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BULLETIN
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MUSEUM OF COMPARATIVE ZOOLOGY
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HARVARD COLLEGE, IN CAMBRIDGE
VOL. 116
CAMBRIDGE, MASS., U. S. A.
1957
TnK Cosmos Press, Inc.
Cambkidiif,, Mass., U.S.A.
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CONTENTS
PACE
No. 1. — Revision of the Chinese Mecoptera. By Fung:
Ying Cheng. (23 plates) March, 1957 ... 1
No. 2. — A Catalogue op the Cerionidae (Mollusca-
Pulmonata). By William J. Clench. April, 1957 119
No. 3. — Studies on New Zealand Elasmobranchii. Part
VI. Two New Species of Etmopterus from New
Zealand. By J. A. F. Garriek. April, 1957 . . 169
No. 4. — Biological Investigations in the Selva Lacan-
dona, Chiapas, Mexico. Raymond A. Paynter, Jr.,
Editor. (1 plate) April, 1957 191
No. 5. — The Genus Tetragnatha (Araneae, Argiopidae)
in Panama. By Arthur M. Chickering. May, 1957 299
No. (i. — The Tenuis and Selenopiiora Groups of the Ant
Genus Ponera (Hymenoptera: Formicidae). By
Edward 0. Wilson. May, 1957 .... 353
No. 7. — The Chinese Caeneressa Species ( Lepidoptera,
Ctenuchidae). By Nicholas S. Obraztsov. (4
plates) June, 1957* 387
No. <S. — A Comparative Morphological Study of the
Proventriculus of Ants (Hymenoptera: Formi-
cidae). By Thomas Eisner. (25 plates) July, 1957 437
No. 9. — The Ixodes Rasts Group of African Ticks with
Descriptions of Four New Species (Ixodoidea,
Ixodidae). By Don. R. Arthur and Colin Burrow.
July, 1957 ' 491
Bulletin of the Museum of Comparative Zoology
A T II A R V A R D COLLEGE
Vol. 116, No. 1
REVISION OF THE CHINESE MECOPTEKA
By Fung Ying Cheng
Harvard University and
National Taiwan University
With Twexty-thkf.e Plates
CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM
March 27, 1957
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Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE
Vol. 116, No. 1
REVISION OF THE CHINESE MECOPTERA
By Fung Ying Cheng
Harvard University and
National Taiwan University
With Twenty-three Plates
CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM
March 27, 1957
No. 1 — Revision of the Chinese Mecoptera
By Fung Yinq Cheng
INTRODUCTION
The Mecoptera, commonly called scorpion flies and hanging
flies, are widely distributed over the Northern Hemisphere and
are well represented in China. However, little collecting of
these insects has actually been done in China, especially in
the interior regions. It was my good fortune to be a member of
the entomological expedition of the Sino-British Committee,
extending from June 1939 to July 1940; this was mainly re-
stricted to Szechwan, Sikang and Shensi Provinces. The present
study is based largely on specimens collected on this expedition.
Since I have been able to examine type specimens in the Museum
of Comparative Zoology, the U. S. National Museum and the
Heude Museum, and also material in the Museum of the Institute
of Zoology, Academia Sinica, Museum of Foochow University,
Chou Collection, Issiki Collection and Maa Collection, I have
included an account of all species described from China.
I wish to express my sincere thanks to all the curators of
these museums and the owners of private collections for the
loan and gift of material and for their kind cooperation which
has made this study possible : Dr. P. J. Darlington, Dr. J. C.
Bequaert, Museum of Comparative Zoology; Dr. A. B. Gurney,
U. S. National Museum ; Dr. Sieien H. Chen, Academia Sinica ;
Father de Cooman, Heude Museum ; Dr. Hsiu Fu Chao, Na-
tional Foochow University ; Dr. Chou, Chang-chia-kang ; Dr. S.
Issiki, Tokyo and Mr. Maa, Taipeh.
Furthermore, I am deeply indebted to Professor F. M. Car-
penter of Harvard University for his encouragement during
the course of my study and for his kindness in reading over
this paper.
HISTORICAL ACCOUNT
The order Mecoptera comprises about three hundred and
thirty described species from the entire world. Until recently,
only a few of them were known from China. In 1921, Esben-
BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Petersen recorded only eight species from there in his mono-
graphic revision of the order. hi 1037, Wu reported nineteen
species in his "Catalogus Insectorum Sinensium," fifteen species
belonging to Panorpidae and four species to Bittacidae. In 1938,
Carpenter listed twenty-eight species of Panorpidae. In the past
ten years, more new species have been described by Carpenter,
Tjeder and myself. The number of described species has thus
been brought to fifty-seven. In the various collections which I
have examined recently, twenty-five additional species have been
found, bringing the total number of known Chinese species to
eighty-two.
Because of its geographical nature, the mainland of China
has been separated into thirty provinces. Scorpion flies and
hanging flies occur in all those (about half) to which expeditions
have been made. Since previous expeditions did not include most
of the mountainous areas of any of the provinces and no expedi-
tions at all have been made for Mecoptera in fifteen of the pro-
vinces, I believe our collections and knowledge of this order in
China are still very incomplete.
SYSTEMATIC ACCOUNT
Of the five existing families of Mecoptera, only two have been
found in China. They can be recognized as follows :
Tarsi with two claws, not raptorial Panorpidae
Tarsi with a single claw, modified for raptorial. . . .Bittacidae
Family PANORPIDAE
Panorpidae Stephens, 1836, 111. Brit. Ent., Enderlein, 1910, Zool. Anz., 25:
385. Esben-Petersen, 1921, Coll. Zool. Selys L'ong., 5(2) : 11. Carpenter,
1931, Bull. Mus. Conip. Zool., 72(6): 209.
Ocelli present; labial palpi two-segmented; abdomen cylin-
drical or nearly so, tapering to a point in the female, without
ovipositor ; 6th to 8th abdominal segments of male narrowly cylin-
drical, apical segment modified, usually enlarged ; tarsi with two
(daws, not modified for grasping ; wings more or less slender,
with primitive venation ; costal space narrow, with few cross-
veins ; Pts originating at about one-third of wing length from
base ; M dividing near the middle of wing.
CHENG : REVISION OF THE CHINESE MECOPTERA S
Three of the six existing genera of the family inhabit China,
namely, Panorpa, Neopanorpa and Leptopanorpa; the fourth
genus, Panorpodcs occurs in Japan, the fifth Apteropanorpa in
Tasmania, and the sixth, Brachypanorpa in North America. Both
Fig. 1. Wing venation and markings of Panorpa: 1A, first anal vein; apb,
apical band; bb, basal band; bs, basal spot ; ms, marginal spot; Ors, origin
of radial seetor; ptb, pterostigmal band.
Panorpa and Neopanorpa were found in most of the Chinese pro-
vinces where expeditions have been made. Leptopanorpa is repre-
sented by a single species found in Hainan, Kwangtung. As a
matter of fact, however, the latter locality is an island, separated
from the mainland bv the sea. The three genera occurring in
A B
Fig. 2. The 6th to 9th abdominal segments of Panorpa: A, kimminsi
Carpenter (diceras group) ; B, obtvsa Cheng (centralis group) ; C, el ado-
rer ca Navas (davidi group).
4 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
China can be distinguished as follows :
1. 1A joins the anal margin of wing far beyond the origin of radial sector
(text-fig. 1) Panorpa
1A joins the anal margin of wing before the origin of radial sector
(text-fig. 3) 2
2. Abdomen in the male only as long as the wings or shorter; 6th to 9th
abdominal segments normal or only a little prolongated (text-fig.
4) Neopanorpa
Abdomen in the male much longer than the wings ; 6th to 9th segments
much prolongated (text-fig. 5) Leptopanorpa
Genus PANORPA Linne
Panorpa Linne, 1758, Syst. Nat., 10:551. Klug, 1836, Abh. Konigl. Akad.
Wiss. Berl., 1836:88. Eambur, 1842, Hist. Nat. Ins. Nevr., 1842:328.
Westwood, 1846, Trans. Ent. Soc. London, 4:184. Loew, 1848, Linn. Ent,,
3:363. Brauer and Low, 1857, Neuropt. Austr., 1857:35. Brauer, 1863,
Verh. Zool.-bot. Ges. Wien, 13:307. Id., 1871, Verh. Zool.-bot. Ges. Wien.
21:109. McLachlan, 1868, Trans. Ent, Soc. London, 1868:209. Klapalek,
1896, Rozp. Ceske Akad. Cisare Frantis. Josef a, 1896:1. Felt, 1896, New
York State Ent. Eep., 10:463. McClendon, 1906, Ent. News, 1906:121,
fig. 14. Stitz, 1908, Zool. Jahrb., 26:537. Mjoberg, 1909, Ent. Tidskr.,
1909:160. Enderlein, 1910, Zool. Anz., 35:389. Miyake, 1912, Journ. Coll.
Agric. Imp. Univ. Tokyo, 4:137. Id., 1913, Journ. Coll. Agric. Imp. Univ.
Tokyo, 4:335. Banks, 1913, Trans. Am. Ent. Soc. 1913:232. Esben
Petersen, 1921, Coll. Zool. Selys Long., 5(2) :13. Carpenter. 1931, Bull.
Mus. Comp. Zool., 72(6) :213.
Aulops Enderlein, 1910, Zool. Anz., 35:390.
Estenella Navas, 1912, Rev. Russe d'Ent., 12:356.
Rostrum long and slender; tarsal claws serrated on inner
margins; wings are fully developed, rather broad, 1A long, ex-
tending to the anal margin of wing beyond origin of the radial
sector; abdomen in both sexes not longer than the wings. 6th
to 9th abdominal segments of male usual, not much prolongated ;
genital bulb of male not pedunculate basally.
Genotype: Panorpa communis Linne.
This is the largest genus of Mecoptera, including one hundred
and sixty-two known species in the whole world. Thirty species
have been already recorded in China and ten new ones are de-
scribed below, making a total of forty. They are distributed
throughout eleven provinces. As suggested by Carpenter (1938),
these species can be conveniently divided into three groups, de-
CHENG : REVISION OP THE CHINESE MECOPTERA 5
pending upon the structure of the 6th abdominal segment of
the male — that is, whether the anal horn is present, absent, or
represented by two similar processes. In the first or diceras
group, with double anal horn, belong diceras McLachlan, tjederi
Carpenter, stotzneri Esben-Petersen and kimminsi Carpenter ; in
the second or centralis group, with a single anal horn, belong
centralis Tjeder, flavipennis Carpenter, emarginata Cheng and
obtusa Cheng; in the third or davidi group, without anal horn,
belong davidi Navas, stigmalis Navas, waongkehzengi Navas,
difficilis Carpenter, fructa Cheng, cladocerca Navas, trifasciata
Fig. 3. Wing venation of Neopanorpa: 1A, first anal vein; Ors, origin
of radial sector.
n. sp., ftavicorporis n. sp., cheni n. sp., obliqua Carpenter, baoh-
washana n. sp., typicoides Cheng, fukiensis Tjeder, curva Car-
penter, aurea n. sp., coomani n. sp., japonica Thunberg, tetrazonia
Navas, sexspinosa Cheng, tincta Navas. Other species, implicata
n. sp., lutea Carpenter, klappcrichi Tjeder, semifasciata Cheng,
leei Cheng, grahamana n. sp., carpenteri n. sp., statura Cheng,
pusilla Cheng, pieli n. sp. and bonis Cheng, are known only from
the female, so that their position in the above grouping is not
determinable.
Other characteristics which have been used under each group
of Panorpa are the wing markings and the general aspect of
both male and female genitalia. The former characteristic some-
times appears to be similar from one species to another and in
some cases passes through a little range of variation, while
the latter remain perfectly stable even in minute details. In
identification, therefore, I use the wing markings to assist in
the first determination of the species and then the characteristics
of both male and female genitalia for the final decision.
b BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
In the wings of Panorpa, the color of the membrane is a con-
venient feature for general classification. In some species, the
wings are clear and colorless (kimminsi, fig. 271), in others they
are distinctly yellow (flavipennis), and in some others, they are
faintly yellow (aurea, fig. 281). The markings of the wings are
readily grouped into several categories, depending upon how
heavy the markings are. In some species, in which the wing is
Fig. 4. Male abdominal segments of Neopanorpa tienmushana n. sp.
heavily marked (japonica, fig. 286), there are three transverse
bands and two spots : these are designated the pterostigmal band,
the basal band, the apical band, the basal spot and the marginal
spot (text-fig. 1). In other species the wing is lightly marked,
the bands are very narrow, the basal spot and the marginal spot
are usually absent, and sometimes both pterostigmal and apical
Fig. 5. Male abdominal segments of Leptopanorpa javanica (Westwood).
CHENG: REVISION OF THE CHINESE MECOPTERA 7
bands are interrupted or incomplete (seniifasciata, fig. 274). In
some other species, no markings are present, except for a slight
suspicion of shadow at the wing apex (obtusa, fig. 272) . Curiously
enough, all species bearing the anal horn belong to the latter
category, the only exception being flavipennis, which has a dis-
tinctly yellow wing membrane and ver}r narrow basal, pterostig-
mal and apical bands. It is interesting to note that the single
anal horn of this species is exceptionally short (fig. 13), quite
different from the other single or double horn species. As worked
out by Carpenter in his "Revision of the Nearctic Mecoptera''
(1931), the venational characteristics of Panorpa are purely
individualistic and could not be used for the classification of the
species. The same statement applies to all the Chinese Panorpa
from Pukien — the only locality from which sufficient specimens
have been collected to allow a conclusion.
The male genital segment (or 9th segment), modified into a
bulb, consists of an upper tergite (preepiproct), a lower sternite
(hypandrium) and a pair of two-segmented structures in be-
tween, the basal U-shaped coxopodites and the distal hooked
harpagones. In some species there is a papilla at the inner distal
margin of each of the coxopodites (carva, fig. 52, davidi, fig. 57).
and in others a number of black spines at the same place (sex-
spinosa, fig. 87, centralis, fig. 10). The harpagones vary much
in shape and degree of development; in most species there is a
concave area on the inner basal surface of each harpagone (cen-
tralis, fig. 10, davidi, fig. 57). In others, instead of a concave
area, there is an expansion or lobe at the same place. This lobe
may be very small (japonica, fig. 90), greatly enlarged (baoh-
washana n. sp., fig. 83), or tooth-shaped (diceras, fig. 7), or it
may arise at a different level (trifasciata n. sp., fig. 49, cheni n.
sp., fig. 86). The preepiproct is visible as a single external ap-
pendage from a dorsal aspect of the bulb. In most of the species,
it has a pair of distal lobes (diceras etc., fig. 2), but in waong-
kehzengi, the lobes are so inconspicuous that the apex of the
preepiproct appears only slightly emarginated (fig. 43), and in
japonica, no lobes occur at all, the preepiproct terminating as
a rounded posterior margin (fig. 85). The hypandrium appears
as a basal plate from a ventral view of the bulb ; this is usually
produced into two long lobes, the hypovalvae. In some species
8 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
the hypandrium is long and stalk-like {japonica, fig. 90, stigmalis,
fig. 33) ; in many others, it is inconspicuous {emarginata etc.,
fig. 31). The hypovalvae are generally slender and flattened
{flavipennis, fig. 11), while in baohwashana n. sp., they are
broadened towards the apex with upwardly folded outer margins
(fig. 83). In the interior of the bulb, between the bases of the
coxopodites arises an irregular aedeagus from the wall of the
genital pouch. This usually has a pair of prominent apical proc-
esses and a pair of lateral processes. In some species, the apical
processes are very short and inconspicuous {tjederi, fig. 6) ; in
others, they are very long {curva, fig. 52), and in some others
they have greatly enlarged distal ends {difficilis, fig. 46). At
PCV
Fig. 6. Wing venation of Bittacus : 1A, first anal vein; Av, anal cross
vein ; Cuv, cubital crossvein ; Ors, origin of radial sector ; Pcv, pterostigmal
crossvein ; Scv, subcostal crossvein.
the sides of aedeagus are two slender parameres supported on
a U-shaped bar in the ventral wall of the genital chamber proxi-
mal to the base of the aedeagus. The parameres exhibit a re-
markable series of modifications in different species. They are
strongly sclerotized and very stiff, usually blackish brown in
color and with a cluster of spines, resembling the barbs in a
bird's feather, on their inner margins and their distal ends. In
some species the parameres are reduced to a very minute linear
thread without spine {japonica, fig. 90), while in most others
they are long with prominent barbs {centralis group, fig. 10,
davidi etc., fig. 57). In some species, they are Y-shaped {aurea
n. sp., fig. 62), in others they have 3-4 branches {diceras group,
fig. 7). Curiously enough, even the extending direction and the
approximate number of the spines of this appendage are constant
in the species.
CHENG : REVISION OF THE CHINESE MECOPTERA 9
The female genital segment (or 9th segment) consists of an
upper normally developed tergite and a lower subgenital plate
which has a free posterior margin forming an opening for the
genital cavity. In some species, the subgenital plate is very
broad and its lateral margins overlap the sides of the 9th ter-
gite (tjederi, fig. 12) ; others are narrow and their lateral mar-
gins are enclosed by the well developed 9th tergite (semifasciata,
etc., fig. 122). The posterior margin of the subgenital plate
differs from species to species. In tetrazonia, etc., it is narrowed
towards the apex (fig. 93), in leei, etc., it is rounded (fig. 125),
in most of the species, it is slightly emarginated (kimminsi, fig.
16) and in japonica, it has a V-shaped distal incision. Inside
the genital cavity and just above the subgenital plate, there is
a highly sclerotized internal skeleton. The latter shows a desired
specific variability and individual constancy and has been used
for identification by several authors for the past ten years. This
plate is often supported by an inner pair of very dark pig-
mented rods or an axis. In most species, the axis is long and ex-
tends beyond the plate of the internal skeleton (flavipennis,
etc., fig. 23), in others, it is less developed and confined by the
plate (tjederi, etc., fig. 17), and in some others the axis is greatly
reduced or absent (japonica, etc., fig. 100). Curiously enough,
all the females of the diceras group have a small axis, i. e.,
slender and not extending beyond the plate of the internal
skeleton. The above mentioned features of both male and female
genital structures are constant in the individuals of each species,
and show simultaneous modifications in the different species.
They therefore serve as excellent characteristics for identifica-
tion. The following keys to both male and female Panorpa are
based mainlv on these characteristics mentioned.
mi
Key to the Males of Panorpa
The males of the following species are unknown : lutea Car-
penter, semifasciata Cheng, leei Cheng, statura Cheng, pusilla
Cheng, pieli n. sp., bonis Cheng, klapperichi Tjeder, implicata
n. sp., guttata Navas, grahamana n. sp., carpenteri n. sp.
1 . Sixth abdominal segment with anal horn 2
Sixth abdominal segment without anal horn (text-fig. 2, C)
(davidi group) 9
10 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
2. Sixth abdominal segment with double anal horn (text-fig. 2, A) ....
(diceras group ) 3
Sixth abdominal segment with single anal horn (text-fig. 2, B)
(centralis group) 6
3. Hypovalvae greatly broadened distally 4
Hypovalvae slightly broadened distally 5
4. Tips of hypovalvae rounded; proximal lobes of the harpagones pointed
( fig. 7 ) diceras
Tips of hypovalvae truncated; proximal lobes of the harpagones rounded
(fig. 8) stotzneri
5. Parameres four branched; preepiproct with only shallow distal incision
(figs. 1, 6) tjederi
Parameres three branched; preepiproct with wide U-shaped distal in-
cision (figs. 4, 9) kimminsi
6. Genital bulb rounded ; the outer margins of harpagones not concave
a t the middle 7
Genital bulb less rounded; the outer margins of harpagones slightly
concave at the middle 8
7. Coxopodites with a group of very strong black spine-like hairs on their
inner apical margins; parameres simple, broad basally, abruptly
tapering into a slender and acute distal portion (fig. 10) . . . .centralis
Coxopodites without strong black spine-like hairs on their inner apical
margins; parameres simple, stout, spindle-shaped (fig. 34) . . . .obtusa
8. Inner margins of harpagones without true lobes; hypovalvae rather
broad, with slightly concave median outer margins; parameres long,
with pointed apex, reaching to the distal part of harpagones (fig. 31)
emarginata
Inner margins of harpagones with true lobes ; hypovalvae very slender,
with even outer margins; parameres short, with blunt apex, reaching
only to the base of harpagones (fig. 11) flavipennis
9. Wing membrane deeply yellow 10
Wing membrane faintly yellow or hyaline 12
10. Wing markings distinct; basal band absent or represented by two
separated spots 11
Wing markings indistinct ; basal band well developed, extending from
subcostal to the anal margin of wing, broadened towards its posterior
end; 6th to 8th abdominal segments as in figure 78 tincta
1 1. Pterostigmal band absent; apical band broad, smoky brown; harpagones
long and slender; hypandrium conspicuous, slender (fig. 33)
stigmalis
Pterostigmal band present, with complete basal branch and separated
apical branch; apical band interrupted; harpagones short; hypan-
drium conspicuous, broad (figs. 48, 277) waongkeh.zengi
CHENG : REVISION OF THE CHINESE MECOPTERA 1 1
12. Pterostigmal band present but interrupted 13
Pterostigmal band present, not interrupted 14
13. Hypandrium rather conspicuous ; apical processes of aedeagus very long,
with greatly enlarged distal ends (fig. 46) difficilis
Hypandrium inconspicuous ; apical processes of aedeagus long, with
narrow tooth-like distal ends and double sinuous outer margins (fig.
38 ) fructa
14. Pterostigmal band with complete basal branch; apical branch absent
15
Pterostigmal band with both basal branch and apical branch 19
15. Basal band complete, not interrupted 16
Basal band interrupted, represented by two spots 17
16. Marginal spot present; inner margin of harpagones with a smooth
median angle, a basal true lobe and a very large basal concave area
(figs. 51, 284) oladocerca
Marginal spot absent ; inner margin of harpagones with a sharp mediaxi
angle and a basal lobe which does not arise at the same level as the
former (figs. 49, 283) trifasciata n. sp.
1 7. Marginal spot present, and prominent ; inner margin of harpagones
with a smooth median angle and a broad basal lobe (fig. 59)
flavicorporis n. sp.
Marginal spot greatly reduced or absent; inner margin of harpagones
with a sharp median angle and a basal lobe which arises at a different
level or no true basal lobe at all 18
18. Parameres branched basally, the outer branch long, broadened towards
apex; the inner branch short, with a bundle of long brown stiff hairs
(fig. 86) dheni n. sp.
Parameres simple, long and slender (fig. 57) davidi
19. Apical branch of pterostigmal band interrupted, separated from ptero-
stigmal band itself 20
Apical branch of pterostigmal band not interrupted, connected with
pterostigmal band itself 23
20. Apical branch of pterostigmal band connected with the apical band to
form a hyaline window which encloses a faintly brown spot; genital
bulb as in figure 45 obliqua
Apical branch of pterostigmal band not connected with the apical band
21
21. Harpagones very long and slender, inner margin with a broad basal
lobe ; hypovalvae narrow basally, very broad and robust distally ;
apex of preepiproct usually folded downward, with very minute distal
incision (figs. 83, 84) baohwashana n. sp.
Harpagones short, no true basal lobe present ; hypovalvae not broadened
distally; apex of preepiproct with very wide U-shaped distal incision
22
12 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
22. Parameres spindle-shaped distally ; inner margin of harpagones with a
sharp median tooth; apical processes of aedeagus simple (fig. 58)
fukiensis
Parameres long and slender ; inner margin of harpagones with a smooth
median angle; apical processes of aedeagus finger-shaped, their distal
inner margins produced inwards to form a small nipple-shaped plate
(figs. 36, 39) typieoides
23. Apical branch of pterostigmal band broad, nearly the same width as
the basal branch 24
Apical branch of pterostigmal band narrow, much narrower than the
basal branch 26
24. Parameres simple, not branched ; the distal inner margin of coxopodites
with a very prominent papilla (fig. 52) curva
Parameres Y-shaped; the distal inner margin of coxopodites without
a papilla 25
25. Wing membrane light yellow, markings yellowish brown; harpagones
with a long and large basal concave area; genital bulb slender, not
elliptical (fig. 62) aurea n. sp.
Wing membrane hyaline, markings sooty brown ; harpagones with no
concave area, but with median toothed basal lobe ; genital bulb ellipti-
cal (fig. 63) coomani n. sp.
26. Hypandrium conspicuous, appearing as a long narrow stalk; preepiproct
with rounded apex; harpagones long and slender; parameres very
small, short rod-like (figs. 85. 90) japonica
Hypandrium inconspicuous; preepiproct with distal incision; harpagones
not very long; parameres well developed 27
27. Inner margin of harpagones with prominent lobes ; distal inner margin
of coxopodites without prominent spines ; parameres widening
abruptly and giving rise to a long curved process which bears numer-
ous long barbs (fig. 91) tetrazonia
Inner margin of harpagones without true lobes, but with basal concave
areas ; distal inner margin of coxopodites with six prominent spines ;
parameres narrow and slender, without barbs (fig. 87). . . .sexspinosa
Key to the Females of Panorpa
The females of the following species are unknown: diceras McLachlan,
obtusa Cheng, davidi Navas, curva Carpenter, difficilis Carpenter, fructa
Cheng, coomani n. sp., stigmalis Navas, tincta Navas.
1. Wing membrane deeply yellow 2
Wing membrane faintly yellow or hyaline or lacteous 5
2. Pterostigmal band complete, forked posteriorly 3
Pterostigmal band not complete, not forked posteriorly, apical branch
CHENG: REVISION OF THE CHINESE MECOPTEEA 13
of pterostigmal band absent; apical band broad, with a large window;
wing length less than 9 mm. (fig. 277) waonglcehzengi
3. Markings blackish brown, prominent; basal spot present; internal
skeleton with a very short axis (fig. 112) lutea
Markings gray or brown ; basal spot absent 4
4. Wing broad ; markings brown ; apical band large with a small hyaline
spot (fig. 279) statura
Wing narrow; markings gray; apical band reduced to a few faint spots
flavipennis
5. Pterostigmal band absent 6
Pterostigmal band present 9
6. Internal skeleton with short axis, not extending beyond the plate
7
Internal skeleton with long axis, usually extending beyond the plate
8
7. Subgenital plate not emarginated distally; internal skeleton as in figure
18, with somewhat converging posterior arms; wing membrane with
faintly yellowish tinge; tip of wing bordered with strong yellowish
brown stotzneri
Subgenital plate slightly emarginated distally; internal skeleton as in
figure 19, with somewhat diverging posterior arms; wing membrane
dusky hyaline; tip of wing bordered with grayish yellow (fig. 271)
Tcimminsi
8. Internal skeleton as in figure 24, axis extending beyond the plate one
half its length emarginata
Internal skeleton as in figure 20, axis extending beyond the plate less
than one half its length centralis
9. Pterostigmal band not prominent, represented only by a spot posterior
to the pterostigma 10
Pterostigmal band prominent 11
10. Wings subobtuse; apical band absent; pterostigma smoky; internal
skeleton as in figure 17, with posterior arms extending parallel to
each other tjederi
Wings narrow and slender ; apical band present, interrupted, represented
by three separated spots; pterostigma yellow guttata
11. Pterostigmal band with only basal branch 12
Pterostigmal band with both basal branch and apical branch 19
12. Basal branch of pterostigmal band narrow 13
Basal branch of pterostigmal band the same width as the pterostigmal
band itself 15
13. Both basal band and marginal spot present ; subgenital plate long and
slender; internal skeleton small, as in figure 69 flavicorporis
Both basal band and marginal spot absent 14
14 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
14. Basal branch of pterostigmal band interrupted; internal skeleton as in
figure 114 semifasciata
Basal branch of pterostigmal band not interrupted; internal skeleton
as in figure 127 leei,
15. Subgenital plate with less sclerotized lateral plates; both sides of the
subgenital plate enclosed by the lateral borders of the 9th tergite
16
Subgenital plate without lateral plates; both sides of the subgenital
plate are free 17
16. Rostrum brownish black in color, with an orange median longitudinal
stripe; the anterior arms of the internal skeleton slightly outwardly
curved as in figure 115 grdhamana n. sp.
Rostrum uniformly reddish brown, no median longitudinal stripe; the
anterior arms of the internal skeleton folded transversely at its median
portion as in figure 116 carpenteri n. sp.
17. Subgenital plate slender, narrow basally; internal skeleton as in figure
68 cladocerca
Subgenital plate somewhat triangular, broad basally 18
18. Subgenital plate with slightly emarginated apex; internal skeleton as
in figure 96 cheni n. sp.
Subgenital plate with rounded apex; internal skeleton as in figure 65
trifasoiata n. sp.
19. Apical branch of pterostigmal band interrupted, separated from ptero-
stigmal band itself 20
Apical branch of pterostigmal band not interrupted, connected with
pterostigmal band itself 27
20. Apical branch of pterostigmal band large, united with the apical band
to form one or two hyaline spots 21
Apical branch of pterostigmal band small, simple, separated from the
apical band 22
21. Wing membrane faintly yellow; subgenital plate with rounded posterior
margin; internal skeleton as in figure 98, with short axis . . . .ohliqua
Wing membrane hyaline, subgenital plate with slightly concave posterior
margin; internal skeleton as in figure 103, with very long axis. . . .
implicata n. sp.
22. Wing membrane light yellow; fore wing length less than 11 mm. . .23
Wing membrane hyaline ; fore wing length more than 12 mm 24
23. Wing markings sooty brown; internal skeleton with small plate as in
figure 118 pusilla
Wing markings gray; internal skeleton with large plate as in figure 117
pieli n. sp.
CHENG: REVISION OF THE CHINESE MECOPTERA 15
24. Fore wing length more than 18 mm.; subgenital plate with prominent
V-shaped distal incision; internal skeleton as in figure 105, with nearly
straight axis baohwas'hana n. sp.
Fore wing length less than 16 mm. ; subgenital plate with no prominent
V-shaped distal incision ; internal skeleton with outwardly curved axis
25
25. Internal skeleton with long axis, extending nearly two-thirds its length
beyond the plate as in figure 67 typicoides
Internal skeleton with short axis, extending not more than half its length
beyond the plate 26
26. Apical band prominent, large; basal band complete; apex of subgenital
plate not emarginated ; axis of internal skeleton simple, as in figure 71
fulciensis
Both apical band and basal band interrupted ; apex of subgenital plate
slightly emarginated ; distal ends of the axis of internal skeleton
branched as in figure 106 bonis
27. Apical branch of pterostigmal band broad, nearly the same width as
the basal branch 28
Apical branch of pterostigmal band narrow, much narrower than the
basal branch 29
28. Wing membrane light yellow, markings yellowish brown; internal skele-
ton large, as in figure 77, with long outwardly curved axis extending
beyond the plate one half its length aurea n. sp.
Wing membrane hyaline, markings sooty brown; internal skeleton small,
as in figures 120, 121, with very short axis not extending beyond the
plate lclapperichi
29. Internal skeleton as in figure 100, no axis present japonica
Internal skeleton with well developed axis 30
30. Wing membrane hyaline ; internal skeleton as in figure 124, posterior
arms short, pointed towards apex sexspinosa
Wing membrane with slightly yellowish tinge; internal skeleton as in
figure 94, posterior arms long, with rounded apex tetrazonia
Descriptions of Species of Panorpa
1'axorpa diceras McLachlan
Figures 2, 7
Panorpa diceras McLachlan, 1894, Ann. Mag. Nat. Hist., (6)13:423. Esben-
Petersen, 1921, Coll. Zool. Selys Long., 5(2) :36. Tjeder, 1936, Ark. for
Zool., 27 A (33) :9, pis. 4, 5, 7, figs. 4, 5. Carpenter, 1938, Proc. Ent.
Soc. Washington, 40(9): 270, figs. 2, 10, 11. Id., 1948, Psyche, 55(1):
28, fig. 1.
Panorpa grahami Carpenter, 1938, Proc. Ent. Soc. Washington. 40(9) :272.
figs. 5, 9.
1 6 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Body light brown ; vertex with a black spot around ocelli ;
rostrum with black stripe on each side ; thorax and abdomen with
a lateral black stripe ; sixth abdominal segment of male with two
short anal horns. Fore wing: length, 11 mm.; width, 3.5 mm.;
membrane hyaline without markings except for a slight suspicion
of gray at the apex ; neuration distinct, Sc extends to the ptero-
stigmal area, R2 is forked and R1? R3, R4, Rg are simple, usually
one crossvein occurring between Rj and R2. Hind wing: similar
to the fore wing. Male genitalia : genital bulb rounded ; coxopo-
dites rather long ; harpagones slender, the outer margin slightly
concave at the middle, the inner margin with a small distal lobe
and a larger pointed proximal lobe ; hypandrium inconspicuous,
hypovalvae long, reaching to the base of the harpagones, nar-
rowed proximally, much broadened distally, with rounded tips;
parameres complex, each consisting of a narrow basal stalk, which
gives rise to three branches, pectinately arranged, the basal
branch being the shortest, the others about equal ; preepiproct
with prominent distal lobes, the incision being almost quadrate.
Female unknown.
Type ( S ) : Tachienlu, Sikang ; in McLachlan Collection.
Distribution: Tachienlu, Sikang; 9 miles south of Tachienlu,
Sikang (8500-13,000 ft.), June 25-27, 1923 (D. C. Graham).
This species, which has double anal horns in the male, belongs
to the diceras group. I have not seen the type. According to
Carpenter's drawing, this species resembles stotzneri Esben-
Petersen in general appearance. However, the rounded tips of
the hypovalvae and the pointed proximal lobes of the harpagones
make its recognition easy.
Panorpa tjederi Carpenter
Figures 1, 6, 12, 14, 17
Panorpa tjederi Carpenter, 1938, Proc. Ent. Soc. Washington, 40(9) :271.
Panorpa diceras Tjeder (vec McLachlan), 1936, Ark. for Zool., 27 A (33) :9,
pis. 4, 5, pi. 7, figs, i, 5.
According to Tjeder, the body characters of the male of this
species agree wholly with McLachlan 's original description for
Panorpa diceras McLachlan. In the characteristics in size, color
of head, body and legs, and shape of wings, the 5 agrees per-
fectly with the 8 . The color of the wings is distinctly different.
CHENG : REVISION OP THE CHINESE MECOPTERA 17
Besides the smoky pterostigma, there is a subtriangular smoky
brown oblique spot, extending from the pterostigma down into
the cellula between Rg and M1 + 2 (just before the fork-point).
The additional side plates of the 7th to 8th abdominal segments
are wholly absent, the tergites are comparatively narrow while
the sternites are very broad (with their lateral margins reaching
up to the dorsal surface), and the pleural regions are very nar-
row and situated latero-dorsally. The detail drawings of both
$ and 9 genitalia were given by Tjeder. According to these
drawings, the genital bulb of $ genitalia is oval; coxopodites
rather long, U-shaped; harpagones very broad at the base but
strongly tapering into a very acute and hooked, curved apex, the
outer margin slightly concave at the middle, the inner margin
with a median small tooth-like process and a large basal tooth
directed inwards; hypandrium inconspicuous; hypovalvae slen-
der, slightly broadened and diverging towards each other in the
distal half, reaching to the base of the harpagones ; parameres
complex, each consisting of a narrow basal stalk, which gives
rise to four branches ; one of these arises inwardly from about
the middle of the stalk, the others arise further distally and
continue nearly straight ; preepiproct broad, narrowed towards
apex, which is broadly emarginated ; aedeagus smoothly emar-
ginated distally, both apical and lateral processes inconspicuous.
Female genitalia : the subgenital plate is very broad, and its
lateral margins overlap the sides of the 9th tergite ; at the hind
margin the plate appears broad and tongue-like, extending
through this tongue-like process farther backwards than the 9th
tergite ; internal skeleton very broad, posterior arms of the plate
blunt and stout, strongly folded, leaving between them proxi-
mally a smoothly rounded space ; axis short, lying wholly within
the plate and ending behind the basal flaps of the plate, which
are very large, and lie close to one another, with their apical ends
directed upwards.
Types ( $ , 9 ) : Yunnan (George Forrest) ; in K. J. Morton
Collection, Edinburgh.
Distribution : same as types.
This species belongs to the diceras group. I have not seen
the types. According to Tjeder 's drawings, it is closest to diceras
McLachlan in general appearance. However, the hypovalvae axe
18 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
not broadened towards the apex as in those of diceras. The para-
meres of this species are apparently four-branched, whereas
those of diceras are only three-branched. According to Tjeder,
the wings of the female of this species show great similarity to
those of Panorpa bicornuta McLachlan (from Japan), as figured
by Esben-Petersen, but the female genitalia of this species are
very different from those of bicornuta.
Panorpa stotzneri Esben-Petersen
Figures 8, 15, 18
Panorpa stotzneri Esben-Petersen, 1934, Vidensk. Medd. Dansk Naturli.
Foren., 97:211, figs. 1, 2. Tjeder, 1936, Ark. for Zool., 27 A (33) :11,
pi. 6.
Body mostly lemon-colored ; vertex with a black spot, enclos-
ing the ocelli ; rostrum with two brownish-black longitudinal
streaks ; thorax with a black longitudinal streak along each mar-
gin ; 1st to 5th abdominal segments of male lemon-colored, with
a black streak along each margin, last few abdominal segments
with indication of a darker color ; the hind border of third ter-
gite produced in a short lobe ; sixth segment somewhat swollen,
and its hind margin above produced in two club-shaped anal
horns; abdomen of female yellowish, with a black streak along
each side. Fore wing: length, 17 mm., membrane faintly yellow-
ish tinged, tip of the wings strongly bordered with yellowish
brown ; venation blackish brown, Sc reaching to the pterostigma.
which is prominent. Hind wing: length, 15 mm., similar to the
fore except that Sc reaches only half way to the pterostigma.
Male genitalia: according to Esben-Petersen 's drawing, the geni-
tal bulb is rounded ; coxopodites rather short, harpagones slen-
der, each with pointed and inwardly curved apex, the outer
margin very slightly concave at the middle, the inner margin
with a sharp median tooth and a rounded proximal lobe ; hypan-
drium inconspicuous; hypovalvae greatly broadened towards
truncated apex, and nearly reaching to the base of the harpa-
gones ; parameres apparently three-branched according to Esben-
Petersen 's drawdng, but unfortunately the structure is not com-
pletely shown. The female genitalia have been worked out by
Tjeder. The additional side-plates of the 7th segment are large
CHENG: REVISION OF THE CHINESE MECOPTEKA 19
aud apically grown together with the 7th tergite. The additional
side-plates of the 8th segment are wholly grown together with
the 8th tergite, which thus appears much broader (in lateral
view) behind the spiracle than before it; the 7th and 8th ster-
nites are comparatively narrow and appear in lateral view as
narrow borders only; the 9th tergite is longer than the 8th seg-
ment; subgenital plate appears broadly oval with tip slightly
produced, smoothly rounded ; distally the plate bears several
strong black setae ; internal skeleton rather large, the plate
broadened near the base, posterior arms well developed, long,
rounded and somewhat converging, the axis very short and
situated wholly within the plate.
Type ( $ ) : Kwanhsien, Szechwan (W. Stotzner) ; in Esben-
Petersen's Collection, Silkeborg. Type ( 9 ) : Wolungkwan,
Szechwan, 55 km. west of Kwanhsien, Szechwan and 150 km.
northeast of Tachienlu, Sikang (\Y. Stotzner) ; in Staatliches
Museum fur Tier- und Volkerkunde, Dresden.
Distribution : same as types.
This species belongs to the diceras group. I have not seen
the types. According to Esben-Petersen's and Tjeder's drawings,
it is closest in general appearance to Panorpa diceras McLachlan.
However, the hypovalvae, though broadened distalby as in diceras,
are truncated distally, whereas those of diceras have a rounded
apex. The proximal lobes of the harpagones of this species are
rounded, not pointed as in diceras. Unfortunately, Esben-Peter-
sen's drawing of the male type does not show details of the
structure of the parameres or of the aedeagus.
Panorpa kimminsi Carpenter
Figures 4, 9, 16,19,271
Panorpa kimminsi Carpenter, 1948, Psyche, 55(1): 29.
Panorpa diceras Carpenter (nee McLachlan), 1938, Proc. Ent. Soc. Washing
ton, 40(9) :270, figs. 2, 10, 11.
Body yellow ; vertex with black area in the region of the ocelli
and between the antennal bases; rostrum with a black stripe
along each side ; eyes grayish brown ; thorax and abdomen yellow,
with a black stripe laterally, 6th abdominal segment of male
with a pair of short anal horns, as in diceras. Fore wing : length,
15-18 mm. ; width, 4 mm. ; membrane dusky hyaline, with grayish
20 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
yellow at the apex ; pterostigma deep yellow, prominent ; neura-
tion as in diceras. Hind wing : length, 13-15.5 mm. ; width, 4 mm. ;
similar to fore wing. Male genitalia: genital bulb oval; coxopo-
dites U-shaped ; harpagones slender, the outer margins smoothly
curved, inner margins with two teeth, a small distal one and a
long, slender, proximal one ; hypandrium inconspicuous, hypo-
valvae prominent, only very slightly broadened distally, reaching
to the base of the harpagones; parameres complex, consisting of
a slender basal stalk, which gives rise to three branches, a small
one on the inner side, a long middle branch, which extends well
up between the harpagones, and a shorter outer branch, which
curves inwardly in back of (i.e., above) the second branch;
preepiproct broad, with long distal lobes ; aedeagus with rounded
apical processes and rounded lateral processes. Female genitalia :
posterior border of subgenital plate rounded, slightly emargi-
nated; internal skeleton small, the axis very small, not project-
ing beyond the plate ; the plate attached to a chitinous, hood-
shaped membrane, which extends slightly beyond the axis.
Holotype ( $ ) : Mt. Omei (11,000 ft.), Szechwan, July, 1936
(D. C. Graham) ; in U. S. National Museum. Allotype ( 9 ) :
near Tachienlu (5000-8500 ft,), Sikang, June 18- July 12, 1923
(D. C. Graham) ; in U. S. National Museum.
Distribution : Vicinity of Tachienlu, 30 miles north of Ta-
chienlu (12,000 ft.), 9 miles southwest of Tachienlu (9000-11,000
ft.), Sikang, June 5-27, 1923; west of Yachow (2000-7500 ft.),
Sikang, June 14-18, 1922; Omei (11,000 ft.), Szechwan, July,
1936 ; Suifu, Szechwan, Oct., 1929.
This species belongs to the diceras group. It is the most widely
distributed of all the double anal horn species. It differs from
the other species by the curved harpagones and the peculiar
configuration of the parameres of the male. It should be noted
that the structure of both the subgenital plate and the internal
skeleton of the female is very similar to that of stotzneri Esben-
Petersen.
Panorpa centralis Tjeder
Figures 5, 10, 20, 21
Panorpa centralis Tjeder, 1936, Ark. for ZooL, 27 A (33) :3, pis. 1, 2, 7,
figs. 1, 2.
CHENG : REVISION OF THE CHINESE MECOPTERA 21
Body mostly black; vertex and frons deep black; rostrum
shining dark piceous with somewhat lighter margins, apically
a little darker j prothorax black with narrow yellowish side-mar-
gins, meso- and metathorax deep-black dorsally, pleura and
underside of thorax reddish yellow ; 1st to 6th abdominal seg-
ments of male deep black, last few abdominal segments fuscous;
anal horn present, single, brownish, directed obliquely upwards ;
the third tergite in the middle of its hind-border develops into a
very small rounded lobe ; abdominal segments of female deep
black, 7th and 8th segments with long narrow additional anal
plates in the pleural region behind the spiracle. Fore wing :
length, 13 mm. ; width, 3 mm. ; membrane hyaline, with a faintly
whitish tinge in male, more yellowish brown in female ; wings of
male without markings, while those of female with spots between
R5 and Mx and at the place where Cux reaches the hind margin;
pterostigma whitish, faintly indicated in male, but distinct and
light-brown in female ; neuration distinct ; Sc extends to the
pterostigmal area, R2 is forked, and Rx, R3, R4, R5 are simple;
usually one crossvein between Rx and R2. Hind wing: length,
12 mm. ; width, 3 mm. ; similar to the fore. Male genitalia : genital
bulb rounded ; coxopodites long, with a group of strong and black,
spine-like hairs on their inner apical margins; harpagones short,
ending in an acute apex, bent slightly inwards, the outer margin
not concave at the middle, the inner margin having a dilatation
midway between base and apex ; the under surface shows a dis-
tinctly rounded excavation; hypandrium inconspicuous; hypo-
valvae rather narrow, not reaching to the base of the harpagones,
their borders straight and running parallel; along their inner
margins they bear a row of long and strong black hairs, directed
inwards ; parameres simple and short ; in their proximal part
they are very broad, but a short distance behind the middle of
their length they suddenly taper into a slender and acute distal
portion, which on the inner margin bears a dense row of lamellae,
directed obliquely inwards; preepiproct narrowed towards apex
with a U-shaped distal incision; aedeagus with very long and
acute apical processes, lateral processes short, blunt tooth-like.
Female genitalia : subgenital plate oval with smoothly rounded
side-margins, which overlap the lower margins of the 9th tergite
to a very little extent ; its apex has a very small excision ; in-
22 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
ternal skeleton large, but rather narrow, with a pair of short,
blunt, posterior arms; on each side of the proximal part of the
plate, there are three lamellae directed forwards and inwards,
ending acutely and having the apices of the two inner pairs bent
somewhat outwards ; the axis is long, extending more than one
third its length beyond the plate, the proximal ends almost un-
pigmented, deeply cleft and directed slightly outwards.
Holotype ( $ ) : Pao-ning-fu (400 m.), northeast Szechwan,
May 20, 1930 (D. Hummel) ; in Stockholm Museum. Allotype
(?*): Tjeggala (3700 in.), Sept. 4, 1930 (D. Hummel); in
Stockholm Museum.
Distribution: same as types.
This species, which has a single anal horn in the male, belongs
to the centralis group. It resembles in general appearance Pan-
orpa cornigera McLachlan, which belongs to the cornigera group,
as established by Issiki. However, many parts of the male and
female genitalia and especially the shape of the parameres and
that of the apical processes of the aedeagus make it easily recog-
nized as a distinct species.
Panorpa emarginata Cheng
Figures 24, 25, 27, 31, 32, 273
Panorpa emarginata Cheng, 1949. Psyche, 56(4) :140, figs. 1, 11. 12, 24, 26,
29.
Vertex entirely black; rostrum grayish brown anteriorly, yel-
lowish brown laterally ; thorax yellowish brown laterally, pro-
notum blackish brown, meso- and metanotum entirely pitchy
black ; the 1st to 6th abdominal segments pitchy black dorsally
and ventrally, last few abdominal segments yellowish brown ; 6th
abdominal segment of male with a single anal horn, yellowish
brown in color. Fore wing : length, 14 mm. ; width, 3.5 mm. ;
membrane hyaline, without markings except for a slight suspi-
cion of gray at the apex ; pterostigma prominent, indicated by
light brown color; the dorsal hind margin of wing slightly emar-
ginated. Hind wing : length. 12.5 mm. ; width, 3.5 mm. ; similar
to fore wing. Male genitalia: genital bulb less rounded; co-
xopodites long, broadened towards its apex ; harpagones short.
the outer margin slightly concave at the middle, inner margin
CHENG : REVISION OF THE CHINESE MECOPTERA 23
with a median small triangular tooth and a large basal concave
area; hypandrium inconspicuous; hypovalvae long, with slightly
concave median outer margins, extending nearly to the base of
the harpagones; parameres simple and long, usually reaching to
the distal part of harpagones, each consisting of a single stalk,
which broadens at the middle and becomes very long and sharp
distally, bearing a series of long barbs at its distal inner margin ;
preepiproct narrowed towards apex, with nearly straight sides
and a narrow U-shaped distal incision; aedeagus with very long
apical processes and prolonged lateral processes, the distal inner
margin of the former usually with a broad triangular plate.
Female genitalia : subgenital plate elongated, emarginated pos-
teriorly, the incision being very small ; internal skeleton large,
the plate concave at the middle, with a pair of sharp distal
posterior arms and two pairs of small basal side plates ; the axis
long, extending beyond the plate one-half its length.
llolotype ( £ ) : Mt. Hwa, Shensi, June, 1942 (Io Chou) ; in
Museum of Comparative Zoolog3r. Allotype ( 9 ) : same collect-
ing data as holotype ; in Cheng Collection, Taipeh .
Distribution : same as types.
This species, possessing a single anal horn, belongs to the
centralis group, with the wing membrane transparent as in
centralis Tjeder. The wing apex of centralis Tjeder is colorless,
whereas that of emarginata is maculated with a slight suspicion
of gray. The male genitalia differ from those of centralis by
the less rounded genital bulb and the longer and sharper para-
meres.
Panorpa obtusa Cheng
Figures 28, 34, 37, 272
Panorpa obtusa Cheng, 1949, Psyche, 56(4) :142, figs. 2, 25, 27, 3U.
Vertex entirely black ; rostrum reddish brown, with a weakly
defined grayish stripe on each side ; thorax reddish brown later-
ally, entirely black dorsally ; the 1st to 6th abdominal segments
black dorsally and ventrally, last few abdominal segments of
male reddish brown; the hind border of the third tergite of male
prolonged into a small semicircular process, 6th abdominal seg-
ment furnished with a single anal horn, reddish brown in color.
24 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
Fore wing : length, 14 mm. ; width, 3.55 mm. ; membrane light
grayish brown, without markings except for a slight suspicion
of grayish brown at the apex ; pterostigma prominent, indicated
by grayish brown color; the wing apex obtuse, broader than in
the preceding species. Hind wing : length, 13 mm. ; width, 3.5
mm. ; similar to fore wing. Male genitalia : genital bulb rounded,
coxopodites long; harpagones short and stout, the outer margin
smoothly curved, the inner margin with a greatly reduced median
tooth (which cannot be seen from ventral view) and a large
basal concave area ; hypandrium inconspicuous ; hypovalvae
rather straight, reaching nearly to the base of the harpagones;
parameres simple and stout, usually not extending beyond the
tips of coxopodites, each consisting of a single spindle-shaped
stalk, formed by the outer strongly sclerotized part ; the distal
inner margins of parameres furnished with a series of long
barbs ; preepiproct slightly narrowed towards apex, with a wide
U-shaped distal incision; aedeagus with small lateral processes
and a pair of long apical processes, the inner margins of the
latter nearly parallel to each other.
Female unknown.
Holotype ( $ ) : Mt. Taipai, Shensi, July 14, 1943 (Chuan
Lung Lee) ; in Cheng Collection, Taipeh.
Distribution : same as types.
This species belongs to the centralis group, having the same
wing markings as emarginata, but the body color and the struc-
ture of the male genitalia, especially the short parameres. make
its recognition easy.
Panorpa flavipennis Carpenter
Figures 3, 11, 13, 22, 23
Panorpa flavipennis Carpenter, 1938, Proe. Ent. Soc. Washington, 40(9) :268,
figs. 3, 7, 12.
Body black ; vertex entirely black ; rostrum reddish brown ;
last few abdominal segments of male reddish brown, single anal
horn on 6th abdominal segment very short, 7th segment moder-
ately incised above. Fore wing : length, 13-14 mm. ; width, 3
mm. ; membrane deep yellow, markings gray ; pterostigmal band
usually complete or nearly so ; basal band interrupted ; apical
CHENG : REVISION OF THE CHINESE MECOPTERA 25
band reduced to a few faint spots ; both basal and marginal spots
absent ; pterostigma dark red, very prominent ; crossveins not
margined. Hind wing: similar to fore wing, except that basal
band is entirely lacking. Male genitalia : genital bulb less
rounded ; coxopodites rather long ; harpagones slender, with the
outer margins slightly concave proximally, and a pair of longer
lobes distally, the ventral lobe having a smoothly curved margin,
the dorsal one an abruptly curved margin; hypandrium incon-
spicuous, hypovalvae very slender, almost reaching to the base
of the harpagones ; parameres simple, each consisting of a single
broad flat process bearing on its inner distal margin a number of
distinct barbs ; preepiproct with curved sides and a moderately
deep, U-shaped incision distally ; aedeagus with rather sharp
horn-like apical processes and rounded lateral processes. Female
genitalia : subgenital plate slender, slightly emarginated distally ;
internal skeleton large, the axis extending beyond the plate for
nearly half its length.
Holotype ( $ ) : Beh-luh-din (6000 ft.), 30 miles north of
Chengtu, Szechwan, Aug. 11, 1933 (D. C. Graham) ; in U. S.
National Museum. Allotype ( 9 ) : same collecting data as holo-
type except for date : Aug. 8, 1933 ; in U. S. National Museum.
Distribution: Beh-luh-din (6000 ft.), 30 miles north of Cheng-
tu, Szechwan ; July 12 to Aug. 27.
This species, possessing a single anal horn, belongs to the cen-
tralis group. At present, there are four known species in this
group. The wing membrane of flavipennis is deep yellow, where-
as that of the other three species is colorless, lacteous or grayish
brown. The male genitalia differ from those of the other three
species in the more slender hypovalvae and the possession of
distinct lobes on the harpagones.
Panorpa davidi Navas
Figure 57
Panorpa davidi Navas, 1908, Mem. Eeal. Acad. Gene. Bare, 1908: 415, figs.
19a, b. Esben-Petersen, 1921, Coll. Zool. Selys Long., 5(2): 29, figs. 21
23. Carpenter, 1945, Psyche, 52(1-2) :73, pi. 10, fig. 4.
Body mostly black ; vertex black, with a reddish-brown spot
behind, touching the eyes ; rostrum reddish brown ; thorax black,
26 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
with a longitudinal median reddish streak; 1st to 6th abdominal
segments of male blackish dorsally and ventrally, last few abdo-
minal segments testaceous; the 6th segment with a small, pale,
hairy notch on the dorsal tip ; no true anal horn present ; 7th
and 8th abdominal segments much narrower than 6th and of
equal length. Fore wing : length, 13 mm. ; membrane hyaline,
with a feeble yellowish tinge ; veins browTnish, the greater part
of crossveins in the apical half of the wing pale; pterostigmal
band complete, with only basal branch; basal band indicated by
two spots; apical band rather large, with a separated spot near
the hind margin of the wing; both basal and marginal spots
absent ; pterostigma rather prominent. Hind wing : length 11.5
mm. ; similar to fore wing, except that the separated spots of
the apical bands are located near the pterostigma. Male geni-
talia : genital bulb slender ; coxopodites long, U-shaped, at the
inner distal end of coxopodites giving rise to a prominent pap-
illa, along the inner surface of -which there is a row of stout
hairs; harpagones small, the outer margin not concave at the
middle, the inner margin no true lobe ; hypandrium inconspic-
uous ; hypovalvae slender, not reaching to the base of the har-
pagones; parameres slender and long, the inner surface bearing
a series of short barbs.
Female unknown.
ilolotype ( 6 ) : Mou-pin, Sikang, 1870 (David) ; in Museum
National d'Histoire Naturelle, Paris.
Distribution : same as holotype.
This species, which has no anal horn in the male, differs from
the others of the davidi group by its elliptical genital bulb, with
long and slender parameres. [ have not seen this species. The
drawing of the genital bulb (fig. 57) is based upon Carpenter's
drawing which he made at the Museum National in Paris in
1938. The wing markings resemble Panorpa cheni n. sp. How-
ever, the structure of the genital bulb is quite different from
that of the latter.
CHENG: REVISION OP THE CHINESE MECOPTERA 27
Panorpa stigmalis Navas
Figures 30, 33
Panorpa stigmalis Navas, 1908, Mem. Real. Acad. Cienc. Bare, 190S: 406.
fig. 20. Esben-Petersen, 1921, Coll. Zool. Selys Long., 5(2) :31, figs. 27-30.
Body mostly black; vertex brown, with a blackish spot en-
closing the ocelli; rostrum blackish at the apex; thorax black
on the dorsum and sides; 2nd and 3rd abdominal segments black
above. 2nd segment also black on the venter, the other abdominal
segments brownish red ; median process of the third tergite rather
long, extending posteriorly and in contact with the conical pro-
jection on the caudal portion of the fourth tergite ; the hind
border of the 6th segment furnished with a group of short hairs
dorsally and ventrally. Fore wing : length, 17 mm. ; very narrow ;
membrane with a strong yellowish tinge, markings smoky brown ;
pterostigmal band absent ; basal band represented by a narrow
streak, extending from the place where 1A joins the hind margin
to Cux ; apical band broad ; both basal and marginal spots absent ;
pterostigma prominent, red in color. Hind wing : length, 16 mm. ;
similar to the fore, except that the interrupted basal band is
entirely absent. Male genitalia : genital bulb oval ; coxopodites
not very long ; harpagones long and slender, the outer margin
concave near the middle, inner margin with a basal process; hy-
pandrium very slender, conspicuous ; hypovalvae slender, with
rather pointed tips, not reaching to the base of the harpagones;
preepiproct slender, with a deep U-shaped distal incision.
Female unknown.
Type ( $ ) : Mou-pin, Sikang (David) ; in Museum National
d'Histoire Naturelle, Paris.
Distribution : same as type.
This species, belonging to the davidi group, is close to the
kongosana group, as established by Issiki (this group comprises
two known species, both of which are continental, i.e., Korean).
However, the wing markings are quite different from those of the
latter group. I have not seen this species. The drawings of the
genital bulb and of the preepiproct are based on Esben-Peter-
sen's, which he made at the Museum National in Paris. Un-
fortunately, his drawings of the male type show nothing of the
structure of the parameres and that of the aedeagus.
28 BULLETIN : MUSEUM OF COMPAEATTVE ZOOLOGY
Panorpa waongkehzengi Navas
Figures 43, 47, 48, 72, 73, 277
Panorpa waongkehzengi Navas, 1935, Notes d'Ent. Chin. Mus. Heude, 2(5) :
98, fig. 62.
Body mostly shining reddish yellow; vertex with a black spot
between ocelli ; antennae blackish brown ; thorax and abdomen
slightly darker dorsally. Fore wing: length, 8.3-9 mm.; width,
2.5 mm., with rounded wing apex ; membrane yellowish, mark-
ings deep brown; pterostigmal band complete, with broad basal
branch and a separated apical branch ; basal band interrupted,
represented by two prominent spots ; apical band also inter-
rupted, consisting of two anterior spots and a narrow shadow
along the wing apex ; basal spot absent ; marginal spot pro-
minent ; pterostigma prominent, deep yellow. Hind wing : length,
7.7-8.3 mm.; width, 2.4 mm.; similar to fore wing, except that
the basal band is represented by only one posterior spot. Male
genitalia : genital bulb slender ; coxopodites long, with narrow
and truncated apex; harpagones very short, the outer margin
slightly concave at the middle, inner margin swollen medially
with a large cylindrical basal lobe which is concave apically to
form the shape of an ear ; hypandrium short, broader distally ;
hypovalvae narrowed towards their rounded apex, not extending
near the base of the harpagones; parameres simple, each con-
sisting of a single stalk, which is distinctly twisted and bears a
row of short barbs along its inner margin; preepiproct broad
basally, abruptly narrowed towards its apex, which is slightly
emarginated and bears a row of long black bristles; aedeagus
large, apical processes very long with enlarged boot-shaped apex ;
lateral processes not prominent, slightly protruding to form a
pair of triangular plates. Female genitalia : subgenital plate
broad, narrowed towards apex which is slightly concave; in-
ternal skeleton large, the plate narrow basally ; the posterior
arms of the plate stout, pointed, the axis long, strongly sclero-
tized, extending beyond the plate for exactly half its length.
Types ( $ , 5 ) : Ruling, Kiangsi, Aug. 28, 1935 (Piel) ; in
Heude Museum, Shanghai.
Distribution : same as types.
CHENG : REVISION OF THE CHINESE MECOPTERA 29
This species, belonging to the davidi group, differs from other
described Panorpa by its short harpagones and broad hypan-
driura of the $ genitalia. The peculiar structure of the internal
skeleton of 9 genitalia also makes its recognition easy. The
description and drawings of both the male and the female of
this species are based upon specimens labeled as paratypes,
which were loaned to me by the Ileude Museum, Shanghai.
Panorpa difficilis Carpenter
Figures 42, 46
Panorpa difficilis Carpenter, 1938, Proc. Ent. Soc. Washington, 40(9) :269,
figs. 4, 6.
