdomen longer than width across segment V (59:42); sides
subparallel. Connexival segment II short, triangular, semifused
with III; latter very long, more than twice as long as II (15:7);
other connexival segments normal. Postero-exterior angles not pro-
truding on II to IV; slightly protruding on V and VI; rounded on
VII. Hypopygium conical, almost as long as its width at base
(9:10). Paratergites wide, flat, curved, tips reposing on the sides
of hypopygium. Spiracles II to V ventral, placed far from border;
VI to VIII lateral, visible from above.

Cotor.—Head, pronotum, with exception of anterior angles,
scutellum, hindhalves of connexival segments III to VI, middle of
tergum VII, and hypopygium, ferruginous. Antennae, anterior
angles of pronotum, connexival segment II, and paratergites ochre
brown. Forehalves of connexival segments III to VI and almost
entire VII, yellow. Ventral side of the body ferruginous, with
yellow spots on connexivum.

MEASUREMENTS.—Total length 4.36 mm.; width of pronotum
1.48 mm.; width of abdomen 1.68 mm.

Hototyepr.—Male, Chapada, Brazil; September (USNM type
67592).

Notapictinus platyceps, new species, is related to N. breviceps
(Champion) but is larger, has antennae with different proportions
of segments; anterior angles of the pronotum produced forward as
long lobes; and paratergites in the form of a hook.

Notapictinus ornatus, new species

FIGURES 22, 23

Matzt.—Elongate ovate, regularly widening backward, then
narrowing; macropterous.

Head almost as long as width across the eyes (16:16.5). Anterior
process long and slender, cleft anteriorly; genae much longer than
clypeus, reaching apical two-fifths of antennal segment I. Anten-
niferous tubercles long, dentiform, slightly divergent, reaching basal
fourth of antennal segment I. Eyes moderately large, protruding.
Postocular tubercles small, dentiform, slightly produced beyond
outer border of the eyes. Vertex with V-shaped row of granules, and
laterad of this with two (1+1) callosites; latter each with a thin,
longitudinal carina. Antennae strong, twice as long as head (31.5:16).
Proportions of antennal segments 8:5.5:11:7. Rostrum reaching
hindborder of rostral groove; latter wide, closed posteriorly.

Pronotum much shorter than width across humeri (19:36). Collar
wide, slightly sinuate anteriorly, clearly separated from disc. Fore-

22 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

lobe narrower than hindlobe (29:36). Anterior angles rounded,
directed somewhat inward, produced beyond collar. Lateral borders
of forelobe convex, rounded; those of hindlobe less convex, crenulate.
Hindborder slightly convex medially and laterally. Foredise with a
deep median silcus, laterad with two (1+1) large, flat callosities.
Inner borders of callosities carinate, discs each with two oblique
carinae. Four (2+2) curved carinae between callosities and lateral
borders. Hind-dise roughly granulate.

Scutellum shorter than width at base (12:19); all borders carinate;
lateral borders sinuate medially. Median carina thin, but prominent;
disc roughly, transversely rugose.

Hemelytra almost reaching hindborder of tergum VII. Baso-
lateral border of corium reflexed; apical angle produced into a short
point, reaching basal fourth of connexival segment III; apical border
concave, forming two angles.

Abdomen longer than width across segment V (59:44). Connexi-
val segments all of about the same length. Postero-exterior angles
from IV to VI slightly protruding; on VII forming small rounded lobes.
Paratergites small, clavate, reaching middle of hypopygium. Latter
cordate, shorter than width at base (11:13.5); upper surface with a
stout, subtriangular ridge not reaching hindborder of hypopygium.
Spiracles JI to V ventral; VI ventral but placed near margin; VII
to VIII lateral.

Cotor.—Light ferruginous; connexival segments yellow brown,
posterior borders yellow. Paratergites and tibiae ochre brown.
Membrane light brown.

MBASUREMENTS.—Total length 4.32 mm.; width of pronotum
1.44 mm.; width of abdomen 1.76 mm.

Houotyer.—Male, Rio de Janeiro, Brazil; October (USNM type
67593).

Notapictinus ornatus, new species, is related to N. kjellanderi
Kormilev (1964, p. 472) from Peru, but the anterior angles of the
pronotum are directed forward and inward; lateral borders of the hind-
lobe are less convex; postero-exterior angle VII forms small, rounded
lobes; hypopygium more elongate, with sinuate lateral borders;
antennal segment II much shorter than I (5.5:8), whereas in N.
kjellanderi it is as long as I.

References

Kormi.evy, N. A.
1956. Notas sobre Aradidae Neotropicales VI (Hemiptera). Anales Soc.

Cient. Argentina, vol. 162, pp. 148-159.

1964. Notes on Aradidae in the Naturhistorijka Rijksmuseum, Stockholm:
Hemiptera-Heteroptera. Ark. Zool., vol. 16, no. 23, pp. 463-479,
21 figs.

ParsHLey, H. M.
1921. Essay on the American species of Aradus (Hemiptera). Trans.

American Ent. Soc., vol. 47, pp. 1-106, 7 pls.
UsinGER, R. L.
1950. The origin and distribution of the apterous Aradidae (Hemiptera-
Heteroptera). Eighth Intern. Congr. Ent. Stockholm, pp. 174-179.
Usincer, R. L., and Marsupa, R.
1959. Classification of the Aradidae (Hemiptera), vol. 7, 410 pp., 102 figs.
23

24 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Ficures 1-15.—New species: Aradus saileri, male: 1, head and pronotum. A. ovatus,
female: 2, head and pronotum, 3, tip of abdomen. J. barberi, male: 4, head and pro-
notum. Aneurus pygmaeus: 5, male, head, pronotum, and scutellum; 6, male, tip of
abdomen; 7, female, tipofabdomen. A. nasutus, male: 8, head, pronotum, and scutellum;
9, tip of abdomen. Proxius peruvianus, female: 10, head, pronotum and scutellum;
11, tip of abdomen. Placogenis clarket, female: 12, head and pronotum; 13, tip of
abdomen. WNotapictinus uruguayensis, female: 14, head and pronotum; 15, tip of abdomen.

No. 3548 ARADIDAE—-KORMILEV Zo

Ficures 16-23.—New species, male: Muiorrhynchus angulatus: 16, head and pronotum;
17, tip of abdomen. M. undulatus: 18, head and pronotum; 19, tip of abdomen. WNo-
tapictinus platyceps: 20, head and pronotum; 21, tip of abdomen. WN. ornatus: 22,
head and pronotum; 23, tip of abdomen.

U.S. GOVERNMENT PRINTING OFFICE: 1966

Proceedings of

the United States

National Museum

SMITHSONIAN INSTITUTION + WASHINGTON, D.C.

Volume 119 1967 Number 3549

THE FREEZE-DRY PRESERVATION
OF BIOLOGICAL SPECIMENS

By Rottanp O. Hower '
Office of Exhibits

Introduction

Visitors today to the United States National Museum see specimens
of small animals that accurately represent the form and color they
possessed in life. This improved representation is the direct result of
the new method of specimen preservation that has been brought about
by the process of freeze-drying. By this process, the specimen
retains most of its original characteristics without further need for
preservation.

This new technique is based upon sublimation of frozen fluids.
One of the earliest papers detailing the phenomenon of sublimation
was delivered to the Royal Society of London in 1813 by William
Hyde Wollaston, a physicist. Wollaston’s commentary (1813) on
water passing from the frozen to the gaseous state, apparently by-
passing the liquid phase, discussed what was already a well-known
phenomenon. It was not until the 1890’s, however, that the removal

1 As far as is known, Mr. Hower is the first to apply the freeze-dry process to
museum work. Developing more sophisticated apparatus and establishing time
schedules for the freezing of specimens, he is now investigating freeze-dry problems
of color retention, fat stabilization, and the freezing of larger specimens.

1

2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

of fluid from tissue by sublimation at low temperatures had been
accomplished by Dr. Richard Altman of Leipzig (Glick, 1949, p. 4).

Only in recent years, with the advent of modern refrigeration and
vacuum techniques and equipment that is widely available, has the
process become practical for commercial or professional purposes.
The process of freeze-drying, now greatly refined, is used currently
for food processing and for the preservation of pharmaceuticals,
human bone, tissue, and blood plasma.

The freeze-drying techniques utilized at the Smithsonian Institu-
tion for the preservation of biological specimens began in the late
1950’s (Hower, 1962). There is now a well-established freeze-dry
program to preserve many types of natural history specimens for
exhibit. Current exhibits include freeze-dried rodents, reptiles,
crustaceans, insects, and fishes.

Reflecting on events of past years that led to work on the freeze-dry
process, I wish to express my gratitude to Dr. H. T. Meryman of the
Biophysics Division of the Navy Medical Research Institute, who
pioneered many of the techniques and has made himself always
available for consultation.

Special thanks also are extended to James C. Nyce, who contributed
invaluable aid in the preparation of this paper, to Mrs. Constance
Minkin, for her writing contributions, and to John Hackmaster, who
prepared the illustrations.

Principles of the Freeze-dry Concept

Freeze-drying consists bascially of dehydrating tissue while it is
frozen. Whereas some tissue dried from a nonfrozen state becomes
shrunken and consequently distorted in the process, tissue dried from
a frozen state virtually retains its original appearance; thus, the process
is not only valuable in preserving the quality of food but also in
maintaining the appearance of museum specimens. An additional
advantage is that the dehydrated tissue is not subject to decay.

Compared to other preservation methods, the shrinkage and dis-
tortion of freeze-dried museum specimens is minimal, but there is some
distortion in certain types of animals, particularly some fishes. In
most cases, however, the distortion is negligible or correctable. Furry
or feathery specimens and those with exoskeletons appear completely
natural when freeze-dried, provided they have been properly prepared.
Such preparation includes the positioning of the specimen before
dehydration, the actual dehydration, installation of artificial eyes,
and the painting of mucous membranes or exoskeletons (mucous
membranes become white after dehydration and crustaceans lose their
coloration).

NO. 3549 FREEZE-DRY PRESERVATION—HOWER 3

SuBLIMATION.—If biological tissue is first frozen to give it mechanical
rigidity and water is then removed by sublimation, most tissue can be
dehydrated without apparent physical change.

Consider this process in terms of plain frozen water: ice. Each ice
crystal is a geometrically sound structure, made up of myriads of
water molecules that are retained in their positions by the gravitational
field of the surrounding molecules. Within the restricting confines of
this lattice, each molecule moves randomly, and there is a possibility
that one of the motions of a surface molecule will be great enough to
propel it out of the confines of the lattice. The greater the crystal
mass, the greater the probability of such escapes. As temperature is
increased and molecular motion becomes accelerated, the probability
of escape becomes greater; thus, the average number of water molecules
that will escape from a given mass at a given temperature can be
statistically predicted. Ice within a biological specimen behaves in
nearly an identical manner.

Sublimation begins at the outer surface of a specimen and continues
at the boundary between the frozen and the dried tissue. This
boundary recedes toward the center of the specimen as drying pro-
ceeds. As water molecules continue to escape from ice crystals on the
boundary, they move about at high velocity, colliding constantly with
other molecules and with the structure of the dried tissue surrounding
them. (As they are buffeted from collision to collision, they are virtu-
ally independent of external forces.) There is a heavy concentration
of water-vapor molecules at the sublimation boundary, due to the
great number of molecules escaping; consequently, there are more colli-
sions, which ricochet molecules along the line of least resistance toward
the outer shell of the specimen. The force of the molecules’ collisions
following their escape from the ice crystals on the sublimation
boundary supports their drive through the dried tissue of the speci-
men and into the atmosphere beyond its outer shell. For ice to
sublime efficiently, the vapor pressure within the specimen chamber
must be lower than the vapor pressure of ice within the specimen
itself.

VAPOR PRESSURE.—If the temperature of a vacuum chamber con-
taining ice is maintained at —10° C., evacuated with a vacuum pump,
and then valved off so that no external air can enter, molecules will
begin to escape from the ice crystals within the chamber (fig. 1).
Some of these molecules will ricochet about, colliding with one another
and with the sides of the chamber, while others will relocate them-
selves upon other ice crystals.

As the concentration of the vapor formed by these free molecules
reaches a specific point (which is dependent upon the temperatures of
their atmosphere), the rate at which molecules return to the ice will

4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

become equal to the rate at which they depart, and the water vapor
is then said to be in a state of equilibrium (fig. 1). The vapor pres-
sure at which this equilibrium occurs is referred to as equilibrium
vapor pressure. With the temperature of the chamber maintained
at —10° C., the pressure indicated on its vacuum gauge will be 1.950
mm. Hg. This is the equilibrium vapor pressure (consequently the
vapor pressure) of water at —10° C.

Water-vapor molecules may be removed from the area immediately
surrounding the ice in a vacuum chamber, thereby upsetting the
equilibrium and permitting more molecules to escape and allowing
fewer toreturn. This principle applies to biological specimens as well
as to ice.

The most effective method of continuously removing water-vapor
molecules from a specimen chamber is to create a lower vapor pressure
elsewhere. ‘This can best be done by establishing a colder surface
(condenser) nearby. Water vapor will diffuse to a colder surface
and recondense to form new ice crystals.

Ficure 1.—Water-vapor equilib-
rium (4=refrigerated chamber;
B=vacuum gauge; C=cut-off
valve; D=vacuum pump).

A refrigerated condenser serving as an efficient water pump is
employed for this purpose. As previously stated, when the tempera-
ture of a specimen chamber is —10° C., the vapor pressure of ice
within it is 1.950 mm. Hg. If the temperature of the nearby con-
densing surface is —40° C., the vapor pressure is 96 micron Hg.,
creating a vapor-pressure differential of 1.854 mm. Hg. Water
molecules will collect on the cold condenser surface. With this

|

NO. 3549 FREEZE-DRY PRESERVATION—HOWER Oo

much vapor-pressure difference, the only limitation to the effective-
ness of a condenser is the surface area (fig. 2).

PRESSURE REDUCTION.—When vapor transfer occurs at atmospheric
pressure, water molecules are impeded by collisions with air molecules.
At atmospheric pressure, the mean free path (the average distance a
vapor molecule can travel before colliding with another molecule) is
approximately .005 microns (510-® mm.). The mean free path of
a water-vapor molecule with relation to pressure is as follows:

pressure mean free path
10 mm. Hg. 0.0034 mm.
1 mm. Hg. 0.034 mm.
100 » Hg. 0.34 mm.
10 uw Hg. 3.4 mm.
1» Hg. 34.0 mm.

The transfer of water molecules from an ice crystal to a condenser
can be accelerated by increasing the length of their mean free paths;
this can be done by reducing the pressure in the chamber with a

8
ar

Ficure 2,—Water-vapor differential (4=specimen chamber; B=condenser chamber; C=
cut-off valve; D=vacuum pump.

vacuum pump. Not all vapors, however, are condensable; therefore,
a pump must be used to remove these vapors so that chamber pres-
sure will remain low enough to permit efficient removal of the water
to its condenser.

6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119
Instrumentation of the Freeze-dry System

The basic equipment necessary to establish a freeze-dry apparatus
consists of a specimen chamber, a condenser chamber, and a vacuum
pump. The Smithsonian system also incorporates refrigeration con-
trol components that are very desirable for maintaining the depend-
ability of the system.

Figure 3,—Continuous copper tubing (5 in.) connected at alternate ends by return bends
within the specimen chamber.

SPECIMEN CHAMBER.—The basic requirements for the specimen
chamber are that (1) it must be vacuum tight; (2) it must be refrig-
erated; and (3) it must have a large access opening.

The ultimate vacuum within the chamber can be as low as 10
microns (1107? mm. Hg.) of pressure, producing a differential
between internal and external pressure of approximately 15 pounds

NO. 3549 FREEZE-DRY PRESERVATION—HOWER a

per square inch. Sturdy construction is required if a chamber is to
withstand such force.

At the Smithsonian, the structural requirements for the chambers
were met by the selection of a 60-gallon paint-spray pressure tank
for the specimen chamber and a 30-gallon tank for the condenser.
These tanks required modification but proved both satisfactory and
economical.

The Smithsonian specimen chamber was mounted horizontally on
a base shaped to the contour of its walls, with allowance for the
thickness of an insulated outer plastic shell. The door on the Smith-
sonian chamber is two feet in diameter and withstands a total ex-
ternal pressure of 6780 pounds. The chamber itself, two feet in
diameter and 36 inches long, withstands a total external pressure
of a little more than 20 tons.

Figure 4.—Detachable refrigeration assembly: 4, outside view; B, inside view.

The vapor line, a piece of steel tubing with an inside diameter of
3% inches, was welded into a 3%-inch hole midway down the side of
the chamber.

The inside of the chamber was lined with %-inch continuous copper
refrigeration tubing, filimg one end and running the length of the
walls, where it was joined at alternate ends with 2-inch return bends.
The tubing was soldered to the steel wall of the chamber, and a
fillet of metallic plastic compound was added to improve heat con-
duction between wall and tubing (fig. 3).

8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Due to the difference in the coefficients of expansion of the two
materials at operating temperatures, however, fracturing of the steel-
to-copper attachments developed. To overcome this difficulty,
the refrigeration line was soldered to the outer surface of a copper
sleeve, negating the need for attachment between the copper tubing
and the steel wall (fig. 4). This innovation not only solved the frac-
turing problem but also established the refrigeration line as a de-
tachable unit within the system, allowing easy removal from the
chamber for maintenance and repair.

Since the refrigeration line entered and left the chamber through
wo small openings in the rear of the tank, the openings around the
tubing were sealed with soft solder. When installation of the re-
frigeration line was completed, the chamber was tested thoroughly
for vacuum leaks.

REFRIGERATION OF THE SPECIMEN CHAMBER.— Within an evacuated
system, heat is transferred principally by radiation. Radiation
travels in a line-of-sight path between a specimen and surrounding
surfaces. There is no heat produced within the specimen chamber;
therefore, the only appreciable load on the refrigeration system is
ambient heat that leaks in through the insulation. This load can be
calculated in BTU’s per hour from the information given (p. 17).

Any refrigeration compressor (q.v.) serving the specimen chamber
should be capable of producing controlled temperatures at the chamber
walls of —15° to —30° C. (after heat leak has been taken into account).

When selecting a refrigeration compressor, it is desirable to double
the calculated heat load (primarily heat leak) to allow a sufficient
margin for error (table 1).

TaBLE 1.—Refrigeration compressor capacity in BTU’s per hour (based on a 90°
F. ambient temperature; F-12 (freon-12)—Dichlorodifluoromethane; F-—22
(freon-22)—Chlorotrifluoromethane)

Horsepower Refrigerant br OE. —10° F. —25° F. —40° F.
0. 25 F-12 1900 1700 1300 750 350
0233 F-12 2275 2050 1550 950 445
0. 50 F-12 3500 3175 2450 1550 1000
0.75 F-12 5100 4600 3550 2250 1400
il F-12 7200 6500 5100 3350 2300
1 F-22 5590 4350 2890 1920

CoNDENSING CHAMBER.— The physical requirements for the condens-
ing chamber are similar to those of the specimen chamber.

NO. 3549 FREEZE-DRY PRESERVATION—HOWER 9

At the Smithsonian, a length of copper tubing, 3% inches in diameter
(later replaced with steel), was brazed into an opening, 3% inches in
diameter, near the top of the 30-gallon tank. The opposite end of the
tubing was connected to the specimen-chamber vapor line. At the
base of the tank was an opening into which a 1}-inch steel pipe was
welded, connecting it to the vacuum pump.

Inside the condensing chamber, a large complex of coils within coils
was fashioned from 200 feet of copper refrigeration line % inches in
diameter. The expansion valve was mounted inside the chamber and
the refrigeration line was brought into and out of the chamber through
openings in the side of the chamber, which were sealed with soft sol-
der as in the specimen chamber (see fig. 4).

REFRIGERATION OF THE CONDENSER.—The force underlying the
motion of water vapor from the ice, through the specimen chamber to
the refrigerated condenser, is the vapor-pressure difference. This
force is produced by the difference in temperature between the ice
crystals within the specimen and the refrigerated condenser. Table 2

Table 2.—Relationship between temperature (° C) and vapor pressure (mm. Hg.)

aC Vp © Vp °C Vp oO Vp

0 4. 579 —16 | 15132 —34] .1873 — 66 . 00349
ae 4,217 =—17 | 1.031 — 36 L507 — 68 . 00261
care 3. 880 — 18) |" 0: 939 — 38 . 1209 —10 . 00194
cams 3. 568 —19 | 0. 854 — 40 . 0966 S172 . 00143
—4 3. 280 20 30! 406 — 42 . 0768 4 . 00105
ae 3. 013 —21 | 0. 705 — 44 . 0609 — 0 . 00077
a0) 2. 765 —22 | 0. 640 — 46 . 0481 —18 . 00056
—7 2. 537 —23 | 0. 580 —48 | . 0378 — 80 . 00040
ae 2. 326 —24 | 0. 526 — 50 . 0295 — 82 . 00029
ae 2. 131 —25 | 0. 476 —§2 . 0230 — 84 . 00020
—10 1. 950 —26 | 0. 430 —54 . 0178 — 86 . 00014
—11 1. 785 —20 | 0.389 — 56 . 0138 eS . 00010
yy 1, 632 —28 | 0.351 —58 . 0106 a) . 00007
Sales 1. 490 —29 | 0.317 — 60 . 0080 — 94 . 00003
—14 1. 361 — S05!) eooo — 62 . 0061 — 98 . 000015
Se 1, 241 —32 . 2318 —64 | .0046

demonstrates the fixed relationship between temperature and the
vapor pressure of water. The temperature difference between the
specimen and the refrigerated condenser need not be extreme in order
to produce a substantial difference in water-vapor pressure; however,
it should be noted that lowering the condenser temperature produced
less gain logarithmically in vapor pressure difference.

219-947 —66—2

10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

It is clear, therefore, that any effort to produce extremely low
condenser temperature is not economical since the cost per BTU
of refrigeration increases rapidly as temperature is reduced, but the
vapor-pressure differential increases only moderately. A tempera-
ture of —40° C. is satisfactory and is within the range of conventional
freon-12 or freon-22 refrigeration units.

Another factor to be considered in designing a condenser is that
the rate of water loss from the specimen will be very low and, thus,
the heat load on the condenser’s refrigeration unit will be slight.

The heat leak through the insulation of the condenser chamber
should be calculated exactly, as for the specimen chamber.

Temperature Control: Temperature may be controlled by asimple
mercury control switch with a 2- or 3-degree low-temperature differ-
ential. The control employed in the Smithsonian system is a sealed
mercury switch, triggered by a vapor-pressure-actuated Bourbon
tube. Exterior adjustments with visible settings over a calibrated
dial are employed.

Figure 5.—Schematic of Smithsonian freeze-dry apparatus (4=specimen chamber; B=
condensing chamber; C=recompression valve; D=vacuum pump; £, F=refrigeration
compressors; G= vacuum gauge).

Vacuum pump.—The vacuum pump, very important to the freeze-
dry system, should be selected with care. There are many suitable
pumps available.

When a refrigerated condenser is employed as a vapor trap, the
only function of the vacuum pump is to reduce air pressure mechani-
cally within the chamber and evacuate the noncondensable gases
released from the specimens during the drying process.

The capacity of a pump is described in terms of displacement.
Displacement, expressed in liters per second or cubic feet per minute,
is a measure of a pump’s capability to produce a vacuum in a given
time, as well as to keep the system evacuated as gases are released

NO, 3549 FREEZE-DRY PRESERVATION—HOWER iT

from volatile materials within the system. Pump displacement or
pumping speed must be calculated according to the volume of the
system and pressure desired. The time required to pump the system
down to the ultimate pressure (about 10 to 100 microns Hg.) should be
established. The pump-down chart (fig. 6) may be used to calculate

nee aor
1 5 11 13 15 16

SSCS EEE Est oe peed
ie eal

1004

eee 2
tH} +--+} +14}
= SS eS SS SS SS SS SSE SS eS ee

10mm
CTT

Piaf A fof 4 aff Wt alee
= = a

ABSOLUTE PRESSURE

100 mm

1000 mm

Figure 6.—Pump-down factor chart.

displacement requirements. By way of illustration, the following
two examples are cited:

(1) To determine the required pumping displacement to evacuate
a system with a total volume of 15 cubic feet to a pressure of 100
microns in 5 minutes, reference to the pump-down chart shows that
at 100 microns the factor is 10.9. Multiply 10.9 by the volume
(15 cu. ft.) to obtain the total number of cubic feet to be pumped
(163.5). To determine the required pump displacement, divide this
total by the number of minutes (5) allowed for the evacuation.

10.9 X15=163.5

a = 32.7 cu. ft./min. or 55,563.6 liters/sec.

12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

(2) To determine how long it will take a pump with a displace-
ment of 27 cubic feet per minute to evacuate a system with a volume
of 160 cubic feet to a pressure of 200 microns (at 200 microns the
factor is 9.50), calculate as follows:

9.50 160=152

Le = 56.29 min.
Under certain conditions the refrigerated condenser can be
eliminated and water vapor can be removed directly through the
vacuum pump, provided the pump is capable of removing the vapor
at a rate equal to its release from the specimen. To determine
whether the condenser can be omitted, the approximate quantity of
water vapor being released each minute by the specimens within the
chamber must be computed.

If, for example, a specimen chamber contains three squirrels, two
flickers, a toad, and a garter snake (at various drying stages and all
of average weight), the average daily release of water vapor will be
approximately 12 grams.

If 150 microns of pressure were maintained within the chamber, it
would be necessary to pump 200 cubic feet of water vapor for each
eram of water removed from the system, or 2400 cubic feet per day
(1.66 cu. ft./min.). If the vacuum pump cannot handle this volume
of vapor, the pressure within the specimen chamber will slowly rise;
and, as it rises, the volume occupied by a unit weight of water vapor
decreases. At a pressure of 300 microns, two grams of water will
occupy the same volume as would one gram at half that pressure.
It is obvious, therefore, that overloading a chamber or providing an
inadqeuate vacuum pump will cause an increase in operating pressure
in the specimen chamber. To avoid this, a refrigerated condenser
should be used.

Since the differential between the water-vapor pressure within the
chamber and the vapor pressure of the ice within the specimen provides
the driving force for the movement of water vapor through the system,
an increase in pressure within the chamber causes a decrease in
efficiency.

An increase in chamber pressure from 150 to 300 microns with a
specimen temperature of —10° C. would result in a vapor pressure
differential of from 1.80 to 1.65 mm. Hg., or a reduction of a little
more than 8 percent.

While this example demonstrates that increasing water-vapor load
is not as deleterious as expected, it nevertheless establishes a relation-
ship between water-vapor load and vacuum-pump capacity and

PROG. U.S. NAT. MUS., VOL. 119 HOWER—PLATE 1

os

Refrigerating apparatus for the specimen and condenser chambers (Smithsonian photo).

PROC. U.S. NAT. MUS., VOL. 119 HOWER—PLATE 2

Interior of the specimen chamber; dehydration is almost complete (Smithsonian photo).

HOWER—PLATE 3

119

PROC. U.S. NAT. MUS., VOL.

*(oyoyd uvruosyyurg) snqieivddy oy} Jo IoJaWOUTIOYIa]e1 oy Burysoyo Joy ny

7

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HOWER—PLATE 4

119

PROC. U.S. NAT. MUS., VOL.

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NO. 3549 FREEZE-DRY PRESERVATION—HOWER 13

indicates that a vacuum-pump limitation can limit the number of
specimens that can be produced in a reasonable span of time.

A 2-stage gas-ballast vacuum pump is recommended for all freeze-
dry use. Gas ballast on the pump enables it to remove condensable
vapors such as water vapor from a vacuum chamber with a minimum
of internal condensation.

In the gas ballast, a valve provides a means for controlling the entry
of air into the exhaust stage of the pump’s operation. The ballast of
air acts as a transporting medium carrying diluted vapors through the
exhaust part to the atmosphere.

Such pumps are generally rated with ultimate pressure ranges from
1 to 10 microns; however, pumping speed usually begins to decrease
at about 100 microns, which is considered a satisfactory pressure for
freeze-drying.

Oil should be changed frequently in any pump used in a freeze-dry
system. Even though the pump is gas-ballasted, a certain amount of
water vapor will contaminate the oil and damage the pump.

TaBLe 3.— Vacuum table

Terminology Pressure Range
Ultrahigh vacuum pressures below 1X10-§ mm. Hg. or .001 mi-
crons Hg.
High vacuum Ei — tx 10 mmakig:
Fine vacuum 1 mm. Hg. 1X10-3 or 1 micron Hg.
Rough vacuum 760 mm. Hg. to 1 mm. Hg.
1 mm. Hg. 1000 microns
.00132 atmosphere
.01934 p.s.i.
.0393 in. Hg.
1 micron .001 mm. Hg.
1X10-3 mm. Hg.
Atmospheric pressure 760 mm. Hg.
14.7 p.s.i.
29.921 in. Hg.

The possibility of future expansion of the freeze-dry system should
also be considered when selecting a vacuum pump. Under certain
conditions the addition of a second chamber may require doubled
pumping capacity (see tables 3-5).

VACUUM-LINE DIMENSIONS.—The diameter of vacuum lines affects
the efficiency of a freeze-dry system since a pump is effective only
if the vacuum lines are large enough to handle the vapor transfer.

Conductance (the volume of flow of vapor through a line) is a
well-investigated function, and there are a variety of ways to calcu-

14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 119

TABLE 4.— Metric conversion table

Aino" 002.5400 em. 025.4000 mm.

latte 000.3048 m. 000.0304 cm.

1 in.2 006.4520 cm.2 645.2000 mm.2

1 ft.? 929.0000 em.2

1 in.3 016.3870 em.3 016.3870 ml.3

1 ft.3 000.0283 m.3 000.2830 cm.
.283 ml.

10 mm. 1 cm. 0.3937 in.

010 em. 1 dm. 3.9370 in.

100 mm.?2 1 em.2

100 em.2 1 dm.?

010 ml. 1 el. 0.3380 fl. oz.

010 el. eae 6.1025 cu. in.

010 dl. ime 1.0567 fl. qt.

010 1. 2.64 gal.

ee

TaBLE 5.—Conversion multipliers (e.g., liters per second times the factor equals
cubic feet per minute)

ee ee

From To Factor
a ee
liters per second cubie feet per minute 02.1200
liters per minute cubic feet per minute 00.0353
liters per second liters per minute 60.0000
cuhic centimeter per second liters per second 00.0010
cubic feet per minute liters per minute 28.3200

eS ee eee

late the vacuum-line dimensions required to permit adequate
conductance.

The flow of fluids (or vapors) within a vacuum system is usually
either molecular flow or viscous flow. Molecular flow occurs at
pressures at which the molecule’s mean free path is greater than the
average diameter of the tubing through which it flows.

Since the mean free path of a molecule does not approach the
diameter of any large vacuum line at above 1-micron pressure, we
are concerned only with viscous flow.

It should be noted that restriction of tubing to the viscous flow of
vapors is significant only when the volumes to be moved are very
large or the pressure differential is small. The law that expresses the
viscous flow of fluids through a tube was first deduced by Jean L. M.
Poiseuille, the French physicist (Daniels et al, 1949, p. 71). This law
establishes relationships between the coefficient of viscosity (vis-
cosity=poises/dyne sec./em.’), the volume of the fluid flowing through

No. 3549 FREEZE-DRY PRESERVATION—HOWER 15)

the whole section of the tube in unit time (seconds, as applied here),
the pressure differential at each end of the tube, the radius, and the
length of the tube.

V=Volume (cm.°/sec.)

p= Pressure (dynes/cm.’) Te pr*
r = Radius (in em.) Sr
£L=Length (in cm.)

The formula states that the volume of vapor transferred through a
tube in a specific period of time is proportional to the fourth power of
the tube radius. This means that a small increase in tube diameter
will produce a considerable increase in vapor conductance.

To utilize the formula, it is necessary to know the coefficient of
viscosity, which for air is approximately 1.7 10-‘ poises at a pressure
of 1 dyne/em.*=7.510-* mm. Hg. 1.

For example, to calculate the conductance of vapor through a
tube that is 193 cm. long, 7.62 cm. in diameter, and with a pressure
differential between the specimen and the condensing surface of
2000 microns, the following values are:

V=cm.3/sec.
p=2666.7 dynes/cm.” (2000 pressure differential)
r=3.81 cm.
L=193 cm.
v=1.7X10~* poises
= (3.14) (2666.7) (3.81)* _1737861.723
()G@9s\iC017) im 261248
V=66209 cm.?/sec.

TESTING FOR LEAKS.—The overall system must be tested for
vacuum leaks. Leaks in a vacuum system can be detected easily with
a Peroni pressure gauge used in conjunction with acetone. The
vapor pressure of acetone is so high that, when joints or surfaces are
brushed with it, any leak will cause a marked increase of pressure
within the system. The gauge head should be installed at a point
near the vacuum pump and then watched for pressure increases after
the system has been evacuated and tested.

Another test is to inject freon under slight pressure into the system
and locate the leaks with a refrigeration leak detector. When the
system is initially pumped down, the pressure in the system may be
higher than anticipated due to gases being given off by zine coating
or other materials of high vapor pressure that may be inside the sys-
tem. The pressure can be effectively reduced by allowing air to
reenter the system and by pumping it down repeatedly until these
vapors have been evacuated.

16 PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 119

Repairs should be made while the system is evacuated because the
vacuum helps a sealant penetrate a leak. Unless the pressure is
unusually high when the pump is operating, it may be assumed that
a leak is small and can be sealed with glyptol varnish or lacquer
applied by brush. The instant that either material reaches the
vacuum through a pinhole leak, it dries and plugs the leak. If the
pressure remains high, the leak is large and should be plugged with
soft solder.

REFRIGERATION COMPRESSORS.—The major consideration in refrig-
eration is that constant temperature be maintained.

The hermetically sealed refrigeration unit is composed of a motor
and compressor shaft of 1-piece construction. The motor (cooled by
the flow of refrigerant gas) and compressor assembly are within a
gas-tight housing that is welded shut. This method of construction
eliminates the need for certain parts (pulley, belt, compressor fly-
wheel, and compressor seal) found in an open unit and, of course,
avoids the servicing and replacing of those parts.

One objection to this type of unit is that, under freeze-dry operating
conditions, there is some danger of the motor’s burning out because
of the very small amount of refrigerant being circulated.

The Smithsonian’s compressors, however, have not overheated
despite continuous operation for several years in a room with an
average temperature of about 90° F. If overheating should occur, it
could be remedied by installing a water-cooled condenser in the
discharge side of the compressor.

A second type, the air-cooled compressor, usually operates with a
belt and pulley; the motor is in the open, where it is cooled by air
circulating around it.

Operating characteristics of both types of retrigeration compressors
are, otherwise, essentially the same.

The selection of a fractional-horsepower refrigeration compressor
should be based upon calculation of heat load on thermal insulation
(see p. 17 and also table 1).

EXPANSION VALVES.—F or maximum vaporization of the refrigerant,
it is important to select thermostatic expansion valves of correct
capacity. It is equally important that the valves be installed at the
proper locations, since both factors can influence the success or failure
of the entire system.

The thermostatic expansion valve selected for the condensing
chamber should be a type designed to control temperatures below
—40°C.

Although the thermostatic expansion valves in either chamber may
be mounted in any position, they should be installed as near the

NO. 3549 FREEZE-DRY PRESERVATION—HOWER 17

evaporator inlet as possible. The valves in the Smithsonian system
are mounted inside the chambers, where maximum efficiency is gained.

Bulb Location: For satisfactory expansion-valve control, good
thermal contact between the bulb and the suction line is essential.
The bulb, which controls the expansion valve, should be fastened
securely with two metal straps to a clean section of the suction line
inside the chamber. The bulb should be located near the midpoint of
the line around the coil. It should not be near the bottom of the line
because a refrigerant-and-oil mixture is usually present there, which
would result in incorrect control of the expansion valve.

Filter and Drier: These should be installed in the liquid line ahead
of the thermostatic expansion valve.

Sight Glass: Further protection is easily and inexpensively provided
with a sight glass through which the refrigerant level can be deter-
mined by the presence or absence of bubbles in the liquid line. Bub-
bles indicate that the refrigerant level is low.

THERMAL INSULATION.—Insulation material is required to sub-
stantially reduce heat leak, which loads the refrigeration unit. This
material may be glass wool, with a K factor of 0.29, rock wool, with
a K factor of 0.26, compressed cork, with a K factor of 0.30, or other
similar material. The Smithsonian freeze-dry apparatus is insulated
with a foam-in-place plastic (polyurithane). Foam-in-place plastics
range in density from 2 to 10 pounds per cubic foot; they offer K fac-
tors of from 0.02 to 0.24. These plastics can be used without great
difficulty and can be made to conform to any contour. Such proper-
ties make plastic foam an ideal material to use as insulation.

Calculation for the determination of thermal impedence and heat
load is as follows:

K=insulation factor (BTU/hr./ A=Ambient temperature (dif-

sq. ft.) ference between the inside
S=surface area of insulation and outside of chambers in
(sq. ft.) °F)
IT=thickness of insulation (in.)
ES) (0)24 =BTU/per hr.

TEMPERATURE READING AND RECORDING.—Temperatures through-
out the drying process should be carefully watched and accurately
recorded. The record is indispensable for establishing drying times
and for determining ideal temperatures for various types of specimen
material. An automatic temperature-recording device is valuable
for indicating the temperature variations as the refrigeration com-
pressors are cycling, and it may also call attention to problems that
would otherwise go unnoticed, such as a faulty temperature-control
switch.

18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

The system at the Smithsonian incorporates a telethermometer
that utilizes a thermistor probe as a sensing element and provides
temperature readings directly from a temperature scale. The tele-
thermometer’s range is from —40° to 150° C. Both surface and
air probes are used and the probe leads are connected through a
multiple-selector switch to permit convenient temperature reading
at several places.

7

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By
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i
4
8
a
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i
i
I
i
5
i
1
5
1
L

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Ficure 7.—Simple freeze-dry apparatus (4=specimen chamber; B=commercial deep-
freeze; C=recompression valve; D=vacuum pump; E=wood-spacer insert).

ReEcorpER.—Any versatile general-purpose D.C. voltage and cur-
rent recorder may be used to record temperatures continuously.
One with a low-to-high-level range (200 microvolts to 500 volts and
200 millimicroamperes to 100 microamperes) is suitable for freeze-
dry purposes. It is important that the recorder be compatible with
the temperature device in scale range and voltage output.

Construction of a Freeze-dry System

The simplest apparatus suitable for freeze-drying consists of a
commercial deep-freeze containing a small chamber connected to a
gas-ballasted vacuum pump (fig. 7).

NO. 3549 FREEZE-DRY PRESERVATION—HOWER 19

The main requirement for the deep-freeze chest is that it maintain
a constant temperature of —15° to —20°C. The only alteration needed
is removal of its hinges and latch and the insertion of a wood spacer
between the chest and its lid to permit passage of a vapor line between
them. The vapor line must pass through the wood spacer.

The hinges and latch should be relocated for the raised top; the
thickness of the wood spacer is determined by the diameter of the
vapor line that will pass through it.

A small specimen chamber may be made from a 5-gallon paint-spray
pressure tank roughly 18 inches in diameter and 21 inches high. A
hole must be cut in the side of the chamber for a vacuum line, which
can be brazed into place.

The major consideration in selecting a gas-ballasted vacuum pump
is the volume occupied by the water vapor at the temperature and
pressure being used. Assuming that the chamber temperture is
— 20° C., the vapor pressure is 0.8 mm. Hg. Ata vapor pressure of 0.8
mm., one gram of water occupies a volume of 1200 litters. If the
pressure gradient is approximately 0.6 mm., we find one gram of water
will occupy approximately 1400 liters. A gas-ballasted vacuum pump
with a 25-liter-per-minute capacity will take approximately one hour
to pump one gram of water. A pump with a capacity of 79 liters per
minute will remove approximately one gram of water in 18 minutes.

If the vapor line is 150 cm. (5 ft.) long, the diameter is calculated
to be approximately 2.5 cm. or 1 inch. Tubing 1 inches in diameter
will allow a sufficient operating margin when using the larger capacity
pump.

The chamber must not be loaded beyond pumping capacity. Ina
system of this size, it is suggested that a single specimen be placed in
the chamber and a second one added two or three days later, followed
by a third specimen on the completion of the first, and so on, in this
pattern. Observations of drying times during experimental drying
cycles will determine the capacity of each individual system. The
drying cycle can be interrupted, provided specimens are not permitted
to thaw.

The apparatus described above is limited by two basic characteris-
tics of its construction: the vapor-handling capacity and the size of
the specimen chamber (fig. 8). As stated, the most efficient device
for removing water vapor from the specimen chamber is a refrigerated
condenser. If an external refrigerated condenser of sufficient size is
used, the only limitation to the size of the specimen chamber is the
size of the deep-freeze unit that holds the specimen chamber.

A refrigerated condenser can be constructed within a paint-spray
pressure tank similar to the one used for the specimen chamber. The
condensing surface may be a coil of %-inch copper refrigeration tubing

20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

with an expansion valve mounted at the top or inlet side of the coil.
The coil should have as many turns as space permits. A coil-within-
a-coil arrangement is most desirable (fig. 9).

Ficure 8.—Simple freeze-dry apparatus utilizing a condenser surface (4=specimen
chamber; B=commercial deep freeze; C=condenser chamber; D=vacuum pump).

A vacuum-tight drain should be installed at the bottom of the
condensing chamber to conveniently remove the condensation from
the chamber during defrosting. The condensing chamber should be
encased in an insulated box with openings in appropriate positions
for vapor lines and refrigeration tubing.

An important requirement in any freeze-dry system is a valve (see
fig. 5) that will permit atmospheric air to enter the system in order

Figure 9.—Condensing coil.

that the chamber can be opened. This valve should be so located as
to insure that the condensing chamber is situated between the valve
and the specimen chamber, thereby assuring condensation of atmos-
pheric moisture on the condensing coil rather than on the specimen.
The valve must be vacuum tight when closed and need not be large;
a ¥-inch intake is sufficient to shut down the system. It is recom-

NO. 3549

FREEZE-DRY PRESERVATION—HOWER

21

TaBLE 6.—Freezing mixtures (PS=primary substance and its proportion;
SS=secondary substance and its proportion; TS=tertiary substance and its
proportion; A=temperature (° C throughout) of substance before mixture;
B=temperature of mixture; C=total reduced temperature; D=temperature
when all snow is melted (when snow is used); H=heat absorbed (calories when
A is grams); *=lowest temperature obtained. Table modified from ‘‘Smith-
sonian Physical Tables,’”’ 1959, 9th rev. ed., by W. E. Forsythe.

PS

NaC2H302(cr)
NH;Cl

NaN 0O3
NaS203 (er)
KI

CaCly (er)
NHiNO3

(N Hy) 2504
NHi:Cl
CaCly

KNO3
NaSO,4
NaNO3
K2S0O4

NaC 03 (cr)
KNO3
CaCly
NHiCl
NHiN 03
NaN O3
NaCl
H2S01+H20
(66.1% H2SO)

Alcohol at 4° C
“é
Chloroform
Ether
Liquid SO,
NHiNO3
“ec

85
30
75
110
140
250
60
25

RSE Noe WNW WH YH
woo or or or or or

bt co
ao

wo =
RRB BB BB BBB Be BPP BP RP POO

a
va

Be eee Bee eee

SS TS

H20 100

“ ““

NHiND3 25

“ “cc “ “
“ “c “c “
NHgCl 25
“ “ “ “

“cc “ “cc “

sf - 61
ss . 70
s 81
coe 1583
See onaG
ee eon
“ce 73
CO: solid

“ “

snow “‘
H20 1.20
snow “
H20 1.31
snow ‘“
H20 3.61
snow ‘

A

10.7
13.3
13.2
10.7
10.8
10.8
13.6

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alt
=il
ill

erties hess Ten Seale ctor Vie Ua
en

ooooqocooccqo &

Co

me bho wy

i e
qooooqoooqcoo aoc ©

_

B

4.7
= Dik
—5.3
a0

1rd
—12.4
—13.6

—1.9

—2.0

—2. 85
—10.9
—15.4
—16.75
—17.75
—21.3
—37.0
—36. 0
—35. 0
—30.
—25.
—20.
= LO.

oooo

—30.
—72.
—T7.
—T71.
—82.

on
ero Oo Ovo Oo er a oreo 17

Cc

15.4
18.4
18.5
18.7
22. 5
23. 2
27.2
26. 0
22.0
20. 0
20. 0
19. 0
17.0

1.0

1.85

O89
14, 4
15.75
16. 75
20.3
36. 0
35. 0
34. 0
29. 0
24,0
19.0
15.0

—37.0
—30. 2
—25.0
—12.4
Ta
oak

oak

—19.7
—39. 0
—54, 9*
—40.3
—21.5
950)
—4.0

17.0
27.0
133. 0
273.0
553. 0
967.0
62.1
49.5
40.3
30.0
46.8
88. 5
192.3
392.3

22 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

mended that this valve be opened while the pump is still running to
prevent atmospheric air from forcing the pump to run backward or
forcing oil from the pump into the condensing chamber.

Figure 8 illustrates assembly of the modified apparatus using a
refrigerated condenser.

Preparation of Specimens

The freeze-dry process will not improve a poor specimen. If a
specimen is emaciated, slightly deteriorated, or otherwise inferior
when it enters the chamber, it will be in no better condition when it
leaves. Sagged tissue, however, can often be reshaped with subcu-
taneous injections of water, and sagged body cavities can be restored
with cotton before freezing.

The art and skill of the taxidermist will help decide the success or
failure of the freeze-dry technique. Many of the same tools and
methods of conventional taxidermy can be employed advantageously
in the freeze-dry process. Wires, supports, and other tricks of the
trade are useful. Also rapid freezing with liquid nitrogen or freezing
mixtures will hold a specimen in position.

INITIAL FREEZING OF THE SPECIMEN.—Of all compounds in animal
tissue, water is the most abundant. For the average, it constitutes
70 percent of the animal’s total weight. Water is found in cellular
and vascular spaces, and, in small quanties, in protein and carbohy-
drates. Of the total liquid content, 20 percent is usually in extracel-
lular fluid; approximately 25 percent of this extracellular fluid is plasma,
and the remainder is interstitial fluid, mostly water. About 76 per-
cent of muscle tissue is water. The Rowntree data (see Harrow, 1951)
relating to the biochemistry of man is a good general guide to the
distribution of tissue water in most mammals.

Water in tissue is found almost always in combination with naturally
occurring salts; for this reason, freezing should be brought about in
the shortest possible time. Slow freezing invariably leads to the
formation of eutectics through the concentration of salts. As freezing
water separates itself from a solution by the process of ion diffusion, the
salts become more highly concentrated than they were in the original
solution.

Kutectics have lower freezing points and lower vapor pressures than
water; thus, their formation during the process slows down drying.
Due to unfrozen saturated fluid in the tissue and resulting surface
tension, shrinkage occurs. Rapid freezing, which reduces the
formation of eutectics, can be accomplished by using the freezing
mixtures listed in table 6, liquid nitrogen, or a freezing chest with a

NO. 3549 FREEZE-DRY PRESERVATION—HOWER 23

temperature of less than —25°C. Rapid freezing also creates smaller
ice crystals, causing less tissue distortion during the process.
TECHNIQUES FOR FASTER DRYING.—The greatest amount of water
is removed early in the drying cycle when a specimen’s dried shell is
thinnest and offers the least impedence to the escape of water vapor.
The rate of weight loss due to the escape of vapor approaches zero
with the completion of drying. If a specimen chamber is loaded with
a great number of fresh specimens, the water-vapor atmosphere
will be very great during the first few days, possibly exceeding the
capacity of the system. If, however, specimens are introduced at
spaced intervals, the same number of specimens can probably be
processed without exceeding the system’s vapor-removing capacity,
and a constant level of water-vapor removal may be approached.

By drilling holes in the back or bottom portions of specimens, their
drying surfaces are increased and the area of their epidermal layers
is decreased, allowing water vapor to escape more rapidly. Skinning
one side of a specimen serves a similar function in that it removes
epidermal tissue that would otherwise act as a vapor barrier.

Evisceration of a specimen also reduces its water content consid-
erably; however, the removal of water-laden organs requires incision,
removal of the viscera, filling the body cavity with cotton or similar
material, all of which requires time and involves some of the less
desirable features of taxidermy.

A record should be maintained of each specimen’s weight since
observing the weight reduction (by removing the specimen from the
chamber and weighing it) is the best method for determining when a
specimen is completely dry. A specimen should be left in the chamber
for one or two days after the loss of weight ceases to be apparent.
This is especially true for rodents such as mice or rats with a scaly
epidermis on their tails. The tails dry slowly and it is therefore
advisable to perforate their undersides and keep the specimens in
the chamber beyond their apparent drying times.

Conclusion

As technology advances, it becomes apparent that freeze-dry
will be an ever-growing field. Plans are already underway for
increasing the Smithsonian facilities. Theoretically, there is no
limitation to the ultimate size or numbers of specimens that at one
time could be processed in this manner. It is hoped that this paper
will help to encourage the growth and exchange of further ideas on
the freeze-drying process.

24 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

References
Brappick, H. J. J.
1954. The physics of experimental method. New York: John Wiley and
Sons.
DanieEts, F.; Matuews, J. H.; Wiuuiams, J. W., et al
1949. Experimental physical chemistry, 4th ed. New York: McGraw-Hill
Book Co.
Guick, Davin.
1949. Techniques of histo- and cytochemistry.
Harrow, B.
1951. Textbook of biochemistry. Philadelphia and London: W. B.
Saunders Co.

Hower, R. O.
1962. Freeze-drying biological specimens. Smithsonian Inst. Inf. Leafl.,
no. 324.
1964, Freeze-drying biological specimens. Mus. News Techn. Suppl., vol.
no. e

Merryman, H. T.
1960. The preparation of biological museum specimens by freeze-drying.
Curator, vol. 3, no. 1.
1961. Biological museum specimens. Naval Medical Research Institute,
Bethesda, Md.
1961. The preparation of bilogical museum specimens by freeze-drying, 2:
Instrumentation. Curator, vol. 4, no. 2.
STRONG, J.
1953. Procedures in experimental physics. New York: Prentice-Hall.
Wo.LLASTON, WILLIAM HypE
1813. On a method of freezing at a distance. Roy. Soc. Phil. Trans.,
pp. 71-74.

U.S. GOVERNMENT PRINTING OFFICE: 1967

;

Proceedings of
the United States

National Museum

SMITHSONIAN INSTITUTION +« WASHINGTON, D.C.

Volume 119 1966 Number 3550

CATALOG OF TYPE SPECIMENS OF THE DARTERS
(PISCES, PERCIDAE, ETHEOSTOMATINI)!

By Bruce B. Cotiterre and Lestre W. Knapp’

Introduction

The darters are a tribe of small freshwater fishes restricted to North
America. Some 212 specific and subspecific names have been pro-
posed for the approximately 100 valid described species. About a
dozen species await description. As part of a long-term study of
these fishes, we have prepared this catalog of type specimens. We
hope that our efforts will be of value in furthering systematic research
and stabilizing the nomenclature of this most fascinating group of
North American fishes.

In preparing this catalog we have attempted to examine or at
least to verify the location of the type specimens of all nominal forms
of the three presently recognized genera of darters: Percina, Ammo-
crypta, and Htheostoma. By type specimens we mean holotypes,
lectotypes, syntypes, paralectotypes, and paratypes. Each form is

1 Fifth paper in a series on the systematics of the Percidae by the senior author.

2 Collette: Assistant Laboratory Director, Bureau of Commercial Fisheries
Ichthyological Laboratory, Division of Fishes, U.S. National Museum; Knapp:
Supervisor in charge of vertebrates, Oceanographic Sorting Center, Smithsonian
Institution.

1

2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

listed in alphabetical order by the generic and specific name used in
the original description. If subgeneric allocation was made in the
original description, that name has been placed in parentheses between
the genus and the species. Heolotypes or lectotypes are listed before
paratypes or paralectotypes.

The following data appear after each name: (1) citation of original
description; (2) catalog number; (3) number of specimens and range in
standard length in mm. are in parentheses; sex is given in a few cases;
(4) locality; (5) collector; (6) date of collection; (7) determination of
current status with reference to pertinent literature.

All original descriptions were consulted to help determine whether
a collection was actually type material. The generic and subgeneric
arrangement is modified from that presented by Bailey (in Bailey,
Winn, and Smith, 1954, and in Bailey and Gosline, 1955). The use
of the tribe Etheostomatini for the darters follows Collette (1963).

Lectotypes were selected in some instances in order to stabilize
nomenclature. The following people have assisted in the selection of
lectotypes: Dr. Charles F. Cole (subgenus Boleosoma); Mr. Donald
A. Distler (E. spectabile); Dr. Robert V. Miller (Z. blennioides); Dr.
Edward C. Raney (P. notogramma, P. crassa, subgenus Nothonotus);
Dr. William J. Richards (EF. swannanoa, E. thalassinum); Dr. Chu-fa
Tsai (E. histrio, E. rupestre, and EF. zonale). Brief data are presented
for each lectotype herein selected and the following abbreviations are
used: D=dorsal spines (Roman numerals) and rays (Arabic nu-
merals) ; A=anal spines (Roman) and rays (Arabic) ; LL=number of
lateral line scales; POM=number of preoperculomandibular pores;
and INF=number of infraorbital pores. If the lateral line is incom-
plete, the number of pored scales is given first, separated from the
number of unpored scales by a plus sign; if the infraorbital canal is
incomplete, the number of pores in the anterior portion is separated
from the number in the posterior portion by a plus sign.

Five nominal species are replacement names. No original material
was saved of the five species of darters described by Constantine
Rafinesque. Type material of 21 nominal species could not be located.
In instances where a type series was divided among several institu-
tions, we usually located some of the specimens. However, when all
of the types of a form were deposited in one institution we were
occasionally unable to locate any specimens.

About 1860, a considerable number of darters, including types of
16 species described by Charles Girard, were sent from the U.S.
National Museum to the Museum of Comparative Zoology at Har-
vard University for Frederick Putnam to study. Many of these were
cataloged into the MCZ collection (table 1), and Jordan and others
subsequently believed them lost. Representatives from a number of

ee ea eee ee SS ee Se

NO. 3550

DARTERS—COLLETTE

AND KNAPP 3

these series at the MCZ were later transferred to the University of
Michigan Museum of Zoology by Carl L. Hubbs.

Another problem concerns the type material studied by Jordan at
Indiana University. These specimens first received a lot number and

TABLE 1.—Darter types transferred from USNM to MCZ

First Present
USNM MCZ MCZ
No. No. No. Species and describer Locality
740 269 24599 Pileoma carbonaria Baird and Tex., Rio Salado
Girard.
743 87 24625 Diplesion fasciatus Girard Mex., Chihuahua R.
744 82 24575 Boleosoma lepida Baird and Tex., Rio Leona
Girard.
745 84 24567 Boleichthys elegans Girard Tex., Piedrapainte
748 83 24580 Oligocephalus leonensis Girard Tex., Rio Leona
1157 = 130 24546 Hadropterus maculatus Girard Md., Potomac R.
1175 257 246386 Pileoma zebra Agassiz Lake Superior
1179 1179 1179 Aplesion potsi Girard Mex., Chihuahua R.
1182 85 24576 Oligocephalus grahami Girard Tex., Devil’s R.
1187 22 24652 Catonotus fasciatus Girard IN-Y<, Grass) Re
1188 101 24589 Boleosoma fusiforme Girard Mass., Charles R.
1192 87 24625 Diplesion fasciatus Girard Mex., Chihuahua R.
1197 122 24603 Hadropterus nigrofasciatus Ala., near Mobile
Agassiz.
1203 16 24634 Oligocephalus humeralis Girard Va., James R.
1203 16 385975 Oligocephalus humeralis Girard Va., James R.
1206 87 24625 Dziplesion fasciatus Girard Mex., Chihuahua R.
1283 93 24570 Microperca punctulata Putnam Wis., Oconomowoc R.
1288 94 24690 Microperca punctulata Putnam Mich., Pt. Huron
1294 96 24582 Microperca punctulata Putnam Mich., Detroit R.
1303 129 24525 Hadropterus maculatus Girard Md., Potomac R.
1311 270 24616 Pleurolepis pellucidus Agassiz Ohio, Black R.
1827 __..- 32952 Boleichthys warreni Girard N. Dak., Cannon
Ball R.
1328) 113 113 Boleosoma gracile Girard Tex., Rio Seco
1333 41 24569 Boleichthys whipplii Girard Okla., Coal Cr.
1334 ___. 32950 Oligocephalus pulchellus Girard Okla., Gypsum Cr.
1336 114 24728 Boleichthys exilis Girard N. Dak., Little
Muddy R.
1365 2 24658 Catonotus Kennicotti Putnam Tll., Union Co.
1377 95 24566 Microperca punctulata Putnam IIil., Calumet R.
1393 21 24656 Catonotus fasciatus Girard N.Y., Grass R.
1801 1 24637 Catonotus lineolatus Agassiz Tll., near Quincy
1801 1 24657 Catonotus lineolatus Agassiz Til., near Quincy
1811 37 24665 Catonotus Kennicotti Putnam Til., Union Co.
1865 237 24602 Hyostoma Newmanii Agassiz Ala., Huntsville

later a regular catalog number.

We have recorded both numbers.

Some of these specimens are now at Stanford; many were transferred

to the California Academy of Sciences.

Later, some of the CAS types

4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

were transferred to the University of Michigan. However, a number
are still missing.

In addition to listing the type specimens, we have compiled a list
of the genus-group names proposed for the darters. The original
description for each was verified. We have attempted to discover
the first subsequent selection of type-species for those nominal genera
which lacked originally designated type-species. The list is divided
into available and unavailable genus-group names.

To facilitate using this catalog, we have also compiled an alpha-
betical index of nominal forms, using the original spelling. The
author and the genus in which the species was originally described
follow the specific name. Abbreviations of museums presently hous-
ing type material of each form are given following the name. Mu-
seums with the holotype, lectotype, or most syntypes are listed first.
A museum abbreviation with a question mark indicates where the
types were probably originally deposited even though we could not
locate them. A question mark indicates we could not locate any
types and do not know where they might be. An ‘“‘X”’’ is used when
there was no type material, or it is almost certainly lost.

We express our deep appreciation to the curators and staffs of the
museums and collections which house the type material of the darters.
Type material is or has been in the collections listed below. Collec-
tions marked with an asterisk have been visited by one or both of the
authors (BBC, LWK). Other initials represent those who kindly
verified material in a few collections, with small holdings of darter
types, which we were unable to visit.

*ANSP Academy of Natural Sciences, Philadelphia. Dr. James E. Bohlke.
[BBC, LWK.]
*BF U.S. Bureau of Fisheries. Collection now merged with USNM.

*BMNH British Museum (Natural History). Dr. P. H. Greenwood, Mr.
Peter J. Whitehead. [BBC.]

*CAS California Academy of Sciences, San Francisco. Mr. W.I. Follett,
Mrs. Lillian Dempster. [BBC, LWK.]

*CM Charleston Museum, §8.C. Mr. E. Milby Burton. [BBC]

*CU Cornell University, Ithaca, N.Y. Dr, Edward C. Raney, Mr.
Timothy Zorach, Mr. Robert E. Jenkins. [BBC, LWK.]

*DU Duke University, Durham, N.C. Dr. Joseph R. Bailey, Dr.

Walter R. Courtenay, Jr. [BBC.]
*FMNH Field Museum of Natural History, Chicago. Mrs. Marion Grey,
Miss Pearl Sonoda. [BBC, LWK.]

*INHS Illinois Natural History Survey, Urbana. Dr. Philip W. Smith
[BBC, LWK.]

*ISC Iowa State University, Ames. Dr. Kenneth D. Carlander. Type
darters transferred to USNM. [BBC.]

*IU Indiana University, Bloomington. Specimens now at CAS or

UMMZ. [BBC, LWK.]
*KU Kansas University, Lawrence. Dr. Frank B. Cross. [LWK, BBC].

NO. 3550 DARTERS—COLLETTE AND KNAPP 5

*MCZ Museum of Comparative Zoology, Harvard. Dr. Giles W. Mead,
Mrs. Myvanwy Dick. [BBC, LWK.]

*MNHN Muséum National d’Histoire Naturelle, Paris. Mme. M. L.
Bauchot. [LWK, BBC.]

*NCSM North Carolina State Museum, Raleigh. Mr. William M. Palmer.
Darter types transferred to USNM. [BBC, LWK.]

OAM-OSU Oklahoma Agricultural and Mechanical College, now Oklahoma
State University, Stillwater. Dr. Rudolph J. Miller. [RJM.]

*OSM Ohio State Museum, Columbus. Dr. Milton B. Trautman. [BBC,
LWK.]

*SU Stanford University, Division of Systematic Biology. Dr. George
S. Myers, Dr. Warren C. Freihofer, Mr. Stewart Springer.
[BBC, LWK.]

TAMC Texas Agricultural and Mechanical University, College Station.
Dr. Richard J. Baldauf. [RJB.]

TNHC Texas Natural History Collections, University of Texas, Austin.
Dr. Clark Hubbs. [CH.]

aw Tulane University, New Orleans. Dr. Royal D. Suttkus, Dr.
John S. Ramsey. [JSR.]

*TUC Tulsa University Collections. Dr. Albert P. Blair. Darter
types transferred to USNM. [BBC.]

UAIC University of Alabama Ichthyological Collections. Mr. William
M. Howell. [WMH.]

UG University of Georgia, Athens. Dr. Donald C. Seott. Darter

types transferred to CU. [Dr. William Richards, LWK.]
*UMMZ University of Michigan, Museum of Zoology, Ann Arbor. Dr.
Reeve M. Bailey, Dr. Robert R. Miller. [BBC, LWK.]
UOMZ University of Oklahoma, Museum of Zoology, Norman. Dr. Carl
D. Riggs. [CDR.]

UR University of Richmond, Richmond, Va. Dr. William S. Wool-
cott. [WSW.]

*USNM U.S. National Museum, Washington. [BBC, LWK.]

UW University of Wisconsin, Milwaukee. Dr. Carroll R. Norden.
[CRN.]

VET Virginia Polytechnic Institute, Blacksburg. Dr. William 8. Wool-
cott. [WSW.]

In addition to those already mentioned who have selected lecto-
types or who have permitted us to examine material housed in their
institutions, we wish to thank Drs. Robert H. Gibbs, Jr., James A.
Peters, and W. Ralph Taylor of the U.S. National Museum, for
critically commenting upon the manuscript.

Available Genus-Group Names

Allohistium Bailey in Bailey and Gosline, 1955, Misc. Publ. Mus.
Zool. Univ. Michigan, no. 93, p. 6. Type-species Htheostoma
cinerea Storer, 1845, by original designation.

Alvordius Girard, 1859, Proc. Acad. Nat. Sci. Philadelphia, vol. 11,
p. 68. Type-species Alvordius maculatus Girard, 1859, by
monotypy.

6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Ammocrypta Jordan, 1877, U.S. Nat. Mus. Bull. 10, p. 5. Type-
species Ammocrypta beanit Jordan, 1877, by original designation.

Anemoces Jordan, 1929, Manual of Vertebrates, 13th ed., p. 156.
Type-species Ulocentra gilbertt Evermann and Thoburn in
Jordan and Evermann, 1896, by original designation.

Arlina Girard, 1859, Proc. Acad. Nat. Sci. Philadelphia, vol. 11, p.
64. Type-species Arlina effulgens Girard, 1859 (=—Etheostoma
olmstedi Storer, 1842), by monotypy.

Asproperca Heckel in Canestrini, 1860, Verh. Zool.-Bot. Wien, vol.
10, p. 311. Type-species ‘‘Asproperca zebra Heck.” (=Ptleoma
zebra Agassiz, 1850), by monotypy.

Astatichthys Vaillant, 1873, Nouv. Arch. Mus. Hist. Nat. Paris, vol.
9, p. 106. Type-species Htheostoma caeruleum Storer, 1845, by
original designation.

Austroperca Hubbs, 1936, Occ. Pap. Mus. Zool. Univ. Michigan, no.
341, p.1. A replacement name for Torrentaria Jordan and Ever-
mann, 1896, preoccupied by Yorrentaria (Hodgson MS) Gray,
1863, Aves. Takes the same type-species as Torrentaria Jordan
and Evermann, Etheostoma australe Jordan, 1884.

Belophlox Fowler, 1947, Not. Nat. Acad. Nat. Sci. Philadelphia, no.
191, p. 1. Type-species Belophlox mariae Fowler, 1947, by
original designation.

Boleichthys Girard, 1859, Proc. Acad. Nat. Sci. Philadelphia, vol. 11,
p. 103. Type-species Boleichthys exilis Girard, 1859, by original
designation.

Boleosoma DeKay, 1842, Nat. Hist. New York Zool., vol. 4, p. 20.
Type-species Boleosoma tessellatum DeKay, 1842 (=EHtheostoma
olmstedi Storer), by monotypy.

Catonotus Agassiz, 1854, Amer. Journ. Sci. and Arts, vol. 17, p. 305.
Type-species Catonotus lineolatus Agassiz, 1854 (=Etheostoma
flabellaris Rafinesque, 1819), by monotypy.

Claricola Jordan and Evermann, 1896, U.S. Nat. Mus. Bull. 47, p.
1093. Type-species Etheostoma juliae Meek, 1891, by original
designation.

Copelandellus Jordan and Evermann, 1896, U.S. Nat. Mus. Bull. 47,
p. 1100. Type-species Poecilichthys quiescens Jordan, 1884 (=
Boleosoma fusiforme Girard, 1854), by original designation.

Cotiogaster Putnam, 1863, Bull. Mus. Comp. Zool., vol. 1, p.4 Type-
species Boleosoma tessellatum Thompson, 1853, not of DeKay (=
Rheocrypta copelandi Jordan, 1877), by monotypy.

Crystallaria Jordan and Gilbert in Jordan 1885, Ann. Rep. Comm.
Fish and Fish. for 1885, p. 866. Type-species Pleurolepis
asprellus Jordan, 1878, by original designation.

NO. 3550 DARTERS—COLLETTE AND KNAPP 7

Diplesion Rafinesque, 1820, Ichthyologia Ohiensis, p. 37. Type-
species Htheostoma blennioides Rafinesque, 1819, by subsequent
designation of Girard, 1859, p. 101 and Jordan and Gilbert, 1877,
p. 86.

Doration Jordan, 1929, Manual of the Vertebrates, 13th ed., p. 156.
Type-species Boleosoma stigmaeum Jordan, 1877, by original
designation.

Ericosma Jordan, 1877, U.S. Nat. Mus. Bull. 10, p. 8. Type-species
Alvordius evides Jordan and Copeland, 1877, by original desig-
nation.

Estrella Girard, 1859, Proc. Acad. Nat. Sci. Philadelphia, vol. 11,
p. 65. Type-species Estrella atromaculata Girard, 1859 (=Etheo-
stoma olmstedi Storer, 1842), by monotypy.

Etheostoma Rafinesque, 1819, Journ. Phys., Chim., Hist. Nat., vol. 88,
p. 419. Type-species Htheostoma blennioides Rafinesque, 1819, by
subsequent designation of Agassiz (1854, p. 305). Verified by
Opinion 14, Int. Comm. Zool. Nomen., 1910, Smithsonian
Inst. Publ., 1938, pp. 25-27. Not Etheostoma flabellaris Rafi-
nesque, 1819, as designated by Jordan and Gilbert, 1877, p. 85.

Hadropterus Agassiz, 1854, Amer. Journ. Sci. and Arts, vol. 17, p.
305. Type-species Hadropterus nigrofasciatus Agassiz, 1854, by
monotypy.

Hololepis Agassiz in Putnam, 1863. Bull. Mus. Comp. Zool., vol. 1,
p. 4. Type-species Boleosoma Barratti Holbrook, 1885 (=Boleo-
soma fusiforme Girard, 1854), by subsequent designation of
Jordan and Gilbert, 1877, p. 93.

Hyostoma Agassiz, 1854, Amer. Journ. Sci. and Arts, vol. 17, p. 305.
Type-species Hyostoma Newmaniw Agassiz, 1854 (=—EHtheostoma
blennioides Rafinesque, 1819), by monotypy.

Hypohomus Cope, 1870, Proc. Amer. Philos. Soc., vol. 11, p. 449.
Type-species Cottogaster aurantiacus Cope, 1869, by monotypy.

Imostoma Jordan, 1877, Proc. Acad. Nat. Sci. Philadelphia, vol. 29,
p. 49. Type-species Hadropterus shumardi Girard, 1859, by
original designation.

foa Jordan and Brayton, 1878, U.S. Nat. Mus. Bull. 12, p. 88. Type-
species Poecilichthys vitreus Cope, 1870, by original designation.

Litocara Bailey, 1948, Copeia, 1948, p. 79. Type-species Poecilichthys
sagitta Jordan and Swain, 1883, by original designation.

Microperca Putnam, 1863, Bull. Mus. Comp. Zool., vol.1,p.4. Type-
species Microperca punctulata Putnam, 1863 (=Etheostoma
microperca Jordan and Gilbert, in Jordan 1888), by monotypy.

Nanostoma Putnam in Jordan, 1877, U.S. Nat. Mus. Bull. 10, p. 6.
Type-species Poecilichthys zonalis Cope, 1868, by original
designation.

8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Niviperca Whitley, 1951, Proc. Royal Zool. Soc. New South Wales,
1949-50, p. 68. <A replacement name for Nivicola Jordan and
Evermann preoccupied by Nivicola Hodgson, 1844, Aves. Takes
the same type-species as Nivicola Jordan and Evermann, Poe-
cilichthys borealis Jordan, 1844 (= Boleichthys exilis Girard, 1859).

Nothonotus Agassiz in Putnam, 1863, Bull. Mus. Comp. Zool., vol. 1,
p. 3. Type-species Htheostoma maculatum Kirtland, 1840, by
subsequent designation of Jordan and Gilbert 1877, p. 93.

Oligocephalus Girard, 1859, Proc. Acad. Nat. Sci. Philadelphia, vol. 11,
p. 67. Type-species Boleosoma lepida Baird and Girard, 1853,
by subsequent designation of Vaillant (1873, p. 90).

Percina Haldeman, 1842, Journ. Acad. Nat. Sci. Philadelphia, vol. 8,
p. 330. Type-species Perca nebulosa Haldeman, 1842 (=Sciaena
caprodes Rafinesque, 1818), by monotypy.

Pileoma DeKay, 1842, Nat. Hist. New York Zool., vol. 1, pt. 4, p. 16.
Type-species Pileoma semifasciata DeKay, 1842 (=Sciaena
caprodes Rafinesque, 1818), by monotypy.

Plesioperca Vaillant, 1873, Nouv. Arch. Mus. Hist. Nat. Paris, vol. 9,
p. 36. Type-species Plesioperca anceps Vaillant, 1873 (= Hadrop-
terus nigrofasciatus Agassiz, 1854), by monotypy.

Poecilichthys Agassiz, 1854, Amer. Journ. Sci. and Arts, vol. 7, p. 304.
A replacement name for Poecilosoma Agassiz, 1850, preoccupied
by Poecilosoma Huebner, 1819, Lepidoptera; Poecilosoma Stephens
1829, Hemiptera; Poecilosoma Newman, 1838, Coleoptera. Takes
the same type-species as Poecilosoma Agassiz, Etheostoma variatum
Kirtland, 1838.

Psychromaster Jordan and Evermann, 1896, U.S. Nat. Mus. Bull. 47,
p. 1099. Type-species Etheostoma tuscumbia Gilbert and Swain,
1887, by original designation.

Rafinesquiellus Jordan and Evermann, 1896, U.S. Nat. Mus. Bull. 47,
p. 1082. Type-species Aplesion pottsiit Girard 1859, by original
designation.

Rheocrypta Jordan, 1877, U.S. Nat. Mus. Bull. 10, p. 9. Type-species
Rheocrypta copelandi Jordan, 1877, by monotypy.

Rhothoeca Jordan, 1885, Ann. Rept. Comm. Fish and Fish. for 1885,
p. 868. Substitute name for Nanostoma Putnam erroneously
regarded as preoccupied by Nannostomus Giinther and taking the
same type-species Poecilichthys zonalis Cope, 1869.

Richiella Coker, 1927, Copeia, no. 162, p. 18. Substitute name for
Richia Coker preoccupied by Richia Grote, 1887, Lepidoptera.
Takes the same type-species as Richia Coker, Richia brevispina
Coker, 1926 (=Etheostoma flabellaris Rafinesque, 1819).

NO. 3550 DARTERS—COLLETTE AND KNAPP 9

Serraria Gilbert, 1884, Proc. U.S. Nat. Mus. vol. 7, p. 205. Type-
species Hadropterus scierus Swain, 1883, by original designation.

Swainia Jordan and Evermann, 1896, U.S. Nat. Mus. Bull. 47,
p. 1040. Type-species Etheostoma squamatus Gilbert and Swain,
1887, by original designation.

Ulocentra Jordan, 1878, Manual of Vertebrates, 2nd ed., p. 223. Type-
species Arlina atripinnis Jordan, 1877, by monotypy.

Vaillantia Jordan in Jordan and Brayton, 1878, U.S. Nat. Mus. Bull.
12, p. 89. Type-species Boleosoma camurum Forbes, 1878
(=Vaillantia chlorosomum Hay, 1880), by original designation.

Vigil Jordan, 1919, Proc. Acad. Nat. Sci. Philadelphia, vol. 70, p. 344.
Substitute name for Pleurolepis Agassiz, 1863, preoccupied by
Pleurolepis Quenstedt, 1852, Pisces. Takes the same type-species
as Pleurolepis Agassiz, Pleurolepis pellucidus Agassiz in Putnam,
1863.

Villora Hubbs and Cannon, 1935, Mise. Publ. Univ. Michigan Mus.
Zool., no. 30, p. 11. Type-species Villora edwint Hubbs and
Cannon, 1935, by original designation.

Unavailable Genus-Group Names

Nivicola Jordan and Evermann, 1896, U.S. Nat. Mus. Bull. 47, p. 1082.
Type-species Poecilichthys borealis Jordan 1884, (=Boleichthys
exilis Girard, 1859), by original designation; preoccupied by
Nivicola Hodgson, 1844, Aves.

Pleurolepis Agassiz in Putnam, 1863, Bull. Mus. Comp. Zool., vol. 1,
p. 5. Type-species Pleurolepis pellucidus Agassiz in Putnam,
1863, by monotypy; preoccupied by Pleurolepis Quenstedt, 1852,
Semionotidae.

Poecilisoma Agassiz, 1850, Lake Superior, p. 299. 'Type-species
Etheostoma variata Kirtland, 1838, by subsequent designation of
Agassiz, 1854, p. 306. Name preoccupied by Poecilosoma
Huebner, 1819, Lepidoptera; Poecilosoma Stephens, 1829, Hemip-
tera; and Poecilosoma Newman, 1838, Coleoptera.

Richia Coker, 1926, Bull. Bur. Fish., vol. 42, p. 106. Type-species
Richia brevispina Coker, 1926 (=Etheostoma flabellaris Rafinesque,
1819), by original designation; preoccupied by Lichia Grote,
1887, Lepidoptera.

Torrentaria Jordan and Evermann, 1896, U.S. Nat. Mus. Bull. 47,
p. 1080. Type-species Etheostoma australe Jordan, 1884, by
original designation; preoccupied by Torrentaria (Hodgson MS)
Gray, 1863, Aves.

10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Nominal Species

Alvarius fonticola Jordan and Gilbert, 1886, Proc. U.S. Nat. Mus., |

vol. 9, p. 23. NRedescribed in more detail as ‘‘Htheostoma
(Alvarius) fonticola Jordan and Gilbert, sp. nov.’”’ in Gilbert
(1887, p. 63).

Lectotype: USNM 36523 (07, 27), Texas, San Marcos R. at San

Marcos; D. S. Jordan and C. H. Gilbert; September 1884; figured in |

Jordan and Evermann (1900, fig. 470); herein selected. D VII-10;

A I, 6; LL 2+30=32; INF 3+1; POM 8; supratemporal canal |

interrupted; nape, cheeks, and breast naked; breeding tubercles on
posterior anal rays.

Paralectotypes: USNM 198004 (3, 21-24), removed from USNM
36523.

=Etheostoma (Microperca) fonticola (Jordan and Gilbert).

Alvordius aspro Cope and Jordan in Jordan, 1877, Proc. Acad. Nat.
Sci. Philadelphia, vol. 29, p. 51, based on the figure but not the
description of ‘‘Htheostoma blennioides”’ in Kirtland, 1841, Boston
Journ. Nat. Hist., p. 3, pl. 6, fig. 1.

Types: This name was proposed as ‘‘Alvordius aspro, Cope and
Jordan, nom. sp. nov.,’”’ a replacement name for Etheostoma blennioides
of Kirtland’s figure. However, the name clearly applies to a new
species based on Kirtland’s misidentified figure. Cope and Jordan
did not provide any description, did not mention any localities, and
gave no indication of having any material before them. ‘Therefore,
the type of A. aspro must be the specimen from the Mahoning River,
Ohio, from which Kirtland drew his figure. We have not been able
to locate this specimen. However, there are two specimens collected
by Jordan from the White River in Indiana. One, labeled ‘‘paratype,”’
is USNM 20137, and the other, labeled ‘‘cotype,” is BMNH
1880.1.21.84. In the same year that A. aspro was described, Jordan
(1877, p. 375) listed A. aspro as ‘frequent’ in collections made by
Copeland and Jordan in the White River around Indianapolis. The
USNM and BMNH specimens should not be considered as type
material because the original description was based only on the
references to Kirtland’s figure.

= Percina (Alvordius) maculata (Girard).

Alvordius crassus Jordan and Brayton, 1878, U.S. Nat. Mus. Bull.
12(A); pri2:

Lectotype: USNM 23459 (<7, 63), South Carolina, Saluda R., near
Greenville; D. S. Jordan and A. B. Brayton; summer, 1877; labeled
“drawn’’; herein selected by Dr. Edward C. Raney. D XII-13; A
II,10; LL 54, complete; cheeks and anterior portion of nape naked;

NO. 3550 DARTERS—COLLETTE AND KNAPP 11

breast with one enlarged scale; another enlarged scale between pelvic
fin bases; opercle covered with scales.
Paralectotypes: USNM 31111 (8, 46-50), same data as lectotype.
=Percina (Ericosma) crassa crassa (Jordan and Brayton).

Alvordius evides Jordan and Copeland in Jordan, 1877, Proc. Acad.
Nat. Sci. Philadelphia, vol. 29, p. 51.

Lectotype: USNM 17872 (<7, 45), Indiana, Marion Co., White
R., near Indianapolis; D. S. Jordan; 1874; herein selected. D XII-13;
A 11,9; LL 64; INF 8; POM 10; cheeks with a small patch of scales
under junction of lateral and infraorbital canals; opercles scaly; nape
with scattered scales; breast naked except for one modified scale at
center; subocular bar present.

Paralectotypes?: MCZ 24399 (2 o&@, 42-46), same locality as
lectotype. We are not certain that MCZ 24399 contains the other
two syntypes on which the original description was based.

The original description was based on three specimens. Jordan
and Copeland (in Jordan, 1877, p. 8) utilized additional specimens
when they described Ericosma to contain the previously described
A. evides. These additional specimens were incorrectly labeled as
types: USNM 20119, MCZ 35957, and BMNH 1880.1.21.12-14.

=Percina (Ericosma) evides (Jordan and Copeland).

Alvordius maculatus Girard, 1859, Proc. Acad. Nat. Sci. Philadelphia,
vol. 11, p. 68.

Syntypes: USNM 1346 (2, ?), Fort Gratiot on Lake Huron; Maj.
B. Alvord; listed as thrown away in USNM catalog and we have
been unable to find them.

=Percina (Alvordius) maculata (Girard).

Ammocrypta beanii Jordan, 1877, U.S. Nat. Mus. Bull. 10, p. 5.
Holotype: USNM 17833 (<7, 52), Louisiana, Notalbany R., near
Tickfaw; T. H. Bean; December 1876.
= Ammocrypta (Ammocrypta) beanii Jordan.

Ammocrypta clara Jordan and Meek, 1885, Proc. U.S. Nat. Mus.,
vol. 8, p. 8.

Lectotype: USNM 35828 (1, 42), Iowa, Des Moines R. at Ottumwa;
D. S. Jordan and S. E. Meek; August 1884. Jordan and Evermann
(1896, p. 1063) listed USNM 35828, then containing four specimens,
as “type.” Linder (1959) selected one specimen as lectotype and
removed the other three.

Paralectotypes: USNM 164166 (3, 33-51), out of USNM 35828.
Linder overlooked paralectotypes SU 1500 (1, 46) and UMMZ
187510 (2, disintegrated), orig. IU 753, then IU 7323, that are also
from the original type series.

12 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

The original description was based on the Ottumwa specimens and
is followed by “specimens were also obtained in Red River at Fulton,
Arkansas, and in the Sabine River at Longview, Texas” (italics ours).
The Sabine River specimens (USNM 36488) and the Red River speci-
mens (USNM 36337) were labeled paratypes in the USNM collection,
but we do not feel they should be considered as such because they are
listed in the fashion of ‘other material.”

= Ammocrypta (Ammocrypta) clara Jordan and Meek. See Linder
(1959).

Ammocrypta gelida Hay, 1881, Proc. U.S. Nat. Mus., vol. 3, p. 490.
Syntypes: USNM 27425 (2, 44-45), Mississippi, Clarke Co.,

Chickasawha R. at Enterprise; O. P. Hay; March—April 1880.
=Ammocrypta (Ammocrypta) beanvi Jordan.

Ammocrypta vivax Hay, 1883, Bull. U.S. Fish Comm., vol. 2, p. 58.
Holotype: USNM 32213 (1, 39), Mississippi, Pearl R. at Jackson;
O. P. Hay; summer, 1881.
=Ammocrypta (Ammocrypta) vivax Hay.

Aplesion pots Girard, 1859, Proc. Acad. Nat. Sci. Philadelphia,
vol. 11, p. 102; potsit a misprint for pottsit.
Holotype: MCZ 1179 (1, 50), Mexico, Chihuahua R. and tribs.;
J. Potts; orig. USNM 1179.
= Ktheostoma (Oligocephalus) pottsii (Girard).

Arlina atripinnis Jordan, 1877, U.S. Nat. Mus. Bull. 10, p. 10.
Holotype: USNM 20483 (7, 56), Tennessee, trib. of Cumberland
R., near Nashville; A. Winchell; in poor condition; frenum present,
dorsal fin pattern clearly that of Ulocentra.
= KHtheostoma (Ulocentra) atripinne (Jordan).

Arlina effulgens Girard, 1859, Proc. Acad. Nat. Sci. Philadelphia,
Vole wy p: Go:
Holotype: USNM 1253 (7, 56), Washington, D.C., Rock Cr.,
trib. of Potomac R.; 8S. F. Baird; 1851.
= Etheostoma (Boleosoma) olmstedi olmstedi Storer. See Cole (1958).

Belophlox mariae Fowler, 1947, Notulae Naturae, no. 191, fig. 1.
Holotype: ANSP 71731 (, 55), North Carolina, Moore Co., trib.
of outlet of Watson’s Lake, about 3 mi. W. of Southern Pines, Little
Pee Dee drainage; H. W. Fowler; Oct. 12, 1946.
= Ktheostoma (Oligocephalus) mariae (Fowler).

Boleichthys elegans Girard, 1859, Proc. Acad. Nat. Sci. Philadelphia,
Vo it p04.

Syntypes: MCZ 24567 (2, 35-40), Texas, Piedrapainte; J. H. Clark

under Col. J. D. Graham; U.S. and Mexican Boundary Survey;

NO. 3550 DARTERS—COLLETTE AND KNAPP ites

orig. USNM 745, then MCZ 84. The USNM catalog gives four for
the number of specimens in USNM 745, but we have been unable to
find any types other than MCZ 24567.

= Etheostoma (Oligocephalus) grahami (Girard).

Boleichthys eos Jordan in Nelson, 1876, Bull. Illinois Mus. Nat. Hist.,
no. 1, p. 35. NRedescribed in more detail as ‘‘Boleichthys eos,
Jordan and Copeland, n. s.”’ in Jordan (1877, p. 46).

Syntypes: USNM 23460 (1, 42), Indiana, La Porte Co., Clear Lake;
Jordan; summer, 1875. USNM 17848 (9, 27-42), Indiana, Wabash
R.; Jordan. USNM 20117 (1, 34), Indiana, Wabash R.; Jordan; in
very poor condition.

=LHtheostoma (Oligocephalus) exile (Girard). See Hubbs (1926,
p. 67).

Boleichthys exilis Girard, 1859, Proc. Acad. Nat. Sci. Philadelphia,
VOL tl. p, 103:

Syntypes: USNM 1336 (24, 23-33), North Dakota, Little Muddy
R., trib. of upper Missouri R.; G. C. Suckley. MCZ 24728 (6, 27-33),
same data, all but the largest in poor condition; orig. USNM 1336,
then MCZ 114. UMMZ 86492 (1, 31) out of MCZ 24728.

=Ktheostoma (Oligocephalus) exile (Girard). See Hubbs (1926.
pp. 64-68).

Boleichthys warrent Girard, 1859, Proc. Acad. Nat. Sci. Philadelphia,
VOLuIt ps Or.

Holotype: MCZ 32952 (1, 42), North Dakota, Cannon Ball R.,
Hayden under Lt. G. K. Warren, exploration of the upper Missouri
and Yellowstone; September 1856. The USNM catalog lists USNM
1327 as the type of B. warreni, but we have been unable to find a
jar bearing this number. Jordan (1885, p. 869) and Hubbs (1926)
wrote that the type was lost. The original description was based on
only one specimen, and MCZ 32952 fits it very well.

=Etheostoma (Oligocephalus) exile (Girard). See Jordan (1885) and
Hubbs (1926, p. 67).

Boleichthys whipplit Girard, 1859, Proc. Acad. Nat. Sci. Philadelphia,
Ole di hesps LOS

Syntypes: MCZ 24569 (1, 39), ‘‘Ark.,’’ Coal Cr. [=trib. of Arkansas
R. in E. Oklahoma, Hubbs and Black, 1941]; party under Lt. A. W.
Whipple, Pacific Railroad Survey, orig. USNM 1333, then MCZ 41.
USNM. 1333 is listed in USNM catalog as containing two speci-
mens, but we have been able to find only the specimen now cataloged
as MCZ 24569.

=Etheostoma (Oligocephalus) whipplit whipplii (Girard). See
Hubbs and Black (1941).

14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Boleosoma aesopus Cope, 1870, Proc. Amer. Philos. Soc., vol. 11,
p. 270.

Syntypes: ANSP ?, Pennsylvania, Lycoming Co., Loyalsoc [now
Loyalsock] Cr. in the Allegheny region; A. H. Smith. Types of
other species described by Cope in this paper were deposited at
ANSP, but Fowler (1907) did not mention types of this form, and we
have not found them.

=IHtheostoma (Boleosoma) olmstedi olmstedi Storer.

Boleosoma Barratti Holbrook, 1855, Journ. Acad. Nat. Sci. Phila-
delphia, ser. 3, vol. 2, p. 56, pl. 6, fig. 3.

Lectotype: MCZ 24571 (o, 46), Florida, Holbrook; orig. MCZ
97; herein selected. D X-10; A II, 7; LL 26+24=—50; INF 3+1;
POM 9; interorbital, breast, and cheeks scaly.

Paralectotypes: MCZ 43090 (4, 37-40) and UMMZ 36598 (3%,
47), removed from MCZ 24571.

=Etheostoma (Hololepis) fusiforme barrattti (Holbrook). See Hubbs
and Cannon (1935) and Collette (1962).

Boleosoma camurum Forbes in Jordan, 1878, Bull. Illinois State Lab.
Nat. Hist:-vol.. 1,.nos25p.40:

Syntypes: USNM 27894 (2 9, 39), Illinois, Peoria Lake; S. A.
Forbes; Oct. 27, 1878. SU 1683 (9, 44), Illinois. USNM 23455
(9, 44), Illinois, Illinois R., figured in Jordan and Evermann (1898,
fig. 452). MCZ 25053 (9, 38), Illinois, Peoria Lake. UMMZ
187505 (1, 39), Illinois, Pekin, orig. IU 506, then IU 4707. There
were originally additional syntypes from ‘‘Cache R. and Clear Cr.,
Union Co.; Johnson Co.” according to the original description, but
we have not been able to locate them. Smith and Bridges (1960)
reported that there were no syntypes in the collection of the Illinois
Natural History Survey which inherited much of the material of the
old Illinois State Laboratory of Natural History.

=Etheostoma (Boleosoma) chlorosomum (Hay) because camurum
Forbes is preoccupied in Etheostoma by Poecilichthys camurus Cope.

Boleosoma fusiforme Girard, 1854, Proc. Boston. Soc. Nat. Hist., vol.
5, p. 41.

Lectotype: USNM 1188 (9, 33), Massachusetts, trib. of Charles
R. at Framingham; 8S. F. Baird; selected by Hubbs and Cannon
(1935, p. 78).

Paralectotypes: USNM 94686 (25, 23-36); MCZ 24589 (4, 27-36);
orig. USNM 1188, then MCZ 101. UMMZ 86582 (1, 37); out of
MCZ 24589.

=Etheostoma (Hololepis) fusiforme fusiforme (Girard). See Hubbs
and Cannon (1935) and Collette (1962).

NO. 3550 DARTERS—COLLETTE AND KNAPP 15

Boleosoma gracile Girard, 1859, Proc. Acad. Nat. Sci. Philadelphia,
1859, p. 103.
Lectotype: USNM 1328 (<7, 35), Texas, Rfo Seco near Ft. Inge;
C. B. Kennerly; selected by Hubbs and Cannon (1935, p. 44).
Paralectotypes: MCZ 113 (1, 36), orig. USNM 1328. USNM
1329 (1, 30), Texas, Rio Leona near Ft. Inge; Kennerly.
=Etheostoma (Hololepis) gracile (Girard). See Hubbs and Cannon
(1935) and Collette (1962).

Boleosoma lepida Baird and Girard, 1853, Proc. Acad. Nat. Sci.
Philadelphia, vol. 6, p. 388.

Syntypes: USNM 744 (6, 32-45), Texas, Rio Leona at Uvalde, trib.
of Nueces R.; J. H. Clark. MCOZ 24575 (8, 42-46), same data; orig.
USNM 744, then MCZ 82. UMMZ 86335 (1, 41), out of MCZ 24575.

=Etheostoma (Oligocephalus) lepidum (Baird and Girard).

Boleosoma maculaticeps Cope, 1870, Proc. Amer. Philos. Soc., vol. 11,
p. 269.

Lectotype: ANSP 13862 (07, 45), North Carolina, upper waters of
Catawba R.; E. D. Cope; selected and illustrated by Fowler (1907,
p. 523, fig. 6). All 15 syntypes were still in the same bottle. The
next to the largest specimen is closest (55.4 mm.) in total length to
the size indicated by Fowler for the ‘‘type”’ (57.0 mm.), and it has the
correct number of fin rays: D [X-13; A IIJ,8. These are also the counts
given by Cope in the original description. The lectotype has 43
lateral line scales, close to Fowler’s figure which has about 45. This
specimen herein confirmed as lectotype. INF 8; POM 10; pectoral
rays 12-12.

Paralectotypes: ANSP 13863-76 (14, 23-48), same data as lecto-
type.
=FEtheostoma (Boleosoma) nigrum maculaticeps (Cope). See Cole
(1958).

Boleosoma maculatum Agassiz, 1850, Lake Superior, p. 305, pl. 4,
he. 3.

Lectotype: USNM 1840 (0, 43), Ontario, Lake Superior, Ft.
William and “the Pic” at the mouth of the Pic R.; L. Agassiz and
party; July 13-20, 1848; herein selected by Dr. Charles F. Cole.
D IX-138; A 1,8; pectoral 12-13; LL 45+3=48; infraorbital pores
4-++2; nape, cheeks, and breast naked; opercles partially scaled.

Paralectotype: USNM 197633 (9, 41), removed from USNM 1840.

=LHtheostoma (Boleosoma) nigrum nigrum Rafinesque.

Boleosoma mutatum Vaillant, 1873, Nouv. Arch. Mus. Hist. Nat.
Paris, p. 88, a substitute name for Boleosoma maculatum Agassiz
erroneously thought by Vaillant to be preoccupied in Boleosoma
by Etheostoma maculatum Kirtland.

16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Types: The types of B. mutatum are the same as those of Boleosoma
maculatum Agassiz.
=Etheostoma (Boleosoma) nigrum nigrum Rafinesque.

Boleosoma nigrum eulepis Hubbs and Greene, 1935, Trans. Wisconsin
Acad. Sci., Arts, Lett., vol. 29, p. 96, pl. 3, figs. 1-3.

Holotype: UMMZ 77225 (1, 45), Wisconsin, Dodge Co., trib. of
West Branch Rock R., 2 mi. N. of Atwater; C. W. Greene, Stuart,
Creaser, and Griffin; Aug. 27, 1927.

Paratypes: UMMZ 100771 (79, 26-47), out of UMMZ 77225.
USNM 74927 (2, 33-53), Wisconsin, Turtle Cr. at Delavan; N.
Hollister; Oct. 11, 1910. USNM 117505 (8, 45-46), Wisconsin,
Marquette Co., Welsh Cr.; Greene and H. S. Jones; Sept. 1, 1925;
orig. UMMZ 73880. UMMZ 101528 (9, 25-52), Iowa, Dickinson
Co., Spirit L.; J. C. Salyer; July 27, 1932. UMMZ 94832 (22, 26-48),
Minnesota, Mississippi R. between Cass L. and Bemidji; Hubbs and
Schultz; June 5, 1926. UMMZ 66681 (9, 24-28), Minnesota, Hub-
bard Co., L. Laselle and outlet; P. A. Moody; Aug. 11, 1925. UW 64
(17, 24-40), Wisconsin, Columbia Co.; Sand (Rock) Cr.; Greene and
Jones; Aug. 18, 1925. UW 466 (5, 19-37), Wisconsin, Rock Co.,
Yahara R. at Indian Ford; G. Wagner; June 23, 1906. UW 467
(2, 40-41), Wisconsin, Walworth Co.; Geneva L.; Wagner; June 26,
1906. UW 473 (22, 24-59), Wisconsin; Manitowoc Co., Mud Cr. at
Reedville; Wagner; July 17, 1906. UW 474 (18, 25-31), Wisconsin,
Calumet Co., cr. at Brillion; Wagner; July 18, 1906. UW 479 (2,
38-43), Wisconsin, Vilas Co., Trout Lake Cr.; Wagner; July 10, 1907.
UW 480 (8, 44-48), Wisconsin, Oneida Co., cr. between Squirrel L.
and Diamond L.; Wagner; July 20, 1907.

The following paratypes are all UMMZ specimens from Wisconsin:
72370 (13, 29-54), Polk Co., Apple R., 2 mi. W. of Turtle L. on rt. 14;
P. Okkelberg; Aug. 26, 1928. 72382 (13, 23-48), same data. 72414
(60, 25-57), Vilas Co., SE. side of Big Arbor Vitae L.; Metzelaar and
Langlois; Sept. 19, 1925. 72447 (9, 36-50), Oneida Co., Kamaguesaga
L., W. of Minoigua; Metzelaar and Langlois; Sept. 20, 1925. 72467
(2, 32-37), Vilas Co., Trout L.; Metzelaar and Langlois; Sept. 20,
1925. 72495 (14, 39-53), Vilas Co., Little Arbor Vitae L.; Metzelaar
and Langlois; Sept. 21, 1925. 72502 (18, 32-52), Vilas Co., St.
Germain L.; Metzelaar and Langlois; Sept. 21, 1925. 73442 (15,
30-49), Columbia Co., Fox R., 10 mi. E. of Portage on rt. 33; Greene
and Jones; Aug. 18, 1925. 73458 (1, 42), Columbia Co., French
Cr., below dam; Greene and Jones; Aug. 19, 1925. 73464 (35,
32-56), Columbia Co., French Cr., 1 mi. above mill pond; Greene
and Jones; Aug. 19, 1925. 73472 (4, 46-56), Columbia Co., Spring
Branch Cr., above co. rd.; Greene and Jones; Aug. 19, 1925. 73475
(3, 48-53), same data. 73563 (11, 31-51), Marquette Co., trib. of

NO. 3550 DARTERS—COLLETTE AND KNAPP 17

Neenah Cr., 1.5 mi. from Neenah Cr.; Greene and Jones; Aug. 21,
1925. 73571 (5, 37-47), Columbia Co., outlet of Spring L., Pardee-
ville; Greene and Jones; Aug. 22, 1925. 73579 (28, 23-48), Columbia
Co., old channel, below Park L. Dam; Greene and Jones; Aug. 22,
1925. 73586 (11, 34-45), Columbia Co., Park L.; Greene and Jones;
Aug. 22, 1925. 73627 (89, 27-55), Marquette Co., Neenah Cr., 2 mi.
NE. of Briggsville on rt. 23; Greene and Jones; Aug. 24, 1925.
73640 (2, 40-41), Columbia Co., trib. of Neenah Cr., 6 mi. E. of
Briggsville; Greene and Jones; Aug. 24, 1925. 73670 (66, 30-48),
Marquette Co., Belle Fountaine Cr., 4 mi. N. of Dalton; Greene and
Jones; Aug. 25, 1925. 73683 (11, 31-49), Columbia Co., Neenah Cr.,
near mouth; Greene and Jones; Aug. 26, 1925. 73693 (33, 30-44),
Columbia Co., French Cr. at mouth; Greene and Jones; Aug. 26, 1925.
73706 (7, 32-41), Columbia Co., Fox R., below mouth of French Cr.;
Greene and Jones; Aug. 26, 1925. 73728 (4, 33-47), Columbia Co.,
Fox R., opposite Lock 25; Greene and Jones; Aug. 26, 1925. 73735
(6, 41-55), Columbia Co., Fox R. at Portage; Greene and Jones;
Aug. 26,1925. 73744 (47, 28-45), Fond du Lac Co., Grand R., above
Fairwater Millpond; Greene and Jones; Aug. 5, 1925. 73810 (14,
34-49), Green Lake Co., Grand R., below Kingston Dam; Greene and
Jones; Aug. 29, 1925. 73821 (4, 40-47), Greene Lake Co., Grand R.
at Kingston; Greene and Jones; Aug. 29, 1925. 73831 (5, 27-44),
Green Lake Co., trib. of Grand R., 4 mi. SW. of Uthey; Greene and
Jones; Aug. 31, 1925. 73835 (85, 31-51), Fond du Lac Co., trib. of
Grand R., 2 mi. S. of Fairwater; Greene and Jones; Aug. 31, 1925.
73862 (30, 39-51), Green Lake Co., Grand R., 3 mi. S. of Marquette;
Greene and Jones; Sept. 1, 1925. 73874 (26, 27-48), Marquette Co.,
trib. of Grand R., 2 mi. W. of Kingston; Greene and Jones; Sept. 1,
1925. 73877 (8, 38-51), Marquette Co., trib. of Grand R., 1.5 mi.
S. of Welsh Cr.; Greene and Jones; Sept. 1, 1925. 73888 (57, 28-55),
Marquette Co., trib. of Grand R., Welsh Cr.; Greene and Jones;
Sept. 1, 1925. 73955 (34, 26-54), Fond du Lac Co., Rush Cr., near
Waupun; Greene and Cuchio; July 4, 1926. 77054 (33, 25-53),
Columbia Co., Crawfish R., 2 mi. NW. of Fall R.; Greene and Stuart;
Aug. 21, 1927. 77073 (35, 30-48), Dodge Co., Beaver Dam Cr., 5 mi.
S. of Beaver Dam; Greene and Stuart; Aug. 21, 1927. 77082 (18,
32-53), Columbia Co., trib. of Crawfish R., 4 mi. NW. of Columbus;
Greene and Stuart; Aug. 21, 1927. 77088 (15, 28-44), Columbia Co.,
trib. of Crawfish R. just N. of Columbus on rt. 16; Greene and
Stuart; Aug. 22, 1927. 77096 (3, 33-40), Columbia Co., Crawfish R.
at Columbus; Greene and Stuart; Aug. 22, 1927. 77110 (5, 32-39),
Dodge Co., trib. of Crawfish R., 10 mi. W. of Watertown on rt. 19;
Greene and Stuart; Aug. 23, 1927. 77125 (8, 31-46), Dodge Co.,
trib. of Beaver Dam Cr., 2 mi. E. of Lowell on rt. 16; Greene and
219-948—66——2

18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Stuart; Aug. 23, 1927. 77135 (6, 28-39), Dodge Co., W. Branch of
Rock R., above millpond at Hustisford; Greene and Stuart; Aug. 27,
1927. 77153 (2, 37-41), Dodge Co., Beaver Dam Cr., below dam at
Fox L.; Greene and Stuart; Aug. 26, 1927. 77165 (11, 32-45), Dodge
Co., Crawfish R., 8 mi. N. of Watertown; Greene and Stuart; Aug. 26,
1927. 77208 (22, 27-44), Dodge Co., Old Mill Cr., 1 mi. S. of At-
water; Greene, Stuart, Creaser, Griffin; Aug. 27, 1927. 77232 (44,
25-49), Fond du Lac Co., W. Fk. of Upper W. Br. of Rock R., 4 mi.
NW. of Waupun; Greene, Stuart, Creaser, Griffin; Aug. 29, 1927.
77592 (32, 21-44), Jefferson Co., Rock R., below Ft. Atkinson;
Schultz and Tarzwell; Aug. 7, 1928. 77779 (30, 26—56), St. Croix Co.,
Apple R., above Somerset; Schultz and Tarzwell; Aug. 11, 1928.
78384 (6, 24-40), Rock Co., Rock R., 2.5 mi. N. of Beloit ; Schultz and
Tarzwell; Aug. 27, 1928. 78415 (48, 24-37), same data. 95933 (3,
26-49), Sawyer Co., Grindstone L., 10 mi. SE. of Haywood; Creaser
and Jones; Sept. 2, 1928. 96111 (9, 24-48), Washburn Co., Trough
L., 14 mi. SE. of Spooner on rt. 53; Creaser and Jones; Sept. 7,
1928. 96234 (15, 30-61), Polk Co., Apple R. at Amery; Creaser and
Jones; Sept. 11, 1928. 96254 (13, 28-40), Polk Co., Balsam L. at
Balsam City; Creaser and Jones; Sept. 12, 1928. 96570 (11, 43-52),
Vilas Co., Trout L.; Wright; Aug. 6, 1927. 96573 (7, 20-48), Vilas
Co., Trout L.; Wright; Aug. 4, 1927. 96703 (4, 48-47), Vilas Co,.
Trout L.; Wright; Aug. 4, 1927. 100202 (2, 37-45), Vilas Co., Silver
L.; Schneeberger; July 19, 1932.

=? Etheostoma (Boleosoma) nigrum nigrum Rafinesque. See Lagler
and Bailey (1947) and Underhill (1963). We agree with Underhill
that recognition of the scaly populations of H. nigrum as a subspecies
does not aid in understanding the complex distributional pattern of
the two morphological types present in the Great Lakes and upper
Mississippi regions.

Boleosoma olmstedi var. brevipinnis Cope, 1868, Journ. Acad. Sci.
Philadelphia, ser. 2, vol. 6, p. 214.

Lectotype: ANSP 22653 (o, 46), Pennsylvania, Kiskiminitas R.,
trib. of Allegheny R.; E. D. Cope; number selected by Fowler (1907,
p. 522 misprinted as “25,563 (type) to 22,567’), but all syntypes
found still mixed together. Largest specimen herein selected as
lectotype. D VIII-11; A 1,9; LL 38+4=42; INF 4-+3; pectoral
rays 12-12; nape, preopercle, and breast naked; anterior part of belly
naked.

Paralectotypes: ANSP 22654-67 (14, 34-40), same data as lecto-
type.

Cope mentioned two additional specimens of this form from the
Holston River. Fowler did not list these as types. We feel it is best

NO. 3550 DARTERS—COLLETTE AND KNAPP 19

not to consider these specimens as paralectotypes and formally re-
strict the type locality to the Kiskiminitas River.
=Etheostoma (Boleosoma) nigrum nigrum Rafinesque.

Boleosoma perlongum Hubbs and Raney, 1946, Misc. Publ. Mus.
Zool. Univ. Michigan, no. 65, p. 17, pl. 1, fig. 3.

Holotype: UMMZ 138475 (0, 71), North Carolina, Columbus Co.,
N. shore of Lake Waccamaw; E. C. Raney, E. A. Lachner, and R. A.
Pfeiffer; Mar. 30, 1941; orig. CU 9285.

Paratypes: UMMZ 138476 (16, 44-75), same data as holotype.

=Ktheostoma (Boleosoma) perlongum (Hubbs and Raney). See
Cole (1958).

Boleosoma phlox Cope, 1880, U.S. Nat. Mus. Bull. 17, p. 30.
Holotype: ANSP 20447 (1, 34), Texas, Trinity R. at Fort Worth;
E. D. Cope. We have found no other type material and cannot
determine if the original description was based on one or more speci-
mens. Thelabel says ‘‘type.”” Fowler (1907) referred to ANSP 20447
as ‘“‘type,’”’ not cotype, so we assume this specimen is the holotype.
=Etheostoma (Oligocephalus) lepidum (Baird and Girard).

Boleosoma stigmaeum Jordan, 1877, Ann. New York Lyceum Nat.
Bist? vol. 12, p. sit.

Lectotype: ANSP 20645 (<7, 39), Georgia, Floyd Co., Etowah R.,
trib. of Coosa R. near Rome; D. S. Jordan and C. H. Gilbert; summer,
1876; selected by Bailey, Winn, and Smith (1954, p. 142).

Paralectotypes: ANSP 20646-48 (3, 36-39), same data as lectotype.
One syntype was figured by Fowler (1907, fig. 5).

Although labeled as a syntype of B. stigmaeum, USNM 31108 is a
specimen of Etheostoma (Ulocentra) coosae (Fowler).

=Etheostoma (Boleosoma) stigmaeum (Jordan).

Boleosoma susanae Jordan and Swain, 1883, Proc. U.S. Nat. Mus.,
vol. 6, p. 249.

Lectotype: USNM 36501 (07, 41), Kentucky, Whitley Co., trib.
of Clear Fk., Cumberland R. near Pleasant View; D. S. Jordan and
J. Swain; May 1883; herein selected by Dr. Charles F. Cole.
D VIII-12; A 1,9; pectoral 11-11; LL 40+4=44; infraorbital canal
412; preoperculomandibular pores 9; cheeks, opercles, nape, breast,
and anterior half of belly naked.

Paralectotypes: USNM 197634 (7, 25-30), removed from USNM
36501.

=Etheostoma (Boleosoma) nigrum susanae (Jordan and Swain),
apparently a valid subspecies.

Boleosoma tessellatum DeKay, 1842, Nat. Hist. New York Zool., vol.
1, pt. 4, p. 20, pl. 20, fig. 57.

20 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

Syntypes: Description and figure based on specimens then in
New York State collection. Dr. E. M. Reilly, Curator of Zoology,
has informed us that the types of this species are no longer present
in the New York State Museum.

= Etheostoma (Boleosoma) olmstedi olmstedi Storer. See Cole (1958).
Described after July 1, 1842, because that is the date on the preface
to part 4, ‘“‘Fishes,”’ of the ‘“‘Zoology of New York.” Thus Etheostoma
olmstedi Storer, January 1842 has priority.

Boleosoma vexillare Jordan, 1880, Proc. U.S. Nat. Mus., vol. 2, p. 235.

Holotype: ?(o, 2%in. TL), Virginia, Rappahannock R. at Warren-
ton. Jordan and Evermann (1896, p. 1058) wrote that the type was
in the Charles Hallock collection. Cole was unable to find the type
during the course of his thesis work on the subgenus Boleosoma, and
we have also been unable to find it.

=Etheostoma (Boleosoma) olmstedi vevillare (Jordan). See Cole
(1958).

Catonotus fasciatus Girard, 1859, Proc. Acad. Nat. Sci. Philadelphia,
vol lie piGs:

Lectotype: USNM 1345 (o’, 60), New York, Grass R., near
Madrid; E. O. Dayton; 1852; herein selected. D VIII-14; A II,7;
LL 32+21=538; INF 4+2; POM 9; supratemporal canal moderately
interrupted; about 10 dark vertical bars on side of body.

Paralectotypes: USNM 197997 (o, 53), removed from USNM
1345. USNM 1393 (4, 27-41), New York, Madrid; Dayton; 1852.
MCZ 24652+24656 (6, 21-58), same locality; 1853. MCZ 24652
(“‘2””) was originally USNM 1187, then MCZ 22. MCZ 24656 (‘‘6’’)
was originally USNM 1393, then MCZ 21.

= KHtheostoma (Catonotus) flabellare flabellare Rafinesque.

Catonotus Kennicotti Putnam, 1863, Bull. Mus. Comp. Zool., vol. 1,
Dp. o:

Lectotype: USNM 1365 (<7, 35), Illinois; Union Co., a rocky
brook; R. Kennicott; April 1857; herein selected. D VII-14; A IJI,7;
LL 25+28=53; INF 4+2; POM 10; supratemporal canal moderately
interrupted; cheeks, opercles, and breast naked; nape with few,
scattered, imbedded scales; branchiostegal membranes moderately
connected; tips of dorsal spines with bony knobs; about 10 horizontal
lines on body.

Paralectotypes: USNM 198002 (20, 29-41), removed from USNM
1365. USNM 154713 (7, 27-42), same data; orig. USNM 1365,
then Wesleyan Uniy. 554; specimens badly dried out. MCZ 24658
(6, 29-50), same data; orig. USNM 1365, then MCZ 2. UMMZ
86347 (1, 42), out of MCZ 24658. MCZ 24665 (0%, 52; 39, 31-47),
same locality; April-May 1856; orig. USNM 1811, then MCZ 37.

NO. 3550 DARTERS—COLLETTE AND KNAPP 2A

USNM 154784 (1, 42), same data; orig. USNM 1811, then Wesleyan
Univ. 558.
= KHtheostoma (Catonotus) kennicottt (Putnam).

Catonotus lineolatus Agassiz, 1854, Amer. Journ. Sci. and Arts, vol. 17,
p. 305.

Lectotype: MCZ 24637+24657 (<7, 46), Illinois, small creeks near
Quincy; L. Watson; 1853; orig. USNM 1801, then MCZ 1; herein
selected. D VII-14; A II,8; LL 24+29=53; INF 4+2; POM 10;
supratemporal canal incomplete; interorbital pores present; head and
breast naked; humeral spot prominent; tips of dorsal spines widened ;
about seven rows of fine longitudinal stripes on the body above the
lateral line; branchiostegal membranes broadly connected.

Paralectotypes: MCZ 43093 (5, 30-39) and UMMZ 86481 (20,
41-52) removed from MCZ 24637+24657. MNHN 2785 (0, 48),
same locality, orig. MCZ 859.

= EK theostoma (Catonotus) flabellare lineolatum (Agassiz).

Cottogaster aurantiacus Cope, 1868, Journ. Acad. Nat. Sci. Philadelphia,
SEE. 2, vol. 6, p. 211, color pl: 24, fig: 6.

Lectotype: ANSP 13789 (1, 55), Virginia, N. Fork of Holston R.;
E. D. Cope; 1867; number selected by Fowler (1907, p. 521); specimen
herein selected. D XV-14; A II,10; LL 91.

Paralectotype: ANSP 13790 (1, 55), same data as lectotype.

= Percina (Hypohomus) aurantiaca (Cope).

Cottogaster cheneyi Evermann and Kendall, 1898, Bull. U.S. Fish
Comm., vol. 17, p. 129, pl. 8, fig. 8.

Holotype: USNM 48781 (<7, 51), New York, St. Lawrence Co.,
Racket R. [now Raquette R.], near Norfolk; B. W. Evermann and
B. A. Bean; July 18, 1894; figured in Jordan and Evermann (1900,
fig. 445).

Paratypes: USNM 48782 (4, 39-47); USNM 126978 (4, 40-47),
orig. BF 527; and SU 9048 (4, 40-49), orig. given as SU 5774; all with
same data as holotype.

=Percina (Cottogaster) copelandi (Jordan).

Cottogaster putnami Jordan and Gilbert, 1883, U.S. Nat. Mus. Bull.
16, p. 498.

Lectotype: USNM 1314 (<7, 45), New York, Westport Br., Lake
Champlain; S. F. Baird; September 1853; orig. no. 55; herein selected.
D XI-12; A IJ,9; LL 48; INF 8; POM 10; cheeks naked; breast
naked except for a large modified scale in the center and another
between the bases of the pelvic fins; nape naked anteriorly; opercles
scaly. This name was proposed as ‘nom. sp. nov.” for Boleosoma
tessellatum of Thompson (1853), Putnam (1863), and others, not of

22 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

DeKay. Cottogaster putnami was actually being proposed as the
name of a new species and not as a substitute name, so the collection
on which the description was based, USNM 1314, becomes the type
material.

Paralectotypes: USNM 198000 (27, 35-49), removed from USNM
1314 + USNM 1193. USNM 1193, with the same locality and
collector as USNM 1314, is now mixed with 1314 and may have been
so at the time of the original description. All the specimens are the
same species. At least one syntype of C. putnami out of USNM 1314
was included in a gift to the University of Cincinnati, dated Oct. 30,
1884.

=Percina (Cottogaster) copelandi (Jordan).

Diplesion fasciatus Girard, 1859, Proc. Acad. Nat. Sci. Philadelphia,
vol/ 11>. 100:

Lectotype: MCZ 24625 (<7, 45), Mexico, Chihuahua R.; J. Potts;
orig. USNM 743 (6)+USNM 1192 (3)+-USNM 1206 (1), then MCZ
87; herein selected. D X-12; A I,7; LL 46+13=59; INF 9; POM
about 10; interorbital, parietal, nape, cheeks, opercle, and breast
naked; anal spine very strong.

Paralectotypes: MCZ 43092 (o', 42; 9, 42) and UMMZ 86349
(9, 41), removed from MCZ 24625.

=Etheostoma (Austroperca) australe Jordan because fasciatus is
preoccupied in Htheostoma by Catonotus fasciatus Girard.

Estrella atromaculata Girard, 1859, Proc. Acad. Nat. Sci. Philadelphia,
volali; p66:

Types: We have been unable to find type material of this form.
During the course of his thesis work on the subgenus Boleosoma,
Cole was also unable to locate any types. The original description
was based on specimens from ‘‘the Potomac River, in the neighborhood
of Washington.”

=Etheostoma (Boleosoma) olmstedi atromaculatum (Girard). See
Cole (1958).

Etheostoma (Nothonotus) acuticeps Bailey, 1959, Occ. Pap. Mus. Zool.
Univ. Michigan, no. 603, pl. 1.

Holotype: UMMZ 159014 (<7, 58), Tennessee, Sullivan Co., S.
Fork of Holston R., 0.5 mi. above S. Holston Dam, 7 mi. SE. of
Bristol; R. M. Bailey, G. P. Cooper, and J. D. Kilby; June 23, 1949.

Paratypes: UMMZ 159013 (3, 49-51), same data as holotype.
UMMZ 157585 (2, 45-52), same locality, R. M. Bailey, R. W.
Eschmeyer, and J. Chance; Sept. 23, 1947.

= Etheostoma (Nothonotus) acuticeps Bailey.

NO. 3550 DARTERS—COLLETTE AND KNAPP 23

Etheostoma arcus-celestis Crevecoeur, 1903, Trans. Kansas Acad. Sci.,
vol. 18, pp. 177-178.

Syntypes: USNM?, Kansas, Pottawatomie Co., French Cr., trib.
of Vermillion R. near Onaga; F. F. Crevecoeur; April-May 1902.
The types were sent to the U.S. National Museum in 1907 along
with specimens of Campostoma anomalum and Semotilus atromaculatus.
We have located the minnows (USNM 72474 and 70457) but have
been unable to find the types of E. arcus-celestis.

=LHtheostoma (Oligocephalus) spectabile pulchellum (Girard) accord-
ing to Mr. Donald A. Distler (pers. comm.).

Etheostoma aubeenaubei Evermann, 1900, Rept. U.S. Fish Comm.,
vol, 25, p. 367, pl. 17.

Holotype: USNM_ 65248 (9, 41), Indiana, Marshall Co.,
Aubeenaubee Cr., E. inlet of Lake Maxinkuckee; Evermann and
Scovell; Aug. 23, 1899; erroneously given as USNM 49379 in original
description.

Paratypes: All from Aubeenaubee Cr., Indiana. USNM 65255 (2,
38-39), Aug. 23, 1899. USNM 65258 (10, 34-40), Aug. 4, 1899.
USNM 65260 (69, 35-40), Aug. 23, 1899. Two paratypes out of
USNM 65260 were given to the Lebanon City Schools, Lebanon,
Wis., Apr. 5, 1911. USNM 65249 (10, 25-30); Long Point; Aug. 11,
1899. USNM 65263 (5, 30-34), Aug. 8, 1899.

=Kitheostoma (Oligocephalus) exile (Girard). See Hubbs (1926,
p. 68).