Body black ; vertex with a narrow transverse black band en-
closing ocelli ; rostrum light brown ; anal horn absent ; last few
abdominal segments brown. Fore wing: length, 11 mm.; width
3 mm. ; membrane hyaline, markings grayish brown ; pterostig-
mal band broad but interrupted, basal band reduced to a small
spot at the origin of Rs (in type) ; apical band much broken;
both basal spot and marginal spot absent ; crossveins not mar-
gined. Hind wing: similar to the fore wing. Male genitalia:
genital bulb rounded ; coxopodites long ; harpagones small, the
outer margins slightly concave, apices abruptly curved, no true
lobes present ; hypandrium rather conspicuous, hypovalvae of
moderate width, not reaching to the base of the harpagones;
parameres simple, each consisting of a single stalk, broader dis-
tally than proximally, and with a cluster of barbs near the tip ;
the interior cavity of the genital bulb is unusually narrow ; pre-
epiproct slender, with a deep U-shaped distal incision; apical
processes of aedeagus very long with greatly enlarged distal end,
lateral processes more or less rounded.
Female unknown.
Holotype ( $ ) : Chengtu, Szechwan, 1936 (D. C. Graham) ;
in U. S. National Museum.
Distribution: same as holotype.
This species, belonging to the davidi group, resembles Panorpa
curva Carpenter superficially, but differs in the broader genital
bulb and especially in the absence of the prominent papilla at
the distal inner margin of the coxopodites.
30 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Panorpa fructa Cheng
Figures 29, 35, 38
Panorpa fructa Cheng, 1949, Psyche, 56(4) :144, figs. 5, 6, 7.
Body mostly sooty black, last few abdominal segments of male
reddish brown, vertex black anteriorly, deeply reddish brown
posteriorly ; rostrum uniformly reddish brown. Fore wing :
length, 11.5 mm.; width, 3 mm.; membrane hyaline, markings
light brown, ill-defined, fragmentary; pterostigmal band incom-
plete, with a spot-like basal branch; basal band represented by
two spots ; apical band appearing as a light suspicion of brown
at the apex ; basal spot very small ; marginal spots large ; ptero-
stigma not very prominent. Hind wing : length, unknown ; width.
3 mm. ; similar to fore wing, except that the basal band and the
basal spot are entirely absent. Male genitalia: genital bulb very
rounded ; eoxopodites long, stout, with four spine-like hairs and
a series of short hairs in the distal inner portions; harpagones
slender, the outer margin rather straight, inner margin with a
median angle and a rather large basal concave area ; hypandrium
inconspicuous; hypovalvae rather short, far from reaching to
the base of the harpagones ; parameres simple, long and twisted,
the distal half well developed, with rounded apex furnished with
a short spine-like tip; preepiproct rather short, broad at the
base, narrowed towards apex, with a broad U-shaped distal in-
cision; aedeagus with long apical and lateral processes, the
former with rather straight inner margins and double sinuous
outer margins.
Female unknown.
Holotype ( $ ) : Wakiakeng (50 miles west of Tachienlu).
Sikang, Sept. 9, 1939 (F. Y. Cheng, Io Chou and Tein Ho Hei) ;
in Cheng Collection, Taipeh.
This species belonging to the davidi group, resembles Panorpa
difficilis Carpenter superficially, but differs in the structure of
the male genitalia. In difficilis, the apical processes of the aedea-
gus have greatly enlarged distal ends, whereas in this species,
the distal ends of the apical processes of the aedeagus are narrow
and rather pointed.
i:HENG: REVISION OF THE CHINESE MECOPTERA 31
Panorpa cladocerca Navas
Figures 41, 51, 68, 74, 284
Pavorpa cladocerca Navas, 1935, Notes d 'Ent. Chin. Mus. Heude, 2(5) :95,
figs. 59, 60.
Vertex shining black anteriorly, chestnut brown posteriorly ;
rostrum uniformly shining reddish yellow ; thorax black dorsally,
deep yellow laterally ; 1st to 5th abdominal segments of male
black dorsally, reddish yellow ventrally, 6th abdominal segment
black anteriorly, chestnut brown posteriorly, last few abdominal
segments reddish brown; anal horn absent; abdominal segments
of female black dorsally, chestnut brown ventrally. Fore wing :
length, 12.5-14.5 mm. ; width, 3.6 mm. ; membrane hyaline, mark-
ings sooty brown ; pterostigmal band broad, with same broad
basal branch, but no apical branch; basal band complete; apical
band large, slightly interrupted posteriorly ; basal spot absent ;
marginal spot very small, not extending to the anterior margin
of the wing; pterostigma prominent. Hind wing: length, 11.3-
12.7 mm. ; width, 3.4 mm. ; similar to fore wing, except that the
small marginal spot is absent. Male genitalia : genital bulb oval ;
coxopodites short, with truncated apex ; harpagones slender, the
outer margin slightly convex at the middle, inner margin with
a median angle and a true basal lobe ; basal concave area very
large, same width as the harpagones and occupying two-thirds
of the whole length of the latter; hypandrium inconspicuous;
hypovalvae long and slender, slightly crooked at the middle,
diverging from each other distally, reaching to the base of the
harpagones; parameres simple, each consisting of a single stalk,
with broad base and inwardly curved apex, the inner margins
usually bearing a row of short barbs ; preepiproct slender, with
nearly straight sides and a deep U-shaped distal incision ; aedea-
gus prominent, apical processes short with rounded apex, lateral
processes very slender, extending upward behind the basal lobe
of the harpagones. Female genitalia : subgenital plate slender,
slightly constricted at the middle, with wedge-shaped apex ; in-
ternal skeleton small, the median portion of the plate less sclero-
tized, with a pair of narrow vertically erected lobes, the pos-
terior arms of the plate rather long, pointed towards apex.
Types ( $ , 9) -. Kuling, Kiangsi, Sept. 4-7, 1934 (Piel) : in
32 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Heude Museum, Shanghai.
Distribution : same as types.
This species, belonging to the davidi group, differs from other
described Panorpa by its unforked pterostigmal band and the
complete basal band in the wing markings. The peculiar struc-
ture of both the male and the female genitalia also makes its
recognition easy. The description and drawings of both the male
and the female of this species are based upon specimens labeled
as paratypes which were loaned to me by the Heude Museum,
Shanghai.
Panorpa trifasciata n. sp.
Figures 40, 49, 50, 64, 65, 283
Body mostly black ; vertex sooty brown with black mark en-
closing ocelli ; rostrum uniformly grayish brown ; thorax light
brown laterally, sooty brown dorsally; the 1st to 5th abdominal
segments of male piceous black dorsally, 6th segment twice as
long as preceding segment, uniformly black, last few abdominal
segments nearly same length as the 6th, reddish brown in color,
the hind border of the third tergite slightly produced ; the 1st
to 6th abdominal segments of female blackish brown dorsally
and ventrally, last few abdominal segments reddish brown. Fore
wing: length, $ , 13.2 mm., $ , 15 mm.; width, $ , 3.1 mm., $ ,
3.6 mm. ; membrane hyaline, markings sooty brown ; pterostigmal
band broad, basal branch of the pterostigmal band well developed,
as broad as pterostigmal band so as to form a broad transverse
band ; apical branch of pterostigmal band absent ; basal band
complete ; apical band prominent ; both basal and marginal spots
absent ; pterostigma very prominent. Hind wing : length, $ , 12
mm., $ , 13.5 mm. ; width, <3 , 3 mm. ; $ , 3.5 mm. ; similar to
fore wing, except that the basal band is not so developed. Male
genitalia : genital bulb oval ; coxopodites long and stout with
truncated apex and a row of long spine-like bristles on its dis-
tal inner margin ; harpagones slender, the outer margin slightly
smoothly curved, inner margin with a sharp median angle and
a basal lobe which does not arise at the same level as the former ;
hypandrium short; hypovalvae straight, slender, with rounded
apex, not extending near to the base of harpagones; parameres
CHENG : REVISION OF THE CHINESE MECOPTERA 33
simple, narrowed towards apex, their whole inner margin bearing
dense and long blackish brown, spine-like bristles; the median
portion of parameres usually covered by the aedeagus in ventral
view; preepiproct broad, abruptly narrow at the apical portion
with deep U-shaped distal incision; aedeagus large, the apical
processes tooth-like, lateral processes well developed, long, rod-
shaped with swollen apex, extended upward and reaching to
the median portion of the harpagones. Female genitalia : sub-
genital plate rather long, narrowed towards apex with folded
margins on its basal portion ; internal skeleton small, the median
portion of the plate swollen to form a pair of thick margins;
posterior arms of the plate narrowed towards apex; no axis
present.
Holotype ( $ ) : Ta-chu-lan, Shaowu Hsien, Fukien, Nov. 7,
1943 (Maa) ; in Museum of National Foochow University, Foo-
chow. Allotype ( 2 ) : Sien-feng-ling, Shaowu Hsien, Fukien,
Oct. 15, 1941 (Maa); in Maa Collection. Paratypes: 4 5 9,
same collecting data as holotype ; 2 $ $ , 1 2 , Kuatun, Chungan
Hsien, Fukien, Aug. 22-Oct. 29, 1942-1945 (Maa and Lin), in
Maa Collection ; 1 $ , 12, same collecting data as holotype, in
Museum of Comparative Zoology ; 2 $ $ , 3 2 2 , same collecting
data, in Cheng Collection, Taipeh.
This species belongs to the davidi group and resembles Panorpa
cladocerca Navas in the wing markings. However, in cladocerca,
the wing has more extensive markings and the marginal spot is
always present, whereas in trifasciata, the three bands of the
wing are much narrower than those of cladocerca and the mar-
ginal spot is absent. The genitalia of trifasciata differ from those
of cladocerca by the peculiar basal lobes of the harpagones of
the male and the triangular-shaped subgenital plate of the
female.
Panorpa flavicorporis n. sp.
Figures 59, 60, 69, 75, 80
Body yellowish brown ; vertex deep brown with slightly black-
ish-brown mark enclosing ocelli ; rostrum uniformly light brown ;
thorax slightly blackish-brown dorsally, meso- and metanotum
with a very broad, light brown streak ; the 1st to 5th abdominal
34 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
segments of male slightly blackish brown dorsally, last few ab-
dominal segments yellowish brown; the 1st to 6th abdominal
segments of female blackish brown dorsally, last few segments
yellowish brown. Fore wing : length, 6 , 12.3 mm., 9 , 14.2 mm. ;
width, $ , 3.1 mm., 9 , 3.5 mm. ; membrane hyaline, markings
sooty brown; pterostigmal band complete with narrow basal
branch, apical branch not developed ; basal band much restricted
medially, separated into two spots ; apical band large, inter-
rupted posteriorly ; basal spot very small ; marginal spot pro-
minent; pterostigma not very prominent. Hind wing: length.
$ , 11 mm., 9 , 12.5 mm. ; width, $ , 3.1 mm., 9 , 3.5 mm. ; simi-
lar to fore wing, except that the basal band is represented only
by one spot and both basal and marginal spots are entirely lack-
ing. Male genitalia : genital bulb slender ; coxopodites long, with
truncated apex ; harpagones slender, the outer margin smoothly
curved, inner margin with a median angle and a broad basal
lobe ; hypandrium short ; hypovalvae with narrow, blunt apex,
reaching far beyond to the base of the harpagones; parameres
simple, very long with broad base and twisted apical portion,
extending nearly to the apex of harpagones ; preepiproct broad,
narrowed towards apex with broad U-shaped distal incision ;
aedeagus large, the apical processes horn-like, extending out-
ward, lateral processes appearing as a pair of lobes. Female
genitalia : subgenital plate long and slender, with folded, nar-
row, membranous margins and wedge-shaped distal end ; internal
skeleton very small, the plate rather broad, the posterior arms
of the plate narrowed towards apex, the axis small, slightly pro-
jecting beyond the plate.
Holotype ( S ) : Ta-chu-lan, Shaowu Hsien, Fukien, Oct. 21,
1943 (Maa) ; in Museum of National Foochow University, Foo-
chow. Allotype ( 9 ) : Hwang-keng, Kienyang Hsien, Fukien,
Oct. 11, 1943 (Maa); in Maa Collection. Paratypes: 8 S $, 16
9 9 , same collecting data as holotype ; 1 $ , 2 9 9 , Kienyang
Hsien. Fukien, Oct. 17-31, 1942 (Maa), in Maa Collection; 2$ $ ,
2 9 9, Pciu-tun, Kienyang Hsien, Fukien, Oct. 15, 1941 (Maa
and Lin), in Museum of Comparative Zoology; 2$ $, 4 9 9,
same collecting data, in Cheng Collection, Taipeh.
This species, belonging to the davidi group, resembles Panorpa
davidi Navas in the wing markings. However, in this species,
CBLENG : REVISION OF THE CHINESE MECOPTERA 35
the marginal spot of the wing is prominent, whereas that of
davidi is absent. The harpagones of the male genitalia of this
species have broad basal lobes, whereas those of davidi are absent.
The yellowish body color and the peculiar structure of the fe-
male genitalia also make its recognition easy.
Panorpa cheni n. sp.
Figures 56, 86, 88, 95, 96
Body brown; vertex blackish brown, with small black mark
enclosing ocelli; rostrum uniformly light brown; thorax blackish
brown dorsally, light brown laterally ; the 1st to 5th abdominal
segments of male blackish brown dorsally, light brown ventrally.
6th segment long, cylindrical, blackish brown, last few segments
light brown, the hind border of the third tergite slightly pro-
duced ; 1st to 6th abdominal segments of female black dorsally
and ventrally, last few segments light brown. Pore wing: length.
S, 11.7 mm., 9, 13.5 mm.; width, $, 2.9 mm., 9, 3.5 mm.;
membrane hyaline, markings sooty brown; pterostigmal band
complete, basal branch of pterostigmal band broad, the outer
margin of the latter with hyaline area, apical branch absent;
basal band represented by two narrow spots; apical band pre-
sent ; basal spot absent ; marginal spot very small and sometimes
absent; pterostigma not very prominent. Hind wing: length.
£ , 10.7 mm., 5 , 12.5 mm. ; width, 6 , 2.8 mm., 9 , 3.5 mm. ;
similar to fore wing, except that the basal band is entirely lack-
ing. Male genitalia : genital bulb oval ; coxopodites long, with
truncated apex ; harpagones slender, the outer margin slightly
concave at the middle, inner margin with a prominent tooth-like
process and a large true basal lobe which does not arise at the
same level ; hypandrium inconspicuous ; hypovalvae narrow and
slender, divergent, not extending near the base of the harpa-
gones ; parameres branched basally, the outer branch long, broad-
ened towards apex, with knob-shaped distal end ; the inner branch
short, with a bundle of long, stiff brown hairs; preepiproct broad,
slightly narrowed towards apex, with U-shaped distal incision ;
aedeagus with prominent ventral processes and twisted rod-
shaped apical processes, lateral processes absent. Female geni-
talia : subgenital plate triangular, with slightly emarginated
36 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
apex; internal skeleton small, the median part of the plate a
little swollen, less sclerotized with a very short axis, the posterior
arms of the plate long and slender, narrowed towards the apex.
Holotype ( S ) : Tien-mu-shan, Chekiang, July 27, 1936 ; in
Museum of Institute of Zoology, Academia Sinica, Shanghai.
Allotype ( 2 ) : Tien-mu-shan, Chekiang, Aug. 17, 1935; same
type location as holotype. Paratypes : 1 2 , same collecting data
and same type location as allotype ; 1 2 , same collecting data, in
Museum of Comparative Zoology ; 1 $ , 1 2 , same collecting data,
in Cheng Collection, Taipeh.
I take the liberty to name this species in honor of Dr. Sieien
H. Chen who has been so kind as to loan me the material from
the Museum of the Institute of Zoology, Academia Sinica, Shang-
hai.
This species, belonging to the davidi group, resembles Panorpa
davidi Navas and P. flavicorporis n. sp. in the wing markings,
but is quite different in both the male and the female genitalia.
The parameres of the male genitalia of both davidi and flavicor-
poris are simple, whereas those of cheni are branched. The sub-
genital plate of the female genitalia of flavicorporis is long and
slender, whereas that of cheni is triangular.
Panorpa obliqua Carpenter
Figures 44, 45, 97, 98
Panorpa obliqua Carpenter, 1945, Psyche, 52(1-2) :70, pi. 10, figs. 2, 3, 8;
pi. 11, fig. 9.
Body brown ; vertex dark brown ; thoracic nota and abdominal
tergites very dark brown ; anal horn of male absent. Fore wing :
length, 12 mm. ; width, 3.5 mm. ; membrane faintly yellow, mark-
ings blackish brown; pterostigmal band complete with rather
narrow basal branch and a separated apical branch, the latter
connected with the apical band to form a hyaline window which
encloses a faint, brown spot ; basal band represented by two
spots, the large one, situated on the anal margin of wing, is very
large with a hyaline window; apical band large, usually united
with the apical branch of pterostigmal band, and within its
area, there are two hyaline bands and one large hyaline win-
dow; basal spot appears as a transverse band along the
CHENG : REVISION OF THE CHINESE MECOPTERA 37
anal margin of the wing; marginal spot elongated, band-like,
united with the basal branch of pterostigmal band to form a
V-shaped marking; pterostigma prominent; crossveins not mar-
ginated. Hind wing : similar to fore wing in markings, except
that the left "arm" of the "V" is usually interrupted. Male
genitalia : genital bulb rounded, coxopodites long, U-shaped ;
harpagones short, outer margins slightly concave ; prominent
lobes on inner margin of the harpagones near the base ; hypan-
drium rather long, conspicuous; hypovalvae short, close to each
other at the middle of the genital bulb, and extending slightly
beyond the base of the harpagones; parameres simple, each con-
sisting of a slightly flattened process, with a few short barbs on
the inner surface distally, and terminating in a longer series of
barbs directed inwards; preepiproct much narrower distally,
with a pair of broad terminal lobes, close together. Female
genitalia: subgenital plate with rounded posterior margin; in-
ternal skeleton large, with two long, slender distal processes and
a pair of large, ear-like flaps laterally; the usual axis is short,
but there is a very slender median process extending posteriorly.
Holotype ( $ ) : Hong San, Southeast Kiangsi, June 28, 1936 ;
in Museum of Comparative Zoology. Allotype ( 9 ) : Hong San,
Southeast Kiangsi, June 29, 1936 ; in Museum of Comparative
Zoology.
Distribution : same as types.
This strikingly marked species, belonging to the davidi group,
is easily recognized by the oblique stripe traversing the middle
of the wing, and extending posteriorly and distally from the
anterior margin. The short hypovalvae of the male genital
bulb are unique among the known Chinese species of Panorpa,
but are very much like those of certain Japanese, Siberian and
Formosan species (e.g., Panorpa preyeri McLachlan, wormaldi
McLachlan, and peterseana Issiki), to which obliqua Carpenter
is undoubtedly closely related.
Panorpa implicata n. sp.
Figures 99, 103, 280
Body mostly black; vertex entirely black; rostrum rather
short, uniformly reddish brown; thorax pale-white laterally,
38 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
sooty brown dorsally; the 1st to 9th abdominal segments of
female deep brown dorsally and ventrally. Fore wing : length,
13.5 mm. ; width, 3.2 mm. ; membrane hyaline, markings sooty
brown, similar to the preceding species; pterostigmal band
complete with broad basal branch and a separate large apical
branch, the latter with a hyaline spot and connected with the
apical band to form another hyaline spot ; basal band represented
by two spots ; the one situated on the anal margin of wing is very
large and includes a hyaline spot; apical band large, usually
united with the apical branch of pterostigmal band, and
within its area there are one large hyaline spot and two hyaline
bands; basal spot appears as a transverse band along the anal
margin of wing ; marginal spot elongate, band-like, united with
the basal branch of pterostigmal band to form a V-shaped mark-
ing ; pterostigma prominent. Hind wing : length, 12 mm. ; width,
8.1 mm. ; similar to the fore. Female genitalia : subgenital plate
broadened medially with slightly concave apex; internal skeleton
large, the plate bearing paired, folded, side plates; the posterior
arms of the plate crooked, narrowed towards apex, the axis long,
extending beyond the plate for exactly half its length.
Male unknown.
Holotype ( 9 ) : Ta-chu-lan, Shaowu Hsien, Fukien, May 15,
1945 (Maa) ; in Museum of National Foochow University, Foo-
chow. Paratypes : 2 9 5 , same collecting data as holotype ; 1 9 .
Tsi-li-chiao, Chungan Hsien, Fukien, May 1, 1943 (Maa), in Maa
Collection ; 1 9 , same collecting data as holotype, in Museum of
Comparative Zoology ; 1 9 , Kuatun, Chungan Hsien, Fukien,
May 11, 1942 (Maa), in Cheng Collection, Taipeh.
This species resembles Panorpa obliqua Carpenter in the wing
markings, but differs in the female genitalia. The subgenital
plate of obliqua has a rounded posterior margin, whereas that
of implicata is slightly concave. The axis of the internal skeleton
of obliqua is short, whereas that of implicata is long. The posi-
tion of this species in the above grouping is not determined.
Panorpa baohwashana n. sp.
Figures 82, 83, 84, 101, 105, 285
Body mostly black; vertex uniformly deep black; rostrum uni-
CHENG : REVISION OF THE CHINESE MECOPTERA 39
formly brownish black ; thorax mostly black dorsally, reddish
brown laterally; the 1st to 9th abdominal segments of both sexes
black dorsally and ventrally, the hind border of the third ter-
mite of male produced into a short process, and in contact with
the small, very sharp conical projection on the median fourth
tergite^ the (Jth to 8th segments prolonged, usually longer than
the 5th segment. Fore wing : length, 18-20 mm. ; width, 5 mm. ;
membrane hyaline, markings brownish black; pterostigmal band
complete, with broad basal branch and very narrow and separated
apical branch; basal band represented by two spots; apical band
very large; basal spot absent; between the basal area and the
basal band, there are two additional small spots; marginal spot
small, not extending to the anterior margin of wing; pterostigma
not very prominent. Hind wing: length, 16.5-18.5 mm.; width,
4.5 mm. ; similar to fore wing, except that the basal band is
represented by only one spot, the two additional spots between
the basal area and the basal band are entirely absent, and the
marginal spot not well developed. Male genitalia: genital bulb
slender; coxopodites long, with truncated apex; harpagones very
slender, outer margin deeply concave basally, smoothly curve!
distally, inner margin with three small lobes; the outer margin
folded upward, the apex folded inward to form a rounded distal
end which reaches to the base of the harpagones; parameres
simple, each consisting of a single stalk, which is broader distally
than basally and bears on its distal inner margin a row of short
barbs; preepiproct slender, the basal portion broad, oval; the
distal portion narrow, the apex with very minute distal incision
usually folded downward ; aedeagus with prominent ventral
processes, apical processes long with folded apical portion, ex-
tending above the basal lobes of harpagones, lateral processe.->
finger-like, extending upward. Female genitalia : subgenital plate
broader distally than basally, with small V-shaped distal incision,
the median longitudinal line of the subgenital plate less sclero-
tized; internal skeleton with long and stout axis, the plate lonp:
and narrow, occupying the area between the two axes, the pos-
terior arms of the plate well-developed, with sharp apex.
Holotype ($): Bao-hwa-shan, Kiangsu, July 16, 1942; in
Museum of Institute of Zoology, Academia Sinica, Shanghai. Al-
lotype (?':>: Same collecting data and same type location as
40 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
holotype. Paratypes : 4 S 5,5$ $ , same collecting data as holo-
type, in same type location as holotype ; 2 $ 5,2$ $ , same col-
lecting data as holotype, in Museum of Comparative Zoology;
2$ 5 , 3 5 5 , same collecting data ; 3 5 $ , 2 5 $ , N. Han-nu-
shan, Jul}7 16, 1940, in Cheng Collection, Taipeh.
This species, belonging to the davidi group, resembles Panorpa
approximata Esben-Petersen, which is in the amurensis group,
as established by Issiki (this group comprises four known
species, all of which are known as continental, i.e., E. Siberia
and Korea), but differs in several respects. Esben-Petersen 's
drawing of the male genitalia of approximata shows that the
hypandrium is rather long and the distal end of the preepiproct
is only slightly emarginated. The distal part of the hypandrium
of this new species is deeply cleft to form the narrow bases of the
hypovalvae ; the distal end of the preepiproct has a narrow
U-shaped distal incision. Unfortunately, according to Esben-
Petersen, the type of approximata ( 5 ) lacks the parameres, so
I cannot compare the genital structures of these two species in
detail.
Panorpa typicoides Cheng
Figures 26, 36, 39, 66, 67, 276
Panorpa typicoides Cheng, 1949, Psyche, 56(4): 143, figs. 3, 13, 14, 28, 31.
Body mostly black ; vertex black anteriorly, brown posteriorly ;
rostrum entirely brown ; thorax black dorsally, yellowish brown
laterally, meso- and metanotum with a broad brown median band ;
1st to 6th abdominal segments of male black dorsally and ven-
trally, last few abdominal segments reddish brown, anal horn
absent; the hind border of third tergite slightly prolonged be-
hind, and in contact with the small, sharp conical production on
the median axis of the 4th tergite ; abdominal segments of female
entirely black. Fore wing : length, 12.5 mm. ; width, 3 mm. ;
membrane hyaline, markings sooty brown; pterostigmal band
complete, with a broad basal branch and a separated narrow
apical branch ; basal band interrupted, represented by two large
spots; apical band broad, with a large hyaline spot posteriorly;
basal spot very small ; marginal spot large, not extending beyond
the vein P^ ; pterostigma brown, very prominent. Hind wing :
length, 11.5 mm. ; width, 3.3 mm. ; similar to fore wing, except
that the basal spot and the anterior part of the basal band are
CHENG : REVISION OP THE CHINESE MECOPTERA 41
entirely lacking. Male genitalia : genital bulb rounded ; coxo-
podites long, U-shaped, furnished with a series of long hairs
at the distal inner portion ; harpagones slender, the outer margin
slightly concave at the middle, inner margin with a median
angle and a small basal concave area; hypandrium inconspicu-
ous; hypovalvae rather long, reaching to the base of the harpa-
gones ; parameres simple and slender, each consisting of a single
stalk, which is distinctly twisted and pointed at its apex ; pre-
epiproct slender, slightly narrowed towards apex, with a deep
U-shaped distal incision ; aedeagus with finger-shaped apical
processes and slightly prolonged lateral processes, the distal inner
margins of the former usually produced inwards to form a small
nipple-shaped plate. Female genitalia : subgenital plate elon-
gated, broadened at the middle ; internal skeleton long, the plate
narrowed towards its base with a pair of sharp posterior arms,
the axis very long, extending nearly two-thirds its length beyond
the plate.
Holotype ( 9 ) : Tachienlu (5000-8500 ft.), Sikang, Aug. 27,
1939 (F. Y. Cheng, Io Chou and Tein Ho Hei) ; in Museum of
Comparative Zoology. Allotype ( 9 ) : same collecting data as
holotype ; in Cheng Collection, Taipeh.
Distribution : same as types.
This species, belonging to the davidi group, resembles the com-
mon European species Panorpa communis Linne and P. fukiensis
Tjeder in the wing markings, but differs in the structure of the
male genitalia. The parameres of communis are very broad, lobe-
shaped; those of fukiensis are spindle-shaped, whereas those of
typicoides are narrow and slender, thread-like.
Panorpa fukiensis Tjeder
Figures 54, 58, 70, 71
Panorpa fukiensis Tjeder, 1950, Bonn Zool. Beitr., 1950 (2-4) :2S6, figs. 1, 2.
Head blackish brown ; rostrum yellowish brown ; thorax black-
ish brown dorsally, yellowish brown laterally, the meso- and
metanotum with light brown across their hind parts; 1st to 5th
abdominal segments of male blackish brown dorsally; 6th seg-
ment blackish brown in the upper part and faintly yellow in the
lower part, no anal horn present ; 7th and 8th segments long and
narrow, of the same length ; abdomen of female long and slender,
42 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
blackish brown, a little lighter at the basal portion; apex of the
abdomen light brown ; additional side plates of the 7th and 8th
segments very narrow, 9th tergite long. Fore wing : length, 6 .
14 mm. ; 9 , 16 mm. ; width, £ , 3.3 mm. ; 9 , 3.8 mm. ; membrane
hyaline, markings blackish brown ; veins whitish ; pterostigmal
band distinct, with broad basal branch and separated narrower
apical branch ; basal band broad, complete ; apical band broad, in-
terrupted posteriorly, connected with the pterostigmal band an-
teriorly ; basal spot absent ; marginal spot present ; pterostigma
rather prominent. Hind wing : length, S , 12.5 mm. ; $ , 14 mm. ;
width, $ , 3 mm. ; $ , 3.5 mm. ; similar to fore wing. Male geni-
talia : genital bulb elliptical ; coxopodites long, with more or less
truncated apex ; harpagones slender, the outer margin smoothly
curved, inner margin with broad tooth-like projection, the
median tooth located ventrally; hypandrium narrow, conspicu
ous ; hypovalvae short, reaching about the middle of the coxo-
podites, slightly diverging from each other; parameres simple,
spindle-shaped (list ally, each with eleven long, strong barbs on
its inner edge and the same number of slightly narrower barbs
at its apex ; preepiproct narrow, with almost parallel edges and
a deep U-shaped distal incision; aedeagus with long finger-like
apical processes and ax-shaped lateral processes. Female geni-
talia: subgenital plate long, narrowed towards apex; internal
skeleton large, the outer margins slightly concave at the middle,
with a pair of sharp posterior arms; axis well-developed, out
wardly curved at their proximal ends, extending beyond the plate
for about one-third its length.
Holotype ( S ) : Kwangtseh, Fukien, Sept. 25, 1937 (J. Klap
perich) ; in Zool. Keichsinstitut and Museum A. Koenig. Allotype
( 9 ) : Kwangtseh, Fukien, Oct. 9, 1937 (J. Klapperich) ; same
type location as holotype.
Distribution: same as types
This species, belonging to the davidi group, resembles Panorpa
typicoides Cheng in the wing markings, but the shape of the
genital segments of both the male aud the female make it a dis-
tinct species.
CHENG: REVISION OF THE CHINESE MECOPTEBA 43
Panorpa curva Carpenter
Figures 52, 53
Panorpa curva Carpenter, 1938, Proc Ent. Soc. Washington, 40(9) :269.
tie?. 1. 8.
Body black ; vertex with a transverse black band, enclosing
the ocelli; rostrum light reddish brown; anal horn absent; last
few abdominal segments reddish brown. Fore wing: length, 7
mm. ; width. S mm. ; membrane hyaline, markings grayish brown ;
both basal and marginal spots absent ; basal, pterostigmal and
apical bands complete, the last with a few interrupted spots ;
erossveins not margined. Hind wing: similar to the fore wing
^Male genitalia: genital bulb rather long; coxopodites very long,
deep U-shaped; at the inner distal margins of coxopodites, there
is a very prominent papilla bearing a number of black hairs and
giving rise proximally to a large black spine; harpagones small,
the outer margins slightly concave near the middle, the apices
abruptly curved ; no true lobes present ; hypandrium inconspicu-
ous; hypovalvae rather slender, not reaching to the base of the
harpagones; parameres simple, each consisting of a single stalk,
which is distinctly twisted and bears distally a cluster of short
barbs; preepiproct slender, with nearly straight side and a deep
U-shaped distal incision ; apical processes of aedeagus very long
and slender, the outer margins abruptly convex near the middle,
the lateral processes short, horn-like.
Female unknown.
Holotype ( $ ) : 0-er (9000 ft., 26 miles north of Li-fan),
Sikang, Aug. 6, 1916 (D. C. Graham) ; in U. S. National Museum.
This species, belonging to the davidi group, resembles Panorpa
davidi Navas superficially. However, the parameres of this
species are distinctly twisted distally, whereas those of davidi are
not so. The outer margins of the harpagones of curva are slightly
eoncave at the middle, whereas those of davidi are not concave at
all.
Panorpa aurea n. sp.
Figures 55, 61, 62, 76, 77, 281
Body yellowish brown; vertex brown anteriorly with sooty
brown mark enclosing ocelli, yellowish brown posteriorly with
44 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
three narrow longitudinal streaks; rostrum uniformly yellow;
thorax yellow laterally, meso- and metanotum deep brown an-
teriorly, yellowish brown posteriorly; the 1st to 5th abdominal
segments of male and female brown dorsally, last few abdominal
segments yellowish brown, the hind border of the third abdominal
tergite of male slightly produced. Fore wing : length, $ , 15.5 mm.,
9 , 16.3 mm. ; width, $ , 4 mm., 5 , 4.2 mm. ; membrane light yel-
low, markings yellowish brown ; pterostigmal band complete with
same broad basal branch and apical branch; basal band broad,
complete ; apical band large, with a faint and small window ; both
basal and marginal spots are very small ; pterostigma not prom-
inent. Hind wing: length, $ , 14.5 mm., 5 , 14.7 mm. ; width, S ,
3.7 mm., 9 , 4 mm. ; similar to fore wing except that both basal
and marginal spots are entirely lacking. Male genitalia : genital
bulb slender ; coxopodites long with truncated apex ; harpagones
slender, the outer margin slightly concave basally, smoothly
curved distally, inner margin with a median small angle and a
long and large basal concave area ; hypandrium short ; hypo-
valvae slender with rounded apex, not extending near the base
of the harpagones; parameres long, Y-shaped, the outer branch
crooked, the inner branch straight; preepiproct long with deep
U-shaped distal incision ; aedeagus elongated, the apical processes
long and sharp, lateral processes short, lobe-shaped. Female geni-
talia : subgenital plate long with wedge-shaped distal end ; inter-
nal skeleton large, the plate constricted medially to form the
proximal and distal oval portions, the posterior arms of the
plate narrowed towards apex, the axis long, with broad base,
extending beyond the plate for exactly half its length.
Holotype ( $ ) : Kuatun, Chungan Hsien, Fukien, Oct. 28, 1942
(Maa) ; in Maa Collection. Allotype ( 2 ) : Kwantseh Hsien,
Fukien, Sept. 23, 1943 (Maa) ; in Museum of National Foochow
University, Foochow. Paratypes : 1 9 , same collecting data as
allotype, in Museum of National Foochow University ; 4 9 9 ,
same collecting data as holotype ; 4 9 9 , Ta-chu-lan, Shaowu
Hsien, Fukien, Oct. 14-28, 1942 (Maa), in Maa Collection; 1$ ,
1 9 , Ta-chi-lan, Shaowu Hsien, Fukien, Sept. 2-Nov. 28, 1942
(Maa), in Museum of Comparative Zoology; 1 $ , 1 9 , same col-
lecting data, in Cheng Collection, Taipeh.
This species, belonging to the davidi group, differs from other
CHENG : REVISION OF THE CHINESE MECOPTERA 45
described Panorpa by its golden body color and the peculiar
structures of both the male and the female genitalia.
Panorpa coomani n. sp.
Figures 63, 79, 282
Body dull brown ; vertex deep brown, with black mark en-
closing ocelli ; rostrum dull brown, on each of its sides a black
longitudinal stripe which narrows towards distal end ; thorax
deep brown dorsally, meso- and metanotum with black marking
on each side ; the 1st to 3rd abdominal segments of male slightly
blackish brown dorsally, the rest of segments reddish brown,
6th to 8th segments much prolonged, the hind border of the 3rd
tergite slightly produced. Fore wing: length, 11.3 mm.; width,
3 mm.; membrane hyaline, markings sooty brown; pterostigmal
band complete, with broad basal branch and apical branch;
basal band interrupted medianly ; apical band large, with a large
hyaline spot ; both basal and marginal spots present ; pterostigma
prominent. Hind wing: length, 10.5 mm.; width, 2.8 mm.; simi-
lar to fore wing, except that the basal spot is not so well de-
veloped. Male genitalia : genital bulb elliptical ; coxopodites long,
with truncated apex ; harpagones slender, the outer margin not
concave at the middle, inner margin with a median angle and a
median toothed lobe ; hypandrium very short ; hypovalvae nar-
row and slender, not extending near the base of the harpagones ;
parameres Y-shaped, the inner branch very narrow, the outer
branch broader and longer with a row of short barbs on its distal
inner margin; preepiproct broad basally, narrow distally, with
deep V-shaped distal incision ; aedeagus elongated, the apical
processes long, finger-like, lateral processes short, extended
downward, just opposite to the direction of the apical processes.
Female unknown.
Holotype ( $ ) : Ku-ling, Kiangsi, Sept. 18, 1945 ; in Heude
Museum, Shanghai.
I take the liberty to name this species in honor of Father De
Cooman, who has been so kind as to loan me the material from
the Heude Museum.
This species, belonging to the davidi group, differs from other
described Panorpa by its small body size and the peculiar struc-
ture of the male genitalia.
[6 BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY
Panorpa japonica Thunberg
Figures 85, 90, 100, 104, 286
Panorpa japonica Thunberg, 1784, Nov. Ins. Sp. Dissert., 3:67, fig. 9. Bur
meister, 1839, Handb. Ent.. 2:957. Westwood, 1846, Trans. Ent. Soc
London, 1846:188. MeLachlan, 1868, Journ. Linn. Soc, 9:256. Id.,
1875, Trans. Ent. Soc. London, 1875: 183. Miyake, 1908, Bull. Coll.
Agr. Imp. Univ. Tokyo, 1908:1. Id., 1913, Journ. Coll. Agr. Imp. Univ.
Tokyo, 1913: 347, pi. 30, fig. 14, pi. 3-",, figs. 1, 2, 3, 4, 5, 6. Esben
Peterson, 1921, Coll. Zool. Selys Long. 5 (2) :43, figs. 45, 46.
Panorpa macrogastcr MeLachlan, 1868, Journ. Linn. Soc, 1868:257. Id..
1875, Trans. Ent. Soc London, 1875:184.
Panorpa Icucothyria Navas, 1908, Mem. Real. Acad. Cienc. Barcelona,
1908:414.
Panorpa dyscola Navas, 1908, Mem. Real. Acad. Cienc. Barcelona, 1908:420.
Panorpa rectifasciata Miyake, 1908, Bull. Coll. Agr. Imp. Univ. Tokyo.
1908:5, pi.. 1, figs. 10, 10a, 10b. Id., 1913, Journ. Coll. Agr. Imp. Univ.
Tokyo, 1913:350, pi. 30, fig. 16, pi. 35 figs. 7, 8.
Panorpa niphonensis Miyake, 1908, Bull. Coll. Agr. Imp. Univ. Tokyo,
1908:7, pi. 1, figs. 3, 3a, 3b.
Panorpa pulchra Miyake, 1908, Bull. Coll. Agr. Imp. Univ. Tokyo, 1908:8,
pi. 1, fig. 4. Id., 1913, Coll. Agr. Imp. Univ. Tokyo, 1913:349, pi. 30,
fig. 17, pi. 35, figs. 4, 7, 9.
Panorpa sinanoensis Miyake, 1909, Bull. Coll. Agr. Imp. Univ. Tokyo,
1909:4, pi. 1, figs. 7, 7a, 7b.
Panorpa liagrni Navas, 1909, Rev. Russe d'Ent., 9:276.
Panorpa irregularis Miyake, 1910, Journ. Coll. Agr. Imp. Univ. Tokyo,
1910:198, pi. 11, figs. 7, 7a, 7b.
Panorpa japonica subsp. macrogastcr Miyake, 1913, Journ. Coll. Agr. Imp.
Univ. Tokyo. 1913:348.
Aulops intcrrupta Navas, 1913, Kev. Russe d'Ent., 13:283, fig. 11.
Body mostly deeply black, often shining ; the hind border of
third abdominal tergite of male produced into a short and broad
lobe, 6th abdominal segment cylindrical, 7th segment as long as
6th, but thinner, and its posterior angles somewhat produced,
forming a triangular tooth ; 8th segment much longer than 7th,
slightly thickened towards the apex, which is obliquely truncate
above; no anal horn present. Fore wing: length, 15-19 mm.;
broad, with rounded tips; membrane with slightly yellowish
tinge, markings sooty black ; pterostigmal band complete, with a
very broad basal branch and a very narrow apical branch; in
CHENG : REVISION OF THE CHINESE MECOPTERA 47
some specimens, the latter is absent, or present either as a com-
plete, curved streak or as a spot at the hind margin ; apical band
very broad, the inner margin somewhat concave; in some speci-
mens the pterostigmal band and the apical band may be tra-
versed longitudinally by a pale line between each of the longi-
tudinal veins ; in the apical band these pale lines are divided by
the darker crossveins; basal band either as a complete, oblique
band or as one or two separated spots ; basal spot mostly absent,
but in some strongly marked specimens, a basal spot, sometimes
isolated and sometimes connected with the basal band ; mar-
ginal spot present mostly, but usually a little separated from
the margin ; pterostigma not very prominent ; veins blackish
brown. Hind wing: length, 14-18 mm.; similar to the fore. Male
genitalia : genital bulb oval ; coxopodites not very long ; harpa-
gones long and slender, the outer margin smoothly curved, inner
margin more or less uneven ; the median tooth is very close to
the base of the harpagones, the basal lobe very small and tri-
angular ; hypandrium conspicuous, appearing as a long narrow
stalk; hypovalvae narrow, short and thick, usually divergent
from each other, nearly reaching to the base of the harpagones ;
parameres simple, short rod-like ; preepiproct tongue-shaped,
with rounded apex; aedeagus with a pair of peculiar, weakly
sclerotized, hairy, flattened structures and a pair of strongly
sclerotized club-shaped processes; lateral processes of the aedea-
gus not distinct. Female genitalia : subgenital plate rather long,
with V-shaped distal incision; internal skeleton small, long
U-shaped, the plate not highly sclerotized, with a rounded ante-
rior margin, axis not present.
Type: Japan; in Zool. Mus. Univ. Upsala.
Distribution : Tien-tseun, China ; Gifu, Japan, April-May
1886; Yokoama, Japan; Higo, 1906, Japan; Kumamoto, Japan,
April 17, 1913.
This species, belonging to the davidi group, is very common in
Japan. Issiki has established a japonica group which included
this species, Panorpa klugi MeLachlan, P. nipponensis Navas and
P. obscura Miyake. Apparently japonica resembles in general
appearance these three Japanese species. However, the wing
membranes of klugi and nipponensis are strongly yellowish and
that of obscura is ochraceous yellow, whereas that of japonica is
48 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
only slightly yellow. The male genitalia of japonica also show
some differences from those of the other three allied species. It
is interesting to note that the aedeagns of the male and the
internal skeleton of the female of japonica are similar to those of
most Neopanorpa.
Panorpa tetrazonia Navas
Figures 91, 92, 93, 94
Panorpa tetrazonia Navas, 1935, Notes d'Ent. Chin. Mus. Heude, 2(5) :96,
fig. 61. Carpenter, 1945, Psyche, 52(1-2) :71, pi. 10, figs. 1, 5, 6; pi.
11, fig. 10.
Body light to dark brown ; vertex black anteriorly, light brown
posteriorly ; the thoracic nota and abdominal tergites being some-
what darker than the rest of the body ; anal horn of male absent.
Fore wing : length, 12-13 mm. ; width, 3-3.5 mm. ; membrane
faintly yellow, markings brown ; pterostigmal band complete,
with broad basal branch and a narrow apical branch ; apical
band interrupted posteriorly and usually with a few small clear
spots around the crossveins; basal band complete but slender;
basal spot present ; marginal spot elongate ; crossveins not mar-
gined. Hind wing : similar to the fore, except that the basal band
is interrupted and the basal spot is absent. Male genitalia :
genital bulb oval ; coxopodites long, U-shaped ; harpagones mod-
erately long, the outer margin not concave, inner margin with
prominent lobe ; hypandrium inconspicuous ; hypovalvae broad
and short, not extending as far as the bases of the harpagones;
parameres conspicuous, each arising from a very slender stalk
which widens abruptly and gives rise to a long curved process;
the wide head of the stalk and the curved process bear numerous
long barbs; preepiproct with a shallow distal concavity; apical
processes of aedeagus rather long, with narrower distal ends;
lateral processes very short. Female genitalia : subgenital plate
slender; internal skeleton with broad plate and short axis, pos-
terior arms slender, slightly convergent distally.
Holotype ( $ ) : Killing, Kiangsi; in Heude Museum, Shanghai.
Distribution: Kuling, Kiangsi; Taiping-shien, Anhwei, Oct.
1932 (G. Liu) ; Huang-shan (few miles southwest of Taiping-
shien, Anhwei).
CHENG : REVISION OF THE CHINESE MECOPTERA 49
This species, belonging to the davidi group, differs from all
the formerly described Panorpa by its genital structures. The
parameres of the male are most unusual, as is also the form of
the internal skeleton of the genital segment of the female.
Panorpa sexspinosa Cheng
Figures 81, 87, 89, 123, 124, 278
Panorpa sexspinosa Cheng, 1949, Psyche, 56(4) :145, figs. 4, 8, 9, 15, 16.
Vertex yellowish brown, with four black spots on its anterior
region, one small spot enclosing the median ocelli anteriorly, one
around the other two ocelli posteriorly, the other two are on both
sides of the former two spots; rostrum uniformly yellowish
brown ; thorax blackish brown dorsally, light yellow laterally,
meso- and metanotum as a rule with broad median light yellow-
ish streaks; abdominal segments blackish brown dorsally, light
brown ventrally, the hind part of 6th abdominal segment of male
and its last few abdominal segments yellowish brown, the hind
border of the third tergite with a band-like prolongation. Fore
wing : length, 12 mm. ; width, 3 mm. ; membrane hyaline, mark-
ings blackish brown; pterostigmal band complete, with a broad
basal branch and a narrow apical branch ; basal band unusually
broad ; apical band complete, with a hyaline spot ; basal spot very
small ; pterostigma not very prominent. Hind wing : length,
10.8 mm. ; width, 3 mm. ; similar to fore wing, except that the
basal spot is lacking. Male genitalia: genital bulb rounded;
coxopodites long, with six spines on its distal inner margin;
harpagones slender, the outer margin smoothly curved, inner
margin with a reduced median angle and a large basal concave
area ; hypandrium inconspicuous ; hypovalvae rather short, not
nearly reaching to the base of the harpagones; parameres nar-
row and slender, each consisting of a single stalk which is some-
what twisted and pointed at its tip ; preepiproct slender, the dis-
tal incision being almost quadrate ; apical processes of aedeagus
somewhat prolonged on its distal outer margins, lateral processes
well-developed. Female genitalia : subgenital plate elongate,
slightly emarginate posteriorly ; internal skeleton large, the plate
distinctly concave at its base, with a pair of sharp posterior arms
and a pair of anterior side plates ; axis well-developed, extending
beyond the plate for nearly one-third its length.
50 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Holotype ( $ ) : Mt. Taipai, Shensi, June, 1942 (Io Chou) ; in
Cheng Collection, Taipeh. Allotype ( 9 ) : same collecting data
as holotype; in Museum of Comparative Zoology.
Distribution : same as types.
This species, belonging to the davidi group, differs from the
formerly described Pernor pa in its wing markings ; the basal band
is as broad as in Panorpa dado cere a Navas, but its pterostigmal
band is quite different. The structure of the male genitalia,
especially the six spines on the distal coxopodites, makes its
recognition easy.
Panorpa tincta Navas
Figure 78
Panorpa tincta Navas, 1931, Rev. Acad. Cienc. Madrid, 26:75, fig. 13.
Vertex black; rostrum dull yellow; thorax black; abdomen
tawny yellow ventrally, black dorsally with fine dull yelloAV
hairs on the straight posterior border of the tergites; 6th ab-
dominal segment cylindrical, brownish yellow, somewhat nar-
rowed towards apex where it is obliquely cut off dorsally; 7th
segment narrow at the base, cylindrical and black in color
dorsally, with a sudden swelling near the middle, the upper
border of the swollen part being slightly concave, the tip oblique,
the lower border convex basally and somewhat concave towards
the middle; 8th segment, with narrow cylindrical base, gradually
enlarges, the upper border being at first concave and then
slightly convex, while the lower border is almost straight. Fore
wing: length, 15 mm.; apex elliptieally rounded; membrane yel-
low, markings dark rust colored, indistinct ; pterostigmal band
complete with connecting basal branch and apical branch ; basal
hand well-developed, extending from the subcostal to the anal
margin of the wing and broadened towards the posterior end ;
apical band complete, sinuous on its inner margin ; basal spot
absent ; marginal spot present ; veins black in color ; pterostigma
rather prominent, dirty yellowish. Hind wing : length, 13.5 mm. :
similar to the fore, except that the basal band and the marginal
spot are absent. The £ genitalia have not been worked out.
However, according to Navas' original description, the pre-
epiproct (''upper cerei" of Navas) is long, slender and bowed;
CHENG: REVISION OP THE CHINESE MECOPTEKA 51
the hypovalvae ("lower cerci" of Navas), oblong, with rounded
lips, are almost in contact with each other and have nearly
parallel margins.
Female unknown.
Type ( $ ) : Hweihsien, Kansu; M. II.; in Hamburg Museum.
Distribution : same as type.
This species, having a deep yellow wing membrane, belongs
to the davidi group. It differs from the other species with yellow
wing-membranes by its indistinct wing markings and the well-
developed basal band. The shape of the 6-8th abdominal seg-
ments also makes its recognition easy. I have not seen this
species. The above description is based upon Navas' original
description.
Panorpa lutea Carpenter
Figures 107, 112
Panorpa lutea Carpenter, 1945, Psyche, 52(1-2) :72, pi. 10, fig. 7, pi. 11.
fig. 11.
Body reddish brown; darker brown on vertex, thoracic nota
and abdominal tergites. Fore wing : length, 15 mm. ; width, 3.5
mm. ; membrane deep yellow or orange, markings blackish brown;
pterostigmal band complete, with broad basal branch and broad
apical branch ; apical band separated by a wide hyaline stripe
into a large anterior apical area and a small posterior spot ; basal
band complete and very broad ; basal spot present ; marginal
spot rectangular ; crossveins not margined ; Hind wing : similar to
fore wing. Female genitalia : subgenital plate broad ; internal
skeleton small, with a very short axis and convergent posterior
processes.
Male unknown.
Holotype ( 9 ) : Huang-shan, Anhwei (G. Liu) ; in Museum of
Comparative Zoology.
Distribution : Huang-shan, Anhwei ; Kinhua-shan, Anhwei,
Oct. 1932.
This species, which is not determined in the above grouping,
is unlike any other described Panorpa in the deep yellow color
of the wings, which have the basal spot present and a complete,
forked pterostigmal band.
52 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Panobpa klapperichi Tjeder
Figures 120, 121, 126
Panorpa TclapperioM Tjeder, 1950, Bonn Zool. Beitr., 1950 (2-4) :289, figs.
3, 4.
Head blackish brown, rostrum light brown; pronotum black-
ish brown, meso- and metanotum light brown with darker spots
at the bases of the fore and hind wing ; abdomen blackish
brown dorsally, slightly lighter ventrally, the apex of the abdo-
men yellowish brown; additional side plates of the 7th and 8th
segments large; 9th tergite long and broad, its lateral borders
bent ventrad to embrace the side borders of the subgenital plate.
Fore wing : length, 13 mm. ; width, 3.5 mm. ; membrane hyaline,
markings blackish brown ; veins yellowish ; pterostigmal band
complete, with broad basal branch and narrower apical branch
apical band large, with two or three indistinct small windows
basal band narrow; both basal spot and marginal spot present
pterostigma prominent. Hind wing : length, 11.5 mm. ; width,
3.5 mm. ; similar to fore wing, except that the basal band is in-
terrupted, represented only as a spot at the hind margin of the
wing and the basal spot is entirely absent. Female genitalia:
according to Tjeder 's drawings, the subgenital plate is long and
narrow, concave at the middle, the apex slightly concave; in-
ternal skeleton very small, located at the posterior half of the
subgenital plate, posterior arms narrowed towards apex, the axis
extremely small and obliquely placed, so that their proximal ends
point obliquely upward, the proximal ends not extending beyond
the plate.
Male unknown.
Holotype ( 9 ) : Kwangtseh, Fukien, Oct. 9, 1937 (J. Klap-
perich) ; in Museum A. Koenig, Bonn.
This species, which I have not seen, differs from all the
formerly described species by the internal skeleton, which is very
small in proportion to the subgenital plate and also by the
peculiar small axis. The position of this species in the above
grouping is not determined.
CHENG : REVISION OF THE CHINESE MECOPTERA 58
Panorpa semifasciata Cheng
Figures 113, 114, 122, 274
Panorpa semifasciata Cheng, 1949, Psyche, 56(4) :146, figs. 19, 20, 21, 53.
Body entirely sooty black; vertex black; rostrum uniformly
black ; the middle part of the 8th abdominal tergite slightly pro-
longed into a band-like extension, the 9th tergite very broad, its
lateral borders bent ventrad to embrace the posterior part of
the subgenital plate in ventral view. Fore wing : length, 14 mm. ;
width, 3.5 mm.; membrane light yellow, markings sooty brown;
pterostigmal band incomplete, with an interrupted narrow basal
branch; apical band small, with two hyaline spots; pterostigma
prominent. Hind wing : length, 12.8 mm. ; width, 3 mm. ; similar
to fore wing, except that the basal branch of pterostigmal band
is greatly reduced. Female genitalia: subgenital plate broad,
with strongly sclerotized median part and less sclerotized narrow
borders, apex of subgenital plate protruded, rounded, less sclero-
tized, furnished with several long hairs ; the sides of the plate are
enclosed by the well-developed 9th tergite as mentioned above;
internal skeleton flattened, the plate very small, less sclerotized;
the posterior arms of the internal skeleton very long, sharp and
strongly sclerotized, the anterior arms flattened, joined with the
posterior arms and extending a little beyond the plate.
Male unknown.
Holotype ( 9 ) : Jihti (30 miles east of Tachienlu), Sikang,
Sept. 1, 1939 (F. Y. Cheng, Io Chou and Tein Ho Hei) ; in Cheng
Collection, Taipeh.
This species differs from all the formerly described species by
its black body color, reduced wing markings and the peculiar
shape of the genital segment of the female. The position of
this species in the above grouping is not determined.
Panorpa leei Cheng
Figures 125, 127, 275
Panorpa leei Cheng, 1949, Psyche, 56(4) :147, figs. 17, 18, 54.
Vertex black; rostrum reddish brown, with a short and deep
brown stripe on each side of its upper portion; thorax black
dorsally, yellowish brown laterally; 1st to 6th abdominal seg-
54 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
ments black dorsally and ventrally, the 7th to 9th abdominal seg-
ments very small, reddish brown. Fore wing : length, 14 mm. ;
width, 4 mm. ; membrane hyaline, markings sooty brown ; ptero-
stigmal band broad, with a complete basal branch and a greatly
reduced spot-shaped apical branch ; apical band small, including a
prominent narrow band and some faintly smoky spots; ptero-
stigma prominent. Hind wing : length, 13 mm. ; width, 3.55 mm. ;
similar to fore wing, except that the basal branch of pterostigmal
band is greatly reduced. Female genitalia : subgenital plate
small, narrowed posteriorly, apex rounded ; internal skeleton
long, the plate abruptly narrow at the base, with a pair of sharp
posterior arms, the axis extending for nearly half its length
beyond the plate.
Mabj unknown.
Holotype ( 9 ) : Mt. Taipai, Shensi, July 14, 1943 (Chuan
Lung Lee) ; in Museum of Comparative Zoology.
This species differs from all the formerly described species by
its wing markings and the peculiar shape of the genital segment
of the female. The position of this species in the above grouping
is not determined.
Panorpa graham ana n. sp.
Figures 108, 115
Body mostly black ; vertex black ; rostrum brownish black, with
an orange median longitudinal stripe ; the 9th abdominal tergite
very broad, its lateral borders bent ventrad to embrace the
margins of the subgenital plate in ventral view. Fore wing :
length, 13 mm. ; width, 3.3 mm. ; membrane hyaline, markings
sooty brown ; pterostigmal band broad, with a broad basal branch,
but no apical branch ; basal band interrupted, represented as a
large spot, extended to the hind margin of the wing ; apical band
broad, a little interrupted anteriorly; both basal and marginal
spots absent; pterostigma rather prominent. Hind wing: length,
12 mm. ; width, 3.2 mm. ; similar to fore wing, except that the
basal band is entirely absent. Female genitalia : subgenital plate
slender, tongue-shaped, with strongly sclerotized median part
and less sclerotized narrow lateral plates ; apex of subgenital
plate rounded, less sclerotized, furnished with some prominent
CHENG : REVISION OF THE CHINESE MECOPTERA 55
hairs; the sides of the plate enclosed by the well-developed 9th
tergite; internal skeleton flattened, with long posterior arms
which are narrowed towards apex, anterior arms of the plate
slightly outwardly curved.
Male unknown.
Holotype ( 9 ) : Suifu, Szechwan, (D. C. Graham) ; in Museum
of Comparative Zoology.,
This species is named in honor of D. C. Graham. It is close
to Panorpa semifasciata in the female genitalia, but differs
greatly in the wing markings. The apex of the subgenital plate
of this species is broadly rounded, whereas that of semifasciata
is protruded and narrowly rounded. The position of this species
in the above grouping is not determined.
Panorpa carpenteri n. sp.
Figure 116
Body mostly black ; vertex black ; rostrum uniformly reddish
brown ; the 9th abdominal tergite very broad, its lateral borders
bent ventrad to embrace the margins of the subgenital plate in
ventral view. Fore wing: length, 13.2 mm.; width, 3.2 mm.;
membrane hyaline, markings soot}7 brown; pterostigmal band
broad, with a broad basal branch, but no apical branch; basal
band represented as a small spot ; apical band broad, interrupted
posteriorly; both basal spot and marginal spot absent; pterostig-
ma rather prominent. Hind wing : length, 12 mm. ; width, 3.2 mm. ;
similar to fore wing, except that the basal branch of pterostigmal
band is narrower than that of the fore wing and the basal band
is entirely absent. Female genitalia: subgenital plate slender,
tongue-shaped, with strongly sclerotized median part and less
sclerotized narrow lateral plates, apex of subgenital plate
rounded, less sclerotized, furnished with some prominent hairs ;
the si. it's of the plate enclosed by the well-developed 9th tergite;
internal skeleton flattened, the plate greatly reduced, with very
long and sharp posterior arms and well-developed anterior arms ;
the latter are folded transversely right at its median portion.
Male unknown.
Holotype ( 9 ) : foot of Mt. AVa (6000-7000 ft.), Szechwan,
July 27, 1925 (D. C. Graham) ; in Museum of Comparative
Zoology.
56 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
This species is named in honor of Professor F. M. Carpenter,
who has been so kind to me. It resembles the preceding species
in both body color and wing markings, but differs in the color
of the rostrum and the anterior arms of the internal skeleton of
the female genitalia. The position of this species in the above
grouping is not determined.
Panorpa statura Cheng
Figures 109, 110, 279
Panorpa statura Cheng, 1949, Psyche, 56(4) :148, figs. 32, 33, 34, 57.
Vertex blackish brown anteriorly, with a black mark within
the ocelli, light brown posteriorly, with a median and a pair of
longitudinal bands ; rostrum uniformly reddish brown ; thorax
entirely brown laterally, prothorax blackish brown dorsally,
meso- and metanotum uniformly blackish brown ; 1st to 4th
abdominal segments of female blackish brown dorsally, brown
ventrally, last few abdominal segments entirely brown. Fore
wing: length, 16.5 mm.; width, 4.55 mm.; membrane deeply
yellowish brown, markings deep brown ; pterostigmal band com-
plete, with a broad basal branch and a broad apical branch ; basal
band interrupted; apical band large, with a hyaline spot; basal
spot absent; marginal spot very small; pterostigma not very
prominent. Hind wing : length, 15 mm. ; width, 4.2 mm. ; similar
to fore wing, except that the small marginal spot is lacking.
Female genitalia: subgenital plate elongated, narrowed pos-
teriorly, shallowly emarginated at its apex, its lateral borders
bent to form a narrow ridge; internal skeleton long, the plate
concave on its median sides with a pair of short tooth-like
posterior arms; the axis long, extending beyond the plate for
exactly half its length.
Male unknown.
Holotype ( 5 ) : Mt. Taipai, Shensi, July 14, 1943 (Chuan
Lung Lee) ; in Cheng Collection, Taipeh.
This species, having a yellowish brown wing membrane, differs
from Panorpa flavipennis Carpenter by its very long wing and
the markings of the apical band. The peculiar shape of the
genital segment of the female makes its recognition easy. The
position of this species in the above grouping is not determined.