Etheostoma (Etheostoma) australe Jordan, 1889, Proc. U.S. Nat. Mus.
vol. 11, p. 862, replacement name for Diplesion fasciatus Girard,
preoccupied in Etheostoma by Catonotus fasciatus Girard.

Lectotype: MCZ 24625 (7, 45). See Diplesion fasciatus.
Paralectotypes: MCZ 43092 and UMMZ 86349.
=KHtheostoma (Austroperca) australe Jordan.

Etheostoma blennioides Rafinesque, 1819, Journ. Phys., Chim., Hist.
Nat., Arts, vol. 88, p. 419.

Types: As Call (1899) has noted, Rafinesque did not preserve any
type material. Type locality Ohio River. Examination of a photo-
eraph of a sketch of this species from Rafinesque’s notebook (fig. 1)
confirms the attribution of the name blennioides to the species cur-
rently known by this name.

=F theostoma (Etheostoma) blennioides blennioides Rafinesque. See
Miller (1964).

Etheostoma (Rhothoeca) blennius Gilbert and Swain in Gilbert, 1887,
Proc: U:S. Nat. Mus., vol. 10, p. 55.

QA PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Holotype: USNM 36187 (1, 60), Alabama, Cox’s Cr., trib. of
Tennessee R. near Florence; C. H. Gilbert and J. Swain; summer, 1884.

Paratype: SU 1955 (1, 47), Alabama, Shoal Cr., trib. of Tennessee
R. near Florence; Gilbert and Swain; summer, 1884.

= Htheostoma (Etheostoma) blennius Gilbert and Swain. See Hubbs
and Black (1940).

Etheostoma caerulea D. H. Storer, 1845, Proc. Boston Soc. Nat.
Hist., vol. 2, p. 47. Original description repeated almost ver-
batim, with the addition of color changes in alcohol, in Storer
(1846).

Holotype: MCZ 32949 (9, 53), Illinois, Fox R.; S. C. Clark.
=LKtheostoma (Oligocephalus) caeruleum caeruleum Storer. See

Trautman (1930) and Knapp (1964).

Etheostoma cinerea D. H. Storer, 1845, Proc. Boston Soc. Nat. Hist.,
vol. 2, p. 49. Original description repeated almost verbatim in
Storer (1846).

Syntypes: MCZ ?, Alabama, Florence; received from C. A. Hentz.

We have been unable to find any types.

=LKtheostoma (Allohistium) cinereum Storer. See Bailey and
Gosline (1955).

hi ree ee yy FAI
Coser hae ae

Pe os 4 SS BWP Dat te
yy aren ot > syd R ee

Figure 1.—Etheostoma blennioides Rafinesque (sketch from Rafinesque’s notebook).

Etheostoma (Hololepis) collis lepidinion Collette, 1962, Tulane Stud.
Zool., vol. 9, no. 4, p. 194, figs. 17, 18.

Holotype: USNM 179847 (, 40), Virginia, Charlotte Co., trib. of
Horsepen Cr., 2.4 mi. NW. of Wyliesburg on Va. 907; B. B. Collette,
C. F. Cole, J. G. New, E. C. Raney, C. R. Robins; Mar. 31, 1956.

Paratypes: CU 29992 (8, 30-38), same data as holotype. USNM
100215 (8, 29-31), Virginia, Charlotte Co., Wards Fork, trib. of
Roanoke R., S. of Madisonville; S. Abraham; Apr. 23, 1935. USNM
101330 and 101334 (12, 22-29), Virginia, Charlotte Co., Wards Fork,
Roanoke Cr., below mill dam between Madisonville and Cullen;
G. S. Myers and Abraham; Sept. 15, 1935. CU 34544 (7, 29-40),
Virginia, Charlotte Co., Wards Fork Cr., 6.7 mi. SSW. of Madisonville

No. 3550 DARTERS—COLLETTE AND KNAPP 25

on Va. 47; Collette and J. S. Ramsey; Sept. 16, 1959. DU uncat.
(co, 38), North Carolina, Granville Co., Beech Cr., 3 mi. NNE. of
_ Cornwall (now flooded by Kerr Dam); J. R. Bailey 52-7; spring, 1952.
DU uncat. (59, 31-36), North Carolina, Durham Co., Lick Cr. on
rt. 264, 4.5 mi. E. of Oak Grove; Bailey and Walter; Apr. 6, 1950.
DU uncat. (0, 36), North Carolina, Orange Co., Eno R., 2 mi. W. of
Hillsboro; J. R. Bailey 49-9; Mar. 20, 1949. DU uncat. (9, 38),
North Carolina, Orange Co., Eno R. at ford N. of Hillsboro near
Skipper Wright’s; J. R. Bailey 55-3; Apr. 21, 1955.

= Etheostoma (Hololepis) collis lepidinion Collette.

Etheostoma cragini Gilbert, 1885, Bull. Washburn College Lab. Nat.
Hist., vol, 1, no. 3, p. 99.

Lectotype: USNM 38320 (1, 33), Kansas, Finney Co., small brook
- leading from the “Lake” at Garden City to Arkansas R.; F. W. Cragin;
March-April 1885; USNM 38320 listed by Jordan and Evermann
(1896, p. 1092) as “‘type” then containing three specimens; lectotype
herein selected. D IX—12; A 11,6; LL 19+32=51; POM 10; INF 7
| pores on right, 8 on left; supratemporal canal widely interrupted;
cheeks and opercles naked; breast and prepectoral area with few, im-
_ bedded scales.

Paralectotypes: USNM 197994 (2, 29), removed from USNM
38320. USNM 125109 (1, 23), same data, orig. BF 309.

Additional syntypes from SU 2141 became mixed with SU 694 from
Canon City, Colo. and so were not considered for selection of a lecto-

type.

=Ktheostoma (Oligocephalus) cragini Gilbert.

Etheostoma cumberlandicum Jordan and Swain, 1883, Proc. U.S. Nat.
Mus. vol. 6, p. 251.

Lectotype: USNM 36502 (0, 41), Kentucky, Whitley Co., Wolf
Cr., trib. of Clear Fork, Cumberland R., near Pleasant View; D. S.
Jordan, J. Swain, C. H. Gilbert; May 1883; herein selected. This
specimen was found alone in a bottle with three labels, each reading
“type”. Four additional specimens were found in another bottle, also
labeled USNM 36502, Etheostoma cumberlandicum. The single speci-
men is in as good condition as the largest of the four, so it is selected
lectotype. D VII-11; A II,7; LL 25+19=44; INF 4+1; POM 10;
supratemporal canal interrupted; preopercle smooth; cheeks, opercles,
nape, and breast naked.

Paralectotypes: USNM 197992 (4, 20-42), removed from USNM
36502.

=Etheostoma (Catonotus) flabellare cumberlandicum Jordan and
Swain. See Ross and Carico (1963, p. 12).

2°26 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

Etheostoma (Hadropterus) cymatotaenia Gilbert and Meek in Gilbert,
1887, Proc. U.S. Nat. Mus., vol. 10, p. 51. i
Lectotype: USNM 36215 (<7, 68, labeled “‘drawn’’), Missouri, Ni- |
angua R., near Marshfield; C. H. Gilbert and S. E. Meek; summer,
1884; herein selected; (fig. 2). D XII-12; A 11,10; LL 66; branchi-
ostegal membranes slightly connected; opercles, cheeks, breast, and
belly scaly; midventral belly scales slightly modified; thin flangelike
midventral keel on lower edge of caudal peduncle near caudal base.

Paralectotypes: USNM 198090 (<7, 63), removed from USNM
36215. USNM 36260 (2 &, 63-68), Missouri, Osage Fork of Gas-
conade R., near Marshfield; Gilbert and Meek; summer, 1884; erro-
neously given as USNM 38260 in original description and in Jordan
and Evermann (1896, p. 1042). USNM 36308 (5, 40-44), Missouri,
Sac R., near Greenfield; Gilbert and Meek; summer, 1884. SU 2603
(4, 39-54), same data as USNM 36308.

Additional syntypes from SU 3949, with the same data as USNM
36215, became mixed with nontype specimens from Kentucky, col-
lected by A. J. Woolman as a result of the 1906 earthquake.

=Percina (Hypohomus) cymatotaenia (Gilbert and Meek).

Ficure 2.—Etheostoma cymatotaenia Gilbert and Meek (previously unpublished drawing of
lectotype drawn in October 1884).

Etheostoma (Ulocentra) davisoni Hay, 1885, Proc. U.S. Nat. Mus., vol.
8, p. 554.

Holotype: USNM 37365 (1, 41), Florida, Santa Rosa Co., Yellow
R., near Chaffin; H. T. Mann and D. M. Davison; April 1885.

—=Ltheostoma (Boleosoma) davisoni Hay. Etheostoma davisoni has
long been considered a synonym of E. stigmaeum. However, E.
davisoni has recently been recognized as a distinct species by Mr.
William Mike Howell, who is presently studying the taxonomy and
systematics of this group. His validation of /. davisoni will appear
elsewhere.

NO. 3550 DARTERS—COLLETTE AND KNAPP 20

_ Etheostoma (Oligocephalus) ditrema Ramsey and Suttkus, 1965, Tulane

Stud. Zool., vol. 12, no. 3, p. 66, figs. 1-3.
Holotype: TU 35703 (0, 34) Georgia, Chattooga Co., trib. to

Mills Cr., trib. to Chattooga R., 4.3 airline mi. W. of Lyerly; July 18,
| 1962.

Paratypes: TU 29153 (21, 15-35), out of TU 35703. The following

| are all from the type locality. TU 26086 (8, 24-31), Apr. 19, 1962;

TU 27566 (9, 18-34), May 30, 1962; TU 32762 (34, 19-42), June 1,
1964; TU 32981 (43, 20-39), June 23, 1964. Five paratypes out of TU
32981 were sent to each of the following institutions: USNM 198607
(29-32), MCZ 43123 (29-37), ANSP 101231 (26-32), CU 47872
(30-32), SU 62401 (27-36), and UMMZ 187501 (29-36). Also ANSP
20649 (1, 23), Georgia, Floyd Co., near Rome; summer, 1876; D. S.
Jordan.
=FEtheostoma (Oligocephalus) ditrema Ramsey and Suttkus.

Etheostoma duryi Henshall, 1889, Journ. Cincinnati Soc. Nat. Hist.,
vol. 11, p. 32.
Syntypes: Cincinnati Soc. Nat. Hist. (3 o, 2 9, 2-3 in.), Tennessee,

small trib. of Tennessee R. at Whiteside on Memphis and Charleston
_RR.; C. Dury. We have been unable to locate any of the syntypes.

=KHtheostoma (Ulocentra) duryi Henshall.

Etheostoma flabellaris Rafinesque, 1819, Journ. Phys., Chim., Hist.
Nat., Arts, vol. 88, p. 419.

Types: As Call (1899) indicated, Rafinesque did not preserve
any of his type material. Type locality—the Ohio R. The sketch
of this species in Rafinesque’s notebook (fig. 3) leaves no doubt
as to the identity of what Rafinesque had before him.

=Htheostoma (Catonotus) flabellare flabellare Rafinesque.

Etheostoma fontinalis Rafinesque, 1820, Ichthyologia Ohiensis, p. 85.
Types: As Call (1899) noted, Rafinesque did not preserve types.
Type locality—‘“‘in the springs and caves near Lexington.”
=LHtheostoma (Catonotus) flabellare flabellare Rafinesque according
to Jordan and Evermann (1896, p. 1097).

Etheostoma formosa Henshall, 1889, Journ. Cincinnati Soc. Nat.
Est vol tt p. 32.

Holotype: Cincinnati Soc. Nat. Hist. (1, 2% in.), Tennessee, small
trib. of Tennessee R. at Whiteside on Memphis and Charleston RR.;
C. Dury. We have been unable to locate the type.

=Etheostoma (Oligocephalus) caeruleum ssp. See Knapp (1964).

Etheostoma fricksia Hildebrand, 1923, Bull. Bur. Fish., vol. 39, p. 7,
fies L,

28 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Holotype: USNM 82633 (<7, 44), Georgia, Richmond Co., Savan-
nah Dr., small creek on Sanitary Dairy Farm near Augusta; S. F,
Hildebrand; Mar. 26, 1918.

Paratypes: USNM 92818 (o’, 31; 22, 33-47), same data as
holotype.

=Kiheostoma (Oligocephalus) fricksium Hildebrand. See Bailey
and Richards (1963).

Etheostoma gintheri Eigenmann and Eigenmann, 1892, Amer. Nat.,
vol. 26, p. 962. Description much enlarged in EHigenmann
(1894).

Lectotype: BMNH 1892.12.30.151 (oo, 41), Canada, Manitoba,
Red R. of the North at Winnipeg; C. H. Eigenmann; August-September
1892; herein selected. D IX—13; A 11,10; LL 57; moderate frenum
present; posterior rays of anal fin extending beyond posterior end
of depressed second dorsal; black spot at anterior and posterior ends
of first dorsal.

Paralectotypes: BMNH 1892.12.30.152-153 (2, 41-51), same data
as lectotype.

gee hy yes:

———_

,

249 th a 9 ;
Mls eerky 07 fbUelon --C%an feata

“

Figure 3.—Etheostoma flabellare Rafinesque (sketch from Rafinesque’s notebook).

Three other specimens are mentioned, both in the original descrip-
tion and in the 1894 paper. We do not think that these should be
considered as syntypes. USNM 45567 (7, 42) and UMMZ 187522
(2, 31-42), orig. IU 1449, then IU 4506, Iowa, Cedar Rapids; S. E.
Meek.

=Percina (Imostoma) shumardi (Girard). See Hubbs and Greene
(1928, p. 384).

Etheostoma hildebrandtii Evermann and Clark, 1910, Proc. Biol:
Soc. Washington, vol. 23, p. 87, fig. 1.

Holotype: USNM 64616 (1, 43), Indiana, Fulton Co., Fletcher
Lake, 12 mi. N. of Logansport near Cass Co. line; B. W. Evermann and
H. W. Clark; Oct. 6, 1900.

=LHtheostoma (Oligocephalus) exile (Girard).

NO. 3550 DARTERS—COLLETTE AND KNAPP 29

Etheostoma (Ulocentra) histrio Jordan and Gilbert in Gilbert, 1887,
Proc. U.S. Nat. Mus., vol. 10, p. 47.

Lectotype: USNM 363886 (<7, 42), Oklahoma, Poteau R., W. of
Hackett City, Ark.; D. S. Jordan and C. H. Gilbert; summer, 1864;
herein selected by Dr. Chu-fa Tsai. D JX-13; A II,7; LL 53; INF
4+3; POM 9; belly and breast naked; cheek with 6 imbedded scales;
opercle with 2 scales; 6 dorsal blotches.

Paralectotypes: USNM 188972 (3, 24-33), removed from USNM

| 36386. SU 2682 (2, 33-35), same data. USNM 36409 (4, 24-35)

and SU 569 (1, 32), Arkansas, Clark Co., Washita (Ouachita) R. at

_ Arkadelphia; Jordan and Gilbert; summer, 1884. USNM 36448 (3,
_ 30-34), Arkansas, Saline Co., Saline R. at Benton, just above railroad
_ bridge; Jordan and Gilbert; summer, 1884.

=Etheostoma (Etheostoma) histrio Jordan and Gilbert.

Etheostoma (Oligocephalus) hopkinsi binotatum Bailey and Richards,
1963, Occ. Pap. Mus. Zool. Univ. Michigan, no. 630, pl. 1, B—C.
Holotype: CU 44157 (co, 53), South Carolina, Abbeville Co.,
Calhoun Cr., 7.6 mi. E. of Calhoun Falls on S.C. 72; E. C. Raney,
C. R. Robins, R. H. Backus, R. W. Crawford, J. N. Layne, and
R. L. Wigley; ECR 1932; Mar. 27, 1951; orig. CU 19600.
Paratypes: CU 19600 (14, 37-55) and UMMZ 180075 (<7, 48; 3 9,
44-50), same data as holotype. Apparently the ‘additional speci-
mens examined” by Bailey and Richards must also be considered as
paratypes because they form part of the type-series as defined by
Article 72b of the International Code of Zoological Nomenclature and
were not referred to as variants, doubtfully associated with the
nominal form, or expressly excluded from it. CU 48401 (1, 38),
Georgia, Madison Co., S. Fork of Broad R., 2 mi. W. of Comer;
Nov. 1, 1952, Ichthyology Class; orig. UG 269. CU 48399 (1, 33),
Georgia, Madison Co., S. Fork of Broad R., 1-2 mi. S. of Carlton;
Apr. 11, 1954; orig. UG 22E. CU 48400 (1, 46), Georgia, Madison
Co., S. Fork of Broad R., 1-2 mi. S. of Carlton; Apr. 19, 1954; Ichthy-
ology Class; orig. UG 22D. CU 48396 (5, 30-49), Georgia, Ogle-
thorpe-Madison cos., S. Fork of Broad R., Anthony Shoals, near
Carlton; Nov. 1, 1952; Ichthyology Class; orig. UG 22C. CU
43994 (o’, 48), Georgia, Madison Co., S. Fork of Broad R., Anthony
Shoals, near Carlton; Sept. 3, 1950; W. J. Houck. UMMZ 133122
(3, 32-47), Georgia, Oglethorpe Co., Dry Fork Cr.; O. K. Fletcher;
Nov. 3, 1940. UMMZ 133224 (8, 27-42), Georgia, Oglethorpe Co.,
Dry Fork Cr.; Fletcher; Feb. 8, 23, 1941. UMMZ 137773 (81, 27-
43), Georgia, Oglethorpe-Wilkes cos., Dry Fork Cr., about 1 mi. N.
of U.S. 78; Fletcher; Mar. 11, Apr. 13, 1941. CU 48398 (2, 38-46),
Georgia, Oglethorpe Co., Buffalo Cr., 6 mi. E. of Lexington; Apr. 29,
1953; B. Martof; orig. UG 283. CU 48397 (2, 30-41), Georgia,

30 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Columbia Co., W. Fork of Cliatt Cr.; July 24, 1948; orig. UG 90.
CU 48402 (15, 21-51), Georgia, Lincoln Co., Lloyds Cr.; July 23,
1948; Scott and Tyson; orig. UG 80. USNM 179063 (4, 37-49),
South Carolina, Anderson Co., Twenty-three Mile Cr., 0.9 mi. NW.
of Sandy Springs, 11.1 mi. NW. of Anderson on U.S. 76; Mar. 25,
1951; Raney, Robins, Backus, Crawford, Layne, Wigley; orig. CU
19604. CU 43515 (6, 23-39), South Carolina, Edgefield Co., 3 mi.
E. of Edgefield Co. line on Ga. 23; Apr. 3, 1953; J. New.

=Ktheostoma (Oligocephalus) hopkinst binotatum Bailey and |

Richards.

Etheostoma iowae Jordan and Meek, 1885, Proc. U.S. Nat. Mus., vol.
8, p. 10.

Lectotype: SU 1457 (1, 36), Iowa, Chariton R. at Chariton; D. S.
Jordan and S. E. Meek; July-September, 1884; herein selected; one
specimen of FE. nigrum removed and recataloged as SU 62430. D
IX-10; A IJ, 7 or 8; LL 28+31=859, first anal spine much thicker
than second; preopercle, opercle, nape, and belly scaled; breast
naked.

Paralectotype: The original description was based on three speci-
mens, but we have been unable to locate the third.

=F theostoma (Oligocephalus) exile (Girard). See Hubbs (1926, p.
66).

Etheostoma (Nothonotus) jordani Gilbert, 1891, Bull. U.S. Fish Comm.,
vol. 9, p;, 156,-pl. 43, fies2:

Lectotype: USNM 125110 (07, 48), Alabama, Calhoun Co., Choc-
colo Cr. trib. of Coosa R. at Oxford; P. H. Kirsch, W. M. Andrews,
and E. O. Jones; May 23, 1889; herein selected. D X-11; A II,
8; LL (right side) 49; INF 8; POM 10; cheeks, prepectoral area, and
breast naked; opercles, nape, and belly scaled; no regular longitudinal
dark stripes along edges of scale rows.

Paralectotypes: USNM 198107 (8, 27-47), removed from USNM
125110. BMNH 1893.5.5.9-10 (2, 27-39), same data as lectotype.
CAS 22854 (4, 24-38), and UMMZ 61706 (9, 40), same data; orig.
IU 4693; badly dried. We have not located the additional syntypes
from Chestnut Cr., Verbena, Ala.

=KEtheostoma (Nothonotus) jordani Gilbert. See Bailey (1959).

Etheostoma juliae Meek, 1891, Bull. U.S. Fish Comm., vol. 9, p. 130,
pl. 42, fig. 2.

Syntypes: USNM 125116 (2 o&, 41-49) and SU 1010 (0%, 37),
Missouri, Greene Co., James R., trib. of White R., near Springfield;
S. E. Meek, L. Rettger, and F. M. Drew; July 31, 1889; one of syn-
types figured in Jordan and Evermann (1900, fig. 466).

| NO. 3550 DARTERS—COLLETTE AND KNAPP 31

_ Two topotypes of this species were found with LE. spectabile (USNM
63366) that were collected with the types of E. juliae. They have
_ recataloged as USNM 196750.

=Etheostoma (Oligocephalus) juliae Meek.

_ Etheostoma lepidogenys Evermann and Kendall, 1894, Bull. U.S. Fish
womm:, vol. 12, p, 114, pl..35.

Syntypes: USNM 44840 (2 o, 38-42), Texas, Comal Co., Guada-

_ lupe Drainage, Corral Cr. in the largest of the Comal Springs at New

| Braunfels; B. W. Evermann; Dee. 3, 1891.

=Ktheostoma (Oligocephalus) lepidum (Girard).

Etheostoma Linsleyr H. R. Storer, 1851, Proc. Boston Soc. Nat. Hist.,
vol. 4, p. 37.

Syntypes: New York, Wayne Co., streams at Walcott [now Wol-
cott], near Lake Ontario; J. H. Linsley. We have been unable to
find any type material of this nominal form.

=Etheostoma (Catonotus) flabellare flabellare Rafinesque.

Etheostoma (Imostoma) longimana Jordan, 1888, Proc. Acad. Nat.
Sci. Philadelphia, vol. 40, p. 179.

Lectotype: MCZ 24619 (1, 53), Virginia, trib. of James R., S. F.
Baird; about 1855; orig. USNM 1305, then MCZ 150; herein selected
by Dr. Charles F. Cole. Anal spines II; LL 44; INF 8; POM 10;
supratemporal canal complete; pectoral rays 13-12; scales above
lateral line 4; scales below lateral line 8.

Paralectotypes: MCZ 43074 (2, 36-37), removed from MCZ
24619; 5 specimens of E. nigrum removed and recataloged as MCZ
42199. USNM 120258 (2, 42-44) and UMMZ 86603 (3, 33-47),
same data; previously MCZ 24619; 1 specimen of L. nigrum removed
from USNM 120258 and recataloged as USNM 197414.

=KHtheostoma (Boleosoma) longimanum Jordan.

Etheostoma (Etheostoma) luteovinctum Gilbert and Swain in Gilbert,
1887, Proc. U.S. Nat. Mus., vol. 10, p. 58.

Lectotype: USNM 36139 (1, 37), Tennessee, Stone R., near Nash-
ville; C. H. Gilbert and J. Swain; summer, 1884; herein selected.
D IX-13; A JI,7; LL 33+17=50; INF barely incomplete, 4+-5;
POM 10; supratemporal canal complete; cheeks, opercles, and nape
fully scaled; breast partially scaled; 6 dorsal saddles.

Paralectotypes: USNM 198003 (2, 33-35), removed from USNM
36139, and SU 3945 (2, 39-40), same data as lectotype.

=Etheostoma (Oligocephalus) luteovinctum Gilbert and Swain.
Etheostoma lynceum Hay in Jordan, 1885, Ann. Rept. Comm. Fish

and Fish. for 1884, p. 868, a substitute name for Nanostoma
elegans Hay which is preoccupied in Etheostoma by Boleichthys
elegans Girard.

32 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Types: The types of EL. lynceum are the same as those of Nanostoma
elegans, USNM 27445, holotype and SU 627, paratype.

=Ktheostoma (Etheostoma) zonale lynceum (Hay), a valid subspecies
according to Dr. Chu-fa Tsai (pers. comm.).

Etheostoma macrocephalum Cope, 1869, Trans. Amer. Philos. Soc.,
new ser., vol. 13, p. 400.

Lectotype: ANSP 22626 (<7, 56), Pennsylvania, Youghiogheny R.;
E. D. Cope. This specimen herein selected as the lectotype because
it appears to be the only extant syntype. D XIV-12; A 11,10; LL
about 76; opercle naked; few small scales on cheek posterior to eye;
snout long and pointed; eye length longer than snout length.

Fowler (1907) listed ANSP 22626 and 22628 as cotypes. ANSP
22628 (<7, 58) is not part of the type series but a Percina maculata
with LL 62-64, cheek and opercle completely covered with scales,
short blunt snout, eye about equal to snout length. The ANSP
catalog lists this specimen as Alvordius maculatus from the Kiski-
minitas River. Cope based his description on three syntypes, but we
have been unable to locate the other two.

= Percina (Alvordius) macrocephala (Cope).

Etheostoma maculata Kirtland, 1841, Boston Journ. Nat. Hist., vol. 3,
P2io, pla2s fies os
Types: We have not found any extant type material. Type lo-
cality—Mahoning R., Ohio.
=Etheostoma (Nothonotus) maculatum maculatum Kirtland. See
Bailey (1959) and Raney and Zorach (MS).

Etheostoma microperca Jordan and Gilbert in Jordan, 1888, Manual
of the Vertebrates, 5th ed., p. 134, a substitute name for M@-
croperca punctulatum (Putnam), preoccupied in Htheostoma.

Types: The types of EL. microperca are the same as those of M-
croperca punctulatum Putnam.
=LHtheostoma (Microperca) microperca Jordan and Gilbert.

Etheostoma micropterus Gilbert, 1890, Proc. U.S. Nat. Mus., vol. 13,
p. 289.

Holotype: USNM 38245 (9, 40), Mexico, Chihuahua; E. Wilkin-
son; figured in Evermann and Kendall (1894, pl. 35) and Jordan and
Evermann (1900, fig. 459).

=LHtheostoma (Oligocephalus) pottsia (Girard).

Etheostoma (Nothonotus) moorei Raney and Suttkus, 1964, Copeia,
nove ip. Lieshole
Holotype: CU 42883 (co, 43), Arkansas, Cleburne Co., Devils
Fork, 2.5 mi. SW. of Woodrow and 6 airline mi. W. of Drasco; L. W.
Knapp and R. V. Miller; Apr. 26, 1962.

NO. 3550 DARTERS—COLLETTE AND KNAPP ap

Paratypes: CU 41966 (15, 28-42), same data as holotype. USNM
188357 (108, 20-56) and UMMZ 181397 (15, 31-50), type locality;
K. Strawn et al.; Aug. 21, 1962. TU 22697 (41, 26-57), Arkansas,
Cleburne Co., Devils Fork between Stark and Edgemont on rt. 16;
R. D. Suttkus, M. A. Wilkens, and B. I. Sundararaj; Oct. 23, 1959.
TU 26227 (22, 27-50), Arkansas, Van Buren Co., Middle Fork at
Shirley; Suttkus and W. T. Mason; Aug. 17, 1962. TU 26249 (21,
31-44), Arkansas, Van Buren Co., South Fork, 9 mi. SW. of Clinton
on rt. 95; Suttkus and Mason; Aug. 17, 1962. KU 6652 (5, 20-36),
Arkansas, Van Buren Co., South Fork of Little Red R. on Ark. 95,
approx. 2.5 mi. SW. of Scotland; D. A. Distler and W. N. Berg;
Sept. 3, 1960.

=KHtheostoma (Nothonotus) moore: Raney and Suttkus.

Etheostoma nevisense Cope, 1870, Proc. Amer. Philos. Soc., vol. 11,
p- 261.

Holotype: ANSP ?, (1, ?), North Carolina, falls of Neuse R., 8 mi.
E. of Raleigh. Types of other species described by Cope in the 1870
paper were deposited at ANSP, but Fowler (1907) did not mention
the type of this form, and we have been unable to find it.

=Percina (Alvordius) peltata nevisense (Cope) according to Dr.
Edward C. Raney (pers. comm.).

Etheostoma (Hadropterus) nianguae Gilbert and Meek in Gilbert 1887,
Proc. U.S. Nat. Mus., vol. 10, p. 52.

Lectotype: USNM 36214 (<7, 86), Missouri, Webster Co., head-
waters of Niangua R., near Marshfield; C. H. Gilbert and S. E. Meek;
summer, 1884; listed as type by Jordan and Evermann (1896, p. 1043);
selection herein confirmed. D XII-14; A IJ,12; LL 73; caudal
peduncle scale rows 12—2-13.

Paralectotype: SU 5299 (1, 82), same data as lectotype. The
original description says deposited in museum of University of
Cincinnati, but as Béhlke (1953, p. 73) noted, this specimen bears the
proper data and the word ‘“‘type.”’

=Etheostoma (Oligocephalus) nianguae Gilbert and Meek. See
Kuehne and Bailey (1961).

Etheostoma (Hadropterus) nianguae spilotum Gilbert, 1887. Proc. U.S.
Nat. Mus., vol. 10, p. 53.

Lectotype: USNM 38319 (1, 49), Kentucky, Owsley Co., Sturgeon
Cr., trib. of Kentucky R. near Traveler’s Rest; C. H. Gilbert; summer,
1884; listed as type by Jordan and Evermann (1896, p. 1044); selection
herein confirmed. D X-13; A II,11; LL 50+10=60; caudal peduncle
scale rows 10-210.

219-948—66——3

34 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Paralectotypes: SU 4036 (6, 16-67), same data as lectotype; 3
specimens (17-20), apparently Htheostoma (Boleosoma) sp., removed
and recataloged as SU 62429.

= Ktheostoma (Oligocephalus) sagitta spilotum Gilbert. See Kuehne
and Bailey (1961).

Etheostoma nigra Rafinesque, 1820, Ichthyologia Ohiensis, p. 37.
Types: The original description was based on material from the
Green R., Kentucky. As Call (1899) noted, Rafinesque did not
preserve any of his type material.
=Etheostoma (Boleosoma) nigrum nigrum Rafinesque.

Etheostoma nuchale Howell and Caldwell, 1965, Tulane Stud. Zool.,
vol. 12; no. 4p. 101; fiz 1)

Holotype: UMMZ 187523 (co, 39), Alabama, Jefferson Co., Glen
Spring, Bessemer on Co. rt. 20; W. M. Howell and R. D. Caldwell;
Mar. 24, 1964.

Paratypes: UMMZ 187524 (9, 40, allotype), UMMZ 187525 (20,
27-45), USNM 259800-F 1 (20, 22-48), UAIC 1227 (9, 24-43), same
collection as holotype. TU 34591 (20, 17-40), same locality; R. D.
Suttkus, J. S. Ramsey, and F. Rose; Sept. 9, 1964.

=KHtheostoma (Oligocephalus) nuchale Howell and Caldwell.

Etheostoma obeyense Kirsch, 1892, Bull. U.S. Fish. Comm., vol. 10,
p. 292, fig.

Lectotype: USNM 45565 (07, 52), Kentucky, Clinton Co., trib.
of Cumberland R.; P. H. Kirsch; fall, 1889 or spring, 1890; selected
and figured by Jordan and Evermann (1896, p. 1092 and 1900, fig.
463); selection herein confirmed. .

Paralectotypes: Kirsch collected a total of 210 specimens from
four Clinton Co. tributaries of the Cumberland in the fall of 1889
and spring of 1890: Indian Cr., 21; Spring Cr., 36; Smith Cr., 23;
and Albany Branch, 130. We have been able to locate the following:
SU 5116 (41, 28-51), Albany Branch, 2 Htheostoma spectabile removed
by Knapp and recataloged as SU 62132. SU 1918 (88, 32-55),
Spring Cr. This bottle was broken during the earthquake, which
may account for the presence of two more specimens. All are
E. obeyense, so perhaps Kirsch erred in his count of 36. BMNH
1892.12.30.136-139 (4, 35-41) Albany Branch, one small specimen of
E. (Oligocephalus) spectabile removed and recataloged as BMNH
1892.12.30.140.

USNM 46207 (6, 28-43), Wayne Co., Beaver Cr., 1891 were
labeled as types but were collected after the types and at a locality
not mentioned in the original description and so are not types.

=LHtheostoma (Catonotus) obeyense Kirsch.

NO. 3550 DARTERS—COLLETTE AND KNAPP 35

Etheostoma olmstedi D. H. Storer, 1842, Journ. Boston Soc. Nat.
Hist., vol. 4, p. 61, pl. 5, fig. 2.

Syntypes: MCZ 24781 (2c, 46-64), Connecticut, Connecticut
R. at Hartford; C. H. Olmsted; orig. MCZ 151.

=EKtheostoma (Boleosoma) olmstedi olmstedi Storer. See Stone
(1947) and Cole (1958, 1965). Described in January 1842 and so
has priority over Boleosoma tessellatum DeKay, July 1842. See
Storer (1843) and also summary of the meeting of Boston Society
of Natural History for June 21, 1843 (Proc. Boston Soc. Nat. Hist.,
wol. 1, p. 120).

Etheostoma (Hadropterus) ouwachitae Jordan and Gilbert in Gilbert,
1887, Proc. U.S. Nat. Mus., vol. 10, p. 49.

Syntypes: USNM 36449 (207, 40-48; 9, 43), Arkansas, Saline
Co., Saline R. at Benton; Jordan and Gilbert; summer, 1884. UMMZ
187509 (1, 42), same data, orig. IU 779, then IU 4958.

Boéhlke (1953, p. 74) listed SU 552 [(3, 31-47), Arkansas, Ouachita R.
at Arkadelphia; Jordan and Gilbert; summer, 1884] as paratypes,
but we do not feel that they should be considered part of the type
series because only the Saline River locality was listed, both in the
original description and in an advance list (Jordan and Gilbert,
1886, p. 138) where Hadropterus ouachitae appeared as a nomen nudum.

=Percina (Imostoma) uranidea (Jordan and Gilbert). See Hubbs
and Black (1940).

Etheostoma pagei Meek, 1894, Amer. Nat., vol. 27, p. 957.

Lectotype: USNM 45566 (<7, 50); Missouri, spring branch,
trib. of Neosho R. at U.S. Fish Hatchery, Neosho; S. E. Meek; Apr.
15, 1893; figured in Jordan and Evermann (1900, fig. 464); herein
selected. D IX-12; A JI, 7; LL 13+40=53; INF 8; cheeks with
small patch of scales; opercles naked; well-developed tubercles on
the anal fin characteristic of E. cragini (see Collette, 1965).

Paralectotype: USNM 197969 (9, 40), removed from USNM
45566.

=F theostoma (Oligocephalus) cragint Gilbert.

Etheostoma (Oligocephalus) pallididorsum Distler and Metcalf, 1962,
Copeia, no. 3, p. 556, fig. 1.

Holotype: KU 7144 (0, 42), Arkansas, Montgomery Co., Caddo
R., 8.5 mi. W. of Black Springs; Sec. 26, T3S, R27W; Distler and
Metcalf; June 28, 1961.

Paratypes: KU 6921 (21, 20-42); CU 44004 (10, 23-43), trans-
ferred to CU after orig. description; UMMZ 178997 (10, 34-41);
USNM 196547 (10, 33-46); same data as holotype. KU 6158 (18,
29-46), Arkansas, Montgomery Co., Caddo R., 1 mi. W. of Caddo

36 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Gap; Distler and Metcalf; Apr. 6, 1961. KU 6922 (8, 32-36),
Arkansas, Montgomery Co., trib. of Mill Cr., 2.56 mi. W. of Caddo
Gap; Distler and Metcalf; June 29, 1961.

= Ktheostoma (Oligocephalus) pallididorsum Distler and Metcalf.

Etheostoma (Etheostoma) parvipinne Gilbert and Swain in Gilbert,
1887, Proc. U.S. Nat. Mus., vol. 10, p. 59.

Holotype: USNM 36716 (1, 43), Alabama, small spring-branch trib.
of Black Warrior R. at Tuscaloosa; C. H. Gilbert and J. Swain;
summer, 1884.

=FEtheostoma (Oligocephalus) parvipinne Gilbert and Swain. See
Moore and Cross (1950).

Etheostoma peltatum Stauffer in Cope, 1864, Proc. Acad. Nat. Sci.
Philadelphia, vol. 16, p. 233.
Holotype: ANSP 22627 (0, 73), Pennsylvania, Lancaster Co.,
Conestoga Cr.; J. Stauffer.
=Percina (Alvordius) peltata peltata (Stauffer).

Etheostoma phoxocephalum Nelson, 1876, Bull. Illinois Mus. Nat. Hist.,
nos 4," ps3;

Syntypes: UMMZ 187504 (2.7, 49-52), Illinois; S. A. Forbes; orig.
IU 499, then IU 4702. Apparently syntypes but not labeled as such
or listed in the IU catalog as types. No syntypes are in the collection
of the Illinois Natural History Survey according to Smith and Bridges
(1960).

=Percina (Swainia) phorxocephala (Nelson). See Bailey (1941).

Etheostoma (Boleosoma) podostemone Jordan and Jenkins in Jordan,
1889, Proc. U.S. Nat. Mus., vol. 11, p. 359, pl. 45, fig. 11.

Lectotype: USNM 39863 (<’, 45), Virginia, Roanoke R. at Alle-
gheny Springs; Jordan and party; summer, 1888; herein selected by
Dr. Charles F. Cole. D X-14; A II, 8; LL 38, complete; pectoral
12-12; infraorbital canal complete, 8 pores; preoperculomandibular
pores 9; nape, cheeks, breast, and anterior part of belly naked; opercle
partially scaled.

Paralectotypes: USNM 197635 (4, 40-45), removed from USNM
39863.

USNM 125477 (3, 26-42), Virginia, Roanoke R. at Roanoke and
USNM 40305 (11, 24-41), Virginia, Roanoke R. are listed as paratypes
in the U.S. National Museum catalog. Bdohlke (1953, p. 74) con-
sidered SU 622 (8, 36-45), Virginia, Roanoke R. at Roanoke as
paratypes. CAS 22857, formerly IU 7866 (12, 22-44), also from the
Roanoke R. at Roanoke are labeled as paratypes. BMNH 1889.10.
30.45 (1, 43), Roanoke R., U.S. Fish Comm. is labeled as ‘‘one of the
types.”” However, Jordan and Jenkins designated USNM 39863,
containing five specimens, as “type,’’ and we believe this to be a

NO. 3550 DARTERS—COLLETTE AND KNAPP 37

designation of these specimens as syntypes from the type locality of
Roanoke River at Allegheny Springs. The other specimens should
then be considered as material examined and not part of the type
series.

=LEtheostoma (Boleosoma) podostemone Jordan and Jenkins. See
Cole (1958).

Etheostoma quapella Eigenmann and Eigenmann, 1892, Amer. Nat.,
vol. 26, p. 963. Described in more detail in Eigenmann (1894).
Holotype: BMNH 1892.12.30.183 (7, 36), Canada, Manitoba,
Qu’Apelle R. at Fort Qu’Apelle, Red R. system; C. H. Eigenmann;
August-September 1892.
=KHtheostoma (Oligocephalus) exile (Girard). See Jordan and
Evermann (1896) and Hubbs (1926, p. 68).

Etheostoma (Percina) rex Jordan and Evermann in Jordan, 1889, Proc.
Wess Nat. Must; vol. 11, p. 357, pl. 45, fig. 9.

Holotype: USNM 39858 (co, 122), Virginia, Roanoke R., near
Roanoke; Jordan; summer, 1888.

Paratype: Jordan and Evermann mentioned a second smaller
specimen taken with the type, but the whereabouts of this specimen
is unknown.

=Percina (Pereina) rex (Jordan and Evermann).

Etheostoma (Hadropterus) roanoka Jordan and Jenkins in Jordan, 1889,
Proce: U.S. Nat. Mus., vol. 11, p. 358, pl. 45, fig. 10.

Lectotype: USNM 39866 (<7, 48), Virginia; Roanoke R. at Alle-
gheny Springs; Jordan and party; summer, 1888. USNM 39866, then
containing at least five specimens, given as ‘“‘type” in original descrip-
tion; one specimen herein selected as lectotype by Dr. Edward C.
Raney. D XI-12; A II,9; LL 47; cheeks naked; breast naked except
for one large modified scale; nape naked anteriorly, with a few scales
posteriorly; opercles with scales at least on upper portion; suborbital
bar prominent; sides with 10 vertical bars tending to form a lateral
band.

Paralectotypes: USNM 197774 (4 o, 43-48), removed from USNM
39866. MOZ 3106 (7, 40); CU 46469 (3, 43-48); CU 46470 (1, 48),
found with CU 46469 but probably out of MCZ 3106; SU 616 (9, 41—
53), one specimen of Percina peltata removed; and UMMZ 187514
(7, 24-45), orig. IU 1143, then IU 5619, listed in old IU catalog as
type, one specimen of Etheostoma podostemone removed and re-
cataloged as UMMZ 187515; all from Roanoke R. at Roanoke.

In addition to the collections from the Roanoke River at Roanoke
and Allegheny Springs listed above, specimens from two other rivers
are mentioned in the original description. We feel that only the speci-

38 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

mens from the Roanoke River should be considered as types, because
Jordan and Jenkins designated a Roanoke series as “type,” and
because they specifically mentioned that the North Carolina speci-
mens had larger scales than the Roanoke material. Therefore, we
consider the following specimens from the Neuse and Tar rivers in
North Carolina as other material, not as types. SU 17777 (1, 48),
North Carolina, Neuse R. at Raleigh; listed by Béhlke (1953, p. 74)
as a paratype. USNM 125359 (1, 42), same data; listed as a paratype
in the USNM catalog. USNM 40367 (13, 29-48) and ANSP 707434
(2, 33-38), out of USNM 40367; same data, not listed as type in
USNM catalog. UMMZ 187516 (0, 69), ong. IU 1158, then’ 1
7998, part of the same series from the Neuse R.; not listed as type in
IU catalog. UMMZ 187517 (1, 33), North Carolina, Tar R. at
Rocky Mount; orig. IU 1144, then IU 7888, not listed as type in IU
catalog; two specimens of Percina peltata removed and recataloged
as UMMZ 187518.

=Percina (Ericosma) crassa roanoka (Jordan and Jenkins), accord-
ing to Dr. Edward C. Raney (pers. comm.).

Etheostoma (Rhothoeca) rupestre Gilbert and Swain in Gilbert, 1887,
Proc. U.S. Nat. Mus., vol. 10, p. 57.

Lectotype: USNM 36695 (9, 32), Alabama, North R., trib. of
Black Warrior R., near Tuscaloosa; C. H. Gilbert and J. Swain;
summer, 1884; herein selected by Dr. Chu-fa Tsai. D XI-12 AIL:
LL 52; INF 4+3; POM 9; cheeks and breast naked; opercle with 2
imbedded scales; belly scaled anteriorly; 6 dorsal blotches.

Paralectotypes: USNM 188971 (17, 23-34), removed from USNM
36695.

= LEtheostoma (Etheostoma) rupestre Gilbert and Swain.

Etheostoma sciotense Osburn and Williamson, 1898, Ann. Rept. Ohio
State Acad. Sci., vol. 6, p. 17.

Lectotype: USNM 48846 (<7, 29), Ohio, Franklin Co., Big Wal-
nut Cr., trib. of Scioto R., near Columbus; B. C. Osburn and E. B.
Williamson; June 26, 1897; herein selected. D XIT-12; Avt7:
LL 27+19=46; anterior third of belly naked.

Paralectotypes: USNM 197995 (3, 24), removed from USNM
48846. A third syntype was deposited in Jordan’s collection, but
we have been unable to find this specimen.

= LEtheostoma (Nothonotus) tippecanoe Jordan and Evermann. See
Osburn and Williamson (1899).

Etheostoma scovellii Woolman, 1892, Amer. N at., vol. 26, p. 260.
Lectotype: USNM 44155 (1, 33), Mexico, Rio de las Conchos,
Chihuahua; A. J. Woolman; August 1891 ; herein selected. D XI-11;

No. 3650 DARTERS—COLLETTE AND KNAPP 39

A 1,7; LL 44+15=59; INF 8; interorbital pores 2; cheeks, opercles,
nape, and breast naked.

Paralectotypes: USNM 197970 (2, 27-34), out of USNM 44155.
BMNHE 1892.12.30.186-188 (3, 29-32), same data. CAS 22856 (3,
20-25), same data; orig. 1U 4841.

—Etheostoma (Austroperca) australe Jordan. See Hubbs (1936).

Etheostoma (Hadropterus) squamatus Gilbert and Swain in Gilbert,
1887, Proc. U.S. Nat. Mus., vol. 10, p. 50.
Holotype: USNM 36652 (<7, 85), Tennessee, French Broad R. at
mouth of Wolf Cr.; C. H. Gilbert and J. Swain; summer, 1884.
— Percina (Swainia) squamata (Gilbert and Swain).

Etheostoma squamiceps Jordan, 1877, U.S. Nat. Mus. Bull. 10, p. 1.

Lectotype: USNM 1145 (<7, 57), Kentucky, Logan Co., Russell-
ville; Bebb; erroneously given as USNM 1345 in orig. description;
herein selected. D IX-13; A II,7; LL 45+3=48; POM 10; INF
incomplete; cheek and opercle covered with scales; small unpigmented
fleshy bulbs around tips of first dorsal spines.

Paralectotype: USNM 197968 (9, 54) removed from USNM 1145.

—Etheostoma (Catonotus) squamiceps Jordan. See Moore and
Cross (1950, p. 145).

Etheostoma (Nanostoma) swannanoa Jordan and Evermann in Jordan,
1889, Proc. U.S. Nat. Mus., vol. 11, p. 360, pl. 45, fig. 13.

Lectotype: USNM 39861 (9, 59), Virginia, S. Fork of Holston R.
at Holstein Mills; D. S. Jordan and B. W. Evermann; summer, 1888;
USNM 39861 given as type by Jordan and Evermann but then con-
taining two specimens; lectotype herein selected by Dr. William J.
Richards. D XII-12; A II,9; pectoral 16-16; LL 57; INF 8; POM
10; head, cheeks, opercles, and breast naked; vomerine teeth present;
palatine teeth absent.

Paralectotypes: USNM 196777 (9, 56), removed from USNM 39861.
Jordan and Evermann designated USNM 39861 as “‘type,” but they
gave counts of the lateral line scales in 8 of the 10 specimens they
had before them. Therefore, their description was apparently based
on all their material, and the following are also considered paralecto-
types. USNM 125490 (9, 40), North Carolina, S. Fork of Swannanoa
R. at Black Mountain; Jordan and Evermann; August 1888; orig.
BF 772. SU 3507 (9, 41), same locality as USNM 125490. Bohlke
(1953, p. 74) gave the same data for SU 992 2 &%, 54-57) as for USNM
39861, but Jordan and Evermann reported only two specimens from
this locality, so it seems likely that SU 992 is from Middle Fork of
Holston R. at Marion or S. Fork of Swannanoa R. at Black Mountain.
UMMZ uncat. (<7, 49), orig. IU 1169, then IU 8083. Locality in
IU catalog for IU 8083 is N. Fork of Holston R. at Saltville, Va., but

40 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Jordan and Evermann did not report this species from there. Locality
in the IU catalog for 1169 is apparently S. Fork of Holston R. at
Holstein Mills, Va. However, as with SU 992, this specimen must
be from Middle Fork of Holston R. at Marion or 8S. Fork of Swanna-
noa R. at Black Mountain.

—Ktheostoma (Etheostoma) swannanoa Jordan and Evermann.

Etheostoma tessellata D. H. Storer, 1845, Proc. Boston Soc. Nat.
Hist., vol. 2, p. 48. Original description repeated almost ver-
batim in Storer (1846).

Syntypes: Alabama, Florence; received from C. A. Hentz. We
have been unable to find any type material of this nominal form.
=?Percina (Ericosma) evides (Jordan and Copeland) according to

Bailey (1959, p. 2). Although £. tessellata Storer is a senior synonym

of Alvordius evides Jordan and Copeland, 1877, it is not available be-

cause tessellata is preoccupied in Htheostoma by Boleosoma tessellatum

DeKay, 1842.

Etheostoma tippecanoe Jordan and Evermann, 1890, Proc. U.S. Nat.
Mus., vol. 13, p. 3 [with fig. of FE. camurum].

Lectotype: USNM 40080 (1, 29), Indiana, Tippecanoe R. at Marsh-
land; B. W. Evermann; herein selected. D XII-13; A II, 8; LL
37+16=53; cheeks, prepectoral area, nape, and breast naked; opercle
scaled; anterior part of belly naked along midline.

Paralectotype: USNM 197919 (1, 26), out of USNM 40080.

Four specimens are mentioned in the original description of EF.
tippecanoe. The two USNM specimens fit the original description
and the current concept of EH. tuppecanoe. The other two syntypes
apparently are UMMZ 187513 (2, 29-44), orig. IU 1194, then IU
4688. They have the belly completely scaled and lateral line scale
counts of about 54+0=—54 and 66+1=67. The larger one is a speci-
men of £. camurum and the smaller one probably is also. Jordan and
Evermann (1896, p. 1090) wrote that the original description of F.
tippecanoe was accompanied by a figure of EH. camurum “by an error
of printer.” We believe that the specimen figured in Jordan and
Evermann is the smaller specimen in UMMZ 187513. We count the
same number of lateral line scales on the figure and the small specimen.
The original drawing is labeled as a specimen 1%, in. long (total
length) from no. 40080. ‘This is equal to a standard length of about
29mm. ‘Three sets of comments are written on the original drawing.
“The drawing is bad, but it was made from specimens sent to the
National Museum as types of FE. tippecanoe.”’ ‘This is probably E.
camurum. Certainly not E. tippecanoe. Anal fin very bad.” ‘‘Fear-
ful and not taken from our type.”

NO. 3550 DARTERS—COLLETTE AND KNAPP |

=LHtheostoma (Nothonotus) tippecanoe Jordan and Evermann.
See Bailey (1959).

Etheostoma (Psychromaster) trisella Bailey and Richards, 1963, Occ.
Pap. Mus. Zool. Univ. Michigan, no. 630, pl. 1, D-E.

Holotype: UMMZ 180073 (1, 35), Alabama, Cherokee Co.,
Cowans Cr., trib. of Spring Cr. which flows into Coosa R. at Jordans
Farm on U.S. 411, 6.7 mi. SE. of Centre; E. A. Lachner and P. S.
Handwerk; Sept. 13, 1947; orig. CU 18579.

=Ktheostoma (Psychromaster) trisella Bailey and Richards.

Etheostoma (Etheostoma) tuseumbia Gilbert and Swain in Gilbert, 1887,
Proc. U.S. Nat. Mus., vol. 10, p. 63.

Syntypes: SU 1003 (80, 25-48) and BMNH 1894.5.18.26-27
(2, 28-31), large spring at Tuscumbia flowing into Tennessee R.;
C. H. Gilbert and J. Swain; summer, 1884; one of types figured by
Jordan and Evermann (1900, fig. 467). Apparently UMMZ 187512
(44, 22-35), orig. IU 1380, then IU 5666, specimens almost completely
disintegrated, are also syntypes, although they are not listed as types
in the IU catalogs.

USNM 36154 with the same data also originally contained syntypes.
This jar was found completely dried out in March 1937 and only
fragments now remain. We believe that not many specimens were
in the jar at that time because old correspondence of Barton Bean
shows that at least 37 specimens from this series were sent out as
“duplicate sets’ to 37 high schools and other such institutions during
the period Aug. 4, 1902 to Dec. 12, 1903. We assume that Bean was
unaware that USNM 36154 contained types, for he certainly would
not have sent type material to such institutions as the public schools
of Kent, Wis.; Omaha, Nebr.; Windfall, Ind.; Galliopolis, Ohio;
Helena, Mont.; Savanna, IIll.; Northwood, Iowa; Lovington, IIl.;
Wiconisco, Pa.; the Free Public Library of Vinton, Iowa; the San
Antonio Female College, Tex.; and the Ursuline Seminary, New
Rochelle, N.Y.!

Bohlke (1953) and Bailey and Richards (1963) gave USNM 36154
as holotype. Bdéhlke listed SU 1003 as paratypes. From the number
of specimens distributed out of USNM 36154, it is clear that all of
the original type material should be listed as syntypes.

Bohlke erroneously listed SU 2683 (2, 27-28) as paratypes, but
these specimens were collected at Florence, Ala., a locality not men-
tioned in the original description. Bailey and Richards (1963) listed
USNM 43484 and 63115 and UMMZ 43484 as paratopotypes, but
this material was collected by Kirsch in 1889, subsequent to the
original description of H. tuscumbia, and so they are merely topotypes.

42 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

=Ftheostoma (Psychromaster) tuscumbia Gilbert and Swain. See
Bailey and Richards (1963).

Etheostoma (Cottogaster) uranidea Jordan and Gilbert in Gilbert, 1887,
Proc. U.S. Nat. Mus., vol. 10, p. 48.

Syntypes: USNM 36413 (8, 31-46), Arkansas, Washita (Ouachita)
R. at Arkadelphia; Jordan and Gilbert; summer, 1884. One specimen
of Ktheostoma histrio removed and recataloged as USNM 197419.

=Percina (Imostoma) uranidea (Jordan and Gilbert).

Etheostoma variata Kirtland, 1841, Boston Journ. Nat. Hist., vol. 3,
p. 274, pl. 2, fig. 2.
Types: Type locality—Mahoning R., Ohio. We have been unable
to find any extant type material.
= KEtheostoma (Etheostoma) variatum Kirtland.

Etheostoma (Ulocentra) verecundum Jordan and Evermann in Jordan,
1889, Proc. U.S. Nat. Mus., vol. 11, p. 360, pl. 45, fig. 12.
Holotype: USNM 39862 (1, 44), Virginia, Middle Fork of Holston
R., 5 mi. S. of Glade Spring; summer, 1888.
=Etheostoma (Ulocentra) simoterum (Cope).

Etheostoma (Etheostoma) whippler alabamae Gilbert and Swain in
Gilbert, 1887, Proc. U.S. Nat. Mus., vol. 10, p. 62.

Syntypes: USNM 36735 (6, 25-30), Alabama, Black Warrior R..,
near Morris; C. H. Gilbert and J. Swain; summer, 1884. USNM
36687 (2, 29-41), and SU 2318 (2, 35-41), Alabama, North R.., trib.
of Black Warrior R., near Tuscaloosa; Gilbert and Swain; summer,
1884.

=Ktheostoma (Oligocephalus) whipplvi artesiae (Hay). See Hubbs
and Black (1941).

Etheostoma wrightt McCormick, 1892, Bull. Oberlin College Lab., no.
2, p. 30.

Holotype: Oberlin Coll. Mus. (1, 2% in.), Ohio, Lorain Co.,
Vermillion R., trib. of Lake Erie; L. M. McCormick; Apr. 9, 1892.
This specimen was not present in the Oberlin collection in mid-
January 1931, according to Trautman (1957, p. 548), and has not
been located.

=Percina (Imostoma) shumardi (Girard), according to Jordan and
Evermann (1896, p. 1047) and Jordan, Evermann, and Clark (1930,
p. 285). The type locality and original description support this
conclusion, but Trautman (1957, p. 548) has suggested that EL. wrights
may be a hybrid between P. shumardi and P. caprodes or P. maculata,
or between P. caprodes and P. maculata.

Etheostoma zonale arcansanum Jordan and Gilbert, 1886, Proc. U.S.
Nat. Mus., vol. 9, p. 5.

No. 3550 DARTERS—COLLETTE AND KNAPP 43

Lectotype: USNM 36410 (9, 38), Arkansas, Washita (Ouachita)
R. at Arkadelphia; D. S. Jordan and C. H. Gilbert; summer, 1884;
herein selected by Dr. Chu-fa Tsai. D XI-11; A II,7; LL 56; INF
4+3; POM 10; cheeks, opercles, and belly completely covered with
scales; a few scales on breast; 6 dorsal blotches.

Paralectotypes: USNM 188970 (25, 24-36), removed from USNM
36410. UMMZ 187506 (7, 25-36), orig. IU 777, then IU 4543, same
data. USNM 36249 (2, 40-43), Missouri, Spring Fork of Neosho R.,
near Carthage; Gilbert and S. E. Meek. USNM 36275 (1, 41),
Missouri, James Fork of White R., near Marshfield; Gilbert and Meek.
USNM 36399 (2, 30-31), Oklahoma, Poteau R., W. of Hackett City,
Ark.; Jordan and Gilbert; summer, 1884. USNM 36447 (15, 21-35),
Arkansas; Saline R., near Benton, just above railroad bridge; Jordan
and Gilbert; summer, 1884; six specimens of Etheostoma r. radiosum
removed. UMMZ 187507 (4, 22-29), same locality, orig. IU 779,
then IU 4546; one specimen of F. histrio removed and recataloged
as UMMZ 187508.

=KHtheostoma (Etheostoma) zonale zonale (Cope), according to
Dr. Chu-fa Tsai (pers. comm.).

Hadropterus evermanni Moenkhaus, 1903, Bull. U.S. Fish Comm., vol.
22° pe 390, he:

Holotype: Given as IU 9785 (1, 77), Indiana, Lake Tippecanoe in
original description, but we have not located this specimen.

Paratypes: USNM 50834 (1, 48) and USNM 126919 (1, 48); 1899;
orig. BF 2742; Lake Tippecanoe. We have not found the paratype
originally cataloged as IU 9786 nor the specimen said to have been
deposited at Stanford University. There appears to be no record of
a paratype having been deposited in the British Museum, and Collette
could find none among the type material or in the regular collections
in October 1963.

=Percina (Percina) caprodes semifasciata (DeKay) X Percina
(Alvordius) maculatus (Girard). See Hubbs (1926, p. 60). The two
USNM specimens and the figure of the holotype appear to be inter-
mediate between P. caprodes and P. maculatus in such characters as
color pattern, number of lateral line scales, juncture of gill membranes,
nape squamation, and snout shape. Trautman (1948) has also
reported this natural hybrid combination from southwestern Lake
Erie and tributaries that enter the lake in this region.

Hadropterus maculatus Girard, 1859, Proc. Acad. Nat. Sci. Philadelphia,
vol. 11, p. 100.
Lectotype: USNM 1157 (<7, 69), Maryland, Anne Arundel Co.,
eastern trib. of Potomac R.; J. H. Clark; selected herein by Dr.
Edward C. Raney. D XIV-12; A II,9; pectoral rays 14-13; LL 55;

44 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

scales above and below lateral line 7-10; caudal peduncle scales 21;
cheeks sparsely scaled; opercles scaled; preopercles and nape naked;
11 enlarged scales on midline of belly. This specimen is also the
type of Hadropterus notogrammus Raney and Hubbs.