CHENG : REVISION VV THE CHINESE MECOPTEBA 57
Panorpa pusilla Cheng
Figures 118, 119
Panorpa pusilla Cheng, 1949, Psyche, 56(4) :149, figs. 37, 38, 52.
Vertex yellow anteriorly, with a black spot enclosing ocelli,
sooty brown posteriorly, with a median quadrangular plate;
rostrum uniformly yellow ; thorax brownish yellow dorsally, yel-
low laterally, meso- and metanotum with sooty brown markings
on each side; abdominal segments sooty brown dorsally, yellow
laterally and ventrally. Fore wing : length, 10.8 mm. ; width, 2.8
mm. ; membrane light yellow, markings sooty brown ; pterostigmal
band complete, with a complete basal branch and a separated
apical branch ; basal band complete ; apical band represented by
two prominent bands, the inner one narrow, being parallel to
the pterostigmal band, the outer one running along the wing
apex ; basal spot situated on the hind margin of wing ; marginal
spot very large ; pterostigma not very prominent. Hind wing :
length, 9.5 mm. ; width, 2.8 mm. ; similar to fore wing, except
that the basal spot on the hind margin of wing is entirely lack-
ing. The venation of both fore and hind wings identical; Sc,
as usual, does not extend to the pterostigmal area, Ri is forked
and Ro is simple, no crossvein between Rj and R2. Female
genitalia : subgenital plate elliptical, with a slightly distal
emargination ; the plate of the internal skeleton small, the pos-
terior arms of the plate large, twisted at the middle, the axis
short and slender, not extending beyond the plate.
Male unknown.
Holotype ( 5 ) : Mt. Taipai, Shensi, June, 1942 (Io Chou) 5 in
Museum of Comparative Zoology.
This species, having a light yellowish wing membrane differs
from the other described Panorpa by its very small body size,
wing markings and the peculiar shape of the genital segment of
the female. The position of this species in the above grouping
is not determined.
Panorpa pieli n. sp.
Figures 111, 117
Body yellowish white ; vertex brown, with small grayish brown
mark enclosing ocelli ; rostrum yellowish white, with brownish
58 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
yellow longitudinal stripe on each side ; thorax yellowish white,
meso- and metanotum with yellowish brown mark on each side;
the whole abdomen of female brownish yellow dorsally. Fore
wing: broad basally, rounded apically; length, 11 mm.; width
3.2 mm. ; membrane yellowish white, markings gray ; pterostig-
mal band complete, with a basal branch and a separated apical
branch ; basal band represented by two spots ; apical band repre-
sented by an inner spot and an outer band which is enclosed by
the wing apex ; basal spot absent ; marginal spot elongated ; ptero-
stigma not prominent. Hind wing: length, 10 mm.; width, 3.2
mm. ; similar- to fore wing, except that the basal band is repre-
sented by only one spot. The venation of fore and hind wings
identical, in both pairs of wings Sc, as usual, not extending to
the pterostigmal area; Rj is forked and R2 is simple, no cross-
vein between R3 and R2. Female genitalia: subgenital plate
broad, with a wide V-shaped distal incision; internal skeleton
small, the plate rather broad, with long posterior arms which
are pointed towards apex ; the axis short and slender, not ex-
tending beyond the plate.
Male unknown.
Ilolotype ( 9 ) ; Ku-ling, Kiangsi, Aug. 18, 1943 (Piel) ; in
ileude Museum, Shanghai.
The material was collected by Dr. Piel, in honor of whom I
name the species.
This species resembles Panorpa pusilla Cheng in body size,
wing venation and wing markings. It is evident that they are
closely allied ; but the differences in the structure of the geni-
talia and in the color of the vertex and rostrum are so conspicu-
ous that there is guod reason to distinguish it as a good species.
The position of this species in the above grouping is not de-
termined.
Panorpa bonis Cheng
Figures 102, 106
Panorpa bonis Cheng, 1949, Psyche, 56(4) :150.
I'anorpa comigera Tjeder {nee MeLachlan), 1936, Ark. for Zool. 27A
(33): 7, pi. 3, pi. 7, fig. 3.
The body characters of this species agree wholly with the
CHENG: REVISION OF THE CHINESE MECOPTEBA 59
original description for Panorpa cornigera McLachlan accord-
ing to Tjeder. The wing-photo of this species is exactly the same
as that of the 9 type of cornigera (given by Esben-Petersen)
as Esben-Petersen agreed. The detail drawings of 9 genitalia
were given by Tjeder. According to these drawings, the sub-
genital plate is oval with smoothly rounded side-margins, which
slightly overlap the lower margins of the 9th tergite and its apex
is very shallowly emarginated ; the internal skeleton large but
slender, the two inner pairs of the plate prominent, their proxi-
mal part appearing to end straight ; posterior arms of the plate
with acute distal ends; axis long, extending more than one third
its length beyond the plate.
Male unknown.
Ilolotype ( 9 ) : Lu-pa-sze (at river Tao-ho, about 2750 m.)
South Kansu, July 11, 1030 (D. Hummel); in Stockholm Mu-
seum.
Distribution : same as type.
This species resembles Panorpa cornigera McLachlan super-
ficially, but differs in the structure of the female genitalia. The
additional lateral plates of the 7th-8th abdominal segments of
this species are not so slender as those of cornigera. The sub-
genital plate is pointed at its posterior part and shallowly emar-
ginate at its apex, while that of cornigera is rounded and not
emarginate. The internal skeleton of this species is quite distinct
from that of cornigera: the plate of the former is slender with
a small proximal part and short posterior arms, while that of
the latter is broad, with a well-developed oval proximal part and
long pointed posterior arms. The axis of this species extending
beyond the plate is less than half the length of the whole axis,
while that of cornigera usually extends beyond the plate more
than half its length. The position of this species in the above
grouping is not determined.
Panorpa guttata Navas
Panorpa guttata Xavas, 1908, Mem. Real Acad. Cienc. Barcelona, 1908:416.
fig. 19c. Esben-Petersen, 1921, Coll. Zool. Selys Long. 5(2) :32, fig. 32.
Panorpa davidi ( 9 ) Navas, 1908, Mem. Eeal Acad. Cienc. Barcelona, 1908:
415, fig. 19c (nee davidi Navas, figs. 19a, b).
Head and rostrum grayish testaceous; vertex with a blackish
60 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
spot between the ocelli; thorax and abdomen pale castaneous
dorsally, pale grayish yellow ventrally. Fore wing : length, 12.5
mm. ; membrane hyaline with a faint yellowish tinge ; pterostig-
mal band indicated by a spot posterior to the pterostigma ; apical
band represented as three faint spots; pterostigma prominent,
yellowish ; veins brownish. Hind wing : length, 11 mm. ; similar
to fore wing, except that the faint apical band is absent.
Male unknown.
Type ( 5 ) : Mou Pin, Tibet, 1870 (David) ; in Museum Na-
tional d'Histoire Naturelle, Paris.
Distribution : same as type.
This species resembles Panorpa tjederi ( 5 ) in the pterostigmal
band of the fore wing, but differs in the presence of its apical
band. The wings of tjederi are subobtuse, whereas those of gut-
tata are very narrow and slender. However, a thorough exami-
nation of the genitalia of the female type would be highly
desirable. The position of this species in the above grouping
is not determined.
Genus NEOPANORPA Weele
Neopanorpa Weele, 1909, Notes Leyden Mus. 31:4. Esben-Petersen, 1913,
Notes Leyden Mus. 35:226. Id., 1921, Coll. Zool. Selys Long. 5(2) :73.
Campodotecnum Enderlein, 1910, Zool. Anz., 35:391. Id., 1912, Notes Leyden
Mus. 34:235.
Rostrum long and slender; tarsal claws serrated on inner
margins; wings are fully developed, rather narrow, especially
at the base; 1A short, extending to the anal margin of wing
before origin of the radial sector; abdomen in both sexes not
longer than the wings; 6th to 8th abdominal segments of male
normal, not much prolongated ; genital bulb of male not peduncu-
late basally.
Genotype: Neopanorpa angustipennis "Westwood.
This genus, common in southeast Asia, includes forty known
species in the whole world. Nineteen species have been already
recorded in China and eleven new ones are described below,
making a total of thirty. They are distributed throughout nine
provinces. Since no Neopanorpa have been found in North China
(Shensi, Kansu) and Korea, I presume therefore that this genus
is restricted to North Asia. The limit seems to be approximately
along latitude 40°.
CHENG : REVISION OF THE CHINESE MECOPTERA 61
The most obvious difference between Neopanorpa and Panorpa
is the length of 1A, as indicated in the key for the family
Panorpidae. However, differences are also apparent in the 3rd
abdominal tergite and in both male and female genitalia. In the
male of Neopanorpa the median process of the 3rd abdominal
tergite tends to be longer and more slender than that of Panorpa.
In most of the species of Neopanorpa, this process extends to the
middle of the 4th tergite or beyond the hind border of the latter
(the only exception is claripennis, the process of which is not
much prolonged and appears as a semicircular lobe). In most
species of Panorpa, this process is not distinct, only prolonged
into a small semicircular lobe (the only exception is stigmalis,
the process of which extends to the hind portion of the 4th tergite
but not beyond the hind border of the latter). In the male of
Neopanorpa the hypovalvae are broad, mostly overlapping each
other distally, whereas those of Panorpa are slender, never over-
lapping each other distally. The parameres of Neopanorpa are
mostly absent or reduced to a short slender thread-like rod, some-
times branched, without barbs or hairs, and partly or wholly
fused with the aedeagus, whereas those of Panorpa are well de-
veloped, prominent and mostly with barbs or hairs on their inner
margins, absolutely free from the aedeagus. The only exception
is japonica which has reduced rod-shaped parameres and is close
to those of Neopanorpa. The aedeagus of male Neopanorpa is
mostly very small, and the paired apical processes are short,
almost united together, whereas those of Panorpa are very prom-
inent, the two apical processes being very long and wide apart
from each other (except in the diceras group, the aedeagus of
which has united and short apical processes). The preepiproct
of most male Neopanorpa has a rounded distal margin (heii etc.,
fig. 156) ; in some species it is slightly emarginated (translucida
n. sp., fig. 223), and in others it has distal processes which are
directed inward towards the interior of the bulb (pilosa, fig. 192,
taoi, fig. 159). In Panorpa, the preepiproct has a deep U-shaped
distal incision, although in waongkehzengi it is slightly emar-
ginate and in japonica rounded. This again shows that japonica
is close to Neopanorpa. In the female of Neopanorpa, the sub-
genital plate is deeply emarginate distally (except that of
kwangtsehi n. sp., which is truncated), whereas that of Panorpa
62 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
has either a rounded apex or is slightly emarginate; the only
exception is in japonica, in which the subgenital plate is as deeply
emarginate as that of the Neopanorpa.
The larva of Neopanorpa has very short setae, whereas that of
Panorpa has much longer setae. The larval head sutures of Neo-
panorpa are usually accompanied by a broadly sclerotized band,
while those of Panorpa are simple. Moreover, along the posterior
margin of the 10th abdominal segment of the larval Neopanorpa
there are numerous hairs, absent in Panorpa.
The characteristics which have been used for the specific identi-
fication of Neopanorpa are the median process of the 3rd abdom-
inal tergite of the male and the wing markings and both the male
and female genitalia. In some species the median process of the
3rd tergite of the male is very short, far from reaching to the
middle of the fourth tergite (claripennis, fig. 132) ; in others it
is very long, extending far beyond the hind border of the 6th
abdominal segment (choui, fig. 155). The color of wing mem-
brane is useful. In some species, the wings are hyaline {nigritis,
fig. 287), in others, they are deep yellow {caveata n. sp., fig. 290)
and in some others, they are faintly yellow {cavaleriei) . The
markings of the wings are different from species to species. In
claripennis, faoi, pilosa, nigritis and validipennis, there are no
markings at all. In apicata only a shadoAv of dark appears at the
wing apex. In choui and kwangtsehi n. sp.. the markings are
very indistinct, while in some others they are sooty brown and
very extensive (cantonensis n. sp., fig. 302).
As in the genus Panorpa, both the male and the female geni-
talia of N eopanorpa remain perfectly stable even in the minute
details. The general structures of both sexes are just the same as
those of Panorpa. In the male of Neopanorpa, the outer margins
of the harpagones are mostly concave at the middle {caveata
n. sp., etc., fig. 133) ; in others they are distinctly convex (taoi,
fig. 157, pilosa, fig. 193). The hypandrium is usually long and
broad {caveata n. sp., etc., fig. 133), but in nigritis and choui, it
is very inconspicuous. The hypovalvae are mostly short and
broad, usually tending to overlap each other distally {claripen-
nis, etc., fig. 136), but in some species, they are narrow and slen-
der (nigritis, fig. 191, mutabilis n. sp., fig. 141). The parameres
are absent in most of the species (heii etc., fig. 162), in others,
they are small and rod-shaped {claripennis, etc., fig. 136) and in
CHENG : REVISION OF THE CHINESE MECOPTERA 63
some others, they are branched (pilosa, etc., fig. 193). In the
female of Neopanorpa, the subgenital plate is distinctly emar-
ginate distally. This emargination is diversely shaped in differ-
ent species. In parva, etc., it is wide and deep (fig. 184). In
pulchra, etc., it is slightly concave (fig. 182). In translucida n.
sp., etc., it is V-shaped (fig. 197 ) . In banksi, etc., it is U-shaped (fig.
177), and in kwangtsehi n. sp., it is exceptionally truncated, -with-
out emargination at all (fig. 198). The internal skeleton of the
female Neopanorpa is usually U-shaped, with the axis absent,
but in caveata n. sp. nigritis, choui, kwangtsehi n. sp., translucida
n. sp., pielina, mutdbilis n. sp. and maai n. sp., the axis is promi-
nent and projects beyond the plate of the internal skeleton. The
following keys to both male and female Neopanorpa are based
upon the characteristics mentioned above.
Key to the Males of Neopanorpa
The males of the following species are unknown : dimidiata
Xavas, banksi Carpenter, parva Carpenter, pulchra Carpenter,
Jatipennis Cheng, varia Cheng, chaoi n. sp., cantonensis n. sp.,
kwangtsehi n. sp., carpenteri n. sp.
1. Wing membrane deep yellow 2
Wing membrane slightly yellow or hyaline 4
2. Apical band large, with four hyaline spots ; hypovalvae stout, the inner
margins rather straight, the distal portions slightly separated from
each other (figs. 133, 290) caveata n. sp.
Apical band smaller, with one or two hyaline spots ; hypovalvae over-
lapping each other distally 3
3. Preepiproct with a small U-shaped distal incision; the outer margins
of the hypovalvae concave at the middle, inner margins straight, each
with a proximal lobe (fig. 134) tienmushnno, n. sp.
Preepiproct almost truncated ; the outer margin of the hypovalvae
smoothly curved, inner margins without the proximal lobe (fig. 135)
hyangshana n. sp.
4. Wings without color markings 5
Wings with color markings 9
5. Median process of third abdominal tergite very short, not extending to
middle of the fourth tergite ; preepiproct with truncated apex
claripennis
.Median process of third abdominal tergite rather long, usually ex-
tending beyond the middle of the fourth tergite; preepiproct not
truncated at the apex 6
64 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
rt. The outer margins of harpagones convex near the base, basal lobe with
two tooth-like processes; preepiproct with a pair of long distal
processes 7
The outer margins of harpagones not convex at all, basal lobe rounded,
without a tooth-like process ; preepiproct without the long distal
processes 8
7. Parameres simple, leaf -shaped ; the outer margins of hypovalvae con-
vex basally (fig. 157) taoi
Parameres branched, both branches thread-like; the outer margins of
the hypovalvae concave at the middle (fig. 193) pilosa
8. Hypandrium short, inconspicuous ; hypovalvae slender, less sclerotized
and curved inward distally, separating each other, not reaching to
the base of the harpagones (fig. 191) nigritis
Hypandrium long, conspicuous ; hypovalvae broad, overlapping each
other, with slender basal stalk, extending far beyond the base of the
harpagones (fig. 145) validipennis
!». Wing markings represented only by an apical band: no pterostigmal
band present apicata
Both apical band and pterostigmal band present 10
1<>. Wing markings indistinct; median process of third abdominal tergite
extraordinarily long, measuring up to 4.2 mm., divided into two
portions; hypovalvae with an abruptly narrowed apex (figs. 155, 164)
ohoui
Wing markings distinct ; median process of third abdominal tergite not
very long; hypovalvae without abruptly narrow apex 11
11. Wing markings less developed, with a long narrow band and a sus-
picion of dark on the wing apex; wings with dark longitudinal
stripes among their veins and their branches ; 7th abdominal segment
of male with truncated apex (fig. 188) brisi
Wing markings well developed, with broad pterostigmal and apical
bands ; no longitudinal stripes occur among veins and their
branches 12
12. Pterostigmal. band with broad basal branch and a separate, narrow
apical branch; genital bulb as in figure 162 heii
Pterostigmal band with both basal and apical branches, apical branch
not separated from the pterostigmal band itself 13
13. Apical band interrupted posteriorly, without hyaline spots 14
Apical band not interrupted posteriorly, with hyaline spots 15
14. Fore wing length measures 14 mm. ; median process of the third ab-
dominal segment short, with rounded posterior margin, about half
the length of the fourth tergite; genital bulb as in figure 194. .ohelata
Pore wing length measures 16 mm.; median process of the third
abdominal segment narrow and long, almost as long as the fourth
tergite cavaleriei
CHENG : REVISION OF THE CHINESE MECOPTERA 65
15. No additional band occurs between pterostigmal band and apical
band 16
A narrow band occurs between pterostigmal and apical band 18
16. Basal band interrupted, represented by two spots which are separated
from the marginal spot lacunaris
Basal band not interrupted, but irregular, connected with the marginal
spot 17
1 7. Wing markings brown ; harpagones with large square-shaped basal
lobes; apex of hypovalvae rounded (fig. 148) translucida n. sp.
Wing markings sooty brown; harpagones without true lobes; apex of
hypovalvae more or less pointed in ventral view (fig. 151) . . . .pielina
18. Outer margins of hypandrium and hypovalvae slightly concave at the
middle; parameres present, consisting of a narrow stalk, which gives
rise to two branches (fig. 141) mutabilis n. sp.
Outer margins of hypandrium and hypovalvae abruptly concave at the
middle ; parameres absent 19
19. Hypovalvae slender, with rounded apex as in figure 143 . . . .maai n. sp.
Hypovalvae broad and stout, with tooth-like apex in ventral view as
in figure 149 ovata n. sp.
Key to the Females of Neopanorpa
The females of the following species are unknown: cavaleriei
Navas, lacunaris Navas, brisi (Navas) Carpenter, pilosa Car-
penter, validipennis Cheng, taoi Cheng, ovata n. sp.
1. Wing membrane deep yellow 2
Wing membrane slightly yellow or hyaline 4
2. Internal skeleton large, with long and stout axis which extends beyond
the plate nearly one-third its length (fig. 204) caveata n. sp.
Internal skeleton small, axis absent 3
3. Pterostigmal band with narrow apical branch; basal band interrupted;
subgenital plate with deep Y-shaped distal incision; internal skeleton
as in figure 171 tienmushana n. sp.
Pterostigmal band with broad apical branch; basal band complete;
subgenital plate with shallow distal incision; internal skeleton as
in figure 170 haangshana n. sp.
4. Wings without color markings 5
Wings with color markings 6
5. Internal skeleton with long axis as in figure 207 nigritis
Internal skeleton without axis as in figure 172 claripennis
6. Wing markings represented only by an apical band; no pterostigmal
band present apicata
Both apical band and pterostigmal band present 7
7. Wing markings indistinct 8
Wing markings distinct 9
66 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
8. Pterostignial band of fore wing complete; subgenital plate with trun-
cated apex ; internal skeleton as in figure 202, axis long, with rounded
basal ends Icwangtsehi n. sp.
Pterostignial band of fore wing incomplete, represented only by a faint
basal branch; subgenital plate with a narrow U-shaped distal
incision ; internal skeleton as in figure 154, axis long, with abruptly
curved hook-shaped basal ends dlwui
9. Pterostigmal band with same broad basal branch, but no apical branch;
internal skeleton as in figure 181 banksi
Pterostignial band with both basal and apical branches 10
10. Either basal branch or apical branch of pterostignial band inter-
rupted 11
Both basal branch and apical branch of pterostigmal band complete 14
11. Apical branch of pterostigmal band interrupted, very narrow; basal
branch complete 12
Apical branch of pterostigmal band complete; basal branch inter
rupted 13
12. Wing markings slightly gray; between the pterostigmal band and
apical band there is an additional band ; internal skeleton as in figure
200, axis straight maai n. sp.
Wing markings sooty brown, no additional band between pterostignial
band and apical band ; internal skeleton as in figure 212, axis fork
shaped heii
13. Apical band interrupted posteriorly; internal skeleton as in figure 214
varia
Apical band not interrupted posteriorly, with a faint hyaline spot
di/midiata
14. Apical band large, complete, without hyaline spot 15
Apical band more or less interrupted or with prominent hyaline spot. .16
15. Basal spot present; internal skeleton U-shaped as in figure 179
cantoi%ensis n. sp.
Basal spot absent; internal skeleton with widely divergent arms as in
figure 183 pulchra
1<>. Internal skeleton with long and paired axis 17
Internal skeleton without axis or with single, short axis 19
17. Wing markings brown; subgenital plate with deep V-shaped distal
incision; internal skeleton as in figure 201 ; the length of axis is nearly
the same length as the posterior arms translucida n. sp.
Wing markings sooty brown ; distal incision of the subgenital plate not
deep V-shaped ; the axis of the internal skeleton longer than the
posterior arms 18
18. Wing membrane slightly yellow; rostrum shining reddish brown; sub-
genital plate with shallow V-shaped distal incision; internal skeleton
CHENG : REVISION OF THE CHINESE MECOPTERA 6 i
as in figure 206 pielina
Wing membrane hyaline; rostrum deeply grayish brown; subgenital
plate narrow distally, with small U-shaped distal incision; internal
skeleton as in figure 205 mutabUis n. sp.
11'. Wing markings brown; internal skeleton V-shaped as in figure 178 ....
chaoi n. sp.
Wing markings sooty brown ; internal skeleton more or less U-shaped . . 20
20. Internal skeleton with very short median axis as in figure 185 . . . .parva
Internal skeleton without true axis 21
21. Subgenital plate abruptly narrowed posteriorly, with wide U-shaped
distal incision; internal skeleton with a long stalk at its base as in
figure 203 latipennis
Subgenital plate gradually narrowed posteriorly, with shallow V-shaped
distal incision; internal skeleton without long stalk at its base ... .22
•22. Wing length measures up to 16 mm.; wing membrane slightly yellow;
apical band with one hyaline spot posteriorly; outer margins of the
internal skeleton smooth as in figure 180 carpenteri n. sp.
Wing length measures only 1-1 mm.; Aving membrane hyaline; apical
hand interrupted posteriorly, without hyaline spot; outer margins
of the internal skeleton sinuous as in figure 173 chelate
Descriptions of Species of Neopanorpa
Neopanorpa caveata n. sp.
Figures 128, 129, 133, 137, 204, 208, 290
Body light brown ; vertex brown with a blackish brown mark
enclosing ocelli ; rostrum uniformly reddish brown ; thorax yel-
lowish brown laterally with six black spots, the middle of the
meso- and metanotum blackish brown ; in addition to mesonotum,
there is usually a blackish brown streak along its anterior side
margins; the 1st to 5th abdominal segments of male blackish
brown dorsally; 6th abdominal segment blackish brown with
reddish brown hind margin; last few segments reddish brown,
median process of the 3rd tergite short, never extending beyond
the hind margin of the 4th tergite and in contact with the conical
production on the median axis of the latter ; the 1st to 5th abdom-
inal segments of female blackish brown dorsally, last few seg-
ments slightly reddish brown. Fore wing : length, 15-16 mm. ;
width, 3.5 mm. (holotype, length, 15 mm. ; width, 3.5 mm.) ; mem-
brane yellow, markings deep brown ; pterostigmal band complete
with broad basal branch and apical branch ; basal band complete ;
68 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
apical band large, usually joined to the pterostigmal band by
some very narrow additional network-like bands so as to form
several hyaline spots; basal spot absent; marginal spot large,
band like ; pterostigma not very prominent. Hind wing : length,
13-14 mm. ; width, 3.4 mm. ; similar to fore wing, except that the
basal band is represented only by a spot, the network-like addi-
tional bands not so developed and the marginal spot entirely
lacking. Male genitalia : genital bulb slender ; coxopodites long
with truncated apex; harpagones slender; the outer margin
slightly concave at the middle, inner margin with a slightly pro-
truded median portion and a small true basal lobe ; hypandrium
very long ; hypo valvae stout, the outer margin smoothly curved
towards its apex, the inner margin rather straight, the basal
parts widely separated and the distal parts slightly separated
from each other, the apex of hypovalvae usually reaching to the
middle of the harpagones; parameres very short, basal portions
bending inward and then upward, the whole paramere fused
with the basal part of aedeagus; preepiproct abruptly narrowed
at the distal portion with truncated apex ; aedeagus very small,
the apical processes united together, lateral processes tooth-like,
usually bending upward, the base of aedeagus provided with a
pair of sclerotized elongate plates. Female genitalia : subgenital
plate rather broad with V-shaped distal incision ; internal skele-
ton large, the plate mostly occupied by the axis with U-shaped
posterior arms, which are somewhat twisted, the axis very large
and stout, extending beyond the plate nearly one-third its length.
Holotype ( c? ) : Ta-chu-lan, Shaowu Hsien, Fukien, June 13,
1945 (Maa) ; in Museum of National Foochow University, Foo-
chow. Allotype ( 9 ) : same locality as holotype ; June 10, 1944
(Maa) ; in Museum of National Foochow University, Foochow.
Paratypes : 1 $ , 10 9 9 , same locality as holotype, June 3-10,
1943-1945; 1 9 , same locality, Aug. 13, 1943; 1 9 , same locality,
Sept. 29, 1943 ; 1 9 , Tnng-mu-kwan, Chungan Hsien, Fukien,
May 17, 1945 ; 1 9 , San-chiang, Chungan Hsien, Fukien, Aug.
12, 1945 (Lin); 39 9, Chien-men; Kwantseh Hsien, Fukien,
4 5 $ , 74 9 9 , Ta-chu-lan, Shaowu Hsien Fukien, April 28-
Sept. 20, 1942-1945 (Maa) in Maa Collection; 1 $ , 19, Sien-
feng-ling, Chingan Hsien, Fukien, June 3-10, 1943, in Museum
of Comparative Zoology ; 1 S , 3 9 9 , Ta-chu-lan, Shoawu Hsien,
CHENG : REVISION OF THE CHINESE MECOPTERA 69
Fukien, May 8-June 13, 1942-1943 (Maa), in Cheng Collection,
Taipeh.
The color of the wing membrane of this species resembles
that of Neopanorpa ophthalmica Navas and sauteri Esben-Peter-
sen, but the markings are quite different. In ophthalmica and
sauteri, the apical band is completely separated from the ptero-
stigmal band and without rounded hyaline spots, whereas that
of caveat a n. sp. is usually connected with the pterostigmal band
by some network-like bands and forms several rounded hyaline
spots. The shape of the hypovalvae of the male and the very
large axis of the female also make its recognition easy.
Neopanorpa tienmushana n. sp.
Figures 130, 134, 138, 167, 171, 292
Body mostly brown; vertex brown, with a narrow longi-
tudinal median band posteriorly and a blackish brown mark
enclosing ocelli ; rostrum yellowish brown, with a black median
longitudinal stripe on its distal half ; thorax light brown ; meso-
and metanotum with a black median longitudinal streak ; the 1st
to 5th abdominal segments of male black dorsally, last few seg-
ments reddish brown, median process of 3rd tergite short, ex-
tending a little beyond the middle of the 4th tergite ; the 1st to
9th abdominal segments of female uniformly blackish brown.
Fore wing : length, $ , 13-14 mm., $ , 13-14 mm. ; width, $ 2 , 3.4-
3.5 mm. (holotype, length, 13 mm.; width 3.4 mm.) ; membrane
yellow ; markings deep brown ; pterostigmal band complete, with
a broad basal branch and narrow apical branch ; basal band com-
plete, narrow ; apical band large, with a large prominent hyaline
spot and sometimes also with a faint spot; basal spot small,
marginal spot long, narrow, connected with the basal band;
pterostigma not very prominent. Hind wing: length, $ , 11.7-
12.5 mm., $ , 11.4-12.5 mm. ; width, $ , 3-3.5 mm., 5 , 3-3.2 mm. ;
similar to fore wing, except that the basal band is indicated
only by a spot at the hind margin and both basal and marginal
spots are entirely lacking. Male genitalia : genital bulb slender ;
the outer margin slightly concave at the middle, inner margin
with a slightly projecting median portion and a small true basal
lobe ; hypandrium very long ; hypovalvae stout, wide apart at
base, their apical parts overlapping each other, the outer margins
70 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
concave at the middle, the inner margins straight, each with
a proximal lobe, reaching to the middle of the harpagones ; para-
meres very small, Y-shaped, the outer branches simple, con-
nected with the lateral processes of the aedeagus, inner branches
longer, with twisted narrow apices ; preepiproct narrowed to-
wards apex, with a small U-shaped distal incision ; aedeagus with
the two apical processes united together, lateral processes ex-
tending upward, with tooth-like apex. Female genitalia: sub-
genital plate broad, with a wide V-shaped distal incision ; in-
ternal skeleton small, U-shaped, the posterior arms rather sharp
distally, broad basally, with a sclerotized bridge; no axis present.
Holotype (2): Tien-mu-shan, Chekiang, June 6, 1936; in
Museum of Institute of Zoology, Academia Sinica, Shanghai.
Allotype ( 9 ) : Same collecting data and same type location as
holotype. Paratypes : 3 S £ , 3 $ $ , same locality as holotype,
June 12, 1936, in Museum of Institute of Zoology, Academia
Sinica, Shanghai ; 1 $ , 1 $ , same locality as holotype, Aug. 15-
19, 1936, in Museum of Comparative Zoology ; 3 $ $ , 2 $ $ .
same locality, June 6, 1936, 1 6 , 3 $ $ , same locality, July 9-26,
1936, in Cheng Collection, Taipeh.
This species, having deep yellow wing membranes, resembles
the Formosan species, Neopanorpa ophthalmica Navas, in wing
markings, but differs in the shape of the male genitalia. The
length of the hypovalvae of ophthalmica is the same length as
its hypandrium (Esben-Petersen, 1921), whereas that of tien-
mushana is much shorter than its hypandrium. The preepiproct
of ophthalmica is not narrowed toward the apex, and has only
a slightly concave hind margin, whereas that of tienmushana is
narrowed towards the apex, with a small U-shaped distal in-
cision.
Neopanorpa huangshana n. sp.
Figures 135, 139, 166, 170, 291
Body reddish brown ; vertex blackish brown anteriorly, with
a black mark enclosing ocelli ; rostrum reddish brown, with a
deep brown longitudinal stripe on each side ; thorax reddish
brown, meso- and metanotum with black median longitudinal
streak ; the 1st to 5th abdominal segments of male black dorsally,
CHENG: REVISION OF THE CHINESE MECOPTERA 71
6th segment uniformly blackish brown, last few abdominal seg-
ments reddish brown, median process of 3rd tergite short, not
extending beyond the middle of the 4th tergite; the 1st to 9th
abdominal segments of female uniformly blackish brown. Fore
wing: length, $, 12.5 mm., 9, 13.2-1-1 mm.; width, $, 3 mm.;
9 , 3.5 mm. (holotype, length, 12.8 mm.; width, 3.2 mm.) ; mem-
brane yellow, markings deep brown; pterostigmal band com-
plete, with broad basal branch and apical branch ; basal band
complete, broader than that of tienmushana; apical band large,
with small hyaline spot at the hind margin; basal spot small;
marginal spot elongated, connected with the basal band ; ptero-
stigma not very prominent. Hind wing: length, $ , 11 mm., 9 ,
12-13 mm. ; width, i , 3 mm., 9 , 3.3 mm. ; similar to fore wing,
except that the basal band is indicated only by a spot at the
hind margin and both basal and marginal spots are entirely
Lacking. Male genitalia: similar to those of tienmushana, except
that the outer margin of the hypovalvae are not concave, the
inner margin is without the proximal lobe, the apex of the
preepiproct is rather truncated, without the small U-shaped
distal incision, and the two apical processes of the aedeapus are
slightly separated. Female genitalia: similar to those of tien-
mushana, except that the V-shaped distal incision of the sub-
genital plate is wider and the two bases of the posterior arms
are smaller and are separated from each other more than those
of tienmushana.
Holotype ( 6 ) : Huang-shan, Anhwei, June 19, 1936; in the
Museum of Institute of Zoology, Academia Sinica, Shanghai.
Allotype ( 9 ) : same locality as holotype; Aug. 5 ,1936; in same
type location as holotypes. Paratypes: 19, same locality as
holotype, June 21, 1936; in Cheng Collection, Taipeh.
This species resembles Neopanorpa tienmushana n. sp. super-
ficial^, but differs in the shape of the hypovalvae and especially
in the absence of the small U-shaped incision at the distal end
of the preepiproct. This species also differs from Neopanorpa
ophthalmica Xavas by the short hypovalvae of the male ami the
more extensive win?' markings.
72 BULLETIN : MUSEUM OF COMPAKATIVE ZOOLOGY
Neopanorna claripennis Carpenter
Figures 131, 132, 136, 140, 168, 172
Neopanarpa claripennis Carpenter, 1938, Proe. Ent. Soc. Washington,
40(9) :273, figs. 21 24.
Body mostly black ; vertex black ; rostrum uniformly light
brown ; median process of third abdominal tergite of male very
short. Fore wing : length, 13 mm. ; width, 3 mm. ; wing mem-
brane hyaline, slightly smoky, without color markings, although
a few of the specimens have a very slight indication of gray dis-
tally; pterostigma large, dark gray, prominent. Hind wing:
similar to the fore. Male genitalia : genital bulb elongate ; coxo-
podites rather long ; harpagones slender, with a prominent lobe
on the inner margin proximally ; hypandrium conspicuous ;
hypovalvae well-developed, reaching to the base of the harpa-
gones, flattened apically and much broader distally than prox-
imally. Parameres ver}- small, filamentous ; preepiproct with
nearly truncated apex ; aedeagus with short apical processes,
united with each other, lateral processes tooth-shaped, extending
posteriorly. Female genitalia : snbgenital plate broad, with small
V-shaped incision posteriorly ; internal skeleton small, U-shaped,
the axis apparently entirely absent.
Holotype ( $ ) : Beh-luh-din (6000 ft,, 30 miles north of
Chengtu) Szechwan, July 28, 1923 (D. C. Graham) ; in U. S.
National Museum. Allotype ( 9 ) : Chengtu, Szechwan, 1936
(D. C. Graham) ; in U. S. National Museum.
Distribution: Beh-luh-din, Szechwan, July 23-Aug. 28, 1923
and 1933; Chengtu, Szechwan, 1936.
This species, without color markings, resembles Ncopanorpa
apicata Navas. However, according to Carpenter's idea, the tip
of the wing of apicata shows a prominent darkening of the apex
(hence the name apicata), whereas that of claripennis lacks the
darkened apex. Also, in Navas' drawing, the wing of apicata
gives no indication of a prominent pterostigma, whereas that of
claripennis is made very prominent by its color. Unfortunately,
according to Navas, the type of apicata ( S ) lacks the terminal
portion of the abdomen, so we shall never know the genital struc-
ture of the type.
CHENG : REVISION OF THE CHINESE MECOPTERA 73
Neopanorpa taoi Cheng
Figures 157, 158, 159, 160
Neopanorpa taoi Cheng, 1949, Psyche, 56(4): 155, figs. 10, 58, 66, 68.
Body light brown ; vertex entirely black ; rostrum light brown,
with sooty brown stripe on each side ; pronotum sooty brown,
meso- and metanotuin sooty brown on the median portion; the
1st to 5th abdominal segments of male sooty brown dorsally, last
few abdominal segments brown in color, median process of third
abdominal tergite short, with swollen and truncated apex, not
extending beyond the middle of the fourth tergite. Under this
process, there is a small median process and a pair of lateral
processes; the fourth tergite is provided with a concave area on
its anterior portion. Fore wing : length, 17 mm. ; width, 3.8 mm. ;
membrane hyaline with slightly grayish tinge, no markings
present; R2a forked into R2ai an(l R-2a2 ; pterostigma prominent.
Hind wing : length, 15.8 mm. ; width, 3.8 mm. ; similar to the
fore. Male genitalia : genital bulb slender ; coxopodites long, nar-
row distally, bearing many long hairs; harpagones rather short,
the outer margin convex near the base, furnished with a series of
short barbs at the middle, inner margin with a large basal lobe
which bears two tooth-like processes; hypandrium broad, hypo-
valvae wide apart basally, slightly overlapping each other dis-
tally, the basal outer margins greatly convex and strongly
M-lerotized ; parameres simple, leaf-shaped ; preepiproct slender,
with median concave margins, the distal outer portions extended
laterally forming large, distal, tooth-like processes; aedeagus
rather small, both apical processes and lateral processes tooth-
like, extending the same direction and having nearly the same
size.
Female unknown.
Ilolotype ( $ ) : Mt. Lo, Sichang, Sikang, June 10, 1944 (Chia
Chu Tao) ; in Cheng Collection, Taipeh.
This species, without any wing markings, differs from the
other described Neopanorpa by the broadened apex of its median
process of the third abdominal tergite and also by the structure
of its male genitalia, especially the double-toothed basal lobes
of the harpagones and the simple parameres.
i 1 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Xkopanorpa pilosa Carpenter
Figures 192, 193
Neopanorpa pilosa Carpenter, 1945, Psyche, 52(1-2) :75, text-figs. 1, -.
Body light brown, the vertex, thoracic nota and first four
abdominal tergites blackish brown; male with the median process
of the third abdominal tergite well developed, reaching almost to
tbf hind border of the fourth segment. Fore wing: length, 17.5
mm. ; width, 3.8 mm., membrane nearly hyaline, faintly smoky
in appearance ; no markings : pterostigma pale yellow. Male
genitalia: genital bulb slender, coxopodites very long; harpa-
gones rather short, the outer margin concave near the base and
with a large cluster of short black hairs near the middle; each
of the harpagoncs has a prominent lobe on the inner margin near
the base, bearing a number of long black hairs; similar hairs
arise from a short papilla on the distal inner margin of coxo-
podites ; hypandrium conspicuous ; hypovalvae broad and long,
reaching well beyond the base of the harpagones ; each is folded
along the outer margin ; parameres small, branched ; preepiproct
with a pair of thick, distal processes, enlarged distally and
directed inward towards the interior of the bulb; both apical
processes and lateral processes of the aedeagus tooth-like, ex-
tending in the same direction and having nearly the same size.
Female unknown.
llolotype ($)-. Suifu (1000 ft.), Szechwan, Aug.. 1928
(D. C. Graham) ; in U. S. National Museum.
Distribution : same as type.
This species, without any wing markings, resembles Neo-
panorpa taoi Cheng superficially, but differs in the structure of
the male genitalia, especially the parameres. In taoi, the para-
meres are simple, leaf-like, whereas those of pilosa are branched.
This species also bears some resemblance to nigritis Carpenter,
but is much larger and has a lighter body. In nigritis, the outer
margins of the harpagones of the male are normal and smooth,
whereas those of pilosa are distinctly convex at their base.
Neopanorpa nigritis Carpenter
Figures 189, 190, 191, 207, 211, 219, 287
X •opanorpa nigritis Carpenter, 1938, Proe. Ent. Soc. Washington 40(9):
1'74, figs. 17-20.
CHENG : REVISION OF THE CHINESE MECOPTERA 75
Body mostly black ; vertex black ; rostrum light brown, but
with a wide, median black stripe along the anterior surface; the
entire abdomen of male including genital bulb, black, though the
tips of the genital harpagones are reddish brown; median process
of third abdominal tergite well-developed, reaching almost to
the hind border of the fourth segment; the entire abdomen of
female like the male, black. Fore wing: length, 13 mm.; width.
•'3 mm. ; wing membrane nearly hyaline, faintly smoky in appear-
ance ; no markings present ; pterostigma well-developed, reddish
brown. Hind wing : length, 12 mm. ; width, 3 mm. ; similar to the
fore. Male genitalia: genital bulb slender, coxopodites long;
harpagones only of moderate length, with a small lobe proximally
on the inner margin ; hypandrium not very conspicuous ; hypo-
valvae slender, not quite reaching to the base of the harpagones,
nearly membranous distally, their tips bent slightly; parameres
simple, each consisting of a slender stalk, broader distally than
proximally; preepiproct slender, with abruptly broader distal
portion, the apex; slightly emarginate, the distal outer portion
extended laterad to embrace the proctiger as shown in figure 190 ;
aedeagus with short apical processes and tooth-like lateral
processes. Female genitalia : subgenital plate broad, with a
V-shaped distal incision; internal skeleton with the axis pro-
jecting beyond the plate, the two posterior arms being well-
developed and reaching to the tips of the subgenital plate.
Holotype ( $ ) : Mt. Omei (1000 ft.) Szechwan, July 19, 1936
(D. C. Graham) ; in U. S. National Museum. Allotype ( $ ) :
same collecting data and type location as holotype.
Distribution: Mt, Omei" (7000-9000 ft.). Szechwan, July 19,
1936.
This species, without any wing markings, differs from the
other described similar Neopanorpa by its short and slender
hypandrium and hypovalvae. The wings of this species resemble
those of N. claripennis Carpenter, but the body color, the median
stripe on the rostrum, the long median process of the third
abdominal tergite of the male and the large internal skeleton of
the female make its recognition easy.
76 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Neopanorpa validipennis Cheng
Figures 145, 146, 217, 222, 227, 288
Neopanorpa validipennis Cheng, 1949, Psyche, 56(4): 154, figs. 46, 47, 48,
64, 65.
Vertex entirely black ; rostrum deep brown, with a median
longitudinal light brown streak ; thorax sooty brown dorsally,
brown laterally, the 1st to 5th abdominal segments of male dark
brown dorsally, reddish brown ventrally, 6th segment long,
sooty brown, 7th segment reddish brown, 8th segment reddish
brown anteriorly, sooty brown posteriorly; both the 7th and 8th
segments broaden towards apex, the posterior end of the pleural
regions of the 7th segment protruded posteriorly to form two
small processes; median process of the third tergite rather long,
extending nearly to the hind border of the 4th tergite, pointed
at the apex when seen dorsally. Under this median process,
there is another small reddish process, and on both sides of this
median process is a pair of small tooth-like prolongations ; the
median axis of the 4th tergite slightly protruding upward. Fore
wing : length, 14.5 mm. ; width, 3.5 mm. ; membrane slightly gray-
ish brown, no markings present; veins very stout, R^a usually
forked into Roai and Roa2 ; pterostigma not very prominent.
Hind wing: length, 13.5 mm.; width, 3.5 mm.; similar to fore
wing. Male genitalia : genital bulb slender ; coxopodites very
long, abruptly narrow distally, bearing a number of long hairs
on the distal inner margins; harpagones short and slender, the
outer margin slightly concave at the middle, furnished with a
series of short barbs at the basal half, inner margin with a large
lobe basally ; hypandrium long, slightly narrowed towards apex ;
hypovalvae Avith slender basal stalks, wide apart basally, over-
lapping each other, the outer borders extending laterad and being
concave near its middle ; parameres club-shaped with rounded
apex ; preepiproct slender with rounded apex, the distal outer
portion extended laterad to embrace the proctiger, and forming
distal tooth-like processes ; aedeagus very small, the two apical
processes nearly united, lateral processes extended posteriorly,
sharp and tooth-like.
Female unknown.
Holotype ( $ ) : Jihti ( 30 miles east of Tachienlu ) , Sikang,
CHENG : REVISION OF THE CHINESE MECOPTERA 77
Sept. 2, 1939 (F. Y. Cheng, Io Chou and Tein Ho Hei) ; in Cheng
Collection, Taipeh.
Distribution : same as type.
This species, without any wing markings, differs from the
other described Neopanorpa by the rounded basal lobes of the
harpagones and the very long hypandrium and hypovalvae,
which extend far beyond the base of the harpagones. The very
stout veins of this species also make its recognition easy.
Neopanorpa apicata Navas
Neopanorpa apicata Navas, 1927, Rev. Acad. Cienc. Zaragosa, 7:27, fig. 6.
Head deep black ; rostrum yellow ; thorax entirely black ;
abdominal segments mostly black with yellow hairs ; the terminal
portion of the abdomen is lacking in the $ type. However, the
last segment of the remaining abdomen is yellow. Fore wing :
membrane hyaline, iridescent, no marking present, except a dark
shadow at the wing apex; the inner margin of this shadow is
straight, and the shadow disappears gradually posteriorly; be-
tween the veins there are longitudinal indistinct lines formed
by the presence of the black hairs ; veins black ; pterostigma
not prominent. Hind wing : similar to the fore wing. Both $
and 9 genitalia are not known.
Type ( 9 ) : Kweichow (Cavalerie) ; originally in Navas Col-
lection.
Distribution : same as type.
This species, having reduced wing markings, differs from
other described Neopanorpa by the presence of the apical band
and the absence of the other markings. I have not seen this
species. The above account is based upon Navas' original de-
scription.
Neopanorpa choui Cheng
Figures 153, 154, 155, 161, 164, 165
Neopanorpa choui Cheng, 1949, Psyche, 56(4) :151, figs. 22, 23, 43, 44, 45, 62.
Body light brown, the middle part of the thoracic nota sooty
brown ; vertex entirely black ; rostrum yellowish brown ; median
process of the third abdominal tergite of male extraordinarily
long (measuring up to 4.2 mm.), apparently divided into two
7S BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
portions and bearing a series of dense, short stiff hairs on its
ventral surface ; the fourth tergite extremely long, covering
several of the following abdominal segments, somewhat elevated,
and furnished with many short stiff hairs on its surface. Fore
wing : length, 3.5 mm. ; width, 3 mm. ; membrane smoky hya-
line, markings slightly brown, very indistinct; pterostigmal band
incomplete, usually represented only by the faint basal branch
and apical branch ; basal band represented only by two small
spots on the hind margin ; apical band large ; pterostigma brown,
very prominent. Hind wing : length, 12 mm. ; width, 3 mm. ;
similar to fore wing, except that the pterostigmal band and the
basal band are entirely lacking. Male genitalia: genital bulb
slender ; coxopodites long, with truncated apex ; harpagones
slender, the outer margin concave at the middle, inner margin
with a triangular angle and a large basal lobe ; hypandrium
short and broad ; hypovalvae broad and less sclerotized, with an
abruptly narrow apex, extending beyond the base of the harpa-
gones ; parameres modified into a pair of sclerotized rods, greatly
swollen distally and with an incised apex and fused with the
basal part of aedeagus basally ; preepiproct narrow distally,
with truncated and slightly concave apex ; aedeagus rather
small, the two apical processes united together ; lateral processes
extending upward with tooth-like apex. Female genitalia: sub-
genital plate broad basally, narrowed towards apex, with a
narrow U-shaped incision distally ; internal skeleton large, the
plate small, less sclerotized, its posterior arms narrow and slen-
der, sword-shaped, the axis very stout, with abruptly curved
hook-shaped basal ends.
Holotype ( $ ) : Mt. Chowkung, Yaan, Sikang, July 14, 1939
(F. Y. Cheng, lo Chou and Tein Ho Hei) ; in Cheng Collec-
tion, Taipeh. Allotype ( 9 ) : same collecting data and type
location as holotype.
Distribution : same as types.
This species, having very indistinct wing markings, differs
from other described Neopanorpa by its very long median
process of the third abdominal segment and the peculiar struc-
tures of both the male and the female genitalia.
CHENG : REVISION OF THE CHINESE MECOPTERA 79
Neopanorpa brisi (Navas)
Figure 188
Neopanorpa (?) brisi (Navas), Carpenter, 1938, Proc. Ent. Soc. Washing
ton, 40(9) :280.
I.rptnpanorpa brixi Navas, 1930, Notes d'Ent. Chin. Mus. Heucle, 1(6) -A.
fig. 3.
Vertex yellowish brown anteriorly, black posteriorly ; rostrum
slender, brownish yellow with an inverted T-shaped mark at its
base ; thorax black dorsally, yellowish brown ventralby and lat-
erally ; 1st and 2nd abdominal segments black dorsally ; median
process of 3rd tergite with parallel margins and extending to
the hind border of the 4th tergite, bearing a short golden fringe,
4th and 5th segments also black dorsally with a yellowish brown
posterior margin, 6th segment partly cylindrical and partly
conical, brownish yellow dorsally, with indistinct dark lines
running lengthwise, 7th segment narrow, rather short and sub-
cylindrical, narrow basally, obliquely truncated at the distal end ;
the upper lateral corner is bidentate ; seen from above it is
dilated posteriorly and cut off in a bow. Fore wing : length, 17
mm. ; apex rounded, elliptical ; membrane light grayish yellow,
markings brownish yellow; only one long and narrow marking
on the wing, the apex is somewhat darkened ; veins black ; among
the veins and their branches there are dark longitudinal stripes.
Hind wing : length, 15.5 mm. ; similar to the fore, except that
the longitudinal stripes are not so distinct as those of the fore
wing. Male genitalia have not been worked out.
Female unknown.
Type ( $ ) : Yunnan; in Navas Collection.
Distribution: same as type.
This species, having reduced wing markings, differs from other
described Neopanorpa by the dark longitudinal stripes among
their veins. The truncated apex of the 7th abdominal segment
of the male also makes its recognition easy. I have not seen this
species. The above account is based upon Navas' original de-
scription.
80 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Xeopanorpa kwangtsehi n. sp.
Figures 198. 202, 289
Vertex entirely black; rostrum grayish brown, with a pale
white median longitudinal stripe ; thorax yellowish brown, with a
wide sooty brown median longitudinal band dorsally; abdominal
segments of female sooty brown dorsally, yellowish brown later-
ally and ventrally. Fore wing : length, 13.5 mm. ; width, 3.15
mm. ; membrane slightly brown, markings light grayish brown,
indistinct ; pterostigmal band comjjlete, with complete basal
branch and apical branch ; basal band interrupted, not prom-
inent ; apical band large, with three windows ; basal spot absent ;
marginal spot very small ; pterostigma prominent, deep brown.
Hind wing : length, 12.2 mm. ; width, 3 mm. ; no markings pres-
ent, except for a slight suspicion of grayish brown at the apex
of wing and deep brown at the pterostigma. Female genitalia :
subgenital plate broad, with truncated apex ; internal skeleton
large, the plate with concave anterior margin, posterior arms of
the plate U-shaped, axis prominent, but not extending beyond
the plate.
Male unknown.
Holotype ( 9 ) : Chinmen, Kwangtseh, Fukien, Aug. 22, 1945
(Maa) ; in Maa Collection.
This species, having indistinct wing markings, differs from the
other described Neopanorpa by the truncated apex of the sub-
genital plate and the shape of the internal skeleton.
Neopanorpa heii Cheng
Figures 156, 162, 163, 212, 213, 293
Neopanorpa heii Cheng, 194!), Psyche, 56(4) :152, figs. 35, 36, 49, 50, 51.
Vertex entirely black ; rostrum uniformly brown ; thorax sooty
brown dorsally, deep brown laterally ; the 1st to 5th abdominal
segments of male sooty brown dorsally, deep brown laterally
and ventrally, 6th abdominal segment twice the length of 5th
segment, sooty brown in color, last three abdominal segments
also very long, deep brown in color ; median process of the third
tergite short, never extending to the middle of the fourth tergite,
and in contact with the conical projection on the median axis of
CHENG : REVISION OF THE CHINESE MECOPTERA 81
the fourth tergite ; abdominal segments of female sooty brown
dorsally, deep brown laterally and ventrally. Fore wing : length,
S , 12.8 mm. ; 5 , 13.5 mm. ; width, $ , 3.2 mm. ; 9 , 3 mm. ;
membrane slightly brown, markings sooty brown ; pterostigmal
band complete, with a broad basal branch and a greatly re-
duced and separated apical branch ; basal band represented by
a reduced marking on the hind margin ; apical band well de-
veloped ; basal spot very small ; marginal spot consisting of two
reduced spots ; pterostigma prominent. Hind wing : length, $ ,
11.5 mm. ; 9 , 12.2 mm. ; width, $ , 3.2 mm. ; 9 , 3 mm. ; similar
to fore wing, except that apical branch of pterostigmal band,
basal band, basal spot and marginal spot are entirely lacking.
Male genitalia : genital bulb slender ; coxopodites rather long,
with a protruding apex; harpagones very slender, the outer
margin slightly concave at the middle, inner margin with a
smooth angle and a true basal lobe ; hypandrium rather long ;
hypovalvae not flattened, broadend towards the apex, the basal
portion wide apart, the median inner parts greatly prolonged
upward and overlapping each other ; parameres apparently ab-
sent; preepiproct slightly narrowed towards the apex, the distal
portion bent laterad and caudad so as to embrace the proctiger ;
aedeagus very small, both the apical and the lateral processes
tooth-like, the basal part usually covered by a pair of elliptical
membranous plates. Female genitalia ; subgenital plate broad,
with a wide U-shaped distal incision; internal skeleton small, the
plate being band-shaped, transversely elongated, the posterior
arms of the internal skeleton lanceolate, extending laterad and
reaching to the side margins of the subgenital plate ; the axis
small, fork-shaped, the distal portion of the forks joined closely
with the basal posterior arms.
Holotype ( 8 ) : Mt. Chowkung, Yaan, Sikang, July 29, 1939
(F. Y. Cheng, Io Chou and Tein Ho Hei) ; in Cheng Collec-
tion, Taipeh. Allotype ( 9 ) : same collecting data and deposi-
tion as holotype.
Distribution : same as types :
This species superficially resembles Neopanorpa cavaleriei
Xavas in the wing markings, but it can be distinguished by
the smaller wing size, the greatly reduced apical branch of the
pterostigmal band in the fore wing and the entire lack of this
82 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
band in the hind wing. Another difference is the unforked Roa
of this species as compared with the forked R2a in the rede-
scribed figure of cavaleriei by Esben-Petersen (1921, fig. 93).
This species also resembles N. chelata Carpenter, in wing mark-
ings, but these two species are at once distinguished by the forms
op both the male and the female genitalia.
Neopanorpa chelata Carpenter
Figures 169, 173, 186, 194, 195, 216, 299
Xeopanorpa chelata Carpenter, 1938, Proe. Ent. Soe. Washington, 40(9):
274, figs. 13-16.
Body brown and black ; vertex black ; rostrum light brown ;
thorax mostly brown, with a wide median dorsal black stripe;
abdomen black above, the segments beyond the fifth reddish
brown, median process of the third abdominal tergite prominent,
with rounded posterior margin. Fore wing: length, 14 mm.;
width, 3 mm. ; membrane hyaline, markings sooty brown, apical
ban dpresnt, but usually interrupted posteriorly; pterostigmal
band entire, with broad basal branch and very narrow apical
branch; basal band interrupted, represented as three spots;
basal spot absent ; marginal spot present ; pterostigma rather
prominent. Hind wing : length, 13 mm. ; width, 3 mm. ; similar
to fore wing. Male genitalia: genital bulb slender; coxopodites
rather long ; harpagones unusually long and slender, with a
prominent proximal lobe on the inner margin ; hypandrium con-
spicuous; hypovalvae broad, especially distally, each possessing
an outer small apical lobe, reaching to the base of the harpa-
gones; parameres greatly reduced, mostly united with aedeagus;
preepiproct slender, with round distal margin; apical processes
of aedeagus short, united with each other, lateral processes short,
tooth-like. Female genitalia : subgenital plate broad, abruptly
narrowed posteriorly, with a narrow V-shaped distal incision ;
interna] skeleton small, posterior arms U-shaped, axis apparently
absent.
Holotype ( £ ) : Shinkaisi, Mt. Omei, Szechwan, Aug. 16-20.
1934; in U. S. National Museum. Allotype ( $ ) : same collect-
ing data and type location as holotype.
Distribution : same as holotype ; Chengtu, Szechwan, 1936 ;
CHENG : REVISION OP THE CHINESE MECOPTERA 83
foot of Mt. Wa (6000-7000 ft.), Szechwan, July 27, 1925; Kuan-
shien, Szechwan, 1936.
This species resembles Neopanorpa cavaleriei Navas. However,
the median process of the third abdominal tergite of cavaleriei
is narrow and long, almost reaching to the fifth segment, whereas
that of chelata is short and broad. Moreover, the wing size of
cavaleriei is larger than that of chelata.
■ ov
Neopanorpa cavaleriei Navas
Neopanorpa cavaleriei Navas, 1908, Mem. Real. Acad. Cienc. Bare.
1908:417. Esben-Petersen, 1921, Coll. Zool. Selys Long., 5(2) :83, figs.
93, 94. Navas, 1926, Mem. Pont. Aecad. Nuovi Lincei, 9:920. Id., 1930,
Rev. Brot., 24(1):13. Carpenter, 1945, Psyche, 50(1-2) :74, text-figs.
4,7.
Head castaneous ; rostrum reddish brown ; thorax reddish
brown dorsally, sides yellowish brown with some small linear
black spots ; abdomen reddish brown, the hind border of third
tergite narrow and long, almost as long as the fourth segment ;
6th segment cylindrical, a little narrowed towards apex ; 7tb
segment one-fourth shorter than 6th, a little incrassated towards
apex ; 8th as long as 7th, thickened towards apex, which is
obliquely truncated above. Fore wing : length, 16 mm. ; elliptical
at tip ; membrane hyaline, with a faint yellowish tinge, markings
blackish brown ; pterostigmal band complete, with a narrow
basal branch and a narrow apical branch ; basal band indicated
by two small spots ; apical band large, with an oblique prolonga-
tion at the middle of its inner margin, connected along the anterior
margin with the pterostigmal band ; basal spot absent ; marginal
spot very small ; pterostigma not very prominent ; longitudinal
veins and basal crossveins reddish brown, the apical crossveins not
very distinct, Roa forked into Roal and R2a2- Hind wing : length.
14.5 mm. ; similar to the fore, except that the small spot which
represented the marginal spot in the fore wing is entirely absent.
Male genitalia of this species has not been worked out. However,
according to Esben-Petersen, the hypovalvae are rather stout,
the interior margins running close together, but forming a
circular hole at their base ; preepiproct is rounded at tip with
long setae.
Female unknown.
84 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
Type ( $ ) : Kweiyang, Kweichow, 1906 (Cavalerie) ; in Mu-
seum National d'Histoire Naturelle, Paris.
Distribution : same as type ; Tokin, Indo-china.
This species resembles Neopanorpa chelata Carpenter in the
wing markings, but has a larger wing size. The median process
of the third abdominal tergite of this species is narrow and
long, almost reaching to the fifth segment, whereas that of
chelata is short and broad. Unfortunately, the male genitalia
of the type specimen of cavaleriei have not been worked out.
Neopanorpa lacunaris Navas
Xcopanorpa lacunaris Navas, 1930, Notes d'Ent. Mus. Heude, 1(6) :3,
fig. 2.
Head brownish yellow ; vertex black with a deep black spot
within ocelli ; rostrum brownish yellow, with a black stripe along
each side ; thorax brownish yellow, pronotum black, the first
abdominal segment brownish yellow dorsally and ventrally ; 2nd
to 5th segments black dorsally, brownish yellow ventrally; 6th
segment conical, black, with brownish yellow apex ; the latter is
truncated and rounded ; the 8th segment tawny yellow, with the
same shape and same length as in the 7th segment, except that
its posterior border is obliquely truncated ; the 9th segment
globular, rusty brick-color, covered with black hairs ; the median
process of third tergite has parallel sides and extends a little
beyond the tip of the fourth tergite; its apex is covered with
dark hairs. Fore wing : length, 12.5 mm. ; narrow, with rounded,
elliptical apex, membrane hyaline, very light yellow in the basal
third, markings black; pterostigmal band broad, forked pos-
teriorly, both basal and apical branches are broad ; basal band
indicated by two transverse markings, one anterior and the
other posterior; apical band broad, with a small hyaline spot
posteriorly ; basal spot absent ; marginal spot present ; veins
black. Hind wing : length, 11.4 mm. ; similar to the fore, except
that the basal bands are not so distinct. Male genitalia have
not been worked out.
Female unknown.
Type ( S ) : Yunnan ; originally in Navas Collection.
Distribution: same as type.
This species differs from the other described, distinctly
CHENG : REVISION OF THE CHINESE MECOPTERA 85
marked, Neopanorpa bj- the hyaline spot bearing an apical band,
the forked pterostigmal band and the interrupted basal band.
I have not seen the species. The above account is based upon
Xavas' original description.
Neopanorpa translucida n. sp.
Figures 147, 148, 197, 201, 218, 223, 295
Body yellowish brown ; vertex brown with a sooty mark en-
closing ocelli ; rostrum uniformly brown ; thorax yellowish brown
laterally with four black spots, the middle of the meso- and
metanotum deep brown, the anterior portion usually deep brown
in color ; the 1st to 5th abdominal segments of male deep brown
dorsally, 6th segment long, deep brown with somewhat restricted
reddish brown apex, the 7th and 8th segments rather short and
stout, reddish brown in color; median process of the 3rd tergite
rather long, a little extended beyond the hind margin of the
4th tergite; the 1st to 5th abdominal segments of female deep
brown dorsally, last few segments slightly reddish brown. Fore
wing: length, 14.5 mm.; width, 3.7 mm.; membrane slightly
yellowish brown, markings brown; pterostigmal band complete
with broad basal branch and apical branch ; basal band irregu-
lar; apical band very large with two hyaline spots, usually
joined to the pterostigmal band by two very narrow bands ; basal
spot very small; marginal spot represented by an inverted Y-
shaped band ; pterostigma prominent. Hind wing : length, 13
mm. ; width, 3.5 mm. ; similar to fore wing, except that the basal
band and the marginal spot are entirely lacking. Male genitalia :
genital bulb slender ; coxopodites long, with a bundle of 4-5 black
hairs on their distal inner margins and a row of short hairs along
its anterior inner margin; harpagones with broad base and
slender flattened distal portion, the outer margin concave at the
middle, inner margin with a small median angle and a very
large square-shaped basal lobe which is concave ventrally; the
inner margins of this lobe bear a dense row of black hairs, while
its posterior margins bear a row of comb-like brown hairs;
hypandrium rather long; hypovalvae elongated, very broad in
lateral view with rounded apex, extending a little beyond the
base of the harpagones; parameres very short, Y-shaped, the
86 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
inner branches free, the outer branches shorter than the inner
pair, less sclerotized and fused with the basal part of aedeagus;
preepiproct somewhat restricted at its median margins and with
slightly concave distal apex ; aedeagus small, the apical proc-
esses united together, the lateral processes being sharp and
tooth-like, extending outward and upward. Female genitalia :
subgenital plate rather broad with deep V-shaped distal incision ;
internal skeletons rather large, the plate mostly occupied by
the axis with U-shaped posterior arms which are somewhat
twisted, the axis short and stout with sharp anterior end, ex-
tending only a little beyond the plate.
Holotype ( $ ) : Ta-chu-lan, Shaowu Hsien, Fukien, May 18,
1945 (Maa) ; in Museum of National Foochow University, Foo-
chow. Allotype ( 9 ) : Same locality as holotype, Aug. 7, 1915
(Maa); in Maa Collection. Paratypes: 1 $ , Tao-shui, Shaowu
Hsien, Fukien, June 17, 1943 ; 8 $ $ , same locality as holotype ;
May 10-June 10, 1944-1945 (Maa), in Museum of National
Foochow University ; 1 $ , Pen-tien-tung, Changting Hsien, Fu-
kien, April 22, 1941 ; 1 $ , Chien-men, Kwantseh Hsien, Fukien,
Aug. 22, 1945 ; 34 <$ $ , 3 9 9 , same locality as holotype, April
1-Aug. 20, 1942-1945 (Maa), in Maa Collection; 2$ $, 19,
same locality as holotype, May 8-Aug. 19, 1942-1945 (Maa), in
Museum of Comparative Zoology ; 5 $ 5,29 9 , same collecting
data, in Cheng collection, Taipeh.
The wing markings of this species somewhat resemble those of
Neopanorpa caveata n. sp. However, the color of the markings
and the wing membrane are much lighter than those of the
latter. The shape of the harpagones and hypovalvae of the male
and the short axis of the female also enable it to be easily dis-
tinguished as a distinct species.
Neopanorpa pielina Navas
Figures 151, 152, 206, 210, 301
Xeopanorpa pielina Navas, 1936, Notes d 'Ent. Mus. Heude, 3(4):58,
figs. 72, 73.
Vertex reddish brown, with black spot between ocelli ; rostrum
shining reddish brown ; thorax blackish brown dorsally, reddish
brown laterally; the 1st to 5th abdominal segments of male
CHENG : REVISION OP THE CHINESE MECOPTERA 87
blackish brown dorsally, reddish brown ventrally, 6th abdominal
segment black, with narrow reddish brown posterior border, last
three abdominal segments reddish brown ; median process of
the third tergite long and slender, extending to the hind border
of the fourth tergite, the anterior portion of the latter concave
at the middle, with reddish brown color; abdominal segments of
female blackish brown dorsally, reddish brown ventrally. Fore
wing: length, $, 11.5 mm.; 9, 12.5 mm.; width, $, 3 mm.;
9 , 3.2 mm. ; membrane dusky hyaline, markings sooty brown ;
pterostigmal band complete, with broad basal branch and narrow
apical branch; the anterior portion of the pterostigmal band
extends outward and forms a small spot posterior to the ptero-
stigma; basal band represented by two spots; apical band broad,
with a large hyaline spot posteriorly ; basal spot absent ; mar-
ginal spot present ; pterostigma prominent. Hind wing : length,
(5, 10.5 mm.; 9, 11.5 mm.; width, $, 3 mm.; 9, 3.2 mm.;
similar to fore wing, except that the basal band is represented
by only one spot. Male genitalia : genital bulb slender ; coxo-
podites long, with truncated apex and a row of bristles on its
distal inner margins ; harpagones slender, the outer margins
slightly concave at the middle, inner margins with a smooth
median angle and a basal concave area; hypandrium conspicu-
ous; hypovalvae broad, restricted proximally, their outer mar-
gins bent, with broader distal folded portion, extending far
beyond the base of the harpagones; parameres small, Y-shaped,
united with aedeagus and supported by a weakly selerotized
V-shaped bar; preepiproct broad basally with rounded apex,
its distal outer margins bearing a pair of small bent lobes;
aedeagus small, the apical processes united together, lateral
processes lobe-shaped. Female genitalia : subgenital plate broad
at the middle, with a wide V-shaped distal incision ; internal
skeleton large, the outer margins of the plate less selerotized,
the posterior arms of the plate broad at the middle, abruptly
narrowed apically, the axis long, enclosed by the strongly
selerotized extended posterior portion of the plate.
Types (3 9): Killing, Kiangsi, Aug. 13-16. 193.1 (Piel) ; in
lleude Museum, Shanghai.
Distribution : same as types.
88 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
This species, having complete basal and apical bands (with a
hyaline spot), differs from the other described Neopanorpa by
the peculiar structures of both the male and the female genitalia,
i.e., no true lobes in the harpagones of the male and the long
axis of the internal skeleton of the female. The latter usually
occurs with a subgenital plate that has a wide V-shaped distal
incision. The description and drawings of both the male and the
female of this species are based upon the specimens labeled as
paratypes in the Ileude Museum, Shanghai.
Neopanorpa mutabilis n. sp.
Figures 141, 142, 205, 209, 221, 226
Body mostly blackish brown ; vertex deep grayish brown
with sooty brown mark enclosing ocelli ; rostrum uniformly deep
grayish brown; thorax deep grayish brown dorsally, meso- and
metanotum with broad blackish brown streak ; the 1st to 5th
abdominal segments of male blackish brown dorsally, 6th seg-
ment long, blackish brown, 7th and 8th segments shorter than
the 6th, reddish brown in color; median process of the 3rd
tergite short, never extending beyond the hind margin of the
4th tergite ; the anterior median portion of the 4th tergite less
sclerotized, forming a light brown square-shaped area; the 1st
to 6th abdominal segments of female blackish brown dorsally,
last few abdominal segments reddish brown. Fore wing : length,
o , 12.6 mm., 9 , 13.5 mm. ; width, £ , 3.5 mm., 9 , 3.7 mm. ;
membrane hyaline, markings sooty brown; pterostigmal band
complete with broad basal branch and a comparatively narrow
apical branch ; basal band represented by two short bands which
in some individuals connect with each other; apical band large
with a hyaline spot posteriorly; between the apical band and
the pterostigmal band there is usually a narrow additional band
extending from the pterostigmal area to the hind margin of
wing but in some individuals, this band is interrupted, the an-
terior half connected with the apical band to form a large
hyaline window ; basal spot absent ; marginal spot elongated,
band-like; pterostigma not very prominent. Hind wing: length,
3, 11.6 mm., 9, 12.2 mm.; width, $, 3.6 mm., 9, 3.2 mm.;
similar to the fore. Male genitalia: genital bulb slender; coxo-
CHENG : REVISION OF THE CHINESE MECOPTERA 89
podites long with a row of hairs on its distal inner margin;
harpagones slender, the outer margin slightly concave at the
middle, inner margin with a large basal concave area ; hypan-
drium broad ; hypovalvae long, broadened at their median por-
tion, extending far beyond the base of harpagones; parameres
very inconspicuous, weakly sclerotized, consisting of a narrow
stalk, which gives rise to two branches, the inner branch free,
the outer branch united with the lateral process of the aedeagus;
preepiproct narrowed towards its apex with slightly concave
distal margin; aedeagus small, the apical processes united to-
gether, lateral processes tooth-like, extending outward and up-
ward. Female genitalia : subgenital plate broad at the middle
with narrow U-shaped distal incision ; internal skeleton large,
the plate mostly occupied by the long axis ; posterior arms of
the plate swollen at their outer margins.
Holotype ( $ ) : Ta-chu-lan, Shaowu Hsien, Fukien, June 3,
1945 (Maa) ; in Museum of National Foochow University, Foo-
chow. Allotype ( 9 ) : same locality as holotype ; May 25, 1945
(Maa) ; in Museum of National Foochow University, Foochow.
Paratypes : 11 9 9 , same locality as holotype, May 10- June 10.
1945 (Maa), in Museum of National Foochow University, Foo-
chow ; 31 $ $ , 64 9 9 , same locality as holotype, April 23-June
13, 1942-1945; 59 9, Changting Hsien, Fukien, April 30- June
3, 1942-1945 (Maa and Lin), in Maa Collection; 2$ $, 2 9 9.
same collecting data as holotype, in Museum of Comparative
Zoology ; 2 $ £ , 2 9 9 , same collecting data, in Cheng Collection.
Taipeh.
This species differs from previously described Neopanorpa by
its wing markings, especially the presence of the additional band
which extends from the outer part of the pterostigma to the
apical band to form a large hyaline window. The structures of
both the male and the female genitalia are also specific charac-
ters.
Neopanorpa ovata n. sp.
Figures 149, 150, 187, 298
Body deep brown; vertex blackish brown with black mark
enclosing ocelli ; rostrum deep brown ; thorax blackish brown
dorsallv; the 1st to 5th abdominal segments of male blackish
90 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
brown, 6th segment long, slightly blackish brown, last few
abdominal segments deep brown; median process of 3rd tergite
rather long, reaching to the hind border of the 4th tergite, the
median portion of the 4th tergite projecting to form a convex
process which is situated behind the light brown square area of
this tergite. Pore wing: length, 13 mm.; width, 3.4 mm.; mem-
brane hyaline, markings brown ; the wing apex rather rounded ;
pterostigmal band complete, with broad basal branch and nar-
row apical branch ; basal band represented by two large spots ;
apical band complete, with two hyaline spots, the additional
band between the apical band and the pterostigmal band as in
Neopanorpa mutabilis n. sp. ; basal spot absent ; marginal spot
large ; pterostigma prominent. Hind wing : length, 12.2 mm. ;
width, 3.2 mm. ; similar to fore wing, except that the basal band
is represented only by a single spot. Male genitalia : genital
bull) slender; coxopodites long with a row of hairs on their
distal inner margins; harpagones slender, the outer margin
slightly concave at the middle, inner margin slightly convex at
the middle with a well developed basal concave area ; hypan-
drium very broad ; hypovalvae broad and stout, these distal outer
portions usually prolonged to form a broad lobe, which is
folded upward to embrace the hind part of the preepiproct ;
parameres absent ; preepiproct slender, with truncated apex,
rather broad a short distance behind its apex ; aedeagus strongly
sclerotized, with tooth-like apical and lateral processes.
Female unknown.
Ilolotype ( S ) : Pen-tien-tung, Changting Hsien, Fukien.
April 22, 1941 (Maa) ; in Maa Collection, Taipeh.
This species is very similar to Neopanorpa mutabilis n. sp.,
but its rounded wing apex and its very broad hypovalvae enable
it to be easily separated as a distinct species.
Neopanorpa maai n. sp.
Figures 143, 144, 196, 200, 220, 224, 296
Body very weak, yellowish brown ; vertex grayish brown ;
rostrum uniformly light yellowish brown; thorax grayish brown
dorsally, meso- and metanotum with deep grayish brown broad
median longitudinal streak ; the 1st to 5th abdominal segments
of male brown dorsally, 6th segment not so prolonged as in
CHENG : REVISION OF THE CHINESE MECOPTERA 91
Neopanorpa mutdbilis n. sp., brown in color, last few abdominal
segments light reddish brown ; median process of 3rd abdominal
tergite long, extending a little beyond the hind margin of the
4th tergite, the median portion of the 4th tergite less sclerotized,
forming a light brown square area, which is much smaller
than that of mutdbilis; the 1st to 9th abdominal segments of
female uniformly yellowish brown. Fore wing: length, 14 mm.;
width. 3.8 mm. ; membrane hyaline, markings slightly gray ;
pterostigmal band complete with a broad basal branch and a
separate apical branch ; basal band complete, irregular ; apical
hand large with a median hyaline band; between the apical
band and the pterostigmal band there is an additional band, as
in mutabilis, the middle portion of this band being usually con-
nected with the apical branch of the pterostigmal band; basal
snot absent; marginal spot elongated; pterostigma not prom-
inent. Hind wing: length, 12.5 mm.; width, 3.5 mm.; similar
to the fore. Male genitalia: genital bulb slender; coxopodites
long with a row of short hairs on their distal inner margins;
harpagones slender, the outer margin slightly concave at the
middle, inner margin with a rounded angle and a basal concave
area which is not so developed as in mutabilis; hypandrium
broad; hypovalvae shorter than those of mutabilis, extending a
little beyond the base of the harpagones; parameres absent; pre-
epiproct broad at the middle, slender distally with slightly con-
cave distal margin ; aedeagus small, the apical processes united
together, lateral processes tooth-like, short. Female genitalia : sub-
genital plate broad at the middle with deep and narrow V-shaped
distal incision ; the length of axis is the same as that of the
posterior arms of the plate.
Holotype ( $ ) : Ta-shu-lan, Shaowu Hsien, Fukien, April 24,
1944 (Maa) ; in Museum of National Foochow University, Foo-
chow. Allotype ( 9 ) : Li-chia-tun, Kienyang Hsien, Fukien,
April 18, 1945 (Maa) ; in Museum of National Foochow Uni-
versity, Foochow. Paratypes : 2 9 9 , same collecting data as
holotype, in Museum of National Foochow University, Foochow ;
2 cS S ■ , 5 9 9 , same locality as holotype, April 20-May 8, 1942-
1945 (Maa and Lin), in Maa collection; 19, same collecting
data as holotype, in Museum of Comparative Zoology ; 1 $ , 1 9 .
San-chiang, Chungan Hsien, Fulkien, May 8-19, 1943 (Maa).
in Cheng Collection, Taipeh.
92 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
I have named this species in honor of Mr. Maa, who has been
so kind as to allow me the loan of his collection.
This species somewhat resembles Neopanorpa mutabilis n. sp.,
but the gray color of the wing markings, the shorter last few
abdominal segments of the male and the short axis of the
female internal skeleton make its recognition easy.
Neopanorpa banksi Carpenter
Figures 177, 181
Neopanorpa banksi Carpenter, 1938, Proc. Ent. Soc. Washington, 40(9) :
275, figs. 25, 26.
Body mostly black above ; vertex black ; last few abdominal
segments brown, the others black above. Fore wing : length, 15
mm. ; width, 3 mm. ; membrane hyaline ; markings sooty brown ;
pterostigmal band interrupted, with same broad basal branch,
hut no apical branch; basal band reduced to a few spots; apical
band entire, but narrowed posteriorly. Hind wing : similar to the
fore. Female genitalia : subgenital plate broad, with a small
U-shaped distal incision; internal skeleton large, the axis absent,
but with three well developed plates at the base of the long,
posterior arms, which reach to the tip of the subgenital plate.
Male unknown.
Holotype ( $ ) : Suifu, Szechwan (D. C. Graham) ; in U. S.
National Museum.
Distribution : same as holotype.
This species, having distinct wing markings, differs from other
described Neopanorpa by the peculiar shape of the internal
skeleton, which has three plates at the base of the long posterior
arms.
Neopanorpa varia Cheng
Figures 214, 215, 294
Xeopanorpa varia Cheng, 1949, Psj'che, 56(4) :157, figs. 41, 42, 56.
Body light brown, black above, last few abdominal segments
brown; vertex entirely black; rostrum light brown, with black
stripe on each side. Fore wing : length, 14 mm. ; width, 3.2 mm. ;
membrane hyaline, markings sooty brown; pterostigmal band
OHENG : REVISION OF THE CHINESE MECOPTERA 93
complete, with a separated basal branch and a narrow apical
branch ; apical band complete ; pterostigma prominent. Hind
wing: length, 13 mm.; width, 3.3 mm.; similar to fore wing,
except that the basal band is represented by a small marking
on the hind margin. Female genitalia : subgenital plate broad,
with a U-shaped distal incision; internal skeleton large, U-
shaped, posterior arms rather long, obtuse distally, very large
basally, with a narrow sclerotized bridge and a rounded mem-
branous portion between them; axis apparently absent.
Male unknown.
Ilolotype ( 9 ) : Heierhwan (100 miles south of Tachienlu),
Sikang, Sept. 20, 1939 (F. Y. Cheng, Io Chou and Tein Ho Hei) ;
in Cheng Collection, Taipeh.
Distribution: Heierhwan, Sikang, Sept, 20, 1939; Jihti (20
miles east of Tachienlu), Sikang, Sept. 9, 1939; AVantung (50
miles south of Tachienlu), Sikang, Sept. 17, 1939.
This species is somewhat variable with regard to the markings
of the wings. In my collection, there is one individual collected
in AVantung, Sikang, with a greatly reduced pterostigmal band
on both fore and hind wings and without the basal band on
the hind wing.
The wings of this species resemble those of N eopanorpa dimi-
diata Navas superficially. However, in dimidiata the apical
band is well developed, with a faint hyaline spot, whereas that
of varia is interrupted posteriorly and without a hyaline spot.
Moreover, the body color of these two species differs very much.
Neopanorpa cantonensis n. sp.
Figures 175, 179, 302
Body mostly black : vertex blackish brown, with a black mark
enclosing ocelli ; rostrum reddish brown, with blackish brown
longitudinal stripe on each side ; thorax brown laterally, with
some obscurely blackish brown maculations, meso- and metano-
tum brown, with very broad blackish brown median streak; the
1st to 6th abdominal segments of female black dorsally, last few
segments yellowish brown. Fore wing : length, 13 mm., width, 3
ram. ; membrane hyaline, markings blackish brown ; pterostigmal
band very broad, complete, with broad basal branch and apical
!»4 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
branch ; basal band complete ; apical band large, connected with
pterostigmal band at the costal margin ; both basal and marginal
spots present, elongated ; pterostigma not prominent. Hind wing :
length, 12 mm.; width, 2.9 mm.; similar to fore wing, except
that both basal and marginal spots are entirely lacking. Female
genitalia : subgenital plate broad at the middle portion, with
wide V-shaped distal incision ; internal skeleton small, U-shaped,
posterior arms rather sharp distally, their basal outer margins
smoothly curved, connected to each other by a bridge, which is
covered by a rounded large membranous part ; no axis present.
Male unknown.
Holotype (9): Canton, Kwangtung; in Heude Museum,
Shanghai.
This species differs from other described Ncopanorpa by its
very broad wing markings and the presence of the basal spot.
The peculiar structures of the female genitalia also make its
recognition easy.
Neopanorpa dimidiata Navas
Neopanorpa dimidiata Navas, 1930, Notes d 'Ent. Chin. Mus., 1(6) :2, fig. I.
Vertex black, with a rusty yellow line running lengthwise near
the eyes ; rostrum yellowish brown, with a black spot on its
upper surface ; thorax dull yellow, with a deep black median
band dorsally ; 1st to 6th abdominal segments dull yellow, with
a broad median band running lengthwise on the dorsum, the
last few abdominal segments dirty yellow. Fore wing : length,
15.5 mm. ; apex of wing rounded in an elliptical fashion ; mem-
brane appears smudged or very lightly touched with rust ; mark-
ings sooty brown; pterostigmal band broad, with complete apical
branch, which is narrow posteriorly ; basal branch of pterostig-
mal band interrupted, represented by a spot at the anal margin
of the wing ; basal band absent ; apical band broad, complete,
with curved inner border and a faint window posteriorly. Both
basal and marginal spots are absent; veins black and strongly
developed. Hind wing : length, 15 mm. ; similar to the fore.
Female genitalia have not been worked out.
Male unknown.
Type (9): Yunnan; originally in Navas Collection.
Distribution : same as type.
CHENG : REVISION OF THE CHINESE MECOPTERA 95
This species resembles Neopanorpa varia Cheng in the struc-
ture of the pterostigmal band of the wing, but differs in the
apical band. In varia, the apical band is interrupted posteriorly,
without a hyaline spot, whereas that of dimidiata is well de-
veloped, with a faintly hyaline spot posteriorly. I have not seen
this species. The above account is based upon Navas' original
description.
Neopanorpa pulchra Carpenter
Figures 182, 183
.V ropanorpa pulchra Carpenter, 1945, Psyche, 52(1-2) :75, text-fig. 6, pi. 11,
fig. 12.
Body light brown, slightly darker above. Fore wing : length,
14 mm. ; width, 3 mm. ; membrane hyaline, markings grayish
brown; pterostigmal band broad, complete, with broad basal
branch and narrower apical branch ; basal band complete ; apical
band wide and entire, contiguous with pterostigmal band along
costal margin ; basal spot absent ; marginal spot present, small ;
pterostigma not very prominent. Hind wing: length, 12.5 mm.;
width, 3 mm. ; similar to fore wing. Female genitalia : subgenital
plate broad, with a shallow distal notch ; internal skeleton
broader than long, with widely divergent arms and no axis.
Male unknown.
Holotype ( 9 ) : Ta-han, Hainan Island, Kwangtung, June 23.
1935 (L. Gressitt) ; in Museum of Comparative Zoology.
This species has the general wing pattern of Neopanorpa
cantonensis n. sp., but the wing is more slender and the basal
spot is absent. It also resembles N. parva Carpenter super-
ficially, but has more extensive markings. The internal skeleton
of this species differs from that of parva by the widely divergent
arms.
Neopanorpa parva Carpenter
Figures 184, 185
Neopanorpa parva Carpenter, 1845, Psyche, 52(1-2) :73; text figs. 3, 5,
pi. 11, fig. 13.
Body light to dark brown ; vertex, thoracic nota and abdom-
96 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
inal tergites darker. Fore wing : length, 11-13 mm. ; width, 2-2.8
mm. (holotype, length, 11 mm.; width, 2 mm.) ; membrane hya-
line, markings grayish brown ; pterostigmal band wide, with broad
basal branch and narrower apical branch ; basal band inter-
rupted, represented by two narrow bands ; apical band broken
posteriorly ; basal spot absent ; marginal spot appearing as an
elongated narrow band. Hind wing: length, 10 mm.; width, 2
mm. (holotype); similar to the fore. Female genitalia: sub-
genital plate rather broad, with a wide U-shaped distal incision ;
internal skeleton small, with nearly parallel arms and very
short median axis.
Male unknown.
Holotype ( 9 ) : Kwanshien, Szechwan, July 16, 1937 (G. Liu) ;
in Museum of Comparative Zoology.
Distribution s same as holotype.
This species has wing markings resembling those of Neo-
panorpa cavaleriei Navas, but it is much smaller than the latter
and the wing membrane is hyaline, not slightly yellowish as in
cavaleriei. The short median axis of the internal skeleton of the
female makes its recognition easy.
Neopanorpa chaoi n. sp.
Figures 174, 178
Body yellowish brown ; vertex deep brown with a sooty brown
mark enclosing ocelli ; rostrum reddish brown f rontally, yellow-
ish brown laterally ; thorax yellowish brown laterally with few
black spots, deep brown dorsally, meso- and metanotum with a
median longitudinal sooty brown streak; 1st to 6th abdominal
segments of female deep brown dorsally, last few segments red-
dish brown. Fore wing : length, 12.5 mm. ; width, 3.2 mm. ;
membrane hyaline, markings brown ; pterostigmal band com-
plete with broad basal branch and apical branch ; basal band
complete ; apical band prominent with two hyaline spots and
partly united with the pterostigmal band at the pterostigmal
area ; basal spot present, two in number ; marginal spot band-
like, united with the basal band at the median portion of wing;
pterostigma not prominent. Hind wing: length, 11.2 mm.;
width, 3.1 mm. ; similar to fore wing, except that the basal band
CHENG : REVISION OF THE CHINESE MECOPTERA 97
is not so developed and both basal and marginal spots are en-
tirely lacking. Female genitalia : subgenital plate rather broad
with a wide V-shaped distal incision ; internal skeleton small,
V-shaped, posterior arms blade-shaped with rather sharp pos-
terior ends and stout bases which are connected to each other by
a small strongly sclerotized bridge and a large membranous
part ; no axis present.
Male unknown.
Holotype ( 9 ) : Li-chia-tun, Kienyang Hsien, Fukien, Aug. 11,
1915 (Maa) ; in Museum of National Foochow University, Foo-
chow. Paratypes : 3 9 9 , Ta-chu-lan, Shaowu Hsien, Fukien,
May 21-June 9, 1942-1943, 1 9 , Yao-tou, Kienyang Hsien, Fu-
kien, June 11, 1942 (Maa), in Maa collection; 19, Ta-chu-lan,
Shaowu Hsien, Fukien, May 6, 1943 (Maa), in Museum of
Comparative Zoology ; 1 9 , same collecting data, in Cheng
Collection, Taipeh.
I have named this species in honor of Dr. Hsiu Fu Chao, who
has been so kind as to allow me to borrow the material from
the Museum of National Foochow University.
This species, having brown and extensive markings, is easily
recognized by its double hyaline spots of the apical band and
the small V-shaped internal skeleton of the female.
Neopanorpa latipennis Cheng
Figures 199, 203, 297
.V ' .o panorpa latipennis Cheng, 1949, Psyche, 56(4) :156, figs. 39, 40, 55.
Body deep brown, black above, vertex black anteriorly, brown
posteriorly, with a sooty brown marking on the median portion ;
rostrum brown, with a sooty brown median stripe on its lower
portion. Fore wing : length, 14 mm. ; width, 3.53 mm. ; mem-
brane hyaline, markings sooty brown ; pterostigmal band very
broad, with broad basal branch and narrower apical branch;
basal band narrow and uneven, extending to the median portion
of the wing ; apical band large, represented by a big marking
and an inner small Y-shaped band, the latter connected with
the former to form a large hyaline spot ; marginal spots small ;
pterostigma prominent; the wing apex rather broad. Hind wing:
length, 12.55 mm. ; width, 3.5 mm. ; similar to fore wing, except
98 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
that the apical branch of the pterostigmal band, the inner small
Y-shaped band of the apical band and the basal band are
greatly reduced. Female genitalia : subgenital plate abruptly
narrow posteriorly, with a wide U-shaped distal incision; in-
ternal skeleton small, being U-shaped, with a rather long stalk
at its base, the axis apparently absent.
Male unknown.
Holotype ( 9 ) : Moupin, Sikang, July 29, 1941 (Chuan Lung
Lee) ; in Cheng Collection, Taipeh.
This species, having a large hyaline spot in the posterior part
of the apical band of the fore wing, differs from the other de-
scribed Neopanorpa by the shape of the internal skeleton of the
female.
Neopanorpa cabpenteri n. sp.
Figures 176, 180, 300
Xeopanorpa cavalerie\ Navas, Carpenter, 1945, Psyche, 50(1-2) :74, text
figs. 4, 7.
Head chestnut brown, vertex with blackish brown spot be-
tween ocelli ; rostrum chestnut brown with reddish brown tip :
thorax and abdomen blackish brown dorsally, reddish brown
ventrally. Fore wing : length, 15 mm. ; width, 3.7 mm. ; mem-
brane light yellow, markings sooty brown; pterostigmal band
complete, with broad basal branch and narrower apical branch ;
basal band narrow, uneven ; apical band large, with a large
hyaline spot posteriorly and an oblique prolongation in the
middle, not connected along the anterior margin with the
pterostigmal band; basal spot absent; marginal spot appears as
a long and narrow band and is connected to the basal band at
the middle of the wing ; pterostigma prominent ; longitudinal
veins and basal crossveins blackish brown, apical crossveins not
very distinct, Roa simple, not forked. Hind wing : length, 13.5
mm. ; width, 3.8 mm. ; similar to fore wing, except that the apical
band is interrupted posteriorly, the basal band is reduced to
one spot on the hind margin of wing and the marginal spot is
entirely absent. Female genitalia: subgenital plate rather broad,
with shallow V-shaped distal incision; internal skeleton small,
with blade-like and twisted posterior arms, no true axis present.
CHENG: REVISION OP THE CHINESE MECOPTEKA 99
Male unknown.
Ilolotype ( 9 ) : Yim-na-shan, East Kwangtung, June 16, 1936
(L. Gressitt) ; in Museum of Comparative Zoology.
Distribution : same as holotype.
I take the liberty of naming this speeies in honor of Professor
F. M. Carpenter. The species resembles Neopanorpa cavaleriei
in appearance, but differs in the apical band of the fore wing.
In cavaleriei, the apical band is interrupted posteriorly and is
connected along the anterior margin with the pterostigmal band,
whereas that of this species is not interrupted, has a large hya-
line spot and is separated from the pterostigmal band on the
anterior margin of the wing. The R2a vein of cavaleriei is forked
into lUai and K'ja:>, whereas that of this species is simple. More-
over, the basal band of cavaleriei is interrupted, whereas that of
this species is complete and distinct.
Genus LePTOPAXOKPA McLachlan
Leptopanorpa McLachlan, 1875, Trans. Ent. Soc. London, 1875:187. Weele.
1909, Notes Leyden Mus., 31:11. Enderlein, 1910, Zool. Anz., 35:393.
Miyake, 1913, Journ. Coll. Agr. Imp. Univ. Tokyo, 4:381. Esben
Petersen, 1913, Notes Leyden Mus., 35:228. Id., 1921, Coll. Zool. Selys
Long. 5(2) :85. Lieftinck, 1936, Treubia, 15(3) :271.
Himanturella Enderlein, 1910, Zool. Anz., 35:392.
Xeopanorpa Enderlein, 1912, Notes Leyden Mus., 34:237 (nee Weele;.
Rostrum long and slender; tarsal claws serrated on inner
margins; wings are fully developed, slender and narrow, es-
pecially at the base ; 1A short, extending to the anal margin of
wing far before origin of the radial sector ; abdomen very long
and slender in male, much longer than the wings; 6th to 8th
abdominal segments of male much prolongated ; genital bulb of
male with a narrow stalk basally (pedunculate).
Genotype : Leptopanorpa ritsemae McLachlan.
This genus, which inhabits mostly southeast Asia, especially
Japan and Java, is represented in China by only one species.
L. javanica (Westwood) ; the Chinese locality of this species
is Hainan, an island, which is separated from the mainland by
a narrow sea.
The most obvious difference between Leptopanorpa and Neo-
panorpa is the length of the last four abdominal segments of the
100 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
male. However, the genital structures of both the male and the
female of Leptopanorpa are quite close to Neopanorpa. In some
species of Lcptopanorpa, e.g., longicauda, the last four abdom-
inal segments of the male are exceedingly long, in others, e.g.,
erythrura, they are rather short. It seems to me that there is no
distinct difference between Leptopanorpa and Neopanorpa.
Whether or not this characteristic (prolongation of the last
four abdominal segments of the male) is of generic significance
is not certain. A morphological study of the larva of this genus
and that of Neopanorpa would be a great help in settling the
question. A knowledge of the life history and feeding habits of
the Leptopanorpa is also very desirable.
Leptopanorpa javanica (Westwood)
Figures 228, 229
Leptopanorpa javanica (Westwood) Esben-Petersen, 1913, Notes Leyden
Mus. 35:229. Id., 1915, Ent. Medd., 10:231, cat.-no. 9. Id., 1921, Coll.
Zool. Selys Long., 5(2) :89, fig. 100. Lieftinek, 1936. Treubia, 15(3):
315, pis. 6, 7, 10, 12, 14.
Panorpa javanica Westwood, 1846, Trans. Ent. Soc. London, 4:186. Id..
1852, Trans. Ent. Soc. London, 1(2) :5. Walker, 1853, Cat, Nenr. Ins.
Brit. Mus., 1853:460. Weele, 1909, Notes Leyden Mus. 31:6.
Campodotecnum javanicum Enderlein, 1910, Zool. Anz., 35:391. Id., 1912,
Notes Leyden Mus., 34:236.
Head black ; rostrum reddish brown ; thorax black above, sides
grayish testaceous to pitchy black; abdomen of female black
above, terminal segments and the venter paler; 1st to 5th ab-
dominal segments of male black, last few segments dark pitchy
brown ; the hind border of third tergite extending into a slender
cylindrical prolongation which reaches the middle of next seg-
ment, where a tubercle is found ; 6th segment cylindrical, 7th and
8th slender, much thinner than 6th, of equal length and one and
a half times longer than 6th ; their apical part gradually incras-
sate towards the apex, which is obliquely truncated above ; 9th
segment pedunculate. Fore wing : length, 9-10.5 mm. ; slender
with elliptical apex ; membrane whitish, markings sooty black ;
pterostigmal band complete, with a broad basal branch and a
very narrow apical branch ; basal band interrupted, represented
by two spots; apical band broad, with nearly straight inner
CHENG : REVISION OF THE CHINESE MECOPTERA 101
margin, sometimes enclosing a whitish spot in its posterior part ;
along the front margin it is narrowly connected with the ptero-
stigmal band ; basal spot absent ; marginal spot present ; ptero-
stigma not prominent. Hind wing : length, 8-9 mm. ; similar to
the fore, except that the basal band is represented by only one
spot. The male genitalia have been figured out by Lieftinck. Ac-
cording to his drawings, the genital bulb is slender; coxopodites
rather long, with truncated apex ; harpagones long, slender,
smoothly incurved at apices, the outer margin slightly concave
near the base, the inner margin with a basal rounded tooth and
two lobes, opposite each other ; hypandrium conspicuous, broad ;
hypovalvae broad, with rounded apex, reaching beyond the base
of harpagones, the distal portion of hypovalvae coming into
contact with each other; parameres simple, with slender and
twisted stalk and a greatly enlarged apical portion (if I under-
stand Lieftinck 's drawing correctly) ; preepiproct with rounded
tip ; aedeagus with well developed and stout apical processes ;
lateral processes curved backward and inward with broad apex.
The internal skeleton of the female genitalia has been figured by
Lieftinck. According to his drawing, the internal skeleton is
very short and comparatively broad; the basal portion of the
plate is in the form of two wing-like structures, which are rather
twisted and turned dorsad; mesially the two portions converge
and are connected with each other by a thin membrane ; the
distal portion of the plate well demarcated ; posterior arm of the
plate short, with tooth-like apex ; axis not present.
Types ( $ , 9 ) : Java (D. Horsfield) ; in Mus. Soc. Merc. Ind.
Orient.
Distribution : Mt. Wuchi, Hainan, Kwangtung, May 21, 1903 ;
Leito, Burma (Leonardo Pea) ; Carin Chebai, Burma (900-
1100 m.), Dec. 5, 1888 (Leonardo Fea) ; Sumatra (Ericson) ;
Java (Horsfield) ; Banjoemas, Noesa, Mid. Java, no. 10, 1925-
1928 ("teak forest," L. G. E. Kalshoven) ; coastal forest around
Sempoertjondong (Tjidaoen), 100 m., S. W. Java, Sept. 5,
1935 (MaxBartels, Jr.).
This is the only known species in China. It resembles L. peter-
seni Lieftinck (East Java) in body and wing color. However,
according to Lieftinck 's drawings, the male genitalia of this
species are quite different from those of peterseni, especially the
102 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
shape of the hypovalvae. The axis of the female internal skele-
ton of this species is apparently absent, whereas that of peterseni
is well developed.
Family BITTACIDAE
Hittacidae Enderlein, 1910, Zool. Anz., 35:387. Esben-Petersen, 1921, Coll.
Zool. Selys Long. 5(2):115, fig. 126. Carpenter, 1931, Bull. Mus.
Comp. Zool., 72(6): 257.
Ocelli present ; labial palpi two-segmented ; abdomen narrowly
cylindrical; females without ovipositor; terminal segments of
male only slightly modified; legs tenuate, with a single tarsal
claw, modified for grasping ; wings usually subpetiolate, slender ;
costal space narrow, with few crossveins; Rs originating at two-
fifths to one-half the wing length from base ; M dividing near
the middle of the wing.
Bittacus is the only one of the six existing genera of the fam-
ily which inhabits China.
Genus BlTTACUS Latreille
Bittacus Latreille, 1805, Hist. Nat. Crust, et Ins., 8:20. Id., 1807, Gen.
Crust, et Ins., 3:189. Klug, 1836, Abh. Konigl. Akad. Wiss. Berlin.
1836:97. Burmeister, 1839, Handb. Ent., 2:955. Rambur, 1S42, Hist.
Nat, Ins. Nevr., 1812:326. Brauer, 1855, Verh. Zool.-bot. Ges.. 5:707,
pi. 2. Id., 1863, Verb. Zool.-bot. Ges., 21:109, p. 3. Brauer and Low,
1857, Neuropt, Austr., 1857:36. Felt, 1896, New York State Ent.
Rep., 10:463, pis. 3, 4. Hine, 189S, Journ. Colunib. Hortieult. Soc.
12:105, pis. 1, 2. MeClendon, 1906, Ent. News, 1906:121, fig. 15.
Klapalek, 1910, Acta Soc. Ent. Bobem., 7:114. Enderlein, 1910, Zool.
Anz., 35:396. Esben -Petersen, 1913, Revue Zool. Afr., 3:135. Banks,
1913, Trans. Amer. Ent. Soc, 39:233. Lestage, 1917, Revue Zool.
Afr., 5:112. Esben-Petersen, 1921, Coll. Zool. Selys Long., 5(2) :117.
Carpenter, 1931, Bull. Mus. Comp. Zool., 72(6) :257.
Leptobittacim Hine, 1898, Joum. Columb. Hortieult. Soc., 12:108.
Thyridates Navas, 1908, Mem. Real. Acad. Cienc. Art. Barcelona, 1908:412.
Diplostigma Navas, 1908, Mem. Real. Acad. Cienc. Art. Barcelona, 1908:413.
Haplodictyus Navas, 1908, Mem. Real. Acad. Cienc. Art, Barcelona, 1908:
413. Id., 1908, Rev. Russe d'Ent,, 1908:277.
Eyes widely separated below antennae ; basal segment of hind
tarsus longer than fourth segment ; wings present, their mark-
CHENG: REVISION OF THE CHINESE MECOPTEKA 103
ings appearing as spots, without bands ; one costal crossvein ;
1A of hind wing coalescing with Cu2 for a short distance.
Genotype : Bittacus iialicus 0. F. Muller.
This is the second largest genus of Mecoptei'a, including sixty-
two known species in the whole world. Seven species have
already been recorded in China and four new ones are described
below, making a total of eleven. They are widely distributed in
that country. Although only eleven species have been found,
the localities cover the whole mainland of China, that is, from
northern Shensi, Liaoning (one province of Manchuria) to
southern Kwangtung, Yunnan, and from western Sikang to
eastern Kiangsu. Some species, e.g., sinensis Walker, besides
being common in Kiangsu, Chekiang, have also been recorded
in Korea and Japan.
In the classification of the species of Bitiacus, the chief charac-
teristics which have been used are the body structure and wing
eoloration. In some species, the wing membrane is yellow {sinen-
sis, etc.), in others, light brown {sinicus, etc.). The size of the
wing affords some specific characters; in appendiculatus, the fore
wing is shorter than 17 mm., whereas that of sinensis is longer
than 24 mm. The apex of the wing of most species is obtusely
angulated, but in carpenteri n. sp., it forms nearly a right angle,
forming a prominent corner. The wing markings usually appear
as several small spots. They are present in all the species,
except planus and appendiculatus. Most of the venational
eliaracteristics are subject to individual variation. However,
there are some venational features which are of use and value
in the determination of species, especially for the female : the
position of the ending of 1A and that of cubital crossvein (Cuv)
with respect to the fork of media (M) ; the presence or absence
of the anal crossvein ( Av) ; and the number of pterostigmal
erossveins (Pcv).
Another important characteristic is the structure of the male
genitalia, which are not so complex as those of Panorpidae. The
tergum of the 9th abdominal segment is deeply cleft posteriorly,
forming a pair of claspers. The shape of this appendage varies
in different species. Seen from the side, it is triangular in some
species (triangularis, fig. 233) and subquadrangular in others
{sinicus, fig. 245). Some species have a caudal incision in each
104 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
of the claspers (sinensis, fig. 235), whereas others have a single
long process (carpenteri n. sp., fig. 246). In most species, the
posterior parts of the inner surface of the claspers have a patch
of short, stout, sooty-brown bristles (sinensis, etc., fig. 255),
others have no bristles at all (carpenteri n. sp., fig. 256), and
others have two to three bristles which are borne on lobes at-
tached to the median, inner surface of the claspers. The sternum
of the 9th segment is a simple semicircular plate. It is not of
much use for identification. The coxopodites are reduced, mostly
fused with the 9th sternum. The harpagones are also greatly
reduced. However, the shape of this small appendage varies in
different species. In some species, the outer margins of the
harpagones are concave (sinicus, etc., fig. 249) and in others
convex, forming a prominent process (pieli, fig. 250). Some ap-
pear as an inverted boot, others have a long and slender distal
process (planus, fig. 248). Extending upward between the coxo-
podites is a long coiled spiral filament or filum. A pair of
prominent lobes, the aedeagus lobes, project upwards on each
side of the base of the filament. In some species, they are long
and slender (tienmushana n. sp., fig. 268) and in others short
and stout (pieli, fig. 250). Some have a rounded apex (sinensis,
fig. 270), others have the apex truncated (pla?ius, fig. 248). The
proctiger which extends dorso-caudad between the preepiproct
and the coxopodites affords some taxonomic value. In some
species, it is long and slender (carpenteri n. sp., fig. 262) and
in others short and stout (gressitti n. sp., fig. 254). Some have
a pair of lateral lobes at the middle (coreana, fig. 240), while
others have a pair of long processes on the dorsal part of the apex
(tiennmushana n. sp., fig. 263). The lower process of the proctiger
varies much in degree of development. In some species, it is very
long (carpenteri n. sp., fig. 262), in others, it is very short (gres-
sitti n. sp., fig. 254).
The females of Bittacus seem to have lost the internal skeleton
which is so useful in the taxonomy of the Panorpidae. The sub-
genital plate is not well-developed, and no taxonomic value can
be found. Therefore, the identification of females is based only
upon the general body and wing characteristics.
CHENG: REVISION OF THE CHINESE MECOPTERA 105
Key to the Species of Bittacus
1. Wing membrane yellowish 2
Wing membrane light brown or brown 6
2. Fore wing length shorter than 17 mm. or longer than 24 mm 3
3. Wing membrane slightly yellowish ; length of fore wing shorter than
17 mm.; preepiproct extending upward, with pointed dorsal process
and an upwardly curved caudal process appcndiculatus
Wing membrane strongly yellowish ; length of fore wing longer than
24 mm.; preepiproct extending posteriorly, deeply cleft at the apex,
with upper branch and lower branch sinensis
4. Preepiproct cleft at the apex, the upper branch less developed, lower
branch elongated, broadening towards its apex which is curved in-
wards; proctiger with a pair of median lateral lobes coreanus
Preepiproct not cleft at the apex 5
5. Preepiproct triangular when viewed from side; proctiger with a pair
of side lobes located close to its apex triangularis
Preepiproct not triangular when viewed from side, with more or less
truncated posterior margin ; proctiger without paired side lobes ....
planus
6. Prepiproct more or less triangular when viewed from side 7
Preepiproct not triangular when viewed from side 8
7. The posterior part of the ventral portion of preepiproct extends pos
teriorly to form a process; aedeagus lobes long, with rather sharp
tips; dorsal apical half of proctiger with a pair of side lobes
zoensis n. sp.
The posterior part of the ventral portion of preepiproct less extended ;
aedeagus lobes very long, with truncated apex; dorsal apical part of
proctiger with a pair of processes tienmushana n. sp.
8. Preepiproct with prominent posterior process when viewed from side. .9
Preepiproct without posterior process when viewed from side 10
y. Preepiproct irregularly quadrangular when seen from side, with a very
long posterior process at the dorsum of its posterior margin ; both
proctiger and lower process long and slender ; apices of wings appear-
ing as a right angle carpenter i n. sp.
Preepiproct more or less quadrangular when seen from side, with the
caudal margin cleft, the upper branch short, while the lower branch
greatly extends posteriorly to form a process with rounded tip ;
both proctiger and lower process short and stout; apices of wings
not appearing as a right angle gressitti n. sp.
10. Preepiproct subquadrangular, no lobes at its inner side; outer margins
or harpagones not convex siriicu-s
Preepiproct irregular in shape, with two lobes on its median inner side ;
outer margins of harpagones very much convex pieli n. sp.
106 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Descriptions of Species of Bittacus
Bittacus appendiculatus Esben-Petersen
Figure 230
Bittacus appendiculatus Esben-Petersen, 1927, Notul. Ent. 7: 14. figs. '■'•. 4.
Body brown ; vertex with a black spot enclosing ocelli. P'ore
wing : length, 16 mm. ; rather narrow, with smoothly angulated
apex; membrane with yellowish tinge, no markings present;
veins brown, Sc terminating near the middle of the costal margin,
1A very short, ending on the anal margin far before the level
of the fork of M ; cubital crossvein (Cuv) present, located before
the fork of media; no anal crossvein (Av) present; pterostigma
prominent, short, almost triangularly shaped, yellowish brown,
connected with Rs by one pterostigmal crossvein (Pcv). Hind
wing : length. 16 mm. ; similar to the fore, except that the first
anal is a little longer. Male genitalia : I have not seen this
species; however, according to P]sben-Petersen's drawing, the
preepiproct is very broad and short in lateral view, extending
upwards, rather than posteriorly, with a prominent pointed dor-
sal process ; the anterior margin of the preepiproct is strongly
sinuous, the posterior margin straight, with upwardly curved
posterior processes which extend from the lower margin of the
preepiproct; coxopodites prominent, harpagones ax-shaped; both
proctiger and lower process narrowed towards apex.
Type ( S ) : San-nen-kai, Yunnan ; in Esben-Petersen Collec-
tion, Silkeborg.
Distribution ■ same as type.
This species, having light yellowish wing membranes, differs
from the other described Bittacus by the dorsally extended
preepiproct of the male genitalia.
Bittacus sinensis Walker
Figures 235, 237, 255, 270, 303
Bittacus sinensis Walker, 1853, Cat. Neur. Ins. Brit. Mus., 1853:469.
MeLachlan, 18S7, Mitt. Sc-hweiz. Ent. Ges., 1SS7:406. Miyake, 1913,
Journ. Coll. Agr. Imp. Univ. Tokyo, 4:386. Navas, 1913, Notes d'Ent.
Chin. 1(7) :4. Esben-Petersen, 1921, Coll. Selys Long., 5(2):121, fig*.
132, 133. Okamoto, 1925, Bull. Agr. Exp. Sta. Got. Chosen, 2(1) :8.
CHENG: REVISION OF THE CHINESE MECOPTERA 10/
Diplostigma sinense Navas, 1908, Mem. Real Acad. Oiene. Art. Barcelona,
1908:413. Id., Rev. Russe d'Ent.. 1909:277.
Bittacus quatemipunctatus Enderlein, 1910, Zool. Anz., 1910:397. Miyake,
1913, Journ. Coll. Agr. Imp. Univ. Tokyo, 4:387. pi. 33, fig. 6, pi. 37
fig. 10.
fiittncus strategus Navas, 1913. Bull. Mus. d'Hist. Nat., Taris. 1913:442,
figs. 2a. 2h.
Head pale brown, vertex with black spot between ocelli ;
rostrum fuscous with paler tip; thorax and abdomen pale brown.
Pore wing: length, 24-26 mm.; width, 6-6.3 mm.; rather broad
with more or less rounded apex ; membrane strongly yellowish,
with four minute dark brown spots, one at the fork where M
separated from Cu1} one at the base of Rs, one at the subcostal
erossvein (Scv) and one at the first fork of Rs; veins yellowish
brown, 1A terminating at the level of the fork of M, some of
the crossveins in the apical part faintly and narrowly brownish
shaded; both cubital erossvein (Cuv) and anal erossvein (Av)
present, the former a little beyond the fork of media; ptero-
stigma rather prominent, subquadrangular, yellowish, connected
with Rs by one or two pterostigmal crossveins (Pcv). Hind
wing: length, 21-23 mm.; width, 5.5-6 mm.; similar to fore
wings. Male genitalia : the dorsal margin of the preepiproct
when viewed from side, convex, deeply cleft at the tip ; lower
branch of this cleft larger than the upper one, curved inwards,
both lower and upper branches with rounded apex and with a
series of short black bristles on their interior side; caudal end
of coxopoclites concave ; harpagones short, with inner process ;
aedeagus lobes on each side of the base of filum (spiral filament)
narrow and slender; proctiger rather truncated at the apex,
furnished with a bundle of hairs, no lateral lobe present ; lower
process also truncated at the apex.
Type ( 9 ) : Shanghai, China (Saunders 68:3) ; in the type-
series of Walker, Museum of London.
Distribution: same as type: Soochow (Wuhsien), Kiangsu ;
shanghai. Kiangsu ; Chusan, Chekiang, June 3-4, 1934-35; Sui-
gen, Korea, June to July, 1922 ; Shakuofi, Korea, July, 1922 ;
Tokyo, Japan.
This species differs from other described Bittacus by its
strongly yellowish wing membrane. The apex of the preepiproct
of this species is deeply cleft as in B. coreanus Issiki, but the
108 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
upper branch of this species is more developed than that of
coreanus.
BlTTACUS COREANUS Issiki
Figures 234, 240, 247, 251, 309
Bittacus coreanus Issiki, 1929, Trans. Nat. Hist. Soc. Formosa, 19(102):
304, text-fig. 20.
Head brownish yellow, the area between the ocelli blackish ;
rostrum fuscous, with yellowish tip ; thorax and abdomen
yellowish brown or grayish brown, each basal tergite of abdomen
with a very narrow, black, median, transverse band on the hind
border. Fore wing : length, 22 mm. ; width, 5.5 mm. ; broadened
towards the apical area, with obtuse tip ; membrane with yellow-
ish tinge ; markings appear as minute blackish brown spots,
four in number, one at the fork where M separates from Cu,
one at the base of Rs, one at subcostal crossvein (Scv) and one
at the first fork of Rs; besides these, there is a very feeble spot
at the fork of R4 + 5; of all the spots, the first two are more
distinct ; veins brown, some of them yellowish ; 1A ending on
the anal margin a little beyond the fork of M; crossveins in the
apical part of wing faintly and narrowly brownish shaded, cubi-
tal crossvein (Cuv) a little beyond the fork of media, anal
crossvein (Av) between Cu2 and 1A absent; pterostigma not
prominent, connected with Rs by two crossveins (Pcv). Hind
wing : length, 20 mm. ; width, 4.7 mm. ; similar to fore wings.
Male genitalia : the dorsal margin of preepiproct more or less
rounded when viewed from side, deeply cleft at the tip, lower
branch of this cleft elongated, broadened towards apex, curved
inwards, with a series of short black bristles on the interior side
of the tip, upper branch also with short black bristles on the
interior side of its apex ; caudal end of coxopodites with V-
shaped distal incision; harpagones short, with inconspicuous
inner processes ; aedeagus lobes on each side of the base of filum
(spiral filament) short and broad, with rounded tips, reddish
brown; proctiger truncated at the apex, the latter furnished
with a few short hairs on its dorsal corner; at the middle of
proctiger is a pair of lateral lobes, furnished with very short
hairs ; lower process short and narrowed towards apex.
CHENG : REVISION OF THE CHINESE MECOPTERA 109
Type ( $ ) : Keizyo, Korea, June 24, 1926 (Issiki) ; in Issiki
Collection.
Distribution : same as type ; Suigen, Korea, June 23, 1926
(Issiki); Shanghai, Kiangsu, China, June 16-22, 1931-1933;
Zikawei, Shanghai, Kiangsu, China, July 17, 1938 (Piel).
This species has previously been known only from Korea.
Coreanus, having a yellowish wing membrane, differs from
other described Bittacus by the peculiar shape of its preepiproct,
which is deeply cleft at the tip ; the upper branch of this cleft
is short, the lower branch elongate, broadening towards the apex
and curved inwards. The paired median lateral lobes of the
proctiger also make its recognition easy.
Bittacus triangularis Issiki
Figures 233, 241, 252, 267, 310
Bittacus triangularis Issiki, 1929, Trans. Nat. Hist. Soe. Formosa, 19(102) :
30G, text-fig. 21.
Body pale yellowish brown; vertex yellowish brown, with
blackish brown marking between the ocelli and also between
the antennae ; rostrum blackish brown, with yellowish tip ; the
basal segments of the abdomen have very narrow blackish hind
margin above. Fore wing : length, 20-21 mm. ; width, 5.7 mm. ;
the wing apex rather obtuse ; membrane with yellowish tinge ;
markings very small, blackish brown, three in number, one at
the fork where M separates from Cu1} one at the base of R8
and one at the first fork of Rs; veins yellowish brown; 1A
terminating on anal margin near the level of the fork of M,
crossveins in the apical half narrowly and slightly shaded with
brown, subcostal crossvein (Scv) shaded with dark brown, cubi-
tal crossvein (Cuv) considerably beyond the fork of media, anal
crossvein ( Av) absent ; pterostigma not very prominent, con-
nected with Rs by two pterostigmal crossveins (Pcv). Hind
wing : length, 18-19 mm. ; width, 5 mm. ; similar to fore wings.
Male genitalia : preepiproct triangular when viewed from side,
the hind-margin slightly emarginate at the tip, the lower side
of this emargination a little produced, and its inner surface
without black bristles, while the apical part of the upper side
has short black bristles on the inner surface ; distal end of coxo-
1 1 0 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
podites conspicuously produced ; harpagoues comparatively long,
with rounded tips; aedeagus lobes on each side of the base of
filum (spiral filament) grayish yellow, rather long and broad,
with rounded tips; apex of proctiger rounded, with long dense
hairs; very close to the apex there is a pair of side lobes, which
are furnished with soft hairs ; the lower process long, well de-
veloped, with a few inconspicuous hairs on the dorsal surface.
Type ( 8 ) : Moukden (Shenyang), Liaoning, (one province
of Manchuria), July, 1916 (A. Nohira) ; in Issiki Collection.
Distribution: same as type; Keizyo, Korea, June 24, 1926;
Moukden, Liaoning, Aug., 1916.
This species, having a yellowish wing membrane, differs from
the other described Bittacus by its triangular preepiproct in
side view and the peculiar structure of the proctiger.
Bittacus planus Cheng
Figures 239, 244, 248, 258, 305
Bittacus planus Cheng, 1949, Psyche, 56(4) :158; figs. 59, 60, 61, 63, 67.
Body light brown, vertex brown, with a sooty brown marking
enclosing ocelli ; rostrum brown ; mesothorax with two sooty
brown spots on each side dorsally. Fore wing : length, 20.2 mm. ;
width, 5.2 mm. ; the wing apex rather broad, apex obtusely
angulated ; membrane light yellowish brown, without markings;
veins brown, 1A terminating a little before the level of the fork
of M, crossveins very slightly emarginate, cubital crossvein
(Cuv) located beyond the level of the fork of M, no anal cross-
vein (Av) present; pterostigma not very prominent, connected
with Rs by two pterostigmal crossveins (Pcv). Hind wing:
length, 17.5 mm. ; width, 4.2 mm. ; similar to fore wings, except
that there is only one pterostigmal crossvein (Pcv). Male geni-
talia : preepiproct with V-shaped inner margins when seen from
above, with truncated apex ; the apical margins slightly con-
cave, furnished with a series of short black bristles on its inner
sides; posterior end of coxopodites extending upward for a
considerable distance, with smooth apex ; harpagones broad
basally, very narrow and slender distally, with prominent inner
process; aedeagus lobes on each side of the base of filum (spiral
filament) broaden towards apex, furnished with a bundle of
CHENG: REVISION OF THE CHINESE MECOPTERA 111
short hairs; the lower process very long, pointed towards its
apex.
Holotype ( $ ) : Mt. Taipai, Shensi, June, 1942 (Io Chou) ; in
Cheng collection, Taipeh.
This species, having a yellowish brown wing membrane, dif-
fers from the previously described species by the more or less
truncated caudal margins of the preepiproct in lateral view.
The slender harpagones and the broadened apex of the aedeagus
lobes also make its recognition easy.
Bittacus zoensis n. sp.
Figures 231, 253, 257, 265, 306
Body brown, vertex deep brown, with sooty brown marking
enclosing ocelli ; rostrum blackish brown. Fore wing : length,
23.5 mm. ; width, 5.8 mm. ; rather broad, with obtusely angulated
apex; membrane light brown, with six minute blackish brown
spots; one at the fork where M separates from Cux, one at the
base of Rs, one at the first fork of Rs, one at the subcostal cross-
vein (Scv), one at the fork of R4 + 5 and one at the pterostigmal
crossvein (Pcv) ; veins brown; 1A terminating before the level
of the fork of M, crossveins slightly shaded with brown, cubital
crossvein (Cuv) one or two, located in the level of the fork of
M, no anal crossvein (Av) present; pterostigma not prominent,
connected with Rs by one or two pterostigmal crossveins (Pcv).
Hind wing : length, 21 mm. ; width, 5.4 mm. ; similar to fore
wings, except that both cubital and pterostigmal crossveins are
represented by one crossvein. Male genitalia: preepiproct equi-
laterally triangular when seen from side, the caudal margins
slightly convex at the middle, the posterior parts of the ventral
margin extending backward to form a prominent process; at
the inner surface of the angle between the above margins are
short black bristles; coxopodites slightly curved upward with
emarginate apex ; harpagones short, with rounded tips, the
median inner margin with inner processes ; aedeagus lobes on
each side of the base of filum (spiral filament) rather long with
rather sharp tips ; proctiger with truncated apex, furnished with
a bundle of brown hairs; on its apical half there is a pair of
prominent side lobes, furnished with a row of long brown
112 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
bristles ; the lower process rather short, pointed towards its apex.
Holotype ( $ ) : Zo-se, Chungkiang-Hsien, Kiangsi ; June 5,
1934 ; in Heude Museum, Shanghai. Paratypes : 1 9 , same col-
lecting data as holotype; in Museum of Comparative Zoology;
1 $ , same collecting data as holotype ; in Cheng Collection,
Taipeh.
The male of this species has a triangular preepiproct, as seen
in lateral view, as in Bittacus triangularis Issiki, but the more
equilateral form of the preepiproct, the stout harpagones and
the light brown wing membrane make its recognition easy.
Bittacus tienmushana n. sp.