Paralectotype: USNM 197773 (0, 56) removed from USNM 1157.

MCZ 24525 (2 o, 60-64), originally USNM 1308, then MCZ 129,
and MCZ 24546 (2 o&, 54-55), originally USNM 1157, then MCZ
130, have the same data and were probably part of the original series;
however, they were received from the USNM in 1853, but the descrip-
tion was not published until 1859, so it is doubtful if Girard had the
specimens before him at the time of the original description. There-
fore, the four specimens had best be considered only as topotypes,
both of H. maculatus Girard and H. notogrammus Raney and Hubbs.

=Percina (Alvordius) notogramma notogramma (Raney and Hubbs)
because maculatus is preoccupied by Alvordius maculatus Girard. See
Hogarth and Woolcott (1966).

Hadropterus maxinkuckiensis Evermann, 1900, Rept. U.S. Fish Comm.,
VOl. 25. Pp. o00, pl. 17.

Holotype: USNM 49378 (<7, 75), Indiana, Marshall Co., Aubeenau-
bee Cr., E. inlet of L. Maxinkuckee, 0.5 mi. from lake; Evermann and
Scovell; Aug. 4, 1899.

=Percina (Alvordius) maculata (Girard). See Hubbs and Black
(1954).

Hadropterus nasutus Bailey, 1941, Oce. Pap. Mus. Zool. Univ.
Michigan, no. 440, figs. 1, 3.

Holotype: UMMZ 132898 (7, 66), Arkansas, Searcy Co., Middle
Fork, Little Red R., trib. of White R., near bridge on U.S. 65, 1.5 mi.
SE. of Leslie; R. M. Bailey and M. E. Davis; Aug. 17, 1940.

Paratype: USNM 198146 (9, 63), same data as holotype; orig.
ISC 2.

=Percina (Swainia) nasuta (Bailey).

Hadropterus nigrofasciatus Agassiz, 1854, Amer. Journ. Sci. and Arts,
Serv vols tip. 0b:

Lectotype: MCZ 24603 (co, 103), Alabama, near Mobile, A. Stein;
orig. out of USNM 1197, then MCZ 122; herein selected. D XII-12;
A II, 9; LL 50; preopercle without serrae; branchiostegal membranes
slightly connected; nape, cheek, opercle, and breast completely scaled.

Paralectotypes: MCZ 43094 (11, 41-98) and UMMZ 86300 (3,
51-77), removed from MCZ 24603. USNM 1197 (7, 74; 2, 82),
same data.

=Percina (Hadropterus) nigrofasciata nigrofasciata (Agassiz). See
Crawford (1956).

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NO. 3550 DARTERS—COLLETTE AND KNAPP 45

Hadropterus nigrofasciatus westfalli Fowler, 1942, Notulae Naturae,
no. 107, p. 9, fig. 9.
Holotype: ANSP 69980 (o, 44), Florida, Wekiva R., trib. of
St. Johns R.; M. J. Westfall, Jr.; Mar. 8, 1941.
=Percina (Hadropterus) nigrofasciata nigrofasciata (Agassiz). See
Crawford (1956).

Hadropterus notogrammus Raney and Hubbs, 1948, Occ. Pap. Mus.
Zool. Univ. Michigan, no. 512, pl. 1, figs. 1-2; pl. 2, figs. 1-2; a
replacement name for Hadropterus maculatus Girard which was
preoccupied.

Types: Raney and Hubbs designated UMMZ 144696 [(<7, 67),
Virginia, Rockbridge Co., South R., 5 mi. E. of Lexington, L. C.
Pettit; May 17, 1941; orig. CU 9719] as holotype; and USNM
40242; CU 8324, 9708, 10138, 4909; UMMZ 144697 and 95194 as
paratypes. However, these designations are not valid because the
type specimens for a new name must be the same as for the
original name (Art. 72 d, Int. Code Zool. Nomencl.), in this case
USNM 1157, lectotype and USNM 197773, paralectotype of Hadrop-
terus maculatus Girard.

=Percina (Alvordius) notogramma notogramma (Raney and Hubbs).
See Hogarth and Woolcott (1966).

Hadropterus oxyrhynchus Hubbs and Raney, 1939, Occ. Pap. Mus.
Zool. Univ. Michigan, no. 396; pl. 1.

Holotype: UMMZ 118422 (<7, 80), West Virginia, Randolph Co.,
Cheat R. at Cheat Bridge; E. C. Raney; June 25, 1935.

Paratypes: USNM 56862 (<7, 63), West Virginia, Cheat R. at
Raines; W. P. Hay; Sept. 12, 1899. UMMZ 118482 (4, 44-54),
Virginia, Wythe Co., New R., above Jacksons Ferry; C. L. Hubbs
and family; Aug. 21, 1936. UMMZ 119268 (7, 20-33), West Virginia,
New R. at Prince; J. Addair; July 26, 1985. UMMZ 119246 (2,
30-31), West Virginia, New R., 2 mi. below Hinton; Addair; July 18,
1935.

—=Percina (Swainia) oxyrhyncha (Hubbs and Raney). See Bailey
(1941).

Hadropterus palmaris Bailey, 1940, Journ. Washington Acad. Sci.,
vol. 30, Nos 12... 525, fies 1%

Holotype: UMMZ 126179 (<7, 66), Georgia, Lumpkin Co., Etowah
R., trib. of Coosa R., just above bridge on U.S. 19, 4 mi. SW. of
Dahlonega; R. M. and M. K. Bailey; Aug. 25, 1939.

Paratypes: UMMZ 126180 (2, 45-67), USNM 117881 (2 o, 61-68)
and USNM 198145 (co, 63), orig. ISC 1, same data as holotype.
UMMZ 111232 (1, 59) and UMMZ 111183 (22, 31-63), Alabama,
Saugahatchee Cr., trib. of Tallapoosa R.; F. E. Guyton; Oct. 24,

46 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

1930. UMMZ 111234 (4), Alabama, Macon Co., 3 mi. E. of Tuskegee;
Guyton; June 3, 1931; there is a note in the UMMZ ecard file that these
specimens were not located during a check several years ago, and we
were unable to find them in May 1964.

=Percina (Ericosma) palmaris (Bailey). See Crawford (1954).

Hadropterus pantherinus Moore and Reeves, 1955, Copeia, no. 2, p. 90,
pl. de

Holotype: UMMZ 167120 ( 9, 63), Oklahoma, Pushmataha Co.,
Little R., W. of Pickens, sec. 1, T. 2S., R. 20 E.; G. A. Moore and
J. D. Reeves; Mar. 25, 1951.

Paratypes: UMMZ 81185 (1, 59), Arkansas, Polk Co., trib. of
Mountain Fork R.; June 29, 1927; Univ. Oklahoma Mus. Exped.
OAM 2559 (1, 38), Oklahoma, McCurtain Co., Mountain Fork R..,
Beavers Bend State Park, sec. 10, T. 5 S., R. 25 E.; Moore and class;
Aug. 21, 1948. UMMZ 167122 (3, 22-27), same locality as OAM
2559, Moore and F. B. Cross; May 30, 1948. USNM 198108 (i, 35),
Oklahoma, LeFlore Co., Little R., S. of Honobia, sec. 30; TT IN. AR:
22 E.; A. P. Blair; orig. TUC uncat. MCZ 38491 (1, 41), Oklahoma,
McCurtain Co., Mountain Fork R. at Smithville, sec. 13, T.1S., R.
25 E.; Moore and C. C. Rigney; Aug. 17-19, 1948. UMMZ 167121
(6, 48-72), same locality as UMMZ 167120; Moore and Reeves; Mar.
25, 1951. OAM 4948 (1, 52), Oklahoma, McCurtain Co., Glover R.
in sec. 13, T. 3 S., R. 22 E.; Moore and Reeves; Mar. 25, 1951.

=Percina (Alvordius) pantherina (Moore and Reeves).

Hadropterus scierus Swain, 1883, Proc. U.S. Nat. Mus., vol. 6, p. 252.

Lectotype: USNM 34350 (co, 57), Indiana, Monroe Co., Bean
Blossom Cr., trib. of White R., 6 mi. N. of Bloomington; D. S. Jordan
and J. Swain; herein selected. D XIII-14; A II,10; LL complete, 71;
preopercle serrate; branchiostegal membranes moderately joined.

Paralectotypes: Two specimens were mentioned in the original
description, but we do not know if the second syntype is still in
existence.

=Percina (Hadropterus) sciera sciera (Swain). See Hubbs and
Black (1954).

Hadropterus scierus apristis Carl L. Hubbs and Clark Hubbs in Clark
Hubbs, 1954, Amer. Mid. Nat., vol. 52, no. iL pte lle

Holotype: UMMZ 162377 (<7, 68), Texas, Caldwell-Guadalupe
cos., San Marcos R., 0.5 mi. SE. of Prairie Lea; K. C. Jurgens and
R. C. MacEwan; Feb. 11, 1950.

Paratypes: UMMZ 162378 (40, 46-74), same data as holotype.
USNM 160628 (6 9, 48-61; 5 <7, 57-83), Texas, Caldwell-Guadalupe
cos., San Marcos R. at Stairtown; Jurgens and MacEwan ; Feb. 18,
1950, orig. TNHC 255. USNM 160629 (11 9, 50-69, 1 &, 73),

NO. 3550 DARTERS—COLLETTE AND KNAPP AT

Texas, Caldwell-Guadalupe cos., San Marcos R., 0.5 mi. S. of Prairie
Lea; Jurgens and W. Brown; Feb. 8, 1950; orig. TNHC 180. UMMZ
162374 (9, 41-72), Texas, Hays Co., San Marcos R., below cotton gin
near State Fish Hatchery; C. Hubbs and Jurgens; Sept. 26, 1949;
orig. TNHC 21. FMNH 46194 (13, 27-64), Texas, San Marcos R.,
3 mi. SE. of State Fish Hatchery; Jurgens and Hubbs; Sept. 26, 1949;
orig. TNHC 40; specimens partly dried out; erroneously given as
FMNH 46195 in original description. OAM 4707 (8, 42-63), Texas,
Caldwell-Guadalupe cos.; San Marcos R. at Fentress; Jurgens and
Brown; Nov. 18, 1949. TNHC 90 (15, 28-56), Texas, Caldwell-
Guadalupe cos.; San Marcos R., 0.5 mi. N. of Staples; Jurgens; Nov.
11,1949. TNHC 501 (1, 75), Texas, Gonzales Co., Guadalupe R. at
rt. 80, S. of Belmont; Hubbs; Apr. 6, 1950. TNHC 162 Cle 7G):
Texas, Gonzales Co., Guadalupe R., 4 mi. SE. of Gonzales; Jurgens.
TNHC 314 (1, 66), Texas, Comal Co., Guadalupe R. at Hunter;
Jurgens and Brown; Mar. 22, 1950. SU 17311 (3), Texas, Caldwell-
Guadalupe cos.; San Marcos R. at Stairtown; Mar. 25, 1950; Jurgens
and W. Faber; orig. TNHC 333; specimens now completely disinte-
grated. TAMC 1F-13-g-10 (16, 35-78), Texas, Caldwell-Guadalupe
cos., San Marcos R. at Martindale; Jurgens and Brown; Nov. 6, 1949.
TAMC 1F-13-¢-4 (1, ?), Texas, Gonzales Co., Guadalupe R., 0.5 mi.
S. of Gonzales; F. T. Knapp; Nov. 26, 1950; dried out.
—Percina (Hadropterus) sciera apristis (Hubbs and Hubbs).

Hadropterus scierus serrula Jordan and Gilbert, 1886, Proc. U.S. Nat.
Mus., vol. 9, p. 16.

Lectotype: USNM 36481 (3, 50), Texas, Sabine Reb) miesgek
Longview; D. S. Jordan and C. H. Gilbert; summer, 1884; herein
selected. D XII-14; A 11,9; LL complete, 65; preopercular serrae
10=13:

Paralectotypes: USNM 198010 (5, 43-47), removed from USNM
36481.

Jordan and Gilbert had other material of their new variety from the
Trinity River at Dallas, Tex. (USNM 36476) and from the Rio
Lampasas at Belton, Tex. Evermann and Kendall (1894, pp. 88,
113) gave the type locality as the Sabine River at Longview. Jordan
and Evermann (1896, p. 1038) gave the “type” as USNM 36481
and the type locality as the Red River at Fulton, Ark. Hubbs
and Black (1954, p. 205) pointed out that Jordan and Evermann did
not collect serrula in the Red River and no. 36481 is from the Sabine.
Hubbs and Black designated USNM 36481 “as the sole lectotypes.”
We feel that Jordan and Evermann meant USNM 36481 to be the
“types” in the first place, so Hubbs and Black’s action was not neces-
sary. Béhlke (1953, p. 74) listed SU 3934 from Rio Lampasas as

48 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

paratypes and USNM 36481 as holotype. SU 3934 cannot be para-
types because USNM 36481 from the Sabine contains the entire type
series as originally indicated by Jordan and Gilbert and confirmed by
Evermann and Kendall (1894), Jordan and Evermann (1896), and
Hubbs and Black (1954).

=Percina (Hadropterus) sciera ssp. Further study is necessary to
determine whether or not serrula can be recognized as a valid sub-
species. See Hubbs and Black (1954) and Hubbs (1954),

Hadropterus sellaris Radcliffe and Welsh, 1913, Bull. U.S. Bur. F ish.,
VOl. 32, p: 31, pl. 18:
Holotype: USNM 74346 (9, 39), Maryland, Harford Co., Swan Cr,
near Havre de Grace; L. Radcliffe and W. W. Welsh; May 2, 1912.
Paratype: USNM 74347 (9, 39), same data as holotype.
=Ktheostoma (Etheostoma) sellare (Radcliffe and Welsh). See
Hubbs and Black (1940) and Knapp et al. (1963).

Hadropterus shumardi Girard, 1859, Proc. Acad. Nat. Sci. Philadelphia,
voll sp. 100:
Holotype: USNM_ 1330 (@, 48), Arkansas, Arkansas R., near
Ft. Smith; G. C. Shumard.
=Percina (Imostoma) shumardi (Girard).

Hadropterus spillmani Hay, 1881, Proc. U.S. Nat. Mus., vol. 3, p. 491.

Lectotype: USNM 27432 (co, 64), Mississippi, Clarke Co., Chicka-
sawha R. at Enterprise; O. P. Hay; March-April 1880; listed as type
by Jordan and Evermann (1896, p. 1039); selection herein confirmed.
D XII-12; A II,9; pectoral 14-15; LL 54; scales above and below
lateral line 6-11; cheeks, opercles, breast, and nape completely covered
with scales; several serrations on preopercle.

Paralectotypes: The original description was based on “several
specimens” of which some were placed “in Professor Jordan’s collec-
tion” and the remainder in Butler University. These specimens are
apparently no longer in existence.

=Pereina (Hadropterus) nigrofasciata nigrofasciata (Agassiz). See
Crawford (1956, p. 41).

Hadropterus tessellatus Jordan, 1877, U.S. Nat. Mus. Bull. 10) paw.

Lectotype: USNM 1199 (1, 47), Pennsylvania, Allegheny R. at
Foxburg; S. F. Baird; herein selected. D RIMS 12 A TO at
pectoral rays 16; INF 8, fourth pore opening at the end of a dorsally
directed tube beneath the eye; POM 10; four blackish dorsal saddles :
cheeks naked; opercles and breast with few scales.

Paralectotype: USNM 197416 (1, 29), removed from USNM 1199.
This smaller specimen was found in the bottle labeled USNM 1199.
This might not be part of the type material because Jordan referred

NO. 3550 DARTERS—COLLETTE AND KNAPP A9

to ‘a specimen of an Etheostomoid” and “length of type 2% inches”
and so may have examined only the larger specimen. However,
Jordan (1885, p. 163) mentioned two specimens as “the types of
Hadropterus tessellatus.”’

—Etheostoma (Etheostoma) variatum Kirtland. See Jordan (1885)
and Hubbs and Black (1941, p. 15).

Hololepis barratti appalachia J. R. Bailey, 1950, Copeia, no. 4., p. 312.

Holotype: UMMZ 156224 (o, 37), North Carolina, Buncombe
Co., pond S. of mouth of Bent Cr., 300 ft. W. of French Broad R.,
7 mi. SSW. of Asheville; F. Cross and Y. Barber; July 14, 1947.

Paratypes: UMMZ 156225 (48, 13-42), same data as holotype.
CU 18444 (3, 30-40), and DU uncat. (69, 19-40), same locality;
June 7, 1949; J. R. Bailey.

—Etheostoma (Hololepis) fusiforme barratti (Holbrook). See Col-
lette (1962).

Hololepis collis Hubbs and Cannon, 1935, Mise. Publ. Mus. Zool.
Univ. Michigan, no. 30, p. 52, pl. 1, fig. 7.

Holotype: UMMZ 94560 (<7, 40), South Carolina, York Co.,
creek near York; D. Ameel; Noy. 11, 1931.

Paratypes: UMMZ 107085 (9, 40), same data as holotype.
UMMZ 94546 (<7, 36), South Carolina, York Co., Steel Cr., trib. of
Catawba R. at Rock Hill; Ameel; Nov. 11, 1931.

—Etheostoma (Hololepis) collis collis (Hubbs and Cannon). See
Collette (1962).

Hololepis erochrous Cope, 1864, Proc. Acad. Nat. Sci. Philadelphia,
vol. 16, p. 233.

Lectotype: MNHN 4669 (1, 33), New Jersey, Delaware drainage,
Brown’s Mills; herein selected. D IX-10; A II, about 7; LL17+31=
48; cheeks, nape, breast scaly; interorbital naked. Fowler (1907),
Hubbs and Cannon (1935), and Collette (1962) were unable to locate
any types of this form at ANSP. Vaillant (1873, p. 134) wrote
‘“Texemplaire suivant envoyé & la collection du Muséum par cet
auteur. . . .” with reference to its receipt from Cope in 1867.
Figured in Vaillant, pl. 3, fig. 9.

—Etheostoma (Hololepis) fusiforme fusiforme (Girard). See Collette
(1962).

Hololepis fusiformis atraquae Hubbs and Cannon, 1935, Misc. Publ.
Mus. Zool. Univ. Michigan, no. 30, p. 68, pl. 1, fig. 11.
Holotype: UMMZ 107090 (<7, 31), Maryland, Prince Georges-
Charles Co. line, Mattawoman Cr.; C. L. Hubbs, G. 5. Myers, and
E. D. Reid; May 21, 1933.
219-948—66——4

50 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Paratypes: UMMZ 107089 (15, 27-32), USNM 117547 (2, 29-30),
and USNM 92946 (4, 32-34), same data as holotype.

= Etheostoma (Hololepis) fusiforme fusiforme (Girard). See Collette
(1962).

Hololepis fusiformis insulae Hubbs and Cannon, 1935, Mise. Publ.
Mus. Zool. Univ. Michigan, no. 30, p. 83, pl. 1, fig. 15.

Holotype: MCZ 28274 (1, 24), Massachusetts, Nantucket Is.,
Gibbs Pond; H. Allen; ‘“‘rec’d. Aug. 10, 1893.”

Paratypes: UMMZ 86601 (2, 22-25) and MCZ 42078 (8, 16-26),
same data as holotype. The latter was originally MCZ 33547, but
because that number has also been used for a Girardinus, the paratypes
have been recataloged.

=Etheostoma (Hololepis) fusiforme fusiforme (Girard). See Collette
(1962).

Hololepis fusiformis metae-gadi Hubbs and Cannon, 1935, Mise.
Publ. Mus. Zool. Univ. Michigan, no. 30, p. 81, pl. 1, fig. 14.

Holotype: USNM 77860 (<7, 32), Massachusetts, Barnstable Co.,
Osterville, Tempies Pond; V. Edwards; Nov. 12, 1902.

Paratypes: USNM 94683 (86, 27-43) and UMMZ 107093 (2,
31-38), same data as holotype.

=Etheostoma (Hololepis) fusiforme fusiforme (Girard). See Col-
lette (1962).

Hololepis saludae Hubbs and Cannon, 1935, Mise. Publ. Mus. Zool.
Univ. Michigan, no. 30, p. 50, pl. 1, fig. 6.

Holotype: UMMZ 107079 (1, 21), South Carolina, Saluda Co.,
Richland Cr., trib. of Lake Murray, 10 mi. SE. of Saluda; E. M.
Burton; June 21, 1933.

Paratypes: UMMZ 107078 (2, 20-22), CM 33.149.1 (4, 16-22),
and USNM 94685 (1, 21), same data as holotype. FMNH 38331
(1, 18), out of CM 33.149.1. UMMZ 107077 (4, 17-21) and CM
33.139.1 (4, 14-16), South Carolina, Saluda Co., Moore’s Cr., trib.
of Lake Murray, 6 mi. SE. of Saluda; Burton; June 21, 1933.

=LKtheostoma (Hololepis) saludae (Hubbs and Cannon). See
Collette (1962).

Hololepis serrifer Hubbs and Cannon, 1935, Mise. Publ. Mus. Zool.
Unive Michizan, no:°30;.p: 31) .pl.1,enes 2:
Holotype: UMMZ 107053 (, 53), North Carolina, Wake Co.,
Buffalo Cr. at Wendell; C. S. Brimley and Harris; Nov. 19, 1925.
Paratypes: The following are from North Carolina: USNM 93132
(9, 45) and USNM 93133 (27, 43), Wilmington; W. P. Seal; March
1900. USNM 86163 (9, 38), Wilmington; W. W. Welsh; Dec. 26,
1916. UMMZ 107056 (2, 33-36) and USNM 200333 (1, 34), Bertie

No. 3650 DARTERS—COLLETTE AND KNAPP il

Co., Windsor; C. S. Brimley and W. B. Mabee; Oct. 21, 1924.
UMMZ 107061 (1, 33), Martin Co., Williamston; Brimley; Oct. 23,
1924. UMMZ 107065 (<7, 40), Martin Co.; Jamesville; Brimley;
Oct. 23, 1924. UMMZ 107054 (2, 34-38) and USNM 200335 (2, 38),
Wake Co., Little R. at Wendell; Brimley and Mabee; Dec. 28,
1923. UMMZ 107064 (1, 33), same locality; Brimley and Harris;
Nov. 19, 1925. UMMZ 94658 (2, 34-37), same locality; Brimley
and Brady; Nov. 30, 1931. UMMZ 107060 (2, 37-43) and USNM
200330 (1, 38), Wake Co., Buffalo Cr. at Wendell; Brimley and Mabee;
Nov. 30, 1923. UMMZ 107062 (0, 37), same data; Mar. 30, 1925.
UMMZ 107063 (5, 34-39) and USNM 200331 (5, 36-48), Johnston
Co., Buffalo Cr., near Archer; Brimley and Mabee; Nov. 26, 1923.
UMMZ 107066 (1, 37), Harnett Co., Black R. at Dunn; Brimley and
Mabee; Nov. 19, 1923. UMMZ 107055 (8, 37-47) and USNM 200336
(4, 38-41), Johnston Co., Mingo Cr.; Brimley and Mabee; Nov. 19, 1923.
UMMZ 107067 (3, 33-38) and USNM 200337 (3, 30-35), same data,
Dec. 10,1924. UMMZ 107059 (<7, 39) and USNM 200334 (2, 36-39),
Harnett Co., Lillington; Brimley and Mabee; Dec. 11, 1923. UMMZ
107058 (1, 35), Harnett Co., Upper Little R. at Bunlevel; Brimley
and Mabee; Dec. 11, 1923. UMMZ 107057 (1, 32) and USNM
200332 (1, 35), Sampson Co., Clinton; Brimley and Mabee; Nov. 20,
1923. UMMZ 70705 (1, 26), Moore Co., Lakeview; R. E. Coker;
June 12, 1926. UMMZ 107073 (9, 44), same data; Noy. 11, 1934.
The following are from South Carolina: CM 34.215.1 (%, 43), Marl-
boro Co., 6 mi. N. of Bennettsville; E. M. Burton; July 25, 1934.
CM 33.260.1 (2, 32-34), Richland Co., Cedar Cr. on Leesville Rd.;
Burton; Aug. 31, 1933. CM 33.264.1 (1, 32), Richland Co., old dam,
near Wilson’s Mill; Burton; Aug. 31, 19383. CM 34.179.1 (2, 33) and
UMMZ 107075 (2, 32-33), Bamberg Co., Little Salkahatchie R., 3.5
mi. N. of Ehrhardt; Burton; July 11, 1934. CM 34.177.1 (2, 31-34)
and UMMZ 107074 (2, 26-33), Colleton Co., Little Salkahatchie R.,
near Bell’s Crossroads; Burton; July 11, 1934.

=Ktheostoma (Hololepis) serriferum (Hubbs and Cannon). See
Collette (1962).

Hololepis thermophilus Hubbs and Cannon, 1935, Misc. Publ. Mus.
Zool. Univ. Michigan, no. 30, p. 63, pl. 1, fig. 10.
Holotype: UMMZ 107072 (<7, 32), North Carolina, Harnett Co.,
millpond at Kipling; C. S. Brimley and W. B. Mabee; Dec. 11, 1923.
Paratypes: UMMZ 107071 (2, 30) and USNM 200239 (2, 28-30),
same data as holotype. UMMZ 107070 (<0, 31), North Carolina,
Wake Co., Buffalo Cr. at Wendell; Brimley and Mabee; Nov. 26,
1923. UMMZ 107069 (1, 29), same data; Nov. 30, 1923. UMMZ

52 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

107068 (1, 29), North Carolina, Johnston Co., Big Swamp at Kenly;
Brimley and Mabee; Nov. 26, 1923. USNM 52060 (8, 27-34) and
USNM 52071 (6, 27-29), North Carolina, Wilmington; W. P. Seal.
USNM 86165 (9, 26-30), North Carolina, ditch at Wilmington;
W. W. Welsh; Dec. 29, 1916. USNM 86160 (2, 22-24), North Caro-
lina, lake at Wilmington; ‘““Grampus”’; Jan. 3, 1917.

=F theostoma (Hololepis) fusiforme fusiforme (Girard). See Bailey
and Frey (1951) and Collette (1962).

Hololepis thermophilus oligoporus Bailey and Frey, 1951, Journ. Elisha
Mitchell Sci. Soc., vol. 67, no. 2, p. 193, pl. 6, figs. 9-18; pl. 7,
figs. 11-19; pl. 8, figs. 5-11.

Holotype: UMMZ 161969 (<7, 25), North Carolina, Bladen Co.,
Jones Lake; North Carolina lake survey; Sept. 10, 1947.

Paratypes: All from Bladen Co., N.C., collected by the North
Carolina lake survey in 1947. Jones Lake: UMMZ 161968 (44,
15-27), same data as holotype. UMMZ 161970 (2, 21-23), June 17.
UMMZ 161971 (8, 20-24), Aug. 26. UMMZ 161972 (88, 19-27),
Aug. 28. DU uncat. (15, 21-25), Aug. 28. Salters Lake: UMMZ
161974 (46, 19-25), Sept. 9. UMMZ 161973 (2, 21), June 27.
UMMZ 161975 (2, 18-20), July 4. Singletary Lake: UMMZ 161965
(2, 22-25), July 17. UMMZ 161966 (25, 17-27), Sept. 5. UMMZ
161967 (5, 22-25), Sept. 7.

=F theostoma (Hololepis) fusiforme fusiforme (Girard). See Collette
(1962).

Hololepis zonifer Hubbs and Cannon, 1935, Mise. Publ. Mus. Zool.
Univ. Michigan, no. 30, p. 47, pl. 1, fig. 5.

Holotype: UMMZ 88803 (9, 31), Alabama, Montgomery Co.,
pools of Catoma Cr., 5 mi. SW. of Montgomery; E. P. Creaser and
H.R. Becker; Sept. 18, 1929.

Paratype: UMMZ 88822 (9, 25), Alabama, Lowndes Co., Big
Swamp Cr., 25 mi. SW. of Montgomery; Creaser and Becker; Sept. 18,
1929.

=Etheostoma (Hololepis) zoniferum (Hubbs and Cannon). See
Collette (1962).

Hyostoma blennioperca Cope, 1868, Journ. Acad. Nat. Sci. Philadelphia,
sera 2, vol.6,)p. 215.

Lectotype: ANSP 14026 (<7, 85), Virginia; Holston R.; E. D. Cope;
October 1867. Fowler (1907, p. 522) designated number 14026 as
“type” but left it mixed with the other syntypes in the bottle.
However, he mentioned that ‘‘the type as here restricted is evidently
the largest in the collection.’”’ Therefore, the largest specimen is
herein confirmed as lectotype by Dr. Robert V. Miller. D XIV-13;

NO. 3550 DARTERS—COLLETTE AND KNAPP 53

A II, 8; LL 74; pectoral rays 15-15; cheeks and opercle, completely
covered with scales; snout blunt, not overhung.

Paralectotypes: ANSP 14027-35 (8, 47-60), same data as lectotype.
ANSP 14004 (1, 54), Virginia, Wythe Co., trib. of Kanawha R.;
Cope; 1867. ANSP 14006-08 (3, 51-61), Virginia, Wythe Co.,
Walker’s Cr., trib. of Kanawha R.; Cope; 1867. ANSP 14020-25
(6, 54-66), Virginia, Sinking Cr., trib. of Kanawha R.; Cope; 1867.

By selecting ANSP 14026 as lectotype, Fowler made the name
H. blennioperca Cope applicable to the Holston River population of
E. blennioides. H. blennioperca thereby becomes a synonym of EL. b.
newmanit, according to Miller (1964). However, the Kanawha River
specimens represent the upper Ohio River subspecies EF. 6. blennioides,
according to Miller (1964).

=Etheostoma (Etheostoma) blennioides newmanii (Agassiz). See
Miller (1964).

Hyostoma Newmanit Agassiz, 1854, Amer. Journ. Sci. and Arts, vol. 17
p. 305.

Lectotype: USNM 1865 (<7, 93), Alabama, near Huntsville;
Newman; herein selected by Dr. Robert V. Miller. D XIV-12;
A II, 7; pectoral rays 15-15; LL 71; cheeks, opercles, and belly
completely covered with scales.

Paralectotypes: MCZ 24602 (5, 68-106), orig. out of USNM 1865,
then MCZ 237. UMMZ 86764 (1, 106), out of MCZ 24602.

—=LKtheostoma (Etheostoma) blennioides newman (Agassiz). See
Miller (1964).

Hyostoma simoterum Cope, 1868, Journ. Acad. Nat. Sci. Philadelphia,
ser. 2, vol..6, p. 215, color pl. 24, fig. 5.

Syntypes: ANSP 14039-54 (16, 35-51), Virginia, Holston R.;
E. D. Cope; October 1867. ANSP 14036-8 (8, 57-62); ANSP 14499-
503 (5, 40-48); ANSP 5368-74 (7, 37-52); USNM 36604 (<7, 40);
MNHN 4856 (1, 53), all with same data. Fowler (1907, p. 524)
erroneously designated ANSP 14036 as lectotype and ANSP 14037-38
as paralectotypes of Poecilichthys zonalis Cope. These specimens
are syntypes of H. simoterum as is clear from Fowler’s figure 8 (see
P. zonalis). With reference to the MNHN specimen, Vaillant
(1873, p. 100) wrote “Cette espéce est représentée au Museum par un
exemplaire type dt 4 l’obligeance de M. Cope.”

=Etheostoma (Ulocentra) simoterum (Cope).

Ioa vigil Hay, 1882, Bull. U.S. Fish. Comm., vol. 2, p. 59.

Holotype: USNM 32201 (1, 23), Mississippi, Pearl R. at Jackson,
O. P. Hay; July-August 1881.

=Percina sp. Perhaps P. (Imostoma) uranidea (Jordan and
Gilbert). Not a species of Ammocrypta as has been presumed, but

54. PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

a species of Percina with a well-developed ctenoid caducous scale
between the bases of the pelvic fins.

Microperca proeliaris Hay, 1881, Proc. U.S. Nat. Mus., vol. 3, p. 496.
Holotype: USNM 27418 (1, 26), Mississippi, Alcorn Co., trib. of
Tuscumbia R. at Corinth; O. P. Hay; March-April 1880; in poor
condition.
=Etheostoma (Microperca) proeliare (Hay).

Microperca punctulata Putnam, 1863, Bull. Mus. Comp. Zool., vol. 1,
p. 4.

Syntypes: USNM 1377 (<7, 22; 9, 24), Illinois, Calumet R., near
Chicago; R. Kennicott. MCZ 24566 (2 9, 23-24); orig. out of
USNM 1377, then MCZ 95. USNM 1283 (20, 23-30), Wisconsin,
Oconomowac R., Lac la Belle; S. F. Baird; July 1853. MCZ 24570
(9, 24-31) and UMMZ 86457 (8, 24-31), orig. out of USNM 1283,
then MCZ 93. USNM 1288 (28, 15-30), Michigan, Port Huron;
Baird; August 1853. MCZ 24690 (11, 16-28) and UMMZ 86316
(3, 19-28), orig. out of USNM 1288, then MCZ 94; in poor condition.
MCZ 24582 (1, 19), Michigan, Detroit R.; Baird; August, 1853;
orig. USNM 1294, then MCZ 96; in poor condition. Also USNM
1276 (1, ?), Wisconsin, tribs. of Fox R., Vienna; Baird; August 1853;
which we have been unable to locate.

=F theostoma (Microperca) microperca Jordan and Gilbert because
punctulata is preoccupied in Etheostoma by Poecilichthys punctulatus
Agassiz.

Nanostoma elegans Hay, 1881, Proc. U.S. Nat. Mus., vol. 3, p. 493.

Lectotype: USNM 27445 (<7, 43), Mississippi, Clarke Co., shallow
rocky branch of Chickasawha R. at Enterprise; O. P. Hay; March-
April 1880; listed as type by Jordan and Evermann (1896, p. 1075);
selection herein confirmed. D X-12; A IJ, 8; LL 42; branchiostegals
5-5; INF 7; POM 9; cheeks, opercles, and nape scaly; breast naked.

Paralectotype: SU 627 (1, 53), same data as lectotype.

=Etheostoma (Etheostoma) zonale lynceum (Hay) because elegans is
preoccupied in Etheostoma by Boleichthys elegans Girard.

Nanostoma vinctipes Jordan, 1880, Proc. U.S. Nat. Mus., vol. 2,
p. 236.

Lectotype: USNM 23454 (1, 47), Illinois, trib. of Illinois R. at
Naperville; E. R. Copeland; listed as type by Jordan and Evermann
(1896, p. 1075); selection herein confirmed. D XI-10; A II, 6;
LL 46; branchiostegals 6-6; INF 7; POM 10; cheeks, opercles,
breast, and nape scaly.

Paralectotypes: The whereabouts of the other four specimens from
the type series is unknown.

NO. 3550 DARTERS—COLLETTE AND KNAPP 55

=Etheostoma (Etheostoma) zonale vinctipes (Jordan), a valid sub-
species according to Dr. Chu-fa Tsai (pers. comm.).

Nothonotus inscriptus Jordan and Brayton, 1878, U.S. Nat. Mus.,
Bull. 12(A), p. 34.

Lectotype: USNM 31145 (9, 43), Georgia, Hall Co., Oconee R. at
Sulphur Springs; D. S. Jordan and A. W. Brayton; summer, 1877;
herein selected. D X-10; A II,8; LL 49; prevomerine teeth present;
6 dorsal blotches; cheeks, opercles, prepectoral area, and breast
naked; nape completely scaled.

Paralectotype?: USNM 197439 (co, 42), apparently same data
as lectotype.

We select USNM 31145 as lectotype for the following reasons.
The original description was based on two syntypes, male and female.
USNM 31145 has the proper data and is labeled type, so it is clearly
part of the type series. The male that we believe is the other syntype
was discovered in the USNM collection labeled USNM 31132, Etheo-
stoma, Etowah R., Georgia, Jordan and Brayton. WN. inseriptum was
reported only from the type locality in the original description, so it
seems probable that this locality is in error. Under the species
column in the USNM catalog we found the following entry for
USNM 31132: ‘Notropis stilbius Semotilus, etc. (Etheostoma),” and
for the locality ‘Etowah R., Rome, Ga. (chiefly).”’ It is apparent
that the data for the specimen of inscriptus under USNM 31132 are
totally confused. Therefore, in order to avoid further confusion should
other specimens be found bearing this number, we recataloged this
specimen as USNM 197439 and designate USNM 31145 as lectotype,
because it is the specimen that we are certain is part of the type
material.

=Ktheostoma (Etheostoma) inscriptum (Jordan and Brayton).

Nothonotus thalassinus Jordan and Brayton, 1878, U.S. Nat. Mus.
Bulle Algca) %p. 13.

Lectotype: USNM 31122 (<7, 40), South Carolina, Saluda R., near
Greenville; D. S. Jordan and A. W. Brayton; summer, 1877; listed
as type by Jordan and Evermann (1896, p. 1072); selection herein
confirmed. D XIJ-12; A II,8; pectoral rays 15-14; LL 44; INF 8;
POM 10; prevomerine teeth present; cheeks, opercles, prepectoral
area, and breast naked; anterior portion of belly naked; nape scaled;
7 dorsal blotches.

Paralectotypes: SU 987 (9, 46; 5 o, 37-49), South Carolina,
Saluda R. at Farr’s Mills, W. of Greenville; Jordan and Brayton;
summer, 1877. MCZ 24397 (o, 44), South Carolina, Saluda R.
BMNH 1880.1.21.68 (c’, 52), South Carolina, Saluda R., Jordan.

=Etheostoma (Etheostoma) thalassinum (Jordan and Brayton).

56 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

Oligocephalus grahami Girard, 1859, Proc. Acad. Nat. Sci. Philadel-
phia, vol. 11, p. 102.
Holotype: MCZ 24576 (1, 33), Texas, Devil’s R.; J. H. Clark
under Col. J. D. Graham; orig. USNM 1182, then MCZ 85.
=LKHtheostoma (Oligocephalus) grahami (Girard).

Oligocephalus humeralis Girard, 1859, Proc. Acad. Nat. Sci. Phila-
delphia, vol. 11, p. 67.

Lectotype: USNM 1203 (<’, 42), Virginia, James R.; S. F. Baird;
1850; herein selected. D VII-13; A II,8: LL 36+11=47; INF 442;
POM 10; supratemporal canal interrupted; cheeks, opercles, nape,
and breast naked; about 11 vertical bars on body; tips of dorsal spines
enlarged and slightly forked; branchiostegal membranes moderately
joined.

Paralectotypes: USNM 198001 (45, 23-49), removed from USNM
1203. MCZ 24634 (10, 25-54), orig. out of USNM 1203, then MCZ 16.
MCZ 35975 (2, 22-33), orig. out of USNM 1203.

=Etheostoma (Catonotus) flabellare flabellare Rafinesque.

Oligocephalus leonensis Girard, 1859, Proc. Acad. Nat. Sci. Philadelphia,
Vol 4as sp: 102.

Syntypes: MCZ 24580 (2 9, 34-37), Texas, Rio Leona; J. H. Clark;
badly dried; orig. USNM 748, then MCZ 83.

=?Htheostoma (Oligocephalus) lepidum (Baird and Girard). Al-
though most characters suggest E’. lepidum, there are well-developed
ctenoid scales on the opercle, a character of L. grahami which is found
in tributaries of the Rio Grande. The body appears to be more elon-
gate than in either L. lepidum or E. grahami.

Oligocephalus pulchellus Girard, 1859, Proc. Acad. Nat. Sci. Philadel-
phia, vol. 11, p. 102.

Holotype: MCZ 32950 (1, 28), Oklahoma, trib. of Gypsum Cr., trib.
of Canadian R.; Lt. A. W. Whipple and party, exploration of railroad
route near 35th parallel; orig. USNM 1334.

=Ltheostoma (Oligocephalus) spectabile pulchellum (Girard). See
Hubbs and Ortenburger (1929).

Perca minima Haldeman, 1842, Journ. Acad. Nat. Sci. Philadelphia,
vol. 8, p. 330.

Holotype: ANSP 13928 (1, about 47), Pennsylvania, Susquehanna
R.; S. S. Haldeman; a skin preserved in alcohol and reported to be in
poor condition by Fowler (1907).

=LHtheostoma (Boleosoma) olmstedi olmstedi Storer. See Storer
(1846, p. 271).

Perca (Percina) nebulosa Haldeman, 1842, Journ. Acad. Nat. Sci.
Philadelphia, vol. 8, p. 330.

NO. 3550 DARTERS—COLLETTE AND KNAPP 57

Holotype: ANSP 22652 (7, 110), Pennsylvania, Susquehanna R.,
S. S. Haldeman; figured by Fowler (1945, fig. 202).
=Percina (Percina) caprodes semifasciata (DeKay).

Percina bimaculata Haldeman, 1844, Proc. Boston Soc. Nat. Hist., vol.
rp. LT.

Types: USNM 1405, ‘‘Percina, Susquehanna River, S. S. Haldeman,
Percina dried.—ocrig. of Haldeman” may be the type of P. bimaculata,
but we have been unable to find this specimen.

=Percina (Percina) caprodes semifasciata (DeKay).

Percina caprodes burtoni Fowler, 1945, Acad. Nat. Sci. Philadelphia
Monogr., no. 7, p. 134, figs. 218-219.
Holotype: ANSP 70701 (1, 134), North Carolina, Buncombe Co.,
Swannanoa R., near Oteen; KE. M. Burton; Aug. 21, 1934.
Paratype: ANSP 13707 (1, 66), Virginia, Holston R.; E. D. Cope.
=Percina (Percina) caprodes burtoni (Fowler).

Percina (Hadropterus) lenticula Richards and Knapp, 1964, Copeia,
no. 4, p. 695, fig. 1 A-C.

Holotype: CU 43592 (7, 96), Alabama, Bibb Co., Cahaba R. on
Ala. 27, 8.5 mi. N. of Centerville; R. D. and J. S. Suttkus; May 9,
1959.

Paratypes: CU 43593 (9, 95), Alabama, Bibb Co., Cahaba R. on
Ala. 5, 2.2 mi. N. of Centerville; Suttkus and party; July 8, 1958. TU
15291 (7 9, 77-92) and CU 43594 (07, 86), same locality as CU 43593;
Suttkus, R. J. and H. V. Miller, and J. de Abate; Mar. 17, 1957. UG
209 (3 o', 19, 48-63), CU 43435 (1 of, 1 9, 55-58), USNM 197491
(oi, 64), and UMMZ 180074 (<7, 60), Georgia, Cherokee Co., Etowah
R. at the mouth of Canton Town Cr.; C. M. Tarzwell; Oct. 8, 1950.

=Percina (Hadropterus) lenticula Richards and Knapp.

Percina manitou Jordan, 1877, Proc. Acad. Nat. Sci. Philadelphia, vol.
29, p. 53.

Lectotype: USNM 23458 (9, 68), Indiana, Fulton Co., Lake
Manitou; D. S. Jordan; labeled ‘‘drawn”’; herein selected. D XV-14;
A JI,11; LL 86; breast and nape naked; cheeks and opercles scaled;
one enlarged ctenoid scale on midline of anterior part of belly.

Paralectotypes: The original description was based on three speci-
mens in the Indiana State collections. One of these possessed and the
other two lacked “enlarged, mucronate plates.”” We have not been
able to locate the other two syntypes (one male and one female) and
therefore designate the syntype we have found as lectotype.

=Percina (Percina) caprodes semifasciata (DeKay).

Percina nigrofasciata raneyi Crawford, 1956, Tulane Stud. Zool., vol.
4, no. 1, p. 49, fig. 10.

5S PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Holotype: CU 23344 (0, 79), South Carolina, Abbeville Co.,
Calhoun Cr., trib. of Savannah R., 7.6 mi. E. of Calhoun Falls on rt.
72; E. C. Raney, R. W. Crawford, R. H. Backus, C. R. Robins, J. N.
Layne, and R. L. Wigley; Mar. 27, 1951.

Paratypes: CU 19599 (87, 43-78) and USNM 188896 (27, 75-78;
29, 61-63), same data as holotype.

—Percina (Hadropterus) nigrofasciata raneyi Crawford.

Percina (Alvordius) notogramma montuosa Hogarth and Woolcott,
1966, Chesapeake Sci., vol. 7, no. 2, p. 101, fig. 2.

Holotype: USNM 199556 (<7, 51), Virginia, Campbell Co., Beaver
Cr. 4 mi. E. of Lynchburg on US 460; W. T. Hogarth and E. Collins;
Oct. 20, 1964.

Paratypes (all from Virginia): UR 2337 (3, 40-57), Albemarle Co.,
Hog Cr., at jet. with Rockfish R. at jct. of Co. rt. 602 and 722;
Hogarth, C. Beirne, L. Kiss; Apr. 15, 1965. UMMZ 144697 (3,
47-49), Rockbridge Co., North R.; L. C. Pettit; May 17, 1941.
UMMZ 175103 (5, 52-62), Allezhany Co., Potts Cr. above ject. with
Jackson R.; C. R. Gilbert and D. G. Myer; Sept. 13, 1957. UR 2294
(6, 34-59), Albemarle Co., ject. with Rock Castle Cr. and Totier Cr.
1.2 mi. S. Scottsville on Co. rt. 726; W. S. Woolcott and Hogarth;
Apr. 1, 1965. VPI 1006 (2, 51-64), Craig Co., Craig Cr. 6.8 mi. from
New Castle on rt. 311; R. D. Ross, Wakeman, Cook; Aug. 19, 1958.
CU 47322 (1, 48), Craig Co., Craig Cr. 0.25 river mi. downstream
from Pinetop; R. E. Jenkins, J. E. Carico, R. L. Miles; June 6, 1963.
VPI 1014 (1, 68), Rockbridge Co., South R. 4.9 air mi. E. of Lexington;
Ross, Wakeman, Cook, Carroll, Handley; Aug. 21, 1958. VPI 981
(1, 52), Bath Co., Jackson R. 1.5 mi. N. of Greenwood; Ross, Wake-
man, Cook; July 15, 1958. CU 47321 (2, 56-63), Bath Co., Jackson
R., Hidden Valley Farm 4.3 mi. N. Mitchell Town; Ross, Jenkins;
July 20, 1963. WPI 954 (2, 64-67), Bath Co., Cow Pasture R..,
0.4 mi. NE. Nimrod Hall; Ross, Wakeman, Cook; July 3, 1958. UR
2350 (1, 52), Nelson Co., Owens Cr. 2.3 mi. SK. Buffalo Springs on
Co. rt- 626; Hogarth, Beirne, Kiss; Apry 15, 1965: (UK 1797 ie.
53-62), Craig Co., John’s Cr., Co. rt. 632, 2 mi. NE. ject. Co. rt. 632
and 658; Woolcott and Ichthyology Class; June 27, 1963. UR 2321
(12, 33-58), Amherst Co., Partridge Cr., Co. rt. 622 at jct. Co. rt.
624 near Stapleton; Hogarth, Beirne, Kiss; Apr. 15, 1965. VPI 963
(1, 47), Bath Co., Jackson Cr. 8.5 mi. S. Bacova on Co. rt. 603; Ross,
Wakeman, Cook; July 2, 1958. VPI 1005 (1, 55), Craig Co., Craig
Cr., 3 mi. SE. New Castle; Ross, Wakeman, Cook; Aug. 19, 1958.
USNM 199557 (3, 44-62), Campbell Co., Beaver Cr. 4 mi. E. Lynch-
burg on U.S. 460; Hogarth and E. J. Collins; Oct. 20, 1964. UMMZ
175121 (1, 51), Alleghany Co., Potts Cr.; Gilbert and Myer; Sept. 13,

NO. 3550 DARTERS—COLLETTE AND KNAPP 59

1957. UMMZ 95194 (1, 41), Alleghany Co., Dunlap Cr.; C. L. and
L. C. Hubbs; Sept. 10, 1928.

=Percina (Alvordius) notogramma montuosa Hogarth and Woolcott.

Pileoma carbonaria Baird and Girard, 1853, Proc. Acad. Nat. Sci.
Philadelphia, vol. 6, p. 387.

Lectotype: USNM 740 (<7, 85), Texas, Rio Salado, 4 mi. from
San Antonio; J. H. Clark; herein selected. D XV-15; A II, 11;
LL 90; breast naked; cheeks, opercles, and nape completely covered
with scales.

Paralectotypes: USNM 197999 (2 o&, 75-87; 9, 73), removed
from USNM 740. MCZ 24599 (2 &, 88-108) and UMMZ 86594
(os, 86), orig. out of USNM 740, then MCZ 269.

Jordan and Evermann (1896, p. 1027) also listed USNM 742 as
“type.”’ This specimen was taken by Kennerly in San Pedro Creek;
there is no mention of either this locality or collector in the original
description, so the specimen is not part of the type series.

= Percina (Percina) caprodes carbonaria (Baird and Girard).

Pileoma cymatogramma Abbott, 1860, Proc. Acad. Nat. Sci. Phila-
delphia, vol. 12, p. 327.
Holotype: ANSP 14009 (9, 45), no locality.
= Etheostoma (Etheostoma) blennioides blennioides Rafinesque.
See Miller (1964).

Pileoma semifasciatum DeKay, 1842, Nat. Hist. New York Zool.,
vole Iho pte4, p. 16, pl.. 50, fir. 162.

Syntypes: ?New York, Westport on Lake Champlain; J. E.
DeKay; figure based on material then in the N.Y. State collection.
Dr. E. M. Reilly, Curator of Zoology, has formed us that the types
of this species are no longer present in the New York State Museum.

Topotypes are available: MCZ 24613 (5, 41-77), S. F. Baird.

= Percina (Percina) caprodes semifasciata (DeKay).

Pileoma zebra Agassiz, 1850, Lake Superior, p. 308, pl. 4, fig. 4.
Syntypes: MCZ 24636 (7, 51; 2, 60), Ontario, Lake Superior,
“The Pic” at the mouth of the Pic R.; L. Agassiz; about July 13,
1848; orig. USNM 1175, then MCZ 257. UMMZ 86592 (1, 53) out
of MCZ 24636.
= Percina (Percina) caprodes semifasciata (DeKay). See Hubbs
and Brown (1929, p. 47).

Plesioperca anceps Vaillant, 1873, Nouv. Arch. Mus. Hist. Nat.
Parse vol) 9. p. 37, pli 1, fig. 3.
Holotype: MNHN 2778 (<’, 86), locality unknown; donated by
Agassiz; orig. MCZ 851.
= Percina (Hadropterus) nigrofasciata nigrofasciata (Agassiz).

60 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

See Jordan (1880, p. 225) and Crawford (1956).

Pleurolepis asprellus Jordan, 1878, Bull. Illinois State Lab. Nat.
ists, “wol ie mnon2.9 pao:

Holotype: INHS uncat. (1, 90), Illinois, Hancock Co., small
rocky trib. of Mississippi R.; C. K. Worthen.

Paratypes: ?(1, ?), same data as holotype. ? Illinois, Effingham
Co., Little Wabash R.; S. A. Forbes; July 1877. We have been
unable to find any paratypes of this species. The types were orig-
inally in the collection of the Illinois State Laboratory of Natural
History. Smith and Bridges (1960, p. 254) did not list any type
material of this species in the collections of the Illinois Natural
History Survey which is the State Laboratory’s successor, but
Dr. Smith recently found the holotype for us.

= Ammocrypta (Crystallaria) asprella (Jordan).

Pleurolepis pellucidus Agassiz in Putnam, 1863, Bull. Mus. Comp.
Zool. vol 18 p: 5.

Lectotype: USNM 1311 (1, 50), Ohio, Black R., below falls at
Elyria; S. F. Baird; August 1853; orig. no. 17. USNM 1311, then
containing 21 specimens, was listed as type by Jordan and Evermann
(1896, p. 1063); Linder (1959, p. 182) selected 1 specimen as lectotype.

Paralectotypes: USNM 164165 (20, 40-48) removed from USNM
1311 by Linder (1959). MCZ 24616 (9, 30-47) and UMMZ 86489
(3, 47-50) orig. out of USNM 1311, then MCZ 270. Although not
mentioned by Jordan and Evermann or Linder, these collections are
obviously part of the original material and come from the type locality
and so must be considered parlectotypes.

There is no mention of any localities in the original description of
P. pellucidus, but the following specimens were probably part of the
original material USNM 1289 (1, 49), Michigan, Port Huron;
Baird; August 1853. USNM 1295 (1, 39), Mich., Detroit R.; Baird;
August 1853. MCZ 24626 (4, 44-49) and UMMZ 86479 (1, 46),
Ohio, Rockport; Kirtland.

=Ammocrypta (Ammocrypta) pellucida (Agassiz) not A. pellucida
(Baird) as frequently given because the Baird name was listed as a
synonym in Putnam and therefore has no status.

Poecilichthys artesiae Hay, 1881, Proc. U.S. Nat. Mus., vol. 3, p. 494.
Holotype: USNM 27434 (0, 63), Mississippi, Lowndes Co.,
Catawba Cr., trib. of Tombigbee R. at Artesia; O. P. Hay; March—
April 1880.
=Ktheostoma (Oligocephalus) whipplii artesiae (Hay). See Hubbs
and Black (1941).

Poecilichthys asprigenis Forbes in Jordan, 1878, Bull. Illinois State
Lab. Nat. Hist., vol. 1 no. 2, p. 41.

NO. 3550 DARTERS—COLLETTE AND KNAPP 61

Syntypes: SU 2201 (2, 38-39), Illinois, small creek near Pekin.
The original description was based on 14 specimens from the type
locality, but we have been unable to locate the other 12. Smith and
Bridges (1960) did not report any type specimens of this species in the
collections of the Illinois Natural History Survey which inherited the
material of the Illinois State Laboratory.

MCZ 25051 (1, 45), Illinois, Peoria, 1880, is labeled as cotype but
was collected after the original description was published and came
from Peoria rather than from Pekin.

=Etheostoma (Oligocephalus) asprigene (Forbes).

Poecilichthys beani Jordan, 1884, Proc. U.S. Nat. Mus., vol. 7, p. 479.
Holotype: USNM 35754 (9, 36), Missouri, Lafayette Co., Tabo
Cr., trib. of Missouri R., near Lexington.
=LEtheostoma (Boleosoma) nigrum nigrum Rafinesque. Jordan
(1885, p. 548) admitted P. beani was a mutilated individual of Boleo-
soma maculatum (=E. nigrum).

Poecilichthys borealis Jordan, 1884, Proc. U.S. Nat. Mus., vol. 7,
p. 477.

Lectotype: USNM 35747 (<7, 41), Canada, Montreal; T. J. Doran;
herein selected. D IX-11; A II,7; LL 20+34=54; INF 8; POM 10;
supratemporal canal widely interrupted; cheeks, opercles, nape, and
belly covered with scales; few scattered scales on breast and pre-
pectoral region.

Paralectotypes: USNM 197993 (8, 35-48), removed from USNM
35747. The fifth syntype mentioned in the original description is in
SU 5447 but unfortunately became mixed with two specimens from
Gravenhurst, Canada (SU 6261) during the earthquake.

=F theostoma (Oligocephalus) exile (Girard). See Hubbs (1926,
p. 67) and Richardson (1938).

Poecilichthys butlerianus Hay, 1883, Bull. U.S. Fish Comm., vol. 2,
Deol:
Holotype: USNM 32224 (1, 43), Mississippi, Yazoo Co., Big Black
R., near Vaughn’s Station; O. P. Hay; Aug. 20, 1881.
=F theostoma (Hololepis) gracile (Girard). See Hubbs and Cannon
(1935) and Collette (1962).

Poecilichthys camurus Cope, 1870, Proc. Amer. Philos. Soc., vol. 11,
p. 265.

Lectotype: ANSP 13741 (1, 55), Tennesee, headwaters of Cumber-
land R.; herein selected by Dr. Edward C. Raney. D X-13; A IL9;
LL 58.

Paralectotypes: ANSP 13742-44 (3, 49-56), same data as lectotype.

62 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

The types were found mixed in with the types of Poecilichthys sangui-
fluus Cope by Dr. Edward C. Raney.
=Etheostoma (Nothonotus) camurum (Cope). See Bailey (1959).

Poecilichthys coosae Fowler, 1945, Acad. Nat. Sci. Philadelphia,
Monogr., no. 7, p. 356, figs. 226-228.

Holotype: ANSP 71189 (1, 38), Alabama, Cherokee Co., small cr.,
% mi. NE. of Chesterfield, Coosa R. drainage; G. A. Coventry and
C. B. Peterson; July 7, 1942.

Paratypes: All collected by Coventry and Peterson in Cherokee
Co., Ala. ANSP 71190-91 (2, 34-35), same data as holotype. ANSP
71192-95 (4, 32-50), Cowan’s Cr., Jordan’s store; July 3, 1942.
ANSP 71196 (9, 39), cave on Lookout Mt., “Split Rock” at Little
R. Gulch; July 1, 1942.

= LHtheostoma (Ulocentra) coosae (Fowler).

Poecilichthys euzonus erizonus Hubbs and Black, 1940, Occ. Pap.
Mus. Zool. Univ. Michigan, no. 416, p. 17, pl. 1, fig. 2.

Holotype: UMMZ 124597 (7, 66), Missouri, Ripley Co., Current
R. at “The Nook,” T. 23 N., R. 2 E., sec. 9; A. H. Denney; Aug. 10,
1938.

Paratypes: UMMZ 124598 (5, 44-57), same data as holotype.
UMMZ 124595 (8, 60-70), Missouri, Carter Co., Carter County
Hunting and Fishing Club, T. 26 N., R. 1 E., sec. 11 and 12; Denney;
July 25, 1938. UMMZ 124596 (2, 36-62), Missouri, Ripley Co.,
Doniphan Boat Landing, T. 23 N., R. 2 E., sec. 27; Denney; July 26,
1938. UMMZ 124599 (2, 58), Missouri, Carter Co., Current R.,
just above mouth of Pike Cr., T. 27 N., R. 1 W., sec. 23; Denney;
Aug. 18, 1938.

=Ktheostoma (Etheostoma) euzonum erizonum (Hubbs and Black.)

Poecilichthys euzonus euzonus Hubbs and Black, 1940, Occ. Pap. Mus.
Zool. Univ. Michigan, no. 416, p. 23, pl. 1, fig. 3.

Holotype: UMMZ 123548 (<7, 61), Arkansas, Searcy Co., Buffalo
R., 4 mi. SE. of St. Joe; J. D. and R. Y. Black; July 8, 1938.

Paratypes: UMMZ 124594 (4, 28-51), and USNM 117587 (1, 35),
same data as holotype. UMMZ 123391 (2, 42-43), Arkansas,
Madison Co., King’s R., 3 mi. E. of Alabam at Denney Cave; J. D.
and R. Y. Black; June 30, 1938. UMMZ 123504 (1, 45), Arkansas,
Carroll Co., White R. near Busch; J. D. and R. Y. Black; July 8,
1938.