Figures 232, 259, 263, 268, 304
Body blackish brown, vertex brown, with a black marking
within ocelli; rostrum reddish brown. Fore wing: length, 25.5
mm. ; width, 6 mm. ; rather broad, with obtusely angulated
apex; membrane light brown, with three minute blackish brown
spots, one at the fork where M separates from Cu1? one at the
base of Rs and one at the first fork of Rs; veins brown; 1A
terminating at the level of the fork of M, crossveins slightly
shaded with brown, cubital crossvein (Cuv) located in the level
of the fork of M, no anal crossvein (Av) present; pterostigma
not very prominent, connected with Rs by two pterostigmal
crossveins (Pcv). Hind wing: length, 21.5 mm.; width, 5.5 mm.;
similar to fore wings. Male genitalia : preepiproct equilaterally
triangular when seen from side, the caudal parts of the ventral
margins very slightly produced behind; at the inner surface of
the lower area of the caudal margin are short black bristles ; coxo-
podites slightly produced ; harpagones short, inverted boot-
shaped ; aedeagus lobes on each side of the base of filum (spiral
filament) long, with truncated apex and irregular outer margins;
proctiger cone-shaped, with truncated apex; on the dorsal part
of the apex, there is a pair of prominent long processes; the
lower process rather long, pointed towards its apex.
Holotype ( $ ) : Tien-mu-shan, Chekiang ; July 11, 1936 ; in
Museum of Institute of Zoology, Academia Sinica, Shanghai.
Paratypes : 1 $ , same collecting data and same location as holo-
type ; in Museum of Comparative Zoology ; 1 $ , same collecting
data as holotype ; in Cheng Collection, Taipeh.
CHENG : REVISION OF THE CHINESE MECOPTERA 113
This species, having a light brown wing membrane, differs
from Bittacus zoensis n. sp. by its less extended processes in the
caudal-ventral portion of the preepiproct.
Bittacus carpenteri n. sp.
Figures 246, 256, 262, 269, 307
Body light brown; vertex brown, with a deep brown marking
enclosing ocelli; rostrum brown. Fore wing: length, 21.5 mm.;
width, 5.5 mm. ; narrow, with right angulated apex ; membrane
light brown, with several minute dark brown spots, one at the
fork where M separates from Cu1} one at the base of Rs, one
at the first fork of Rs, one at the fork of R4+5 and one at the
cubital crossvein (Cuv) ; veins brown, 1A terminating far
before the level of the fork of M ; the distal end of R5 and most
of the crossveins are heavily shaded with brown; cubital cross-
vein located before the fork of M, no anal crossvein (Av) pres-
ent; pterostigma prominent, connected with Rs by two ptero-
stigmal crossveins (Pcv). Hind wing: length, 17.5 mm.; width,
4.5 mm. ; similar to fore wings. Male genitalia : preepiproct
irregular when seen from side, with a swollen caudal portion
which has a very long caudal process ; coxopodites with concave
apex ; harpagones very small, inwardly bent ; aedeagus lobes on
each side of the base of film (spiral filament) narrowed towards
apex, with truncated tips; proctiger very long, slender at the
middle portion, with enlarged apex which is furnished with a
row of hairs; the lower process very long, pointed towards its
apex.
Holotype ( $ ) : Mt. Omei (11,000 ft.), Szechwan; July 21,
1935; D. C. Graham; in Museum of Comparative Zoology.
I take the liberty to name the species in honor of Professor
F. M. Carpenter, who has allowed me to describe the species.
This species, having a light brown wing membrane, is
easily distinguished from the other described Bittacus by the
apex of the wing forming nearly a right angle and the long
posterior processes of the preepiproct. The very long and slender
proctiger and lower process also make its recognition easy.
114 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
BlTTACUS GRESSITTT 1L sp.
Figures 236, 254, 260, 266, 312
Body deep brown ; vertex blaek anteriorly ; reddish brown
posteriorly ; rostrum blackish brown. Fore wing : length, 23.8
mm. ; width, 5.3 mm. ; membrane light brown, with some ill-
defined reddish brown shadows along the margin of the apical
portion of wing; markings minute, blackish brown, one at the
fork where M separates from Cuj, one at the base of Rs and
one at the first fork of Rs ; veins brown ; 1 A terminating at the
level of the fork of M, crossveins rather heavilv shaded with
brown, cubital crossvein (Cuv) located a little beyond the fork
of M, no anal crossvein (Av) present; pterostigma rather prom-
inent, connected with Rs by two pterostigmal crossveins (Pcv).
Hind wing: length, 21 mm.; width, 5 mm.; similar to fore
wings. Male genitalia : preepiproct U-shaped when seen from
dorsal side, apparently emarginate at the tip, lower parts of this
emargination much produced and not so curved inwards as in
Bittacus chujoi Issiki and Cheng; upper parts short and thick;
both the lower and upper parts are furnished with short black
bristles on the interior sides; harpagones longer than that of
chujoi, with broad bases, rounded tips and small inner processes;
aedeagus lobes on each side of the base of filum (spiral filament)
slender, pointed towards apex in caudal view; proctiger with
its posterior half bent upward ; apex truncated, with only
very minute hairs; close to the apex there is a prominent side
lobe covered with many soft hairs. Lower process short, but
apparently present.
Holotype ( $ ) : Yim-na-shan, E. Kwangtung ; June 12, 1936 ;
L. Gressitt; in Museum of Comparative Zoology. Paratypes:
5 9 9 , same type locality and deposition as holotype ; June 12-17,
1936; L. Gressitt; in Museum of Comparative Zoology.
This species is named in honor of the collector, Dr. L. Gressitt.
It resembles Bittacus corcanus Issiki in the structure of the
preepiproct, but these two species are at once distinguished by
the color of the wing membrane and the body size. This species
is also closely allied to B. chujoi Issiki and Cheng from Formosa.
The preepiprocts of these two species have nearly the same
shape when seen from the side. In gressitti, however, the lower
CHENG: REVISION OF THE CHINESE MECOPTERA 1 1 ~i
margins of the preepiproct are wholly (not just partly, i.e..
caudally, as in chujoi) folded outward and the lower process
is much longer and extends straighter than that of chujoi. Be-
sides this, the bare apex of the proctiger of this species is quite
easy to distinguish from that of chujoi, which has an apical
bundle of long hairs.
*£■>
Bittacus sinicus Issiki
Figures 238, 245, 249, 264, 311
Bittacus sinicus Issiki, 1931, Ann. Ma?. Nat. Hist., (10)7:221, fi«. 2.
Head and rostrum blackish brown, posterior part of vertex
paler; dorsum of thorax blackish brown, meso- and metathorax
with a pale median longitudinal streak, scutella pale; abdomen
blackish brown, becoming blackish towards apex, except the
preepiproct (9th tergite), which is pale brown. Fore wing:
length, 17.5-19.3 mm. ; width, 5 mm. ; rather narrow, dilated
posteriorly, apex obtusely angulated, hind margin conspicuously
sinuate at the end of Cu ; membrane brownish, apical margin
darker ; markings appear as four flecks, one at M, where it sep-
arates from Ciii, one at the base of Rs, one at the first fork of
Rp, and one at ending of Cu2 ; 1A terminating on anal margin
before the level of the form of M, crossveins (except in basal
part of wing) shaded with blackish brown; three of these fall
in a line from fork of Rj + C to near the end of Cu1 ; passing the
fork of M3+!, their shading forms a narrow transverse streak;
cubital crossvein (Cuv) a little before the fork of M, anal cross-
vein (Av) present; pterostigma rather short, not very prom-
inent, connected with R2+3 by two pterostigmal crossveins
(Pcv). Hind wing: length, 15.5-17.5 mm.; width, 4.5 mm.; simi-
lar to fore wing. Male genitalia: preepiproct with deep U-shaped
inner margins, when seen from above, with rounded apex, viewed
laterally, snbquadrangular, upper and lower margin slightly
concave, distal margin slightly convex, oblique and without cleft,
furnished with a series of short black bristles along its inner
sides ; posterior end of coxopodites extending upward for a con-
siderable distance, with emarginate apex ; harpagones broad
basally, narrow distally, with prominent inner process ; aedeagus
lobes on each side of the base of filum (spiral filament) short,
116 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
rather broad, rounded at apex; proctiger narrow, with tooth-
like apex, furnished with a bundle of hairs; the lower process
broad basally, narrowed towards apex.
Type ( $ ) : Mt. Omei (4500 ft.) Szechwan, July 17, 1929
(Collector unknown) ; in Issiki Collection, Tokyo.
Distribution : same as type ; Jihti (30 miles east of Tachienlu),
Sikang, Sept. 2, 1939 (F. Y. Cheng).
This species, having a brownish wing membrane, differs from
other described Bittacus by its subquadrangular preepiproct
(in side view) and its rather small body size.
Bittacus pieli Navas
Figures 242, 243, 250, 261, 308
Bittacus pieli Navas, 1935, Notes d'Ent. Chin. Mus. Heude, 2(5) :99, fig.
63. Id., 1936, Notes, d'Ent. Chin. Mus. Heude, 3(4) :59, fig. 74.
Body dull brown ; vertex with black marking enclosing ocelli ;
rostrum blackish brown. Fore wing : length, 22 mm. ; width,
4.8 mm.; narrow and slender, with obtusely angulated apex;
membrane light brown, markings grayish brown ; of these mark-
ings three are prominent, one at the fork where M separates
from Cuj, one at the base of Rs and one at the first fork of Rs;
the apical portion of the hind margin and the apex of wing
heavily shaded with grayish brown; veins brown; 1A terminat-
ing on anal margin a little before the level of the fork of M, all
the crossveins heavily shaded with grayish brown, cubital cross-
vein (Cuv) located a little before the fork of M, anal crossvein
(Av) present; pterostigma prominent, connected with Es by
two pterostigmal crossveins (Pcv). Hind wing: length, 19.5-20
mm.; width, 4.6 mm.; similar to fore wings. Male genitalia:
preepiproct with irregular dorsal margin and inwardly curved
rounded apex when seen from side; the median inner side of
the preepiproct furnished with two lobes, the dorsal one more
or less elongate, with two or three short black bristles, coxopo-
dites slightly produced upward, with emarginated apex; harpa-
gones short, with rounded tips, the median outer margins slightly
convex ; aedeagus lobes on each side of the base of filum (spiral
filament ) rather short, with gently rounded outer margin and
obtuse tips; proctiger narrowed towards apex, furnished with
CHENG : REVISION OF THE CHINESE MECOPTERA 117
a bundle of brown hairs ; the lower process rather short, broad
basally, pointed towards its apex.
Types ($, 9): Kuling, Kiangsi, Sept. 19, 1934 (Piel) ; in
Heude Museum, Shanghai.
Distribution : same as type.
This species, having a light brown wing membrane, differs
from the other described Bittacus by the peculiar shape of the
preepiproct, which has an irregular dorsal margin and an in-
wardly curved rounded apex when seen from the side. The
slender and heavily marked wing also makes its recognition easy.
BIBLIOGEAPHY
Carpenter, F. M.
1931. Revision of the Nearctic Meeoptera. Bull. Mus. Comp. Zool.,
72:205-277.
1938. Meeoptera from China, with descriptions of new species. Proc.
Ent. Soc. Washington, 40(9) : 267-281.
1940. A new genus of Meeoptera from Tasmania. Pap. and Proc.
Roy. Soc. Tasmania, 1940:51-53.
1945. Panorpidae from China (Meeoptera). Psyche, 52(1-2) : 70-78.
1948. Notes on Chinese Panorpidae (Meeoptera). Psyche, 55:28-30.
Cheng, F. Y.
1949. New species of Meeoptera from Northwest China. Psyche,
56:139-173.
Esben-Petersen, P.
1921. Meeoptera. Collections Zoologiques du Baron Edm. de Selys
Longchamps. Catalogue systematique et descriptif. Bruxelles,
1921:1-172.
1927. New and little-known species of Meeoptera and Neuroptera in
the Zoological Museum of Helsingfors. Notul. Ent., 7:13-18.
1934. Two new species of Panorpa Linn. (Meeoptera). Videnak.
Medd. Dansk Naturh. Foren., 97:211-213.
Issiki, S.
1927. New and rare species of Meeoptera from Corea, Formosa and
Japan. Insecta Matsum., Sapporo, 2:1-12.
1929. Descriptions of new species of the genus Panorpa from Japan
and Formosa. Journ. Soc. Trop. Agric, 1:182-191.
1929. Descriptions of new Meeoptera from the Japanese Empire.
Trans. Nat. Hist. Soc, Formosa, 19(102) :260-314.
1931. Two new species of scorpion flies (order Meeoptera). Ann.
Mag. Nat. Hist., (10)7:219-222.
118 BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY
1933. Morphological studies on the Panorpidae of Japan and adjoin-
ing countries and comparison with American and European
forms. Jap. Journ. Zool., Tokyo, 4:315-416.
Issiki, S. and F. Y. Cheng
1947. Formosan Mecoptera with descriptions of new species. Mem.
Coll. Agric. Nat. Taiwan Univ., 1(4) : 1-17.
Lieftinck, M. A.
1936. Studies in Oriental Mecoptera. I. The genus Leptopanorpa in
Malaysia. Treubia, Buitenzorg, 15:271-320.
McLachan, R.
1894. On two small collections of Neuroptera from Tachienlu, in the
province of Szechwan, western China, on the frontier of Tibet.
Ann. Mag. Nat. Hist., (6)13:421-436.
MlYAKE, T.
1908. A list of Panorpidae of Japan, with descriptions of ten new
species. Bull. Coll. Agric. Tokyo Imp. Univ., 8(1):1-12.
1913. Studies on the Mecoptera of Japan. Journ. Coll. Agric. Univ.
Tokyo, 4:265-400.
Navas, L.
1908. Neuropteros nuevos. Mem. Real. Acad. Cienc. Art, Barcelona,
1908:406-417.
1931. Decadas de insectos nuevos. Rev. Acad. Cienc. Madrid, 26:75.
1934. Nevroptferes et insectes voisins. Notes d'Ent. Chin. Mus. Heude,
2:95-99.
Okamoto, H.
1925. The Mecoptera of Korea. Bull. Agric. Expt. Sta. Gov. Chosen,
Suigen Korea, 2(1) :l-8.
Tjeder, B.
1936. Schwedisch-chinesisehe wissenschaftliche Expedition nach den
nordwestlichen Provinzen Chinas. 51. Mecoptera. Ark. for Zool.,
27(A) -.1-14.
1950. Mecopteren aus Fukien. Bonn Zool. Beitr., 1950(2-4) :286-290.
VVu, C. P.
1937. Catalogus insectorum sinensium. Peiping, 3:1263-1269.
JNDEX OF FAMILIES, GENERA AND SPECIES
Synonyms are printed in italics
angustipennis, 60
apicata, 77
appendieulatus, 106
A ulops, 4
interrupta, 46
a urea, 43
banksi, 92
baohwashana, 38
Bittaeidae, 102
Bittacus, 102
appendieulatus, 106
earpenteri, 113
eoreanus, 10S
gressitti, 114
pieli, 116
planus, 110
quaternipunetatus, J 07
sinensis, lOii
sinicus, 115
strategus, 107
tienmushana, 112
triangularis, 109
zoensis, 111
honis, 58
brisi, 79
( a mpodotecn urn, 60
javanicwm, 10U
cantonensis, 93
earpenteri (Bittaeus), 113
carpenteri ( Xeopanorpa ) , 9s
earpenteri (Panorpa), 55
cavaleriei, 83, 98
caveata, 07
centralis, 20
chaoi, 96
chelate, 82
cheni, 35
choui, 77
cladocerca, 31
claripennis, 72
coomani, 45
eoreanus, 108
cornigera, 58
curva, 43
davidi, 25, 59
diceras, 15, 16, 19
difficilis, 29
dimidiate, 94
Diplostigma, 102
sinensc, 107
tlyscola, 46
emarginata, 22
Estenella, 4
Havicorporis, 33
llavipennis, 24
fructa, 30
f'ukiensis, 41
grahamana, 54
grahami, 15
gressitti, 114
guttata, 59
hageni, 46
Haplodictyus, 102
heii, 80
Rimanturella, 99
huangshana, 70
implicata, 37
interrupta, 46
irregularis, 46
japonica, 46
javanica, 10U
javanicum, 100
kimminsi, 19
klapperiehi, 52
kwangtsehi, 80
lacunaris, 84
latipeunis, 97
leei, 53
Leptobittaeus, 102
Leptopanorpa, 99
brisi, 79
javanica, 100
leucothyria, 46
lutea, 51
maai, 90
macrogaster, 46
mutabilis, 88
Neopanorpa, 60, 99
apicata, 77
banksi, 92
brisi, 79
cantonensis, 93
carpenteri, 98
cavaleriei, 83, 98
caveata, 67
chaoi, 96
chelata, 82
choui, 77
elaripennis, 72
dimidiata, 94
heii, 80
huangshana, 70
kwangtsehi, 80
lacunaris, 84
latipennis, 97
maai, 90
mutabilis, 88
nigritis, 74
ovata, 89
parva, 95
pielina, 86
pilosa, 74
pulchra, 95
taoi, 73
tienmushana, 69
translucida, 85
validipennis, 76
varia, 92
nigritis, 74
niphonensis, 46
obliqua, 36
obtusa, 23
ovata, 89
Panorpa, 4
aurea, 43
baohwashana, 38
bonis, 58
carpenteri, 55
centralis, 20
cheni, 35
cladocerca, 31
coomani, 45
cornigera, 58
curva, 43
davidi, 25, 59
diceras, 15, 16, 19
difficilis, 29
dyscola, 46
emarginata, 22
flavicorporis, 33
flavipennis, 24
fructa, 30
fukiensis, 41
grahami, 15
grahamana, 54
guttata, 59
hageni, 46
implicata, 37
irregularis, 46
japonica, 46
japonica macrogaster, 46
javanica, 100
kimminsi, 19
klapperichi, 52
leei, 53
leucothyria, 46
lutea, 51
macrogaster, 46
niphonensis, 46
obliqua, 36
obtusa, 23
pieli, 57
pulchra, 46
pusilla, 57
r.ectifasciata, 46
semifasciata, 53
sexspinosa, 49
sinanoensis, 46
statura, 56
stigmalis, 27
stotzneri, 18
tetrazonia, 48
tincta, 50
tjederi, 16
trifasciata, 32
typicoides, 40
waongkehzengi, 28
Panorpidae, 2
parva, 95
pieli (Bittacus), 116
pieli (Panorpa), 57
pielina, 86
pilosa, 74
planus, 110
pulchra, 46
pulchra, 95
pusilla, 57
quaternipunctatxis, 107
rectifasciata, 46
semifasciata, 53
sexspinosa, 49
sinanoensis, 46
sinense, 107
sinensis, 106
sinieus, 115
statura, 56
stigmalis, 27
stotzneri, 18
strategus, 107
taoi, 73
tetrazonia, 48
Thyridates, 102
tienmushana (Bittacus), 112
tienmushana (Neopanorpa), 69
tincta, 50
tjederi, 16
translucida, 85
triangularis, 109
trifasciata, 32
typicoides, 40
validipennis, 76
varia, 92
waongkehzengi, 28
zoensis, 111
PLATES
PLATE 1
Fig. 1. Panorpa tjederi Carpenter, preepiproct of $ type after B.
Tjeder in K. J. Morton Collection, Edinburgh.
Fig. 2. Panorpa diceras McLachlan, preepiproct of $ type after F. M.
Carpenter in British Museum (Natural History).
Fig. 3. Panorpa flavipennis Carpenter, preepiproct of $ holotype, in
U. S. National Museum.
Fig. 4. Panorpa kimminsi Carpenter, preepiproct of S paratype in
M. C. Z., Cambridge.
Fig. 5. Panorpa centralis Tjeder, preepiproct of S holotype in Stockholm
Museum.
Fig. 6. Panorpa tjederi Carpenter, genital bulb of $ type after B.
Tjeder in K. J. Morton Collection, Edinburgh.
Fig. 7. Panorpa dic.eras McLachlan, genital bulb of $ type after F. M.
Carpenter in British Museum (Natural History).
Fig. 8. Panorpa stotzneri Esben-Petersen, genital bulb of $ type after
Esben-Petersen in Staatl. Museum fiir Tier- und Volkerkunde, Dresden.
Fig. 9. Panorpa kimminsi Carpenter, genital bulb of $ paratype in
M. C. Z., Cambridge.
Fig. 10. Panorpa centralis Tjeder, genital bulb of $ holotype in Stock-
holm Museum.
Fig. 11. Panorpa flavipennis Carpenter, genital bulb of $ holotype,
in U. S. National Museum.
10
1 1
PLATE 1
PLATE 2
Fig. 12. Panorpa tjederi Carpenter, 9th abdominal segment of 9 typo
(lateral view) after B. Tjeder in K. J. Morton Collection, Edinburgh.
Fig. 13. Panorpa flavipennis Carpenter, 6th abdominal segment (lateral
view) of $ holotype, in U. S. National Museum.
Fig. 14. Panorpa tjederi Carpenter, subgenital plate of 9 type after B.
Tjeder in K. J. Morton Collection, Edinburg.
Fig. 15. Panorpa stotzneri Esben-Petersen, subgenital plate of 9 type
after B. Tjeder in Esben-Petersen Collection, Silkeborg.
Fig. 16. Panorpa Tcimminsi Carpenter, subgenital plate of 9 paratype
in M. C. Z., Cambridge.
Fig. 17. Panorpa tjederi Carpenter, internal skeleton of 9 type after
B. Tjeder in K. J. Morton Collection, Edinburgh.
Fig. 18. Panorpa .stotzneri Esben-Petersen, internal skeleton of 9 type
after B. Tjeder in Esben-Petersen Collection, Silkeborg.
Fig. 19. Panorpa Tcimminsi Carpenter, internal skeleton of 9 paratype in
M. C. Z., Cambridge.
Fig. 20. Panoi-pa centralis Tjeder, internal skeleton of 9 allotype in
Stockholm Museum.
Fig. 21. Panorpa centralis Tjeder, subgenital plate of 9 allotype in
Stockholm Museum.
Fig. 22. Panorpa flavipennis Carpenter, subgenital plate of 9 allotype,
in U. S. National Museum.
Fig. 23. Panorpa flavipennis Carpenter, internal skeleton of 9 allotype,
in U. S. National Museum.
Fig. 24. Panorpa emarginata Cheng, internal skeleton of 9 allotype in
Cheng Collection, Taipeh.
Fig. 25. Panorpa emarginata Cheng, subgenital plate of 9 allotype in
Cheng Collection, Taipeh.
12
13
15
PLATE 3
Fig. 26. Panorpa typicoides Cheng, preepiprcxrt of £ holotype in
M. C. Z., Cambridge.
Fig. 27. Panorpa emarginata Cheng, preepiproct of $ holotype in
M. C. Z., Cambridge.
Fig. 28. Panorpa obtusa Cheng, preepiproct of $ holotype in Cheng
Collection, Taipeh.
Fig. 29. Panorpa fructa Cheng, preepiproct of $ holotype in Cheng
Collection, Taipeh.
Fig. 30. Panorpa stigmalis Navas, preepiproct of $ holotype after Esben-
Petersen in Museum National d'Histoire Naturelle, Paris.
Fig. 31. Panorpa emarginata Cheng, genital bulb of £ holotype in
M. C. Z., Cambridge.
Fig. 32. Panorpa emarginata Cheng, genital bulb of £ holotype, showing
acdeagus, in M. C. Z., Cambridge.
Fig. 33. Panorpa stigmalis Navas, genital bulb of £ holotype after
Esben-Petersen in Museum National d'Histoire Naturelle, Paris.
Fig. 34. Panorpa obtusa Cheng, genital bulb of £ holotype in Cheng
Collection, Taipeh.
Fig. 35. Panorpa fructa Cheng, genital bulb of $ holotype in Cheng
Collection, Taipeh.
Fig. 36. Panorpa typicoides Cheng, genital bulb of $ holotype in
M. C. Z., Cambridge.
Fig. 37. Panorpa obtusa Cheng, genital bulb of £ holotype, showing
aedeagus, in Cheng Collection, Taipeh.
Fig. 38. Panorpa fructa Cheng, genital bulb of 6 holotype, showing
aedeagus, in Cheng Collection, Taipeh.
Fig. 39. Panorpa typicoides Cheng, genital bulb of $ holotype, showing
aedeagus, in M. C. Z., Cambridge.
PLATE
PLATE 4
Fig. 40. Panorpa trifasciata n. sp., preepiproct of $ holotype in
Museum of Foochow University, Foochow.
Fig. 41. Panorpa cladocerca Navas, preepiproct of S paratype in Heude
Museum, Shanghai.
Fig. 42. Panorpa difficilis Carpenter, preepiproct of 6 holotype in IT. S.
National Museum.
Fig. 43. Panorpa waongkchzcngi Navas, preepiproct of $ paratype, in
Heude Museum, Shanghai.
Fig. 4i. Panorpa obliqua Carpenter, preepiproct of o holotype in
M. C. Z., Cambridge.
Fig. 45. Panorpa obliqua Carpenter, genital bulb of 8 holotype in
M. C. Z., Cambridge.
Fig. 46. Panorpa difficilis Carpenter, genital bulb of S holotype in U. S.
National Museum.
Fig. 47. Panorpa waongkehzengi Navas, genital bulb of £ paratype,
showing aedeagus, in Heude Museum, Shanghai.
Fig. 48. Panorpa waonglcehzengi Navas, genital bulb of $ paratype, in
Heude Museum, Shanghai.
Fig. 49. Panorpa trifasciata n. sp., genital bulb of $ holotype in
Museum of National Foochow University, Foochow.
Fig. 50. Panorpa trifasciata n. sp., genital bulb of $ holotype, showing
aedeagus, in Museum of National Foochow University, Foochow.
Fig. 51. Panorpa cladocerca Navas, genital bulb of $ paratype in Heude
Museum, Shanghai.
40
41
42
43
44
45
46
47
48
49
51
PLATE 5
Fig. 52. Panorpa curva Carpenter, genital bulb of $ holotype in U. S.
National Museum.
Fig. 53. Panorpa curva Carpenter, preepiproet of $ holotype in U. S.
National Museum.
Fig. 5-4. Panorpa fuTciensis Tjeder, preepiproet of $ holotype in Museum
A. Koenig, Bonn.
Fig. 55. Panorpa aurea n. sp., preepiproet of S holotype in Maa Collec-
tion, Taipeh.
Fig. 56. Panorpa cheni n. sp., preepiproet of $ holotype in Museum of
Institute of Zoology, Academia Sinica, Shanghai.
Fig. 57. Panorpa davidi Navas, genital bulb of $ holotype after F. M.
Carpenter in Museum National d'Histoire Naturelle, Paris.
Fig. 58. Panorpa fuTciensis Tjeder, genital bulb of $ holotype after B.
Tjeder in Museum A. Koenig, Bonn.
Fig. 59. Panorpa flavicorporis n. sp., genital bulb of $ holotype in
Museum of National Fooehow University, Foochow.
Fig. 60. Panorpa flavicorporis n. sp., genital bulb of $ holotype, show-
ing aedeagus, in Museum of National Foochow University, Foochow.
Fig. 61. Panorpa aurea n. sp., genital bulb of 3 holotype, showing
aedeagus, in Maa Collection, Taipeh.
Fig. 62. Panorpa aurea n. sp., genital bulb of S holotype, in Maa
Collection, Taipeh.
Fig. 63. Panorpa coomani n. sp., genital bulb of $ holotype in Heude
Museum, Shanghai.
52
53
54
55
61
63
PLATE 6
Fig. 64. Panorpa trifasciata n. sp., subgenital plate of 9 allotype in
Maa Collection, Taipeh.
Fig. 65. Panorpa trifasciata n. sp., internal skeleton of 9 allotype in
Maa Collection, Taipeh.
Fig. 66. Panorpa typicoides Cheng, subgenital plate of 9 allotype in
Cheng Collection, Taipeh.
Fig. 67. Panorpa typicoides Cheng, internal skeleton of 9 allotype in
Cheng Collection, Taipeh.
Fig. 68. Panorpa cladocerca Navas, internal skeleton of 9 paratype in
Heude Museum, Shanghai.
Fig. 69. Panorpa flavicorporis n. sp., internal skeleton of 9 allotype in
Maa Collection, Taipeh.
Fig. 70. Panorpa fukiensis Tjeder, subgenital plate of 9 allotype after
B. Tjeder in Museum A. Koenig, Bonn.
Fig. 71. Panorpa fukiensis Tjeder, internal skeleton of 9 allotype after
B. Tjeder in Museum A. Koenig, Bonn.
Fig. 72. Panorpa waongkelizengi Navas, subgenital plate of 9 paratype,
in Heude Museum, Shanghai.
Fig. 73. Panorpa waongkehzengi Navas, internal skeleton of 9 paratype,
in Heude Museum, Shanghai.
Fig. 74. Panorpa cladocerca Navas, subgenital plate of 9 paratype
in Heude Museum, Shanghai.
Fig. 75. Panorpa flavicorporis n. sp., subgenital plate of 9 allotype in
Maa Collection, Taipeh.
Fig. 76. Panorpa aurea n. sp., subgenital plate of 9 allotype in Museum
of National Foochow University, Foochow.
Fig. 77. Panorpa aurea n. sp., internal skeleton of 9 allotype in Museum
of National Foochow University, Foochow.
PLATE 6
PLATE 7
Fig. 78. Panorpa tincta Navas, 6th to 8th abdominal segments of $
holotype after Navas in Hamburg Museum.
Fig. 79. Panorpa coomani n. sp., preepiproct of S holotype in Heude
Museum, Shanghai.
Fig. 80. Panorpa flavieorporis n. sp., preepiproct of $ holotype, in
Museum of National Foochow University, Foochow.
Fig. 81. Panorpa sexspinosa Cheng, preepiproct of <$ holotype in Cheng
Collection, Taipeh.
Fig. 82. Panorpa baohwashana n. sp., genital bulb of $ holotype, showing
aedeagus, in Museum of Institute of Zoology, Academia Siniea, Shanghai.
Fig. 83. Panorpa baohwashana n. sp., genital bulb of $ holotype in
Museum of Institute of Zoology, Academia Siniea, Shanghai.
Fig. 84. Panorpa baohwashana n. sp., preepiproct of $ holotype in
Museum of Institute of Zoology, Academia Siniea, Shanghai.
Fig. 85. Panorpa japonica Thunberg, preepiproct of $ identified speci-
men in Cheng Collection, Taipeh.
Fig. 8G. Panorpa cheni n. sp., genital bulb of £ holotype in Museum of
Institute of Zoology, Academia Siniea, Shanghai.
Fig. 87. Panorpa sexspinosa Cheng, genital bulb of <$ holotype in Cheng
i 'olleetion, Taipeh.
Fig. 8S. Panorpa cheni n. sp., genital bulb of $ holotype, showing
aedeagus, in Museum of Institute of Zoology, Academia Siniea, Shanghai.
Fig. 89. Panorpa sexspinosa Cheng, genital bulb of $ holotype, showing
aedeagus, in Cheng Collection, Taipeh.
Fig. 90. Panorpa japonica Thunberg, genital bulb of 6 identified speci-
men in Cheng Collection, Taipeh.
PLATE
PLATE 8
Fig. 91. Panorpa telrasonia Navas, genital bulb of $ identified speci
men in M. C. Z., Cambridge.
Fig. 92. Panorpa tetrazonia Navas, preepiproct of $ identified specimen
in M. C. Z., Cambridge.
Fig. 93. Panorpa tetrazonia Navas, subgenital plate of 9 identified
specimen in M. C. Z., Cambridge.
Fig. 94. Panorpa tetrazonia Navas, internal skeleton of 9 identified
specimen in M. C. Z., Cambridge.
Fig. 95. Panorpa cheni n. sp., subgenital plate of 9 allotype in Museum
of Institute of Zoology, Academia Sinica, Shanghai.
Fig. 96. Panorpa cheni n. sp., internal skeleton of 9 allotype in Museum
of Institute of Zoology, Academia Sinica, Shanghai.
Fig. 97. Panorpa obliqua Carpenter, subgenital plate of 9 allotype in
M. C. Z., Cambridge.
Fig. 98. Panorpa obliqua Carpenter, internal skeleton of 9 allotype in
M. C. Z., Cambridge.
Fig. 99. Panorpa implicate/, n. sp., subgenital plate of 9 holotype in
Museum of National Foochow University, Foochow.
Fig. 100. Panorpa japonica Thunberg, internal skeleton of 9 identified
specimen in Cheng Collection, Taipeh.
Fig. 101. Panorpa baohwasliana n. sp., subgenital plate of 9 allotype in
Museum of Institute of Zoology, Academia Sinica, Shanghai.
Fig. 102. Panorpa bonis Cheng, subgenital plate of 9 holotype after B.
Tjeder in Stockholm Museum.
Fig. 103. Panorpa implicata n. sp., internal skeleton of 9 holotype in
Museum of National Foochow University, Foochow.
Fig. 104. Panarpa japonica Thunberg, subgenital plate of 9 identified
specimen in Cheng Collection, Taipeh.
Fig. 105. Panorpa baohicashana n. sp., internal skeleton of 9 allotype
in Museum of Institute of Zoology, Academia Sinica, Shanghai.
Fig. 106. Panorpa bonis Cheng, internal skeleton of 9 holotype after B.
Tjeder in Stockholm Museum.
98
102
PLATE 8
PLATE 9
Fig. 107. Panorpa lutca Carpenter, subgenital plate of 9 hole-type
in M. C. Z., Cambridge.
Fig. 108. Panorpa grahamana n. sp., subgenital plate of 9 holotype in
M. C. Z., Cambridge.
Fig. 109. Panorpa statura Cheng, subgenital plate of 9 holotype in
Cheng Collection, Taipeh.
Fig. 110. Panorpa statura Cheng, internal skeleton of 9 holotype in
Cheng Collection, Taipeh.
Fig. 111. Panorpa pieli n. sp., subgenital plate of 9 holotype in Heude
Museum, Shanghai.
Fig. 112. Panorpa lutea Carpenter, internal skeleton of 9 holotype in
M. C. Z., Cambridge.
Fig. 113. Panorpa semifaseiata Cheng, subgenital plate of 9 holotype
in Cheng Collection, Taipeh.
Fig. 114. Panorpa semifaseiata Cheng, internal skeleton of 9 holotype in
Cheng Collection, Taipeh.
Fig. 115. Panorpa grahamana n. sp., internal skeleton of 9 holotype in
M. C. Z., Cambridge.
Fig. 116. Panorpa carpenteri n. sp., internal skeleton of 9 holotype in
M. C. Z., Cambridge.
Fig. 117. Panorpa pieli n. sp., internal skeleton of 9 holotype in Heude
Museum, Shanghai.
Fig. 118. Panorpa pusilla Cheng, internal skeleton of 9 holotype in
M. C. Z., Cambridge.
Fig. 119. Panorpa pusilla Cheng, subgenital plate of 9 holotype in
M. C. Z., Cambridge.
Fig. 120. Panorpa Mapperichi Tjeder, internal skeleton (ventral view)
of 9 holotype after B. Tjeder in Museum A. Koenig, Bonn.
Fig. 121. Panorpa Mapperichi Tjeder, internal skeleton (lateral view)
of 9 holotype after B. Tjeder in Museum A. Koenig, Bonn.
Fig. 122. Panorpa semifaseiata Cheng, ventral view of last few abdominal
segments of 9 holotype in Cheng Collection, Taipeh.
Fig. 123. Panorpa sexspinosa Cheng, subgenital plate of 9 allotype in
M. C. Z., Cambridge.
Fig. 124. Panorpa sexspinosa Cheng, internal skeleton of 9 allotype in
M. C. Z., Cambridge.
Fig. 125. Panorpa leei Cheng, subgenital plate of 9 holotype in
M. C. Z., Cambridge.
Fig. 126. Panorpa Jclapperichi Tjeder, subgenital plate of 9 holotype
after B. Tjeder in Museum A. Koenig, Bonn.
Fig. 127. Panorpa leei Cheng, internal skeleton of 9 holotype in. M. C. Z..
Cambridge.
PLATE 10
Fig. 128. Neopanorpa caveata n. sp., median process of the 3rd abdominal
tergite of $ holotype in Museum of National Foochow University, Foochow.
Fig. 129. Neopanorpa caveata n. sp., preepiproct of $ holotype in
Museum of National Foochow University, Foochow.
Fig. 130. Neopanorpa tienmushana n. sp., preepiproct of $ holotype in
Museum of Institute of Zoology, Academia Sinica, Shanghai.
Fig. 131. Neopanorpa claripennis Carpenter, preepiproct of $ paratype
in M. C. Z., Cambridge.
Fig. 132. Neopanorpa claripennis Carpenter, median process of the 3rd
abdominal tergite of $ paratype in M. C. Z., Cambridge.
Fig. 133. Neopanorpa caveata n. sp., genital bulb of $ holotype in
Museum of National Foochow University, Foochow.
Fig. 134. Neopanorpa tienmushana n. sp., genital bulb of $ holotype in
Museum of Institute of Zoology, Academia Sinica, Shanghai.
Fig. 135. Neopanorpa huangshana n. sp., genital bulb of $ holotype in
Museum of Institute of Zoology, Academia Sinica, Shanghai.
Fig. 136. Neopanorpa claripennis Carpenter, genital bulb of $ paratype
in M. C. Z., Cambridge.
Fig. 137. Neopanorpa caveata n. sp., genital bulb of $ holotype, showing
aedeagus, in Museum of National Foochow University, Foochow.
Fig. 138. Neopanorpa tienmushana n. sp., genital bulb of $ holotype,
showing aedeagus, in Museum of Institute of Zoology, Academia Sinica,
Shanghai.
Fig. 139. Neopanorpa huangshana n. sp., genital bulb of $ holotype,
showing aedeagus, in Museum of Institute of Zoology, Academia Sinica,
Shanghai.
Fig. 140. Neopanorpa claripennis Carpenter, genital bulb of $ paratype,
showing aedeagus, in M. C. Z., Cambridge.
PLATE 11
Fig. 141. Neopanorpa mutabilis n. sp., genital bulb of $ holotype in
Museum of National Foochow University, Foochow.
Fig. 142. Neopanorpa mutabilis n. sp., genital bulb of $ holotype, show
ing aedeagus, in Museum of National Foochow University, Foochow.
Fig. 143. Neopanorpa maai n. sp., genital bulb of £ holotype in
Museum of National Foochow University, Foochow.
Fig. 144. Neopanorpa maai n. sp., genital bulb of $ holotype, showing
aedeagus, in Museum of National Foochow University, Foochow.
Fig. 145. Neopanorpa validipennis Cheng, genital bulb of S holotype
in Cheng Collection, Taipeh.
Fig. 146. Neopanorpa validipennis Cheng, genital bulb of 8 holotype,
showing aedeagus, in Cheng Collection, Taipeh.
Fig. 147. Neopanorpa translucida n. sp., genital bulb of $ holotype,
showing aedeagus, in Museum of National Foochow University, Foochow.
Fig. 148. Neopanorpa translucida n. sp., genital bulb of $ holotype in
Museum of National Foochow University, Foochow.
Fig. 149. Neopanorpa ovata n. sp., genital bulb of $ holotype in Maa
Collection, Taipeh.
Fig. 150. Neopanorpa ovata n. sp., genital bulb of $ holotype, showing
aedeagus, in Maa Collection, Taipeh.
Fig. 151. Neopanorpa pielin-a Navas, genital bulb of S paratype in
Heude Museum, Shanghai.
Fig. 152. Neopanorpa pielina Navas, genital bulb of $ paratype, showing
aedeagus, in Heude Museum, Shanghai.
141
142
143
144
149
150 151
PLATE 11
152
PLATE 12
Fig. 153. Neopanorpa choui Cheng, subgenital plate of $ allotype in
Cheng Collection, Taipeh.
Fig. 154. Neopanorpa choui Cheng, internal skeleton of ? allotype in
Cheng Collection, Taipeh.
Fig. 155. Neopanorpa choui Cheng, median process of 3rd abdominal
tergite (lateral view) of $ holotype in Cheng Collection, Taipeh.
Fig. 156. Neopanorpa heii Cheng, preepiproct of $ holotype in Cheng
Collection, Taipeh.
Fig. 158. Neopanorpa taoi Cheng, genital bulb of o holotype, showing
aedeagus in Cheng Collection, Taipeh.
Fig. 159. Neopanorpa taoi Cheng, preepiproct of 6 holotype in Cheng
Collection, Taipeh.
Fig. 160. Neopanorpa taoi Cheng, median process of 3rd abdominal
tergite (lateral view) of $ holotype in Cheng Collection, Taipeh.
Fig. 161. Neopanorpa choui Cheng, preepiproct of $ holotype in Cheng
Collection, Taipeh.
Fig. 162. Neopanorpa heii Cheng, genital bulb of $ holotype in Cheng
Collection, Taipeh.
Fig. 163. Neopanorpa heii Cheng, genital bulb of $ holotype, showing
aedeagus, in Cheng Collection, Taipeh.
Fig. 164. Neopanorpa choui Cheng, genital bulb of $ holotype in Cheng
Collection, Taipeh.
Fig. 165. Neopanorpa choui Cheng, genital bulb of S holotype, showing
aedeagus, in Cheng Collection, Taipeh.
163 164
PLATE 12
165
PLATE 13
Fig. 166. Neopanorpa huangshana n. sp., subgenital plate of 9 allotype
in Museum of Institute of Zoology, Academia Sinica, Shanghai.
Fig. 167. Neopanorpa tienmushana n. sp., subgenital plate of 9 allotype
in Museum of Institute of Zoology, Academia Sinica, Shanghai.
Fig. 168. Neopanorpa claripennis Carpenter, subgenital plate of 9
paratype in M. C. Z., Cambridge.
Fig. 169. Neopanorpa chelata Carpenter, subgenital plate of 9 paratype
in M. C. Z., Cambridge.
Fig. 170. Neopanorpa huangshana n. sp., internal skeleton of 9 allotype
in Museum of Institute of Zoology, Academia Sinica, Shanghai.
Fig. 171 Neopanorpa tienmushana n. sp., internal skeleton of 9 allotype
in Museum of Institute of Zoology, Academia Sinica, Shanghai.
Fig. 172. Neopanorpa claripennis Carpenter, internal skeleton of 9 para-
type in M. C. Z., Cambridge.
Fig. 173. Neopanorpa chelata Carpenter, internal skeleton of 9 paratype
in M. C. Z., Cambridge.
Fig. 174. Neopanorpa chaoi n. sp., subgenital plate of 9 holotype in
Museum of National Foochow University, Foochow.
Fig. 175. Neopanorpa cantonensis n. sp., subgenital plate of 9 holotype
in Heude Museum, Shanghai.
Fig. 176. Neopanorpa carpenteri n. sp., subgenital plate of 9 holotype
in M. C. Z., Cambridge.
Fig. 177. Neopanorpa banlcsi Carpenter, subgenital plate of 9 holotype
in U. S. National Museum.
Fig. 178. Neopanorpa chaoi n. sp., internal skeleton of 9 holotype in
Museum of National Foochow University, Foochow.
Fig. 179. Neopanorpa cantonensis n. sp., internal skeleton of 9 holotype
in Heude Museum, Shanghai.
Fig. 180. Neopanorpa carpenteri n. sp., internal skeleton of 9 holotype
in M. C. Z., Cambridge.
Fig. 181. Neopanorpa banksi Carpenter, internal skeleton of 9 holotype
in U. S. National Museum.
Fig. 182. Neopanorpa pulchra Carpenter, subgenital plate of 9 holotype
in M. C. Z., Cambridge.
Fig. 183. Neopanorpa pulchra Carpenter, internal skeleton of 9 holotype
in M. C. Z., Cambridge.
Fig. 184. Neopanorpa parva Carpenter, subgenital plate of 9 holotype
in M. C. Z., Cambridge.
Fig. 185. Neopanorpa parva Carpenter, internal skeleton of 9 holotype
in M. C. Z., Cambridge.
170
172
178
179
183 184
PLATE 13
181
185
PLATE 14
Fig. 186. Neopanorpa chelata Carpenter, preepiproct of S paratype
in M. C. Z., Cambridge.
Fig. 187. Neopanorpa ovata n. sp., preepiproct of $ holotype in Maa
Collection, Taipeh.
Fig. 188. Neopanorpa bri-si Navas, last few abdominal segments of $
bolotype by Navas in Navas Collection.
Fig. 189. Neopanorpa 'nigritis Carpenter, preepiproct (dorsal view) of
$ paratype in M. C. Z., Cambridge.
Fig. 190. Neopanorpa nigritis Carpenter, preepiproct (lateral view) of
£ paratype in M. C. Z., Cambridge.
Fig. 191. Neopanorpa nigritis Carpenter, genital bulb of $ paratype
in M. C. Z., Cambridge.
Fig. 192. Neopanorpa pilosa Carpenter, preepiproct of $ holotype in
U. S. National Museum.
Fig. 193. Neopanorpa pilosa Carpenter, genital bulb of $ holotype in
U. S. National Museum.
Fig. 194. Neopanorpa dhelata Carpenter, genital bulb of $ paratype
in M. C. Z., Cambridge.
Fig. 195. Neopanorpa chelata Carpenter, genital bulb of S paratype,
showing aedeagus, in M. C. Z., Cambridge.
193
194
PLATE 14
195
PLATE 15
Fig. 196. Neopanorpa maai n. sp., subgenital plate of 5 allotype in
Museum of National Foochow University, Foochow.
Fig. 197. Neopanorpa translucida n. sp., subgenital plate of 9 allotype
in Museum of National Foochow University, Foochow.
Fig. 198. Neopanorpa Tcwangtschi n. sp., subgenital plate of 9 holotype
in Maa Collection, Taipeh.
Fig. 199. Neopanorpa latipennis Cheng, subgenital plate of 9 holotype
in Cheng Collection, Taipeh.
Fig. 200. Neopanorpa maai n. sp., internal skeleton of 9 allotype in
Museum of National Foochow University, Foochow.
Fig. 201. Neopanorpa translucida n. sp., internal skeleton of 9 allotype
in Museum of National Foochow University, Foochow.
Fig. 202. Neopanorpa Tcwangtsehi n. sp., internal skeleton of 9 holotype
in Maa Collection, Taipeh.
Fig. 203. Neopanorpa latipennis Cheng, internal skeleton of 9 holotype
in Cheng Collection, Taipeh.
Fig 204. Neopanorpa cavcata n. sp., internal skeleton of 9 allotype in
Museum of National Foochow University, Foochow.
Fig. 205. Neopanorpa mutabUis n. sp., internal skeleton of 9 allotype in
Museum of National Foochow University, Foochow.
Fig. 206. Neopanorpa plelina Navas, internal skeleton of 9 paratype
in Heude Museum, Shanghai.
Fig. 207. Neopanorpa 'nigritis Carpenter, internal skeleton of 9 paratype
in M. C. Z., Cambridge.
Fig. 208. Neopanorpa caveata n. sp., subgenital plate of 9 allotype in
Museum of National Foochow University, Foochow.
Fig. 209. Neopanorpa mutabilis n. sp., subgenital plate of 9 allotype in
Museum of National Foochow University, Foochow.
Fig. 210. Neopanorpa pielina Navas, subgenital plate of 9 paratype in
Heude Museum, Shanghai.
Fig. 211. Neopanorpa nigritis Carpenter, subgenital plate of 9 paratype
in M. C. Z., Cambridge.
Fig. 212. Neopanorpa heii Cheng, internal skeleton of 9 allotype in
Cheng Collection, Taipeh.
Fig. 213. Neopanorpa heii Cheng, subgenital plate of 9 allotype in
Cheng Collection, Taipeh.
Fig. 214. Neopanorpa varia Cheng, internal skeleton of 9 holotype
in Cheng Collection, Taipeh.
Fig. 215. Neopanorpa varia Cheng, subgenital plate of 9 holotype in
Cheng Collection, Taipeh.
205
204
202
206
207
209
214
PLATE 15
PLATE 16
Fig. 216. Neo-panorpa ehelata Carpenter, median process of the 3rd
abdominal tergite of $ paratype in M. C. Z., Cambridge.
Fig. 217. Neopanorpa validipennis Cheng, median process of 3rd al>
dominal tergite (dorsal view) of S holotype in Cheng Collection, Taipeh.
Fig. 218. Neopanorpa translucida n. sp., median process of the 3rd ab-
dominal tergite of S holotype in Museum of National Fooehow University,
Fooehow.
Fig. 219. Neopanorpa nigritis Carpenter, median process of 3rd abdominal
tergite of $ paratype in M. C. Z., Cambridge.
Fig. 220. Neopanorpa maai n. sp., median process of 3rd abdominal
tergite of 6 holotype in Museum of National Fooehow University, Fooehow.
Fig. 221. Neopanorpa mutabilis n. sp., median process of 3rd abdominal
tergite of $ holotype in Museum of National Fooehow University, Foo-
ehow.
Fig. 222. Neopanorpa validipennis Cheng, median process of 3rd ab-
dominal tergite (lateral view) of $ holotype in Cheng Collection, Taipeh.
Fig. 223. Neopanorpa translucida n. sp., preepiproet of $ holotype in
Museum of National Fooehow University, Fooehow.
Fig. 224. Neopanorpa maai n. sp., preepiproet of $ holotype in Museum
of National Fooehow University, Fooehow.
Fig. 225. Neopanorpa pielind Navas, preepiproet of S paratype in Heude
Museum, Shanghai.
Fig. 226. Neopanorpa mutabilis n. sp., preepiproet of $ holotype in
Museum of National Fooehow University, Fooehow.
Fig. 227. Neopanorpa validipennis Cheng, preepiproet of 6 holotype in
Cheng Collection, Taipeh.
Fig. 228. Leptopanorpa javanica OVestwood), genital bulb of 6 identi
tied specimen from Noesa Kambangan, Java, after Lieftinck in Esben-
Petersen Collection, Silkeborg.
Fig. 229. Leptopanorpa javanica (Westwood), internal skeleton of 9
identified specimen from Noesa Kambangan, Java, after Lieftinck.
Fig. 230. Bittacus appindiculatus Esben-Petersen, genital segment of $
type after Esben-Petersen in his collection. Silkeborg.
216
217
220
218
221
\
S
219
122
A
223
224
225
226
o
227
228
229
230
PLATE 16
PLATE 17
Fig. 231. Bittacus zoensis n. sp., genital segment (lateral view) of<5
holotype in Heude Museum, Shanghai.
Fig. 232. Bittacus tienmushana n. sp., genital segment (lateral view) of
$ holotype in Museum of Institute of Zoology, Academia Sinica, Shanghai.
Fig. 233. Bittacus triangularis Issiki, genital segment (lateral view) of
3 identified specimen from Keizyo, Korea, in Issiki Collection, Tokyo.
Fig. 234. Bittacus coreanus Issiki, genital segment (lateral view) of 3
identified specimen from Keizyo, Korea, in Issiki Collection, Tokyo.
Fig. 235. Bittacus sinensis Walker, genital segment (lateral view) of 3
identified specimen from Chusan, Chekiang, in Cheng Collection, Taipeh.
Fig. 236. Bittacus gressitti n. sp., genital segment (lateral view) of 3
holotype in M. C. Z., Cambridge.
233
PLATE 18
Fig. 237. Bittacus sinensis Walker, proctiger and lower process of 6
identified specimen from Chusan, Chekiang, in Cheng Collection, Taipeh.
Fig. 238. Bittacus sinicus Issiki, proctiger and lower process of &
identified specimen from Jihti, Sikang, in Cheng Collection, Taipeh.
Fig. 239. Bittacus planus Cheng, proctiger and lower process of &
holotype in Cheng Collection, Taipeh.
Fig. 240. Bittacus coreanus Issiki, proctiger and lower process of &
identified specimen from Keizyo, Korea, in Issiki Collection, Tokyo.
Fig. 241. Bittacus triangularis Issiki, proctiger and lower process of $
identified specimen from Keizyo, Korea, in Issiki Collection, Tokyo.
Fig. 242. Bittacus pieli Navas, proctiger and lower process of $ paratype
in Heude Museum, Shanghai.
Fig. 243. Bittacus pieli Navas, genital segment (lateral view) of 6
paratype in Heude Museum, Shanghai.
Fig. 244. Bittacus planus Cheng, genital segment (lateral view) of 6
holotype in Cheng Collection, Taipeh.
Fig. 245. Bittacus sinicus Issiki, genital segment (lateral view) of $
identified specimen from Jihti, Sikang in Cheng Collection, Taipeh.
Fig. 246. Bittacus carpenteri n. sp., genital segment (lateral view) of
o holotype in M. C. Z., Cambridge.
PLATE 18
PLATE 19
Fig, 247. Bittacus coreanus Issiki, genital segment (caudal view) of &
identified specimen from Keizyo, Korea, in Issiki Collection, Tokyo.
Fig. 248. Bittacus planus Cheng, genital segment (caudal view) of j
hole-type in Cheng Collection, Taipeh.
Fig. 249. Bittacus sinicus Issiki, genital segment (caudal view) of 6
identified specimen from Jihti, Sikang in Cheng Collection, Taipeh.
Fig. 250. Bittacus pieli Navas, genital segment (caudal view) of $
paratype in Ileude Museum, Shanghai.
Fig. 251. Bittacus coreanus Issiki, preepiproct of - identified specimen
from Keizyo, Korea, in Issiki Collection, Tokyo.
Fig. 252. Bittacus triangularis Issiki, preepiproct of 6 identified speci
men from Keizyo, Korea, in Issiki Collection, Tokyo.
Fig. 253. Bittacus soensis n. sp., proctiger and lower process of £ hold
type in Heude Museum, Shanghai.
Fig. 254. Bittacus gressitti n. sp., proctiger and lower process of 6
holotype in M. C. Z., Cambridge.
Fig. 255. Bittacus situ nsis Walker, preepiproct of S identified specimen
from Chusan, Chekiang, in Cheng Collection, Taipeh.
Fig. 256. Bittacus carpenteri n. sp., preepiproct of $ holotype in M. C. Z.,
Cambridge,
Fig. 257. Bittacus soensis n. sp., preepiproct of 6 holotype in Ileude
Museum, Shanghai.
Fig. 258. Bittacus planus Cheng, preepiproct of 6 holotype in Cheng
Collection, Taipeh.
Fig 259. Bittacus tienmushama n. sp., preepiproct of 6 holotype in
.Museum of Institute of Zoology, Academia Sinica, Shanghai.
Fig. 260. Bittacus gressitti n. sp., preepiproct of $ holotype in M. C. Z..
Cambridge.
Fig. 261. Bittacus pieli Xavas, preepiproct of $ paratype in Heude
Museum, Shanghai.
247
248
249
250
PLATE 20
Fig. 262. Bittacus carpenteri n. sp., proctiger and lower process of $
holotype in M. C. Z., Cambridge.
Fig. 263. Bittacus tienmushana n. sp., proctiger and lower process of $
holotype in Museum of Institute of Zoology, Academia Sinica, Shanghai.
Fig. 264. Bittacus sinicus Issiki, preepiproct of $ identified specimen
from Jihti, Sikang, in Cheng Collection, Taipeh.
Fig. 265. Bittacus zoensis n. sp., genital segment (caudal view) of £
holotype in Heude Museum, Shanghai.
Fig. 266. Bittacus gressitti n. sp., genital segment (caudal view) of $
holotype in M. C. Z., Cambridge.
Fig. 267. Bittacus triangularis Issiki, genital segment (caudal view )
of $ identified specimen from Keizyo, Korea, in Issiki Collection, Tokyo.
Fig. 268. Bittacus tienmusluina n. sp., genital segment (caudal view) of
S holotype in Museum of Institute of Zoology, Academia Sinica, Shanghai.
Fig. 269. Bittacus carpenteri n. sp., genital segment (caudal view) of $
holotype in M. C. Z., Cambridge.
Fig. 270. Bittacus sinensis Walker, genital segment (caudal view) of $
identified specimen from Chusan, Chekiang, in Cheng Collection, Taipeh.
263
264
265
266
267
268
269
PLATE 20
270
PLATE 21
Panorpidae, fore wings
Fig. 271. Panorpa Mmminsi Carpenter.
Fig. 272. Panorpa oMusa Cheng.
Fig. 273. Panorpa cmargbnata Cheng.
Fig. 274. Panorpa semifasciata Cheng.
Fig. 275. Panorpa leei Cheng.
Fig. 276. Panorpa typieoides Cheng.
Fig. 277. Panorpa waonglcehzengi Navas.
Fig. 278. Panorpa sexspinosa Cheng.
Fig. 279. Panorpa statitra Cheng.
Fig. 280. Panorpa implicata n. sp.
Fig. 281. Panorpa aurea n. sp.
Fig. 282. Panorpa coomani n. sp.
Fig. 283. Panorpa trifasciata n. sp.
Fig. 284. Panorpa cladocerca Navas.
Fig. 285. Panorpa baohwasJiana n. sp.
Fig. 286. Panorpa japonica Thunheig.
279
272
280
273
282
283
284
277
285
278
286
PLATE 21
PLATE 22
Panorpidae, fore wing!<
Fig. 287. Neopanorpa nigritis Carpenter.
Fig. 288. Neopanorpa validipennis Cheng.
Fig. 289. Neopanorpa Jcwangtsehi n. sp.
Fig. 290. Neopanorpa caveata n. sp.
Fig. 291. Neopanorpa huangshana n. sp.
Fig. 292. Neopanorpa tienmushana n. sp.
Fig. 293. Neopanorpa licii Cheng.
Fig. 294. Neopanorpa varia Cheng.
Fig. 295. Neopanorpa translucida n. sp.
Fig. 290. Neopanorpa rnaai n. sp.
Fig. 297. Neopanorpa latipennis Cheng.
Fig. 298. Neopanorpa ovata n. sp.
Fig. 299. Neopanorpa ohelata Carpenter.
Fig. 300. Neopanorpa carpenteri n. sp.
Fig. 301. Neopanorpa pielvna Navas.
Fig. 302. Neopanorpa cantonensis n. sp.
287
295
288
296
289
290
298
299
292
300
293
PLATE 22
PLATE 23
Bittacidae, fore wings
Pig. 303. Bittacus sim nsis Walker.
Pig. 304. Bittacus tienmusliana n. sp.
Pig. 305. Bittacus planus Cheng.
Pig. 306. Bittacus zoensis n. sp.
Fig. 307. Bittacus carpi nteri n. sp.
Pig. 30S. Bittacus picli Navas.
Fig. 309. Bittacus coreanus Issiki.
Fig. 310. Bittacus triangularis Issiki.
Fig. 311. Bittacus sinicus Issiki.
Pig. 312. Bittacus gressitti n. sp.
303
-7 ?
308
304
309
305
310
306
311
307
312
PLATE 23
Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE
Vol. 116, No. 2
A CATALOGUE OP THE CERIONIDAE
(MOLLUSCA-PULMONATA)
Bv William J. Clench
CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM
April, 19f>7
Publications Issued by or in Connection
with THE
MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE
Bulletin (octavo) 1863 - The current volume is Vol. 115.
Breviora (octavo) 1952 — No. 73 is current.
Memoirs (quarto) 1864-1938 — Publication was terminated with Vol. 55.
Johnsonia (quarto) 1941 -- A publication of the Department of Mollusks.
Vol. 3, no. 35 is current.
Occasional Papers of the Department of Mollusks (octavo) 1945 —
Vol. 2, no. 21 is current.
Proceedings of the New England Zoological Club (octavo) 1899-
1948 — Published in connection with the Museum. Publication terminated
with Vol. 24.
The continuing publications are issued at irregular intervals in numbers
which may be purchased separately. Prices and lists may be obtained on
application to the Director of the Museum of Comparative Zoology,
Cambridge 38, Massachusetts.
Of the Peters "Check List of Birds of the World," volumes 1-3 are out
of print; volumes 4 and 6 may be obtained from the Harvard University
Press; volumes 5 and 7 are sold by the Museum, and future volumes will be
published under Museum auspices.
Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE
Vol. 116, No. 2
A CATALOGUE OF THE CEK10N1DAE
(MOLLUSC A-PULMONATA)
1j5y William J. Clench
CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM
April, 1957
No. 2 — A Catalogue of the Cerionidae (Mollusca-Pidmonata)
By William J. Clench
INTRODUCTION
The Cerionidae, a family of terrestrial pulmonate Gastropoda,
are found on certain islands of the West Indies and the southern
Keys of Florida from Miami south and west to the Dry Tortugas.
It is the only family of land mollusks peculiar to the West
Indies. l
They are halophiles and are seldom found more than a few
hundred feet from the sea. In the Bahamas, however, they may
occur at a much greater distance inland, particularly in lowland
areas where salt spray can reach them from more than one
direction.
This family contains but a single genus with a few subgenera
and a vast number of "species" and "subspecies." Probably
less than 20 per cent of the names now extant actually apply to
valid species or subspecies. The task ahead for anyone attempt-
ing to monograph this group is rather appalling due to the fact
that the characters generally held stable in most other groups
of mollusks are, in this group, wildly rampant. Many recent as
well as early describers, the present author included, are and
were completely oblivious to the remarkable plasticity of this
group of mollusks. I think that, in this genus, nature is isolating
or mixing small elements of Colon populations as effectively as
man has done with his domesticated plants and animals. Cerion
lives mainly along the upper strand line, an exceedingly hazard-
ous area to occupy in any region where hurricanes occur. Here,
for short or long periods of time, they may build up strong,
vigorous colonies. The appalling devastation of a hurricane in
the strand line is quite apparent to even a casual observer. In
such an area, a colony may be greatly reduced or even completely
exterminated. The same storm may move elements of this colony
to a new region and bring in other elements of the genus from
distant places by means of flotsam. This is certainly the way it
1 In essence, the lcnver Florida Keys are mainly West Indian in both
their fauna and flora. I do not know of any permanent colonies of Cerion
living on the Florida mainland.
122 BULLETIN: MUSEUM OK COMPARATIVE ZOOLOGY
appears to those of us fortunate in having had extensive field
experience in the West Indian region. Such a statement is, of
course, difficult to prove, but the facts of distribution still remain
and their haphazard distributional patterns seem to offer no other
reasonable explanation. Like all other land pulmonates, the
larval stages are passed within the egg; there is no "free swim-
ming" stage. Their distribution, beyond their ability to migrate
within a narrow ecological niche, is exceedingly limited as far
as their own mobile power is concerned. Their broader distribu-
tion is brought about entirely by mechanical means. A five-foot
stream would be an absolute barrier without such means of
transport.
The morphological characters of the shell appear to be ex-
ceedingly variable and most of the differences are certainly more
apparent than real. Few, if any, of these characters, such as size,
degree of costation, coloration, position of the apertural teeth,
convexity of the spire, or the ratio between height and width,
are at all stable.
We must take a realistic stand regarding the naming of various
elements in this genus. We are not dealing with a "normal"
group so far as the usual specific characters are concerned, but
rather with a group of mollusks existing under natural conditions
that closely approximate the control and isolation which have
brought about man's domesticated animals and plants.
HISTORICAL SUMMARY
As for most of our widely distributed West Indian molluscan
genera the early work in this group began in the late 18th
century. Surprisingly enough, however, only a very few of the
many named forms that now exist found their way into the
European cabinets prior to 1850. But even at this time, few
names had reached the printed page and these few were the re-
sult mainly of the indefatigable Cuban collector Juan Gundlach.
Much of coastal Cuba was then nearly inaccessible, at least from
the land side, and such named forms were described from local-
ities mainly within easj^ walking distance of the larger coastal
cities. At this time, little was known of the richness of the
Bahama Archipelago. Early monographers, such as Kiister in
the Conchylien Cabinet (1841-50) and Sowerby (1875-76) in
CLENCH: CERIONIDAE 128
the Conehologica Iconica tabulated such species as were then
known without any serious attempt to group them into natural
assemblages. This task was first accomplished by Pilsbry in the
Manual of Conchology, 1901-02. Prior to the work of Pilsbry,
Maynard (1889) started such a study but the several new forms
that he described over the course of many years, from 1889 to
1924, completely submerged his original attempt at such a com-
plete classification. His work was marred by many inaccuracies
of all kinds and his attempts toward a clarification of this com-
plex problem dwindled as the jrears passed, ending in brief de-
scriptions and eventually in a sales catalogue with a few "new
species" described. In fairness to Maynard, however, the com-
mercial side of his venture was not to gain profit for himself
but to realize money to finance additional expeditions in quest
of these mollusks in which he was so deeply interested. It seems
to me that Maynard failed to grasp much of the importance of
his own discoveries. He failed to see that he was actually dealing
with unit populations and not with completely isolated entities
which he had termed "species." Somewhere in his writings he
mentioned that a wagon road on New Providence was a complete
barrier between two of his named ' ' species. ' ' But he overlooked
the caprice of a single hurricane and the consequent mixing of
these two populations. Nevertheless, we owe much to Maynard
for his early exploration of both the Bahama Islands and the
Cayman Islands, as the specimens he collected are still the
only materials available for study from many remote and inac-
cessible localities.
It is most unfortunate that his writings were privately pub-
lished and had a very limited sale. He not only wrote the text,
but cut his own wood blocks, and with a small printing press
set his own type and printed his publications. His collection
was purchased from his daughter jointly by the Museum of
Comparative Zoology and the United States National Museum
in 1931.
After 1900 many students besides Maynard and Pilsbry added
materially to the names in this genus ; Dall, Bartsch, Plate, and
Clench for the Bahama Archipelago and Aguayo, Sanchez Roig,
Jaume and Clench for Cuba. Many names were added by H. B.
Baker to the uva complex of the Dutch West Indies, Curacao
and its associated islands.
124 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Mr. Allison V. Armour's yacht, the TJtowana, made several
trips to various islands in the Bahamas and by this means much
Cerion material was collected for this museum mainly by
T. Barbour and J. C. Greemvay.
In 1936, Mr. J. C. Greenway and his brother Gilbert made
extensive collections in the Bahamas by means of a seaplane.
Islands visited were Andros, Grand Bahama, the Abacos, Great
and Little Inagua. I joined them for the exploration of Grand
Bahama and the Abacos. During this same trip I visited also
Eleuthera Island. A year previously I had explored rather ex-
tensively the northern end of Cat Island and Little San Salvador
or Little Island. I was associated on this trip with Henry D.
Russell and John Huntington. Later, students and associates of
mine visited Long Island (Richard W. Foster, Richard McLean
and John Huntington) and Great and Little Inagua (Richard
McLean and Benjamin Shreve). Each of these various trips
necessitated several days in Nassau, and much time was devoted
to collecting Cerion and other mollusks outside of this city on
New Providence Island.
In 1007, Dr. Plate published upon a few species obtained on
islands in the Exuma group. During the summer of 1930, Dr.
Paul Bartsch of the United States National Museum made an
extensive collecting trip in the southern Bahamas visiting such
island groups as: Cay Sal Bank, Ragged Islands, Crooked Island
group, Little and Great Inagua, Caicos and Turks Islands. More
recently, Mr. and Mrs. George F. Kline of Madison, New Jersey,
have added to our series of Cerion from a few islands in the
Exuma group and from cays in the Ragged Islands, Bahamas.
Many others have figured in the exploration of the Bahamas,
mainly with other interests in mind hut, nevertheless, much data
in the form of material have been collected which will aid in the
eventual solution of this mollusk problem. More data on the
historical side are available in the various studies which are
listed in the bibliography.
There are hut three centers of "speeiation" at the present
time, for this genus: Cuba, the Bahamas and the Cayman Is-
lands. In these islands the greatest number of populations occur.
Elsewhere, such as Hispaniola, Puerto Rico and the Virgin
Islands populations are exceedingly few and all appear to be
very closely related. ( hi both Hispaniola and Puerto Rico they
CLENCH : CERIONIDAE 125
are to be found only on the south coasts. It is quite astonishing
that this genus has failed to invade Jamaica.
At this time only a single group in this complex has been
analyzed (Clench and Aguayo, 1952) : the subgenus Umbonis.
This group is limited to the north coast of Cuba and the Baha-
mas. Distributional patterns for the species in this group are
not at all uniform ; they appear to be hit or miss and based upon
chance introductions.
it is interesting to note that this genus succeeded in invading
the Dutch West Indies. These islands are far removed from the
Greater Antilles and the established species complex on them
is quite different from all others in the genus. It is possible that
this was an early introduction and that since then no other mem-
bers have invaded this area, so that all of the present named
entities are exceedingly close in their relationships and appear
to be but unit populations of a closely-knit species.
COLLECTIONS
Major collections of Cerion are to be found in: the Academy
of Natural Sciences, Philadelphia ; American Museum of Natural
History, New York City; Museum of Comparative Zoology.
Harvard University; Museum of Zoology, University of Michi-
gan; Museo Poey, Universidad de la Ilabana, Habana, Cuba;
and the United States National Museum, Washington, D.C.
Many smaller collections exist in most museums, both in the
Americas and in Europe, but the above institutions contain
collections that are rich both in type material and in geographical
series.
GEOGRAPHIC INDEX
The following list of names is arranged geographically to aid
in locating the species of Cerion which have been described from
any given locality. Names of species now considered synonyms
have been omitted from this list. This does not mean that a' I
the names included below refer to valid species but only that to
date no attempt has been made to restudy most of these forms
since they were originally described. A good example is that of
New Providence, Bahamas, given below. Probably no more than
126
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
live or six species exist on this small island, yet the present list
stands at 82!
Fossil species have been included in this list as most of these
"fossil" forms are not old in any geologic sense.
In the list of names below we have retained the original spell-
ing. As Cerion is neuter, all the adjectival specific names should
end in urn.
BAHAMA ISLANDS
Andros Island
albata
bimarginata
eapraia
eamale
casablanca e
•■era
Columbians
erescentia
etolva
grisea
Berry [stands
albolabra
arbusta
aviaria
balaene
berryense
caduca
eana
candida
eonfusa
interealaria
Bimini Islands
biminiensis
Caicos Islands
caicosense
Castle Island
regia
Cat Island
cxiniea
felis
fordii
helena
irregulare
latisinus
lenticularia
normale
obesum
panda
pepper i
persuasa
pilsbryi
jenneyi
litorea
lobata
mixta
obtusa
pictnrata
plebia
porcina
primordia
proavita
lerneri
papilla
regula
restricta
rhyssum
saurodon
sladeni
stupida
variabile
viaregis
procliva
profunda
rara
relequa
scutata
sylvatica
thayeri
travelii
variata
pillsburyi
r'raternuni
lmutiiisi'toni
platei
russelli
CLENCH : CERIONIDAE
127
Cay Sal Bank
nitcloides
Conception Island
fairchildi
Crooked Island
c-liffordi marmorata
inflata niaitensi
Ei-Ei riiKKA Island
eleutherac hyattii
exigua inconsueta
glans indianorum
hughesi laeve
Exuma Group op Islands
accuminator fulvia
agricola genitiva
albicostata gigantea
aspera grayi
eervrna hedwigiae
i-rassa imperfecta
eyelura inconstans
cylindriata inexpecta
'legenis inornata
dissimila inquita
eburnia intentata
elegantissima leueophera
elongata ruariae
exorta marmorosa
extranea milleri
extrema minuta
ritzgeraldi mitra
fruticosa mutatoria
Fortune Island
submarmoratum
Grand Bahama Island
ehrysaloidea oweni
Great Abaco Island
abaeoensia lucayauorum
liendalli inaynardi
Great Exuma Island
adumbra fragilis
'•aerulescens pauli
exasperata pleginatuni
exumense I ml la
tiamea
niultistriatuni
weinlandi
lilionuii
mossi
inulta
imiformis
navalis
nebula
uormanii
palmata
perantiqua
proeessa
prognata
progressa
pumilia
ritchiei
sampsoni
scalariformis
similaria
stroutii
tenucostata
valida
vet a
leticulatum
venniculum
pusilla
recessa
semipolita
transmutata
128
BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
Great Inagua Island
columna rubicunda
dallii tumerae
rehderi
Green Cay
scalarinoides universa
Joulter Cays
ralla
Little Abaco Island
incisum oweni
Little Inagua Island
baconi sarcostornum
ealearea
Long Island
fernandina
josephinae
malonei
Miraguana Island
armouri
barbouri
Mira For Vos Cay
periculosum
New Providence and Adjacent Islands
valida
viola
mcleani
melanostomum
rnanguanense
shrevei
nuda
stevensoni
piraticus
acceptoria
fineastlei
palidula
affinis
Ha cida
phoenecia
agassizi
fulminea
primigenia
agava
glans
prisca
agava-negleeta
gracila
purpurea
agrestina
gubernatoria
pygmea
ajax
hart-bennetii
rod i viva
albata
hesternia
reiucarnata
albea
larga
repetita
angustalabra
latonia
rosacea
antiqua
leva
rosea
argentia
livida
rubiginosa
avita
macularia
rufimaeulata
caerulea
mayoi
rufula
earlotta
migratoria
salinaria
castra
minima
santesoni
cinerea
mobile
saxitina
cinerea -varia
montana
sparsa
clara
morula
sula
CLENCH : CERIONIDAK
12!)
eoncina
eoryi
crassalabra
eurtissii
degeneri
delicata
devereuxi
eratiea
extensa
Plana Cats
utowana
Ragged Islands
juliae
Rum Cay
alba
Sam ana Cay
greenwayi
Turks Islands
blandi
brevispira
comes
Watling Island
inconspicuum
multa
muralia
mutata
neglecta
nivea
novita
oberholseri
oscula
iirownei
eucosmunn
incanoides
percostatum
lacunorum
tenui
territa
thompsoni
thorndikei
tracta
ultima
vagabunda
vetusta
lentiginosa
regma
swift ii
watlingense
CAYMAN ISLANDS
Cayman Brac
copia
intermedia
glaber
lineota
Grand Cayman
caymanerise
martiniana
Little Cayman
acuta
levigata
f estiva
lineota
fusea
nana
intermedia
liarva
perplexa
nitela
pannosa
picta
CLE A
Camaguey Province
aeutieostatum miramarae
bioscai palmeri
rolumbinus paredonis
euspidata pastelilloensis
sanctacruzense
sanzi
saugeti
seopulorum
130
BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
dorotheae
grilloensis
gundlachi
Habana Province
ceiba
eurystoma
fastigata
ignota
pretiosus
sainthilarius
jaumei
noriae
peracutum
rieardi
sellare
tantilhun
roeai
salvatori
striatissinnim
tridentatum
Isle ok Pines
moi'eleti pineria
Las Villas Province
alealdei ebriolum
arangoi herrerai
bennudezi iostoma
eatherwoodianum macTodon
chaplini poeyi
eyclostomum
Matanzas Province
alealdei
canasiense
cardenense
caroli
dickersoni
guillermi
hologlyptuni
Oriente Province
aguayoi
alberti
alleni
banesense
basistriatum
bequaerti
blanesi
cabocruzense
chaparra
eobarrubia
coutini
crassiusculum
dimidiata
disforme
feltoni
feriai
geophilus
infanda
infandulum
ludovici
magistev
maritima
mierostomuni
minusculum
harringtoni
hessei
humberti
jascoense
josephi
lepiduni
longidens
manatiense
raierodon
moralesi
oriental*'
ornatum
pandionis
parvulum
paucicostatum
paucisculptum
polita
pseudoeyclostomum
saguaense
sanctamariae
strigis
subcostulatum
mumiola
obliterata
sagraiana
scripta
sublaevigatum
valdesi
portillonis
portuspatris
prestoni
proteus
ramsdeni
saetiae
scalar ina
smithii
sueyrasi
tanamensis
tenuilabris
torrei
turgidum
vallei
vanattai
victor
CLENCH : CERIONIDAE
131
I'inar Dei, Rio Province
cabrerai hernandezi
cisnerosi hondana
constrictum Johnson i
dominicanum laurcani
Cuba (without specific locality)
hyperlissum kusteri
incrassata
marieliixuiM
sculpta
sisal
wrighti
venusta
DUTCH WEST INDIES
Aruba
arubanum
Bonaire
bonairensis
kralcndijki
Curacao
ilesculptuni
iljerimensis
knipensis
diablpnsis
liutoensis
FLORIDA
uva
incana
s;n'charimeta
vaccinum
HISPANIOLA
t'enuginea
minor
sallei
saona
tortuga
yumaensis
inonaense
striatella
MONA ISLAND
PUERTO RICO
rudif
VIRGIN ISLANDS
striatella
XOTES ON THE GENUS CERION
The following notes are based upon studies in this genus which
were made during a rehabilitation of certain portions of our
collection.
132 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Subgenus STROPHIOPS Dall
Straphiops Dall 1894, Bull. Mus. Comp. Zool., 25, p. 121 (type species, Pupa
deoumana Ferussac (=C. regium) Castle Island, Bahamas).
Pinguita Maynard 1896, Contributions to Science, 3, p. 30 (type species,
Strophia dimidiatia [sic] Pfeiffer, monotypic).
Cyclocerion Bartsch 1952, Kevista de la Sociedad Malacologica ' ' Carlos
de la Torre," 9. p. 1 (type species, Cerion (Cyclocerion) baconi Bartsch).
Both Pinguita Maynard and C yclocerion Bartsch appear to be
absolute synonyms of Strophiops Dall. This group is widely
spread in the Bahamas and along the northern coast of Cuba.
Cerion sanctacruzense Aguayo and Jaume
Cerion sanctacruzense Aguayo and Jaume 1951, Revista de la Sociedad
Malacologica "Carlos de la Torre," 8, p. 14, pi. 1, fig. 14 (Sabanalamar,
.Santa Cruz del Sur, Camagiiey, Cuba ) .
This species was described from Sabanalamar, which is just
east of Santa Cruz del Sur on the southern coast of Camagiiey,
Cuba. It appears to be rather widespread in the Cayos de Doce
Leguas, a long series of small islands that run Avesterly and just
off the coast from Santa Cruz del Sur. Typical sanctacruzense
are smooth, but numerous colonies from the various islands are
both smooth and strongly ribbed. We have specimens from the
following localities : Sabanalamar ; Santa Cruz del Sur ; Cayo
Caguama; Cayo Cochiboca ; Punta Boca de Piedra and Cayo
Anclitas.
Cerion politum Maynard
Strophia marmorata polita Maynard 1896, Contributions to Science, 3.
p. 14, pi. 3, figs. 3-4 (Cabo Cruz, Cuba).
Cerion politum maisianum Pilsbry 1902, Manual of Conchology, (2) 14,
p. 218, pi. 30, figs. 89-91 (Punta Maisi, Cuba).
Both of the above names apply to the same species. Maynard
was in error in giving the type locality as Cabo Cruz. This
species occurs only at Punta de Maisi, at the extreme eastern
end of Cuba. In color it ranges from nearly pure white to mar-
bled with brownish. Both smooth and finely ribbed forms occur
in different colonies as well as mixed in others.
CLENCH : CERIONIDAE 133
Cerion alleni Torre
Cerion alleni Torre L929, Nautilus, 42. no. 3, pi. 4, figs. 10-11 [no descrip-
tion] (Antilla, Cuba).
Cerion madama Sanchez Eoig 1951, Revista de la Sociedad Malaeol6gica
"Carlos de la Torre," 7. p. 112, pi. IS, fig. 9 (Cayo Madama, Bahia Arroyo
Blanco, Mayarf, Oriente, Cuba).
Cerion migueleti Sanchez Eoig 1951, Revista de la Sociedad Malacologica
•'Carlos de la Torre," 7, p. 113, pi. 19, fig. 5 (Cayo Miguel, Boca de
Vaguaneque, Cananova, Sagua de Tanamo, Oriente, Cuba).
Cerion sanchezi Clench and Aguayo 1953, Torreia, no. 18, p. 3, text figs.
4-5, Univ. Habana (Lengua do Pajaro, Bahia de Lebiza, Mayari, Oriente,
Cuba ).
All of tlie above names apply to but a single species. This ap-
pears to be a species which is limited to the margins of rather
large bays. Its distribution extends from Bahia de Banes east to
Bahia de Yaguaneque along the north coast of Oriente, Cuba.
Cerion saxzi Pilsbry and Vanatta
Cerion. sanzi Pilsbry and Yanatta 1898 [1899], Proc. Acad. Nat. Sci.
Philadelphia, p. 478, text fig. 9 (Confites Key, Nuevitas, Cuba).
Cerion royi Aguayo and Jaume 1951, Revista de la Sociedad Malacologica
"Carlos de la Torre," 8. p. 7, pi. 1, fig. 1 (Cayo Cruz, Camagiiey, Cuba).
Cerion circumscriptum Aguayo and Jaume 1951, Revista de la Sociedad
Malacologica "de la Torre," 8. p. 12, pi. 1, fig. 10 (Guanalito, Cayo Ro-
mano, Camagiiey, Cuba).
Cerion tejedori Sanchez Roig 1951, Revista de la Sociedad Malacologica
"Carlos de la Torre," 7, p. 112, pi. 18, fig. 7 (Punta Arenas, Paso de las
Carabelas, Peninsula de Sabinal, Camagiiey, Cuba).
Cerion guajauaense Sanchez Roig 1951, Revista de la Sociedad Malacolo-
gica "Carlos de la Torre," 7, p. 114, pi. 18, fig. 6 (Cayo Grillo, Isla de
Guajaba, Camagiiey, Cuba).
Cerion circumscriptum tenuicallum Aguayo and Sanchez Roig 1953, Mem.
Soc. Cubana Hist. Nat., 21, p. 288, pi. 32, fig. 17 (Cayo Frances, Caibarien,
Las Villas, Cuba).
Cerion circumscriptum romanoensis Aguayo and Sanchez Roig 1953, Mem.
Soc, Cubana Hist, Nat., 21. p. 289, pi. 32, figs. 12, 14 (Cayo Romano,
Camagiiey, Cuba).
Cerion sanzi Pilsbry and Vanatta appears to be the most
widely distributed Cerion throughout the Archipielago de Cama-
giiey, a long chain of cays and little islands on the northern
coast of Cuba. This chain of cays extends from Cayo Guillermo,
134 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
off Punta Alegre to Cayo Sabinal, a distance of 125 miles (200
kilometers).
The synonyms given above all appear to be this species or else
hybrid colonies in which the sanzi characters are most prominent.
Cerion microdon Pilsbry and Vanatta
Cerian mcrassatum microdon Pilsbry and Vanatta 1896, Proc. Acad. Nat.
Sei. Philadelphia, p. 328, pi. 11, fig. 5 (Cuba).
Cerion tenuilabre pygmaeum Pilsbry and Vanatta 189(5, Proc. Acad. Nat.
Sei. Philadelphia, p. 334, pi. 11, fig. 9, (Gibara, Cuba).
The two names given above appear to be the same species. C.
mierodon is exceedingly variable in size and somewhat in colora-
tion. It occurs on both sides of the harbor of Gibara but is very
rare on the eastern side. Most of the specimens from the west
side were collected dead. At the time Dr. Aguayo and I visited
this locality we found even the dead specimens to be localized
in certain areas only and not broadly distributed throughout the
coastal region of the harbor.
ABBREVIATIONS
A few abbreviations have been found necessary to reduce
needless repetition. Other than these, references are given in full
under each species.
Catalogue. "Supplement to Catalogue of Specimens of the
Family Cerionidae" for sale by Charles J. Maynard, West
Newton, Massachusetts. 192-4. This supplement includes the
descriptions of fourteen new species of Cerion without figures.
Contributions. Contributions to Science, By Charles J. Mayn-
ard, Newtonville, Massachusetts. A three volume series (discon-
tinued after Vol. 3 no. 1). Published from April 18S9 to March
1896. Many new species of Cerion were published in this journal.
M. of C. Manual of Conchology (series 2), Academy of Na-
tural Sciences, Philadelphia.
Memorias. Memorias de la Sociedad Cubana de Historia Na-
tural. Museo Poey, Universidad de la Habana, Habana, Cuba.
Proc. ANSP. Proceedings of the Academy of Natural Sci-
ences, Philadelphia.
CLENCH : CERIONIDAE 135
Records, App. Appendix to Records of Walks and Talks with
Xature by C. J. Maynard, "West Newton, Massachusetts. In the
appendices of volumes 5, 6 and 10 of the above series, there are
described and figured numerous species of Strophiops ( =
Cerion).
Revista. Revista de la Sociedad Malacologica "Carlos de la
Torre" Museo Poey, Universidad de la Habana, Habana, Cuba.
Torreia. Published by Museo Poey, Universidad de la Habana,
Habana, Cuba.
Genus CERION Roding
Cerion Roding 1798, Museum Boltenianum, p. 90 (type species, Turbo uva
Linne, subsequent designation, Dall 1894).
Pupa Lamarck 1801, Animaux sans Vertebres, p. 88 (type species, Pupa
urn Linne, monotypic).
Cerium Link 1807, Besehreibung der Naturalien-Sammlung der Univer-
sitiit zu Rostock, p. 131 [emendation for Cerion] (type species, Cerium uva
Linne, subsequent designation, Pilsbry 1918, M. of C. (2) 24. p. 268).
Puppa Denys de Montfort 1810, Conehyliologie Systematique, 2. p. 298,
Paris (type species, Pupa uva Linne, monotypic).
Puparia Rafinesque 1815, Analyse de la Nature, p. 143 [substitute name
for Pupa Lamarck).
Cochlodonta Ferussac 1821, Prodrome, Tableau Systematique des Lima-
rous, Paris, p. 24 [28] and p. 58 (type species, Turbo uva Linne, here
selected).
Cochlodon Sowerby 1825, Catalogue of the Shells in the Collection of the
Late Earl of Tankerville, London, p. 40 [in part] (type species, Cochlodon
uva Linne, subsequent designation, Pilsbry 1918, M. of C, (2) 24, p. 268).
Strophia Albers 1850, Die Heliceen, Berlin, p. 202 (type species, Pupa
mumia Brugiere, subsequent designation, von Martens 1861; non Strophia
Meigen 1832; Stal 1877).
Pulpa Poey 1858, Memorias sobre la Historia Natural de la Isla de Cuba,
Havana, 2. p. 30 [error for Puj)a Lam.] (type species, Pidpa sculpta Poey,
monotypic).
CATALOGUE OF THE CERIONIDAE
abaooensis Pilsbry and Vanatta, Cerion (Maynardia) : 1895, Proc. ANSP,
p. 209; ibid. 1896, p. 332, pi. 11, tig. 11 (Abaco Island [Bahamas]).
acceptoria Maynard, Strophiops: 1913, Records, App., 5. p. 185 (Low
Bay Cay, east end of Rose Island, New Providence, Bahamas).
136 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
accumulator Mayiiard and Clapp, Strophiops: 1920, Records, App., 10,
p. 124, pi. 20, figs. 1-2 (Long Cay, Exuma Group, Bahamas).
acuta Maynard, Strophia: 1889, Contributions, 1. p. 15, pi. 2, fig. 4 41)
(South side of Little Cayman, Cayman Islands). Is C. levigatum Mayn.,
Pilsbry 1901.
acuticostatum Sanchez Roig, Cerion scalarinum: 1948, Revista, 6. p. 68,
pi. 1, fig. 5 (North coast of Cayo Megano Grande, northern Camagiiey,
Cuba).
ad umbra Maynard, Strophiops: 1924, Catalogue, Suppl., p. 2 (Cay south
of Green Turtle Cut, Gt. Exuma, Bahamas).
aedilii Aguayo and A. de la Torre, Cerion enroll: 1951, Revista, 8, p. 22,
pi. 3, fig. 4 (Boca de Bacunayagua, west of the river, Matanzas Prov., Cuba).
affinis Maynard, Strophiops: 1913, Records, App., 5. p. 184 (Sandy and
Green Cays, Rose Island, New Providence, Bahamas).
agassizi Dall, Cerion (Maynardia) : 1894, Bull. Mus. Comp. Zool., 25,
p. 120, figs. 9-10 (west quarry, top of Nassau Ridge, New Providence,
Bahamas).
agava Maynard, Strophia neglecta: 1894 Contributions, 2, p. 152, fig. 48
(Sisal fields west of Nassau, New Providence, Bahamas). Is C. ooryi Mayn.,
Pilsbry 1902.
agava-neglecta Maynard, Strophiops: 1913, Records, App., 5, p. 192 (Sisal
fields west of Nassau, New Providence, Bahamas).
agrestina Maynard, Strophia: 1894, Contributions, 2, p. 179, fig. 60 (6
miles south of Nassau, New Providence, Bahamas).
agricola Maynard, Strophiops: 1924, Catalogue, Supplement, p. 1 (Farm-
ers Cay, 1 mile S.W. of Gt. Guana Cay [Exuma Group] Bahamas).
aguayoi de la Torre and Clench, Cerion: 1932, Nautilus, 45, p. 89, figs. 6-7
(Road to Caletones, 6 km. west of Gibara, Oriente, Cuba).
ajax Maynard, Strophiops : 1924, Catalogue; p. 5 [new name for gigantea
Maynard and Clapp 1921, non gigantea Maynard 1894].
alba Maynard, Strophia: 1899, Contributions, 1, p. 74, pi. 7, fig. 17a-b
(west coast of Rum Cay, Bahamas).
albata Maynard and Clapp, Strophiops: May 1921, Records, App., 10,
p. 132, pi. 30, figs. 3 4 (Hog Cay [4 miles N.W. of] Morgans Bluff, Andros,
Bahamas).
albata Maynard and Clapp, Strophiops: July 1921, Records, App., 10,
p. 145, pi. 41, figs. 7-8 [labeled vagabunda on plate] (Southern end of Rose
Island, New Providence, Bahamas).
albea Maynard, Strophia: 1S94, Contributions, 2, p. 128, fig. 38 (South
side of Spruce Key, New Providence, Bahamas). Is C. varium Bonnet, Pils-
bry 1902.
alberti Clench and Aguayo, Cerion: 1949, Torreia, no. 14, p. 3, pi. 1,
figs. 1-6 (Punta de "El Fuerte, " entrada de la Bahia de Banes, Peninsula
de Ramon, Antilla, Cuba).
CLENCH : CERIONIDAE 137
albicostata Maynard, Strophiops: 1924, Catalogue, Supplement, p. 2
(Long Key, S.E. of Highborn Key [Exuma Group] Bahamas).
albolabra Maynard and Clapp, Strophiops: 1921, Records, App., 10. p. 12!).
pi. 27, figs. 3-4 (Great Harbor Key, Berry Islands, Bahamas).
alcaldei Aguayo and Sanchez Roig, Cerion arangoi: 1953, Memorias, 21,
p. 294, pi. 32, fig. 15 (Playa el Ingles, Yaguanabo, Cienfuegos, Cuba).
alleni de La Torre, Cerion: 1929, Nautilus, 42, pi. 4, figs. 10-11 [no descrip-
tion] (Antilla [Oriente] Cuba).
alvearia Dillwyn, Turbo: 1817, Descriptive Catalogue of Recent Shells, 2.
p. 862 (Santo Domingo and Guadeloupe). [Not recognisable.]
angustalabra Maynard and Clapp, Strophiops: 1921, Records, App., 10,
p. 143, pi. 39, figs. 9-10 (fossil, cliffs, west side of Rose Island, opp. Green
Key, New Providence, Bahamas).
angustocostata Maynard and Clapp, Strophiops : 1921, Records, App., 10.
p. 141 (fossil, Lower Fleming Key [Eleuthera] Bahamas). Is C. exiguum
Mayn., Clench 1952.
anodonta Dall, Strophia (Eostrophia) : 1890, Trans. Wagner Free Insti-
tute of Science, 3., p. 13, pi. 1, fig. 8c-d (fossil, Oligocene, Ballast Point,
Old Tampa Bay, Florida).
antiqua Maynard, Strophiops: 1913, Records, App., 5,, p. 183 (fossil,
Nassau, New Providence, Bahamas).
antonii Kiister, Pupa: 1847, Conehylien-Cabinet (2), 1. pt. 15, p. 92, pi.
10, figs. 7-8 (Berbice [British Guiana]. [Probably Great Inagua, Bahamas.]
apiarium Roding, Cerion: 1798, Museum Boltenianum, (2) p. 90, [refers
to Turbo uva Gmelin].
arangoi Pilsbry and Vanatta, Cerion iostumum : 1896, Proc. ANSP, p.
.".30, pi. 11, fig. 12 (Cienfuegos, Cuba).
arbusta Maynard and Clapp, Stropliiops: 1921, Records, App., 10, p. 133.
pi. 30, figs. 5-6 (Guana Key, Berry Islands, Bahamas).
argentia Maynard, Stropliiops: 1913, Records, App., 5, p. 191 (Three
Silver Keys, New Providence, Bahamas).
argntea Maynard and Clapp, Strophiops : 1921, Records, App., 10, p. 138
(Middle Silver Key, New Providence, Bahamas). [Error for argentia May-
nard].
armourl Clench, Cerion (Strophiops) : 1933, Proc. New England Zool.
Club, 13, p. 96, pi. 1, fig. 4 (South coast of Miraguana Island, Bahamas).
arubanum H. B. Baker, Cerion uva: 1914, Occ. Papers, Univ. Michigan,
Mus. Zool. no. 152, p. 104, pi. 20 (Baranca Alto, Aruba, Dutch West Indies).
aspera Maynard and Clapp, Strophiops: 1920, Records, App., 10, p. 116,
1>I. 1, figs. 9-10 (South end of Great Guana Cay [Exuma Group] Bahamas).
aviaria Maynard and Clapp, Strophiops: 1921, Records, App., 10, p. 130,
pi. 27, figs. 9-10 (Bird Key, Berry Islands, Bahamas).
avita Maynard, Strophiops: 1913, Records, App., 5, p. 190 (fossil, Silver
Key, W. of Nassau Bar, New Providence, Bahamas).
138 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
baeoni Bartsch, Cerion (Cyclocerion) : 1952, Eevista, 9. p. 1, text fig. 1
(Northwest Point, Little Inagua, Bahama Islands).
balaena Maynard and Clapp, Strophiops : 1921, Records, App., 10, p. 131
pi. 29, figs. 3-4 (Whale Key, Berry Islands, Bahamas).
banesensc Clench and Aguayo, Cerion: 1949, Torreia, no. 14, p. 7, pi. 1
figs. 13-15 (east side of Bahia de Sama, Banes, Oriente, Cuba).
barbouri Clench, Cerion (Strophiops) : 1933, Proc. New England Zool
Club, 13, p. 95, pi. 1, fig. 5 (south coast of Miraguana Island, Bahamas).
basistriatum Pilsbry and Vanatta, Cerion (Paraccrion) : 1895, Proc
ANSP, p. 206; ibid. 1896, p. 335, pi. 11, fig. 28 (Cabo Cruz, Cuba).
bendalli Pilsbry and Vanatta, Cerion abacoense: 1896, Proc. ANSP, p
332, pi. 11, fig. 13 ([Great] Abaco, Bahamas).
bequaerti de la Torre and Clench, Cerion aguayoi: 1932, Nautilus, 45, p
91, pi. 6, fig. 8 (dunes at Lucretia lighthouse, near Banes, Cuba).
bermudesi Aguayo and Jaume, Cerion gundlachi: 1951, Eevista, 8, p. 4
pi. 2, fig. 9 (Punta Brava, Caibarien, Las Villas, Cuba).
berryense Plate, Cerion glans: 1907, Archiv fur Rassen-und Gesell. Biol-
ogie, 4. p. 596, pi. 5, fig. e (Great Harbour Cay, Berry Islands, Bahamas).
bidens Beck, Pupa dhrysalis: 1837, Index Molluseorum, p. 82 [based on
Ferussac 1832, Histoire Naturelle General et Particuliere des Mollusques,
pi. 153, fig. 5. Names such as bidens, edentula, normalis, major, minor, etc.
were not used by Beck in any sense for categories below a species, either as
varieties or subspecies. These were descriptive terms only and were used to
indicate minor variations which existed in the material studied, either as
specimens or as figures. Unfortunately many of these names have been used
in subsequent works as validly introduced names. These should be dis-
carded.]
bidens Roding, Cerion: 1798, Museum Boltenianum, p. 9 [based upon
Turbo bidens Gmelin=A*e?n'a bidens Schweigger (Clausiliidae)].
bimarginata Maynard, Strophia: 1894, Contributions, 2. p. 164, fig. 53
(Green Key, east coast of Andros, Bahamas). Is C. griseum Mayn., Pilsbry
1902.
biminiensis Henderson and Clapp, Cerion: 1913, Nautilus, 27, p. 64, pi. 4,
ligs. 9-10 (southern end of North Bimini Cay, Bahama Islands).
bioscai Aguayo and Jaume, Cerion (Paracerion) : 1951, Revista, 8, p. 14,
pi. 1, figs. 11-12 (Punta de Praticos, Nuevitas, Camaguey, Cuba).
blandi Pilsbry and Vanatta, Cerion: 1896, Proc. ANSP, p. 334, pi. 11,
fig. 7 (Turks Island, Bahamas).
blanesi Clench and Aguayo, Cerion: 1951, Revista, 8, p. 70, pi. 11, fig. 1
(Los Cocos, east side of Bahia de Gibera, Cuba).
bonairensis H. B. Baker, Cerion uva: 1914, Occ. Papers, Univ. Michigan,
Mus. Zool., no. 152, p. 105, pi. 21 (Porta Spaiio, Bonaire, Dutch West
Indies).
CLENCH : CERIONIDAE 139
botrys Roding, Cerion: 1798, Museum Boltenianum, p. 00 [based upon
Lister, pi. 585, fig. 43 = Littorina littorea Linnej.
brevispira Pilsbry and Vanatta, Cerion: 1895, Proe. ANSP, p. 209 (Turks
Island [Bahamas]).
brownei Maynard, Strophia: 1889 [1891], Contributions, 1. p. 196, pi. 16,
fig. 4-a (north side of Rum Key, Bahamas).
brunneum Dull, Cerion (Strophiops) : 1905, Smithsonian Misc. Collections,
47, p. 441, pi. 58, fig. 9 (Governors' Harbor, Eleuthera, Bahamas). Is C.
eximewm Mayn., Clench 1952.
bryanti Pfeiffer, Pupa: 1867, Malakozoologische Blatter, 14. p. 130
(southern [Great] Inagua, Bahamas).
caboeruzense Pilsbry and de la Torre, Cerion: 1943, Nautilus, 57. p. 34,
refers to Manual of Conchology, (2) 14. p. 278, pi. 46, fig. 21, description
and figures only, not the name (i.e. stritelhim "Guerin" Pilsbry, is C.
caboeruzense Pilsbry and de la Torre, uon stritellum "Ferassac" Guerin).
oabrerai Aguayo and Sanchez Roig, Cerion mumia: 1953, Mernorias, 21.
p. 283, pi. 32, fig. 1 (Cayo Hicacos o Ines de Soto, N.W. de Puerto Esper-
anza, Pinar del Rio, Cuba).
caelum Maynard and Clapp, Strophiops: 1921, Records, App., 10. p. 133,
pi. 30, figs. 9-10 (Cabbage Key, Berry Islands, Bahamas).
caertdea Maynard and Clapp, Strophiops : 1915, Records, App., 6. p. 181
(Field north of Fort Charlotte, Nassau, New Providence, Bahamas).
caerulescens Maynard and Clapp, Strophiops: 1920, Records, App., 10.
p. 122, pi. 23, fig. 5 (Key north of Key opposite Roseville, Great Exuma,
Bahamas).
eaicosense Clench, Cerion (Strophiops) : 1937, Proc. New England Zool.
Club, 16. p. 23, pi. 1, fig. 4 (Cockburn Town, South Caicos Island, Caicos
Islands, Bahamas).
calcarea Pfeiffer, Pupa: 1847, Zeitschrift fur Malakozoologie, 4. p. 83
(locality unknown [Little Inagua, Bahamas-Bland 1875]).
cana Maynard and Clapp, Strophiops: 1921, Records, App., 10, p. 137,
pi. 34, figs. 3-4 (Fortune Key, Berry Islands, Bahamas).
canasiense Aguayo and A. de la Torre, Cerion ceiba: 1951, Revista, 8.
p. 22, pi. 3, fig. 3 (West of the Boca del Rio/Canasi, Matanzas, Cuba).
Candida Maynard and Clapp, Strophiops: 1921, Records, App., 10. p. 131,
pi. 28, figs. 5-6 (East Marketfish Key, Berry Islands, Bahamas).
eanonicum Dall, Cerion (Strophiops) : 1905, Smithsonian Misc. Collections,
47, p. 439, pi. 48, fig. 13 (Gun Key [Bimini Islands] Bahamas). Is C.
pillsburyi P. and V., Clench 1942.
capilkiris Beck, Pupa: 1837, Index Molluscorum, p. 82 [nomen nudum].
capraia Maynard and Clapp, Strophiops: 1921, Records, App., 10, p. 131,
pi. 28, figs. 7-8 (North Goat Key, Fresh Creek, Andros, Bahamas).
cardenense Aguayo and Sanchez Roig, Cerion miorodon : 1953, Memorias,
140 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
21, p. 285, pi. 32, fig. 7 (El Genoves, Cayos de los Cinco Leguas, Cardenas,
[Matanzas] Cuba).
carlotta Maynard, Strophia: 1894, Contributions, 2, p. 154, fig. 49 (North
side of hill, Fort Charlotte, Nassau, New Providence, Bahamas). Is C.
coryi Mayn., Pilsbry 1902.
carnale Maynard and Clapp, Strophiops : 1921, Records, App., 10, p. 136,
pi. 33, figs. 5-6 (West [one mile] of Morgan's Bluff, Andros Island, Ba-
hamas).
caroli Aguayo and A. de la Torre, Cerion: 1951, Revista, 8, p. 20, pi. 3,
fig. 2 (Near lighthouse at Punto de Guanos, W. of Punta de Sabanilla,
Matanzas, Cuba).
casablancae Bartsch, Cerion: 1920, Carnegie Inst., Washington, 14, pub.
no. 282, p. 33, pi. 2; pis. 32-47 (White House region, Andros, Bahamas).
castra Maynard and Clapp, Strophiops: 1921, Records, App., 10. p. 147.
pi. 42, figs. 9-10 (field west of Williams St., Nassau, New Providence,
Bahamas).
catherwoodianum Wurtz, Cerion: 1950, Proc ANSP, 102, p. 100, pi. 2,
figs. o-<n (Station 7, Cayo Largo, Banco Jardines, Cuba).
caymanense Pilsbry, Cerion: 1902, M. of C, (2) 14, p. 196, pi. 44, figs.
85 86 (Grand Cayman Island [Cayman Islands]).
ceiba Clench, Cerion: 1948, Revista, 6. p. 49, text figs. 1-3 (north side of
Boca del Rio Jibaeoa, Santa Cruz del Norte, Habana, Cuba).
cera Maynard, Strophia bimarginata: 1894, Contributions, 2, p. 168, fit;.
54 (Green Cay, Andros Island, Bahamas). Is C. grisevm Mayn., Pilsbry 1902.
err rina Maynard and Clapp, Strophiops : 1920, Records, App., 10, p. 121,
pi. 22, fi^s. 8-9 (Refuge Key, east of Normans Pond Key [Exuma Group j
Bahamas).
chaparra Aguayo and Sanchez Roig, Cerion: 1953, Memoriae, 21, p. 292,
pi. 32, fig. 19 (Bahia de Puerto l'adre, Oriente, Cuba).
chaplini Wurtz, Cerion: 1950, Proc. ANSP, 102, p. 99, pi. 2, figs. 1 4
(Station 19, Cayo Largo, Banco Jardines, Cuba).
ehristophei Clench, Cerion: 1937, Proc. New England Zool. Club, 16, p. 24,
pi. 1, ±1 j_c . 2 (Northeast Point (Christoph's Palace) Great Inagua, Bahamas:.
chrysalis " Ferussac " Beck Pupa: 1837, Index Molluscorum, p. 82.
chrysaloides Plate, Cerion: 1907, Archiv fiir Rassen- und Gesell. Biologic,
4, p. 597, pi. 5, fig. a (Eight Mile Rock, Great Bahama Island, Bahamas).
cinerea. Maynard, Strophia: 1894, Contributions, 2, p. 119, fig. 35 (Middle
Bay, Hog Island, New Providence, Bahamas). Is C. ruriutn Bonnet.
Pilsbry 1902.
cinerea-vara Maynard, Strophiops: 1913, Records, App., 5, p. 185 (E. end
of Hog Island, New Providence, Bahamas"). [Corrected to varia, ibid. p.
1 99. |
circumscriptum Aguayo and Jaume, Cerion: 1951, Revista, 8, p. 12, pi. 1.
CLENCH : CERIONIDAE 141
fig. 10 (Guanalito, Cayo Romano, northern Camagiiey, Culm). Is C. sanxi
P. and V. See Notes, this report.
cisnerosi Clench and Aguayo, Cerion: 1951, Revista, 8, p. 72, pi. 11.
tigs. 3-4 (Playa Morrillo, 11 kilometers west of Bahia Honda, Pinar del Rio,
Cuba).
clappii Maynard, Strophiops: 1913, Records, App., 5. p. 198 (Salina north
of Current Settlement, Eleuthera, Bahamas). Is C. laevc Plate, Clench 19512.
clara Maynard, Strophiops: 1924, Catalogue, Supp., ]). 4 (Church, East
Bay St. to Fox Hill, Nassau, New Providence, Bahamas).
clathrata Humphrey, Pupa: 1797, Museum Calonnianum, p. 64. [Names
of Cerion, i.e. Pupa, appearing in this sales catalogue are without descrip-
tion, figure or reference. In addition, it has also been ruled (Opinion 51 i
as not acceptable as a source of names.]
cliff ordi Clench, Cerion (Strophiops) martensi: 1933, Proc. New England
Zool. Club, 13. p. 91, pi. 1, fig 3 0 (Landrail Point, Crooked Island, Ba
ha mas).
coarctata Beck, Pupa uva: 1837, Index Molluscorum, p. 82, [see note under
bidens Beck].
cobarrubia Aguayo and Jaume, Cerion: 1951, Revista, 8. p. 8, pi. 1, fig. 2
(Punta Cobarrubia, between Manati and Puerto Padre, Oriente, Cuba).
Cochlodon Sowerby: 1825, Catalogue of the Shells in the Collection of the
Earl of Tankerville, p. 40, London.
Cochlodonta Ferussac : 1821, Prodrome, Tableau Systematique des Lima
i;ons, Paris, p. 24 [2Sj and p. 58.
coluinbiana Maynard and Clapp, Strophiops: 1921, Records, App., 10,
p. 136, pi. 33, figs. 3-4 (Pigeon Key, Staniard Creek, Andros, Bahamas).
columbimus Sanchez Roig, Cerion: 1951, Revista, 7. p. 117, pi. 18, fig. 8
(North coast of Cayo Paloma, north of Cayo Romano, Camagiiey, Cuba).
columna Pilsbry and Vanatta, Cerion (Maynardia) : 1895, Proc. ANSI'.
47, p. 207 (Turtle Cove, Great Inagua, Bahama Islands).
comes Pilsbry and Vanatta, Cerion regina: 1895, Proc. ANSP, 47. p. 20S
(Turks Island [Bahamas]).
concilia Maynard, Strophiops: 1924, Catalogue, Supp., p. 4, (fossil, crab
holes, St. James Corner, Nassau, New Providence, Bahamas).
confusa Maynard and Clapp, Strophiops: 1921, Records, App., 10. p. 130.
pi. 28, figs. 1-2; p. 136, pi. 34, figs. 1-2 (Little Harbor Key, Berry Islands.
Bahama Islands). [This species was described twice as new, the descrip
tions and figures vary a little.]
constrictum Aguayo and Jaume, Cerion marielinum: 1953, Memorias, 21,
p. 275, pi. 31, fig. 11 (La Puntilla, Bahia del Mariel, Pinar del Rio, Cuba).
conus Beck, Pupa: 1837, Index Molluscorum, p. 82 [nomen nudum].
copia Maynard, Strophia: 1889, Contributions, 1. p. 22, pi. 1, figs. 1-2.
7-12; pi. 2, figs. 8-8b (West end of Cayman Brae and north side of Little
Cayman). Is C. pannosum Mayn., Pilsbry 1901.
142 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
copiosa Pilsbry, Strophia: 1901, M of C, (2) 14. p. 187 (Cayman Brae,
Cayman Islands). [A suggested name to replace S. copra Maynard. It has
no value.]
coryi Maynard, Strophia: 1894, Contributions, 2, p. 129, fig. 39 (extreme
west end of New Providence, Bahamas).
costata Beck, Pupa decumana: 1837, Index Molluseorum, p. 82 [see note
under bidens Beck].
costulata Beck, Pupa decumana: 1837, Index Molluseorum, p. 82 [see
note under bidens Beck].
coutini Sanchez Roig, Cerion: 1951, Revista, 7, p. 119, pi. 19, fig. 6 (Taco
Bay, Baracoa, Oriente, Cuba).
crassa Maynard and Clapp, Strophiops : 1920, Records, App., 10. p. 123,
pi. 21, figs. 9-10 (East Hill, Little Norman Key, [Exuma Group] Bahamas).
erassaldbra Maynard and Clapp, Strophiops: 1921, Records, App., 10. p.
143, pi. 39, figs. 7-8 (fossil, in cliffs, E. side of Rose Island [New Providence]
Bahamas).
crassicostata "Maynard" Pilsbry, Cerion: 1896, Proc. ANSP, p. 323
[listed in the synonymy of Cerion grisenm Maynard] (Andros, Bahamas).
crassilabris "Shuttleworth" Sowerby, Pupa: 1875, Conchologia Iconic;).
20. p. 12, fig. 14; Pilsbry 1943, Nautilus, 57. p. 34 (India [Puerto Rico]).
Is Cerion stritellum Guerin, Pilsbry 1943.
orassiusculum "Torre" Pilsbry and Vanatta, Cerion: 1898 [1899], Proc.
ANSP, p. 477, text figs. 7-8 (Cayo Juin, Baracoa, Cuba).
crescentia Maynard and Clapp, Strophiops: 1921, Records, App., 10. p.
136, pi. 33, figs. 9-10 (Calabash Key, N. Andros, Bahamas).
cumingianurn Pfeiffer, Pupa: 1S52 [1854] Proc Zool. Soc. London, p. 68,
(Locality unknown).
curtissii Maynard, Strophia: 1894, Contributions, 2, p. 107, fig. 33 (Ceme-
tery between Waterloo and Nassau, New Providence, Bahamas). Is C.
rarium Bonnet, Pilsbry 1902.
cu.spidata Aguayo and Sam-hez Koig, Cerion circumscriptum romanoen-sis
form: 1953, Memorias, 21, p. 290 [New name for Cerion muinia gigantea
Sanchez Roig, not Maynard 1894].
Cyclocerion Bartseh: 1952, Revista, 9, p. 1 (type species, Cerion {Cyclo-
ceri&n) baconi Bartseh, monotypie). Is a synonym of Strophiops Ball.
cyclostoma Sowerby, Pupa: 1875, Conchologica Iconica, 20. pi. 19, fig.
179 (Cuba) ; non cyclostomum Kiister.
cyclostomvm Kiister, Pupa: 1841, Conehylien-Cabinet, 1. pt. 15, p. 6,
pi. 1, figs. 5-6 (locality unknown [Cayo Prances, Cuba, Arango]).
cyclura Maynard and Clapp, Strophiops : 1920, Records, App.. 10, p. 119,
pl. 3, figs. 3-5 (Bitter Guana Key [Exuma Group] Bahamas).
cylindriata Maynard and Clapp, Strophiops: 1920, Records, App., 10,
p. 123, pl. 23, figs. 8-9 (first key north of Leward Stocking [Lee Stocking]
Key [Exuma Group] Bahamas).
CLENCH : CEB10NIDAE 143
cylindrica Mayuard, Strophia: 1896, Contributions, 3, p. 34, pi. 7, figs. 3 4
(Matthewstown, Inagua, Bahamas). Is C. rubicundum Menke, Pilsbry 190".
dallii Maynard, Stropliia: 1889, Contributions, 1. p. 128, pi. 16, fig. lb
(Inagua, Bahamas).
deani M. Smith, Cerion: 1943, Nautilus, 57. p. 59, pi. 7, fig. 7 (The Cur-
rent, south tip of Abaeo ["The Current" northern Eleuthera] Bahamas).
Is C. laeve Plate, Clench 1952.
decumana "Ferussac" Pfeiffer, Pupa: Monographia Heliceorum Viven-
tium, 2, p. 320 (St. Thomas; Cuba?) [Castle Island, Bahamas]. Is C.
regium Benson, Pilsbry 1902.
decumanus Ferussac, Cochlodonta: 1821, Tableaux Systematiques des
animaux Mollusques, p. .19 [or 63] (locality unknown) [Nomen nudum].
degencri Clench, Cerion: 1948, Revista, 6. p. 50, text figs. 4-6 (Fleeming
Point, New Providence, Bahamas).
degenis Maynard and Clapp, Strophiops : 1920, Records, App., 10. p. 120.
pi. 4, figs. 6-7 (Harvest [Harvey's] Key [Exuma Group] Bahamas).
delicata Maynard, Strophiops: 1913, Records, App., 5. p. 190 (South Ke.i
off Sound Point, New Providence, Bahamas).
desculptum Pilsbry and Vanatta, Cerion: 1896, Proc. ANSP, p. 328, pi. 11
fig. 1 (Curac,oa [Dutch West Indies]).
detrita " Shuttleworth " Pfeiffer, Pupa: 1854, Malakozoologisehe Blatter,
1. p. 205, pi. 3, figs. 9-10 (Florida). Is C. incanum. Binney, Pilsbry 1902.
devereuxi Maynard and Clapp, Strophiops: 1915, Records, App., 6. p. 181
(Devereux Estate, west end of New Providence, Bahamas).
eliablensis H. B. Baker, Cerion uva: 1914, Occ. Papers, Mus. Zool., Univ.
Michigan, no. 152, p. 100, pi. 18, fig. A2 (Ronde Klip, Curasao, Dutch West
Indies).
DiaceHon Dull: 1894, Bull. Mus. Coinp. Zool., 25, p. 122. Type species,
Strophia dallii Maynard.
diekersoni Richards, Cerion : 1935, Jour, of Paleont., 9. p. 257, pi. 25, fig.
24 [not fig. 25 as given in the text] (Pleistocene. Sand in sea cave near
road to Monserrat, Matanzas, Cuba).
iliekersoni Torre and Bennudez, Cerion: 1951, Revista, 8, p. 8 [MS. name
introduced as a synonym of C. royi Aguayo and Jaume, non C. didkersoni
Hie hards].
dimidiata Pfeiffer, Pupa: 1847, Zeitschrift fur Malakozoologie, 4, p. 16
(Cuba).
dimidiatia "Pfeiffer" Maynard, Strophia: 1896, Contributions, 3. p. 30
[error for dimidiata Pfeiffer].
disforme Clench and Aguayo, Cerion: 1946, Revista, 4, p. 85, text figs.
1-6 (Cercanias do Punta Manolito, Peninsula del Ramon, Antilla, Oriente,
Cuba).
dissimila Maynard and Clapp, Strophiops: 1920, Records, App., 10, p. 119,
pi. 24, figs. 8-9 (East Cistern Key [Exuma Group] Bahamas).
144 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
djerimensis H. B. Baker, Cerion uva Tcnipensis: 1914, Occ. Papers, Mus.
Zoo!., Univ. Michigan, no. 152, p. 103, pi. 19, fig. Al (shore cliffs near
Playa Djerimi, Curasao, Dutch West Indies).
dominicanum Clench and Aguayo, Cerion mariclinum: 1951, Eevista, 8
p. 09, pi. 10, figs. 1-3 (east side of Boca del Eio Dominica, about 10 miles
west of Mariel, Cuba).
dorotheae Aguayo and Jaume, Cerion: 1951, Eevista, 8, p. 10, pi. 1, figs.
5-7 (Punta de Cayo Cocos, northern Camagiiey, Cuba).
drupium Dall, Cerion (Strophiops) eleutherae: 1905, [in] The Bahama
Islands, ed. by G. B. Shattuek, Baltimore, p. 35 (fossil, Station 6, above
Glass Window, Eleuthera, Bahamas). Is C. eleutherae P. and V., Clench
1952.
duplodon Pilsbry and Yanatta, Cerion: 1896, Proc. ANSP, p. 337, pi. 11,
fig. 25 (Bahamas).
ebriolum Aguayo and Jaume, Cerion: 1951, Eevista, 8. p. 6, pi. 2, fig.
6 (Cayo Borracho, east of Cayo Frances, Caibarien, Las Villas, Cuba).
eburnia Maynard, Strophia: 1894, Contributions, 2, p. 144, fig. 45 (U Key,
[Exuma Group] Bahamas).
edentula Beck, Pupa chrysalis: 1837, Index Molluscorum, p. 82 [see note
under bidens Beck].
edentula Beck, Pupa urn: 1837, Index Molluscorum, p. 82 [see note uiid'M-
bidens Beck].
elegans Beck, Pupa: 1837, Index Molluscorum, p. 82 [nomen nudum].
elegantissima Maynard and Clapp, Multicostata : 1920, Eecords, App., 10,
p. 126, jil. 25, figs. 1-2 (Norman's Pond Key [Exuma Group] Bahamas).
eleutherae Pilsbry and Yanatta, Cerion: 1896, Proc. ANSP, p. 333, pi. 1 1 ,
figs. 19-20 (Eleuthera, Bahamas).
elongata Maynard, Strophia: 1894, Contributions, 2, p. 14S, fig. 46 (small
key 1 mile north of U Key [Exuma Group] Bahamas). Is C. eburneum
Maynard, Pilsbry 1902.
Eostrophia Dall: 1890, Trans. Wagner Free Inst. Sci., Philadelphia, 3.
p. 12 [type species, Strophia {Eostrophia) anodonta Dall, here selected.]
erotica Maynard and Clapp, Strophiops: 1921, Eecords, App., 10. p. 147,
pi. 42, figs. 7-8 (Fox Hill Village, New Providence, Bahamas).
euoosmium Pilsbry and Yanatta, Cerion regina: 1895, Proc. ANSP, p. 208,
(no locality given [Turks Island]). Is C. regina Pilsbry and Vanatta, here
considered a synonym.
eurystoma Maynard, Strophia: 1896, Contributions, 3. p. 7, pi. 2, figs. 3 4
(Havana, Cuba). Is C. chrysalis Fer., Pilsbry 1902.
evolva Maynard, Strophia pilsbryi: 1894, Contributions, 2, p. 173, fig. 57
(west end of Goat Key, Middle Bight, Andros, Bahamas). Is C. griseum
Mayn., Pilsbry 1902.
exasperata Maynard and Clapp, Stropliiops: 1920, Eecords, App., 10. p.
123, ]il. 23, figs. 1-2 (Key east of Anna's Tract, Great Exuma, Bahamas).
CLENCH : CEBIONIDA 1 : 14:")
cxigua Maynard, Strophiops: 1913, Records, App., 5, p. 196 (southern
portion of Fleming: Key [near northern Eleuthera] Bahamas).
eximea Maynard, Strophia: 1894, Contributions, 2, p. 177, fig. r>9 (Cat
island, Bahamas).
exvmvwm "Maynard" Pilsbry, Cerion : 1902, M. of C, (2) 14. p. 265, pi.
38, figs. 76-78 [error for eximeum] (Cat Island [Bahamas] I.
exorta Maynard and Clapp, Strophiops: 1920, Records, App., 10, p. 122,
pi. 24, figs. 4-5 (Refuge Key, [Exuma Group] Bahamas).
extensa Maynard, Strophiops: 1924, Catalogue Suppl., p. 2, (Baptist
Chapel, East Nassau, New Providence, Bahamas).
extranea Maynard, Strophiops: 1924, Catalogue, Suppl., p. 2 (Roseville
Key, Exuma Group, Bahamas I.
extrema Maynard and Clapp, Strophiops: 1920, Records, App., 10, p. 118,
pi. 2, figs. 10-11 (south end of Great Guana Key [Exuma Group] Bahamas).
exwmenst Plate, Cerion: 1907, Archiv fiir Rassen- und Gesell. Biol., 4, p.
607, pi. 3, figs, b-e-d (two unnamed islands between Shroud CayT and Conch
Cut and Stocking Island, opposite Georgetown, Great Exuma, Bahamas).
fairchildi Clench, Cerion (Strophiops): 1933. Proc. New England Zool.
Club, 13. p. 97. pi. 1, fig. 6 (Conception Island, Bahamas).
fasdata Binney, Pupa: 1859, The Terrestrial Air-Breathing Mollusks of
the United States, 4, pp. 152, 205, pi. 79, fig. 17 (Key Biscayne, Florida'.