=Etheostoma (Etheostoma) euzonum euzonum (Hubbs and Black).

Poecilichthys gutselli Hildebrand, 1932, Journ. Elisha Mitchell Sci.
Soc., vol. 48, no. 1, p. 78, pl. 3.

NO. 3550 DARTERS—COLLETTE AND KNAPP 63

Holotype: USNM 92402 (<, 71), North Carolina, Swain Co.,
Tuckaseegee R. at Ela; J. S. Gutsell; Aug. 26, 1930.

Paratypes: USNM 197411 (9, 75), same data as holotype. USNM
92580 (2, 37), North Carolina, Jackson Co., Tuckaseegee R., below
Cullowhee; Gutsell; Aug. 29, 1930.

=Etheostoma (Etheostoma) blennioides gutselli (Hildebrand). See
Miller (1964).

Poecilichthys hopkinsi Fowler, 1945, Acad. Nat. Sci. Philadelphia
Monogr., no. 7, p. 249, fig. 206.

Holotype: ANSP 71548 (9, 44), Georgia, Wilcox Co., Osewitchee
Springs, 14 mi. N. of Fitzgerald; M. N. Hopkins, Jr.; Aug. 15, 1942.

Paratypes: ANSP 71567-69 (3, 25-45), same locality and collector
as holotype; June 29, 1944.

=Etheostoma (Oligocephalus) hopkinsi hopkinsi (Fowler). See
Bailey and Richards (1963).

Poecilichthys jessiae Jordan and Brayton in Jordan, 1878, Manual
Vertebrates, 2nd ed., p. 227, and im Jordan and Brayton, 1878,
U.S. Nat. Mus. Bull. 12(A), p. 59.

Syntypes: ? “sev. spec. each about 2’’ long,” Georgia, Chickamauga
R. at Ringgold; D. S. Jordan and A. W. Brayton; summer, 1877.
We have been unable to find any of the type material of this species.
Figure 4 was found in the files of the Division of Fishes labeled as
“Poecilichthys jessiae, Chickamauga River, Ga.’? We feel that it is
probably based on one of the types and so reproduce it here.

=Ktheostoma (Boleosoma) jessiae (Jordan and Brayton).

Poecilichthys kanawhae Raney, 1941, Occ. Pap. Mus. Zool. Univ.
Michigan, no. 434, pl. 1.

Holotype: UMMZ 131837 (&, 72), North Carolina, Ashe Co.,
N. Fork of New R. on rt. 16 at Crumpler; E. C. Raney, E.A. Lachner,
and L. J. Kezer; Apr. 1, 1940.

Paratypes: UMMZ 131838 (7, 39-60) and CU 7663 (13, 39-57),
same data as holotype. CU 7678 (207, 48-53), North Carolina,
Ashe Co., N. Fork of New R., 1 mi. NNE. of Warrensville; Raney,
Lachner, Kezer; Apr. 1, 1940. CU 7830 (8, 42-69) and UMMZ
131834 (2, 35), North Carolina, Ashe Co., N. Fork of New R., 1 mi.
NNE. of Creston; Raney, Lachner, Kezer; Apr. 2, 1940. CU 7688
(5, 44-59) and UMMZ 131836 (2, 50-56), North Carolina, Ashe Co.,
S. Fork of New R.on rt. 16 at Index; Raney, Lachner, Kezer; Apr. 2,
1940. CU 7693 (8, 40-44) and UMMZ 131835 (2, 40-42), North
Carolina, Ashe Co., S. Fork of New R., 1 mi. SW. of Fleetwood;
Raney, Lachner, Kezer; Apr. 2, 1940. UMMZ 95371 (2, 30-44),
Virginia, Carroll Co., Big Reed Island Cr.; C. L. Hubbs and E. P.
Creaser; May 17, 1931. USNM 107679 (8, 46-59), Virginia, Carroll

64 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Co., Crooked Cr., 4 mi. E. of Galax; L. P. Schultz and E. D. Reid;
July 13, 1938.
=Etheostoma (Etheostoma) kanawhae (Raney).

Poecilichthys mesaeus Cope, 1864, Proc. Acad. Nat. Sci. Philadelphia,
vol.16, p.232:

Lectotype: ANSP 13943 (9, 49), Nebraska, Platte R., near Ft.
Kearney; W. Hammond; selected and illustrated by Fowler (1907,
p. 524, fig. 7).

Paralectotypes: ANSP 13944-46 (3), same data as lectotype.
Fowler wrote “Type, No. 13,943, A.N.S.P.” and did not mention
these three specimens.

=Ktheostoma (Boleosoma) nigrum nigrum Rafinesque.

Poecilichthys osburni Hubbs and Trautman, 1932, Ohio Journ. Sci.,
vol, 32, no. 1, p: 34, fig. 2:
Holotype: UMMZ 92409 (<7, 75), West Virginia, Pocohontas Co.,
Stony Cr., trib. of Greenbrier R.; J. Addair; June 2, 1931.

Ficure 4.—Poecilichthys jessiae Jordan and Brayton, Chickamauga River, Ga. (pre-
viously unpublished drawing by Ernest Copeland that probably represents one of the
lost syntypes).

Paratypes: UMMZ 92410 (5, 58-85), and OSM 12946 (4, 56-81),
same data as holotype. USNM 39542 (9, 56), Virginia, South Fork
Reed’s Cr.; M. McDonald. UMMZ 95370 (7, 49-57), and USNM
117588 (9, 64), Virginia, Wythe Co., Reed Cr., trib. of New R., 2-3
mi. below Max Meadows; C. L. Hubbs and E. P. Creaser; May 17,
1931.

=LKHtheostoma (Etheostoma) osburni (Hubbs and Trautman). See
Raney (1941).

Poecilichthys palustris Gilbert, 1884, Proc. U.S. Nat. Mus., vol.
7, p. 209.

NO. 3550 DARTERS—COLLETTE AND KNAPP 65

Lectotype: USNM 34983 (<7, 30), Indiana, Greene Co., Switz City
Swamp; C. H. Gilbert; August 1883; selected by Hubbs and Cannon
(1935, p. 44).

Paralectotype: Gilbert mentioned a second specimen in his original
description which has apparently been lost (Hubbs and Cannon,
1935, p. 44).

=F theostoma (Hololepis) gracile (Girard). See Hubbs and Cannon
(1935) and Collette (1962).

Poecilichthys punctulatus Agassiz, 1854, Amer. Journ. Sci. and Arts,
vol. 17, p. 304.

Lectotype: MCZ 39 (1, 52), Missouri, Osage R.; G. Stolley;
herein selected. D XIJ-14; A II, 8;LL59+13=72; INF 8; POM 10;
supratemporal canal widely incomplete; cheeks and opercles naked;
color pattern completely faded.

Paralectotype: MCZ 43091 (1, 48), removed from MCZ 39.

=Etheostoma (Oligocephalus) punctulatum (Agassiz).

Poecilichthys quiescens Jordan, 1884, Proc. U.S. Nat. Mus., vol. 7,
p. 478.

Holotype: USNM 28509 (<, 36), Georgia, trib. of Altamaha R.
[error for Alapaha R.], a branch of the Suwanee R. at Nashville;
W. J. Taylor.

= Etheostoma (Hololepis) fusiforme barratti (Holbrook). See Hubbs
and Cannon (1935) and Collette (1962).

Poecilichthys radiosus cyanorum Moore and Rigney, 1952, Copeia,
no. 1, p. 10, pl. 1, figs. 3-4.

Holotype: UMMZ 161366 (<7, 68), Oklahoma, Johnston Co., Blue
R., trib. of Red R., on U.S. 99, T. 1 S., R. 6 E.; G. A. Moore; Apr. 4,
1949; orig. OAM 2914.

Paratypes: All from the Blue R., Oklahoma, collected by G. A.
Moore and students. OAM 2913 (96, 27-67), same data as holotype.
UM MZ 161367 (77, 32-70), same locality; Apr. 5, 1947. MCZ 37204
(11, 33-53), Blue R., 10 mi. W. of Wapanucka, 3 mi. S. of rt. 7; Apr.
6, 1947; orig. OAM 1497. UOMZ 26154 (82, 28-53), Little Blue R.,
T.1N.,R.6E., NW. sec. 36; Apr. 5, 1947. UOMZ 26155 (15, 39-49),
Bryan Co., Cedar Cr., 7 mi. E. and 1 mi. S. of Durant; Apr. 15, 1949.
USNM 153532 (49, 38-54), Pontotoc Co., Blue R., near source, SW.
of Ada; orig. OAM 1587.

=Etheostoma (Oligocephalus) radiosum cyanorum (Moore and
Rigney).

Poecilichthys radiosus paludosus Moore and Rigney, 1952, Copeia,
Howie. Wit pl 2, figs. 3-4.
219-948—66——5

66 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119 |

Holotype: UMMZ 161368 (, 47), Oklahoma, Pontotoe Co., Bois |
d’Arc Cr., trib. of Clear Boggy Cr., T. 2 N., R. 6 E.; E. Leonard, J. |
Stevenson, M. Chapman; Apr. 16, 1949; orig. OAM 2933. |

Paratypes: All from Oklahoma. UMMZ 161369 (58, 30-45), same |
data as holotype. MCZ 37205 (41, 36-60), Pontotoc Co., Clear
Boggy Cr.; Moore; Apr. 16, 1947; orig. OAM 1625. UOMZ 26156 !
(19, 28-39), Pontotoc Co., Jack Fork, Clear Boggy system, T. 2 N., |
R. 6 E.; J. M. McDowell, P. White, R. Loomis; Apr. 16, 1949. OAM
1703 (2, 36-41), Kiamichi R., S. of Talahina; Moore et al.; June 7,
1947. USNM 153533 (88, 27-39), Pushmataha Co., small trib., T. 2
N., R. 21 E., sec. 17; A. P. Blair and J. T. Herbelin; Apr. 16, 1949. |
UOMZ 26157 (86, 26-35), Latimer Co., Buffalo Cr., 21 mi. S. of |
Wilburton, T. 3 N., R. 19 E.; Blair and Herbelin; Apr. 17, 1949. |
USNM 198109 (27, 30-39), Pushmataha Co., Kiamichi R., SE. of
Clayton, T. 1 N., R. 19 E., sec. 7; Blair and Herbelin; Apr. 16, 1949;
orig. TUC uncat.

=Etheostoma (Oligocephalus) radiosum paludosum (Moore and
Rigney).

Poecilichthys rufilineatus Cope, 1870, Proc. Amer. Philos. Soc., vol.
it ps 267;

Lectotype: ANSP 13791 (1) North Carolina, Madison Co., Warm
Springs Cr., trib. of French Broad R.; E. D. Cope. Selected and
illustrated by Fowler (1907, p. 525, fig. 9). Still mixed with para-
lectotypes, a total of seven specimens (42-58).

Paralectotypes: ANSP 13792-97 (6), same data as holotype.

= KE theostoma (Nothonotus) rufilineatum (Cope). See Bailey (1959).

Poecilichthys sagitta Jordan and Swain, 1883, Proc. U.S. Nat. Mus.
vol. 6, p. 250.

Holotype: ? (1, 2% in. TL), Kentucky, Whitley Co., Wolf Cr.,
trib. of Clear Fork of Cumberland R., near Pleasant View; D. S.
Jordan, J. Swain, et al.; May 1883. According to the original descrip-
tion, the type should be in the USNM collection, but we have been
unable to find it here or elsewhere.

=LKtheostoma (Oligocephalus) sagitta sagitta (Jordan and Swain).
See Bailey (1948) and Kuehne and Bailey (1961).

Poecilichthys sanguifluus Cope, 1870, Proc. Amer. Philos. Soc., vol.
11, p. 264.

Lectotype: ANSP 13738 (1, 55), Tennessee, headwaters of South
Fork of Cumberland R.; E. D. Cope; herein selected by Dr. Edward
C. Raney.

Paralectotypes: ANSP 13739-40 (2, 48-50), same data as lectotype.
Fowler (1907, p. 524) referred to seven syntypes of P. sanguifluus,

NO. 3550 DARTERS—COLLETTE AND KNAPP 67

ANSP 13738 (type) to 13744. Three are types of Poecilichthys
camurus Cope (q.v.).

=Ltheostoma (Nothonotus) maculatum sanguifluum (Cope) accord-
ing to Raney and Zorach (MS).

Poecilichthys saxatilis Hay, 1881, Proc. U.S. Nat. Mus. vol. 3, p. 495.
Holotype: USNM 27433 (1, 31), Mississippi, Clarke Co., trib. of
Chickasawha R. at Enterprise; O. P. Hay; March-April 1880.
=Etheostoma (Boleosoma) stigmaeum (Jordan). See Bailey, Winn,
and Smith (1954, p. 142).

Poecilichthys spectabilis Agassiz, 1854, Amer. Journ. Sci. and Arts,
vol. 17, p. 304.

Lectotype: UMMZ 86340 (<7, 48), Missouri, Osage R., G. Stolley;
out of MCZ 245034 24529-24722; herein selected by Mr. Donald
A. Distler. D XI-13; A IJ, 6; LL 27+19=46; INF 4+3; pectoral
rays 11; supratemporal canal complete.

Paralectotypes: UMMZ 179528 (6, 27-35), same data as lectotype;
1 specimen of E. caeruleum removed by Distler and recataloged as
UMMZ 179529. MCZ 245034 24529424722 (21, 27-40); 15 of the
specimens in poor condition from drying; 3 specimens of /. caeruleum
removed and recataloged as MCZ 42077. USNM 120284 (5, 32-48),
out of MCZ 24503, 1 specimen of EF. caerulewm removed and recata-
loged as USNM 196738.

=Etheostoma (Oligocephalus) spectabile spectabile (Agassiz). See
Trautman (1930).

Poecilichthys swaini Jordan, 1884, Proc. U.S. Nat. Mus., vol. 7, p. 479.
Holotype: USNM 35308 (<7, 41), Mississippi, Lawrence Co., trib.
of Pearl R. at Monticello.
=Etheostoma (Oligocephalus) swaini (Jordan).

Poecilichthys tetrazonus Hubbs and Black, 1940, Occ. Pap. Mus. Zool.
Univ. Michigan, no. 416, p. 11, pl. 1, fig. 1.

Holotype: UMMZ 111330 (1, 33), Missouri, Dallas Co., Big
Niangua R. at mouth of Greasy Cr., 6 mi. SE. of Buffalo; J. C. Salyer;
Aug. 28, 1931.

Paratype: UMMZ 124600 (1, 22), same data as holotype.

—Etheostoma (Etheostoma) tetrazonum (Hubbs and Black).

Poecilichthys versicolor Agassiz, 1854, Amer. Journ. Sci. and Arts, vol.
17, p. 304.
Lectotype: USNM 1822 (<7, 42), Illinois, small creeks near Quincy ;
L. Watson; herein selected. D IX-14; A II, 7; LL 28+19=47;
INF 4+3; POM 10; supratemporal canal complete.

|
68 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119 |

Paralectotypes: USNM 197493 (5, 34-40), removed from USNM |
1822; 1 specimen that is apparently an aberrant spectabile recata- |
loged as USNM 197443. |

USNM 1816, Missouri, Osage R.; G. Stolley is listed in the USNM |
catalog as type material, but these specimens have apparently been |
mixed with the paralectotypes of Poecilichthys spectabilis Agassiz in |
MOZ 24722.

=Etheostoma (Oligocephalus) spectabile spectabile (Agassiz).

Poecilichthys virgatus Jordan, 1880, Proc. U.S. Nat. Mus., vol. 2, p. |
236.
Lectotype: USNM 23456 (1, 39), Kentucky, Rock Castle R. at
Livingston; D. S. Jordan; selected by Jordan and Evermann (1896, |
p. 1093).
Paralectotypes: Of the ‘numerous typical examples” mentioned
in the original description, we have been able to locate only one |
specimen other than the holotype. UMM2Z 187511 (1, 41), same data, |
orig. IU 451, then IU 4748. A specimen was supposed to have been |
sent to the British Museum, but Collette could find no record of this
having been done and could not find such a specimen in their collection |
in October 1963.
=F theostoma (Catonotus) virgatum (Jordan).

{

Poecilichthys vitreus Cope, 1870, Proc. Amer. Philos. Soc., vol. 11, |
p. 2638.
Holotype: ?, North Carolina, Wake Co., Walnut Cr., trib. of |
Neuse R.; E. D. Cope; late November. Types of other species
described by Cope in the 1870 paper were deposited at ANSP, but |
Fowler (1907) did not mention the type (a female) of this species,
and we have been unable to find it in several thorough searches.
=Etheostoma (Ioa) vitreum (Cope).

Poecilichthys vulneratus Cope, 1870, Proc. Amer. Philos. Soc., vol. 11,
p. 266.

Holotype: ANSP 13798 (o, 44), North Carolina, Madison Co.,
Warm Springs Cr., trib. of French Broad R. Found in bottle con-
taining the types of Poecilichthys rufilineatus Cope by Dr. Edward
C. Raney.

= Etheostoma (Nothonotus) maculatum vulneratum (Cope), according
to Raney and Zorach (MS).

Poecilichthys whipphi montanus Hubbs and Black, 1941, Occ. Pap.
Mus. Zool. Uniy. Michigan, no. 429, p. 15, pl. 1, fig. 2.
Holotype: UMMZ 127777 (, 75), Arkansas, Washington Co.,
near head of Blue Hole Cr., trib. of Clear Cr., 1 mi. S. of Winslow;
J. D. Black and J. Yerton; June 17, 1939.

NO. 3550 DARTERS—COLLETTE AND KNAPP 69

Paratypes: USMM 117589 (7, 17-56), same locality as holotype;
J. D. and R. Y. Black; Aug. 3, 1938; orig. UMMZ 123802. The
remainder of the paratypes are all UMMZ specimens from Arkansas.
128678 (2, 24-48), Crawford Co., Schaberg Cr., trib. of Frog Bayou;
Black, Mills, and Yerton; Aug. 17, 1939. 123806 (100, 16-72),
Washington Co., Blue Hole Cr., Blue Hole, T. 13 N., R. 30 W.; Black
and Yerton; Aug. 3, 1938. 123813 (107, 17-54), Washington Co.,
Railroad Cr., trib. of Frog Bayou, 1.5 mi. S. of Winslow; Black and
Yerton; Aug. 4, 1938. 123821 (53, 17-64), Washington Co., Railroad
Cr., below Cliff Hole, trib. of Frog Bayou, T. 13 N., R. 30 W.; Black
and Yerton; Aug. 4, 1938. 123352 (1, 33), Crawford Co., Jones Cr.,
1 mi. W. of Wintrey, T. 12 N., R. 29 W.; Black and Yerton; June 29,
1938. 123842 (2, 41), Crawford Co., Schaberg Cr., trib. of Frog
Bayou, near Schaberg, T. 12 N., R. 30 W.; Black and party; Aug. 5,
1938. 127778 (7, 27-63), Washington Co., Blue Hole Cr., 1 mi. S.
of Winslow; Black and Yerton; June 17, 1939. 123802 (220, 14-69),
Washington Co., Blue Hole Cr.,1.5 mi. S. of Winslow, T. 13 N.,
R. 30 W.; Black and Yerton; Aug. 3, 1938.

= Etheostoma (Oligocephalus) whipplit montanum (Hubbs and Black).

Poectlichthys whipplii radiosus Hubbs and Black, 1941, Occ. Pap.
Mus. Zool. Univ. Michigan, no. 429, p. 10, pl. 1, fig. 1.

Holotype: UMMZ 123080 (<7, 50), Arkansas, Hot Spring Co.,
Sugar Loaf Cr., trib. of Caddo R., Ouachita R. system; J. D. and R. Y.
Black; June 19, 1938.

Paratypes: USNM 36419 (5, 22-30), Arkansas, Ouachita R. at
Arkadelphia; D. S. Jordan and C. H. Gilbert. USNM 36442 (19,
22-46), Arkansas, Saline R. at Benton; Jordan and Gilbert. USNM
42766 (1, 27), Arkansas, Hot Springs; Meek.

The following paratypes are all UMMZ specimens from Oklahoma:
73056 (1, 45), Kiamichi R., 4 mi. E. of Tuskahoma. 81136 (1, 42),
Le Flore Co., 8 mi. W. of Arkansas line; June 18,1927. 109033 (2,
21-25), Pushmataha Co., Ten Mile Cr., trib. of Kiamichi R., 4 mi.
NW. of Miller; June 26, 1929. 109036 (1, 47), Atoka Co., Little
Boggy Cr., 5 mi. NE. of Atoka; June 28, 1928. 109448 (2, 45),
Pushmataha Co., Walnut Cr., trib. of Kiamichi R., 1 mi. SW. of
Albion; June 19, 1931. 110863 (3, 33-46), Le Flore Co., trib. of Big
Cedar Cr., near NW. end of Rich Mt.; June 26, 1934. 129696 (1, 34),
McCurtain Co., Glover R., one-fourth mi. W. of Glover; W. F. and
P. F. Blair; June 16, 1938. 105393 (1, 27), Big Hell Cr., trib. of
Ouachita R. at Mimoca, 7 mi. N. of Norman; Holloway; Mar. 25,
1937.

The following paratypes are all UMMZ specimens from Arkansas:
123067 (1, 21), Garland Co., Lake Hamilton, T. 3 S., R. 20 W.;

70 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

C. E. Burt; July 6, 1931. 123193 (8, 29-40), Hempstead Co., trib. of
Terre Rouge Cr., 5.5 mi. NE. of Hope on U.S. 67; J. D. and R. Y.
Black; June 22, 1938. 123230 (8, 28), Garland Co., Gulpha Cr. at
jet. with Lake Hamilton, 10 mi. E. of Hot Springs; Black and Black;
June 23, 19388. 127842 (8, 19-38), Lincoln Co., Caney Cr., 1 mi.
N. of Star City; Black and Black; June 20, 1939. 127885 (1, 16),
Bradley Co., Saline R., 5 mi. N. of Warren; Black and Black; June
20, 1939. 128060 (1, 16), Polk Co., trib. of Mountain Fk., 6 mi.
SW. of Mena; Black and Black; July 6,1939. 128106 (1, 36), Howard
Co., Holly Cr., one-half mi. SE. of Dierks; Black and Black; July 7,
1939. 128138 (1, 32), Pike Co., Prairie Cr., Murfreesboro; Black
and Black; July 7, 1939. 128247 (1, 23), Hempstead Co., Middle
Fk. of Bois d’Are Cr., 5.5 mi. NE. of Fulton; Black and Black; July
12, 1939. 128277 (4, 16-24), Garland Co., Gulpha Cr., 6 mi. SE. of
Hot Springs; Black and Black; July 12, 1939.

=Ktheostoma (Oligocephalus) radiosum radiosum (Hubbs and Black).
See Moore and Rigney (1952).

Poecilichthys zonalis Cope, 1868, Journ. Acad. Nat. Sci. Philadelphia,
ser. 2, vol. 6, p. 212, color pl. 24, fig. 1.

Lectotype: ANSP 13799 (0, 42), Virginia, Holston R.; E. D. Cope;
herein selected by Dr. Chu-fa Tsai. D XI-11; A 1,7; LL 55; INF 4
+3; POM 10; branchisotegals 5; chest naked; cheeks, opercles, and
belly completely covered with scales; 6 dorsal blotches.

Paralectotypes: ANSP 13800-1 (9, 38; &, 47), same locality data
as lectotype.

Fowler (1907, p. 524) erroneously designated ANSP 14036 lectotype
and ANSP 14037-88 paralectotypes. ‘These specimens are not types
of P. zonalis but are some of the syntypes of Hyostoma simoterum
Cope. The color plate accompanying Cope’s original description
shows clearly that Cope had both EF. zonale and E. simoterum from
the Holston. Fowler’s figure 8, which is labeled as the type of P.
zonalis, represents a specimen of Hyostoma simoterum.

=Ktheostoma (Etheostoma) zonale zonale (Cope).

Poecilosoma erythrogastrum Kirtland, 1854, Annals of Science, includ.
Trans. Cleveland Acad. Nat. Sci., vol. 2, no. 1, p. 4.
Syntypes: USNM 1815 (?), Ohio, Rocky R., 7 mi. W. of Cleve-
land; J. P. Kirtland. We have been unable to locate this collection.
=Etheostoma (Oligocephalus) caeruleum caeruleum Storer. See
Knapp (1964).

Poecilosoma transversum Abbott, 1860, Proc. Acad. Nat. Sci. Phila-
delphia, vol. 12, p. 326.

Lectotype: ANSP 13810 (1, 38), locality unknown; number selected

by Fowler (1907, p. 525); specimen herein selected. D X-13; A II,7;

NO. 3550 DARTERS—COLLETTE AND KNAPP 71

LL 31+15=46; INF 4+3; supratemporal canal complete. Abbott
noted that there were no data with the specimens and assumed that
they might have come from Lake Superior. This could not be so
because E. caeruleum does not occur there.
Paralectotypes: ANSP 13811-19 (9, 28-42), same data as lectotype.
=Etheostoma (Oligocephalus) caeruleum caeruleum Storer. See
Knapp (1964).

Rheocrypta copelandi Jordan, 1877, U.S. Nat. Mus. Bull. 10, p. 9.

Lectotype: USNM 20143 (<7, 43), Indiana, White R., 5 mi. N.
of Indianapolis; Jordan and Copeland; herein selected. D XI-12;
AII,9; LL 53; INF 8; POM 10; opercles scaled; cheeks naked. Jordan
and Evermann (1896, p. 1046) listed USNM 20143, which contained
two specimens, as type; herein further restricted.

Paralectotypes: USNM 197996 (7, 43), removed from USNM
20143; USNM 23461 (<7, 44); MOZ 24392 (<7, 39), and BMNH 1880.
1.21.36-37 (2 o&, 43-45); all part of Jordan’s material originally “some
thirty in number” from the type locality.

=Percina (Cottogaster) copelandi (Jordan).

Richia brevispina Coker, 1926, Bull. Bur. Fish., vol. 42, p. 106, fig. 1.

Holotype: USNM 87411 (co, 36), North Carolina, Burke Co.,
Paddy’s Cr., just above head of Paddy’s Cr. Lake, part of artificial
Lake James system near Bridgewater, Catawba drainage; R. E.
Coker; August 1922.

Paratypes: USNM 87412 (2, 27-36), same data as holotype.

=Ktheostoma (Catonotus) flabellare ssp. See Hubbs (1927) and
Myers (1927). [= EF. (C.) f. brevispina (Coker) according to Ross
and Carico (1963, p. 13)].

Sciaena caprodes Rafinesque, 1818, Amer. Month. Mag. and Critical
Rev., vol. 3, no. 5, p. 354.

Types: Type locality—the Ohio River. As Call (1899) noted,
Rafinesque did not preserve any of his type material. Examination
of a sketch of this species in Rafinesque’s notebook (fig. 5) shows that
the name is correctly attributed.

=Percina (Percina) caprodes caprodes (Rafinesque).

Ulocentra gilberti Evermann and Thoburn in Jordan and Evermann,
1896, U.S. Nat. Mus. Bull. 47, p. 1049.

Holotype: USNM 47531 (1, 38), Tennessee, Clinch R. at Walker’s
Ford, near Tazewell; B. W. Evermann, J. T. Scovell, and R. R.
Gurley; Oct. 12, 1893; figured in Jordan and Evermann (1900, fig.
446).

Paratypes: USNM 125373 (1, 36), orig. BF 285, and SU 1954
(2, 47), same data as holotype. The original description was sup-

de PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

posedly based on three specimens, so there may have been an error
in the original description. The SU bottle was broken during the
1906 earthquake, so there is also a possibility that another specimen
of the same species became mixed with SU 1954.

=Percina (Cottogaster) copelandi (Jordan).

Ulocentra meadiae Jordan and Evermann, 1898, U.S. Nat. Mus.

Bull. 47, p. 2852.

Holotype: USNM 48903 (<’, 46), Tennessee, Indian Cr., trib. of
Powell R. at Cumberland Gap; R. R. Gurley; Oct. 17, 1893; figured
by Jordan and Evermann (1900, fig. 447).

Paratypes: USNM 125623 (07, 46), same data as holotype; orig.
BF 711. The third paratype was sent to SU according to a U.S.
Fish Comm. distribution list, but Bohlke (1953) did not list it among
the Stanford types, and we were unable to find it there.

=Etheostoma (Boleosoma) jessiae (Jordan and Brayton).

Vaillantia chlorosoma Hay, 1881, Proc. U.S. Nat. Mus., vol. 3, p. 495.

Syntypes: USNM 27428, Mississippi, branch of Tuscumbia R. at
Corinth; O. P. Hay; March—April 1880; completely disintegrated and
dried by 1987. SU 726 (8 9, 37-41), Mississippi, Macon; Hay.
Although not listed as types by Béhlke (1953), the data on the label,
plus a label reading ‘‘cotypes,” lead us to believe this is part of the

Figure 5.—Sciaena caprodes Rafinesque (sketch from Rafinesque’s notebook).

original type material. The original description was based on speci-
mens from the Tuscumbia R. at Corinth, Sandy Cr. at Artesia, and
Horsehunter Cr. at Macon. Some of this material was placed in the
U.S. National Museum, some in Jordan’s collections, and the re-
mainder in the Butler University collection. It seems likely that
SU 726 represents material given to Jordan by Hay.

=Etheostoma (Boleosoma) chlorosomum (Hay).

Villora edwinit Hubbs and Cannon, 1935, Misc. Publ. Mus. Zool.
Univ. Michigan, no. 30, p. 18, pl. 1, fig. 1.

NO. 3550 DARTERS—COLLETTE AND KNAPP 73

Holotype: UMMZ 87892 (<7, 37), Florida, Alachua-Columbia Co.
line, Santa Fe R. at Poe Springs; E. T. Boardman; Feb. 8, 1928.

Paratypes: UMMZ 107047 (3, 30-35) and USNM 94684 (9, 33),
same data as holotype. UMMZ 101680 (2 9, 33-35), same locality;
J. D. Kilby and A. F. Carr; Mar. 19, 1934. UMMZ 88685 (1, 25),
Georgia, Early Co., trib. of Chattahoochee R., 2 mi. NW. of Jakin;
E. P. Creaser and H. R. Becker; Sept. 15, 1929. MCZ 98 (1, ?),
“Ga.”; Barratt; no longer present in MCZ collection.

—Etheostoma (Villora) edwint (Hubbs and Cannon). See Collette
and Yerger (1962).

Villora okaloosae Fowler, 1941, Proc. Acad. Nat. Sci. Philadelphia,
vol. 92, p. 242, fig. 12.

Holotype: ANSP 69159 (9, 24), Florida, Okaloosa Co., Little
Rocky Cr., 7 mi. NE. of Niceville on rt. 218 (now Fla. 285); F. Harper;
June 20, 1939. Holotype redescribed by Collette and Yerger (1962).

—Etheostoma (Villora) okaloosae (Fowler). See Yerger (1960) and
Collette and Yerger (1962).

Checklist of Nominal Species

acuticeps Bailey, Etheostoma. (UMMZ)

aesopus Cope, Boleosoma. (ANSP?)
alabamae—see whipplei alabamae.

anceps Vaillant, Plesioperca. (MNHN)
appalachia—see barratti appalachia.
arcansanum—see zonale arcansanum.
arcus-celestis Crevecoeur, Etheostoma. (USNM?)
artesiae Hay, Poecilichthys. (USNM)

asprellus Jordan, Pleurolepis. (INHS)
asprigenis Forbes, Poecilichthys. (SU)

aspro Cope and Jordan, Alvordius. (X)
atraque—see fusiformis atraque.

atripinnis Jordan, Arlina. (USNM)
atromaculata Girard, Estrella. (?)

aubcenaubei Evermann, Etheostoma. (USNM)
aurantiacus Cope, Cottogaster. (ANSP)

australe Jordan, Etheostoma—see fasciatus, Diplesion.

Barratti Holbrook, Boleosoma. (MCZ, UMMZ)
barratti appalachia Bailey, Hololepis. (UMMZ, CU, DU)

beani Jordan, Poecilichthys. (USNM)

|

beanii Jordan, Ammocrypta. (USNM)

bimaculata Haldeman, Percina. (USNM?)

binotatum—see hopkinsi binotatum.
219-948—66 6

74 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119)

blennioides Rafinesque, Etheostoma. (X)

blennioperca Cope, Hyostoma. (ANSP)

blennius Gilbert and Swain, Etheostoma. (USNM, SU)
borealis Jordan, Poecilichthys. (USNM, SU)
brevipinnis—see olmstedi brevipinnis.

brevispina Coker, Richia. (USNM)

butlerianus Hay, Poecilichthys. (USNM)

burtoni—see caprodes burton.

caerulea Storer, Htheostoma. (MCZ)

camurum Forbes, Boleosoma. (USNM, SU, MCZ, UMMZ)
camurus Cope, Poecilichthys. (ANSP)

caprodes Rafinesque, Sciaena. (X)

caprodes burtont Fowler, Percina. (ANSP)

carbonaria Baird and Girard, Pileoma. (USNM, MCZ, UMMZ)
cheneyt Evermann and Kendall, Cottogaster. (USNM, SU)
chlorosoma Hay, Vaillantia. (USNM, SU)

cinerea Storer, EKtheostoma. (?)

clara Jordan and Meek, Ammocrypta. (USNM, SU, UMMZ)
collis Hubbs and Cannon, Hololepis. (UMMZ)

collis lepidinion Collette, Etheostoma. (USNM, CU, DU)
coosae Fowler, Poecilichthys. (ANSP)

copelandi Jordan, Rheocrypta. (USNM, MCZ, BMNH)
cragini Gilbert, Etheostoma. (USNM)

crassus Jordan and Brayton, Alvordius. (USNM)
cumberlandicum Jordan and Swain, Etheostoma. (USNM)
cyanorum—see radiosus cyanorum.

cymatogramma Abbott, Pileoma. (ANSP)

cymatotaenia Gilbert and Meek, Etheostoma. (USNM, SU)

davisont Hay, Etheostoma. (USNM)

ditrema Ramsey and Suttkus, LHtheostoma. (TU, USNM, MCZ, |

ANSP, CU, SU, UMMZ)
duryi Henshall, Htheostoma. (X)

edwint Hubbs and Cannon, Villora. (UMMZ, USNM, MCZ)
effulgens Girard, Arlina. (USNM)

elegans Girard, Boleichthys. (MCZ)

elegans Hay, Nanostoma. (USNM, SU)

eos Jordan, Boleichthys. (USNM)

EFiZONUS—SeE EUZONUS eEriZONUS.

erochrous Cope, Hololepis. (MNHN)

erythrogastrum Kirtland, Poecilosoma. (USNM?)

eulepis—see nigrum eulepis.

euzonus erizonus Hubbs and Black, Poecilichthys. (UMMZ)
euzonus euzonus Hubbs and Black, Poecilichthys. (UMMZ, USNM)

NO. 3550 DARTERS—COLLETTE AND KNAPP 75

evermannt Moenkhaus, Hadropterus. (USNM)
evides Jordan and Copeland, Alvoridus. (USNM, MCZ)
exilis Girard, Boleichthys. (USNM, MCZ, UMMZ)

fasciatus Girard, Catonotus. (USNM, MCZ)

fasciatus Girard, Diplesion. (MCZ, UMMZ)

flabellaris Rafinesque, Etheostoma. (X)

fonticola Jordan and Gilbert, Alvarius. (USNM)

fontinalis Rafinesque, Etheostoma. (X)

formosa Henshall, Htheostoma. (°?)

fricksia Hildebrand, Htheostoma. (USNM)

fusiforme Girard, Boleosoma. (USNM, MCZ, UMMZ)

fusiformis atraque Hubbs and Cannon, Hololepis. (UMMZ, USNM)

fusiformis insulae Hubbs and Cannon, Hololepis. (MCZ, UMMZ)

fusiformis metae-gadi Hubbs and Cannon, Hololepis. (USNM,
UMMZ)

gelida Hay, Ammocrypta. (USNM)

gilberti Evermann and Thoburn, Ulocentra. (USNM, SU)
gracile Girard, Boleosoma. (USNM, MCZ)

grahami Girard, Oligocephalus. (MCZ)

gintherit Kigenmann and EKigenmann, Etheostoma. (BMNH)
gutselli Hildebrand, Poecilichthys. (USNM)

hildebrandti Evermann and Clark, Etheostoma. (USNM)

histrio Jordan and Gilbert, Etheostoma. (USNM, SU)

hopkinsi Fowler, Poecilichthys. (ANSP)

hopkinsi binotatum Bailey and Richards, Etheostoma. (CU, UMMZ,
USNM)

humeralis Girard, Oligocephalus. (USNM, MCZ)

inscriptus Jordan and Brayton, Nothonotus. (USNM)
insulae—see fusiformis insulae.
iowae Jordan and Meek, Etheostoma. (SU)

gessiae Jordan and Brayton, Poecilichthys. (?)
jordani Gilbert, Etheostoma. (USNM, CAS, BMNH, UMMZ)
juliae Meek, Etheostoma. (USNM, SU)

kanawhae Raney, Poecilichthys. (UMMZ, CU, USNM)
Kennicotti Putnam, Catonotus. (USNM, MCZ, UMMZ)

lenticula Richards and Knapp, Pereina. (CU, TU, USNM, UMMZ)
leonensis Girard, Oligocephalus. (MCZ)

lepida Baird and Girard, Boleosoma. (USNM, MCZ, UMMZ)
lepidinion—see collis lepidinion.

lepidogenys Evermann and Kendall, Etheostoma. (USNM)

76 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

lineolatus Agassiz, Catonotus. (MCZ, UMMZ, MNHN)
Linsleyi Storer, Htheostoma. (?)

longimana Jordan, Htheostoma. (MCZ, USNM, UMMZ)
luteovinctum Gilbert and Swain, Htheostoma. (USNM, SU)
lynceum Hay, Etheostoma. See elegans, Nanostoma for types.

macrocephalum Cope, Etheostoma. (ANSP)

maculata Kirtland, Etheostoma. (?)

maculaticeps Cope, Boleosoma. (ANSP)

maculatum Agassiz, Boleosoma. (USNM)

maculatus Girard, Alvordius. (USNM?)

maculatus Girard, Hadropterus. (USNM)

manitou Jordan, Percina. (USNM)

mariae Fowler, Belophlor. (ANSP)

maxinkuckiensts Evermann, Hadropterus. (USNM)

meadiae Jordan and Evermann, Ulocentra. (USNM)

mesaeus Cope, Poecilichthys. (ANSP)

metae-gadi—see fusiformis metae-gadt.

microperca Jordan and Gilbert, Htheostoma. See punctulata, Micro-
perca for types.

micropterus Gilbert, Etheostoma. (USNM)

minima Haldeman, Perca. (ANSP)

montanus—see whipplii montanus.

montuosa—see notogramma montuosa.

moorei Raney and Suttkus, Htheostoma. (CU, TU, USNM, UMMZ,
UK)

mutatum Vaillant, Boleosoma. See maculatum, Boleosoma for types.

nasutus Bailey, Hadropterus. (UMMZ, USNM)

nebulosa Haldeman, Perca. (ANSP)

nevisense Cope, Etheostoma. (ANSP?)

Newmani Agassiz, Hyostoma. (USNM, MCZ, UMMZ)

nianguae Gilbert and Meek, Etheostoma. (USNM, SU)

nianguae spilotum Gilbert, Etheostoma. (USNM, SU)

nigra Rafinesque, Htheostoma. (X)

nigrofasciata raneyi Crawford, Percina. (CU, USNM)

nigrofasciatus Agassiz, Hadropterus. (MCZ, USNM, UMMZ)

nigrofasciatus westfalli Fowler, Hadropterus. (ANSP)

nigrum eulepis Hubbs and Greene, Boleosoma. (UMMZ,USNM, UW)

notogramma montuosa Hogarth and Woolcott, Percina. (USNM,
UMMZ, CU, UR, VPI)

notogrammus Raney and Hubbs, Hadropterus. See maculatus,
Hadropterus for types.

nuchale Howell and Caldwell, Etheosoma. (UMMZ, USNM, TU,
UAIC)

NO. 3550 DARTERS—COLLETTE AND KNAPP 77

obeyense Kirsch, Etheostoma. (USNM, SU, BMNH)

okaloosae Fowler, Villora. (ANSP)

oligoporus— see thermophilus oligoporus.

olmstedi Storer, Htheostoma. (MCZ)

olmstedi brevipinnis Cope, Boleosoma. (ANSP)

osburnt Hubbs and Trautman, Poecilichthys. (UMMZ, USNM, OSM)
ouachitae Jordan and Gilbert, Htheostoma. (USNM, UMMZ)
oryrhynchus Hubbs and Raney, Hadropterus. (UMMZ, USNM)

pager Meek, Etheostoma. (USNM)

pallididorsum Distler and Metcalf, Htheostoma. (KU, USNM,
UMMZ, CU)

palmaris Bailey, Hadropterus. (UMMZ, USNM)

paludosus—see radiosus paludosus.

palustris Gilbert, Poecilichthys. (USNM)

pantherinus Moore and Reeves, Hadropterus. (UMMZ, OAM, MCZ,
USNM)

parvipinne Gilbert and Swain, Etheostoma. (USNM)

pellucidus Agassiz, Pleurolepis. (USNM, MCZ, UMMZ)

peltatum Stauffer, Htheostoma. (ANSP)

perlongum Hubbs and Raney, Boleosoma. (UMMZ)

phlox Cope, Boleosoma. (ANSP)

phoxocephalum Nelson, Etheostoma. (UMMZ)

podostemone Jordan and Jenkins, Htheostoma. (USNM)

potsti Girard, Aplesion. (MCZ)

procharis Hay, Microperca. (USNM)

pulchellus Girard, Oligocephalus. (MCZ)

punctulata Putnam, Microperca. (USNM, MCZ, UMMZ).

punctulatus Agassiz, Poecilichthys. (MCZ)

putnami Jordan and Gilbert, Cottogaster. (USN M)

quapella Eigenmann and Eigenmann, Etheostoma. (BMNH)
quiescens Jordan, Poecilichthys. (USN M)

radiosus—see whipplii radiosus.

radiosus cyanorum Moore and Rigney, Poecilichthys. (UMMZ, OAM,
MCZ, USNM, UOMZ)

radiosus paludosus Moore and Rigney, Poecilichthys. (UMMZ, MCZ,
UOMZ, OAM, USNM)

raneyi—see nigrofasciata raneyt.

rez Jordan and Evermann, Etheostoma. (USN M)

roanoka Jordan and Jenkins, Etheostoma. (USNM, SU, CU, MCZ)

rufilineatus Cope, Poecilichthys. (ANSP)

rupestre Gilbert and Swain, Etheostoma. (USN M)

sagitta Jordan and Swain, Poecilichthys. (USN M?)

78 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

saludae Hubbs and Cannon, Hololepis. (UMMZ, CM, FMNH)

sanguifluus Cope, Poecilichthys. (ANSP)

saxatilis Hay, Poecilichthys. (USN M)

sclerus Swain, Hadropterus. (USNM)

sclerus apristis Hubbs and Hubbs, Hadropterus. (UMMZ, USNM,
TNHC, FMNH, OAM, TAMC, SU)

scierus serrula Jordan and Gilbert, Hadropterus. (USNM)

sciotense Osburn and Williamson, Htheostoma. (USN M)

scovelliti Woolman, Etheostoma. (USNM, BMNH, CAS)

sellaris Radcliffe and Welsh, Hadropterus. (USNM)

semifasciatum DeKay, Pileoma. (?)

serrifer Hubbs and Cannon, Hololepis. (UMMZ, USNM, CM)

serrula—see scierus serrula.

shumardi Girard, Hadropterus. (USN M)

simoterum Cope, Hyostoma. (ANSP, USNM, MNHN)

spectabilis Agassiz, Poecilichthys. (UMMZ, MCZ, USNM)

spillmani Hay, Hadropterus. (USNM)

spilotum—see nianguae spilotum.

squamatus Gilbert and Swain, Etheostoma. (USNM)

squamiceps Jordan, Etheostoma. (USN M)

stigmaeum Jordan, Boleosoma. (ANSP)

susanae Jordan and Swain, Boleosoma.(USNM)

swaint Jordan, Poecilichthys. (USN M)

swannanoa Jordan and Evermann, Etheostoma. (USNM, SU, UMMZ)

tessellata Storer, Etheostoma. (?)

tessellatum DeKay, Boleosoma. (?)

tessellatus Jordan, Hadropterus. (USNM)

tetrazonus Hubbs and Black, Poecilichthys. (UMMZ)

thalassinus Jordan and Brayton, Nothonotus. (USNM, SU, MCZ,
BMNH)

thermophilus Hubbs and Cannon, Hololepis. (UMMZ, USNM)

thermophilus oligoporus Bailey and Frey, Hololepis. (UMMZ, DU)

tippecanoe Jordan and Evermann, Etheostoma. (USNM)

transversum Abbott, Poecilosoma. (ANSP)

trisella Bailey and Richards, Etheostoma. (UMMZ)

tuscumbia Gilbert and Swain, Etheostoma. (SU, BMNH, UMMZ,
USNM)

uranidea Jordan and Gilbert, Etheostoma. (USNM)

variata Kirtland, Etheostoma. (?)

verecundum Jordan and Evermann, Etheostoma. (USNM)
versicolor Agassiz, Poecilichthys. (USN M)

vextllare Jordan, Boleosoma. (?)

vigil Hay, Joa. (USNM)

NO. 3550 DARTERS—COLLETTE AND KNAPP 79

_ vinctipes Jordan, Nanostoma. (USNM)

virgatus Jordan, Poecilichthys. (USNM, UMMZ)
vitreus Cope, Poecilichthys. (ANSP?)

vivax Hay, Ammocrypta. (USNM)

_ vulneratus Cope, Poecilichthys. (ANSP)

warreni Girard, Boleichthys. (MCZ)

— -westfalli—see nigrofasciatus westfalli.

_whipplei alabamae Gilbert and Swain, Etheostoma. (USNM, SU)
whipplii Girard, Boleichthys. (MCZ)

whipplu montanus Hubbs and Black, Poecilichthys. (UMMZ, USNM)
whipplu radiosus Hubbs and Black, Poecilichthys. (UMMZ, USNM)
wrightt McCormick, Etheostoma. (?)

zebra Agassiz, Pileoma. (MCZ, UMMZ)

zonale arcansanum Jordan and Gilbert, Etheostoma. (USNM, UMMZ)
zonalis Cope, Poecilichthys. (ANSP)

zonifer Hubbs and Cannon, Hololepis. (UMMZ)

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82 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

GILBERT, CHARLES H.
1884. A list of fishes collected in the east fork of White River, Indiana, with
descriptions of two new species. Proc. U.S. Nat. Mus., vol. 7, pp.
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1884. Notes on the fishes of Switz City Swamp, Greene County, Indiana.
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1885. Second series of notes on the fishes of Kansas. Bull. Washburn Coll.
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1887. Description of new and little known etheostomoids. Proc. U.S.
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1890. Description of a new species of Etheostoma (E. micropterus) from
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1891. Report of explorations made in Alabama during 1889, with notes on
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GIRARD, CHARLES
1854. Description of some new species of fish from the state of Massachusetts
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1859. Ichthyological notices, 5-27. Proc. Acad. Nat. Sci. Philadelphia,
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HALDEMAN, SAMUEL S.
1842. Description of two new species of the genus Perca, from the Susque-
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1844. [Percina bimaculata, n. sp. from the Susquehannah.] Proc. Boston
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Hay, Outver P.
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1883. On a collection of fishes from the Lower Mississippi valley. Bull.
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HILDEBRAND, SAMUEL F.
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Ho.prook, Joun E.
1855. An account of several species of fish observed in Florida, Georgia,
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NO. 3550 DARTERS—COLLETTE AND KNAPP 83

Howe Lt, Witu1aM MIKE, and CaLpweELtL, Ricnuarp Dae
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Hupsss, Caru L.
1926. A check-list of the fishes of the Great Lakes and tributary waters,
with nomenclatorial notes and analytical keys. Misc. Publ. Mus.
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Husss, Cart L. and Buack, Joun D.
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Husss, Cari L., and GREENE, C. WILLARD
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Husss, Cart L., and Orrensureasr, A. I.
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Housss, Cart L., and RaNnsy, Epwarp C.
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Husss, Cart L., and Trautman, MiTon B.
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Husss, CLark
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1877. Onthe fishes of northern Indiana. Proc. Acad. Nat. Sci. Philadelphia,
vol. 29, pp. 42-82.

84

1877.

1877.

1878.

1878.

1878.

PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

A partial synopsis of the fishes of upper Georgia. Ann. New York
Lyceum Nat. Hist., vol. 11, pp. 307-369.

Contributions to North American ichthyology based primarily on
the collections of the United States National Museum. 2. A—
Notes on Cottidae, Etheostomatidae, Percidae, Centrarchidae,
Aphododeridae, Dorysomatidae, and Cyprinidae, with revisions
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A catalogue of the fishes of Illinois. Bull. Illinois State Lab. Nat.
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A catalogue of the fishes of the fresh waters of North America.
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A manual of the vertebrates of the northern United States, including
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1879-80. Notes on certain typical specimens of American fishes in the

1880.

1884.

1885.

1885.

1885.

1888.

1888.

1889.

1919.

1929.

British Museum and inthe Museum d’Histoire Naturelle at
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Description of new species of North American fishes. Proc. U.S.
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Descriptions of four new species of Poecilichthys in the United States
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Supplementary notes on North American fishes. Proc. U.S. Nat.

Mus, vol. 7, pp. 545-548.

On the EHtheostoma variatum of Kirtland. Proc. U.S. Nat. Mus.,
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A catalogue of the fishes known to inhabit the waters of North
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A manual of the vertebrate animals of the northern United States,
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Description of a new species of Etheostoma (FH. longimana) from
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Descriptions of fourteen species of fresh-water fishes collected by the
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New genera of fishes. Proc. Acad. Nat. Sci. Philadelphia, vol. 70,
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Manual of the vertebrate animals of the northeastern United States
inclusive of marine species, 13th ed., 446 pp.

JORDAN, Davin S., and Brayton, ALEMBERT W.

1878.

Contributions to North American ichthyology, 3: On the distribu-
tion of the fishes of the Alleghany region of South Carolina, Georgia,
and Tennessee, with descriptions of new or little known species.
U.S. Nat. Mus. Bull. 12(A), pp. 1-95.

No. 3550 DARTERS—COLLETTE AND KNAPP 85

JoRDAN, Davip §., and EvERMANN, Barron W.

1890. Description of a new species of fish from Tippecanoe River, Indiana.
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1896. The fishes of North and Middle America: a descriptive catalogue
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1898,1900. The fishes of North and Middle America: a descriptive cata-
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JORDAN, Davin S., EvERMANN, Barton W., and Cuarx, Howarp W.

1930. Check list of the fishes and fishlike vertebrates of North and Middle
America north of the northern boundary of Venezuela and Colom-
bia. Rept. U.S. Comm. Fish., App. 10, 670 pp.

JORDAN, Davin S., and GILBERT, CHARLES H.

1877. On the genera of North American fresh-water fishes. Proc. Acad.
Nat. Sci. Philadelphia, vol. 29, pp. 83-104.

1883. Synopsis of the fishes of North America. U.S. Nat. Mus. Bull. 16,
1018 pp.

1886. List of fishes collected in Arkansas, Indian Territory, and Texas,
in September, 1884, with notes and descriptions. Proc. U.S. Nat.
Mus., vol. 9, pp. 1-25.

JORDAN, Davin S., and Msrkx, Seru EH.

1885. List of fishes collected in Iowa and Missouri in August, 1884, with
descriptions of three new species. Proc. U.S. Nat. Mus., vol. 8, pp.
1-17.

JORDAN, Davin S., and Swan, JOSEPH

1883. List of fishes collected in the Clear Fork of the Cumberland, Whitley
County, Kentucky, with descriptions of three new species. Proc.
U.S. Nat. Mus., vol. 6, pp. 248-251.

Kirscu, Puinie H.

1892. Notes on the streams and fishes of Clinton County, Kentucky, with
a description of a new darter. Bull. U.S. Fish. Comm., 1890,
vol. 10, pp. 289-292.

KIRTLAND, JARED P.

1841. Descriptions of four new species of fishes. Boston Journ. Nat.
Hist., vol. 3, pp. 273-279.

1841. Descriptions of the fishes of the Ohio River and its tributaries.
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1854. Poecilosoma erythrogastrum. Annals of Science, including Trans.
Cleveland Acad. Nat. Sci., vol. 2, no. 1, pp. 4-5.

Knapp, Lesiie W.

1964. A systematic study of Etheostoma caeruleum Storer. Ph. D. thesis,
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Knapp, Lesuiz W.; Ricuarps, Winu1aM J.; Miuier, Rospert V.; and Fosrsr,
Neat R.

1963. Rediscovery of the percid fish Etheostoma sellare (Radcliffe and

Welch). Copeia, 1963, no. 2, p. 455.
Kurune, Rosert A., and Baitey, Reeve M.

1961. Stream capture and the distribution of the percid fish Htheostoma
sagitta, with geologic and taxonomic considerations. Copeia, 1961,
no. 1; pp. 1-8.

86 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119 |

Lacuer, Karu F., and Battey, RreEvE M.

1947. The genetic fixity of differential characters in subspecies of the percid |
fish, Boleosoma nigrum. Copeia, 1947, no. 1, pp. 50-59.

LInDER, ALLAN D.

1959. The American percid fishes Ammocrypta clara Jordan and Meek and
Ammocrypta pellucida (Baird). Southwest. Nat., vol. 4, no. 4, |
pp. 176-184. |

McCormick, Lewis M.

1892. Descriptive list of the fishes of Lorain County, Ohio. Bull. Oberlin |

Coll. Lab., no. 2, 34 pp.
Meek, Seta E.
1891. Report of explorations made in Missouri and Arkansas during 1889,

with an account of the fishes observed in each of the river basins |

examined. Bull. U.S. Fish. Comm., 1889, vol. 9, pp. 113-141.
1894. A new Etheostoma from Arkansas. Amer. Nat., vol. 28, p. 957.
Miuier, Rosperr V.
1964. A systematic study of the greenside darter, Etheostoma blennioides
Rafinesque. Ph. D. thesis, Cornell University.
Mornxkuavs, WILLIAM J.
1903. Description of a new species of darter from Tippecanoe Lake. Bull. |
U.S. Fish Comm. 1902, vol. 22, pp. 397-398.
Moors, GrorGce A., and Cross, Frank B.
1950. Additional Oklahoma fishes with validation of Poecilichthys parvi- |
pinnis (Gilbert and Swain). Copeia, 1950, no. 2, pp. 139-148.
Moore, Grorce A., and Rerves, Jonges D.
1955. Hadropterus pantherinus, a new percid fish from Oklahoma and
Arkansas. Copeia, 1955, no. 2, pp. 89-92.
Moore, Grorce A., and RiagNry, Cart C.
1952. Taxonomic status of the percid fish Poecilichthys radiosus in Oklahoma
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Myers, Grorce S.
1927. The status of the darter Richiella brevispina (Coker). Copeia,
no. 163, pp. 39-43.

Netson, Epwarp W.
1876. <A partial catalogue of the fishes of Illinois. Bull. Illinois Mus. Nat.
Hist., no. 1, pp. 33-52.

OsBuRN, Raymonp C., and WiuuIAMson, E. B. |
1898. A list of the fishes of Franklin County, Ohio, with a description of a |
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vol. 6, pp. 11-20.
1899. Additional notes on the fishes of Franklin County, Ohio. Ann. Rept.
Ohio State Acad. Sci., vol. 7, pp. 33-34.
Putnam, FREDERICK W.
1863. List of the fishes sent by the museum to different institutions, in
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RavDcuiFrFe, Lewis, and WeisH, WILLIAM W.
1913. Description of a new darter from Maryland. Bull. U.S. Bur. Fish.,
1912, vol. 32, pp. 29-32.

NO. 3550 DARTERS—COLLETTE AND KNAPP 87

RAFINESQUE, CONSTANTINE S.
1818. Discoveries in natural history, made during a journey through the
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1819. Prodrome de 70 nouveaux genres d’animaux découverts dans
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1965. Htheostoma ditrema, a new darter of the subgenus Oligocephalus
(Percidae) from springs of the Alabama River basin in Alabama and
Georgia. Tulane Stud. Zool., vol. 12, no. 3, pp. 65-77.
Raney, Epwarp C.
1941. Poecilichthys kanawhae, a new darter from the upper New River
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Raney, Epwarp C., and Huspss, Cart L.
1948. Hadropterus notogrammus, a new percid fish from Maryland, Virginia,
and West Virginia. Occ. Pap. Mus. Zool. Univ. Michigan, no. 512,
26 pp.
RaneEy, Epwarp C., and Surruks, Royau D.
1964. Htheostoma mooret, a new darter of the subgenus Nothonotus from the
White River system, Arkansas. Copeia, 1964, no. 1, pp. 130-139.
Raney, Epwarp C., and Zoracu, Timotuy
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1964. Percina lenticula, a new percid fish, with a redescription of the sub-
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RICHARDSON, LAURENCE R.
1938. A note on variation in squamation of the cheek operculum in two
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Ross, Ropert D., and Carico, JaAmzs E.
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1947. Notes on two darters of the genus Boleosoma. Copeia, 1947, no.
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1843. Notice of a portion of Dr. Dekay’s report on the fishes of New York.
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88 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

1846. Asynopsis of the fishes of North America. Mem. Amer. Acad. Arts,
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1883. A description of a new species of Hadropterus (Hadropterus scierus)
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U.S. GOVERNMENT PRINTING OFFICE: 1966

Proceedings of

the United States

National Museum

SMITHSONIAN INSTITUTION +« WASHINGTON, D.C.

Volume 119 1966 Number 3551

REVIEW OF NEW WORLD MOTHS
OF GENUS EUCHROMIUS GUENEE
WITH DESCRIPTIONS OF TWO NEW SPECIES
(LEPIDOPTERA: CRAMBIDAE)

By Haun W. Capps!

The purpose of this paper is to provide a means for accurate identi-
fication of the species of Huchromius Guenée occurring in the Western
Hemisphere and to provide names for two undescribed species,
heretofore confused with Euchromius ocelleus (Haworth).

Delineations of genitalia were prepared by A. D. Cushman, scientific
illustrator, Entomology Research Division, Agr. Res. Serv., U.S.
Department of Agriculture, and are not drawn to scale. The male
genitalia are in lateral view, with one harpe omitted and the aedeagus
removed. The female genitalia are in ventral view. Photos of adults
are by J. Scott, staff photographer, Smithsonian Institution, and are
twice natural size.