Is C. incanum Binney, Pilsbry 1902.
faseiata "Maynard" Pilsbry. Cerion: 1902, M. of C, (2) 14. p. 215 (Key
Vaea, Florida;. [This has been quoted by Pilsbry as named by Maynard but
he was only using Binney 's name for this identical form, 1889, Contributions,
1. p. 133.]
fastigata Maynard, Strophia: 1896, Contributions, 3, p. 6, pi. 2, figs. 1 2
( Matanzas [Havana] Cuba). Is C. chrysalis Fer., Pilsbry 1902.
faxoni Maynard, Strophia: 1896, Contributions, 3, p. 32, pi. 7, tigs. 1 2
(Cuba). Is C. johnsoni Pilsbry and Yanatta, Pilsbry 1902.
felis Pilsbry and Yanatta. Cerion (Maynardia) : 1895, Proc ANSP, p.
206 (Cat Island, Bahamas).
feltoni Sanchez Roig, Ct rlon: 1951, Revista, 7, p. 119, pi. 19, fig. 3 (Felton,
Mayari, Cuba).
ferioi Clench and Aguayo, Cerion vulneratum: 19.13, Torreia, no. 18, p. 3,
text fig. 6 (Cayo Largo o de los Muertos, Bahia o Puerto Xaranjo, Oriente,
Cuba I.
feriai de la Torre, Cerion: 1953, Torreia, no. 18, p. 3 [MS. name in the
synonymy of Cerion vulneratum feriai Clench and Aguayo].
fernandinn Clench, Cerion (Strophiops): 1937, Nautilus, 51. p. 21, pi. 3,
ri'„'. 5 (Millers, 8 miles S.E. of Simms, Long Island, Bahamas).
ferrugmea Maynard, Strophia: 1896, Contributions, 3, p. 19, pi. 4, figs. 5-6
(Jeremie, Haiti; .
146 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
f estiva Maynard, atrophia: Contributions, 1. p. 17, pi. 2, fig. 5a-c (Little
Cayman Island [Cayman Islands]). Is C. levigatum Mayn., Pilsbry 1901.
fineastlei Maynard and Clapp, Strophiops : 1921, Eeeords, App., 10. p. 148,
pi. 44, figs. 5-6 (Fort Fineastle, West Nassau, New Providence, Bahamas).
fitsgeraidi Maynard and Clapp, Strophiops: 1921, Kecords, App., 10. p.
127, pi. 2."i, figs. 6-7 (northern end of Little Exuma Key [Exuma Group]
Bahamas).
flacida Maynard and Clapp, Strophiops: 1921, Becords, App., 10. p. 152,
pi. 48, figs. 3-4 (fossil, in rocks above Queen's Staircase, Nassau, New
P ro vidence, Bahamas ) .
flamea Maynard and Clapp, Strophiops: 1921, Becords, App., 10. p. 128,
pi. 25, figs. 8-9 (Southwest Point, Great Exuma, Bahamas).
floriiianuni Dall, Strophia anodonta: 1890, Trans. Wagner Free Inst. Sci.,
Philadelphia, 3. p. 13, pi. 1, fig. 6 (Oligocene, Ballast Point, Florida).
fordii Pilsbry and Vanatta, Cerion: 1897, Proc. ANSP, 49. p. 365, text
figs. 1-2 (Bahamas [Cat Island]).
fragilis Maynard and Clapp, Stropldops : 1921, Becords, App., 10. p. 128,
pi. 26, figs. 3 4 (second westernmost Brigadier Key [Brigantine] [Gt.
Exuma] Bahamas).
fraternum Pilsbry, Cerion eximium: 1902, M. of C, (2) 14. p. 265, pi.
38, figs. 79-80 (San Salvador [Cat Island, Bahamas]).
fruticosa Maynard and Clapp, Strophiops: 1920, Becords, App., 10. p. 125,
pi. 20, figs. 3-4 (Bush Key [Exuma Group] Bahamas).
fulminea Maynard and Clapp, Strophiops: 1915, Becords, App., 6, p. 182
(east of Fort Winton, New Providence, Bahamas).
fulvia Maynard and Clapp, Strophiops: 1920. Becords, App., 10. p. 121,
pi. 4, figs. 4-5 (Pipe Key [z=Fowl Key, Exuma Group] Bahamas).
fusca Maynard, Strophia: 1889, Contributions, 1. p. 77, pi. 7, fig. 19a;
text figs. 12a-b (west end of Little Cayman, [Cayman Islands]). Is C.
pannosum Mayn., Pilsbry 1901.
fusoata Binney, Strophia: 1885, Bull. P. S. Nat. Mus., 28. p. 484 [error
for faseiata Binney |.
fastis Bruguiere, Bulimus: 1792, Encyclopedic Methodique (Vers), 1. p.
348 (Santo Domingo and Guadaloupe) [not recognisable].
genitiva Maynard ami Clapp, Strophiops: 1920, Becords, App., 10. p. 124,
pi. 22, figs. 3-4 (fossil, south end of Ship Channel Key, [Exuma Group]
Bahamas).
gcophUus Clench and Ayuago, Cerion: 1949, Torreia, no. 14, p. 5, pi. 1, figs.
7 12 (Punta de Piedra, Banes, Cuba).
gigantea Sanchez Eoig, Cerion mumia: 1951, Bevista, 7, p. Ill, pi. 18,
fig. 5 (Cayo Bomano, N. Lat. 22° 24'; W. Long. 78° 6', Cuba). [Is Cerion
cuspidata Aguayo and Sanchez Boig.]
gigantea Maynard and Clapp, Strophiops: 1921, Becords, App., 10, p. 152,
CLENCH : CERIONIDAE 147
pi. 47, figs. 9-10 (wall, Village Road, Sherley St., Nassau, New Providence,
Bahamas). [Is S. ajax Maynard, non S. gigantea Maynard, 1894.]
gigantea Maynard, Strophia grayi: 1894, Contributions, 2. p. 141, fig. 44A
(middle part of Highburn Key [Exuma Group] Bahamas).
glaber Maynard, Strojyhia: 1889, Contributions, 1. p. 25, pi. 2, figs. 10-10b
(west end of Cayman Brae, Cayman Islands). Is C. pannosum Mayn., Pils-
bry 1901.
glans Kiister, Pupa: 1844, Conehylien-Cabinet, (2) 1, pt. 15, p. 74, pi. 11,
figs. 1-2 (locality unknown [New Providence and Eleuthera, Bahamas]).
gracila Maynard, Strophiops: 1924, Catalogue, Suppl., p. 3, (Soldiers
Road, l\-2 miles from south shore, Nassau, New Providence, Bahamas).
grayi Maynard, Strophia: 1894, Contributions, 2, p. 138, fig. 42, (north
end of Highburn Key [Exuma Group] Bahamas).
grc.cmcayi Clench, Cerion (Strophiops) : 1934, Proc. Boston Soc. Nat.
Hist,, 40, p. 200, pi. 2, fig. II (Black Booby Cay (West Booby) Atwoods or
Samana Group, Bahama Islands).
grillocnsis Sanchez Roig, Cerion: 1951, Revista, 7, p. 117, pi. 19, fig. 2
(Cayo Grillo, south coast of Isla Guajaba, [Camagiiey] Cuba).
grisea Maynard, Strophia glans: 1894, Contributions, 2, p. 159, fig. 56,
(1 mile north of Fresh Creek, Andros Island, Bahamas).
gruneri Tfeiffer, Pupa: 1847, Zeitschrift fiir Malakozoologic, 4, p. 15
(locality unknown).
guajabaense Sanchez Roig, Cerion : 1951, Revista, 7, p. 114, pi. 18, fig. 6
(Cayo Grillo, north coast of Isla de Guajaba [Camagiiey] Cuba). Is C.
sami Pilsbry and Yanatta, See notes, this report.
gubernatoria Crosse, Pupa: 1869, Jour, de Conch., 17, p. 186; ibid. 1870,
18, p. 105, pi. 2, fig. 4 (New Providence, Bahamas).
guillermi de la Torre, Cerion mimiola: 1954, Revista, 9, p. 40, pi. 15, fig. 2
(Playa de Bueyvaquita, Matanzas, Cuba).
gundlachi Pfeiffer, Pupa: 1852, Zeitschrift fiir Malakozoologie, 9, p. 175,
(Punta de San Juan de los Perros [Camagiiey] Cuba).
harringtoni Aguayo and Sanchez Roig, Cerion paucicostatum : 1953,
Memorias, 21, p. 292, pi. 32, fig. 18 (Cueva de los Indios, La Patana, Maisi,
Oriente, Cuba).
hart-bennetii Maynard and Clapp, Strophiops : Records, App., 10, p. 146,
pi. 42, figs. 3-4 (Potter's Key, New Providence, Bahamas).
hatoensis H. B. Baker, Cerion uva: 1914, Occ. Papers, Mus. Zool., Univ.
Michigan, no. 152, p. 100, pi. 18, fig. F6 (Seroe Spelonk, near Landhuis
Hato, Curasao, Dutch West Indies).
hedwigiae Plate, Cerion: 1907, Arehiv fiir Rassen- und Gesell. Biol., 4.
p. 605, pi. 3, fig. a (Ship Channel Cay, northern end of Exuma Sound,
Bahama Islands).
helena Maynard, Strophiops: 1914, Records, App., 6, p. 177 (directly
south of Fresh Creek, Andros, Bahamas).
148 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
hernandezi Aguayo and Sanchez Roig, Cerion p'meria : 1953, Memoriae,
21. p. 295, pi. 32, fig. 20 (Cayo Real, Cayos de San Felipe, Pinar del Rio,
[south coast] Cuba).
herrerai Aguayo and Jaume, Cerion vulneratum? : 1951, Revista, 8. p. 3,
pi. 2, figs. 4-5 (Cayo Santa Maria, Caibarien, Las Villas, Cuba).
hessei Clench and Aguayo, Cerion: 1949, Torreia, no. 14, p. 8, pi. 1, figs.
19-22 (Balcon de las Damas, Guarda la Vaca, Banes, Cuba).
hesternia Maynard and Clapp, Strophiops : 1915, Records, App., 6, p. 180
(west end of Booby Rock [16 miles NE of New Providence] Bahamas).
heterodon Pilsbry, Cerion rubicundum: 1902, M. of C, (2) 14. p. 275, pi.
45, figs. 96-98 (Inagua [Bahamas]).
hologlyptum Pilsbry, Cerion sagraianum : 1902, M. of C, (2) 14, p. 216, pi.
30, fig. 83 (Cayo Blanco, near Cardenas, Cuba).
hondana Pilsbry, Cerion mumia: 1902, M. of C, (2) 14. p. 299, (Bahia
Honda, Cuba).
hughesi Clench, Cerium: 1952, Revista, 8. p. 107, pi. 15, figs. 1-3 (Sandy
Point, Savannah Sound, Eleuthera Island, Bahamas).
Itumberti Clench and Aguayo, Cerion: 1949, Torreia, no. 14, p. 4, pi. 1,
figs. 16-18 (Ensenada de Jucaro, Bahia de Banes, Banes, Cuba).
huntingtani Clench, Cerion: 1938, Bull. Mus. Comp. Zool., 80, p. 526, pi. 3,
tigs. 13 (Columbus Point, SE tip of Cat Island, Bahamas).
hyattii Maynard, Strophiops: 1913, Records, App., 5. p. 194 (Bar Bay
Settlement, Current Island, [Eleuthera Island] Bahamas).
hypcrlissum Pilsbry and Vanatta, Cerion: 1896, Proc. ANSP, p. 330, pi.
11, fig. 10 (Cuba).
ianthina Maynard, Strophia: 1889, Contributions, 1, p. 69, pi. 2, figs. 13-
13a (south shore of Inagua, 25 miles from Matthewstown, Bahamas). Is
C. rubicundum Menke, Pilsbry 1902.
ignota Maynard, Strophia eurystoma: 1896, Contributions, 3, p. 9 (Ha-
vana, Cuba). Is C. chrysalis Fer., Pilsbry 1902.
imperfecta Maynard and Clapp, Strophiops : 1920, Records, App., 10. p.
118, pi. 3, figs. 1-2 (south end of Great Guana Key [Exuma Group] Ba-
hamas).
inaguense Clench, Cerion (Biacerion) : 1933, Proc. New England Zool.
Club, 13. p. 98, pi. 1, fig. 9 (Northwest Point, Great Inagua Island, Ba-
hamas).
incana Binney, Pupa: 1851, The Terrestrial Air-Breathing Mollusks of
the United States, 1, p. 109 (nomen nudum) ; ibid., 1852, 2, p. 316 (as P.
maritima Pfr.) ; ibid., 1852, 3, pi. 68, figs. 1-4 (Key West, Florida).
incanoides Pilsbry and Vanatta, Cerion (Maynardia) : 1895, Proc. ANSP,
p. 209 (Turk's Island [Bahamas]).
incisum Dall, Cerion oweni: 1905, Smithsonian Misc. Collect., 47, p. 443,
pi. 58, fig. 10 (Stranger Cay, NVv of Little Abaco, Bahamas). Is C. bendalli
P. and V., Clench 1938a.
CLENCH : CERIONIDAE 140
inconspicuum Dall, Cerion (Strophiops) : 1905, Smithsonian Misc. Collect. ,
47, p. 439, pi. 58, fig. 2 (Watling Island, Bahamas).
inconstant Maynard and Clapp, Strophiops: 1920, Records, App., 10, p.
119, pi. 3, figs. 7-8 (Bird Key [Exuma Group] Bahamas).
inconsueta Maynard, Strophiops: 1913, Records, App., 5. p. 193 (south
portion of Great Pimlico Island [Eleuthera] Bahamas).
incrassata Sowerby, Pupa: 1876, Conchologica Iconica, 20, pi. 1, fig. 6
(Cuba).
indianorum Clench, Cerion paucicostatum : 1934, Proe. Boston Soc. Nat.
Hist., 40. p. 210, pi. 2, fig. F (Wemyss Bight, Eleuthera Island, Bahamas).
inexpecta Maynard and Clapp, Strophiops: 1921, Records, App., 10, p. 127,
pi. 25, tigs. 3-4 (fossil, next to westernmost Brigadier [Brigantine] Key
[Exuma Group] Bahamas).
infanda "Shuttleworth " Poey, Pupa: 1858, Memorias sobre la Historia
Natural de la Isla de Cuba, Habana, 2, pp. 29, 60 (no locality given [Punta
Gorda, Matanzas, Cuba] (Arango)).
infandulum Aguayo and A. de la Torre, Cerion: 1951, Revista, 8, p. 19,
pi. 3, tig. 1 (Punta de Sabanilla, north of Matanzas, Cuba).
inflata Maynard, Stropliia: 1889, Contributions, 1, p. 126, pi. 7, figs. 21-a;
30 a-b (Salena Point, Acklin Island, Bahamas).
inoimata Maynard and Clapp, Strophiops: 1920, Records, App., 10, p. 126,
pi. 3, figs. 11-13 (Bell Key [Exuma Group] Bahamas).
inijuita Maynard and Clapp, Strophiops: 1920, Records, App., 10. p. 121,
pi. 4, figs. 2-3 (Fowl Key, [Exuma Group] Bahamas).
intmtata Maynard and Clapp, Strophiops: 1920, Records, App., 10, p. 118,
pi. 2, figs. 6-7 (south end of Great Guana Key [Exuma Group] Bahamas),
is C. proccssum M. and C, Clench and Aguayo 1952.
intercalaria Maynard and Clapp, Strophiops: 1921, Records, App., 10, p.
129, pi. 26, figs. 9-10 (fossil, Goat Key, Great Harbor Key, Berry Islands,
Bahamas).
intermedia Maynard, Stropliia: 1889, Contributions, 1, p. 13, pi. 2, figs.
3 3b (south side of Little Cayman and south side of Cayman Brae [Cayman
Islands]). Is C. pannosum Mayn., Pilsbry 1901.
iostoma Pfeiffer, Pupa: 1854, Malakozoologische Blatter, 1. p. 204 (south
coast of Cuba [Cienfuegos] (Pilsbry)).
irregulare Plate, Cerion glavs: 1907, Archiv fiir Rassen- und Gesell. Biolo-
gie, 4. p. 594, pi. 4, figs, a and c (Nieholstown, north end of Andros, Ba-
hamas).
jaucoense Aguayo and Sanchez Roig, Cerion tenuilaore : 1953, Memorias,
21, p. 293, pi. 32, fig. 16 (Jauco, Baracoa, Oriente, Cuba).
jaumei Clench and Aguayo, Cerion peracutum: 1953, Torreia, no. 18, p. 2,
text fig. 3 (La Jijira, between Boca de Jaruco and Santa Cruz del Norte,
Habana, Cuba).
150 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
jcnneyi Maynard and Clapp, Strophiops : 1921, Records, App., 10. p. 134,
pi. 31, figs. 9-10 (Anderson's Key [Berry Islands] Bahamas).
johnscmi Pilsbry and Vanatta, Cerion (Maynardia) : 1895, Proc. ANSP,
p. 207 (Locality unknown [Mariel, Pinar del Rio, Cuba] (Clench)).
josephi Clench and Aguayo, Cerion: 1949, Torreia, no. 14, p. 6, pi. 1,
figs. 23-25 (Playa de Uvita, oeste de Caletones, Gibara, Cuba).
josephinae Clench, Cerion (Strophiops) : 1935, Nautilus, 49. p. 49, pi. 3,
figs. 1, 4 (Tate's Bay, SE Long Island, Bahamas).
juliac Clench, Cerion (Strophiops) : 1936, Nautilus, 49, p. 112, pi. 8, fig. 6,
(Great Ragged Cay, Ragged Island Group, Bahamas).
Tcnipensis H. B. Baker, Cerion uva: 1914, Occ. Papers, Mus. Zool., Univ.
Michigan, no. 152, p. 102, pi. 19 (valley between Seroes Palomba and Bana
Hoendoe, Curasao, Dutch West Indies).
l-ralendijl'i H. B. Baker, Cerion uva bonairensis : 1914, Occ. Paper, Mus.
Zool., Univ. Michigan, no. 152, p. 106, pi. 21, fig. A2 (south of Kralendijk,
Bonaire, Dutch West Indies).
Jcusteri Pfeiffer, Pupa: 1852 [1854] Proc. Zool. Soc. London, p. 69;
Kiister 1855, Conchylien-Cabinet, 1, pt. 15, p. 165, pi. 20, figs. 3-6 (locality
unknown [Cuba]).
lacunorum Dall, Cerion (Strophiops) inconspicuum : 1905, Smithsonian
Misc. Collect., 47. p. 439, pi. 58, fig. 4 (shores of lagoon, Watling Island,
Bahamas).
laeve Plate, Cerion: 1907, Archiv fur Rassen- und Gesell. Biologie, 4, p.
601, pi. 1, fig. 9 (Current Harbour, Eleuthera, Bahamas).
larga Maynard, Strophiops : 1913, Records, App., 5. p. 184 (Rose Island,
opposite Green Key, New Providence, Bahamas).
latilabris Pfeiffer, Pupa: 1855, Malakozoologische Blatter, 2, p. 103, pi. 5,
fig. 3 (Blessing, St. Croix [Virgin Islands]). Is C. rude Pfr., Pilsbry 1902.
latisinus Pilsbry and Black, Cerion: 1930, Proc. ANSP, 82., p. 292, pi. 22,
fig. 9a-d (Wide Opening, Andros, Bahamas).
laionia Maynard and Clapp, Strophiops: 1921, Eecords, App., 10,, p. 147,
pi. 43, figs. 7-8 (St. Pauls Quarry, Sherley St., Nassau, New Providence,
Bahamas).
laureani Clench and Aguayo, Cerion: 1951, Revista, 8, p. 74, pi. 11, figs.
7-8 (Cabo Corrientes, Peninsula de Guanaliacabibes, south Pinar del Rio,
Cuba).
lembeyei "Torre" Aguayo and Sanchez Roig, Cerion: 1953, Memorias, 21,
p. 293 (in the synonymy of Cerion sanctacrusense poeyi Ag. and S. R.).
lenticularia Maynard and Clapp, Strophiops: 1921, Records, App., 10, p.
135, pi. 32, figs. 9-10 (Staniard Creek, Andros Island, Bahamas).
lentiginosa Maynard, Strophia: 1S89, Contributions, 1, p. 75, pi. 7, figs.
1818a (Rum Key, interior on west side, Bahamas).
lepidum Clench and Aguayo, Cerion vulneratum: 1951, Revista, 8, p. 76,
pi. 11, figs. 10-11 (Laguna, Punta de Mulas, Banes, Cuba).
CLENCH : CERIONIDAE 151
f
lemeri Clench, Cerion exemium: 1956, American Museum Novitates no.
1794, p. 1, text figs. 1-2 (East Bimini, Bimini Islands, Bahamas).
leucophera Maynard and Clapp, Strophiops: 1925, Eecords, App., 10. p.
181, pi. 52, figs. 1-2 (northern end of Great Guana Key [Exuma Group]
Bahamas).
leva Maynard and Clapp, Strophiops : 1921, Records, App., 10. p. 142, pi.
48, figs. 1-2 (rocks above Queen's stair-case, Nassau, New Providence, Ba-
hamas).
levigata Maynard, Strophia: 1889, Contributions, 1. p. 12, pi. 2, figs. 2-2b
(west end of Little Cayman [Cayman Islands]).
liliorum Clench, Cerion: 1938, Bull. Mus. Comp. Zool., 80. p. 527, pi. 2,
figs. 2-4 (Next Point (east coast) 1% miles ENE of Governor's Harbour,
Eleuthera Island, Bahama Islands).
lineota Maynard, Strophia: 1889, Contributions, 1. p. 20, pi. 2, figs. 7-7b
(south side of Little Cayman and south side of Cayman Brae [Cayman
Islands] ). Is C. pannosum Mayn., Pilsbry 1901.
litorea Maynard and Clapp, Strophiops : 1921, Records, App., 10. p. 133,
pi. 30, figs. 7-8 (south border of Guana Key, Berry Islands, Bahamas).
livida Maynard, Strophiops: 1924, Suppl. Sale Catalogue, p. 4 (West
Bay St., oppo. North Silver Key, Nassau, New Providence, Bahamas).
lobata Maynard and Clapp, Strophiops: 1921, Records, App., 10. p. 134,
pi. 30, figs. 7-8 (Eastern Soldier Key [Berry Islands] Bahamas).
Longidens Maynard: 1896, Contributions, 3, \>. 39 (type species, Strophia
pannosa Maynard, original designation).
longidens Pilsbry, Cerion: 1902, M. of C, (2) 14. p. 212, pi. 32, figs. 23-
24 (Cabo Cruz, Cuba).
lucayanorum Clench, Cerion (Strophiops) : 1938, Memorias, 12. p. 326, pi.
25, fig. 2 (NW portion of Mores Island, 32 miles NW of Southwest Point,
Great Abaco Island, Bahama Islands).
ludovid de la Torre, Cerion ceiba: 1954, Revista, 9. p. 41, pi. 5, fig. 4
(coast between Canasi and Playa de Palmarego, Matanzas Province, Cuba).
maorodon Aguayo and Jaume, Cerion: 1951, Eevista, 8. p. 12, pi. 1, fig. 9
(Cayo Borraeho, east of Cayo Frances, Caibarien, Las Villas, Cuba).
macularia Maynard, Strophiops: 1913, Records, App., 5. p. 189 (south
shore of New Providence, west side of first sound to 2 miles west to a salina).
madama Sanchez Roig, Cerion: 1951, Revista, 7. p. 112, pi. 18, fig. 9 (Cayo
Madama, Bahia Arroyo Blanco, Mayari, Oriente, Cuba). Is C. alleni Torre.
See notes, this report.
magister Pilsbry and Vanatta, Cerion mumia: 1896, Proc. ANSI', p. 322,
pi. 11, fig. 4 (Matanzas, Cuba).
maisianum Pilsbry, Cerion politum: 1902, M. of C, (2) 14. p. 218, pi. 30,
figs. 89 91 (Punta Maisi, Cuba). Is C. politum Maynard. Sec notes, this
report.
152 BULLETIN: MUSEUM OK COMPARATIVE ZOOLOGY
major Beck, 1'npa. uva: 1837, Index Molluscorum, p. 82 [see note under
hklens Beck |.
major Kiister, Pupa multieosta: 1845, Gonchylien-Cabinet, (2) 1, pt. 5,
]>. 77, pi. 10, figs. 1-2 (West Indies).
major Pfeiffer, Pupa rnumiola: 1854, Malakozoologisehe Blatter, 1. pi. 3,
fig. 6 (Playa de Indios, Matanzas, Cuba). Not a subspecies but the name
was used to indicate a large form.
malonei Clench, Cerion (Strophiops) : 1937, Nautilus, 51. p. 20, pi. 3,
fig. 6 (3V2 miles SE of Simms, Long Island, Bahamas).
manatiense Aguayo and Jaurae, Cerion: 1951, Revista, 8. p. 9, pi. 1, fig. 4
(Loma Tabaco, SW of Bahia de Manati, Oriente, Cuba).
vianica Lamarck, Pupa: 1830, Encyclopedic Methodique, 2, pt. 2, p. 401
[error for viumia Brug.].
mariae Maynard and Clapp, Strophiops: 1921, Records, App., 10, p. 128,
pi. 24. fig. 10; pi. 25, fig. 5 (Maria Key, Little Exuma, Bahamas).
marielinum Hand. Cerion johnsoni: 1920, Nautilus, 40, p. 38 [nomen
nudum |.
marielinum "Torre" Pilsbry, Cerion johnsoni: 1927, Nautilus, 40, p. 74,
pi. 1, fig. 10 (Mariel, Pinar del Rio, Cuba).
mariguanense Clench, Cerion (Strophiops): 1933, Proc. New England
Zool. Club, 13, p. 94, pi. 1, fig. 3 (south coast of Mariguana Island, Ba-
hamas).
maritima Pfeiffer, Pupa: 1839, Arehiv fur Naturgesch. Wiegmann, 5. pt.
1, p. 353, (Punta de Maya, Matanzas, Cuba).
marmorata Pfeiffer, Pupa: 1847, Zeitschrift fur Malakozoologie, 4, p. 83
(no locality [Fortune Island, Crooked Island Group, Bahamas]).
marmorosu, Maynard and Clapp, Strophiops: 1920, Records, App., 10, p.
125, pi. 24, figs. 6-7 (Well Key a little north of Leward Stocking Key
[Exuma Group] Bahamas).
martensi Weinland, Pupa: 1802, Malakozoologisehe Blatter, 9, p. 194
(Crooked Island, Bahamas).
martiniana Kiister, Pupa: 1844, Conchylien-Cabinet, (2) 1, pt. 15, p. 75,
pi. 11, figs. 3-4 (West Indies [Grand Cayman, Cayman Islands]).
maynardi Pilsbry and Vanatta, Cerion (Maynardia) : 1895, Proc. ANSI',
p. 210 (Abaco Island [Bahamas]).
Maynardia Dall: 1894, Bull. Mus. Comp. Zool., 25, p. 122 (type species
Strophia neglecta Maynard).
mayoi Maynard and Clapp, Strophiops: 1921, Records, App., 10. p. 148, pi.
43, figs. 9-10 (field east of Mackey St., Nassau, New Providence, Bahamas).
mcleani Clench, Cerion (Strophiops) : 1937, Nautilus, 51. p. 22, pi. 3, fig.
7 (1 mile cast of O'Neills, Long Island, Bahamas).
media Maynard Strophia: 1896, Contributions, 3. p. 18, pi. 4, figs. 3-4
(Cuba). Is C. mumia Brug., Pilsbry 1902.
melanostomum Clench, Cerion: 1934, Proc. Boston Soc. Nat. Hist., 40, p.
CLENCH : CERIONIDAE 153
212, pi. 2, figs, a and c (Mortimers, south end of Long Island, Bahama
Islands).
microdon Pilsbry and Yanatta, Cerion incrassatum: 1896, Proe. ANSP,
p. 328, pi. 11, fig. 5 ([Gibara, Oriente] Cuba).
microstomum Pfeiffer, Pupa: 1854, Malakozoologische Blatter, 1. p. 207,
(Punta de Jicaco [Peninsula de Hicacos, Matanzas Prov.] Cuba).
migratoria Maynard and Clapp, Strophiops: 1921, Kecords, App., 10. p.
1-47, pi. 43, figs. 3-4 (Methodist Sunday School grounds, Sherley St., Nassau,
Xew Providence, Bahamas).
miguelete Sanchez Roig, Cerion: 1951, Revista, 7, p. 113, pi. 19, fig. 5
(Cayo Miguel, Boca de Yaguaneque, Cananova, Sagua de Tanamo, Cuba).
Is C. allcni Torre. See notes, this report.
milleri Pfeiffer, Pupa: 1867, Malakozoologische Blatter, 14, p. 129 (Duck
("ay, Exuma Group, Bahama Islands).
minima Maynard, Strophiops: 1924, Catalogue, p. 4 (St. James corner,
East Nassau, New Providence, Bahamas).
minor Kiister, Pupa striatella: 1847, Conehylien-Cabinet, (2) 1, pt. 15,
1). 92, pi. 11, figs. 13-15 (Haiti).
minusGulum Aguayo and A. de la Torre, Cerion ceiba : 1952, Revista, 9.
p. 35, text fig. 1 (east of Boca del Rio Canasi, Matanzas, Cuba).
minuta Maynard and Clapp, Strophiops palmata: 1920, Records, App., 10.
p. 120, pi. 21, fig. 5 (Wax Key [Exuma Group] Bahamas).
miramarae Sanchez Roig, Cerion: 1951, Revista, 7, p. 116, pi. 18, fig. 1
(Miramar, Punta Domingo, Nuevitas, Camaguey, Cuba).
mitra Maynard and Clapp, Strophiops: 1920, Records, App., 10. p. 118,
pi. 2, figs. 4-5 (south end of Great Guana Key [Exuma Group] Bahamas).
mixta Maynard and Clapp, Strophiops: 1921, Records, App., 10, p. 130,
pi. 27, figs. 5-6 (Bonds Key, Berry Islands, Bahamas).
mobile Maynard and Clapp, Strophiops: 1921, Records, App., 10. p. 146,
pl. 41, figs. 9-10 (west end of Rose Island, New Providence, Bahamas).
monaense Clench, Cerion: 1951, Journal de Conchyliologie, 90, p. 274.
(igs. 7-11 (Mona Island, Puerto Rico).
montana Maynard, Strophiops : 1924, Catalogue, Suppl., p. 3 (Sunnyside
estate, East Bay St., Nassau, New Providence Bahamas).
moralesi Clench and Aguayo, Cerion torrei: 1951, Revista, 8, p. 77, pl. 11,
figs. 13-14 (Playa de Morales, 11 kilometers SE of Banes, Oriente, Cuba).
moreleti Clench and Aguayo, Cerion iostomum: 1951, Revista, 8, p. 73,
pl. 11, rig. 6 (Punta del Este, Isle of Pines, Cuba).
mortuorium de la Torre, Cerion: 1953, [MS name in the synonymy of
Cerion vulneratum feriai Clench and Aguayo].
morula Maynard and Clapp, Strophiops: 1915, Records, App., 6, p. 179
(Spruce Key, [4 miles east of Nassau] New Providence, Bahamas).
mossi Clench, Cerion: 1952, Revista, 8, p. 108 [new name for Cerion pau-
cicostatum Clench 1934 non Torre 1929].
1 54 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
multa Maynard, Strophiops: 1913, Records, App., 5. p. 197 (northern
portion of Fleming Key [about 20 miles NE of New Providence] Bahamas).
Is C. exiguum Mayn., Clench 1952.
Multicostata Maynard: 1920, Records, App., 10. p. 69; 126 (type species,
.1/. eximea Maynard). [Is a synonym of Multistrophia Maynard.]
multicostata "Krister" Sowerby, Pupa: 1S75, Conehologia Iconica, 20.
pi. 2, fig. 13 (Cuba) [error for Pupa multirostum Kiisterj.
multicostum Kiister, Pupa: 181"), Conchylien-Cabhu't, (2) 1. pt. 15, p. 77,
pi. 10, figs. 3 4; pi. 11, figs. 6-7 (West Indies).
multistriatum Pilsbry and Yanatta, Cerion: 1896, Pioc ANSP, p. 335,
pi. 11, fig. S (Crooked Island, Bahamas).
Multostrophia Maynard: 1894, Contributions, 2. p. 177 (type species
Strophia eximea Maynard, original designation).
mumia Bruguiere, Bulimus: 1792, Encyclopedic Methodique, 1, p. 384
(Ocean American [Matanzas, Cuba, fide Pilsbry 1902, p. 225] ).
mumia Sowerby, Pupa: 1834, The Genera of Recent and Fossil Shells,
[it. 41, fig. 2 (no locality given). [Is Cerion regivm Benson, won mumia
Bruguiere.]
mumiola Pfeiffer, Pupa: 1839, Archiv fur Naturgesch. Weigmann, 5, pt. 1.
p. 353 (Playa de Indios, Matanzas, Cuba).
muralia Maynard and Clapp, Strophiops: 1921, Records, App., 10, p. 151,
pl. 47, figs. 5-6 (in walls at East Nassau, New Providence, Bahamas).
mufa "Maynard" Batchelder, Strophia: 1951, Jour. Soc. Bibliography
Natural History, 2. pt. 7, p. 238 [error for mutata Maynard].
mutata Maynard, Strophia einerca : 1894, Contributions, 2, p. 125, fig. 37a
(NW part of Long Key [=Athol Id.] NE of Nassau, New Providence, Ba-
hamas). Is C. varium Bonnet, Pilsbry 1902.
mutatoria Maynard and Clapp, Strophiops: 1920, Records, App., 10. p. 11 0,
pl. 1, figs. 3-4 (south end of Great Guana Key [Exuma Group] Bahamas).
nana Maynard, Strophia: 1889, Contributions, 1. p. 27, pl. 2, figs. 11-lld
(west end of Little Cayman [Cayman Islands]).
navalis Maynard and Clapp, Strophiops: 1920, Records, App., 10, p. 124,
id. 20, figs. 8-10 (north end of Ship Channel Key [Exuma Group] Bahamas).
nebula Maynard and Clapp, Strophiops: 1920, Records, App., 10, p. 122,
pl. 24, fig. 1 (first Key north of Stocking Island, Great Exuma, Bahamas).
neglecta Maynard, Strophia: 1894, Contributions, 2, p. 150, fig. 47, (one
mile west of Fort Charlotte, Nassau, New Providence, Bahamas). Is C.
coryi Maynard, Pilsbry 1902.
nitela Maynard, Strophia: 18S9, Contributions, 1. p. 73, pl. 17, figs. 16-16a
(west end of Little Cayman [Cayman Islands]). Is C. Irrigation Maynard.
Pilsbry 1901.
niteloides Dall, Cerion (Maynardia) : 1S90, Bull. Lab. Nat. Hist. State
Univ. Iowa, 4, no. 1, p. 15, pl. 1, fig. 2 (Water Cay, Salt Cay Bank, Ba-
hamas ) .
CLENCH : CERION'IDAE 155
)i.trra Maynard, Strophia curtissii: 1894, Contributions, 2. p. 102, (ceme-
tery between Waterloo and Nassau, New Providence, Bahamas).
nivia Maynard, Strophiops: 1894, Records, App., 5. p. 186 [error for
iiirca Maynard 1894].
noriae Aguayo and Sanchez Roig, Cerion mumia: 1853, Memorias, 21. p.
•285, pi. 32, fig. 8 (La Noria, Cojimar, Cuba).
normale Pilsbry and Black, Cerion sladeni: 1930, Proc. ANSP, 82. p. 292,
pi. 21, fig. 2 (Purser Point, Andros, Bahamas).
normalis Beck, Pupa chrysalis: 1837, Index Molluscorum, p. 82 [see note
under bidens Beck].
normalis Beck, Pupa uva: 1837, Index Molluscorum, p. 82 [see note under
hi <Iens Beck].
normanii Maynard and Clapp, Strophiops : 1920, Records, App., 10. p. 123,
pi. 20, fig. 5 (West Hill, Little Norman Key [Exuma Group] Bahamas).
northropi Dall, Cerion (Strophiops) : 1905, Smithsonian Misc. Collect.,
47, p. 442, pi. 58, fig. 11 (one of the westernmost islets near Gun Cay,
Bahamas). Is C. pillsburyi P. and V., Clench 1942.
novita Maynard and Clapp, Strophiops: 1921, Records, App., 10. p. 148,
pi. 45, figs. 1-2 (Park west of Fort Montague, East Nassau, New Providence,
Bahamas).
nuda Maynard, Strophia: 1889, Contributions, 1. p. 29, pi. 2, figs. 12a-b
(near Clarence Harbor, Long Island, Bahamas).
oberholseri Maynard, Strophiops: 1913, Records, App., 5, p. 193 (South-
west Bay, New Providence, Bahamas).
obesum Dall, Cerion glans: 1905, Smithsonian Misc. Collect., 47, p. 437,
pi. 58, fig. 15 (Long Cay, North Bight of Andros and Mangrove Cay, South
Bight of Andros, Bahamas).
obesum "Torre" Clench and Aguayo, Cerion: 1951, Revista, 8. p. 80 [a
nomen nudum appearing as a synonym under C. paucicostatum Torre].
obliterata Maynard, Strophia scripta: 1896, Contributions, 3. p. 5, pi. 1,
figs. 5-6 (Matanzas, Cuba). Is C. chrysalis Fer., Pilsbry 1902.
obliterata Maynard, Strophiops : 1913, Records, App., 5, p. 197 (East
Booby Rock [16 miles NE of] New Providence, Bahamas). [Is S. snla
Maynard and Clapp, new name for obliterata Maynard 1913, non 1896].
obscura Maynard, Strophia: 1896, Contributions, 3. p. 21, pi. 3, figs. 5-6
(Cayo Birde del Norte, Cuba). Is C. sagraianum Pfr., Pilsbry, 1902.
obtusa Maynard and Clapp, Strophiops : 1921, Records, App., 10. p. 132,
pi. 29, figs. 9-10 (Cat Key, Berry Islands, Bahamas).
orbicularia Maynard, Strophia: 1890, Contributions, 1, pi. 16, fig. 6a-b
[not described; Inagua, Bahamas, label on type specimens].
orbicularis "Maynard" Pilsbry, Strophia: 1902, M. of C, (2) 14. p. 281
[error for orbicularia Maynard].
orientate Clench and Aguayo, Cerion: 1951, Revista, 8, p. 79, pi. 11, fig. 16
(Punta Nigra, 18 kilometers SW of Punta Maisi, Oriente, Cuba).
156 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
ornatula Maynard, Strophiops: 1913, Kecords, App., 5. p. 197 (Current
Settlement, Eleuthera, Bahamas). Is C. laeve Plate, Clench 1952.
ornatula-clappi Maynard, Strophiops : 1913, Records, App., 5, p. 198
(north of Current Settlement, Eleuthera, Bahamas). Is C. laeve Plate,
Clench 1952.
ornntula-rufina Maynard, Strophiops : 1913, Records, App., 5. p. 198 (south
of Current Settlement, Eleuthera, Bahamas).
ornatum Pilsbry and Vanatta, Cerion torrei: 1898 [1899] Proc. ANSP,
p. 476, text figs. 3-4 (Vita, Cuba).
osoula Maynard and Clapp, Strophiops: 1921, Records, App., 10, p. 146.
pi. 42, figs. 5-6 (Old Thompson Place, Bay St., Nassau, New Providence,
Bahamas).
oweni Dall, Cerion: 1905, Smithsonian Misc. Collect., 47, p. 443, pi. 58,
fig. 12 (south side of Little Abaco, opposite Marsh Harbor and [at] Riding
Point, Grand Bahama, Bahamas).
palidula Maynard and Clapp, Stropliiops: 1921, Records, App., 10, p. 145,
pi. 40, figs. 9-10 (east end of Hog Island, Nassau, New Providence, Ba
hamas).
pallida Maynard, Strophia: 1889, Contributions, 1. p. 70, pi. 2, figs. 1414a
(south shore of Inagua, 15-20 miles from Matthewstown, Bahamas). Is
C. rubieundum Menke, Pilsbry 1902.
palmata Maynard and Clapp, Stroj)hiops: 1920, Records, App., 10, p. 120,
pi. 21, figs. 1-2 (Wax Key [Exuma Group] Bahamas).
palmeri Sanchez Roig, Cerion: 1948, Revista, 6, p. 69, pi. 1, fig. 6 (Cayo
Romano (22° 24' N; 76° 6' W) Cuba).
panda Maynard and Clapp, Strophiops: 1921, Records, App., 10, p. 136,
pi. 33, figs. 7-8 (Saddleback Key, north Andros, Bahamas).
pandionis Aguayo and Jaume, Cerion vulneratmn: 1951, Revista, 8, p. 1,
pi. 2, figs. 13 (Cayo Guineho, NE of the bay of Puerto Padre, Oriente,
Cuba).
pannosa Maynard, Stropliia: 1889, Contributions, 1, p. 10, pi. 1, figs. 2, 13
(west end of Little Cayman [Cayman Islands]).
Paracerion Pilsbry and Vanatta: 1895, Proc. ANSP, 47, p. 206. [Two
species are given, tridentatum P. and V. and basistriatum P. and V., type
species, here selected is Cerion tridentatum P. and V.]
paredonis Pilsbry, Cerion cumin gianum: 1902, M. of C, (2) 14, p. 199,
pi. 32, figs. 21-22 (Cayo Paredon Grande, north shore of Puerto Principe
[Camaguey] Cuba).
parva Maynard, Strophia: 1889, Contributions, 1, p. 24, pi. 2, figs. 9-9b
(west end of Cayman Brae [Cayman Islands]). Is C. pannosum Mayn.,
Pilsbry 1-901.
parvulum Aguayo and Jaume, Cerion gundlachi: 1951, Revista, 8, p. 5,
pi. 2, fig. 12 (Mono Ciego, east of Boca de la Bahia Manati, Oriente, Cuba).
CLENCH : CERIONIDAE 157
pastelilloensis Sanchez Roig, Cerion: 1951, Eevista, 7. p. 114, pi. 18, fig. 4
(Pastelillo, Camagiiey, Cuba).
paurieostatuni Clench, Cerion: 1934, Proc. Boston Soc. Nat. Hist., 40. p.
209, pi. 2, tig. E (Miller Hill, southern Eleuthera, Bahama Islands). Is
C. mossi Clench, Clench 1952.
paucicostatum cle la Torre, Cerion: 1929, Nautilus, 42, no. 3, pi. 4, figs. 8-9
[no description] (Cabo Maisi, Oriente, Cuba).
paucisculptum Clench and Aguayo, Cerion (JOmbonis) : 1952, Occasional
Papers on Mollusks, 1. no. 17, p. 425, pi. 54, figs. 1-2 (Punta de Musica,
Bahia de Sama, Banes, Cuba).
pauli M. Smith, Cerion: 1943, Nautilus, 57, p. 60, pi. 7, fig. 8 (Stewart
Manor Hill, Exuma, Bahamas).
pentodon Menke, Helix: 1846, Zeitsehrift fur Malakozoologie, 3, p. 128,
(locality unknown) ; Pfeiffer 1859, Mono. Heliceorum Viventium, 4. p. 154
(Curasao). [Is Cerion uva Linne — only a young shell (Smith 1898)].
pepperi Bartsch, Cerion (Strop)iiops) : 1913, Proc. U. S. Nat. Mus., 46.
1>. 108, pi. 3, figs. 1, 3, 7-12 (2 miles south of Mastic Point, Andros, Ba-
hamas).
peracuta Bartsch, Cerion: 1931, Science (n.s.), 73, p. 419, [nomen nudum].
peracutum Clench and Aguayo, Cerion sagraianum : 1951, Revista, 8. p.
75, pi. 11, fig. 9 (Boca de Jaruco, Habana Province, Cuba).
perantiqu-a Maynard and Clapp, Strophiops : 1920, Records, App., 10. p.
115, pi. 1, figs. 1-2 (south end of Great Guana Key [Exuma Group] Ba-
hamas).
percostatum Pilsbry and Vanatta, Cerion regina: 1895, Proc. ANSP, p.
208 (Turks Island [Bahamas]).
perieulosum Clench, Cerion {Strophiops): 1934, Proc. Boston Soc. Nat.
Hist., 40. p. 215, pi. 2, fig. B (South Cay, Mira Por Vos Group, Bahamas).
perplexa Maynard, Strophia: 1889, Contributions, 1, p. 71, pi. 7, figs. 15-
15a (Cayman Brae, 2 miles from west end [Cayman Islands]). Is C. pan-
nosum Mayn., Pilsbry 1901.
persuasa Maynard and Clapp, Strophiops: 1921, Records, App., 10, p. 131,
pi. 28, figs. 9-10 (along shore north of Fresh Creek, Andros, Bahamas).
phoenecia Maynard and Clapp, Strophiops: 1921, Records, App., 10, p.
149, pi. 45, figs. 3-4 (Waterloo, East Nassau,1 New Providence, Bahamas).
picta Maynard, Strophia: 1889, Contributions, 1, p. 18, pi. 2, figs. 6-6b
(west end of Little Cayman [Cayman Islands]). Is C. levigatum Mayn.,
Pilsbry 1901.
pieturata Maynard and Clapp, Strophiops: 1921, Records, App., 10. p. 135,
pi. 32, figs. 7-8 (fossil, Cabbage Key, Berry Islands, Bahamas).
pillsburyi Pilsbry and Vanatta, Cerion: 1897, Proc. ANSP, 49, p. 366,
text fig. 5, (Gun Cay [Bimini] Bahamas).
pilsoryi Maynard, Strophia: 1894, Contributions, 2. p. 170, fig. 55 (Goat
158 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Key, Middle Bight, Andros, Bahamas). Is C. griseum Mayn., Pilsbry, 1902.
pineria Dall, Cerion (Maynardia) : 1895, Proc. U. S. Nat. Mus., 18. p. 6,
(Isle of Pines [Cuba]).
pinguis Humphrey, Pupa: 1797, Museum Calonnianum, p. 64 [see note
under olathrata Humphrey].
Pinguita Maynard: 1896, Contributions, 3. p. 30 [type speeies, Strophia
dimidiatia [sic] Pfeiffer, monotypie],
piraticus Clench, Cerion: 1937, Proe. New England Zool. Club, 16. p. 64,
pi. 3, fig. 1 (Southeast Point, 12 miles SE of Abrahams Bay, Mariguana
Island, Bahamas).
platei Clench, Cerion (Strophiops) : 1933, Proc. New England Zool. Club,
13. p. 90, pi. 1, figs. 7-8 (Bight road, Cat Island, Bahamas).
plcbia Maynard and Clapp, Strophioiis: 1921, Eecords, App., 10. p. 130,
pi. 28, figs. 3-4 (Lignum Vitae Key, Berry Islands, Bahamas).
plegmatum Dall, Cerion (Strophiops) : 1905, Smithsonian Misc. Collect.,
47. p. 441, pi. 58, fig. 5 (Exuma Island, Bahamas).
poeyi Aguayo and Sanchez Boig, Cerion sanctacruzense : 1953, Memorias,
21. p. 293, pi. 32, fig. 11 (Trinidad, Cuba).
polita Maynard, Strophia marmorata: 1896, Contributions, 3. p. 14, pi. 3,
figs. 3-4 (Cabo Cruz [is Cabo Maisi] Cuba).
pordna Maynard and Clapp, Strophiops: 1921, Becords, App., 10, p. 132,
pl. 29, figs. 7 8 (Frazar's Hog Key, Berry Islands, Bahamas).
portillonis Welch, Cerion ramsdeni: 1934, Nautilus, 47, p. 105, pl. 11, fig. 2
(near Portillo, 10 kilometers east of Ensenada de Mora, Oriente, Cuba).
portuspatris Aguayo and Sanchez Roig, Cerion josephi : 1953, Memorias.
21. p. 291, pl. 32, fig. 10 (Cayo Juan Claro, Bahia de Puerto Padre, Oriente.
Cuba).
praedicta Maynard and Clap]), Strophiops: 1915, Records, App., 6. p. 181,
(fossil, Great Pimlico [30 miles NE of New Providence] Bahamas). Is C.
uniforme Mayn., Clench 1952.
praedivina Maynard, Strophiops: 1913, Records, App., 5, p. 196 (fossi!.
Upper Fleming Key [20 miles NE of New Providence] Bahamas). Is C.
exiguum Mayn., Clench 1952.
praedivina-univrrm Maynard, Strophiops: 1913, Records, App., 5, p. 196
(fossil, upper Fleming Key [20 miles NE of New Providence] Bahamas ).
Is C. exiguum Mayn., Clench 1952.
prestoni Sanchez Roig, Cerion: 1951, Revista, 7, p. 118, pl. 19. fig. 4
(Preston, Nipe, Cuba).
pretiosus Sanchez Roig, Cerion : 1951, Revista, 7, p. 115 pl. 18, figs. 2 2a
(Playa Bonita, f rente al Faro de Maternillos, Camagiiey, Cuba).
primigenia Maynard, Strophiops: 1913, Records, App., 5, p. 184 (fossil
1 1 neath a sand cliff, east end of Salt Key, New Providence, Bahamas).
peimordia Maynard and Clapp, Strophiops: 1921, Records, App., 10, p.
132, pl. 29, figs. 1 2 (Great Stirrup Key [Berry Islands] Bahamas).
CLENCH : CERIONIDAE 159
prisca Maynard and Clapp, Strophiops: 1915, Eecords, App., 6. p. 182
(fossil, east end of Salt Key, New Providence, Bahamas).
proavita Maynard and Clapp, Strophiops: 1921, Eecords, App., 10. p. 129,
pi. 26, figs. 9-10 (fossil, Goat Key, Great Harbor, Berry Islands, Bahamas)
processa Maynard and Clapp, Strophiops : 1920, Records, App., 10. p. 11(5,
pi. 1, figs. 7-8 (south end of Great Guana Key [Exuma Group] Bahamas).
procliva Maynard and Clapp, Strophiops: 1921, Records, App., 10. p. 134.
1)1. 31, figs. 5-6 (Goat Key, Great Harbor [Berry Islands] Bahamas).
profunda Maynard and Clapp, Strophiops : 1921, Records, App., 10. p. 129,
pi. 27, figs. 1-2 (fossil, Goat Key, Great Harbor, Berry Islands, Bahamas).
prognata Maynard and Clapp, Strophiops: 1920, Records, App., 10. p. 118,
pi. 2, figs. 8-9 (south end of Great Guana Key [Exuma Group] Bahamas).
progressa Maynard and Clapp, Strophiops : 1920, Records, App., 10. p.
119, pi. 2, figs. 1-2 (north end of Great Guana Key [Exuma Group] Ba
hamas).
proteus Pfeiffer and Gundlach, Pupa: 1861, Malakozoologische Blatter, 7.
p. 19, (Gibara, Cuba). Is C. dimidiatum Pfr., Pilsbry 1902.
pseudocyclostomum Aguayo and Sanchez Roig, Cerion: 1953, Memorial
21. p. 289, pi. 32, fig. 13 (Cayo Frances, Caibarien, Cuba).
pudicum Pilsbry, Cerion bryanti: 1902, M. of ('.. (2) 14. p. 273, pi. 4''.
figs. 17-18 (no locality given [Great Inagua] ).
pulla Maynard and Clapp, Strophiops : 1920, Records, App., 10. p. 122.
pi. 23, figs. 3-4 (small key opposite Roseville, Great Exuma, Bahamas).
PuJpa Poey: 1858, Memorias sobre la Historia Natural de la Isla de Cuba.
2. p. 30 [error for Pupa; type species, Pulpa sculpta Poey = Cerion sculptum
Poey].
pa in ilia Maynard, Strophia grayi: 1894, Contributions, 2. p. 143, fig. 441!
(NE end of Highburn Key [Exuma Group] Bahamas).
pupa Boding, Cerion: 1798, Museum Boltenianum, p. 90 [based upon Heli.r
pupa Gmel., a species in the genus Mast us].
papilla Ball, Cerion (Strophiops) variabile: 1905, Smithsonian Misc. Col
lect., 47. p. 440, pi. 58, fig. 1 (Red Bay, northwest end of Andros, Bahamas;.
purpurea Maynard, Strophiops: 1913, Records, App., 5. p. 188 (Creek
Settlement and one mile east [East Point Light] New Providence, Bahamas I.
pusilia "Maynard" Batchelder, StropJiiops : 1951. Jour. Soc. Bibliography
Nat. Hist., 2. p. 255 [error for pusilia Maynard and Clapp].
pusilia Maynard and Clapp, Strophiops: 1921, Records, App., 10, p. 127,
pi. 22, fig. 5 (key near Long Rock, Great Exuma, Bahamas).
pygmaeum Pilsbry and Vanatta, Cerion: 1896, Proc. ANSP, p. 334, pi. 11.
fig. 9 (Gibara, Cuba). Is C. microdon P. and V. See Notes, this report.
pygmca Maynard, Strophiops: 1924, Catalogue, Suppl., p. 4 (fossil, crab
holes, St. James Corner, Nassau, New Providence, Bahamas).
ralla Maynard and Clapp, Strophiops: 1921, Records, App., 10. p. 137.
160 BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY
pi. 34, figs. 8-9 (Joulter Keys, north of Andros, Bahamas).
ramsdeni "Torre" Welch, Cerion: 1934, Nautilus, 47. p. 105, pi. 11, fig.
la-e (Playa Rineon, Ensenada de Mora, Oriente, Cuba).
rara Maynard and Clapp, Strophiops: 1921, Records, App., 10. p. 133,
pi. 31, figs. 1-2 (fossil, west side, Cabbage Key, Berry Islands, Bahamas).
reeessa Maynard and Clapp, Strophiops: 1920, Records, App., 10. p. 122,
pi. 24, figs. 2 3 (southern portion of Stocking Island, Great Exuma, Ba-
hamas).
rediviva Maynard, Strophiops: 1913, Records, App., 5. p. 187 (west of St.
Paul quarry, East Nassau, New Providence, Bahamas).
regia Benson, Pupa: 1849, Ann. Mag. Nat. Hist., (2) 4. p. 125 (Nanking,
China [Castle Island, Bahamas]).
regina Pilsbry and Vanatta, Cerion (Maynardia) : 1895, Proc. ANSP, p.
208; ibid., 1896, p. 330, pi. 11, figs. 23-24 (Turks Island [Bahamas]).
regula Maynard, Strophia: 1894, Contributions, 2. p. 161, fig. 52 (Fresh
Creek, near settlement, Andros, Bahamas). Is C. griseum Mayn., Pilsbry
1902.
rehderi Clench and Aguayo, Cerion (Umbonis) : 1952, Occ. Papers on Mol-
lusks, 1, no. 17, p. 422, pi. 57, fig. 2 (Jackline, one mile west of Conch Shell
Point, Great Inagua, Bahama Islands).
reincarnata Maynard and Clapp, Strophiops: 1921, Records, App., 10.
p. 148, pi. 44, figs. 1-2 (Ocean Hole, east of Mackey St., Nassau, New Provi-
dence, Bahamas).
relequa Maynard and Clapp, Strophiops: 1921, Records, App., 10, p. 130,
pl. 27, figs. 7-8 (Holms Key, Berry Islands, Bahamas).
repetita Maynard and Clapp, Strophiops: 1921, Records, App., 10, p. 149,
pl. 45, figs. 5-6 (fields off Kemp's road, East Nassau, New Providence, Ba-
hamas).
restricts Maynard, Strophia: 1894, Contributions, 2. p. 175, fig. 58 (middle
of Goat Key, Middle Bight, Andros, Bahamas). Is C. griseum Mayn., Pils-
bry 1902.
reticulatum Dall, Cerion oweni: 1905, Smithsonian Misc. Collect., 47. p.
143, pl. 58, fig. 8 (Sugar Loaves Rocks, NW of Elbow Cay, Great Abaco,
Bahamas). Is C. hendalli P. and V., Clench 1938a.
rhysstun Dall, Cerion (Strophiops) : 1905, [in] The Bahama Islands, The
Geographic Society of Baltimore, Maryland, ed. by G. B. Shattuck, p. 31,
pl. 12, fig. 46 (fossil in aeolian rock, Station 14, small unnamed key south
of Reids Cay, Middle Bight, Andros, Bahamas).
ricardi Clench and Aguayo, Cerion: 1951, Revista, 8. p. 71, pl. 11, fig. 2
( Punta de Tarara, Habana Province, Cuba).
ritchiei Maynard, Strophia: 1894, Contributions, 2. p. 135, fig. 41 (High-
liurn Key [Exuma Group] Bahamas).
robusta Maynard, Strophia cinerea: 1894, Contributions, 2, p. 121, fig.
CLENCH : CERIONIDAE 161
36a-b (north side of Hog Island, New Providence, Bahamas). Is C. varium
Bonnet, Pilsbry 1902.
rocai Clench and Aguayo, Ccrion tridentatum : 1953, Torreia, no. 18, p. 2,
text fig. 2 (Lagunas Salobres de Boca de Guanabo, Habana, Cuba).
romanocmsis Aguayo and Sanchez Roig, Ccrion circumscriptum: 1953,
Memorias, 21. p. 289, pi. 32, figs. 12-13 (Cayo Romano, Camagiiey, Cuba).
Is C. sanzi P. and V. See notes, this report.
rosacea Maynard and Clapp, Strophiops: 1921, Records, App., 10. p. 13!'.
pi. 35, figs. 7-8 (West [North?] Silver Key, Nassau, New Providence, Ba-
hamas).
rosea Maynard and Clapp, Strophiops : 1921, Records, App., 10. p. 151,
pi. 48, figs. 9-10 (fossil, north east end of Rose Island, New Providence.
Bahamas).
royi Aguayo and Jaume, Ccrion: 1951, Revista, 8. p. 7, pi. 1, fig. 1 (Cayo
Cruz, northern Camagiiey, Cuba). Is C. sansi Pils. and Van. See notes, this
report.
rubicunda Menke, Pupa: 1829, Yerzeiclmis Conchy. -Samml. Malsburg.
Pyrmont, p. 8 (no locality given [Great Inagua]).
rubiginosa Maynard and Clapp, Strophiops: 1921, Records, App., 10. p.
147, pi. 43, figs. 1-2 (field east of Methodist church, Sherley St., Nassau, New
Providence, Bahamas).
rubra Humphrey, Pupa : 1787, Museum Calonnianum, p. 64 [see note under
cAathrata Humphrey].
rudis Pfeiffer, Pupa: 1855, Malakozoologische Blatter, 2. p. 102, pi. 5,
figs. 1-2 (subfossil, Diamond, Blessing and Paradise Plantations, St. Croix.
[Virgin Islands] ).
rufimaculata Maynard, Strophiops: 1913, Records, App., 5. p. 189 (south
shore of NeAv Providence, west side of salina to Sound Point, Bahamas).
rufina Maynard, Stropliiops: 1913, Records, App., 5. p. 198 (south of
Current Settlement, Eleuthera, Bahamas). Is C. hyattii Mayn., Clench 1952.
rufolabris Beck, Pupa uva: Ps37, Index Molluscorum, p. 82, [see note
under bidens Beck].
rufula Maynard, Sitrophiops : 1924, Catalogue, Suppl., p. 3 (west side of
Kemp's Road, St. James Corner, Nassau, New Providence, Bahamas).
russelli Clench, Cerion: 1938, Bull. Mus. Comp. Zool., 80, p. 528, pi. 1,
figs. 5-8 (near Turtle Cove, 4 miles NNE of The Bight, central Cat Island,
Bahama Islands).
saccluirimeta "Blanes" Pilsbry and Vanatta, Cerion ineanum: 1898
[1899] Proc. ANSP, p. 447, text fig. 5 (Sugar Loaf Key, Florida).
saetiac. Sanchez Roig, Cerion: 1948, Revista, 6. p. 67, pi. 1, fig. 7 (Playa
del Cristo-Saetia, Bahia de Nipe, Cuba).
sagraiana Pfeiffer, Pupa: 1847, Zeitschrift fiir Malakozoologie, 4. p. 15,
(Cayo Galindo [Matanzas] Cuba).