1 Entomology Research Division, Agricultural Research Service, U. S. Depart-
ment of Agriculture. Retired September 26, 1964.
1

PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 119

Keys to New World Species of Euchromius

BASED ON COLOR AND DISTRIBUTION

Forewing brownish from base to inner transverse postmedial band, with
little or no irroration (pl. 1: fig. 1) . . . . . galapagosalis, new species
Forewing ochreous white from base to inner transverse postmedial band,
with fuscous irroration variable but distinct (pl. 1: figs. 2-4) . . ... 2
Forewing with whitish area adjacent to blackish terminal dots narrow; a
fine ochreous line about midway between dots and termination of irrorated
areal figs: 2,3)! si sis, ose ae, Pores
Forewing with whitish area broader fa mati ime mel Blaeer: to termination
of the irrorated area than to terminal dots (pl. 1: fig. 4).
californicalis (Packard)
Forewing with two transverse postmedial bands straight or nearly so; thin
line in whitish patch adjacent to blackish terminal dots somewhat sinouus
and weakly denticulate; distribution Argentina (pl. 1: fig. 3).
saltalis, new species
Forewing with two transverse postmedial bands usually sinuate or bent
inwardly; thin line in whitish patch adjacent to blackish terminal dots
rather smooth (pl. 1: fig. 2); distribution otherwise . ocelleus (Haworth)

BasED ON MALE GENITALIA

Harpe conspicuously extenuated and narrowed distally, somewhat finger-like

(fig. 3) . » «+ « es « ~~ Californicalis (Packard)
Harpe Serie Gan 1, 2, Sa Sra es : SaaS
Aedeagus with two elongate patches of anvils ite epinmleg pasa of the

large cornuti (figs. la, 2a) ... ass a 3
Aedeagus with but one elongate patel of pena lender eanles asad of

the large cornuti (fig. 4a) .... . . . . ocelleus (Haworth)

Harpe broad, costal margin strongly ceri ae basal production truncated
distally, apices subequal (fig. 1) .... . . . . Saltalis, new species
Harpe narrower than above, serration of postal margin weak, the basal
production rounded apically, with a single spine adjacent to apex (fig. 2).
galapagosalis, new species

BASED ON FEMALE GENITALIA

With conspicuous hooklike sclerotizations laterad of the ostium (figs.
Bicnk ema be Gf hie bere Braces, aareseeen eae
Without such Rookie eleanor (figs. 6, 3) ois ce re vee
Distance from ostium to origin of ductus seminalis much fen than that from
ductus seminalis to top of bursa copulatrix (fig. 7).
californicalis (Packard)
Distance from ostium to origin of ductus seminalis equal to or greater than
that from ductus seminalis to tip of bursa copulatrix (fig. 5).
saltalis, new species
Ventral margin of ostium rather broad and distinctly undulate; ductus bursae
with some distinct ana ce between ostium and origin of ductus seminalis
GH EAG) ic V.e. otiste . .. . galapagosalis, new species
Ventral margin of esriane narrower When in above, and straight or nearly so;
ductus bursae with conspicuous longitudinal grooves, without Sainniion
(fig. 8) in. aie tenke ae ey ee ERS, en Le euis,. yormienet 6) belie) 1 OCCHEUA Eta worth)

NO. 3551 EUCHROMIUS—CAPPS 3

Euchromius ocelleus (Haworth)
Ficurss 4, 8; PLats 1 (Fia. 2)
Palparia ocellea Haworth, 1811, Lepidoptera Britannica, vol. 3, p. 486.
Eromene texana Robinson, 1870, Ann. Lyc. Nat. Hist. New York, vol. 9, p. 155.

Mats (pl. 1: fig. 2).—Alar expanse 18-24 mm. Forewing: Upper
surface coloration somewhat variable; sordid white, ochreous white
to pale brown, with brownish or fuscous irroration distinct; two
ochreous transverse postmedial bands, sinuate or concave inwardly,
margined with silvery white or gray; two short, oblique bars of similar
color adjacent to apex; terminal dots blackish, margined outwardly
with silvery metallic scales; a narrow whitish patch adjacent to ter-
minal dots with a thin brownish line midway between the dots and
termination of the irrorated area. Hind wing subhyaline, sordid
white or with some pale ochreous brown suffusion, without markings,
or with a thin testaceous terminal line.

Genitalia (fig. 4): Uncus rather slender, length about equal to that
from receptacle to tip of the gnathos. Harpe broad; costa serrate,
basal process triangular distally, hook from near base of the process.
Aedeagus with a rather extensive series of short, stout, spinelike
cornuti and basad an elongate patch of small, slender spines.

Frmate.—Alar expanse 17-26 mm. Similar to male in color and
maculation.

Genitalia (fig. 8): Ostium rather narrow, weakly scobinate, ventral
margin straight or nearly so; ductus bursae long, with longitudinal
grooves, spinulation inconspicuous; distance from ostium to origin of
ductus bursae about equal that from ductus seminalis to tip of the
bursa copulatrix; two subequal signa, elongate, scobinate, and ridge-
like.

Larva.—Arrangement of body setae typical of a crambid. Pro-
thorax with two setae on prespiracular shield; group VI bisetose.
Meso- and metathorax with group VI unisetose. Abdominal proleg-
bearing segments (3-6) with seta IV approximate to V, on same
pinaculum and under the spiracle. Ninth abdominal segment with
paired setae II on same pinaculum; setae I approximate to seta III
and on same pinaculum; setae IV and V absent, only seta VI of this
group present. Crochets on abdominal prolegs in a complete ring,
irregularly triordinal in length, weaker outwardly.

Mature larva 18-20 mm. long. Head: Pale amber with some
brownish reticulation; ocellar pigmentation blackish, extending from
ocellus I to ocellus V, posterior margin of pigmentation tangent to
anterior margin of ocellus VI; without fuscous pigmentation at lateral
incision of hind margin of head. Thorax: Prothoracic shield amber,
with some brownish suffusion and a few dark fuscous patches, the
most conspicuous one posterior to and slightly below level of base of

A PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 119

seta Ib, the large one somewhat reniform and the other narrow, with
long axis transverse. Body color sordid white, pinacula amber,
rather large, round or nearly so and moderately sclerotized, except
that on dorsum of ninth abdominal segment which is subrectangular.
Spiracles: Rim black, central area sordid white; on abdominal segments
1-7, round or but slightly oval, small, with diameter but slightly
larger than ring at base of the seta dorsad; on prothorax and abdominal
segment 8, distinctly oval and much larger than on abdominal segments
1-7, three to four times larger. Anal shield broadly rounded, with a
few inconspicuous brownish patches.

Typr.—Male, in British Museum (Nat. Hist.), ocelleus; sex and
location uncertain, tezana.

Typr Locauitins.—Near London, England, ocelleus; Texas, terana.

Foop pLANnt.—Corn and milo maize, on roots.

SPECIMENS EXAMINED. —167.

NEW WORLD DISTRIBUTION.—United States: Tennessee, Alabama,
Texas, New Mexico, Colorado, Utah, Arizona, California, and Wash-
ington. Mexico: Sonora, Sinaloa, Durango, Federal District, and
Vera Cruz. Panama: Corozal. French Guiana: Cayenne.

Remarks.—EL. ocelleus is of wide distribution, occurring in many
parts of the world. The distribution cited herein is for the Western
Hemisphere and is restricted to localities represented by material
studied. For additional synonymy and distribution, see Bleszynski
and Collins (1962).

Heretofore, ocelleus has been associated with stored products and
dead vegetation (Beirne, 1952; Corbet and Tams, 1943; Hinton, 1943;
Meyrick, 1928). The species is not a feeder on such material and
doubtless its association with these was accidental.

Euchromius californicalis (Packard)
Ficures 3, 7; PLATE 1 (Fria. 4)

Eromene californicalis Packard, 1873, Ann. Lye. Nat. Hist. New York, vol. 10,
p. 264.

Mate (pl. 1: fig. 4).—Alar expanse 20-23 mm. Resembling
ocelleus in color and maculation but with the whitish patch adjacent
to blackish terminal dots somewhat broader; and the thin brownish
line within, closer to termination of irrorated area than to the terminal
dots.

Genitalia (fig. 3): Uncus stout. Gnathos with lateral, flaplike
production at base and two conspicuous, dorsal, toothlike productions
at receptacle. Harpe much narrower distally, fingerlike; process from
near base of costa hooklike. Aedeagus (fig. 3a) slender, with a few
coarse distal cornuti and an elongate patch of weaker spinules some-
what basad.

No. 3551 EUCHROMIUS—CAPPS 5

Frema.Le.—Alar expanse 14-22 mm. Similar to male in color and
maculation.

Genitalia (fig. 7): Hookhke sclerotization laterad of ostium greatly
expanded basally. Ductus bursae short; distance from ostium to
origin of ductus seminalis about one-third that from ductus seminalis
to tip of bursa copulatrix. Two subequal signa, narrow, ridgelike.

Lectrotypr.—Male, in collection of Museum of Comparative
Zoology, MCZ type no. 14297, present designation.

TypE LocaLity.—California.

Foop pLant.—Unknown.

SPECIMENS EXAMINED.—54.

DistriputTion.—California, Washington, Idaho, Montana, Utah,
and Colorado.

Remarks.—E. californicalis was based on three males. Only two
of the syntypes were located, both at the Museum of Comparative
Zoology. One is without an abdomen; that with the abdomen is
made lectotype of the species. The genitalia of the lectotype have
been examined by brushing.

The illustration of Corbet and Tams (1943) of californicalis female
genitalia does not agree with those of specimens in the U.S. National
Museum. Their figure is definitely not of californicalis but of an
abnormal specimen of ocelleus or of a species unknown to me.

Euchromius galapagosalis, new species
Ficurss 2, 6; Puats 1 (ria. 1)

Eromene ocellea (Haworth).—Schaus, 1923, Zool. Sci. Zoologica, vol. 5, no. 2,
p. 29.

Maue.—Alar expanse 16-21 mm. Resembling ocelleus, but differ-
ing from it as follows: Facies somewhat darker. Upper surface of
forewing smoother in appearance, the whitish ground color heavily
overlaid with brown from base of wing to inner transverse postmedial
band and with little or no irroration; irroration beyond outer trans-
verse postmedial band more blackish and denser; the ochreous trans-
verse postmedial bands straight or bent outwardly. Hind wing with
subterminal line brownish and more diffuse.

Gentialia (fig. 2): Uncus moderately stout, somewhat dilated
between base and tip. Harpe with costal serration weak; basal
production of costa rounded at apex, the hooklike spine from below
apex. Aedeagus (fig. 2a) with an elongate series of short, stout
spines and two basal patches of small, slender spinules.

FEMALE (pl. 1: fig. 1).—Alar expanse 16-22 mm. Similar to male
in color and maculation.

Genitalia (fig. 6): Ventral margin of ostium rather broad, undulate,
strongly sclerotized, and finely spinulate. Ductus bursae long,

6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

longitudinal grooves weaker than in ocelleus but spinulation stronger.
Two subequal signa, narrow, elongate, ridgelike.

Typre.—Female, in collection of the U.S. National Museum,
USNM type no. 67634, genitalia slide HWC 12,115.

TYPE LocALITYy.—South Seymour, Galdpagos Islands.

PaRatTyPes.—Galipagos Islands: Isla Santa Cruz, Academy Bay,
Darwin Research Station 13407, 1782 (Jan. 22 to Feb. 27, 1964;
R.O. Schuster and D.Q. Cavagnaro); Isla Santa Cruz, Bella Vista
Trail 2° (Feb. 11, 1964; D.Q. Cavagnaro); Isla Pinzon, Summit
and Upper Calderas Areas 1007, 39 (Feb. 7, 1964; D.Q. Cavagnaro).
Paratypes in collections of the U.S. National Museum and the
California Academy of Sciences.

Foop pLrant.—Unknown.

Remarks.—I am greatly indebted to Dr. C. Don MacNeill, Assist-
ant Curator of Insects, Department of Entomology, California Acad-
emy of Sciences, for making available for study the large series
collected by their representatives on a recent expedition to the
Galipagos.

Efforts to locate the remainder of the series examined by Dr. Schaus
were unsuccessful; however, in view of the absence of ocelleus in the
extensive series of the California Academy of Sciences, it appears
that ocelleus does not occur in the Galapagos and that galapagosalis
is an endemic species, not occurring elsewhere.

Euchromius saltalis, new species

Ficorss 1, 5; PLats 1 (Fa. 3)

Mate (pl. 1: fig. 3)—Resembling californicalis in color and
maculation but forewing more cream colored from base to inner
transverse postmedial band; and whitish patch, adjacent to terminal
blackish dots, with the thin line more ochreous and located midway
between the dots and termination of the irrorated area.

Genitalia (fig. 1): Uncus moderately stout. Harpe with costa
strongly serrate, basal production rather broad, straplike and truncate
distally with apices subequal, one strong, hooklike and the other
minute. Aedeagus (fig. 1a) with a series of short, stout spines and
two elongate patches of slender spinules.

Fremautz.—Alar expanse 20 mm. Similar to male in color and
maculation.

Genitalia (fig. 5): Ostium with two lateral adzlike sclerotizations;
ductus bursae long, somewhat longitudinally rugose and spinulation
within rather coarse and conspicuous; origin of ductus seminalis
about midway between ostium and tip of bursa copulatrix, two
subequal signa, narrow, scobinate, ridgelike.

NO. 3551 EUCHROMIUS—CAPPS Pl

Typr.—Male, in collection of the U.S. National Museum, USNM
type no. 67635, genitalia slide HWC 12,110.

TYPE LOCALITY.—Salta, Argentina.

PARATYPE.—Female, in the U.S. National Museum.

Foop pLraAnt.—Unknown.

Literature Cited

BEIRNE, B. P.
1952. British pyralid and plume moths, 208 pp.
BLESZzYNSKI, S., and Couuins, R.
1962. A short catalogue of the world species of the family Crambidae
(Lepidoptera). Acta Zool. Cracoviensia, vol. 7, pp. 197-389.
Corset, A. S., and Tams, W. H. T.
1943. Keys for the identification of the Lepidoptera infesting stored food
products. Proc. Zool. Soc. London, vol. 113, pp. 55-148.
Hinton, W. E.
1943. The larvae of the Lepidoptera associated with stored products.
Bull. Ent. Res., vol. 34, pp. 163-244.
Meyrick, E.
1928. A revised handbook of the British Lepidoptera, 914 pp.

8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119 |

Ficures 1-4.—Euchromius, male genitalia: 1, 1a, saltalis, new species; 2, 2a, galapagosalts,
new species; 3, 3a, californicalis (Packard); 4, 4a, ocelleus (Haworth).

EUCHROMIUS—CAPPS

NO. 3551

SM

Ficures 5-8.—Euchromius, female genitalia: 5, saltalis, new species; 6, galapagosalis,

new species; 7, californicalis (Packard); 8, ocelleus (Haworth).
U.S. GOVERNMENT PRINTING OFFICE:1966

CAPPS—PLATE

PROC. U.S. NAT. MUS. VOL. 119

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Proceedings of

the United States

National Museum

SMITHSONIAN INSTITUTION + WASHINGTON, D.C.

Volume 119 1967 Number 3552

PRELIMINARY REVISION OF BUTTERFLIES
OF THE GENUS CALYCOPIS SCUDDER

(LYCAENIDAE: THECLINAE)

By Wii.1am D. FreLp

Associate Curator, Department of Entomology

The genus of butterflies treated in this paper contains 21 species,
of which 6 are here described as new. Nineteen of these species are
neotropical in distribution; one species is entirely nearctic and one is
both nearctic and neotropical.

In Calycopis the forewings of the males lack scent pads. Upper
surfaces of hindwings have some blue or purplish scales and sometimes
these wings are almost entirely blue or purplish. Hindwings with
tails at end of veins Cu, and Cuz, the tail at end of Cuz being much the
longest. Undersurface of hindwing with a submarginal ocellate spot
below vein Cu, and sometimes with one or more of these spots above
this vein. The area between the large blue lunule in interspace Cuz
and the postmedian line is always darker than the ground color and is
usually some shade of red.

In the male genitalia the uncus is divided dorsally into two lateral
elements, often completely separated. Aedeagus usually with a
single terminal, spinelike cornutus, very rarely with a small second

1

2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

terminal thornlike cornutus. The aedeagus lacks the ventral keel
found in some Theclinae and is sometimes upturned distally. Harpes
fused for more than one-half of their lengths from base and terminally
truncate, or if bluntly pointed, with ends slightly upturned. Forearm
of gnathos often with carina along inner surface, this carina sometimes
laminate or dentate.

Female genitalia with ductus bursae a simple, lightly sclerotized
tube, funnel-shaped at ostium bursae and with rounded or sharply
pointed lobes above and posterior to ostium bursae. Bursa copulatrix
with two large platelike signae, each ornamented with a centrally
placed, rose-thorn shaped spine and with anterior margin deeply
dentate.

On the basis of male genitalia characters the species of Calycopis
are divided into five species groups. ‘These groups are included in
the key and are also separately treated in the body of this paper.

The figures were drawn by the author except figures 1, 4, 6, 9, 10,
and 17, which were drawn by Mrs. Amy Awl. All photographs were
made by Mr. Jack Scott, staff photographer, and are reproduced here
(figs. 35-126) twice natural size.

Key to the Species of Calycopis

(Unless otherwise stated, all comparisons in this key pertain to the male genitalia
in lateral view and to the female genitalia in ventral view.)

MALES
ie Gnathos extending well beyond lower margin of lateral lobe of uncus;
lower distal angle of this lobe not greatly expanded (figs. 1-9) . . 2

Gnathos not extending beyond lower margin of lateral lobe of uncus;

lower distal angle of this lobe aa pct a (figs. 10-19),

partunda group) .. s SEE IGE

2. (1) Lateral lobe of uncus ats ae ee concave or straight: ventral
margin of uncus concave or straight in front of lower distal angle

(figs: 1-8) asa 3 ce es 5 Sak Uap hte taeengtes

Lateral lobe of uncus mite ciate erent convex; vectra margin of

uncus convex in front of lower distal angle (fig. 9), (cecrops

(AOD) oo oo. e Syitr. eto

3. (2) Fused harpes wide in entra view, SSE a nid cal is More than
two-thirds their length; distal end of aedeagus distinctly bent

Whonranel (Girass Tots) 6 "4! oan e ah Ro iss ee re oe EE

Fused harpes more narrow in Ventral view, Hea a width that is not

more than two-thirds their length, usually much less; aedeagus

straight or only ae upcurved distally (figs. 1-6), (janetrica

group) .. 5

4 (8) Aedeagus in oral view enero peat eee to the right Aad bifur-
cate; gnathos with dentate carina (fig. 7), (torqueor group).

C. torqueor

Aedeagus in dorsal view not as above; carina laminate (fig. 8),

(xenela group) 20 oe ee OK eG Aloe Coin chee ce Re ee OES

NO.

10.

11.

14.

3552

(3)

(5)

(6)

(7)

(7)

(2)

(1)

~ (10)

. (12)

(11)

(14)

GENUS CALYCOPIS—FIELD 3

Upper distal angle of lateral lobe of uncus greatly produced, forming a
fingerlike process; basal end of aedeagus upturned (fig. 6). C. lerbela
Upper distal angle of lateral lobe of uncus not so greatly produced,
without fingerlike process; basal end of aedeagus slightly down-

turned or straight, not upturned (figs. 1-5)... ....2.2.2.2. 6
Harpes gradually tapered, forming long narrow apices; distal margin
of lateral lobe of uncus straight or nearly so (fig. 1) . . . C. bactra
Harpes constricted before but expanded at apices; distal margin of
lateral lobes of uncus concave (figs; 2-5) . . 2. 1... 4... 1 0
Aedeagus with two terminal cornuti; harpes fused for more than
three-fourths their length (figs. 4, on aorta) fe CAE ELS
Aedeagus with a single terminal cornutus; free ficed less than
three-fourths their length (figs. 2,3) ...... Rae A ao

Upperside of hindwing with a large area of blue er a much more
narrow fuscus border; upper distal a of lateral lobe of uncus
slightly produced (fig. 4c) . . . . 3 \ . . . C. janeirica

Upperside of hindwing with a eealier! area of aalue and a broader
border of fuscus; upper distal angle of lateral lobe of uncus more
greatly ode (Ass De) see. Pies als eae, | “Cl chacona

Highth tergite of male abdomen with posterior concavity broader and
proportionately more shallow and not grooved mesially (fig. 20b);
blue disc on upperside of hindwing large and lustrous (fig. 114).

C. nicolayi

Highth tergite of male abdomen with posterior concavity narrow and
deep and distinctly grooved mesially (fig. 20c); blue disc on upper-
side of hindwing smaller and not so lustrous (fig. 115). C. caulonia

Hindwing beneath having marginal ocellate spot in interspace Cu
with black center much larger than the red or gray lunule that
partially surroundsit ..... - .. . . C. cecrops

Hindwing beneath having mereiel pcelintel spot in interspace Cu;
with black center smaller than surrounding red lunule . C. isobeon

Harpes fused for three-fourths or less their lengths (figs. 10, 11,

May itt le se omamunelie
Harpes fused ion more than) i fourth cient jeneiasl (ape 12, 18,
15-19) . sh : meters ate einy Jain ome

Lower margin ae lateral fake of uncus nearly ene sliphtty concave
anteriorly; forearm of gnathos without carina (fig. 10) . C. susanna
Lower margin of lateral lobe of uncus undulate; forearm of gnathos

wit laminate carina (figs. EIs TA) oo. ke cy sk Se Smells
Saccus longer than harpes; aedeagus straight; harpes fused much less
than three-fourths their length (fig. 11)... . .. . C. drusilla
Saccus not as long as harpes; aedeagus upeurved; harpes fused nearly
three-fourths their length (fig. 4)........ ... C. bellera
Upper distal angle of lateral lobe of uncus greatly produced, forming
an acute fingerlike process (figs. 16-19)... . CN) si ietitorae Lit
Upper distal angle of lateral lobe of uncus slightly nrodteed but not as
an acute fingerlike process (figs, 12-15) . 2 2. eae + 2 es 8 2S

Forearm of gnathos without carina; dorsal surface of tegumen greatly
uplifted; distal end of harpe bluntly rounded (fig. 15) . C. partunda
Forearm of gnathos with carina; dorsal surface of tegumen smoothly
rounded; distal end of harpe slightly upturned (figs. 12, 18) . . 16

16.

17.

18.

19.

(15)

(14)

(17)

(17)

(1)
(2)

(3)

(2)

(1)

PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

Forearm of gnathos with sharply pointed dentate carina near apex; |
aedeagus nearly straight; distal margin of lateral lobe of uncus)
straight or slightly convex (fig. 18) ........ .. C. vitruvia))

Forearm of gnathos with dentate carina not sharply pointed and near)
middle in position; aedeagus in dorsal view sharply bent to the!
right; distal margin of lateral lobe of uncus slightly concave |
(fle Mare see ls . . . . C, vibulena}

Finger-like process at ‘distal eed ‘of Been aivecen upward Com
18, 19). eoie 18 |

Finger-like process a distal end oe pedeaeus pean cn 16, 17) . 19 |

Lateral lobes of uncus connected dorsally; upper distal angle of lateral |
lobe of uncus greatly produced; forearm of gnathos without carina;)
aedeagus very ee more than four times the length of harpe
(fig. 19) . ems : ... . C. fractunday

Lateral lobes. of uncus dentro pene upper distal angle of lateral \\7
lobe of uncus produced less than one-half as much as above; forearm ||
of gnathos with carina; aedeagus shorter than above, less en four
times the length of harpe (fig. 18)... ... .. C. anastasia)

Lateral lobes of uncus narrowly connected dorsally; aedeagus with two (
cornuti, the larger one straight; dorsal margin of harpe greatly |)
constricted before apex (fig. 17). ......... C. anfractm

Lateral lobes of uncus entirely separate; aedeagus with a single large |
cornuti which is bent near middle; dorsal margin of harpe not as
greatly constricted before apex as in species above (fig. 16). C. indigo:

FEMALES
Length of ductus bursae plus bursa copulatrix less than ng times the
width of ostium bursae (figs. 28, 81-33) ..... sd
Length of ductus bursae plus bursa copulatrix 4 or more times the |
width of ostium bursae (figs. 22-27, 29, 30, 34) . vee on
Posterior ostium bursae lobes sharply pointed. ........ 8)
Posterior ostium bursae lobes rounded. ... . a) |

Posterior ostium bursae lobes large and pase length an ductus

Jonacerey (Gres SS) 5 4 6 6 5 ae
Posterior ostium’ bursae lobes nel aoe eae conan two teeth; |
length of dultus bursae plus bursa copulatrix about 3 times the |
width of ostium bursae (fig. 32)... .. ... . C. ampliagg
Hindwings beneath with reddistinct, not seed with fuscous, par- —
ticularly in the area between the large blue lunule in interspace }
Cu, and the postmedian line and with a submarginal red spot
above vein Cu,..... . . . C. anfracta
Hindwings beneath with red pace ppeemeed ae fuscous, partic- |
ularly in the area between the large blue lunule in interspace Cu,
and the postmedian line and with a submarginal red spot lacking |
above vein Cu,;..... ... . . C.indigas
Ventral plate of ductus purses ke pea ise broadly rounded |’
laterally (fig. 28) . wx ... . . C. drusilla@
Ventral plate of Baccus ee at agin pares sharply angled, not |
rounded laterally (fig. 31) . ae . . C. partunda |
Forewing above with large enmee area nie anoe below cell. . 7 |
Forewing above without this spot, or if present it is pale blue in color,
or forewing dark metallic blue over most of wing except outer
matgin . WUC gh ah a A ee ee a

eS ee

|
|

|.NO. 3552 GENUS CALYCOPIS—FIELD 5
7. (6) MHindwing beneath with postmedian black line above vein Cu, bordered
outwardly with white and inwardly with red; spurs of posterior

ostium bursae lobes set close together (fig. 29). . . . C. vitruvia

Hindwing beneath with postmedian black line lacking red inner

border; spurs of posterior ostium bursae lobes set further apart

}

than in wiruwvia. .. . 3.) Sig) 88 CY fractunda
' 8. (6) Forewing dark metallic Blac) peer alone oaies margin. . C. bellera
Forewing not as above, dark brown sometimes with bright blue below
2dA and sometimes pale blue over cell and base of wing. . . 9
/ 9. (8) Total length of ductus bursae and bursa copulatrix 5 times the width
of ostium bursae. .. . PGR a aeLO
! Total length of ductus honnene aa pares copileientes jee than 5 times
} the width of ostium bursae... . Se dered
110. (9) Hindwing above with a wide brown border nt site. blue area more
restricted (ie. 49) 0. 2 5 . i. ws GC. Chacona
Hindwing above with brown pore: more narrow and with blue area
Hangers (tore 5) een et en te eh ee Ex ganeirice
'11. (9) Total length of ductus bursae oad Buen Copnlaie 4144 mm. or slightly
| THIOLE Ween 1 Sh felon coe pales

Total length of deine bareee ara aa ponaleae 3 a mm.orless . 13

12. (11) Inner red element of postmedian line on hindwing beneath broad,
: nearly 1 mm.in width. .... Je), sue cecrops
Inner red element of postmedian oe on Pind yin beneath not nearly
as broad, about 4 mm. in width. . . . . C. isobeon, C. susanna

13. (11) Posterior ostium bursae lobes sharply pointed. ........ 14
Posterior ostium bursae lobes rounded. ..... . . C. lerbela
14, (13) Ductus bursae sharply bent ventrally just before attachment of ductus
seminalis; area around the opening of ductus seminalis nonsclerotized

| C. bactra
| Ductus bursae only gradually bent toward the left just before attach-
: ment of ductus seminalis; area around opening of ductus seminalis
| Releto tized np suis wits By co ey cds eat irs tei eS poqaes ue meCaULOMa

The janeirica Group

Tn this group the lower angle of uncus in lateral view is only slightly
produced and much less so than the upper angle of uncus, (the ventral
“Margin is not nearly as long as the distal or caudal margin). The
aedeagus is fairly straight, ending distally in a straight or upward
projected point. The fused harpes in ventral view are not nearly as
wide as long.

The species in this group are all very closely related and most of
them are usually more easily distinguished upon wing habitus and color
characters, than by genitalic characters. In the male sex, C. bactra
is distinguished at once from all the others in having a wider post-
median band on the wings underneath and in having the cell bar closer
to it, being separated from it by less than twice the width of the
band. C. lerbela is the only species in this group that has two black
pupilated lunular spots on the outer margin of hindwing below. Of
the remaining four species, C. nicolay? is the most distinct in having the

6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119 |}

blue color on the upperside of hindwing more lustrous and covering a/
larger area, and in having a dark gray ground color below. The re-|
maining three species: caulonia, janeirica, and chacona are most
closely related. They can be distinguished except for a few genital |
characters, only by color characters on the upper surfaces of the hind- |
wings. C. chacona has a broad black outer border that completely
obliterates the marginal black spots and consequently has a much
smaller discal blue area; it has a marginal bluish white line in inter-.
space Cup only. C. janeirica has the outer border narrower, containing |
two large marginal black spots, nearly confluent but outlined by a |
trace of blue. Because of the narrow border there is a larger blue disc '
in janeirica. This species also has a marginal bluish-white line extend- |
ing above vein Cu;. C. caulonia has the marginal spots much smaller |
and more isolated than in janeiwica, with marginal bluish-white line |
as in janeirica.

Calycopis bactra (Hewitson), new combination
Figures 1, 20a, 22, 35-42

Thecla bactra Hewitson, 1877, Illustrations of diurnal Lepidoptera: Lycaenidae,
vol. 1, p. 194; vol. 2, pl. 77, figs. 619, 620.

Thecla beon (Cramer).—Godman and Salvin [not Cramer, a misidentification,
in part], 1887, Biologia Centrali-Americana, Lepidoptera-Rhopalocera,
vol. 2, pp. 74, 75.

Mate (figs. 35, 36).—Forewing above dark brown, nearly black,
sometimes with a faint and dull bluish-purple irridescence over base;
fringe paler brown. Hindwing above with costal and outer margin |
dark brown, nearly black; base and cell to outer dark border lustrous |
blue with a slight purplish tinge; abdominal margin below vein 2dA |
thickly covered in middle with pale grayish-purple scales; lobe of |
anal angle with a small orange-red spot, surrounded by the dark |
ground color; immediately above this on abdominal margin a small
nearly triangular white spot; above anal angle along outer margin a
pale bluish-white submarginal line between veins 2dA and M;; tips
of tails white; fringe at base of tail Cu, white. Undersides of both
wings with ground color pale brown; with two very faint parallel
bars slightly darker than ground color crossing the end of each dis-
coidal cell, area between these bars dirty white and sometimes con-
taining a few scattered red scales; fringe slightly darker than ground
color and on hindwing containing a few bluish-white scales below
vein Cu,. Forewing above vein Cuz and hindwing with a narrow
tricolored postmedian line or band; this band white along outside,
black in middle and orange red along its inner side. Below vein Cu;
this band has a W-shape; the V-shaped bar, present in some species
of Calycopis in interspace M; and on vein Cu, is absent or only slightly

NO. 3552 GENUS CALYCOPIS—FIELD 7

indicated in C. bactra. Forewing underneath with costal margin
orange red; outer margin slightly darker than ground color; a faint
double row of slightly darker submarginal bars along outer margin
above vein Cuz. Hindwing beneath with anal lobe black; immedi-
ately inward from this on abdominal margin an orange-red bar; a
few white scales between bar and anal lobe; a black or brown line
extending from anal lobe around outer margin, becoming lighter
brown near costal margin; inward from and adjacent to this line a
white line fading into pale brown above vein M;; interspace Cu, with
a submarginal orange-red lunular marking, partially enclosing a small
black spot along its outer edge; between the lunular marking and the
anal lobe a submarginal dark brown lunular spot suffused with bluish-
white scales; inward from this lunular spot and between middle
elements of W-shaped portion of postmedian band an orange-red
lunule; a double row of very faint and irregularly-shaped dirty white
submarginal spots above vein Cu.

Length of forewing 11-12.5 mm.

Male genitalia, as illustrated by figure 1, with aedeagus short,
under 3 mm. in length, nearly straight, slightly upturned distally and
with base slightly downturned; lateral lobe of uncus with distal
margin nearly straight and lower distal angle not greatly expanded,
lower margin of uncus concave; harpes fused ventrally nearly seven-
eights of their length, in lateral view gradually tapering and forming
along narrow apex. LHighth tergite (fig. 20a) with posterior concavity
broad and shallow.

FrEmMat.s (figs. 37-42).—Wings on both surfaces much like the male,
but frequently with some pale blue in base of forewing above and
with blue of hindwing above much less brilliant; differing on the
hindwing underneath in usually having a little orange red above Cu,
next to the submarginal black pupillated lunule.

Length of forewing 11-12 mm.

Female genitalia, as illustrated by figure 22, with ductus bursae
and bursa copulatrix relatively short, being about 3.5 mm. in length
and just over 4 times the width of ostium bursae, posterior ostium
bursae lobes sharply pointed.

Typr-Locaity.—‘‘Nicaragua.”

ADDITIONAL TYPE DATA.—C. bactra was originally described from a
single female specimen, the holotype, from the collection of W. C.
Hewitson, now in the collection of the British Museum (Natural
History) and labeled type number Rh. 915 with female genitalia
preparation no. R. 1948-14 (N.H.B.).

DisTRIBUTION.—Guatemala south through Central America, north-
ern South America, Trinidad, northern Brazil and Peru.

8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

GUATEMALA: Department of Quezaltenango, Voledn Santa Maria. HonpurRas:
Department of Cortés, La Cumbre. Nuicaracua: no specific locality. PANAMA:
Province of Panama, La Chorrera (May). Canau Zone: Culebra (Nov.); Ancén
(May, June, October); New Culebra (February); Balboa (April, May); Taboga
Island; La Boca (April); Barrow Colorado Island (March); La Cruces Trail, nr.
Corozal (June). CotomBia: Department of Valle del Cauca, Cali District (3200
ft., February). Pru: no specific locality. WaNnrzvuELA: State of Carabobo,
Puerto Cabello. Trinipap: St. Andrew County, Matura (September), Oropuche
(April); Coroni County, Gran Couva (April). Surinam: District of Paramaribo,
Paramaribo (March); District ?, Sint [sic] Barbara Plan [sic] (Surinam River,
April). FrencH Gur1ana: Colonie de la Guyana, Cayenne, Mouth of Kourou
River (July, December). Brazriu: Territory of Rio Branco, Mt. Roraima (4200
ft., December).

MATERIAL STUDIED.—EKighteen males, 20 females, including their
genitalia.

Calycopis nicolayi, new species
Fiaures 2, 20b, 113, 114

Mats (figs. 113, 114).—This species differs from all others in the
janeirica group in having ground color of forewings and costal and
outer borders of hindwings very much darker and purplish black in
color. It is similar to bactra and thus differs from caulonia, janeirica,
and chacona in having a very large blue disc on hindwing. This blue
as in bactra is more lustrous than in the above-mentioned species and
it lacks the purplish tinge found in bactra. The ground color under-
neath is very dark gray, not brown as in the other species. The
markings below are as in caulonia and janeirica with postmedian red
lunule of interspace Cup slightly suffused with black and sometimes
almost entirely black.

Length of forewing 10-12 mm.

Male genitalia, as illustrated by figure 2, with aedeagus about the
length and shape of that found in bactra, somewhat more slender and
less upturned at distal end; lateral lobe of uncus as in janeirica; harpes
fused ventrally about as much as in caulonia, in vental view shaped
about like that species and bactra, but in lateral view shaped like
caulonia and janeirica. Eighth tergite (fig. 206) with posterior con-
cavity broader and proportionately more shallow than in any of the
other species except chacona.

Frmate.—Unknown.

TypE-LocaLiry.—Joao Pessoa, State of Paraiba, Brazil.

ADDITIONAL TYPE DATA.—Described from the holotype, male (lo-
cality as given above; June 29, 1953; USNM type 34897; ex. S. S.
Nicolay Collection; collected by Jorge Kesselring; male genitalia
preparation WDF 5291, 1953), and two male paratypes from the same
locality (taken June 21 and June 27, 1953).

NO. 3552 GENUS CALYCOPIS—FIELD 9

Location oF TypES.—Holotype in United States National Museum,
paratypes in the collection of Lt. Colonel Stanley S. Nicolay.

DistrisutTion.—Known only from the type-locality.

MATERIAL stupDIED.—Three males including their genitalia.

Calycopis caulonia (Hewitson), new combination
Fieurss 3, 20c, 23, 55, 56, 115, 116

Thecla caulonia Hewitson, 1877, Illustrations of diurnal Lepidoptera: Lycaenidae,
vol. 1, p. 188; vol. 2, pl. 75, figs. (9) 587, 588.

Matz (figs. 115, 116).—Very similar to bactra except that post-
median band on undersurfaces of wings contains less red along inner
side and the bars at the end of the cell in both wings on this surface
are separated by a greater distance from the postmedian bands. Very
close to janeirica in this sex, hardly differing except in size.

Length of forewing 10-12 mm.

Male genitalia, as illustrated by figure 3, with aedeagus short as in
bactra, being about 3 mm. in length, straight, not upturned distally
and with distal end not prolonged as it is in bactra and not blunt as
it is In janeirica; harpes and uncus about like janeirica but smaller.
Eighth tergite (fig. 20c) similar to janeirica but with posterior con-
cavity more narrow and proportionately deeper.

Frmatre.—Close to females of bactra, differing chiefly in having the
blue on hindwing more restricted, being nearly absent in interspaces
M, and Mz opposite the end of cell. Wings below as in male with
slightly less red along inner side of postmedian band and with cell bars
further from this band than in bactra.

Length of forewing 10.5-11.5 mm.

Female genitalia, as illustrated by figure 23, with posterior ostium
bursae lobes smaller and with spurs much closer together and nearer
center of upper margin of ostium bursae than in bactra; with ductus
bursae curving to left before entrance of ductus seminalis and with
sclerotized area around opening of ductus seminalis (in bactra the area
is nonsclerotized and the ductus bursae strongly and abruptly curved
ventrally before nonsclerotized area).

TyYpPE-LocALIry.—‘‘Rio Janeiro” [Rio de Janeiro, Brazil].

ADDITIONAL TYPE DATA.—Caulonia was originally described from a
single female specimen, the holotype, W. C. Hewitson collection, now
in the collection of the British Museum (Natural History) and is
labeled type number Rh. 916. According to Mr. D. 8S. Fletcher of
the staff of that institution, this type lacks its abdomen. Another
specimen, a topotype, was carefully compared with the holotype by
Mr. Fletcher and upon being identified as that species was loaned to
me for study. This specimen is the one illustrated in figures 23, 55, 56.

10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Distrisution.—This species is found along the central eastern
coast of Brazil from the State of Rio de Janeiro north to the state
of Paraiba.

Braziu: State of Rio de Janeiro, Rio de Janeiro; State of Pernambuco, Recife
(January), Santa Cruz (July), Bonita (January); State of Paraiba, Joio Pessoa
(December, January, February).

MATERIAL STUDIED.—Nine males and nine females including their
genitalia.

Calycopis janeirica (C. Felder), new combination
Fiaures 4, 20d, 24, 43-46

“‘Thecla beon Cram. var. janeirica’’ C. Felder, 1862, Verh. Kais.-K6énigl. Zool.-
bot. Ges. Wien, vol. 12, p. 474.

Thecla vibulena Hewitson [in part], 1877, Illustrations of diurnal Lepidoptera:
Lycaenidae, vol. 2, figs. 599, 600 (only).

Mate (figs. 43, 44).—Similar to bactra, differing in being slightly
larger and in having dark borders of hindwings wider on the upper
surface, especially so near apex. Forewing below differing from bactra
in lacking the red along inner side of postmedian band. Hindwing
below with less red along innerside of this band and with elements of
this band forming three distinct, connected V-shaped bars below
vein M3.

Length of forewing 11-14 mm.

Male genitalia, as illustrated by figure 4, with aedeagus longer than
in bactra or caulonia, usually nearly 4 mm. in length and nearly
straight, the distal end similar to cawlonia, not upturned as in bactra;
lateral lobe of uncus with distal margin concave and lower distal angle
not greatly expanded but slightly more so than in bactra or caulonia;
lower margin concave. Harpes fused ventrally just barely over three-
fourths of their length; in lateral view with apices slightly expanded,
not long and tapering. Eighth tergite (fig. 20d) with posterior
concavity narrow and much deeper than in bactra but not as narrow
as in caulonia.

FEMALE (figs. 45, 46).—Similar to the male, forewing above slightly
bluish at the base. Hindwing above with border below vein Cu,
more broken, with dark spots more isolated and with blue over disc
not quite as brilliant. Wings underneath not different from those
of the male and differing from bactra in the same ways that the male
does.

Length of forewing 11-13 mm.

Female genitalia, as illustrated by figure 24, with ductus bursae
and bursa copulatrix long, being about 5 mm. in length and just over
5 times the width of ostium bursae, with the pair of sharply pointed
ostium bursae lobes similar to those of caulonia, but slightly larger,

)

|

NO. 3552 GENUS CALYCOPIS—FIELD 1a

ductus bursae only slightly curved, or if greatly so, this curve near
middle.

TyprE-LocaLiry.—‘‘Rio” [Rio de Janeiro, Brazil].

ADDITIONAL TYPE DATA.—This species was described from the female
sex and the number of specimens in the original series was not indi-
cated. Although the C. Felder collection of Lepidoptera supposedly
went to the British Museum (Natural History), the type of this species
could not be located in that collection by Mr. D.S. Fletcher. I am
not suggesting that a neotype be designated at present because of the
possibility of the type being in some other European collection.

Disrrisution.—Widely distributed in Brazil from Amazonas and
Pernambuco south to Santa Catarina.

Braziu: State of Amazonas, Manaos; State of Pernambuco, Espirito Santo,
Recife; State of Baia, Tijuca; State of Rio de Janeiro, Petropolis, Rio de Janeiro,
Nova Friburgo; State of Santa Catarina, Joinville (April, August, September).

MATERIAL STUDIED.—Twelve males and 10 females, including
their genitalia.

Calycopis chacona (Jérgensen), new combination
Ficures 5, 20e, 21, 21h-7, 25, 47-52
Thecla poeas chacona Jorgensen, 1932, Deutsche Ent. Zeit. (Iris), vol. 46, pp. 44, 45.

Mats (figs. 47-48, 51-52).—Closely allied to janeirica, differing in
having the brilliant blue of hindwing covering a much smaller area and
thus leaving a very much broader border on this wing. Differing from
janeirica on the undersurfaces in having the ground color a warmer
brown and with all red markings more of an orange red. The red of
the postmedian band on the forewing below is frequently present
but is very faint.

Length of forewing 11-14 mm.

Male genitalia (figs. 5, 21, 214-7) with uncus, harpes and aedeagus
in lateral view very much as in janeirica; lateral lobes of uncus in
dorsal view further apart and more expanded, forming a deeper middle
notch than in janeirica and harpes fused ventrally for a greater dis-
tance than in that species. Figures 21, 21h~7 show variation of
harpes in lateral view. Eighth tergite (fig. 20e) with posterior
concavity as deep or deeper than in janeirica and much broader.

FEMALE (figs. 49, 50)—Most specimens differ from janeirica in
the same way that the males differ, in having a wider brown border
with a smaller blue disc on the upper surfaces of the hindwings and
in having the red of the postmedian band on the forewing below present
but very faint. In janeirica this red is usually absent. I am unable
to distinguish some female specimens from the state of Parana, Brazil
from janeirica; however, all males I have seen from Parana are typical
chacona.

12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Length of forewing 11-13 mm.

Female genitalia, as illustrated by figure 25, not separable from
janeirica. Ductus bursae usually sharply curved immediately before ©
ductus seminalis. Figures 24 and 24a of janeirica and figure 25 of
chacona were not included to show differences between these two
species. Both species have ductus bursae like figures 24a and 25 and,
although I have not seen chacona like that of figure 24, I do not doubt
that it occurs.

TypE-LocaLiry.—“‘Formosa, Argentina.”

ADDITIONAL TYPE DATA.—This species was originally described
from a series of several specimens (=syntypes), the exact number not
being stated. These syntypes are presumably in the Jérgensen
collection in the Museo Argentino de Ciencias Naturales, Buenos
Aires; however, a lectotype cannot be designated at the present time
because I have been unable to obtain any information concerning these
syntypes. My identification is based upon the original description,
upon a topotype specimen and upon this being the southernmost
species in the genus.

DistRIBuTION.—This species is found in the state of Parana in
Brazil west and south through Paraguay to the provinces of Salta,
Cordoba and Buenos Aires in Argentina.

Braziu: State of Parana, Castro, Londrina (April, September). Paraauay:
Villarica (September). ARGENTINA: Province of Misiones, Iguasu, Santa Maria;
Province of Correntes, Corona near Goya; Province of Entre Rios, La Soledad,
Gualeguaychu (March), Isla Los Cisnes (Parana delta); Province of Buenos Aires,
Islas, Punta Lara, Buenos Aires; Territory of Formosa, Formosa; Province of
Salta, Salta; Province of Tucuman, Tucuman (May); Province of La Rioja, La
Rioja; Province of Cordoba, Cordoba, Cosquin.

MATERIAL STUDIED.—Twenty-six males and 20 females, including
their genitalia.

Calycopis lerbela, new species
Ficurss 6, 20f, 26, 57-60

Mate (figs. 57-58).—Differing from bactra and related species in
having the brilliant blue on hindwings above restricted to lower half
of wing below cell. Anal lobe without red spot or at most with only
a few red scales. Forewing below similar to janeirica, differing from
that species on the hindwing in having a second submarginal black
pupillated red spot, this spot in interspace M; and smaller than
similar spot in interspace Cu,.

Length of forewing 10.5-12.5 mm.

Male genitalia, as illustrated by figure 6, with lateral lobe of uncus
having distal margin concave, with lower margin straight or nearly
so and with upper distal angle produced, forming a short finger-like

NO. 3652 GENUS CALYCOPIS—FIELD 13

process; aedeagus very distinctive, having basal end upturned; harpe
somewhat similar to bactra but slightly more blunt at apex. Highth
tergite (fig. 20f) very much as in chacona and slightly smaller.
FrEmMa.e (figs. 59, 60).—Differing from the male by having a more
extensive blue area in the base of the hindwing on the upper surfaces
and in having the brown border on lower half of this wing more
broken into isolated black spots. Similar to bactra female with blue

somewhat brighter and extending over much of base of forewing.

Wings underneath not differing from those of the male.

Length of forewing 11.5 mm.

Female genitalia, as illustrated by figure 26, with posterior ostium
bursae lobes only slightly produced, rounded and lacking the pair
of spurs usually present and thus very similar to cecrops and some
specimens of susanna. It differs from the latter species in having a
shorter ductus bursae and bursa copulatrix.

TypE-LOcALITy.—Obidos, State of Amazonas, Brazil.

ADDITIONAL TYPE pDATA.—Described from the holotype, male
(locality as given above; USNM type 34898; E. A. Smyth Collection;
male genitalia preparation WDF 2725, 1948); allotype, female (same
locality; Oct—Nov.; M. de Mathan; Rothschild bequest; female
genitalia preparation WDF 3468, 1951) and from ten male paratypes
from Obidos and from other localities in the state of Amazonas,
Brazil.

LocaTIoN oF TypEs.—Holotype and two male paratypes in the
United States National Museum. Allotype and eight male paratypes
in the British Museum (Natural History).

DistRIBUTION.—State of Amazonas in Brazil north into French
Guiana and Surinam. Note: Material from French Guiana and
Surinam was not included in the type series because of the remoteness
of those countries from the type locality.

Braziu: State of Amazonas, Obidos (October-November), Itutuba to Obidos
(January to April), Juhuity (April), Manaos. FreNcH GuraNna: Colonie de la
Guyane, Cayenne. Surinam: District?, Sint [sic] Barbara Plan [sic] (Suriname
River, April).

MATERIAL STUDIED.—Fifteen males and one female, including their
genitalia.

The torqueor Group

In this group the lower angle of uncus is about as in the janetrica
group. The aedeagus is sharply upturned distally and curved to the
right (in dorsal view), the distal portion of the aedeagus bifurcate in
dorsal view, and bluntly pointed in lateral view. The fused harpes
in ventral view are not nearly as wide as long. One species is assigned
here.

14 PROCEEDINGS OF THE NATIONAL MUSEUM voL. 119 |

Calycopis torqueor (Druce), new combination
Ficurss 7, 20g, 111, 112
Thecla torqueor Druce, 1907, Proc. Zool. Soc. London, 1907, p. 608.

Matz (figs. 111, 112).—Wings above similar to wdigo with border |

on hindwing about the same as in that species but with two long

brilliant blue rays on hind margin of forewing, one above and one |

below vein 2dA. Wings below hardly distinguishable from anfracta
and as in that species with red markings more distinct than in indigo

and with red lunule adjacent to postmedian band in interspace |

Cu, containing less fuscous than in anfracta.
Length of forewing 14 mm.

Male genitalia, as illustrated by figure 7, with aedeagus sharply |

bent upward and to the right at distal end; saccus as long as harpe;
vinculum bent greatly to the rear; dorsal margin of tegumen (in
lateral view) greatly produced in the middle; uncus with lateral
elements similar to bactra; the lower margin short, with lower distal
lobe small and distal margin nearly straight; harpes (in lateral view)
of the common type, quite like janeirica, partunda, and several other
species in the genus, with basal two-thirds fused (in ventral view),
unusually broad near middle, narrowing rapidly toward apex and with
free apical one-third widely separated; gnathos with a distinct tooth-
like carina on inner surface of forearm.

Frmate.—Unknown.

Typr-LocaLiry. —“‘Carimang River [sic], British Guiana.”’

ADDITIONAL TYPE DATA.—Described from a series of males all
from the type-locality. Druce designated a holotype as is indicated
in the original description with the statements: ‘“T'ype Mus. Godman,”
and ‘‘Whitely obtained a series of specimens which do not differ.”
The holotype, now in the collection of the British Museum (Natural
History) is labeled type no. Rh 903 and its genitalia preparation
number is 1951-489. Note: One of the paratypes was dissected and
compared with the holotype by Mr. D. S. Fletcher and upon being
found identical was loaned to me for study. This compared specimen
is the one illustrated (figs. 7, 20g, 111, 112).

DistTRIBUTION.—

British GuIANA: Province of Essequibo, Caramang River.

MATERIAL STUDIED.—The only specimen available to me.
The xeneta Group

Uncus as in the janeirica and torqueor groups. The aedeagus is
gradually upcurved distally, ending with a downward directed apex.
One species is assigned here.

NO. 3552 GENUS CALYCOPIS—FIELD 15

Calycopis xeneta (Hewitson), new combination

Ficurss 8, 20h, 107, 108, 117, 118

Thecla xeneta Hewitson, 1877, Illustrations of diurnal Lepidoptera: Lycaenidae,
vol. 1, p. 193; vol. 2, pl. 77, figs. (o*) 611, 612.

Thecla devia Méschler, Schaus [not Méschler, a misidentification], 1920, Ent.
News, vol. 31, p. 176.

Matz (figs. 107, 108, 117, 118).— Both forewing and hindwing with
outer margins more nearly straight than in the other species of the
genus. Forewing more distinctly purplish blue than in any other
species and this blue lighter and more brilliant in color. Hindwing
with a very narrow black outer border. Wings underneath much
darker, being a smoky brown and with red markings very faint or
lacking and red color replaced by dark brown or black. I divide this
species into two subspecies (see below).

Length of forewing 9-14 mm.

Male genitalia, as illustrated by figure 8, with aedeagus distinctly
curved upward before apex and with apex directed downward; both
saccus and harpe in lateral view very short but with saccus over one-
half length of aedeagus; vinculum bent greatly to the rear; elements of
uncus quite similar to bactra and torqueor in having lower margin short
with lower distal angle small and with distal margin nearly straight;
harpes in lateral view blunt at apices, in ventral view very broad near
middle, being almost as broad as long; gnathos with a short ridgelike
carina on inner surface well forward from distal end.

FrmaLe.—Unknown in both subspecies described below.

DistrisuTion.—Guatemala south into Costa Rica and French
Guiana, British Guiana, Brazil, Peru, and Paraguay.

Calycopis xeneta xeneta (Hewitson)
Fiaurss 8, 20h, 107, 108

Mats (figs. 107, 108).—This subspecies differs from C. xeneta devia
(see below) in having the ground color a pale gray on the undersur-
faces, with a dark cloudlike spot in the central area of both forewing
and hindwing and a pale area above and below vein 2dA on forewing.

Length of forewing 12-14 mm.

Male genitalia as described above in the description of the species.

TYPE-LOCALITY.—Department of Chontales, Nicaragua.

ADDITIONAL TYPE DATA.—Described from at least two males as
Hewitson mentions Brazil and Nicaragua for the range of his zeneta.
The male labeled type in the Hewitson collection in the British Mu-
seum (Natural History) is the specimen from Nicaragua. It bears the
labels “BM Type No. Rh. 899,” and genitalia preparation number
“BM 1951-491.” I designate this specimen as the lectotype of xeneta.

16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

A second syntype from Brazil belongs to C. xzeneta devia. A topotypic
male specimen apparently from the type lot was dissected and com-
pared with the lectotype by Mr. D. S. Fletcher and upon being found
identical was sent to me for study. This specimen is the basis for my
identification of xeneta.

DistTRIBUTION.—Guaala south into Costa Rica.

GUATEMALA: Department of Izabal, Cayuga (May). Nicaragua: Depart-

ment of Chontales. Costa Rica: Province of Limén, La Florida (500 ft., March),
Gudpiles (800 ft., May).

MarTERIAL stupieEpD.—Fifteen males, including their genitalia.

Calycopis xeneta devia (Méschler)

Fiaures 117, 118

Thecla devia Méschler, 1883, Verh. Kais.-K6nigl. Zool.-bot. Ges. Wien, vol. 32,
p2 311, spl ly, fie. 7.

Matz (figs. 117, 118).—This subspecies differs from C. xeneta xeneta
as described above. Specimens from Peru have the ground color of
wings below slightly paler than specimens from other areas and have
red markings faintly indicated on hindwings. These specimens are
not here considered distinct enough to warrant separation.

Length of forewing 9-13 mm.

Male genitalia, not illustrated, not differing at all from some speci-
mens of C. xeneta xeneta and differing from other specimens of that
subspecies in having lateral elements of uncus in dorsal view slightly
more produced and farther apart, forming a wider dorsal notch.

TYPE-LOCALITY.—‘‘Surinam.”

ADDITIONAL TYPE DATA.—Described from a single male, the holo-
type, and according to the original description, in the collection of the
“Museum Stuttgart.” I have no additional information on the type
specimen; however, the original figure, original description, and type-
locality indicate that devia is the subspecies of xeneta here described.

DistrIBuTION.—The Guianas south and west through Brazil to
Peru and Paraguay.

FRENcH GuIaANa: Colony of Guyane, Cayene, Penintential Territory, St. Jean
(Maroni River). British Guiana: Province of Essequibo, Kamakusa (Novem-

ber). Braziu: State of Pard, Igarapé-Agu. Prru: Department of Loreto,
Achinamiza (January). Paraguay: Villarica, Sapucai.

MATERIAL STUDIED.—Twelve males, including their genitalia.

The cecrops Group

Uncus in lateral view undulate on ventral margin; aedeagus and
harpes as in the janeirica group. Two species are assigned to this

group.

NO. 3552 GENUS CALYCOPIS—FIELD 7

Calycopis cecrops (Fabricius)
Fiaures 9, 207, 61-70

Hesperia cecrops Fabricius, 1793, Entomologia systematica emendata et
AICTE EVOls os Dts lemp.2 AO:

Rusticus poeas Hubner, 1811, Sammlung exotisher Schmetterlinge, vol. 1, pl.
(101), figs. 1-4.—Scudder, 1876, Bull. Buffalo Soc. Nat. Sci., vol. 3, p. 108.

Strymon beon (Cramer).—Hiibner [not Cramer, a misidentification], 1816, Verze-
ichniss bekannter Schmettlinge [sic], p. 75.

Polyommatus cecrops (Fabricius).—Godart, 1819, in Latreille and Godart, vol 9
in Encyclopedie méthodique, p. 636.

Thecla poeas (Hiibner)—Boisduval and LeConte, 1833, Histoire Générale et
iconographie des Lepidoptéres et des chenilles de ’ Amérique Septentrionale,
pp. 111, 112, pl. 25, figs. 1-4.—Morris, 1860, Catalogue of the described
Lepidoptera of North America, p. 103; 1862, Synopsis of the described
Lepidoptera of North America, p. 103.—Kirby, 1871, Synonymic catalogue
of diurnal Lepidoptera, p. 395.—Edwards, 1877, Trans. American Ent. Soc.,
vol. 6, p. 41; 1884, Trans. American Ent. Soc., vol. 11, p. 298—French,
1886, Butterflies of the eastern United States, pp. 270, 271.

Thecla cecrops (Fabricius)—Westwood, 1857, im Doubleday and Westwood,
Genera of the diurnal Lepidoptera, vol. 2, p. 485.—Kirby, 1871, Synonymic
catalogue of diurnal Lepidoptera, p. 386.—Strecker, 1878, Butterflies and
moths of North America: Complete synonymical catalogue . . . , pp. 86,
87, 185.—Skinner, 1898, Synonymic catalogue of the North American
Rhopalocera, p. 48.—Holland, 1898, Butterfly book, pp. 246, 247, pl. 29,
fig. 18, pl. 30, fig. 7—Draudt, zm Seitz, 1920, Die Gross-Schmetterlinge der
Erde, vol. 5, p. 795, pl. 158, figs. b-1, b-2.—Holland, 1931, Butterfly book,
rev., p. 239, pl. 30, fig. 7, pl. 29, fig. 18.

Tmolus cecrops (Fabricius).—Butler, 1869, Catalogue of diurnal Lepidoptera
described by Fabricius in the collection of the British Museum, p. 189.
Thecla beon (Cramer).—Kirby [not Cramer, as a synonym of 7’. cecrops], 1871,
Synonymic catalogue of diurnal Lepidoptera, p. 395.—Godman and Salvin
{not Cramer], 1887, Biologia Centrali-Americana, Lepidoptera-Rhopalocera,

vol. 2, p. 74.

Calycopis cecrops (Fabricius).—Scudder, 1876, Bull. Buffalo Soc. Nat. Sci., vol.
3, p. 108; 1889, Butterflies of the eastern United States and Canada with
special reference to New England, vol. 3, pp. 1821, 1823.—Dyar, 1903,
Bull. U.S. Nat. Mus., vol. 52, p. 38—Comstock, J. H., and Comstock, A.
B., 1917, How to know the butterflies, p. 231, pl. 34, figs. 10-12 [10=9@ not
o, 11, 12= &].— Field, 1940, Bull. Univ. Kansas, Biol. Ser., vol. 39, no. 10, p.
142; 1940, Bull. Brooklyn Ent. Soc., vol. 35, no. 4, pp. 134, 185; 1941, Journ.
Kansas Ent. Soc., vol. 14, no. 2, pp. 66-68.—Clench, zn Ehrlich, P. R., and
A. H. Ehrlich, 1961, How to know the butterflies, p. 199.

Calycopis beon (Cramer).—Scudder [not Cramer, as a synonym of C. cecrops],
1876, Bull. Buffalo Soc. Nat. Sci., vol. 3, p. 108.

Thecla hugon (Godart).—Godman and Salvin [not Godart, a misidentification],
1887, Biologia Centrali-Americana, Lepidoptera-Rhopalocera, vol. 2, p. 75.

Strymon cecrops (Fabricius —Barnes and McDunnough, 1917, Check list of the
Lepidoptera of Boreal America, p. 14.—Barnes and Benjamin, 1926, Bull.
So. California Acad. Sci., vol. 25, pt. 1, p. 17.—Clark, A. H., 1932, U.S.
Nat. Mus. Bull. 157, p. 249, pl. 23, figs. 3, 4—-McDunnough, 1938, Check
list of the Lepidoptera of Canada and the United States of America, 1:
Macrolepidoptera, p. 24.—Hessel, 1948, Journ. New York Ent. Soc., vol. 56,

219-950—_67——2

18 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

pp. 243-244.— Rawson and Hessel, 1951, Bull. Brooklyn Ent. Soc., vol. 46,
pp. 79-84, 5 figs. [of eggs, larvae, pupae].—Klots, 1951, Field guide to
butterflies of North America east of the Great Plains, pp. 133, 144, plate 15
fig. 12.—Clark, A. H., and L. F. Clark, 1951, Smithsonian Mise. Coll.,
vol. 116, no. 7, pp. 78, 79, pl. 12 h.—dos Passos, 1964, Synonymic list of Ne-
arctic Rhopalocera, p. 56.

Sirymon cecrops ab. gotischalki Clark, A. H., and Clark, L. F., 1938, Proc. Biol.
Soc. Washington, vol. 51, p. 3 (new synonymy); 1951, Smithsonian Misc.
Coll., vol. 116, no. 7, p. 78, frontispiece fig. 7.

Mate (figs. 61-64 [summer form], 67, 68 [spring form]).—Wing®
above usually entirely dark brown, sometimes with a small suffusion
of blue in interspace Cu; and Cuz on hindwing; a pale submarginal
bluish-white line below vein Cuz on hindwing, this line sometimes
extending above through part of interspace Cu,; anal lobe with a
few red scales and a minute white bar at indentation above this lobe
on abdominal margin. Wings below drab or hairbrown in color
with markings similar to the other species of Calycopis; base of costal
margin of forewing orange red, and with two parallel bars slightly
darker than ground color crossing the end of each discoidal cell, these
bars frequently red in color, and with area between these bars dirty
white. C. cecrops differs from the other species, particularly in
having the orange red along inner side of postmedian band much
broader on the under surfaces of hindwing; in having the black
pupilated submarginal spots in interspaces M3; and Cu, gray, only
rarely orange red; in having the lunule adjacent to outerside of
W-shaped portion of postmedian band usually dark brownish gray,
only rarely red or with a slight red suffusion and in having a larger
black spot on the anal lobe.

Length of forewing 11-15 mm.

Male genitalia, as illustrated by figure 9, with distal margin of
lateral lobe of uncus convex and with lower margin of lobe undulate,
convex near base and concave before lower distal angle. Eighth
tergite (figure 202) very similar to that of janeirica and with posterior
concavity slightly broader than in dsobeon.

Frema.e (figs. 65, 66, 69 [spring form], 70 [summer form]).—Wings
above sometimes entirely like the male but usually differing in having
a great deal of blue over disc of hindwing. Wings below quite like
those of the male.

Length of forewing 11-14 mm.

Female genitalia, not illustrated, and not different from C. susanna,
and as in that species having the posterior ostium bursae lobes either
rounded or armed with two spines.

VARIATION.—Spring specimens of both sexes differ from summer
specimens in having more blue on the wings above with smaller
fuscous submarginal spots on the hindwing. On the undersurfaces

|

NO. 3662 GENUS CALYCOPIS—FIELD 19

the ground color in spring specimens is frequently darker and the
black pupillated marginal lunules of the hindwing usually contain
orange or red scales. One aberration has been described and named
and is placed in the synonymy (ab. gottschalki Clark) as it is only an
individual variant differing in having all red of undersurfaces replaced
by yellow.

TYPE-LOCALITIES.—The type-localities for cecrops and poeas are
unknown, although they undoubtedly are both from one of the
states along the eastern coast of the United States between Virginia
and Georgia, and probably the latter. Fabricius states that cecrops
is found ‘in Indiis’’; however, this is entirely false. The type-locality
for the synonym gottschalki is Fort Lewis Mt., Roanoke Co., Virginia.

Location oF TypPES.—I have been unable to locate either the
types of cecrops or of poeas. The holotype of C. cecrops ab. gottschalkr
is in the United States National Museum, type 52256.

DistriputTion.—Eastern United States from Long Island south to
Florida and west to Kansas and Texas.

New York: Long Island, East Marion (August). Nrw Jerssy: Cape May
Co., Reeds Beach (August, September); Monmouth Co., Manasquan (June).
Maryuanp: St. Mary Co., Pt. Lookout (August); Prince George Co., Beltsville
(May). Virernta: Norfolk Co., Dismal Swamp (April) ; Nansemond Co., Suffolk
(October); Princess Anne Co., Virginia Beach; Roanoke Co., Salem (April) ;
Montgomery Co. (May); Bedford Co., Apple Orchard Mt. (4,000 ft.), Mons;
Nelson Co.; Accomac Co., Dahl Swamp (July); King and Queen Co., Aylett
(July); Highland Co., Buckeye (July). Norra Caronina: Henderson Co.,
Tucedo (August) ; Buncombe Co., Asheville (August); Bertie Co., Windsor (May,
August). Sours Caro.ttna: Charleston Co., Charleston (September) ; Clarendon
Co. (September). Grorcta: Okefenokee Swamp (July); White Co., Yonah Mt.
Fiorma: Dade Co., Miami (March, May, June, July); Broward Co., Hollywood
(March); Lee Co., Ft. Myers (April); Palm Beach Co., Palm Beach; Polk Co.,
Ft. Meade; Pasco Co., Dade City, Hudson (July) ; Seminole Co., Sanford; Valusia
Co., De Land (August), Glenwood; Levy Co., Cedar Keys (July) ; Colombia Co.,
Lake City. West Viretnia: Kanawha Co., Charleston (May). TENNESSEE:
(no specific locality). Mussourt: Barry Co., Roaring River (August). Kansas:
Crawford Co. ARKANSAS: Carroll Co. (August, September). Lovuistana: Caddo
Co., Caddo Park (August). Texas: Bowie Co., Maud (June); Dallas Co., Dallas
(August, September); Cameron Co., Brownsville, Esperanza Ranch.

MATERIAL stTupIED.—Fifty-five males and 49 females, including
the genitalia of 14 males and 14 females.

Calycopis isobeon (Butler and Druce), new combination

Figures 20j, 71-78

Tmolus isobeon Butler and Druce, 1872, Cistula entomologica, vol. 1, p. 108.—
Butler, 1873, Lepidoptera exotica, p. 161, pl. 57, fig. 2.

Thecla beon (Cramer)—Godman and Salvin [not Cramer, a misidentification],
1887, Biologia Centrali-Americana, Lepidoptera-Rhopalocera, vol. 2, p. 75;
ibid., vol. 3, pl. 57, figs. 4-6—Draudt, 1920, in Seitz, Die Gross-
Schmetterlinge der Erde, vol. 5, p. 795, pl. 158, figs. b3, b4.

20 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 119

Calycopis beon (Cramer).—Field [not Cramer, a misidentification], 1940, Bull.
Univ. Kansas, Biol. Ser., vol. 39, no. 10, pp. 142-143; 1940, Bull. Brooklyn
Ent. Soc., vol. 35, no. 4, pp. 134-135; 1941, Journ. Kansas Ent. Soc., vol. 14,
no. 2, pp. 66, 68-69.—Clench, in Ehrlich, P.B., and Ehrlich, A. H., 1961,
How to know the butterflies, p. 199.

Strymon beon (Cramer).—Klots [not Cramer, a misidentification], 1951, Field
guide to butterflies of North America east of the Great Plains, p. 1384.—dos
Passos, 1964, Synonymic list of the Nearctic Rhopalocera, p. 56.

Mate (figs. 71-74).—Ground color of wings above dark brown,
sometimes dark metallic blue; differing from the summer form of
C. cecrops on the hindwing in having a considerable amount of blue
in lower half of wing below cell, occasionally with some blue in the
cell and thus similar to the spring form of cecrops. On the under-
surfaces it differs from both forms of cecrops by having the postmedian
tricolored band of the hindwing very much narrower at its broadest
point, being less than a millimeter in width and becoming even more
narrow toward the costal margin; in the forewing the red element of
this band is no wider than the black and white elements; on this
surface of the hindwing the middle section of the W-shaped mark is
more sharply angled than in cecrops; the black spot on anal lobe is
smaller than in cecrops and the red above this spot is more distinct;
the lunule adjacent to the W-shaped mark is always red in isobeon not
dark gray as in cecrops, and the black pupillated submarginal spots in
interspaces M; and Cu, are usually orange red (only rarely gray) with
smaller black pupils.

Length of forewing 10-14 mm.

Male genitalia, not illustrated, hardly differing from cecrops, except
for a slightly shorter aedeagus. In isobeon the average length of the
aedeagus is 3.24 mm. with the extremes of 2.33 and 3.75 mm., while in
cecrops this average is 3.75 mm. with the extremes of 3.50 and 4 mm.
In isobeon the aedeagus becomes progressively shorter toward the
southern limit of its distribution so that in Texas the average length
is 3.37 mm., in Mexico 3.19 mm., in Costa Rica 3mm. and in Panama
2.66mm. Eighth tergite (fig. 207) with posterior concavity usually
more narrow than in cecrops.

Ferma. (figs. 75-78).—Differing from the male in having more blue
on wings above, the blue extending through discal cell of hinawing
and on base of forewing. On the undersurfaces it differs from cecrops
females in the same way that the males differ from that species.

Length of forewing 10-13 mm.

Female genitalia, not illustrated, not different from either cecrops or
susanna

TypE-LocaLity.—‘‘Cartago, Costa Rica.”’

ADDITIONAL TYPE DATA.—Originally described from both sexes, and
although the number of specimens was not stated, only two specimens

NO. 3552 GENUS CALYCOPIS—FIELD 21

in the collection of the British Museum (Natural History) can be
positively identified as belonging to the original series. The male
labeled BM type no. Rh. 913 with the abdomen lacking is hereby
designated the lectotype. Mr. D. S. Fletcher of that institution
kindly compared it with a male from Irazti, Costa Rica (near the
type locality) and this male is the basis for the identification of
C. isobeon in the present work. The female type was dissected and
studied by Mr. Fletcher and is the female of quite another species
not belonging in the genus Calycopis.

DistripuTIion.—Texas south through Mexico and Central America
into Panama.

UNITED STATES: TEXAS: Bowie Co., Maud; Dallas Co., Dallas (September) ;
McLennon Co., Waco (March), Kerr Co., Kerrville; Uvalde Co., Concan; Neuces
Co., (March); Guadalupe Co., Sequin; Bexar Co., San Antonio (March, April,
June); Hidalgo Co., Donna (October), Progresso; Cameron Co., Brownsville
(May, June, July, October), San Benito (May, June July), Esperanza Ranch.
Mexico: State of Nuevo Leon, Monterey (April, 1800 ft.); State of San Luis
Potosi, Huichihud4yan (September), Tamazunchale (April); State of Vera Cruz,
Jalapa, Presidio, Vera Cruz (June), Cordoba, Orizaba (June, 2000 ft.), Portin
(June, 1600 ft.), Ojo de Agua (1600 ft.), Coatepee (May); State of Oaxaca, San
Geronimo (June). GUATEMALA: Department of Retalhuleu, San Sebastian;
Department of Guatemala, Guatemala City (October); Department of Zacapa,
Zacapa (October). Costa Rica: Province of Cartago, Cartago (5000 ft.), Volean
Trazi (6-7000 ft.). Panama: Province of Chiriqui, El Vole4n Chiriqui (February).

MATERIAL STUDIED.—Sixty-four males and 51 females including

the genitalia of 32 males and 29 females.

The partunda Group

Uncus in lateral view with lower angle greatly produced and with
ventral margin greater than, equaling, or nearly equaling length of
distal margin. Aedeagus fairly straight, ending distally in a straight
or upward projected point. One species (C. vibulena), with genitalia
characters somewhat resembling C. torqueor (see the torqueor
eroup), has the aedeagus sharply curved to the right (in dorsal view)
but with the distal end of aedeagus not bifurcate and also with an
uncus typical of the partunda group.

Calycopis susanna, new species
Fiaures 10, 20k, 21e-g, 27, 83-90

Mate (figs. 83-86).—Wings above indistinguishable from C.
isobeon. Undersurfaces of wings of some specimens indistinguishable
from C. isobeon but usually with red of tricolored postmedian band on
forewing a little less distinct. Specimens found in the northern part
of the range (Mexico to Costa Rica) tend to have a second submarginal
black pupillated red spot on the hindwing below; however, this also
occurs in some southern specimens.

22, PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Length of forewing 11-14 mm.

Male genitalia, as illustrated by figure 10, with upper distal angle
of lateral lobe of uncus rounded, lower distal angle also rounded and
somewhat produced (often greatly so), causing distal margin to be con-
cave. Variation in the unci of a single species is unusual in Calycopis
and at first two species were thought to be represented here. One
showing a greatly extended lower distal angle (fig. 21/) is particularly
common in Costa Rica but also occurs in Mexico, Guatemala, Hon-
duras, British Honduras and Panama. The other with this angle not
greatly extended (fig. 21¢) is also common in Costa Rica and is found as
well in Mexico, Honduras and French Guiana. Intermediates (fig. 10)
have not been found in the northern part of its range but do occur in
the Canal Zone, Panama, Colombia and Venezuela. Because of the
presence of intermediates and because of the fact that there is no
correlation between these differences in the unci and the slight habitus
differences reported above, I treat these forms as variations of a
single species. Eighth tergite (fig. 20k) with posterior concavity more
narrow and deeper than in C. isobeon.

FrEmMa ez (figs. 87-90).—Not distinguishable from females of isobeon.
This sex can be identified only by association with males.

Length of forewing 9-13 mm.

Female genitalia, as illustrated by figures 27a-d, very variable and
not differing from isobeon or cecrops.

Typxr-LocaLity.—Territory of Guyane, Cayenne, French Guina.

ADDITIONAL TYPE DATA.—Described from the holotype, male (lo-
cality as given above; USNM type 34899; William Schaus Collection;
male genitalia preparation WDF 2712, 1948), allotype, female (same
data as the holotype; female genitalia preparation WDF 2707, 1948)
and from five male and two female paratypes from Cayenne and from
Maroni River, French Guiana. Note: Many additional specimens
of both sexes were studied but not included in the paratype series
because of the variation in the genitalia. The association of the
allotype and two female paratypes with the French Guiana males is
thought to be accurate because isobeon is not known to occur in
French Guiana.

Location oF TYPES: Holotype, allotype, three male and one female
paratypes in the United States National Museum. One male para-
type and one female paratype in the British Museum (Natural
History). One male paratype in the American Museum of Natural
History.

DisTRIBUTION.—Mexico south through Central America to Ecu-
ador, Colombia, Venezuela, and French Guiana.

Mexico: State of San Luis Potosi, El Bonito (June), El Pujol (July); State of
Vera Cruz, Jalapa, Santa Rosa (April), Presidio (April, June); State of Nayarit,

NO. 3552 GENUS CALYCOPIS—FIELD 23

Compostela (October); State of Colima, Colima (February, March, April, De-
cember); State of Yucatan. GuaTEMALA: Department of Retalhuleu, San
Sebastian. British Honpuras: District of Toledo, Rio Grande (June). Hon-
puRAS: Department of Atldntida, La Ceiba (November). Costa Rica: Prov-
ince of Limén, La Florida (May), Gudpiles (March), Puerto Limén (January,
May), Santa Clara; Province of Cartago, Turrialba (May), Tuis; Province of San
Jose, El Rodeo (June). Canat Zone: Ancon (June, August), Empire (Septem-
ber), Balboa (May, July), Barro Colorado Island (January), La Cruces Trail near
Corozal (June), Corozal (February), La Boca (January), Culebra (December).
Cotomsia: Department of Cundinamarca, Bogotdé; Department of Caldas;
Department of Valle del Cauca, Cali District (February, 3260 ft.), Hormiguero
(January, 3260 ft.). Ecuapor: Province of Bolivar, Balzapamba (700 meters,
June). VENEZUELA: State of Carabobo, Puerto Cabello (April), San Esteban
(September) ; State of Yaracuy, Aroa. FrReNcuH GuraAna: Territory of Guyane,
Cayenne; Territory of L’Inini, 60 miles up Maroni River. Note: One specimen
labeled ‘‘Ponce, Porto Rico” is probably mislabeled, as no member of Calycopis is
known from the West Indies.

MaTERIAL STUDIED.—One hundred five males and 98 females, in-

cluding their genitalia.

Calycopis drusilla, new species
Fiaures 11, 201, 28, 81, 82, 91, 92

Mate (figs. 81-82)—Wings above most similar to C. janeirica
with forewing darker and with a dull metallic bluish-purple iridescence
over base and remainder of wing except along outer margin. Hind-
wing above with bluish purple more brilliant and with outer border
not or seldom breaking up into individual black spots below vein
Cu,. Wings underneath similar to janeirica but hindwing always
with two (instead of one) submarginal black pupillated lunular-shaped
red spots, one below vein Cu, and the second, smaller, adjacent to
this and below vein M3.

Length of forewing 13-14 mm.

Male genitalia, as illustrated by figure 11, with saccus extremely
long, longer than harpes, and with foreward bend in vinculum
extremely elevated, lateral lobe of uncus with distal margin concave,
lower margin convex in the middle and with upper distal angle greatly
produced; inner surface of gnathos arm with carina long and ridgelike.
Eighth tergite (fig. 202) with posterior concavity very shallow and
broad.

Fremaze (figs. 91, 92).—Wings above dark brown with blue ray
on hind margin of forewing and with hindwing blue in the cell and
below vein M;, the blue similar to caulonia in its placement but more
brilliant in tone being nearly as much so as in drusilla males. Wings
below quite like those of the male.

Length of forewing 11—12.5 mm.

Female genitalia, as illustrated by figure 28, with combined length

24 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

of ductus bursae and bursa copulatrix less than 3% times the width of |
ostium bursae; posterior ostium bursae lobes rounded, without spurs. |

TypE-Locatiry.—Province of Limén, La Florida, Costa Rica.

ADDITIONAL TYPE DATA.—Described from the holotype, male
(locality as given above, elevation 500 feet; March; Collection
William Schaus; USNM type 34900; male genitalia preparation |
WDF 2731, 1948), allotype, female (Cabima, Panama; May; August |
Busck, female genitalia preparation WDF 2820, 1948) and from five
male and six female paratypes from various localities in Costa Rica,
Mexico, and the Panama Canal Zone.

Location oF tTypEs.—Holotype, allotype and three male paratypes
in the United States National Museum, two male and six female
paratypes in the American Museum of Natural History, New York
City.

DistTRIBUTION.—Mexico south through Central America into the
Panama Canal Zone.

Mexico: State of Vera Cruz, Santa Rosa (August), Misantla (June,
September); State of Durango, Presidio (April). Costa Rica: Province of
Limén, La Florida (500 feet, March); Gudpiles (December). Canau ZONE:
Barro Colorado Island (March), Ancon Hill (February), Empire (June), Cabima
(May).

MATERIAL STUDIED.—Six males and seven females, including their
genitalia.

Calycopis vibulena (Hewitson), new combination
Figures 12, 20m, 93, 94

Thecla vibulena Hewitson, 1877, Illustrations of diurnal Lepidoptera: Lycaenidae,
vol. 1, p. 190; vol. 2, pl. 76, figs. (co) 601, 603.

Ma tz (figs. 93, 94).—Similar to C. drusilla with entire forewing,
except for a narrow black border, dull and dark metallic bluish purple
and with hindwing having costal and outer borders very dark and
wider than in C. drusilla. In C. vibulena the brilliant bluish purple
is restricted to a smaller area in base of wing. Hindwings underneath
darker smoky brown than in ©. drusilla and lacking the red along
innerside of postmedian band, lunule adjacent to outerside of W-shaped
portion of postmedian band in interspace Cuz, fuscous not red in
color but sometimes with a few scattered red scales. A second
submarginal, black pupillated, lunular-shaped red spot absent or
sometimes barely indicated by a little red in interspace M,; this
red never with a distinct black pupil.

Length of forewing 11-13 mm.

Male genitalia, as illustrated by figure 12, with aedeagus in dorsal
view sharply bent to the right at distal end; with saccus short, much
shorter than harpe; with foreward bend of vinculum lower than in

| NO. 3552 GENUS CALYCOPIS—FIELD 25

— drusilla. Lateral lobe of uncus with distal margin slightly concave

and lower margin slightly concave near base; with upper and lower

distal angles not greatly produced, lower angle less produced than
in drusilla and vitruvia; inner surface of gnathos with pointed carina.
_ Eighth tergite (fig. 20m) similar to drusilla, somewhat more cordate
- in shape.

Frmate.—Unknown.

Typr-LocaLiry.—Belém (‘‘Paré”’), State of Para, Brazil.

ADDITIONAL TYPE DATA.—This was described from both sexes from
“Brazil, the Amazon (Pardé), and New Granada” and the total
number of specimens in the series was not stated in the original
description. According to Mr. D. S. Fletcher of the British Museum
(Natural History) ‘‘there are eight specimens from MHewitson’s
original series in the collection” (in litt.). A study of these specimens
reveals that there are three or possibly four species in addition to
vibulena in this series. ‘Two males are partunda, one male and two
females are janeirica, one male is lerbela and the male with missing
abdomen is probably susanna. Only one of these vibulena specimens
was given a type number and I designate this specimen the lectotype.
It is labeled ‘Para, Brazil” and “BM Type No. Rh. 912.” Mr.
Fletcher dissected this lectotype and compared with it a topotype
which was then lent to me. This compared topotype is the basis for
the identification of C. vibulena in this paper and is the one illustrated
(figs. 12, 20m, 93, 94).

RemarKks.—Three of the five figures included by Hewitson in his
original description (1877) are of species other than mbulena as fol-
lows: figure 599 is of a male C. janeirica, figure 600 is of a male C.
lerbela and figure 602 is of a female, probably C. janeirica although it
looks more like female susanna.

DisrrisutTion.—Known only from the State of Paré in Brazil.

Braziu: State of Pard, Belém, Igarapé-Agu.
MATERIAL STUDIED.—Five males, including their genitalia.

Calycopis vitruvia (Hewitson), new combination
Figures 138, 20n, 29, 109, 110, 119, 120

Thecla vitruvia Hewitson, 1877, Illustrations of diurnal Lepidoptera: Lycaenidae,
vol. 1, p. 193; vol. 2, pl. 77, fig. (co?) 613 (two figures).
Thecla fortuna Druce, 1907, Proc. Zool. Soc. London, 1907, p. 608.

Mate (figs. 109, 110).—Wings above dull metallic bluish purple
in color, similar to those of C. vibulena, with the dark borders on
costal and outer margins more narrow, with brilliant blue color of
forewing extending over base, below the cell and vein Cu, and with
a streak of blue in lower half of the cell. Wings below similar to

26 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119 |

vibulena, not as dark brown in ground color and with red along inner |
side of postmedian band absent as in that species but with the lunular |
marking adjacent to outer side of W-shaped portion of postmedian |
band in interspace Cu, dark red with very little fuscous.

Length of forewing 11.5-12 mm.

Male genitalia, as illustrated by figure 13, with aedeagus nearly |
straight, slightly upturned at distal end; saccus not as long as harpe; |
foreward bend of vinculum below middle; lateral lobe of uncus with
distal margin nearly straight, only slightly convex and shortened, |
being about same length as lower margin and forming nearly a right
angle with lower margin, which is slightly concave; inner surface of
enathos arm with carina pointed and much nearer distal end than in
vibulena. Eighth tergite (fig. 20n) with posterior margin undulate
with a very shallow posterior concavity.

FEMALE (figs. 119, 120)——Wings above similar to male, the blue
color almost as brilliant and restricted in hindwing to the base, the
cell, and area below vein M3; the blue in the forewing, as in the male,
covering base of wing below cell and vein Cu, with a few scattered
blue scales in lower half of cell and in interspace Cu, near cell; hind-
wing with a submarginal white to bluish-white line below vein M3
(this line absent in the male) ; dark border broken into three individual
spots below vein M;. Wings beneath similar to male with red
markings larger.

Length of forewing 12 mm.

Female genitalia, as illustrated by figure 29, with combined length
of ductus bursae and bursa copulatrix five times the width of ostium
bursae, the posterior ostium bursae lobes forming spurs that are closer
to one another than in C. fractunda; ductus seminalis from near middle
of combined ductus bursae and bursa copulatrix.

TYPE-LOCALITIES.—For vitruvia: Belém (‘‘Pard’’), State of Para,
Brazil. For fortuna: “Yurimaguas,’”’ Department of Loreta, Peru.

ADDITIONAL TYPE DATA.—Vitruvia was apparently originally
described from a single male, the holotype (locality as given above;
Hewitson Collection; British Museum (Natural History) type No.
Rh. 900; genitalia preparation number NHB 1951 484.) Fortuna
was apparently described from at least two male specimens as the
localities given in the original description are ‘‘Yurimaguas, Peru’
and ‘“Topajos, Amazona.’’ The specimen from Peru bears a type
label and the British Museum (Natural History) type number Rh.
902. I designate this specimen the lectotype.

NovrE on syNonyMy.—Mr. D. S. Fletcher dissected and compared
the types of both vitruva and of fortuna and found the latter to be a
synonym. A third specimen compared with these types and found

NO. 3552 GENUS CALYCOPIS—FIELD Dik

to be identical was sent to me for study. This is the specimen illus-
trated in figures 13, 20n, 109, 110.

Distripution.—Brazil and Peru.

Braziu: State of Pard, Belém, (Pard). Prru: Department of Loreto,
Yurimaguas, Iquitos, Department ?, Achinamiza.

MATERIAL STUDIED.—T wo males and one female, including their
genitalia.

Calycopis bellera (Hewitson), new combination
Ficures 14, 200, 30, 79, 80, 95, 96
Thecla bellera Hewitson, 1877, Illustrations of diurnal Lepidoptera: Lycaenidae,
vol. 1, p. 194; vol. 2, pl. 77, fig. (9) 618.
Thecla origo Godman and Salvin, 1887, Biologia Centrali-Americana, vol. 37
Lepidoptera-Rhopalocera, vol. 2, p. 73; ibid., vol. 3, pl. 56, figs. 32, 33.

Mats (figs. 95, 96).—Wings above most similar to drusilla, differmg
by the broader outer margin above vein Cu, which gradually broadens
toward costa. Wings below most similar to wmbulena but with a
small amount of red along inner side of postmedian band on hind-
wing.

Length of forewing 13.5 mm.

Male genitalia, as illustrated by figure 14, with aedeagus bent
downward at base and slightly and gradually upturned distally; saccus
relatively short, much shorter than harpe; foreward bend of vinculum
prominent and low (much below middle); lateral lobe of uncus with
distal margin strongly concave, lower distal angle as prominent as
upper distal angle, lower margin slightly undulate; inner surface of
enathos arm with carina ridgelike and pointed; harpe in lateral view
broad at base, gradually tapering and suddenly expanding before apex
and spatula-like in shape; harpes, in ventral view, with apices widely
separated. Eighth tergite (fig. 200) not cordate in shape with
posterior concavity broad and deep.

FreMate (figs. 79, 80).—Wings above darker than the females of
most other species in the genus, the forewing, except outer border,
dull metallic blue and hind margin below vein 2dA, brilliant blue;
hindwing with base of wing, cell, and area below vein M; of same
brilliant blue; a blue submarginal line between vein M; and the black
anal lobe, and immediately inward from this line a series of three
black lunular-shaped marks. Wings below similar to those of male,
with ground color slightly paler brown or gray brown; hindwing with
red along inner side of postmedian band more distinct and red markings
in lower half of wing slightly larger.

Length of forewing 10—-12.5 mm.

Female genitalia, is illustrated by figure 30, with combined length
of ductus bursae and bursa copulatrix slightly over four times the

28 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

width of ostium bursae and with lobes posterior to ostium bursae
only slightly pointed; ductus seminalis distinctly beyond middle of
combined length of ductus bursae and bursa copulatrix.

TyprE-LocaLity.—For bellera: ‘The Amazon.” For origo: Belém
(“Pardé’’), State of Pard, Brazil.

ADDITIONAL TYPE DATA.—C. bellera was described from both sexes
although the number of specimens was not stated. I designate as the
lectotype, the female specimen which is labeled type number Rh. 914
in the collection of the British Museum (Natural History). The male
is apparently lost. C. origo was evidently described from at least
three male specimens since the localities given in the original descrip-
tion are Chiriqui, Panama, Panama City, and the “Amazons Valley.”
The specimen labeled ‘“Amazon, Para,” is the one that bears a red
type label and the British Museum type number Rh. 909. I designate
this specimen the lectotype. A topotype compared by Mr. D. S.
Fletcher with the lectotype of origo and found to be a perfect match
in both maculation and color as well as the genitalia is the basis both
for identification and for synonymizing with C. bellera. This is the
male illustrated in figures 14, 200, 95, 96.

Distrisution.—Brazil and Bolivia.

Braziu: State of Pard, Belém (Pard). State ?: ‘Amazon.’ Boutvra: De-
partment ?, Guanay (1300 ft., April).

MATERIAL STUDIED.—One male and two females, including their
genitalia.

Calycopis partunda (Hewitson), new combination
Ficures 15, 21a, 31, 97, 98

Thecla partunda Hewitson, 1877, Illustrations of diurnal Lepidoptera: Lycaenidae,
vol. 1, p. 206; vol. 2, pl. 82, figs. (1) 685, 686.

Matz (figs. 97, 98)—Forewings above similar to those of bellera
in the ground color with a decidedly dull metallic purplish sheen;
differing from bellera in having a small ray of bright purplish blue in
base of forewing below 2dA and in having the dark border of hindwing
very wide throughout and thus restricting more to the base the bright
purplish blue; wings below as in witruvia entirely lacking the red
usually found along inner side of postmedian band. The two syntypes
of C. partunda were compared by Mr. D. S. Fletcher of the staff of
the British Museum (Natural History) with Hewitson’s figure of the
undersurface (fig. 686). Mr. Fletcher reports that “the area between
the elements of the postmedian arch below vein Cu, should be rusty red
in this figure’ (in litt.).

Length of forewing 13 mm.

Male genitalia, as illustrated by figure 15, with aedeagus nearly
straight; saccus shorter than harpe; dorsal margin of tegumen (in

NO. 3552 GENUS CALYCOPIS—FIELD 29

lateral view) terminating abruptly; lateral lobe of uncus with lower
and distal margins concave, upper distal angle produced, lower distal
angle rounded; inner surface of gnathos arm without carina; harpe in
lateral view with apex blunt; harpes in ventral view with apices not
widely separated. Eighth tergite (fig. 21a) cordate in shape and simi-
lar to that of torqueor except for the posterior concavity being more
like that of drusilla.

Frmate.—Not illustrated. Similar to bellera on both surfaces of
wings, differing only in having the ground color a slightly darker blue
above and darker gray below.

Length of forewing 11.5 mm.

Female genitalia, as illustrated by figure 31, with combined length
of ductus bursae and bursa copulatrix between three and three and
one-half times width of ostium bursae; posterior ostium bursae lobes
rounded, and thus very similar to drusilla, but with these lobes more
flat than in that species. Ductus seminalis from near middle of
combined ductus bursae and bursa copulatrix.

Type LtocaLtity.—‘“The Amazon.”

ADDITIONAL TYPE DATA.—Partunda was described from two males
from Bolivia and the Amazon in the collection of W. C. Hewitson
now in the British Museum (Natural History). I designate the male
specimen bearing the type label number Rh. 910 as the lectotype.
This is the specimen from the Amazon. Mr. D. S. Fletcher, who
reports that this specimen lacks its abdomen, has kindly compared
with it a specimen labeled ‘‘Amazonas” which he found to be identical.
This compared specimen is the basis for the identification of the name
C. partunda in this paper and is the male illustrated in figures 15, 21 A,
95, 96.

DistriputTion.—French Guiana, Brazil and Bolivia.

FrencH Guiana: Territory of Guyane, Cayenne. Braziu: State ?, the
Amazon; State of Pard, Parad; State of Paraiba, Jofo Pessoa (June). Bonivia:
no specific locality.

MATERIAL STUDIED.—F our males and one female, including their
genitalia.

Calycopis amplia (Hewitson), new combination

Fieurss 32, 121, 122

Thecla amplia Hewitson, 1877, Illustrations of diurnal Lepidoptera: Lycaenidae
vol. 1, pl. 195; vol. 2, pl. 77, figs. (?) 621, 622.—Godman and Salvin, 1887
Biologia Centrali-Americana, Lepidoptera-Rhopalocera, vol. 2, p. 75; ibid.,
vol. 3, pl. 57, figs. [female not male] 7, 8.

Mate.—Unknown.
FEMALE (figs. 121, 122).—Wings above most similar to those of
bellera and indigo, but with less blue on hindwing than in bellera and

30 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

with this color indistinct and confined entirely to area near base;
wings below differing from all of its known relatives in having a
crescent-shaped bar below vein Cuz on forewing placed inward from
postmedian line; otherwise similar to wmdigo except that the light
markings are orange not red and with orange lunule adjacent to
postmedian line in interspace Cu, much obscured with fuscous; in
addition to the submarginal black pupillated orange lunule in inter-
space Cu, there is a similar but smaller lunule in interspace M; and
sometimes a trace of a third one in interspace Mp.

Length of forewing 10.5-11 mm.

Female genitalia, as illustrated by figure 32, with combined length
of ductus bursae and bursa copulatrix much the shortest of any species
in the genus, being more than two and one-half and just under three
times the width of ostium bursae; bursa copulatrix shorter than
ductus bursae plus ostium bursae lobes; teeth on posterior ostium
bursae lobes very short.

TypE-LocaLiry.—“Nicaragua (Chontales).”

ADDITIONAL TYPE DATA.—Described from a single female, the
holotype (locality as given above, Hewitson Collection, British
Museum (National History) type No. Rh. 917). The abdomen being
missing from this type, Mr. D.S. Fletcher compared two topotypes
(including one figured by Godman and Salvin in the “Biologia”) and
four other specimens with the holotype. Finding these specimens to
be conspecific with amplia they were loaned to me and are the basis
for the identification of that name in this paper.

DistTRIBUTION.—Nicaragua, Costa Rica, Columbia and Ecuador.

Nicaracua: Department of Chontales. Costa Rica: Province of Limén,

Gudpiles (March, July). Cotumsra: State of Cundinamarca, Cananche (Sep-
tember). Ecuapor: Province ?, Paramba (April).

MATERIAL STUDIED. —Six males including their genitalia.

Calycopis indigo (Druce), new combination
Ficures 16, 21b, 33, 105, 106, 123, 124
Thecla indigo Druce, 1907, Proc. Zool. Soc. London, 1907, pp. 608, 609.

Mate (figs. 105, 106).—Wings above quite similar to those of par-
tunda in having a much narrower border on hindwing than in most
species and in having a longer ray of brilliant blue below 2dA of
forewing. Wings below hardly distinguishable from partunda dif-
fering only in having a smaller submarginal black pupillated lunule
below vein Cu, on hindwing.

Length of forewing 12 mm.

Male genitalia, as illustrated by figure 16, with aedeagus nearly
straight, the base slightly bent downward; saccus nearly as long as

SS ae aa ee ee

NO. 3552 GENUS CALYCOPIS—FIELD 31

harpes; lateral element of uncus with distal margin concave just

_ before upper distal angle, which is greatly produced in lateral view,

almost spurlike, and with lower margin nearly straight, or only slightly
concave; lower distal angle of uncus only slightly rounded; arm of
gnathos with a small ridgelike carina on inner surface; apices of
harpes in lateral view slightly pointed and in ventral view close
together. Highth tergite (fig. 216) most like C. bactra in shape with
posterior concavity very shallow.

FEMALE (figs. 123, 124).—Wings above dark brown with a slight

_ dull purplish-blue sheen in base of hindwing. Wings below like the

male except that the ground color is lighter.

Length of forewing 12 mm.

Female genitalia, as illustrated by figure 33, with combined length
of ductus bursae and bursa copulatrix less than two and one-half times
the width of ostium bursae and with a pair of prominent spurlike teeth
on posterior ostium bursae lobes; ductus seminalis near middle of
combined ductus bursae and bursa copulatrix.

TypE-LocaLity.—‘‘Chapada Campo,”= Plain near Serra da Chapa-
da, ‘‘Brazil.”

ADDITIONAL TYPE DATA.—Originally described from a single male,
the holotype (locality as given above; H. H. Smith; British Museum
(Natural History) type No. Rh. 901; genitalia preparation number
NHB 1951-485). This holotype was compared by Mr. D. S. Fletcher

_ with a specimen from Mato Grosso, Brazil and found to be conspecific.

This compared specimen is the basis for the identification of the name

indigo in this paper and is the one illustrated in figures 16, 21b, 105,

and 106.

DistriputTion.—Brazil and Argentina.

Braziu: State of Mato Grosso, Serra da Chapada (January); State ?, Amazon
ARGENTINA: Territory of Misiones, Puerto Aquirre.

MATERIAL STUDIED.—Two males and two females, including their
genitalia.

Calycopis anfracta (Druce), new combination

Fieures 17, 21c 53, 54, 101, 102
Thecla anfracta Druce, 1907, Proc. Zool. Soc. London, 1907, p. 609.

Mate (figs. 101, 102)——Wings above with dark border slightly
broader than in indigo but not nearly as broad as in C. partunda.
Forewing with ground color having more of a purplish-blue sheen,
sometimes with a greenish overcast and with brilliant blue ray below
vein 2dA shorter, confined to base. Wings beneath similar to those
of C. indigo, differing only in the red markings on hindwing being
more distinctly red.

Length of forewing 12.5-13 mm.

32 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Male genitalia, as illustrated by figure 17, with aedeagus nearly
straight and with base less downward bent than in indigo; saccus
distinctly shorter than harpes; lateral lobe of uncus with distal and
lower margins concave; upper distal angle of uncus greatly produced,
more so than in indigo; lower distal angle broadly rounded; gnathos
arm with more distinct carina than in indigo; harpe in lateral view
distinctly constricted before apex forming a small apical lobe. Eighth
tergite (fig. 21¢) much broader than that of indigo but otherwise
similar.

FEMA ez (figs. 53, 54) —Wings above quite similar to indigo, below |
paler than in the male and as in that sex with markings more distinctly
red than in indigo. Red lunule next to postmedian line below vein
Cuz on undersurface of hindwing less obscured with fuscous.

Length of forewing 10.5 mm.

Female genitalia, not illustrated, not different from indigo.

TyPE-LocALity.—‘“‘Chancamayo,’’=Chanchamayo, “Peru.”

ADDITIONAL TYPE DATA.—Originally described from a single male,
the holotype (locality as given above; H. Whitely; British Museum
(Natural History) type No. Rh. 908; male genitalia preparation num-
ber NHB 1949 150). Mr. D.S. Fletcher sent me a topotype specimen
that he compared with the holotype and found to be conspecific. This
compared specimen is the basis for the identification of the name
anfracta in this paper.

Distripution.—Colombia, Peru and Bolivia.

Cotomsia: Department of Cundinamarca, Bogoté. Prru: Department of
Junin, Chanchamayo; Satipo. Bo.tivia: Department of Santa Cruz, near Santa
Cruz.

MATERIAL STUDIED.—Nine males and one female, including their
genitalia.

Calycopis anastasia, new species
Ficurss 18, 99, 100

Mate (figs. 99, 100).—Wings above similar to those of anfracta
except that the borders on the hindwings are wide as in partunda;
wings underneath with slightly darker ground color than in anfracta
but with markings almost exactly as in that species.

Length of forewing 11.5 mm.

Male genitalia, as illustrated by figure 18, with aedeagus nearly
straight, base shightly bent downward and with distal end elongated,
upceurved, and fingerlike; saccus distinctly shorter than harpes; lat-
eral lobe of uncus with distal margin concave and with lower margin
much more concave than in anfracta; upper distal angle of uncus
greatly produced asin anfracta; lower distal angle somewhat truncate;
enathos arm with carina placed further back from distal end than in

NO. 3552 GENUS CALYCOPIS—FIELD ao

anfracta; harpe in lateral view gradually narrowed to apex which
is rounded forming a lobe that is not as distinct asin anfracta. Eighth
tergite not illustrated and shaped about as in partunda (fig. 21a).

Fremaue.—Unknown.

TyprE-LOCALITY.—Province of Chapare, Department of Beni,
Bolivia.

ADDITIONAL TYPE DATA.—Originally described from the holotype,
male (locality as given above; USNM type 34901; male genitalia
preparation WDF 3432, 1951) and two male paratypes from Santiago
del Estero and Santa Cruz de la Sierra, both in the department of
Santa Cruz de la Sierra, Bolivia.

Location oF TyPES.—Holotype and one paratype in United States
National Museum, one paratype in British Museum (Natural His-
tory).

DistrisuTion.—Known only from Bolivia.

MATERIAL STUDIED. —Three males, including their genitalia.

Calycopis fractunda, new species
Ficures 19, 21d, 34, 103, 104, 125, 126

Mate (figs. 103, 104).—Wings above similar to those of indigo
with brilliant blue on the hindwing decidedly purplish and with fore-
wing a dull metallic bluish purple except along outer margin where
the color is dark brown; wings below not different from partunda
and as in that species with the bars of postmedian band in interspaces
M; and Cu, nearly straight.

Length of forewing 14 mm.

Male genitalia, as illustrated by figure 19, with aedeagus nearly
straight, distal end fingerlike, upcurved, and greatly elongated, much
more so than in anastasia; saccus nearly as long as harpes; lateral
lobes of uncus connected across the top and in lateral view with distal
and lower margins concave; upper distal angle of uncus greatly pro-
longed, much more so than in any other species of Calycopis; lower
distal angle broadly rounded and similar to that of anfracta; gnathos
arm without carina; harpe in lateral view similar to anfracta, in
ventral view similar to anastasia. Eighth tergite (fig. 21d) similar
to that of anfracta, slightly more cordate in shape.

Frmaun (figs. 125, 126).—Wings above and below very similar
to those of vitruvia, differing only in the hindwing underneath having
a smaller black pupil in submarginal red lunule of interspace Cu,
in having the bars of postmedian band in interspaces M; and Cu;
nearly straight and with postmedian black line above Cuz lacking
red inner border.

Length of forewing 10.5 mm.

219-950-673

34 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

Female genitalia, as illustrated in figure 34, differing from wtruvia
in having spurs of ostium bursae lobes set further apart and with
base of these lobes more produced medially.

Typxr-LocaLity.—Achinamiza (Department of Loreto), Peru.

ADDITIONAL TYPE DATA.—Described from the holotype, male (lo-
cality as given above; September; H. Bassler Collection; male geni-
talia preparation WDF 3443, 1951) and from the allotype, female
(Iquitos, Department of Loreto, Peru; October; collection of E. I.
Huntington; female genitalia preparation WDF 5069, 1951).

LocaTIon oF TyPES.—Holotype and allotype in American Museum
of Natural History, New York City.

DistRIBUTION.—Known only from Peru.

MATERIAL STUDIED.—One male and one female.

NO. 3552 GENUS CALYCOPIS—FIELD 35

/\\

Ficures 1-3.—Male genitalia (a=harpe, tegumen, uncus, vinculum, saccus and gnathos
in lateral view; b,=fused harpes in ventral view; c=uncus and tegumen in dorsal view;
d,—aedeagus in lateral view): 1, Calycopis bactra (Hewitson), from preparation 2887
(WDF); 2, C. nicolayi Field, holotype; 3, C. caulonia (Hewitson), from preparation 5310

(WDF).

36 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Ficures 4-5.—Male genitalia (see figs. 1-3 for key): 4, C. janeirica (C. Felder), from prepa-
ration 2697 (WDF); 5, C. chacona (Jorgensen), from preparation 2696 (WDF).

No. 3552 GENUS CALYCOPIS—FIELD 37

Ficures 6-7.—Male genitalia (see figs. 1-3 for key): 6, C. lerbela Field, holotype; 7, C.
torqueor Druce, from preparaion BM 1951 490.

38 PROCEEDINGS OF THE NATIONAL MUSEUM von. 119

Ficures 8-9.—Male genitalia (see figs. 1-3 for key): 8, C. xeneta (Hewitson), from prepa-
ration BM 1951 492; 9, C. cecrops (Fabricius), from preparation 2729 (WDF).

NO. 3552 GENUS CALYCOPIS—FIELD 39

Ficures 10-11.—Male genitalia (see figs. 1-3 for key): 10, C. susanna Field, from prepa-
ration 2683 (WDF); 11, C. drusilla Fiela, holotype.

40 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Figures 12-13.—Male genitalia (see figs. 1-3 for key): 12, C. vibulena (Hewitson), from
preparation BM 1949 165 A; 13, C. vitruvia (Hewitson), from preparation BM 1951 488.

NO. 3552 GENUS CALYCOPIS—FIELD 41

15

Ficures 14-15.—Male genitalia (see figs. 1-3 for key): 14, C. bellera (Hewitson), from
preparation BM 1949 147; 15, C. partunda (Hewitson), from preparation BM 1949 148.

42 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

1'7 a

Ficures 16-17.—Male genitalia (see figs. 1-3 for key): 16, C. indigo (Druce), from prepara-
tion BM 1951 497; 17, C. anfracta (Druce), from preparation 2700 (WDF).

NO. 3552 GENUS CALYCOPIS—FIELD 43

.

19

Ficures 18-19.—Male genitalia (see figs. 1-3 for key): 18, C. anastasia Field, holotype;
19, C. fractunda Field, holotype.

44 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Cr wu
CHER ie
tC

ee
Sa

Ficure 20.—Eighth tergite of male abdomen (scale in fig. h): a, C. bactra (Hewitson), from
preparation 2887 (WDF); b, C. nicolayi F ield, holotype; c, C. caulonia (Hewitson), from
preparation 5310 (WDF); d, C. janeirica (C. Felder), from preparation 2697 (WDF);
e, C. chacona (Jorgensen), from Preparation 2696 (WDF); f, C. lerbela Field, holotype;

&, C. torqueor (Druce), from preparation BM 1951 490; h, C. xeneta (Hewitson), from

preparation BM 1951 492; i, C. cecrops (Fabricius), from preparation 2729 (WDF);

j, C. isobeon (Butler and Druce), from preparation 2839 (WDF); &, C. susanna Field,

holotype; 1, C. drusilla F ield, holotype; m, C. vibulena (Hewitson), from preparation 3436

(WDF); 1, C. vitruvia (Hewitson), from preparation BM 1951 488; 0, C. bellera (Hewitson),

from preparation BM 1949 147,

NO. 3552 GENUS CALYCOPIS—FIELD 45

WU.

VY
a /
aN

yamm s

bb S

:

Ficures 21-23.—Fighth tergite of male abdomen: 21a, C. partunda (Hewitson), from
preparation BM 1949 148; b, C. indigo (Druce), from preparation BM 1951 497; ¢, C.
anfracta (Druce), from preparation 2700 (WDF); d, C. fractunda Field, from holotype.
Lateral views of unci and harpe showing variations in C. susanna Field: 21e, from prepa-
ration 2833 (WDF); f, from preparation 2681 (WDF); g, from preparation 2681. Lateral
views of harpes showing variations in C. chacona (Jorgensen): 21h, from preparation 2923
(WDF); i, from preparation 2925 (WDF); j, from preparation 2920 (WDF). Female
genitalia in ventral view: 22, C. bactra (Hewitson), holotype; 23, C. caulonta (Hewitson),
from preparation BM 1949 145.

46 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

WW

Ficures 24-26.—Female genitalia in ventral view (except 24a, which shows portion of
bursa copulatrix and ductus bursae in lateral view): 24, C. janetrica (C. Felder), from
preparation 2765 (WDF); 25, C. chacona (Jorgensen), from preparation 2766 (WDF);
26, C. lerbela Field, allotype.

ne et ae att ee a

NO. 3552 GENUS CALYCOPIS—FIELD 47

Ficures 27-29.—Female genitalia in ventral view: 27a, C. susanna Field, from preparation
2780 (WDF); 6, from preparation 2792 (WDF); c, from preparation 3467 (WDF); d,
allotype; ¢, from preparation 2896 (WDF); 28, C. drusilla Field, allotype; 29, C. v1truvia
(Hewitson), from preparation 5070 (WDF).

48 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

3 =.

Ficures 30-34.—Female genitalia in ventral view: 30, C. bellera (Hewitson), holotype;
31, C. partunda (Hewitson), from preparation 5319 (WDF); 32, C. amplia (Hewitson),
from preparation 5091 (WDF); 33, C. indigo (Druce), from preparation 3185 (WDF);
34, C. fractunda Field, allotype.

U.S. GOVERNMENT PRINTING OFFICE: 1967

PLATES

(Figures 35-126)

aD.
36.
Mis
38.
39.
40.
41.
42.

43.
44.
45.
46.

47.
48,
49,
50.

Ficures 35-50

C. bactra (Hewitson)

Male, upper surface, Ancon, C.Z., Panama.
Lower surface of 35.
Female, upper surface, Balboa, C.Z., Panama.
Female, upper surface, La Cumbre, Honduras, compared with holotype.
Female, lower surface, Cauca Valley, Cali District, Colombia.
Upper surface of 39.
Lower surface of 37.
Lower surface of 38.

C. janetrica (C. Felder)
Male, upper surface, Rio de Janeiro, Brazil.
Lower surface of 43.
Female, upper surface, Rio de Janeiro, Brazil.
Lower surface of 45.

C. chacona (Jorgensen)
Male, upper surface, Castro, Parana, Brazil.
Lower surface of 47.
Female, upper surface, Castro, Parana, Brazil.
Lower surface of 49,

Sie
52:

Doe
54.

De
56.

Sis
58.
59.
60.

61.
62.
63.
64.
65.
66.

Ficures 51-66

C. chacona (Jorgensen)
Male, upper surface, Villarica, Paraguay.
Lower surface of 51.

C. anfracta (Druce)

Female, upper surface, Puerto Aquirre, Missones, Argentina.
Lower surface of 53.

C. caulonia (Hewitson)
Female, upper surface, Santa Cruz, Brazil, compared with holotype.
Lower surface of 55.

C. lerbela Field

Male, upper surface, holotype.
Lower surface of 57.
Female, upper surface, allotype.
Lower surface of 59.

C. cecrops (Fabricius)
Male, upper surface, Salem, Va.
Lower surface of 61.
Male, upper surface, Miami, Fla.
Lower surface of 63.
Female, upper surface, Montgomery Co., Va.
Lower surface of 65.

67.

69.
70.

Ti

els
74.
15%
76.
is

72:

80.

81.
82.

Ficures 67-82

C. cecrops (Fabricius)
Male, upper surface, Dismal Swamp, near Suffolk, Va.
Lower surface of 67.
Female, upper surface, Montgomery Co., Va.
Female, upper surface, Miami, Fla.
C. isobeon (Bulter and Druce)
Male, upper surface, San Benito, Tex.
Lower surface of 71.
Male, upper surface, Vulcan Irazi, Costa Rica, compared with lectotype.
Lower surface of 73.
Female, upper surface, San Benito, Tex.
Lower surface of 75.
Female, upper surface, San Benito, Tex.
Lower surface of 77.
C. bellera (Hewitson)
Female, upper surface, Belém, Brazil, compared with lectotype.
Lower surface of 79.
C. drusilla Field
Male, upper surface, holotype.
Under surface of 81.

Ficures 83-98

C. susanna Field
83. Male, upper surface, holotype.
84. Lower surface of 83.
85. Male, upper surface, Port Limon, Costa Rica.
86. Lower surface of 85.
87. Female, upper surface, allotype.
88. Lower surface of 87.
89. Female, upper surface, Guapiles, Costa Rica.
90. Lower surface of 89.

C. drusilla Field
91. Female, upper surface, allotype.
92. Lower surface of 91.

C. vibulena (Hewitson)

93. Male, upper surface, Belém, Para, Brazil, compared with lectotype.
94. Lower surface of 93.

C. bellera (Hewitson)
95. Male, upper surface, Belém, Para, Brazil, compared with lectotype of the synonym
origo (Godman and Salvin).
96. Lower surface of 95.
C. partunda (Hewitson)
97. Male, upper surface, Amazonas, Brazil, compared with lectotype.

98. Lower surface of 97.

99:
100.

101.
102.

103.
104.

105.
106.

107.
108.

109.

110.

iu

2:

13:
114.

Ficures 99-114

C. anastasia Field
Male, upper surface, holotype.
Lower surface of 99.
C. anfracta (Druce)
Male, upper surface, Chanchamayo, Peru, compared with holotype.
Lower surface of 101.
C. fractunda Field
Male, lower surface, holotype.
Upper surface of 103.
C. indigo (Druce)
Male, upper surface, Mato Grosso, Brazil, compared with holotype.
Lower surface of 105.
C. xeneta xeneta (Hewitson)
Male, lower surface, Nicaragua, compared with lectotype.
Upper surface of 107.
C. vitruvia (Hewitson)
Male, lower surface, Iquitos, Peru, compared with holotype of vitruvia and lectotype
of fortuna (Druce).
Upper surface of 109.
C. torqueor (Druce)
Male, upper surface, Caramang River, British Guiana, a paratype compared with
holotype.
Lower surface of 111.
C. nicolayi Field
Male, lower surface, holotype.
Upper surface of 113

LS:
116.

117.
118.

19:
120.

Ficures 115-126

C. caulonia (Hewitson)
Male, upper surface, Joao Pessoa, Brazil.
Lower surface of 115,
C. xeneta devia (Moschler)
Male, upper surface, St. Jean, Maroni, French Guiana.
Lower surface of 117.
C. vitruvia (Hewitson)
Female, upper surface, Achinamiza, Peru.
Lower surface of 119.
C. amplia (Hewitson)
Female, lower surface, Guapiles, Costa Rica.
Upper surface of 121.
C. indigo (Druce)
Female, upper surface, Puerto Aquirre, Missones, Argentina.
Lower surface of 123.
C. fractunda Field
Female, lower surface, allotype.
Upper surface of 125.

119
123°

Eroceedings of
the United States

National Museum

SMITHSONIAN INSTITUTION - WASHINGTON, D.C.

Volume 119 1967 Number 3553

TYPE-SPECIMENS OF POLYCHAETES DESCRIBED BY
EDITH AND CYRIL BERKELEY (1923-1964)

By Martian H. PETTIBONE

Associate Curator, Division of Worms

The late Edith Berkeley, working as a volunteer investigator at
the Pacific Biological Station at Nanaimo, British Columbia, pub-
lished some 11 papers between 1923 and 1961, chiefly on the poly-
chaetes from the Nanaimo district. In these publications 1 new
genus, 15 new species, and 8 new varieties (some subsequently raised
to species) were described, and 1 new name was proposed. Edith
Berkeley was later joined by her husband Cyril in her polychaete
studies, and they jointly published 34 papers between 1932 and 1964.
In addition to material from British Columbia, their collection in-
cluded specimens from Alaska, the Bering Sea, the western Canadian
Arctic, southern California, the west coasts of Mexico and Panama,
Peru, Hudson Bay, and the east coast of Canada. These studies
resulted in the erection of 4 new genera, 43 new species, 1 new sub-
species, 7 new varieties, and 1 new name. Cyril Berkeley alone
published two short papers dealing with polychaetes (1942, 1956).
“Canadian Pacific Fauna” includes two papers by the Berkeleys:
“Polychaeta Errantia” (1948) and ‘Polychaeta Sedentaria” (1952a).
Their publications serve as the primary source for information on
the polychaetes of western Canada and nearby areas.

In 1964, Cyril Berkeley kindly donated the Berkeley polychaete
collection to the Smithsonian Institution. I went to Nanaimo to

1

2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

help in the transfer of the collection, Some type material of the
Berkeley species had been set apart from the general collection.
However, a good deal of type material, including syntypes and para-
types, was not so segregated.

The purpose of this paper is to indicate the change in the location
of the Berkeleys’ material from their private collection to that of the
U.S. National Museum and to provide some pertinent information
on the type material.

For each of the species described by the Berkeleys, I have given
the original reference; subsequent references to that species by the
Berkeleys are also listed. I have also included selected references to
publications of other authors, in which additional] figures or descrip-
tions may be given, or in which the Berkeley species are referred to
other genera or species. The type material is documented by the
Museum Catalog Number (USNM), the locality, and number of
specimens. In extracting type material from the general collection,
I have been guided by the date and information given in the original
reference. Type material was found for all but two species, Prio-
nospio multibranchiata E. Berkeley, 1927, and Caulleriella viridis
var. pacifica EK. Berkeley, 1929. Dr. R. W. Sims of the Annelid
Section at the British Museum (Natural History) kindly furnished
me with the list of polychaetes given to the British M useum by Edith
Berkeley. The syntypes and paratypes (referred to as cotypes)
of the Berkeley species that are deposited in the British Museum are
so indicated (BMNH).

A complete list of the papers dealing with polychaetes by Edith
Berkeley, by Edith and Cyril Berkeley, and by Cyril Berkeley is
included, together with a list of the new taxa described in each paper.

I wish to express my appreciation to Mr. Cyril Berkeley for his
gift of the Berkeley collection to the Smithsonian Institution and for
his help in making what might have been a rather tedious task of
transferring a large collection, such a pleasant one. My sister Mrs.
Katherine Swanson was particularly helpful, far beyond the call of
duty, in serving as typist and general helper.

Family Polynoidae

Lepidasthenia longicirrata E. Berkeley
Lepidasthenia longicirrata E. Berkeley, 1923, p. 214, figs. 8-13.—Pettibone, 1953,
p. 51, pl. 26, figs. 229-236.
Lepidametria longicirrata (E. Berkeley).—RF. and C. Berkeley, 1948, p. 18, fig. 22.
Type material: Holotype, USNM 32853, off Jesse Island, British
Columbia, from sponge-bed, 15 fms., June 9, 1920 (middle piece and
posterior piece with regenerating posterior end, anterior end missing).
Metatype, USNM 32852, Port Albion, Ucluelet, British Columbia,

NO. 3553 POLYCHAETES—PETTIBONE 3

July 12, 1945, Barraclough, collector (selected as type material by
E. Berkeley). Metatype, USNM 32851, Deep Bay, Maple Point,
British Columbia, 20 fms., August 1934, commensal form (specimen
in 2 pieces, selected as type material by E. Berkeley).

Malmegrenia nigralba E. Berkeley
Malmgrenia nigralba E. Berkeley, 1923, p. 213, figs. 5-7.—E. and C. Berkeley,
1948, p. 10.
Malmgrenia lunulata (Delle Chiaje).—Pettibone, 1953; peo, pis. 40) fl:

Type material: Syntypes, USNM 32875 and 32876, Piper’s Lagoon,
Vancouver Island, British Columbia, sand bed, very low tide, June
20, 1920, 3 specimens (anterior piece of 17 segments and posterior
piece of 22 segments; 2 anterior pieces of 14 and 16 segments and
2 middle pieces of 9 and 19 segments). Syntypes, BMNH 1935.
12.19.33/35, Piper’s Lagoon, Vancouver Island, British Columbia.

Family Pareulepidae
Eulepethus mexicanus E, and C, Berkeley

Eulepethus mexicanus E. and C. Berkeley, 1939, p. 328, figs. 4-7.
Pareulepis fimbriata (Treadwell).—Hartman, 1959, p. 123.
Mexieulepis mexicanus (E. and C. Berkeley).—Rioja, 1961, p. 237.
Type material: Holotype, USNM 32893, Isle Grande Bay, Mexico,
6 fms., fine sand, Apr. 8, 1937, W. Williams, collector, M. S. Stranger.

Family Sigalionidae
Sigalion lewisii E. and C. Berkeley

Sigalion lewisit E. and C. Berkeley, 1939, p. 326, figs. 2, 3.
Eusigalion lewisii (EK. and C. Berkeley).—Hartman, 1944a, p. 13.
Thalanessa lewisii (E. and C. Berkeley) —Hartman, 1959, pp. 119, 122.

Type material: Holotype, USNM. 32896, Espiritu Santo Island,
Gulf of California, 24°22’ N., 11°11’ W., 11 fms., bottom of coralline
algae, coral and shell, Apr. 18, 1937, F. E. Lewis, collector, M. Ss.
Stranger (specimen in 2 pieces). Paratype, BMNH 1938. 11.29.4,
Espiritu Santo Island, Gulf of California.

Sthenelanelia atypica E. and C. Berkeley
Sthenelanella atypica E. and C. Berkeley, 1941, p. 26, pl. 5, figs. 1-3.

Type material: Syntypes, USNM 32849 and 32850, off Corona
del Mar, Calif., 12-17 fms., July 21, 1937, G. E. MacGinitie, collector,
2 specimens (1 complete, 1 incomplete posteriorly, with pharynx
extended). Syntypes, USNM 32848, off Balboa, Calif., November
and December 1932 and February 1933, G. E. MacGinitie, collector,

33 specimens.
Remarks: Hartman (1961, p. 54) questionably referred S. atypica

to Sthenelanella uniformis Moore, 1910.

4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Family Amphinomidae
Eurythoe complanata var. mexicana E. and C. Berkeley
Eurythoe complanata (Pallas).—E. and C. Berkeley, 1939, p. 323, fig. 1 (part).
Eurythoe complanata var. mexicana E. and C. Berkeley, 1960a, p. 358.

Type material: Holotype, USNM 32884, Espiritu Santo Island,
Gulf of California, littoral, 1937, F. H. Lewis, collector. Paratypes,
USNM 32883, San Carlos Bay, Gulf of California, Sept. 5, 1959,
H. Arai, collector, 2 specimens (anterior and posterior portions).

Family Tomopteridae

Tomopteris elegans E. Berkeley

Tomopteris elegans E. Berkeley, 1924a, p. 289, pl. 1, figs. 1, 2. (Not Chun, 1888.)

Tomopteris renata KE. Berkeley, 1930, p. 76. [New name for T’. elegans E. Berkeley,
1924, preoccupied.]—E. and C. Berkeley, 1948, p. 26, figs. 31, 32.

Tomopteris (Johnstonella) renata E. Berkeley.—Uschakov, 1952, p. 106, figs.
3-5.—E. and C. Berkeley, 1957, p. 576; 1960b, p. 791.

Type material: Holotype, USNM 32879, west of Entrance Island,
British Columbia, 200 fms., July 13, 1915, C. M. Fraser, collector.
Paratypes, USNM 32880, north of Five Fingers, British Columbia,
200 fms., July 10, 1915, C. M. Fraser, collector, 4 specimens. Para-
types, BMNH 1938. 11.29.6/8, Gulf of Georgia, British Columbia.

Remarks: Dales (1955, p. 440; 1957, p. 141) referred 7. renata to
T. (Johnstonella) pacifica Izuka, 1914. Hartman (1959, p. 182) in-
dicated that T. renata is perhaps the same as 7. pacifica. The Berke-
leys (1957) consider the two species to be distinct.

Family Pilargidae
Ancistrosyllis longicirrata E. and C. Berkeley

Ancistrosyllis longicirrata E. and C. Berkeley, 1961, p. 658, figs. 5-7.
Podarmus ploa Chamberlin, 1919.—Pettibone, 1966, p. 199, figs. 24-26. [Family
Polynoidae.]

Type material: Holotype, USNM 30990, South Pacific off Peru,
5°55’ S., 82°02’ W., from plankton, P. N. Sund, collector.
Loandalia fauveli E. and C. Berkeley

Loandalia fauveli E. and C. Berkeley, 1941, p. 30, pl. 5, figs. 4-6.—Hartman,
1960, p. 89.—Pettibone, 1966, p. 196, figs. 22, 23.

Type material: Holotype, USNM 32891, Newport Bay, Calif., mud
flats, Mar. 3, 1935, G. M. MacGinitie, collector (specimen in 8 pieces).

Family Syllidae

Amblyosyllis lineata var. alba E. Berkeley

Amblyosyllis lineata Grube var. alba E. Berkeley, 1923, p. 208.—E. and C. Berk-
eley, 1948, p. 81, fig. 121.

No. 3553 POLYCHAETES—PETTIBONE 5

Type material: Syntypes, USNM 32860, 32861, 32862, Jesse to
Newcastle Island, Nanaimo region, British Columbia, 20 to 25 fms.,
in cavities of siliceous sponges, Sept. 4, 1919, 4 specimens (in 12
pieces: anterior end of 5 setigerous segments, middle piece with 7
segments, posterior piece with 1 setigerous and 1 achaetous segment;
anterior end with 5 segments, middle piece of 2 segments, posterior
piece with 6 setigerous and 1 achaetous segments; 2 anterior ends each
with 5 segments, 4 middle pieces consisting of 3, 2, and 2 segments,
and single parapodium).

Autolytus magnus E. Berkeley
Autolytus magnus BE. Berkeley, 1923, p. 210, pl. 1, figs. 3, 4—E. and C. Berkeley,
1938a, p. 47; 1945, p. 318; 1948, p. 70.—Hartman, 1948, p. 24.—Imajima
and Hartman, 1964, p. 93, pl. 18, figs. a-e.
Type material: Holotype, USNM 32897, Departure Bay, Nanaimo,
British Columbia, Feb. 23, 1920, swarming at station float, sacconereis

phase.
Autolytus trilineatus E. and C. Berkeley

Autolytus aurantiacus—E. and C. Berkeley, 1938a, p. 48. [Not Claparéde,
1868.]

Autolytus trilineatus E. and C. Berkeley, 1945, p. 318, fig. 1; 1948, p. 69, fig. 100.

Autolytus prismaticus (Fabricius).—Pettibone, 1954, p. 249, fig. 29, a, b.

Type material: Syntypes, USNM 32881 and 32882, Newcastle
Island, Nanaimo, British Columbia, June 18, 1943, 5 specimens
(complete specimen showing characteristic color markings, head of
bud indicated between setigerous segments 13 and 14; complete
specimen, 2 anterior ends, 3 middle pieces, and 3 posterior ends).

Exogone lourei E. and C. Berkeley
Exogone loureit E. and C. Berkeley, 1938a, p. 44, figs. 6-12; 1948, p. 79, fig. 117
Ezxogone dispar (Webster).—Pettibone, 1954, p. 259, fig. 28, k.
Type material: Holotype, USNM 32895, False Narrows, Nanaimo,
British Columbia, May 21, 1936.

Odontosyllis parva E. Berkeley
Odontosyllis parva E. Berkeley, 1923, p. 208, pl. 1, figs. 1, 2.—E. and C. Berkeley,
1941, p. 30; 1948, p. 83, fig. 123.
Type material: Holotype, USNM 32892, False Narrows, Nanaimo,
British Columbia, on bryozoan, May 30, 1919 (specimen in 3 pieces,
anterior end of 15 segments, 2 middle pieces each with 6 segments).

Odontosyllis phosphorea var. nanaimoensis E. Berkeley

Odontosyllis phosphorea Moore var. nanaimoensis EK. Berkeley, 1923, p. 206; 1961,
p. 1321.—E. and C. Berkeley, 1942, p. 191; 1948, p. 82, fig. 122.

Type material: Syntypes, USNM 32857 and 32859, Departure
Bay, Nanaimo, British Columbia, swarming from end of July to end

6 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

of November, 32 specimens. Syntypes, USNM 32858, Departure
Bay, Nanaimo, British Columbia, swarming, Aug. 13, 1918, 15
specimens. Syntypes, BMNH 1935. 12.19.39/42, Departure Bay,
Nanaimo, British Columbia.

Syllis harti E. and C. Berkeley
Syllis harti E. and C. Berkeley, 1938a, p. 35, fig. 2; 1942, p. 190; 1945, p. 317;
1948, p. 77, fig. 114.
Typosyllis harti (EK. and C. Berkeley).—Hartman, 1959, pp. 227, 234.

Type material: Holotype, USNM 32671, off Nootka light, west
coast Vancouver Island, British Columbia, about 45 fms., E. G. Hart,
collector (pharynx is loose in vial). Paratype, BMNH 1938. 11.29.-
10, Princess Louise Inlet, British Columbia, 25 fms., J. F. L. Hart,

collector.
Syllis pulchra E. and C. Berkeley

Syllis pulchra E. and C. Berkeley, 1938a, p. 34, fig. 1; 1948, p. 73, fig. 105.
Typosyllis pulchra (E. and C. Berkeley).—Hartman, 1944b, p. 250; 1948, p. 22.

Type material: Holotype, USNM 32847, False Narrows, Nanaimo,
British Columbia, May 30, 1919. Paratypes, USNM 32846, between
Jesse Island and Five Fingers, Nanaimo, British Columbia, 8 fms.,
July 17, 1918, 8 specimens. Paratypes, USNM 32845, Newcastle,
Nanaimo, British Columbia, July 9, 1919, 4 specimens. Paratypes,
BMNH 19388. 11.29.9, Departure Bay, British Columbia.

Syllis spenceri E. and C. Berkeley

Syllis spenceri E. and C. Berkeley, 1938a, p. 38, fig. 4; 1948, p. 75, fig. 112.
Typosyllis adamanteus (Treadwell).—Imajima and Hartman, 1964, p. 132.

Type material: Holotype, USNM 32872, Departure Bay, Nanaimo,
British Columbia, May 24, 1925, G. J. Spencer, collector. Paratypes,
USNM 32871, Departure Bay, Nanaimo, British Columbia, May 24,
1925, G. J. Spencer, collector, 20 specimens. Paratypes, BMNH
1938. 11.29.11/14, Departure Bay, Nanaimo, British Columbia.

Syllis stewarti E. and C. Berkeley
Syllis stewarti E. and C. Berkeley, 1942, p. 191; 1948, p. 72.
Typosyllis stewarti (EK. and C. Berkeley).—Hartman, 1948, p. 22.
Type material: Holotype, USNM 32899, Estevan Point, west
coast Vancouver Island, British Columbia, littoral, 1934, E. G. Hart,
collector, William J. Stewart.

Family Nereidae

Micronereis nanaimoensis E. and C. Berkeley

Micronereis variegata.—Ramsay, 1914, p. 243, figs. 2-5, 7—E. Berkeley, 1924a,
p. 290.—E. and C. Berkeley, 1948, p. 60, fig. 89. [Not Claparéde, 1863.]
Micronereis nanaimoensis E. and C. Berkeley, 1953a, p. 85, pls. 1, 2.

NO. 3553 POLYCHAETES—PETTIBONE 7

Type material: Syntypes, USNM 32854, 32855, and 32856, De-
parture Bay, British Columbia, Apr. 23 to May 23, 1947, 41 specimens.

Nereis (Ceratonereis) scotiae E. and C. Berkeley
Nereis (Ceratonereis) scotiae E. and C, Berkeley, 1956a, p. 267, figs. 1, 2.
Ceratonereis scotiae (E. and C. Berkeley).—Hartman, 1959, pp. 239, 256.

Type material: Holotype, USNM 32889, Mitchell Bay, Nova
Scotia, July 18, 1939, littoral, W. S. Hoar, collector. Paratypes,
USNM 32890, Mitchell Bay, Nova Scotia, July 18, 1939, littoral,
W.S. Hoar, collector, 2 specimens.

Nereis (Eunereis) wailesi E. and C. Berkeley
Nereis (Eunereis) wailest E. and C. Berkeley, 1954a, p. 456, figs. 1, 2.
Eunereis wailesi (E. and C. Berkeley).—Hartman, 1959, p. 241.
Type material: Holotype, USNM 32894, Cachalot, west coast
Vancouver Island, British Columbia, G. H. Wailes, collector.

Nereis (Neanthes) mancorae E. and C. Berkeley
Nereis (Neanthes) mancorae E. and C. Berkeley, 1961, p. 656, figs. 1-4.

Type material: Holotype, USNM 32887, Mancora, Peru, inter-
tidal, W. L. Klawe, collector. Paratypes, USNM 32888, Mancora,
Peru, intertidal, W. L. Klawe, collector, 5 specimens.

Nicon peruviana E. and C. Berkeley
Nicon peruviana EK. and C. Berkeley, 1964, p. 128, figs. 1, 2.

Type material: Holotype, USNM 32822, South Pacific off Peru,
approximately 5° S., 81° W., Apr. 10, 1959, at night-light, P. N. Sund,
collector (male heteronereid). Paratype, USNM 32821, South
Pacific off Peru, approximately 5°S., 81° W., Apr. 10, 1959, at night-
light, P. N. Sund, collector (male herteronereid).

Family Nephtyidae
Nephthys cornuta E. and C. Berkeley

Nephthys sp.? Weese, 1933, p. 20.
Nephthys cornuta E. and C. Berkeley, 1945, p. 328, figs. 2-4; 1948, p. 50, figs. 72, 73.
Nephtys cornuta.—Hartman, 1959, p. 106.—Clark, 1955, p. 547; 1958, p. 206.
Type material: Holotype, USNM 32873, Fishing Bay, East Sound,
Orcas Island, Washington, 11-14 fms., soft mud, 1929, A. O. Weese,
collector. Paratype, USNM 32874, Princess Louise Inlet, British
Columbia, 20 fms., May 13, 1929. Paratypes, Fishing Bay, East
Sound, Orcas Island, Washington. 11-14 fms., soft mud, 1929, A. O.
Weese, collector, 4 specimens (in collection of R. B. Clark, University
of Bristol, sectioned and stained for histological purposes; dissected
for anatomical work on brain structure).

8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Family Sphaerodoridae

Sphaerodorum biserialis E. and C. Berkeley
Sphaerodorum biserialis E. and C. Berkeley, 1944, p. 3, figs. 1-3.

Type material: Holotype, USNM 32867, Dease Strait, western
Canadian Arctic, 69° N., 106°25’ W., 45 fms., 1936-37, H. A. Larsen,
collector, R.C.M.P. St. Roch. Paratype, USNM 32868, Dease
Strait, western Canadian Arctic, 69° N., 106°25’ W., 45 fms., 1936-37 .
H. A. Larsen, collector, R.C.M.P. St. Roch,.

Remarks: According to the revision by Pettibone (1963, p. 208),
S. biserialis should be referred to Ephesiella Chamberlin, thus be-
coming Ephesiella biserialis (E. and C. Berkeley).

Family Goniadidae

Glycinde picta E. Berkeley
Glycinde picta E. Berkeley, 1927a, p. 412.—E. and C. Berkeley, 1942, p. 194;
1948, p. 35, fig. 48 —Hartman, 1948, p. 29.

Type material: Holotype, USNM 32870, False Narrows, British
Columbia, May 20, 1920. Paratypes, USNM 32879, False Narrows,
May 20, 1920; Piper’s Lagoon, Apr. 20, 1920; Departure Bay Beach,
June 27, 1918, low tide; 6 specimens. Paratypes, BMNH 1938.
11.29.18/19, Departure Bay, British Columbia.

Family Onuphidae

Onuphis eremita var. parva E. and C. Berkeley
Onuphis eremita Audouin and Milne-Edwards var. parva E. and C. Berkeley,
1941, p. 35.
Type material: Holotype, USNM 32886, off Corona del Mar,
Calif., 12-17 fms., G. E. MacGinitie, collector. Paratype, USNM
32885, off Corona del Mar, Calif., 12-17 fms., G. E. MacGinitie,

collector.
Onuphis zebra E. and C. Berkeley

Onuphis zebra E. and C. Berkeley, 1939, p. 337, figs. 9-10.—Hartman, 1944c, p.
71, pl. 3, figs. 55-60.
Type material: Holotype, USNM 32898, Punta Gorda, Lower
California, 23°04’ N., 109°35’ W., 14 fms., fine sand, Apr. 24, 1937,
W. Williams, collector, M. S. Stranger.

Rhamphobrachium longisetosum E. and C. Berkeley

Rhamphobrachium longisetosum EB. and C. Berkeley, 1938b, p. 428, figs. 1-8.—
Hartman, 1944¢c, p. 48, pl. 1, figs. 1-8.

Type material: Syntype, USNM 32865, off mouth of Santa Ana
River, southern California, 17 fms., G. E. MacGinitie, collector

NO. 3553 POLYCHAETES—PETTIBONE 9

(anterior end of 9 segments showing characteristic long anterior
setae). Syntypes, USNM 32866, off Balboa, southern California,
33 fms., May 19, 1933, G. E. MacGinitie, collector, 2 specimens
(nearly complete specimen of 34 segments and middle fragment of 18
segments, also portion of tube).

Family Lumbrineridae

Lumbrinereis ligulata E. and C. Berkeley
Lumbrinereis ligulata E. and C. Berkeley, 1941, p. 38.
Lumbrineris ligulata.—Hartman, 1959, p. 335.

Type material: Syntypes, USNM 32877 and 32878, off Corona del
Mar, southern California, 12-17 fms., July 21, 1937, G. E. Mac-
Ginitie, collector, 9 specimens (9 anterior ends, 2 middle pieces, 1
posterior end, plus jaws dissected out in small vial).

Lumbrinereis luti E. and C. Berkeley
Lumbrinereis luti E. and C. Berkeley, 1945, p. 332, fig. 6; 1948, p. 100, fig. 159.
Lumbrineris luti—Hartman, 1959, p. 335.

Type material: Holotype USNM 32864, Northumberland Channel,
British Columbia, 10-20 fms., dense mud, Apr. 16, 1937 (specimen
in 2 pieces, anterior and middle). Paratypes, USNM 32863, Nanoose
Bay, British Columbia, Aug. 28, 1918; Northumberland Channel,
Apr. 16, 1937; east coast Vancouver Island, 10-20 fms., dense mud;
head of Seymour Inlet, 36 fms.; 31 specimens (31 anterior ends, 25
middle pieces).

Family Dorvilleidae

Dorvillea pseudorubrovittata E. Berkeley
Dorvillea pseudorubrovittata E. Berkeley, 1927a, p. 409.—E. and C. Berkeley,
1948, p. 87.—Hartman, 1948, p. 29.

Type material: Syntypes, USNM 32674 and 32675, Round Island
to Mudge Island, east coast Vancouver Island, British Columbia, 25
fms., June 11, 1920, 7 specimens. Syntypes, USNM 32676, between
Jesse and Russel Islands, east coast Vancouver Island, British
Columbia, June 5, 1924, 8 specimens.

Family Orbiniidae

Aricia macginitii E. and C. Berkeley

Aricia macginitii E. and C. Berkeley, 1941, p. 40, pl. 5, figs. 7-10.
Phylo ornatus (Verrill).—Hartman, 1957, p. 265, pl. 24, figs. 1-10.

Type material: Holotype, USNM 32717, Newport Bay, southern
California, mud flats, Dec. 26, 1932, G. E. MacGinitie, collector.

219-951—66——2

10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Family Paraonidae

Aricidea longicornuta E. and C. Berkeley
Aricidea longicornuta E. and C. Berkeley, 1950, p. 53, fig. 2; 1952a, p. 38, figs.
68, 69.
Aricidea uschakovi Zachs.— Hartman, 1957, p. 321, pl. 43, fig. 5.

Type material: Holotype, USNM 32721, Pender Island, Malaspina
Channel, Nanaimo, British Columbia, 75 fms., Jan. 19, 1929 (specimen
in 2 pieces, posterior end missing). Paratypes, USNM 32720, Deep
Hole, outside Departure Bay, Nanaimo, British Columbia, 230 fms.,
2 specimens (in 4 pieces, posterior ends missing).

Aricidea lopezi E. and C. Berkeley
Aricidea lopezi E. and C. Berkeley, 1956), p. 542, figs. 1-3.

Type material: Holotype, USNM 32718, off Upright Head, Lopez
Island, San Juan Archipelago, Washington, 12 fms., Aug. 23, 1955,
R. I. Smith, collector (complete specimen). Paratypes, USNM
32719, off Upright Head, Lopez Island, San Juan Archipelago,
Washington, 12 fms., Aug. 23, 1955, R. I. Smith, collector, 2 specimens
(1 complete, 1 in 2 pieces, posterior end missing).

Family Spionidae
Boccardia columbiana E. Berkeley

Boccardia columbiana E. Berkeley, 1927a, p. 416, pl. 1, figs. 4-8—Woodwick,
1963a, p. 132, fig. 1.

Polydora (Boccardia) columbiana (KE. Berkeley).—E. and C. Berkeley, 1952a,
p. 16, figs. 22, 23.

Type material: Holotype, USNM 32714, Newcastle Island,
Nanaimo, British Columbia, Aug. 3, 1918, boring in shaly rock (in
6 pieces). Paratypes, USNM 32713, Newcastle Island, Nanaimo,
British Columbia, Aug. 3, 1918, boring in shaly rock; off Round
Island, Aug. 19, 1918; in old wood of old float, Nov. 10, 1919; Piper’s
Lagoon, in shaly rock, May 4, 1923, 9 specimens (1 complete, 8 anterior
ends, 5 middle pieces, 4 posterior ends). Paratypes, BMNH 1938.
29.29. Departure Bay, British Columbia.

Boccardia uncata E. Berkeley
Boccardia uncata E. Berkeley, 1927a, p. 418, pl. 1, figs. 9-13.
Polydora (Boccardia) uncata (E. Berkeley).—Okuda, 1937, p. 238, figs. 16, 17.—E.
and C. Berkeley, 1952a, p. 14, figs. 18-21.

Type material: Holotype, USNM 32712, Piper’s Lagoon, British
Columbia, in frail sandy tube, May 4, 1923. Paratypes, USNM
32711, Piper’s Lagoon, British Columbia, in frail sandy tubes, May 4,
1923, 3 specimens (3 anterior ends and 2 middle pieces). Paratypes,
BMNH 1938. 11.29.27/28, Piper’s Lagoon, British Columbia.

NO. 3553 POLYCHAETES—PETTIBONE 11

Neopygospio EK. and C. Berkeley

Neopygospio laminifera E. and C. Berkeley
Neopygospio laminifera E. and C. Berkeley, 1954a, p. 462, figs. 6, 7.

Type material: Holotype, USNM 32715, Rathtrevor Beach,
Nanaimo, British Columbia, in small sandy tube, July 1951. Para-
types USNM 32716, Rathtrevor Beach, Nanaimo, British Columbia,
in small sandy tubes, July 1951, 3 specimens (2 complete, one with
posterior end missing).

Polydora caeca var. magna E. Berkeley
Polydora caeca (Oersted) var. magna E. Berkeley, 1927a, p. 419.
Polydora magna FE. Berkeley.—E. and C. Berkeley, 1936, p. 473; 1952a, p. 21.

Type material: Holotype, USNM 32710, False Narrows, Nanaimo,
British Columbia, low tide (specimen in 2 pieces, anterior and posterior
ends). Paratypes, BMNH 1938. 11.29.24, Rocky Bay, British
Columbia.

Polydora cardalia E. Berkeley
Polydora cardalia EB. Berkeley, 1927a, p. 418, pl. 1, fig. 14—E. and C. Berkeley,
1952a, p. 21, figs. 38, 39.

Type material: Holotype, USNM 32709, Cardale Point, Nanaimo,
British Columbia, July 6, 1921. Paratypes, USNM 32708, Cardale
Point, Nanaimo, British Columbia, July 6, 1921; dredged off Round
Island, Aug. 19, 1920; False Narrows, May 20, 1920; Rocky Bay,
May 16, 1920; 21 specimens (2 complete, 19 anterior ends, 9 middle
pieces, and 6 posterior ends). Paratypes, BMNH 1938. 11.29.25,
Departure Bay, British Columbia.

Polydora ciliata var. spongicola E. and C. Berkeley
Polydora ciliata (Johnston) var. spongicola E. and C. Berkeley, 1950, p. 52, fig. 1;
1952a, p. 20, fig. 35.
Polydora spongicola E. and C. Berkeley —Woodwick, 1963b, p. 212, fig. 2.

Type material: Holotype, USNM 32706, False Narrows, Nanaimo,
British Columbia, in sponge encrusting rock of “reef” (specimen
with anterior end only). Paratypes, USNM 32707, Northumberland
Channel, British Columbia, in sponge on shells of Pecten hindsi,
July 1943, 48 specimens (23 complete, 25 anterior ends, 3 middle
pieces, 4 posterior ends).

Polydora socialis plena E. and C. Berkeley
Polydora socialis (Schmarda) subsp. plena E. and Cc: ee 1936. p. 468;
1952a, p. 22.
Type material: Holotype, USNM 32705, Departure Bay, N anaimo,
British Columbia, clean sand beds, littoral, May 1, 1935. Paratypes,
USNM 32704, Departure Bay and Piper’s Lagoon, Nanaimo, British

12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Columbia, clean sand beds, littoral, May 1 and 20, 1935, 28 specimens
(15 complete, 13 anterior ends, 1 posterior end).

Paraprionospio tribranchiata E. Berkeley
Paraprionospio tribranchiata E. Berkeley, 1927a, p. 415, pl. 1, figs. 2, 3—Weese,
1933, p. 19.—E. and C. Berkeley, 1936, p. 476.
Prionospio pinnata Ehlers.—HE. and C. Berkeley, 1941, p. 42; 1942, p. 196; 1952a,
p. 30, figs. 56, 57; 1963b, p. 149.

Type material: Holotype, USNM 32701, Nanoose Bay, Nanaimo,
British Columbia, Aug. 28, 1918 (anterior end only; has been dry).
Paratypes, USNM 32702, Nanoose Bay, Nanaimo, British Columbia,
Aug. 28, 1918, 2 specimens (anterior ends; have been dry). Para-
types, USNM 32703, Nanoose Bay, Nanaimo, British Columbia,
Sept. 6, 1918, 2 specimens (anterior ends, with proboscis everted).
Paratypes, BMNH 1938. 11.29.30, Comox Spit, Vancouver Island,
British Columbia.

Prionospio multibranchiata E. Berkeley
Prionospio multibranchiata EK. Berkeley, 1927a, p. 414, pl. 1, fig. 1.
Prionospio cirrifera Wirén.—E. and C. Berkeley, 1942, p. 196; 1952a, p. 28,
figs. 52, 53.
Type material: No type material was found in the collection.
Type locality: Station Flat, Nanaimo, British Columbia, low tide.

Prionospio ornata E. and C. Berkeley
Prionospio ornata E. and C. Berkeley, 1961, p. 660, figs. 8-12; 1963b, p. 149;
1964, p. 132.

Type material: Holotype, USNM 32698, South Pacific off Peru,
8°17’ S., 79°09’ W., from plankton, 0 to 15 m., Sept. 25, 1958, P. N.
Sund, collector, larva. Paratypes, USNM 32697, South Pacific off
Peru, 8°17’ S., 79°09’ W., from plankton, 0 to 15 m., Sept. 25, 1958,
P. N. Sund, collector, 35 larvae.

Spio butleri E. and C. Berkeley
Spio butleri E. and C. Berkeley, 1954a, p. 461, figs. 3, 4.

Type material: Holotype, USNM 32695, McIntyre Bay, Queen
Charlotte Islands, British Columbia, swimming in tide-pool on sandy
beach, May 27, 1953, T. H. Butler, collector. Paratypes, USNM
32696, McIntyre Bay, Queen Charlotte Islands, British Columbia,
swimming in tide-pool on sandy beach, May 27, 1953, T. H. Butler,
collector, 6 specimens (one complete, 5 anterior ends, 3 middle pieces,
1 posterior end).

Spio martinensis var. pacifica E. Berkeley

Spio martinensis Mesnil var. pacifica E. Berkeley, 1927a, p. 413.
Spio filicornis (Miiller) var. pacifica E. Berkeley —E. and C. Berkeley, 1936, p.
475; 1952a, p. 25.

NO. 3553 POLYCHAETES—PETTIBONE 13

Spio filicornis (Miiller).—Hartman, 1948, p. 36. (Includes S. martinensis, var.
pacifica.)

Type material: Holotype, USNM 32700, False Narrows, Nanaimo,
British Columbia, May 30, 1919. Paratypes, USNM 32699, Nanaimo
district, British Columbia; Horsewell Point, May 3, 1923; between
Horsewell and Plantas, May 22, 1924; end of Newcastle, May 30,
1923; False Narrows, May 30, 1919; Cardale Point, July 6, 1921;
53 specimens (32 complete, 21 anterior ends, 1 middle piece, | posterior

end).
Family Chaetopteridae

Leptochaetopterus E. Berkeley

Leptochaetopterus pottsi E. Berkeley
Leptochaetopterus pottsi E. Berkeley, 1927b, p. 441, figs. 1-3.
Telepsavus costarum Claparéde.—Monro, 1933, p. 1052—E. and C. Berkeley,
1942, p. 196; 1952a, p. 63, figs. 127-1380.

Type material: Syntypes, USNM 32691 and 32693, Piper’s Lagoon,
Nanaimo, British Columbia, Apr. 20, 1920, 17 specimens (17 anterior
ends, 5 middle pieces). Syntypes, USNM 32692, Departure Bay
Beach, Nanaimo, British Columbia, Apr. 20, 1920, 7 specimens (7
anterior ends, 2 middle pieces). Syntypes, USNM 32694, off Mudge
Island, Nanaimo, British Columbia, 25 fms., in the crevices of a
rotten piece of wood, June 11, 1920, 2 specimens (anterior ends).

Mesochaetopterus rickettsii E. and C. Berkeley

Mesochaetopterus rickettsii E. and C. Berkeley, 1941, p. 43, pl. 5, figs. 15, 16.

Type material: Holotype, USNM 32686, Newport Bay, southern
California, G. E. MacGinitie, collector (anterior end). Paratypes,
USNM 32685, Newport Bay, southern California, Jan. 19, 1932,
E. F. Ricketts, collector, 8 specimens (have been dry). Paratype,
32684, Newport Bay, southern California, Miar,°22)¢1933. (Gare
MacGinitie, collector, 1 specimen (in 13 pieces).

Family Cirratulidae

Caulleriella viridis var. pacifica E. Berkeley
Caulleriella viridis (Langerhans) var. pacifica E. Berkeley, 1929, p. 307.
Caulleriella alata (Southern).—E. and C. Berkeley, 1950, p. 57; 1952a, p. 36;
1958a, p. 405.—Hartman, 1961, p. 108.
Type material: No type material was found in the collection. Type
locality: False Narrows, Nanaimo, British Columbia.

Chaetozone spinosa var. corona E. and C. Berkeley

Chaetozone spinosa Moore var. corona E. and C. Berkeley, 1941, p. 45.
Chaetozone corona E. and C. Berkeley —Hartman, 1961, p. 109.

14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

Type material: Syntypes, USNM 32689 and 32690, off Corona del
Mar, Calif., 12-17 fms., July 21, 1937, G. E. MacGinitie, collector,
4 specimens (one complete).

Dodecaceria fewkesi E. and C. Berkeley, new name

Sabella pacifica Fewkes, 1889, p. 132, pl. 7, figs. 1, 2. [Not Grube, 1859.]

Dodecaceria pacifica (Fewkes).—E. Berkeley, 1929, p. 308.—E. and C. Berkeley,
1932b, p. 314; 1941, p. 46; 1942, p. 197; 1952a, p. 33.

Dodecaceria fistulicola Ehlers (1901).—Reish, 1952, p. 103, pl. 20.

Dodecaceria fewkesit E. and C. Berkeley, 1954b, p. 326, figs. 1-7; 1956, p. 544.
[New name for Sabella pacifica Fewkes, preoccupied.]

Remarks: According to Reish (1952), Sabella pacifica Fewkes (1889)
(preoccupied by Grube, 1859) from Santa Barbara, Calif. is identical
with D. fistulicola Ehlers (1901) from Chile. If so, D. fewkesi E. and C.
Berkeley, a new name for Sabella pacifica Fewkes, should also be
referred to D. fistulicola. Hartman (1959, p. 408) lists both D. fewkesa
and D. fistulicola as valid names.

Tharyx multifilis var. parvus E. Berkeley

Tharyx multifilis Moore var. parvus E. Berkeley, 1929, p. 307.—E. and C.
Berkeley, 1952a, p. 35.
Tharyx parvus E. Berkeley. —Hartman, 1961, p. 113.

Type material: Holotype, USNM 32687, Piper’s Lagoon, Nanaimo,
British Columbia, May 31, 1923. Paratypes, USNM 32688, Piper’s
Lagoon, Nanaimo, British Columbia, May 31, 1923, 10 specimens
(have been dry).

Family Flabelligeridae

Stylarioides negligens E. and C. Berkeley
Stylarioides eruca.—K. Berkeley, 1930, p. 69. [Not Claparéde, 1870.]
Stylarioides negligens E. and C. Berkeley, 1950, p. 58, fig. 4; 1952a, p. 10, fig. 10.
Pherusa negligens (E. and C. Berkeley).—Hartman, 1959, pp. 418, 421.
Type material: Holotype, USNM 32683, Mitlenatch Island, Gulf
of Georgia, British Columbia, 100 fms., D. Williamson, collector
(specimen in 2 pieces).

Family Capitellidae

Heteromastus filobranchus E. and C. Berkeley
Heteromastus filobranchus E. and C. Berkeley, 1932a, p. 671, figs. 5-7; 1952a,
p. 108, figs. 211, 212—Hartman, 1947, p. 428, pl. 53, figs. 1-4.

Type material: Holotype, USNM 32682, Nanoose Bay, Nanaimo,
British Columbia, 15-25 fms., Feb. 16, 1929 (specimen in 5 pieces
has been dry). Paratypes, USNM 32681, Ship Bay, Rosario Inlet,
Washington, 5-6 fms., sandy mud, A. O. Weese, collector, 6 specimens
(in 17 pieces). Paratypes, BMNH 1938. 11.29.32, Saanich Inlet,
Vancouver Island, British Columbia.

NO. 3553 POLYCHAETES—PETTIBONE 15

Notomastus pallidior var. parvus E. Berkeley
Notomastus pallidior Chamberlin var. parvus E. Berkeley, 1929, p. 312.
Notomastus parvus E. Berkeley.—Hartman, 1959, p. 445.

Type material: Syntypes, USNM 32679 and 32680, north end
Newcastle Island, Departure Bay, British Columbia, May 21, 1923,
7 specimens (9 pieces, one complete).

Remarks: N. pallidior, var. parvus is not included in E. and C,
Berkeley, 1952a; a label added to the type material indicates that it
was considered later by the Berkeleys to be the young of Notomastus
tenuis Moore.

Notomastus variegatus E. and C. Berkeley
Notomastus pallidior.—E. and C. Berkeley, 1942, p. 198. [Not Chamberlin, 1918.]
Notomastus variegatus E. and C. Berkeley, 1950, p. 59; 1952a, p. 103.
Type material: Syntypes, USNM 32677 and 32678, Estevan
Point, west coast Vancouver Island, 45 fms., E. G. Hart, collector,
11 specimens (17 pieces).

Family Arenicolidae

Arenicola glasselli E. and C. Berkeley
Arenicola glasselli E. and C. Berkeley, 1939, p. 340—Wells, 1962, p. 344.

Type material: Holotype, USNM_ 32835, San Felipe, Gulf of
California, littoral, June 18, 1936,S. A. Glassell, collector. Paratypes,
USNM 32844, San Felipe, Gulf of California, littoral, June 18, 1936,
S, A. Glassell, collector, 6 specimens. Paratypes, BMNH 1938.
11.29.34/37, San Felipe, Gulf of California.

Protocapitella E. and C. Berkeley

Protocapitella simplex E. and C. Berkeley
Protocapitella simplex E. and C. Berkeley, 1932a, p. 669, figs. 1-4.
Branchiomaldane vincenti Langerhans.—E. and C. Berkeley, 1950, p. 60; 1952a,
p. 99, figs. 203-205.

Type material: Holotype, USNM 32808, Wreck Bay, west coast
Vancouver Island, British Columbia, 1921 (specimen on slide).
Paratypes, BMNH 1938. 11.29.33, Wreck Bay, west coast Van-
couver Island, British Columbia.

Family Maldanidae

Axiothella rubrocincta var. complexa E. and C. Berkeley
Aziothella rubrocincta (Johnson) var. complexa E. and C. Berkeley, 1941, p. 49.
Type material: Holotype, USNM 32818, Newport Bay, southern
California, mud flats, November 1933, G. E. MacGinitie, collector.
Paratypes, USNM 32819, Newport Bay, southern California, mud

16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 119

flats, November 1933, G. E. MacGinitie, collector, 2 specimens (in
3 pieces). Paratypes, USNM 32820, Newport Bay, southern Cali-
fornia, mud flats, November 1933, G. E. MacGinitie, collector, 4
specimens (in 6 pieces).

Clymene (Euclymene) grossa var. newporti E. and C. Berkeley

Clymene (Euclymene) grossa Baird var. newporti E. and C. Berkeley, 1941, p. 49;
1960a, p. 360.

Euclymene grossa (Baird) var. newporti (E. and C. Berkeley).—Hartman, 1959,
pp. 453, 456.

Type material: Holotype, USNM 32826, Newport Bay, southern
California, Nov. 12, 1936, G. E. MacGinitie, collector. Paratype,
USNM 32827, LaJolla, southern California, Jan. 13, 1938, rocky
shore with coarse sand beneath, G. E. MacGinitie, collector, 1 speci-
men (anterior and posterior ends).

Clymene (Euclymene) papillata E. and C. Berkeley
Clymene (Euclymene) papillata KE. and C. Berkeley, 1939, p. 340, figs. 11, 12.
Euclymene papillata (E. and C. Berkeley).—Hartman, 1959, pp. 453, 457.
Type material: Holotype, USNM 32839, Punta Pefiasco, Sonora,
Mexico, S. A. Glassell, collector (specimen in 2 pieces, anterior and
posterior ends). Paratype, BMNH 1938. 11.29.38, Punta Pefiasco,

Sonora, Mexico.
Leiochone columbiana E. Berkeley

Leiochone columbiana E. Berkeley, 1929, p. 315, pl. 1, figs. 1-9.—E. and C.
Berkeley, 1952a, p. 44, figs. 81-84.
Clymenura columbiana (E. Berkeley).—Hartman, 1959, pp. 456, 458.
Type material: Holotype, USNM 32837, Departure Bay, Nanaimo,
British Columbia, 8-10 fms. Paratype, BMNH 1938. 11.29.42,
Departure Bay, Nanaimo, British Columbia.

Praxillella affinis var. pacifica E. Berkeley
Prazillella affinis (Sars) var. pacifica E. Berkeley, 1929, p. 313.—E. and C.
Berkeley, 1942, p. 198; 1952a, p. 49, figs. 97-100.

Type material: Syntypes, USNM 32832 and 32833, Nanoose
Bay, British Columbia, Aug. 28, 1918, 10-15 fms., 4 specimens (1
complete, 3 anterior ends—1 with regenerating head, 1 middle piece,
2 posterior ends).

Family Ampharetidae

Pseudosabellides E. and C. Berkeley
Pseudosabellides lineata E. and C. Berkeley

Sabellides octocirrata.—E. and C. Berkeley, 1942, p. 201. (Not Sars, 1835.)
Pseudosabellides lineata E. and C. Berkeley, 1943, p. 131; 1944, p. 3; 1952a, p.
71, fig. 147.

NO. 3553 POLYCHAETES—PETTIBONE 17

Asabellides stbirica (Wirén).—Pettibone, 1954, p. 318, fig. 36, e.
Asabellides lineata (H. and C. Berkeley).—E. and C. Berkeley, 1956c, p. 241.

Type material: Syntypes, USNM 32809 and 32813, Nanoose Bay,
British Columbia, 25 fms., Aug. 28, 1918, 3 specimens. Syntypes,
USNM 32812, Nanoose Bay, British Columbia, sponge bed, Jan.
11, 1931, 3 specimens. Syntypes, USNM 32811, Port Burwell,
Ungava, Hudson Bay, 10-15 fms., sandy mud, Sept. 16, 1927, 2
specimens. Syntypes, USNM 32810, Nash Harbour, Nunivak
Island and Spiriden Bay, Kodiak Island, Alaska, 8-12 fms., 7
specimens.

Pseudosabellides littoralis EK. and C. Berkeley

Pseudosabellides littoralis E. and C. Berkeley, 1943, p. 131; 1952a, p. 72.

Type material: Holotype, USNM 32817, Departure Bay, Nanaimo,
British Columbia, littoral, Apr. 21, 1929. Paratypes, USNM 32815,
Departure Bay, Nanaimo, British Columbia, littoral, Apr. 21, 1929
and Apr. 5, 1934, 3 specimens. Paratypes, USNM 32816, Portage
Lagoon, Nanaimo, British Columbia, littoral, May 4, 1935, 9 speci-
mens. Paratypes, USNM 32814, Diana Bay, Hudson Bay, littoral,
Aug. 4, 1928, 18 specimens.

Remarks: Pseudosabellides littoralis should be referred to Asabellides
Annenkova, thus becoming Asabellides littoralis (EK. and C. Berkeley).

Family Terebellidae

Novebranchus E, and C. Berkeley
Novobranchus pacificus E. and C. Berkeley
Novobranchus pacificus E. and C. Berkeley, 1954a, p. 465, figs. 8-13.

Type material: Holotype, USNM 32838, Buccaneer Bay, British
Columbia, 25 fms., Sept. 20, 1918.

Pista moorei E. and C. Berkeley

Pista moorei E. and C. Berkeley, 1942, p. 204, figs. 4, 5; 1952a, p. 81, fig. 166.

Type material: Holotype, USNM 32925, Trincomali Channel,
British Columbia, 25-30 fms., August 1934. Paratypes, USNM
32824, Malaspina Channel, Pender Island, British Columbia, 25 fms.,
Jan. 1, 1929, 2 specimens. Paratype, USNM 32822, Departure Bay,
British Columbia, July 11, 1940. G. E. MacGinitie, collector, 1
specimen.

Pista pacifica E. and C. Berkeley

Pista elongata.—E. and C. Berkeley, 1935, p. 773. (Not Moore, 1909.)
Pista pacifica E. and C. Berkeley, 1942, p. 202, figs. 1-3; 1952a, p. 80, figs.
163-165.—Hartman, 1944b, p. 273, pl. 22, figs. 37-41, pl. 25, figs. 61, 62.
Type material: Syntypes, USNM 32841 and 32843, west coast
Vancouver Island, British Columbia, June 27, 1934, 5 specimens

18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 119

(anterior ends and portion of tube). Syntypes, USNM 32834,
Elkhorn Slough, southern California, G. E. MacGinitie, collector, 2
specimens (anterior ends and middle piece).

Scionella estevanica E. and C. Berkeley

Scionella estevanica BH. and C. Berkeley, 1942, p. 205; 1943, p. 130; 1952a, p. 87.

Type material: Holotype, USNM 32831, Estevan Point, west coast
Vancouver Island, 76 fms., 1934, E. G. Hart, collector (anterior end).
Paratype, USNM 32820, Estevan Point, west coast Vancouver
Island, 76 fms., 1934, E. G. Hart, collector (anterior fragment, gills
missing).

Family Sabellidae

Branchiomma burrardum E. Berkeley

Branchiomma burrardum E. Berkeley, 1930, p. 71, fig. 1, A-E.—E. and C.
Berkeley, 1941, p. 55; 1942, p. 201; 1952a, p. 110, figs. 227-229.
Megalomma splendida (Moore).—Hartman, 1956, p. 298.

Type material: Holotype, USNM 32828, Burrard Inlet, British
Columbia, dredged, Sept. 15, 1928 (specimen in two pieces). Para-
types, USNM 32829, off Five Fingers, British Columbia, taken on
setline, June 5, 1929, 2 specimens. Paratypes, BMNH 1935.
12.19.106, Dodd’s Narrows, Vancouver Island, British Columbia.

Distylia volutacornis var. pacifica E. and C. Berkeley

Distylia volutacornis (Montagu) var. pacifica E. and C. Berkeley, 1954a, p. 468,
fig. 16.

Bispira volutacornis (Montagu) var. pacifica (E. and C. Berkeley).—Hartman,
1959, pp. 537, 542.

Type material: Holotype, USNM 32840, off Clarke Rock, Gulf of
Georgia (specimen in 2 pieces, with tube).

Fabricia minuta E. and C. Berkeley

Fabricia minuta E. and C. Berkeley, 1932b, p. 315, fig. 1, A-E; 1952a, p. 120,
figs. 245, 246.

Oridia minuta (EK. and C. Berkeley).—Hartman, 1951, p. 390.

Oriopsis? minuta (EK. and C. Berkeley).—Banse, 1957, p. 88.

Type material: Syntypes, USNM 32806, Long Bay, west coast
Vancouver Island, British Columbia, among algae, 1931, 2 specimens
(on slide).

Fabricia pacifica E. and C. Berkeley

Fabricia sabella (Ehrenberg).—E. Berkeley, 1930, p. 73, fig. 1, H (part).
Fabricia pacifica EK. and C. Berkeley, 1950, p. 66; 1952a, p. 121, fig. 249.
Fabriciola berkeleyi Banse, 1956, p. 429.

Type material: Holotype, USNM 32807, Newcastle Island, Depar-
ture Bay, Nanaimo, British Columbia, on stones at low tide, Aug. 8,
1918 (specimen on slide). Paratypes, USNM 32836, Newcastle

NO. 3553 POLYCHAETES—PETTIBONE 19

Island, Departure Bay, Nanaimo, British Columbia, on stones at
low tide, Aug. 8, 1918, 7 specimens.

Publications on Polychaetes by Edith and/or Cyril Berkeley
with New Taxa Described
BERKELEY, EDITH
1923. Polychaetous annelids fromthe Nanaimo district, 1: Syllidae to Sigalio-
nidae. Contr. Canadian Biol.,n.s., vol. 1, pp. 203-218, 1 pl. [Odon-
tosyllis phosphorea var. nanaimoensis; Odontosyllis parva; Amblyo-
syllis lineata var. alba; Autolytus magnus, Malmgrenia nigralba;
Lepidasthenia longicirrata.]
1924a. Polychaetous annelids from the Nanaimo district, 2: Phyllodocidae
to Nereidae. Contr. Canadian Biol., n.s., vol. 2, pp. 285-294,
1pl. [Tomopteris elegans.]
1924b. On a new case of commensalism between echinoderm and annelid.
Canadian Field-Nat., vol. 38, p. 193.
1927a. Polychaetous annelids from the Nanaimo district, 3: Leodicidae to
Spionidae. Contr. Canadian Biol., n.s., vol. 3, pp. 407-422, 1 pl.
[Dorvillea pseudorubrovittata; Glycinde picta; Spio martinensis var.
pacifica; Prionospio multibranchiata; Paraprionospio tribranchiata;
Boccardia columbiana; Boccardia uncata; Polydora_ cardalia;
Polydora caeca var. magna.]
1927b. A new genus of Chaetopteridae from the N.E. Pacific: With some
remarks on allied genera. Proc. Zool. Soc. London, pp. 441-
445, 3 figs. [Leptochaetopterus; Leptochaetopterus potts?.]
1929. Polychaetous annelids from the Nanaimo district, 4: Chaetopteridae
to Maldanidae. Contr. Canadian Biol., n.s., vol. 4, pp. 305-316,
1 pl. [Tharyx multifilis var. parvus; Caulleriella viridis var. pacifica;
Notomastus pallidior var. parvus; Prazillella affinis var. pacifica;
Leiochone columbiana.|
1930. Polychaetous annelids from the Nanaimo district, 5: Ammocharidae
to Myzostomidae (Appendix: Some pelagic forms from the Straits
of Georgia and the west coast of Vancouver Island). Contr.
Canadian Biol., n.s., vol. 6, pp. 65-77, 8 figs. [Branchiomma bur-
rardum; Tomopteris renata (new name).]
1935. Swarming of Odontosyllis phosphorea Moore, and of other Polychaeta
near Nanaimo, B.C. Nature, vol. 136, p. 1029.
1936a. Plankton of the Bermuda oceanographic expeditions, 5: Notes on
Polychaeta. Zoologica, vol. 21, pp. 85-87.
1936b. Occurrence of Saccocirrus in western Canada. Nature, vol. 137,
p. 1075.
1961. Swarming of the polychaete Qdontosyllis phosphorea Moore, var.
nanaimoensis Berkeley, near Nanaimo, B.C. Nature, vol. 191,
p. 1321.
BERKELEY, Epiru, and BERKELEY, CYRIL
1932a. Some Capitellidae (Polychaeta) from the N.E. Pacific: With a
description of a new genus. Proc. Zool. Soc. London, pp. 669-675.
[Protocapitella; Protocapitella simplex; Heteromastus filobranchus.]
1932b. On a collection of littoral Polychaeta from the west coast of Van-
couver Island. Contr. Canadian Biol., n.s., vol. 7, pp. 309-318.
[Fabricia minuta.]
1935. Some notes on the polychaetous annelids of Elkhorn Slough, Mon-
terey Bay, California. Amer. Midl. Nat., vol. 16, pp. 766-775.

20

PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 119

1936. Notes on Polychaeta from the coast of western Canada, 1: Spionidae.
Ann. Mag. Nat. Hist., ser. 10, vol. 18, pp. 468-477, 1 fig. [Polydora
soctalis plena; Polydora magna.]|

1938a. Notes on Polychaeta from the coast of western Canada, 2: Syllidae.

Ann. Mag. Nat. Hist., ser. 11, vol. 1, pp. 33-49, 12 figs. [Syllis
pulchra; Syllis harti; Syllis spenceri; Exogone lourei.]

1938b. Rhamphobrachium longisetosum, n. sp., with some observations on

the regeneration of the specialized anterior setae. Ann. Mag.
Nat. Hist., ser. 11, vol. 1, pp. 428-435, 8 figs. [Rhamphobrachium
longisetosum.|
1939. On a collection of Polychaeta, chiefly from the west coast of Mexico.
Ann. Mag. Nat. Hist., ser. 11, vol. 3, pp. 321-346, 12 figs. [Sigalion
lewisii; Eulepethus mexicanus; Onuphis zebra; Arenicola glasselli;
Clymene (Euclymene) papillata.]
1941. On a collection of Polychaeta from southern California. Bull.
Southern California Acad. Sci., vol. 40, pp. 16-60, 18 figs. [Sthene-
lanella atypica; Loandalia fauveli; Onuphis eremita var. parva;
Lumbrinereis ligulata; Aricia macginitii; Mesochaetopterus rickettsii;
Chaetozone spinosa var. corona; Clymene (Euclymene) grossa var.
newportt, Axiothella rubrocincta var. compleza.]
1942. North Pacific Polychaeta, chiefly from the west coast of Vancouver
Island, Alaska, and Bering Sea. Canadian Journ. Res., sec. D,
vol. 20, pp. 183-208. [Syllis stewarti; Pista pacifica; Pista moore;
Scionella estevanica.]
1943. Biological and oceanographical conditions in Hudson Bay, 11:
Polychaeta from Hudson Bay. Journ. Fish. Res. Bd. Canada,
vol. 6, pp. 129-132. [Pseudosabellides; Pseudosabellides littoralis;
Pseudosabellides lineata.]
1944. Polychaeta from the western Canadian Arctic. Canadian Journ.
Res., sec. D, vol. 22, pp. 1-5, 1 pl. [Sphaerodorum biserialis.]
1945. Notes on Polychaeta from the coast of western Canada, 3: Further
notes on Syllidae and some observations on other Polychaeta
errantia. Ann. Mag. Nat. Hist., ser. 11, vol. 12, pp. 316-335,
6 figs. [Autolytus trilineatus; Nephthys cornuta; Lumbrinereis luti.]
1948. Canadian Pacific fauna, 9; Annelida; 9b: (1) Polychaeta Errantia.
Fish. Res. Bd. Canada, pp. 1-100, 160 figs.
1950. Notes on Polychaeta from the coast of western Canada, 4: Poly-
chaeta Sedentaria. Ann. Mag. Nat. Hist., ser. 12, vol. 3, pp. 50-69,
8 figs. [Polydora ciliata var. spongicola; Aricidea longicornuta;
Stylarioides negligens; Notomastus variegatus; Fabricia pacifica.]
1951. A second record of the polychaetous annelid Potamethus elongatus
(Treadwell). Journ. Washington Acad. Sci., vol. 41, pp. 333-334,
4 figs.
1952a. Canadian Pacific fauna, 9: Annelida; 9b: (2) Polychaeta Sedentaria.
Fish. Res. Bd. Canada, pp. 1-139, 292 figs.

1952b. Re-discovery of the polychaete worm T'rypanosyllis ingens Johnson.
Journ. Fish. Res. Bd. Canada, vol. 8, pp. 488-490, fig. 1.

1953a. Micronereis nanaimoensis sp. n.: With some notes on its life-history.
Journ. Fish. Res. Bd. Canada, vol. 10, pp. 85-95, pls. 1, 2. [Micro-
nereis nanaimoensis.|

1953b. Swarming of Nereis succinea (Leuckart) off the east coast of Canada.

Nature, vol. 171, p. 847.

NO. 3553 POLYCHAETES—PETTIBONE 21

1954a. Additions to the polychaete fauna of Canada, with comments on
some older records. Journ. Fish. Res. Bd. Canada, vol. 11,
pp. 454-471, 16 figs. [Nereis (Hunereis) wailesi; Spio butleri;

Neopygospio; Neopygospio laminifera; Novobranchus; Novobranchus
pacificus; Distylia volutacornis pacifica.]
1954b. Notes on the life-history of the polychaete Dodecaceria fewkesi
(nom. n.). Journ. Fish. Res. Bd. Canada, vol. 11, pp. 326-334,
7 figs. [Dodecaceria fewkesi (new name).]

1956a. A new species and two new records of Polychaeta from eastern
Canada. Canadian Journ. Zool., vol. 34, pp. 267-271, 3 figs.
{Nereis (Ceratonereis) scotiae.]
1956b. Notes on Polychaeta from the east coast of Vancouver Island and
from adjacent waters, with a description of a new species of
Aricidea. Journ. Fish. Res. Bd. Canada, vol. 13, pp. 541-546,
6 figs. [Aricidea lopezi.]

1956c. On a collection of polychaetous annelids from the northern Banks
Island, from the south Beaufort Sea, and from northwest Alaska;
together with some new records from the east coast of Canada.
Journ. Fish. Res. Bd. Canada, vol. 13, pp. 233-246.

1957. On some pelagic Polychaeta from the northeast Pacific north of
latitude 40° N. and east of longitude 175° W. Canadian Journ.

Zool., vol. 35, pp. 5738-578, 2 figs.
1958a. Some notes on a collection of Polychaeta from the northeast Pacific
south of latitude 32° N. Canadian Journ. Zool., vol. 36, pp.
399-407.

1958b. Polychaeta of the western Canadian Arctic. Journ. Fish Res. Bd.
Canada, vol. 15, pp. 801-804.

1960a. Notes on some Polychaeta from the west coast of Mexico, Panama,
and California. Canadian Journ. Zool., vol. 38, pp. 357-362.
[Hurythoe complanata var. mexicana.]

1960b. Some further records of pelagic Polychaeta from the northeast
Pacific north of latitude 40° N. and east of longitude 175° W.,
together with records of Siphonophora, Mollusca, and Tunicata
from the same region. Canadian Journ. Zool., vol. 38, pp.
787-799.

1961. Notes on Polychaeta from California to Peru. Canadian Journ.
Zool., vol. 39, pp. 655-664, 12 figs. [Nereis (Neanthes) mancorae;
Ancistrosyllis longicirrata; Prionospio ornata.]

1962. Polychaeta from British Columbia; with a note on some western
Canadian Arctic forms. Canadian Journ. Zool., vol. 40, pp. 571-
577, 2 figs.

1963a. The proboscis of Lycastopsis catarractarum Feuerborn. Canadian

Journ. Zool., vol. 41, pp. 907-908, 1 fig.
1963b. Neoteny in larvae of two species of Spionidae. Canadian Journ.
Zool., vol. 41, pp. 149-151.

1964. Notes on some pelagic and some swarming Polychaeta taken off
the coast of Peru. Canadian Journ. Zool., vol. 42, pp. 121-134,
2 figs. [Nicon peruviana.]

BERKELEY, CyrRIL

1942. Occurrence of Ctenodrilus in the Pacific. Nature, vol. 149, p. 248.

1956. Epidiopatra hupferiana Augener from the northeast Pacific. Nature,
vol. 178, p. 748.

apd PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 119

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1919. The Annelida Polychaeta. Mem. Mus. Comp. Harvard, vol. 48, pp.

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NO. 3553 POLYCHAETES—PETTIBONE 23

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1964. The polychaetous annelids of Japan, pts. 1 and 2. Allan Hancock

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