162 BULLETIN": MUSEUM OF COMPARATIVE ZOOLOGY
saguaensfi Aguayo and Sanchez Roig, Cerion sagraianum : 1953, Memorias,
21, p. 286, pi. 32, fig. 9 (Cayo Roteno, Sagua la Grande [Las Villas] Cuba).
sainthilarius Sanchez Roig, Cerion: 1951, Revista, 7. p. 115, pi. 18, fig. 3
(Fuerte San Ililario, Sabinal, Nuevitas, Camagiiey, Cuba).
salimaria Maynard, Strophiops : 1913, Records, App., 5, p. 184 (Salt Key,
New Providence, Bahamas).
sallci Pilsbry and Vanatta, Cerion crassilabre : 1896, Proc. ANSP, p. 325
(San Domingo). Is C. yumaense P. and V., here considered a synonym.
salvatori "Torre" Pilsbry, Cerion: 1927, Nautilus, 40, p. 74, pi. 1, fig. 11
(Jaimanitas [Habana] Cuba).
sampsoni Maynard and Clapp, Strophiops : 1920, Records, App., 10, p. 121,
pi. 4, figs. 1, 10 (Sampson's Key [Stanyard Cay on chart — 3^ miles NYV
of Great Guana Cay, Exuma Group] Bahamas).
sanchezi Clench and Aguayo, Cerion: 1953, Torreia, no. 18, p. 3, text
figs. 4-5 (Lengua de Pajaro, Bahia de Lebiza, Mayari, Oricnte, Cuba). Is
C. alleni Torre. See notes, this report.
sanetacrusense Aguayo and Jaume, Cerion: 1951, Revista, 8. p. 10, pi. 1,
fig. 14 (Sabanalamar, Santa Cruz del Sur, Camagiiey, Cuba).
sanetamariae Aguayo and Jaume, Cerion: 1951, Revista, 8, p. 13, pi. 1,
fig. 13 (Cayo Santa Maria, XE of Caibarien, Las Villas, Cuba).
santesoni Maynard and Clapp, Strophiops: 1921, Records, App., 10, p. 139,
pi. 36, figs. 3-4; pi. 15, fig. 5 (north shore of New Providence, west of
Nassau, Bahamas).
sanzi "Blanes" Pilsbry and Vanatta, Cerion: 1898 [1899] p. 478, text fig.
it (Confites Key, Nuevitas [Camagiiey] Cuba).
saona Vanatta, Cerion: 1924, Proc. ANSP, 75. p. 360, text fig. 3 (Saona
Island, Santo Domingo). Is C. yumaense P. and V., here considered a
synonym.
sarcostomum Pilsbry and Vanatta, Cerion: 1896, Proc. ANSP, p. 331.
pi. 11, fig. 16, (Little Inagua, Bahamas).
saugeti Aguayo and Jaume, Cerion inanatUnse : 1951, Revista, 8, p. 9.
pi. 1, fig. 3 (SW of Bahia de Nuevas Grandes, about 3V& miles from its
mouth, Camagiiey, Cuba).
saurodon Dall, Cerion (Strophiops) raruibile: 1905, Smithsonian Misc.
Collect., 47, p. 440, pi. 58, fig. 14 (Red Bay, NW end of Andros Island, Ba
hamas).
saxitina Maynard and Clapp, Strophiops : 1921, Records, App., 10. p. 145,
pi. 41, figs. 1-2 (Hog Island, east of Three Bays, New Providence, Bahamas i.
scalariformis Maynard and Clapp, Strophiops : 1920, Records, App., 10,
p. 116, pi. 1, figs. 5-6 (south end of Great Guana Key [Exuma Group]
Bahamas). Is C. asperum M. and C, Clench and Aguayo 1952.
scalarina "Gundlach" Sowerby, Pupa: 1875, Conchologia Iconica, 20, pi.
17, fig. 153 [figure and description is for Granopitpa scalaris Benoit from
CLENCH : CERIONIDAE 1 ft]
Sicily ; the reference is to Cerion scalarinum Pfeiffer and Gundlach from
Cuba | .
scalarina Pfeiffer and Gundlach, Pupa: 1860, Malakozoologische Blatter,
7. p. 19 (Gibara [Oriente] Cuba).
soalarinoides Plate, Cerion (flans: 1907, Arehiv fur Rassen- und Gesell.
Biologic, 4, p. 595, pi. 4, fig. f (Green Cay [east of Tongue of the Ocean
and west of Exuma bank] Bahama Islands).
seopulorum Aguayo and Jaume, Cerion: 1951, Revista, 8. p. 11, pi. 1, fig.
8, (Punta SE of Cayo Megano Grande and NE of Cayo Cruz, northern
Camagiiey, Cuba).
scripta Maynard, Strophia: 1896, Contributions, 3. p. 3, pi. 1, figs. 3-4
(Cardenas [Matanzas] Cuba). Is C. chrysalis Fer., Pilsbry 1902.
seulpta Poey, Pulpa [sic] : 1858, Memorias sobre la Historia Natural de
la Isla de Cuba, 2, p. 30, pi. 2, fig. 22 ([northern coast of Pinar del Rio]
Cuba).
scutata Maynard and Clapp, Strophiops : 1921, Records. App., 10. p. 133,
1>1. 31, figs. 3-4 (Petit Key [Berry Islands] Bahamas).
sellare Aguayo and Sanchez Roig, Cerion sansi: 1953, Memorias, 21, p.
291, pi. 32, fig. 6 (Silla de Cayo Romano, Camagiiey, Cuba).
semipolita Maynard and Clapp, Strophiops: 1920, Records, App., 10, p.
123, pi. 23, figs. 6-7 (3rd Key south of Roseville, Great Exuma, Bahamas).
Seniculus Maynard: 1896, Contributions, 3, p. 17 (type species, Strophia
mumia Bruguiere, original designation). [Is a synonym of Strophi-a Albers.]
shrcici Clench and Aguayo, Cerion (Umbonis) : 1952, Oec. Papers on Mol
lusks, 1, no. 17, p. 436, pi. .17, fig. 4 (near North West Point, Little Inagua,
Bahamas).
simihiria Maynard and Clapp, Strophiops: 1921, Records, App., 10. p. 128,
pi. 26, figs, ."it) (westernmost Brigadier [Brigantine, Exuma Group] Ba-
hamas ).
similaris "Maynard" Batehelder, Strophiops: 1951, Jour. Soc. Bibliog
iaphy Natural History, 2, p. 255 [error for similaria Maynard].
sisal Clench and Aguayo, Cerion {Umbonis): 1952, Occ. Papers on Mol
lusks, 1. no. 17, p. 427, pi. 57, fig. 3 (east side — Boca de Mosquito, Mariel,
Pinar del Rio, Cuba).
sladeni Pilsbry and Black, Cerion: 1930, Proc. ANSP, 82, p. 290, pi. 21.
fig. 1 a-1 (Mastic Cay, in Middle Bight. Andros, Bahamas).
smithii "Blanes" Pilsbry, Cerion crassiusculum : 1902, M. of C, (2) 14,
p. 202, pi. 32, fig. 38 (Sagua de Tanamo [Oriente] Cuba).
sparsa Maynard, Strophiops: 1924, Catalogue, Suppl., p. 3 (St. James
Corner, East Nassau, Bahamas).
stevensoni Dall, Cerion: 1900, Nautilus, 14, p. 65 (Long or Berry Island
[Long Island] Bahamas).
striata Schumacher, Pupa: 1817, Essai Nouveau Systeme Vers Testaces
Copenhagen, p. 230. Refers to Chemnitz 1780, Conchylien-Cabinet, (1) 4,
164 BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY
pi. 153, fig. 1439a-b. Is C. mumia Bruguiere, Pilsbry 1902.
striatella "Ferussac" Guerin-Meneville, Pupa: 1829?, Iconographie du
Regno Animal de G. Cuvier, Mollusques, p. 16, pi. 6, fig. 12 (The Antilles
[Puerto Rico] ).
striatissimum Aguayo and Jaume, Cerion salvatorl: 1953, Memorias, 21,
p. 274, pi. 31, fig. 8 (Playa de Santa Fe, Habana, Cuba).
strigis Aguayo and Sanchez Roig, Cerion herrerai: 1953, Memorias, 21,
p. 287, pi. 32, figs. 2, 4 (Cayo Brujas, Caibarien [Las Villas] Cuba).
stritella Humphrey, Pupa: 1797, Museum Calonnianum, p. 64 [see note
under clathrata Humphrey].
strobilus Beck, Pupa: 1837, Index Molluscorum, p. 82, [nomen nudum].
Atrophia Albers: 1850, Die Heliceen, Berlin, p. 202 [type species, Pupa
mumia Bruguiere, v. Maidens 1861, subsequent designation; non Strophia
Meigen 1825; Stal 1877].
Strophiops Dall: 1894, Bull. Mus. Comp. Zool., 25, p. 121 [type species,
Pupa dccumana Ferussac, original designation].
stroutii Maynard and Clapp, Strophiops : 1920, Records, App., 10. p. 120,
pl. 21, figs. 1-2 (Little Strout [Shroud]; Strout [Shroud]; East and West
Hawksbill and Cistern Keys [Exuma Group] Bahamas).
stupida Maynard and Clapp, Strophiops: Records, App., 10, p. 135, pl. 33,
figs. 1-2 (North Key, Staniard Creek, Andros, Bahamas).
subcostulatum Aguayo and Sanchez Roig, Cerion herrerai : 1953, Memorias,
21„ p. 287, pl. 32, fig. 3 (northern part of Cayo Santa Maria, Caibarien
[Las Villas] Cuba).
subcylindrica Beck, Pupa uva: 1837, Index Molluscorum, p. 82, [see note
under bidens Beck].
sublaevigatum "Pfeiffer" Pilsbry and Vanatta, Cerion (Maynardia) :
1895, Proc. ANSI', p. 209 (Matanzas, Cuba).
submarmoratum Pilsbry and Vanatta, Cerion: 1897, Proc. ANSP, 49. p.
365, text tigs. 3-4 ([Fortune Id.] Bahamas). Is C. fordii P. and V., Clench
1938.
sueyrasi "Blanes" Pilsbry and Vanatta, Cerion: 1898 [1899], Proc.
ANSP, p. 477, text fig. 6 (Vita, Cuba).
sula Maynard and Clap]), Strophiops: 1915, Records, App., 6. p. 180 [new
name for obliterafa Maynard 1913, not obliterata Maynard 1896J.
sulcata "Lamarck" Sowerby, Pupa: 1834, The Genera of Recent and Fos-
sil Shells, pt. 41, figs. 3-4 (no locality). Is C. mumia Bruguiere, Pilsbry 1902.
swift ii Pilsbry and Vanatta, Cerion regina: 1895, Proc. ANSP, p. 208
(Turks Island [Bahamas]). Is C. rer/ina P. and V., here considered a
synonym.
sylvatiea Maynard and Clapp, Strophiops : 1921, Records, App., 10, p. 137,
pl. 34, fig. 7 8 (Chub Point Key, Berry Islands, Bahamas).
tabida Maynard, Strophiops: 1913, Records, App., 5. p. 199 (near Cur-
CLENCH : CERIONIDAE 165
rent Settlement, Eleuthera, Bahamas). Is C. hyattii Mayn., Clench 1952.
tanamensis Sanchez Roig, Cerion: 1951, Revista, 7, p. 120, pi. 19, fig. 1
(Punta de Piedra, Yaguaeque, Sagua de Tanamo, Cuba).
lantUlum Aguayo and Jaume, Cerion gundiachi : 1951, Revista, 8, p. 5, pi.
2, fig. 11 (Cayo Guillermo, north of Punta Alegre, Camagiiey, Cuba).
tejedori Sanchez Roig, Cerion: 1951, Revista, 7, p. 112, pi. 18, fig. 7
(Punta Arenas, Paso de las Carabelas, Peninsula de Sabinal, Camagiiey,
Cuba). Is C. sanzi P. and V. See notes, this report.
ttviuco.stata Maynard and Clapp, Strophiops : 1920, Records, App., 10, p.
121, pi. 4, figs. 3, 9 (Sampson's Cay [Stanyard Cay, 3y2 miles XW of
Great Guana Cay, Exuma Group] Bahamas).
tenui Maynard and Clapp, Strophiops : 1915, Records, App., 6. p. 182
(east end of New Providence, Bahamas).
terwicallwm Aguayo and Sanchez Roig, Cerion circumscriptum: 1953,
Memorias, 21, p. 288, pi. 32, fig. 17 (Cayo Frances, Caibarien, Las Villas,
Cuba). Is C. sanzi P. and V. See notes, this report.
tenuilaoris "Gundlach" Pfeiffer, Pupa: 1870, Malakozoologische Blatter,
17, p. 91 (Barigua, Mata [Baraeoa] Cuba).
trrrita Maynard and Clapp, Strophiops: 1921, Records, App., 10, p. 147,
pi. 15, fig. 2; pi. 43, figs. 3-4 (Methodist Churchyard, Nassau, New Provi-
dence, Bahamas).
thaycri Maynard and Clapp, Strophiops: 1921, Records, App., 10, p. 137,
pi. 34, figs. 5-6 (east end of Thompson's Key, Berry Islands, Bahamas).
thompsoni Maynard and Clapp, Strophiops : 1915, Records, App., 6, p. 179
(south shore of Hog Island, New Providence, Bahamas).
thorndikei Maynard, Strophia: 1894, Contributions, 2, p. 116, fig. 34
(Cemetery between Waterloo and Nassau, New Providence, Bahamas). Is
C. varium Bonnet, Pilsbry 1902.
tibida Maynard, Strophiops : 1921, Records, App., 10, p. 152 [error for
tabida Maynard].
torrei "Blanes" Pilsbry and Vanatta, Cerion: 1898 [1899] Proc. ANSP,
p. 476, text figs. 1-2 (Port of Vita, Cuba).
tortuga Pilsbry and Vanatta, Cerion: 1928, Proc. ANSP, 80, p. 476, pi.
27, figs. 15-17 (Tortuga Island, Haiti).
tracta Maynard, Strophia cinerea: 1894, Contributions, 2, p. 123, fig. 37
(eastern point of Hog Island, New Providence, Bahamas). Is C. varium
Bonnet, Pilsbry 1902.
transitoria Maynard, Strophiops : 1913, Records, App., 5, p. 194 (northern
portion of Great Pimlico Island [Eleuthera] Bahamas). Is C. uniforme
Mayn., Clench 1952.
transmutata Maynard and Clapp, Strophiops: 1921, Records, App., 10,
p. 127, pi. 26, figs. 1-2 (Muddy Point Key, Great Exuma, Bahamas).
travelii Maynard and Clapp, Strophiops: 1921, Records, App., 10, p. 135,
pi. 32, figs. 3-4 (Bridgewater Key, Berry Islands, Bahamas).
166 BULLETIN : MUSEUM OV COMPARATIVE ZOOLOGY
tridentatum Pilsbry and Vanatta, Cerion (Paracerion) : 1895, Proc. ANSP,
p. 206; ibid, 1896, p. 336, pi. 11, fig. 27 (Cuba [Eincon de Guanabo, Ha-
hana, Cuba] ).
Tridcntistrophki Maynard : 1896, Contributions, 3. p. 9 [type species,
Strophia striatella Ferussac, original designation]. Is a synonym of Para-
cerion Pilsbry and Vanatta.
tumida Sowerby, Pupa: 1876, Conehologiea Iconica, 20, pi. 1, fig. 6 (Cuba)
[a MS name changed to incrassata].
tumidula Deshayes, Pupa: 1851, Deshayes [in] Ferussac, Histoire Nat-
urelle Generale et Particuliere des Mollusques, 2. pt. 2, p. 207 (Cuba) [is
C. mumia Bruguiere].
turgidum Torre and Welch, Cerion ramsdeni: 1934, Nautilus, 47, p. 106,
pi. 11, fig. 3a-d (hill west of Toro Eiver, 1 km. from beach or "Ojo del
Toro" west of Ensenada de Mora, Oriente, Cuba).
turnerae Clench and Aguayo, Cerion (JJmbonis) : 1952, Occ. Papers on
Mollusks, 1. no. 17, p. 423, pi. 53, figs. 4-7 (Lydia Point, Great Inagua, Ba-
hama Islands).
typica "Pfeiffer" Pilsbry, Cerion maritimum: 1902, M. of C, (2) 14, p.
213 [nomen nudum]. This was not intended to be a name introduced by
Pfeiffer but only a descriptive term.
ultima Maynard, Strophiops: 1913, Records, App., 5, p. 190 (Southwest
Key, New Providence, Bahamas).
Umbonis Maynard: 1896, Contributions, 3, p. 28 [type species, Strophia
scalarina Pfeiffer and Gundlach, monotypic].
uniformis Maynard, Strophiops: 1913, Records, App., 5, p. 194 (Little
Pimlico Island [Eleuthera] Bahamas).
unirersa Maynard, Strophiops: 1913, Records, App., 5, p. 196, (fossil,
Green Key [Rose Island, New Providence] Bahamas). Is C. uniforme Mayn.,
Clench 1952.
utowana Clench, Cerion {Strophiops) : 1933, Proc. New England Zool.
Club, 13. p. 92, pi. 1, figs. 1-2 (East Plana Key, Bahamas).
utrioulus Menke, Pupa: 1829, Verzeichnis Conchy. Samml. Malsburg, Pyr-
mont, p. 8 (locality not given).
uva Linne, Turbo: 1758, Systema Naturae, ed. 10, p. 765, (locality un-
known [Curasao]). [Refers to Gualtieri 1742, Index Testarum, pi. 58,
fig. D].
vaccinum Pilsbry, Cerion incanum: 1902, M. of C, (2) 14, p. 215, pi. 29,
fig. 51 (east end of Key Vaccas [Vaea] Florida).
vagabunda Maynard and Clapp, Strophiops: 1925, Records, App., 10, pi.
41, figs. 7-8 (southern end of Rose Island, New Providence, Bahamas). [New
name for albata Maynard and Clapp July 1921, not albata Maynard and
Clapp May 1921].
CLENCH: CERIONIDAE 167
valdesi de la Torre, Cerion c< iba: 1954, Revista, 9. p. 43, pi. 5, fig. 5 (Abra
Ventura, east of Canasi, Matanzas, Cuba).
valida Maynard and Clapp, Strophiops: 1920, Records, App., 10. p. 124,
pi. 22, figs. 1-2 (near well on west coast of Ship Channel Key [Exuma
Group] Bahamas).
valida Pilsbry and Vanatta, Cerion (Maynardia) columna: 189."), Proc.
ANSP, 47, p. 207 (Inagua [Bahamas]).
vallei Aguayo and Jaume, Cerion vulneratum: 1951, Revista, 8, p. 2, pi. 2,
fig. 7 (north coast of Cayo Puerco, bay of Puerto Padre, Oriente, Cuba).
vanattai Clench and Aguayo, Cerion: 1951, Revista, 8, p. 78. pi. 11, fig. 12
(I'laya Larga, Boca de Jauco, Baracoa, Oriente, Cuba).
vannostrandi Pilsbry and Vanatta, Cerion ritohiei: 1896, Proc. ANSP, p.
323 (locality unknown).
varius Bonnet, Pupa: 1846, Revue et Magasin de Zoologie (2) 16, p. 71,
pi. 6, figs. 3-4 (Tasmania [New Providence, Bahamas] ).
rariahile Dall, Cerion {Strophiops) : 1905, Smithsonian Misc. Collect., 47.
p. 440, pi. 58, fig. 6 (Red Bay, NW end of Andros Island, Bahamas).
varia-nivia Maynard, Strophiops: 1913, Records, App., 5. p. 186 (Eastern
Cemetery, Sherley St. to St. Paul quarry, Nassau, New Providence, Ba-
hamas).
varia-piirpuiui Maynard, Strophiops: 1913, Records, App., 5. p. 188 (Bay
St., east to Creek Settlement, Nassau, New Providence, Bahamas).
varia-thorndikei Maynard, Strophiops: 1913, Records, App., 5, p. 186
(Cemetery east of Nassau, New Providence, Bahamas).
variata Maynard and Clapp, Stropliiops : 1921, Records, App., 10. p. 134,
pi. 32, figs. 1-2 (Crab Key, Berry Islands, Bahamas).
varicgata Pfeiffer, Pupa incana: 1868, Monographia Heliceorum Viven-
tium, 6. p. 289 [based upon W. G. Binney 1859, Terrestrial Air-Breathing
Mollusks of the United States, 4. pi. 70 [79], fig. 17 (Florida)]. Is C. in-
canum Binney, Pilsbry 1902.
variegata Kiister, Pupa rubicunda: 1844, Conchylien-Cabinet, (2) 1. pt.
15, p. 76 (West Indies [Great Inagua, Bahamas] ).
ventricosior Beck, Pupa uva : 1837, Index Molluscorum, p. 82 [see note
under bidens Beck].
venusta Poey, Pupa: 1858, Memorias sobre la Historia Natural de la Isla
de Cuba, 2, p. 30 (Cuba).
vermiculum Dall, Cerion oweni: 1905, Smithsonian Misc. Collect., 47, p.
443, pi. 58, fig. 3 (Mathews Point, south side of Great Abaco, Bahamas).
Is C. bendalli P. and V., Clench 1938a.
veta Maynard and Clapp, Strophiops: 1920, Records, App., 10. p. 120,
pi. 21, figs. 3-4 (fossil, Strout's [Stroud] Key, [Exuma Group] Bahamas).
vetusta Maynard, Strophiops: 1913, Records, App., 5. p. 191 (fossil, Silver
Keys of Nassau bar, Nassau, New Providence, and Pimlico Keys, Eleuthera,
Bahamas). Is C. inconsuetum Mayn., Clench 1952.
168 BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY
vetusta-praedevina Maynard, Strophiops: 1913, Records, App., 5, p. 195
(fossil, Great Pimlico Key, [Eleuthera] Bahamas). Is C. inconsuetvm
Mayn., Clench 1952.
viaregis Bartsch, Cerion: 1920, Carnegie Institution of Washington, 14.
no. 282, p. 13, pi. 5; figs. 7-31 (King's Road, Bastian Point, northeast side
of South Bight, Andros Island, Bahamas).
victor de la Torre, Cerion: 1929, Nautilus, 42. pi. 4, figs. 12-13 [no de-
scription] (Caleta de Ovando, Oriente, Cuba).
viola Maynard, Strophia: 1890, Contributions, 1, pi. 16, fig. 5a-b [no de-
scription] (no locality given but Inagua, Bahamas on original label).
rulgar.e Roding, Cerion: 1798, Museum Boltenianum, (2) p. 90, refers to
Knorr, 6. pi. 25, fig. 4 (no locality). [Is C. uva Linne.]
vulnerata Kiister, Pupa: 1855, Conchylien-Cabinet, (2) 1. pt. 15, p. 161,
pi. 19, figs. 16-18 (locality unknown [Oriente, Cuba]).
watlingense Dall, Cerion (Strophiops): 1907, Smithsonian Misc. Collect.,
47, p. 438, pi. 58, fig. 7 (Watling Island, Bahamas).
weinlandi "Kurr." v. Martens, Pupa: 1860, Malakozoologische Blatter,
6. p. 207, pi. 2, fig. 1, (Crooked Island, Bahamas).
wrighti Aguayo and Sanchez Roig, Cerion mumia: 1953, Memorias, 21, p.
284, pi. 32, fig. 5 (Cuba; Charles Wright. [Northern coast of Pinar del Rio,
Cuba]).
ywmaensis Pilsbry and Yanatta, Cerian (Maynardia) : 1895, Proc. ANSP,
p. 210 (Yuma River, Haiti [Santo Domingo]).
zebra "Weinland" Sowerby, Pupa: 1875, Conchologia Ieonica, 20. Pupa
p. 12, fig. 12a-b (Bahamas).
REFERENCES
Clench, W. J.
1938. Origin of the Land and Freshwater Mollusk Fauna of the
Bahamas, With a List of the Species Occurring on Cat and
Little San Salvador Islands. Bull. Mus. Comp. Zool., 80: 481-
541, 3 plates.
Clench, W. J.
1938a. Land and Freshwater Mollusks of Grand Bahama and the
Abaco Islands, Bahama Islands. Mem. Soc. Cnbana Hist. Nat.,
12: 303-333, 2 plates.
Clench, W. J.
1942. Land Shells of the Bimini Islands, Bahama Islands. Proc. New
England Zool. Club, 19: 53-67.
Clench, W. J.
1952. Land and Freshwater Mollusks of Eleuthera Island, Bahama
Islands. Rev. Soc. Malacologiea "Carlos de la Torre," 8: 97-116,
3 plates.
CLENCH : CEBIONIDAE 1(i!)
Clench, W. .1. and 0. G. Aguayo
1952. The Sealarinum Species Complex (Umbonis) in the Genus
Cerion. Occasional Papers on Mollusks, Harvard University, 1:
413-440. 7 plates.
KuSTER, 11. C.
1841- Conehylien-Cabinet, 1. pt. 15: 1 96, 9 plates.
1847.
M \VX LED, C. J.
1889- Contributions to Science, Newtonville, .Mass. Vols. 1-3.
1896.
Mayxakd, C. J.
1913- Appendices to Records of Walks and Talks with Nature, Wesl
1926. Newton, Mass. Vols. 5. 6. and 10.
Maynabd, C. J.
1924. Catalogue of Specimens of the Family Cerionidae for Sale by
C. J. Maynard, West Newton, Mass., pp. 1-6.
PlLSBRY, II. A.
1901- Manual of Conchology, (2 I 14: 174-281',, 21 plates.
1902.
PlLSBRY, II. A.
1943. Xoie on Cerion striatellum ("Per." Guerin). Nautilus, 57:
34-35.
PlLSBRY, H. A.
1946. Land Mollusca of North America. Acadamy of Natural Sciences
Philadelphia, Monographs 3, 2, pt. 1: 158-169, 5 text figures.
Plate, L.
1907. Die Variabilitat und die Artbildung nach dem Prinzip geog-
raphiseher Fornienketten bei den Cerion-Land Schnecken der
Bahama-Inseln. Arehiv fiir Eassen- und Gesellschafts-Biologie,
4: 433-614, 5 plates.
Smith, E. A.
1898. On the Land-Shells of Curagoa and the Neighbouring Islands.
Proc. Malacological Soc. London, 3: 113-116, 2 text-figures.
Sowkkby, G. B.
1875- Conchologia Iconica, 20. Pupa, text and 3 plates.
1876.
Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE
Vol. 116, No. 3
STUDIES ON NEW ZEALAND ELASMOBRANCHII. PART
VI. TWO NEW SPECIES OF ETMOPTERUS FROM
NEW ZEALAND
By J. A. F. Garrick
Zoology Department, Victoria University College.
Wellington, New Zealaml
CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM
April, 1957
Publications Issued by or in Connection
with THE
MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE
Bulletin (octavo) 1863 - - The current volume is Vol. 115.
Breviora (octavo) 1952 — No. 73 is current.
Memoirs (quarto) 1864-1938 — Publication was terminated with Vol. 55.
Johnsonia (quarto) 1941 -- A publication of the Department of Mollusks.
Vol. 3, no. 35 is current.
Occasional Papers of the Department of Mollusks (octavo) 1945 -
Vol. 2, no. 21 is current.
Proceedings of the New England Zoological Club (octavo) 1899-
1948 — Published in connection with the Museum. Publication terminated
with Vol. 24.
The continuing publications are issued at irregular intervals in numbers
which may be purchased separately. Prices and lists may be obtained on
application to the Director of the Museum of Comparative Zoology,
Cambridge 38, Massachusetts.
Of the Peters "Check List of Birds of the World," volumes 1-3 are out
of print; volumes 4 and 6 may be obtained from the Harvard University
Press; volumes 5 and 7 are sold by the Museum, and future volumes will be
published under Museum auspices.
Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE
Vol. 116, No. 3
STUDIES ON NEW ZEALAND ELASMOBRANCHII. PART
VI. TWO NEW SPECIES OF ETMOPTERUS FROM
NEW ZEALAND
By J. A. F. Garrick
Zoology Department, Victoria University College,
Wellington, New Zealand
CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM
April, 1957
No. 3 — Studies on New Zealand Elasmobranchii. Part VI.
Two New Species of Etmopterus from New Zealand 1
By J. A. F. Garrick
Zoology Department, Victoria University College,
Wellington, New Zealand
Experimental line-fishing- off New Zealand, for the purpose of
adding to our knowledge of the deeper-water shark fauna of this
region, has resulted in the capture of three specimens of Etmop-
terus representing two species apparently new to science. These
specimens were caught off Kaikoura on the east coast of the
South Island, by Mr. Richard Baxter, who, fishing from a 16 foot
dinghy, collected one large brown specimen from 500 fathoms,
and a small grey-black specimen from 200 fathoms in November,
1955. In February, 1956, a further grey -black specimen was
caught in 100 fathoms. All captures were made close inshore, the
submarine topography of the Kaikoura region being such that
water 500 fathoms deep is found within 3 miles of the coast.
Although squaloid sharks are fairly well represented in the
New Zealand fauna, no specimens of Etmopterus have been
known, and geographically the nearest member of the genus is
E. molleri (Whitley) 1939 of southern Australia. It is therefore
of considerable interest that the two species of these luminescent
sharks now known to be present should represent what are more
or less the extremes of morphological diversity in the genus.
The large brown specimen, here proposed as Etmopterus baxteri
n.sp., in honour of Mr. Richard Baxter, is akin to E. princeps
of the North Atlantic in its size — which exceeds that of most
other species ; in the uniform but random arrangement of the
dermal denticles ; in the noticeably small and rounded pectoral
fins ; in the rather plain colouration and inconspicuous pelvic
flank mark ; and in the high number of cusps on the upper teeth.
The grej^-black specimens, named here as Etmopterus abemethyi
n.sp., for Mr. Fred Abernethy who has contributed greatly to
the collection of New Zealand elasmobranchs, are closely allied
to the Pacific species E. lucifer, E. brachyurus and E. molleri,
and like them are small ; with dermal denticles arranged linearly
1 This study has been assisted by a grant from the Research Grants Committee
of the Universitv of New Zealand.
172 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
on the sides and upper surface of the trunk; with large and
more angular pectoral fins; with an obvious colour pattern and
conspicuous flank marks; and with a smaller number of cusps
on the upper teeth.
Comparison of E. baxteri and E. abernethyi with other species
of the genus, of which twelve are listed by Bigelow, Schroeder
and Springer (1953, p. 238), though E. molleri (Whitley) 1939
should be added to these, has been greatly facilitated by the
availability of specimens of E. spinax, E. polli and E. princeps
provided by Drs. H. B. Bigelow and W. C. Schroeder, to whom
I am also especially indebted for their generosity in supplying
access to their manuscript key to the species.
Etmopterus baxteri n.sp.
Figures 1 and 2
Study Material. Holotype, mature female, 742 mm. total
length, Dominion Museum No. 1950, lined from 500 fathoms
seven miles south of Kaikoura, New Zealand, by Mr. R. Baxter,
in November, 1955.
Description. Proportional measurements in per cent of total
length:
Trunk at pectoral origin : breadth, 12.7 ; height, 10.8
Snout length in front of : outer nostrils, 1.7 ; mouth, 8.4.
Eye : horizontal diameter, 4.0 ; vertical diameter, 2.7.
Mouth : breadth, 8.4 ; height, 1.4.
Nostrils : breadth (between inner corners), 3.1.
Labial furrow lengths : upper, 3.5 ; lower, 1.7.
Gill-opening lengths: 1st., 2.7; 3rd., 1.7; 5th., 1.7.
First dorsal fin : vertical height, 3.1 ; length of base, 5.7.
Second dorsal fin: vertical height, 4.0; length of base, 8.1.
Caudal fin: upper margin, 18.4; lower anterior margin, 10.2.
Pectoral fin : anterior margin, 7.5 ; width, 6.1.
Pelvic fin: anterior margin, 5.7; distal margin, 6.2; posterior
margin, 2.2.
Distance from snout to : eye, 5.4 ; 1st gill-opening, 16.2 ; 5th
gill-opening, 20.7; 1st dorsal, 33.4; 2nd dorsal, 65.5; upper
caudal, 81.8 ; pectoral, 21.7 ; pelvic, 57.4.
Interspace between : 1st and 2nd dorsals, 26.0 ; 2nd dorsal and
GARRICK : NEW ZEALAND ELASMOBRANCHII 173
caudal, 10.1 ; pelvic and subcaudal, 14.3.
Distance from origin to origin of : pectoral and pelvic, 35.7 ;
pelvic and subcaudal, 22.2.
Head depressed, wide, compact, and very large-eyed; trunk
moderately stout, and compressed posterior to the pectorals.
Height of trunk at origin of pectorals 7.5 in the length from
snout tip to origin of subcaudal. Length of body measured to
the cloaca, 62 per cent of the total length. Caudal peduncle little
compressed and slender, and without lateral keels or precaudal
pits.
Dermal denticles small, numerous, and in the form of conical
thorns, slightly curved and directed posteriorly, and borne on
four-angled bases. Each denticle carries six ridges, four of
which are continuous with the ridges arising from the angles of
the base, while two are intermediate ridges on the anterior face
of the denticle and do not extend on to the base. Denticles from
the head and fins similar to those from the trunk. The denticles
are distributed uniformly but sparsely so that there are con-
siderable interspaces between them where the skin is visible.
Their arrangement is random, at least on the anterior two-thirds
of the trunk, though towards the caudal peduncle and on the
tail they are in more or less regular longitudinal rows. The
pectoral, pelvic, dorsal and caudal fins are denticle covered
almost to their margins, except for the ventral surface of the
pelvic which has a wide, naked zone distally, and the web of the
second dorsal on which the denticles are very sparse. Other naked
regions of the body include the ventral surface of the tip of the
snout ; the upper and lower lips ; the axil of the pectoral where
the naked area is large and ovoid in outline, extending along the
trunk well posterior to the fin when the latter is laid back, and
also continued on to the upper surface of the fin itself as a wide
band along the posterior margin ; the axil of the pelvic and the
entire upper surface of the base of this fin; the axil of the first
dorsal where the naked area is small ; and the axil of the second
dorsal where the naked area is very extensive, reaching from in
front of the origin of the fin to behind its posterior free tip. In
all cases, the naked regions correspond with the lighter coloured
areas on the trunk and fins. Within the dark area encompassed
by the pelvic flank mark, the denticles are noticeably smaller
174
BULLETIN" : MUSEUM OF COMPARATIVE ZOOLOGY
Figure 1. Etmopterus baxteri n.sp., holotype, 742 mm. total length. A,
lateral view and insets of sections through snout and peduncle; B, dorsal
view of head showing prominent lateral line pores; C, ventral view of
peduncle; B, ventral view of head with pores as in B ; E, left nostril;
F-E, upper teeth from right side, row numbers indicated above; I, lower
teeth, right side; J, 10th lower tooth, right side; K, 6th upper tooth, right
side, (c = level of cloaca.)
GARRICK : NEW ZEALAND ELASMOBRANCHII 175
than the adjacent ones, and also the tips of these denticles are
directed ventrally rather than posteriorly as are the majority of
the trunk denticles.
Head measured to first gill-opening 6.2 in the total length,
and just less than half the distance from snout tip to first
dorsal origin. Head noticeably broad, its greatest width at the
level of the first gill-opening where it is 1.6 times the least fleshy
interorbital width, the latter being equal to the preoral distance.
Width of the head at the level of the nostrils is only slightly
narrower than the interorbital width, so that the contours of the
head between these levels are almost parallel. The snout tip is
broadly rounded, and each nostril forms an abrupt step in the
contour. The snout is thick, slightly wedge-shaped in profile,
strongly depressed, and flat above as is the greater part of the
head to the level of the spiracles. Length of snout measured to
eye, 3.0 in the head. Eye very large, ovoid, 1.5 times as long as
high, its horizontal diameter 1.3 in the snout. Spiracle large, its
length 4.0 in the horizontal diameter of the eye, and placed just
above the level of the dorsal margin of the eye, and behind it by
a distance equal to about twice its own length. Gill-openings of
moderate size and slightly oblique; each gill-opening is deeply
emarginate, especially the first in which the tips of the gill-
filaments are visible. Lengths of the gill-openings decrease from
the first to the fourth, but with the fifth equal to the third.
Length of the first gill-opening 1.7 times that of the fourth, and
1.5 in the horizontal diameter of the eye. Interspaces between
the gill-openings decrease posteriorly, that between the first and
second almost twice that between the fourth and fifth. Nostrils
large, oblique, and well anterior on the venter of the snout. Each
nasal aperture subdivided into an anteriorly directed, circular,
anterolateral aperture and an ovoid posteromedial aperture by
triangular nasal flaps. The anterior nasal flap is large, pointed,
and external to the shorter, fleshy posterior flap. The postero-
medial aperture is also margined in front and behind by a low
membrane. Mouth broad, and only slightly arched, its width just
greater than the preoral distance, and 1.9 in the length of head.
The upper labial furrows moderately long, and deeply incised
anteriorly, their length 1.4 in the distance from their anterior
extremity to the symphysis of the upper jaw. The lower labial
176 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
furrows are shallowly incised and short, their length about half
that of the upper furrows.
Teeth 2g_27 , dissimilar in the two jaws. The upper teeth
erect, each with a long, sharply-pointed, awl-shaped, smooth-
edged major cusp flanked on each side by up to four small lesser
cusps, and borne on a longitudinally-striated bifid base. Most of
the upper teeth have three lesser cusps on each side of the major
cusp, with the middle cusp of these three considerably larger
than the others though not more than one-third the length of the
major cusp. A few teeth near the centre of the jaw have four
lesser cusps on each side, with the largest lesser cusp separated
from the major cusp by two small lesser cusps, while in the teeth
towards the angle of the jaw there is a reduction in the number
of lesser cusps to one or two on each side. Three series of upper
teeth functional at the centre of the jaw, two towards the angles.
The lower teeth blade-like, each with a smooth, little-sculptured,
rectangular base almost twice as high as broad, and bearing a
single, smooth-edged, triangular cusp. Each cusp is sharply
notched laterally, strongly oblique, and overlaps the adjacent
cusp so that an almost continuous cutting edge is formed. There
is no median tooth, and the base of the first tooth on the left
side overlaps that of the first tooth on the right. A single series
of lower teeth functional.
First dorsal small, short-based, and brush-shaped, its distance
from snout tip 33.4 per cent of the total length. Height of first
dorsal 1.9 in its base, and the latter 4.6 in the interspace be-
tween the first and second dorsals. Length of the posterior mar-
gin 1.5 in the length of the base; the posterior tip sharply
pointed. The first dorsal spine short and almost straight, its
length less than half the distance from its origin to the first
dorsal apex. Interspace between the first and second dorsals
equal to the distance from snout tip to the axil of the pectoral.
Second dorsal considerably larger than the first, and originating
above the posterior insertion of the pelvic base. Height of the
second dorsal 2.0 in its base, and the latter 3.2 in the interspace
between the dorsals. The second dorsal spine strongly curved
and long, its length 2.5 times that of the first dorsal spine. Inter-
space between second dorsal and caudal 3.0 in that between first
and second dorsals. Caudal measured from hypural origin 5.0 in
GABKICK : NEW ZEALAND ELASMOBRANCHII
177
the total length. Height of the epiural 4.6 in its length, and its
margin slightly sinuous. The terminal lobe with a convex margin.
Height of the hypural 1.7 times that of the epiural, and its lower
anterior margin almost straight. The apex acutely angled, and
the posterior margin deeply concave. The pectorals short and
wide, their length 2.0 in the head measured to the first gill-open-
ing, and their width 1.4 in their length. The anterior margin
almost straight, the posterior margin slightly convex and contin-
ued without a distinct angle into the distal margin. Pelvics
originating anterior to the second dorsal origin by a distance
equal to the length of the pectoral. Length of pelvic base equal
to that of second dorsal base ; posterior margin short, its length
2.5 in the horizontal diameter of the eye. The posterior tip of
pelvic pointed, and terminating just anterior to the origin of the
second dorsal spine.
I-Or
del J ATC
Figure 2. FA Diopter as baxteri n.sp., holotype, 742 mm. total length. A,
external view of dermal denticles from high on side at level of 1st dorsal ;
B, lateral view.
Colour. The overall colour of the specimen is an almost uni-
form, medium dusky brown, slightly darker on the ventral sur-
face and on the fins, but considerably lighter in the regions which
are smooth and free of denticles, such as the axils of the fins, and
the lips. There is also a vertical white band devoid of pigment
on the anterior surface of the outer part of each gill-arch, though
this is visible only on the first arch where the anterior edge of
the first gill-opening is strongly emarginate. Despite the ap-
178 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
parent uniformity of colour of the specimen, close examination
reveals the presence of well-defined, darker regions which con-
tribute to a pattern similar to that described for other etmop-
terids. These darker regions are characterised in this speci-
men chiefly by the presence of numerous small, black pits,
rather than by an increase in the number of typical chromato-
phores. The black pits are not, however, confined to the
darker regions, but are distributed more sparsely over the en-
tire head, trunk and fins. The most prominent dark region is
a longitudinal flank mark above the pelvic fin, and of the shape
shown in Figure 1A. As described above, within the area en-
compassed by this flank mark, the dermal denticles are smaller
and directed more ventrally than those outside it, so that they
also contribute to its definition. The pelvic flank marks are con-
nected on the ventral surface of the caudal peduncle where they
form a pattern as in Figure 1C. The ventral surface of the ab-
domen is also a distinct dark region, well delineated on the flanks
by a denser concentration of black pits along its edges than
elsewhere on the ventral surface. Anteriorly it is continued
under the head and snout, but apart from an ill-defined trans-
verse band under the head, its extent is not clear. Other dark
marks present are a narrow, curved band on the lower surface
of the pectoral fin, extending from the origin of the fin to its
insertion; a short streak on the anterior margin of each dorsal
fin, close to the tip of each dorsal spine ; and a prominent, dark
line near the tip of the caudal axis, parallel to and just below
the terminal portion of the lateral line where the latter is in the
form of a naked groove. The last-mentioned mark is contributed
to not only by black pits, but also by a very thin black streak
along each edgo. of the lateral line groove. A very few similar
black streaks are also present sporadically on the sides of the
trunk, where they are short and appear to be derived from the
fusion of contiguous black pits. The lining of the mouth is a
light dusky brown, as is the lining of the body cavity.
Luminescence. The black pits which contribute most of the
colour pattern to the specimen, and the few black streaks which
are present, appear to be identical with those of E. spinax in
which thev are known to be luminescent. However, Mr. Baxter
did not notice any luminescence on the specimen when it was
GARRICK : NEW ZEALAND ELASMOBRANCHII 179
first taken from the water, though this was during daytime when
such luminescence might not be obvious.
Maturity. The adult condition of the specimen is evidenced
by the extrusion of two embryos during its transport from Kai-
koura to Wellington. The embryos are two inches long, devoid
of pigment, and only part way through development. They were
attached to large yolk-sacs, though these were ruptured and
could not be measured. At least two other intact yolk-sacs can
be felt within the animal, and possibly others may be present
but ruptured.
Discussion. The thorn-like denticles of E. baxteri readily dis-
tinguish it from those species of Etmopterus with truncate denti-
cles, i.e. frontimaculatus, pusillus and granulosus, as they do
also from paessleri which is described as having denticles with
a larger central spine surrounded by several lesser spines. Of
the remaining etmopterids, all of which have denticles with a
single spine, four more are separable from baxteri on denticle
characters — namely, virens, in which the denticles are thorn-
like but very short and low, and spinax, hillianus and schultzi,
which have elongate, bristle-like denticles; though as the dif-
ferences between these species and baxteri in this respect are less
distinctive than those between baxteri and the species with
truncate or multispinose denticles mentioned above, it is perhaps
better not to rely on them alone as primary specific characters.
Compared with virens, baxteri is heavy-bodied and short-
tailed (the distance from pelvic origin to tip of caudal 42.6 per
cent of the total length in baxteri, 53 per cent in virens). From
villosus, baxteri differs in the very much shorter predorsal length
(equal to less than the distance from origin to origin of the first
and second dorsals in baxteri, but reaching from first dorsal
origin to almost the upper caudal origin in villosus). The two
noticeably short-tailed Pacific species, brachyurus from the
Philippines and molleri from southern Australia, differ from
baxteri not only in their short-tailedness (the length of the upper
caudal margin two-thirds and three-fourths of the distance from
the rear ends of the pelvic bases to the lower caudal origin in
brachyurus and molleri respectively, but more than one and a
third times in baxteri) but also in the linear arrangement of
the denticles on the sides of their trunks; their proportionately
longer pectoral fins (reaching to the first dorsal when laid back
1 80 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
in brachyurus and molleri, but falling well short of this level
in baxteri) ; and their more attenuate pelvic flank marks. The
same differences apply between lucifer and baxteri, though in
lucifer the length of the upper caudal margin is proportionately
longer than in brachyurus or molleri but still considerably
shorter than in baxteri.
Of the etmopterids with bristle-like denticles, i.e. schultzi, hil-
lianus and spinax, schultzi is clearly distinct from baxteri not
only in its very elongate caudal fin (the upper margin of the
caudal almost equal to the distance from snout tip to the tip
of the pectoral when the latter is laid back in schultzi, but
reaching only to the second gill-opening in baxteri) but also in
its peculiar frayed and fringe-like fin margins which differ
from those of all other etmopterids. E. hillianus has a greater
peduncular length than baxteri (the distance from the rear ends
of the pelvic bases to the lower caudal origin as long as the
distance from snout tip to pectoral origin in hillianus, but reach-
ing only midway between spiracle and first gill-opening in
baxteri) ; while spinax is readily separable from baxteri by its
much narrower head (head width equal to the preoral distance
in spinax, but more than one and a half times this distance in
baxteri) • its shorter and less concave gill-openings ; and the shape
of the dark markings on the ventral surface of the trunk and
peduncle.
The remaining etmopterids to be compared with baxteri, i.e.
polli and princeps, both agree with this species in being more or
less plain and dark coloured, and in having thorn-like denticles
which are in random but uniform arrangement. E. polli, how-
ever, is distinctive in having a short interdorsal space (reaching
much less than the distance from snout tip to first gill-opening
in polli, but extending to the axil of the pectoral in baxteri) and
a longer pectoral which reaches behind the base of the first
dorsal spine when laid back. In baxteri and princeps the pec-
torals are short, their tips failing to reach the first dorsal origin,
while further agreement between these species is seen in the
noticeable broadness of the head, the long and concave gill-
openings (which expose the lamellae of the first gill-arches), and
the vertical white markings on the outer part of the gill-arches.
E. baxteri differs from princeps in having a shorter tail (the
length from pelvic origin to tip of caudal equal to distance from
GARRK'K : NEW ZEALAND ELASMOBRANCITII 181
snout tip to posterior tip of the first dorsal fin in baxteri, but
reaching to midway between first dorsal tip and pelvic origin in
princeps) ; a shorter caudal fin (the upper caudal margin reach-
ing from snout to second gill-opening in baxteri but from snout
tip to pectoral origin in princeps) ; a more conspicuous and dif-
ferently shaped pelvic flank mark; less oblique nostrils; and
upper teeth mostly with 7 or 9 cusps rather than the 5 cusps in
princeps. Moreover the arrangement of the lesser cusps of the
upper teeth in baxteri, where a very small lesser cusp is sand-
wiched between a larger lesser cusp and the major cusp, differs
from that in not only princeps but also all other etmopterids
where the lesser cusps usually diminish uniformly in size from
the major cusp outwards.
Etmopterus abernethyi n. sp.
Figures 3 and 4
Study Material. Holotype, immature male, 338 mm. total
length, Dominion Museum Xo. 1951 ; and paratype, female, 278
mm. total length, Mus. Comp. Zool. Xo. 39714; both lined by Mr.
Richard Baxter from 7 miles south of Kaikoura, the holotype
from 200 fathoms in November, 1955, the paratype from 100
fathoms in February, 1956.
Description. Proportional measurements in per cent of total
length : Holotype and paratype.
Trunk at pectoral origin : breadth, 10.4-11.1 ; height, 8.6-8.6.
Snout length in front of : outer nostrils, 2.7-2.5 ; mouth, 10.9-
11.5.
Eye : horizontal diameter, 4.6-5.0; vertical diameter, 2.7-2.9.
Mouth : breadth, 5.9-6.1 ; height, 1.2-1.1.
Nostrils : breadth ( between inner corners), 3.0-3.2
Labial furrow Lengths : upper, 3.3-3.2 ; lower, 1.5-1.4.
Gill-opening lengths: 1st, 1.3-1.3; 5th, 1.0-1.1.
First dorsal fin : vertical height, 3.3-4.0 ; length of base, 5.6-6.5.
Second dorsal fin; vertical height. 5.6-6.1; length of base.
8.0-8.6.
Caudal fin: upper margin, 22.5-23 0; lower anterior margin.
10.9-10.4.
Pectoral fin : anterior margin, 9. 5-!). 7 ; width, 8.3-9.3
182 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Pelvic fin : anterior margin, 7.1-6.8 ; distal margin, 6.8-9.0.
Distance from snout to : eye 6.5-7.2 ; 1st gill-opening, 18.7-19.0 ;
5th gill-opening, 21.9-23.0; 1st dorsal, 29.6-32.0; 2nd dorsal.
57.0-57.9; upper caudal, 77.5-77.0; pectoral, 22.2-23.4; pel-
vic, 48.2-49.4.
Interspace between: 1st and 2nd dorsals, 21.6-19.8; 2nd dorsal
and caudal, 12.7-11.9.
Distance from origin to origin of : pectoral and pelvic, 26.3-
27.1 ; pelvic and subcaudal 27.1-26.3.
Head depressed, long, and very large-eyed ; trunk moderately
slender, and compressed posterior to the pectorals. Height of
trunk at origin of pectorals 8.7 in the length from snout tip to
origin of subcaudal. Length of body measured to the cloaca, 53
per cent of the total length. Caudal peduncle little compressed
and slender, and without lateral keels or precaudal pits.
Dermal denticles small, slender and thorn-like, borne on four-
angled bases and with their tips directed slightly posteriorly.
Each denticle is six-ridged, as in E. baxteri, though the ridges
are less steep and fail to extend to the tip of the denticle. The
denticles are numerous, and well-spaced ; arranged in random on
the ventral surface of the head and trunk, but in distinct parallel
longitudinal rows on the sides and upper surface, the fins, and
the venter of the peduncle. Above the lateral line, the rows are
oblique, sloping posterodorsally on the head and in front of the
first dorsal fin, but with their slope flattening out and reversing
behind the latter level. Below the lateral line, the rows are hori-
zontal. The line of demarcation between the linear arrangement
of the denticles on the sides and the random arrangement below,
is sharp, and parallels that of the dark colour pattern; it skirts
the lateral margin of the snout, follows round the lower edge of
the eye, runs beneath the gill-openings, and is especially prom-
inent from the axil of the pectoral to the origin of the pelvic.
Within the lateral pelvic flank mark, the denticles are more
sparsely distributed, are noticeably smaller, and have their tips
directed ventrally rather than posteriorly. The distal parts of
the webs of the dorsal, pectoral and pelvic fins are naked, as are
the terminal and hypural lobes of the caudal fin. Other naked
areas include the upper and lower lips; the interspaces between
the gill-openings ; and the axils of the pectoral, pelvic and dorsal
fins (though none of the latter is as extensive as in E. baxteri).
GARRICK : NEW ZEALAND ELAKMOBRANCIIII
183
•Ji'"*-1-,/
184 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Head measured to first gill-opening 5.3 in the total length, and
about two-thirds of the distance from snout tip to first dorsal
origin. Head long, flat above, and with little change in width
from the level of the spiracles to the level of the nostrils. Inter-
orbital width 1.4 in the preoral distance. The snout contours
rapidly taper at the level of the nostrils so that the snout tip is
prominently pointed. In lateral view the snout is of almost equal
thickness from the hind level of the eyes to the nostrils, though
anterior to the latter the profile angles steepen to form the
bluntly wedge-shaped snout tip. Length of snout, measured to
eye, 2.9 in the head. Eye large, ovoid, nearly twice as long as
high ; its horizontal diameter 1.4 in the snout. Spiracle large, its
length just less than one quarter of the horizontal diameter of the
eye, and placed just above the eye and behind it by a distance
equal to one and a half times its own length. Gill-openings small,
each vertical but concave, their lengths subequal and about 4.0
in the eye. Interspaces between the gill-openings decrease slightly
posteriorly. Nostrils large, oblique, and well anterior on the
venter of the snout. Each nasal aperture subdivided by tri-
angular nasal flaps into a circular, anterolateral aperture facing
to the anterior, and an ovoid posteromedial aperture which is
margined in front and behind by a low membrane. The anterior
nasal flap is attenuate, sharply pointed, and external to the short
fleshy posterior flap. Mouth broad and little arched, its width
1.5 in the preoral distance, the latter 1.7 in the head. The upper
labial furrows deeply incised anteriorly, their length equal to the
distance from their anterior extremities to the symphysis of the
upper jaw, and arranged so that one-third is anterior to the
angle of the jaw and two-thirds is posterior. The lower labial
furrows are shallowly incised and short, their length less than
half that of the upper furrows.
Teeth 1g_^7 in the male of 338 mm., -fipnr ^n ^ne fema-le °f
278 mm., dissimilar in tbe two jaws. The upper teeth erect, multi-
cusped, each with a long, sharply-pointed, awl-shaped, smooth-
edged major cusp flanked on each side by one or two lesser cusps,
and borne on a longitudinally-striated base. Most of the upper
teeth have two lesser cusps on each side of the major cusp, with
the outer cusp of these two very much smaller than the inner
which is one-third to one-half of the length of the major cusp.
GABRICK : NEW ZEALAND ELASMOBRANCHIT 185
The teeth towards the angle of the jaw have only one or no lesser
cusps on each side. Three series of upper teeth functional at the
centre of the jaw, two towards the angles. The lower teeth blade-
like, each with a smooth-faced, subrectangular, laterally-rounded
base, bearing a single, smooth-edged triangular cusp. Each cusp
is sharply notched laterally, very strongly oblique, and overlaps
the adjacent cusp so that an almost continuous cutting edge is
formed. There is no median tooth, and the base of the first tooth
on the left side overlaps that of the first tooth on the right. A
single series of lower teeth functional.
First dorsal small, brush-shaped, originating just anterior to
the tip of the pectoral when the latter is laid back. Distance from
snout tip to first dorsal origin 29.6 per cent to 32.0 per cent of the
total length. Height of first dorsal 1.7 in its base, and the latter
3.8 in the interspace between the first and second dorsals. Length
of the posterior margin 1.3 in the length of the base, and the
posterior tip pointed. The first dorsal spine short and almost
straight, its length less than half the distance from its origin to the
apex of the fin. Interspace between the dorsals equal to or less than
the distance from snout tip to pectoral origin. Second dorsal much
larger than the first, originating just posterior to the rear inser-
tion of the pelvic base. Height of the second dorsal 1.5 in its
base, and the latter 2.7 in the interspace between the dorsals. The
second dorsal spine curved and long, reaching two-thirds of the
distance from its origin to the apex. Interspace between second
dorsal and caudal 1.8 in that between first and second dorsals.
Caudal measured from hypural origin 4.0 in the total length.
Height of the epiural 6.5 in its length, and its margin straight
along most of its length but convex distally. The terminal lobe
with a convex margin. Hypural originates well anterior to the
epiural, its height 1.6 times that of the latter. Anterior margin
of hypural straight, the apex right-angled but rounded, and the
posterior margin concave. Pectorals noticeably wide, their width
1.2 in their length, and the latter 2.3 in the head. Anterior and
posterior margins convex, distal margin straight, and the pos-
terior angle smoothly rounded. Pelvics originating well anterior
to the second dorsal, the interspace between first dorsal tip and
pelvic origin equal to the length of the pelvic base. Anterior
and distal margin straight, and the apex prominent but rounded.
186 BULLETIN" : MUSEUM OP COMPARATIVE ZOOLOGY
The posterior tip sharply pointed and terminating at the level
of the second dorsal spine. Claspers on holotype cylindrical in
section, tapering to a point posteriorly, and showing no sign of
the external features which might be expected in a mature ani-
mal.
Colour. Dusky dark brown above, black below, though a heavy
coating of mucus gives the specimens a greyish cast. Dorsal,
pectoral and pelvic fins pale and translucent, as is the lower
posterior margin of the caudal. A large, ovoidal pale area covers
the greater part of the top of the head, and posteriorly is con-
tinued as a wide band along the mid-dorsum of the trunk and
peduncle, though it is interrupted at the dorsal fins, the bases of
which are dusky brown. There is also a pale supraorbital streak
on each side of the head ; a narrow indistinct pale band along
the lateral line ; and an elongate pale mark above and anterior to
the pelvic origin. In the female of 278 mm., the pale markings
are more extensive than in the holotype ; the mid-dorsal band
extending well down the sides of the peduncle, and the epiural
lobe as well as the hypural lobe is pale, though the terminal lobe
and the apex of the hypural are darker than elsewhere on the
caudal axis as is also the case in the holotype.
Microscopic examination shows that the darkness of the ventral
surface is due not only to a greater number of chromatophores
compared with the condition on the sides and upper surface, but
also to the presence of numerous small black pits, as in E. baxteri.
The dark regions thus characterised include the undersurface of
the snout, head and trunk (the demarcation line between the
dark region below and the lighter region above well delineated by
a greater concentration of black pits than elsewhere on the under-
surface— see Figure 4C) ; the venter of the peduncle where a
pattern is present as in Figure 3C, though this pattern is not
developed as clearly in the female of 278 mm. ; an attenuate pelvic
flank mark of the shape shown in Figure 3A ; a long, narrow
caudal streak parallel to and below the naked, grooved portion
of the lateral line ; a large curved streak on the underside of
the pectoral, and a short arc on its upper surface ; and a short
streak on the upper surface of the pelvic base. There is also a
prominent row of black pits and streaks along the mid-dorsal
line, while others are scattered over most of the head and the
OARRIOK : NEW ZEALAND ELASMOBRANCIIII
187
trunk where their concentration and arrangement is similar to
that known in E. lucifcr. Lining of mouth dusky grey; lining
of body cavity black.
Luminescence. It is not known if E. abernethyi is luminescent,
for although the black pits and streaks present resemble closely
those of luminescent species of Etmopterus, no luminescence was
observed in the specimens when they were caught.
m
*«**
»-w<"« -* * *. J»*
* * * » .» » * » « » . * ■» «*
dei.JA.FG.
05 mn
20n
Figs. A-B
Figure 4. Etmopterus abernethyi n.sp., holotype, 338 mm. total length.
A, external view of denticles from high on side at level of 1st dorsal; B,
lateral view; C, external view of skin from lower part of side of trunk,
showing three rows of denticles arranged linearly, and others below
arranged at random. Note greater concentration of chromatophores in lower
half, and black pits which are most numerous at demarcation line between
light and dark regions.
Maturity. The claspers on the holotype lack the external spurs
which might be expected in a mature specimen; the female of
278 mm. has not been examined for its state of maturity though
its small size in comparison with the immature male suggests that
it, too, is immature.
Discussion. As in E. baxteri, the thorn-like denticles of E.
abernethyi provide a ready character for the separation of this
species from front imaculat us, pusillus and granulosus which have
truncate denticles, and from paessleri in which the denticles are
188 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
multispinose. The slender thorns of abernethyi are also obviously
distinct from the short, low denticles of virens, though in other
respects including the general proportions of the body, abernethyi
is strikingly similar to this species. It differs from virens in the
relatively shorter peduncular length (the distance from the rear
ends of the pelvic bases to the lower caudal origin equal to the
distance from snout tip to first gill-opening in abernethyi. but
reaching to the pectoral origin in virens) ; in the lack of the
transverse pale markings on the abdomen ; in the presence of the
conspicuous mid-dorsal pale band ; and in the shape of the dark
pelvic flank mark and the ventral peduncular dark marks. E.
abernethyi differs from villosus in the much shorter predorsal
length (just greater than the distance from origin to origin of
the first and second dorsals in abernethyi, but reaching from first
dorsal origin to almost the upper caudal origin in villosus).
The etmopterids with bristle-like denticles differ less from
abernethyi in their denticle shape than they do from baxteri
which has shorter and less slender denticles than abernethyi. But
schultzi with its very elongate caudal (the upper margin of which
is about equal to the distance from snout tip to tip of pectoral
when the latter is laid back) and its fringed fins cannot be con-
fused with abernethyi whose upper caudal margin is just greater
than the length of head measured to the pectoral and whose
fins are not frayed more than is usual in other etmopterids. E.
hillianus differs from abernethyi not only in its bristle-like
denticles, as does spinax, but also in its greater peduncular length
(distance from rear ends of bases of pelvics to origin of lower
caudal equal to distance from snout tip to pectoral origin in
hillianus, but only to first gill-opening in abernethyi) ; the shape
of the pelvic flank mark and the ventral peduncular mark; and
the prepelvic transverse pale band which is lacking in abernethyi.
E. spinax has a peduncular length similar to abernethyi, but
differs in its random arrangement of bristle-like denticles, and its
colour patterns including the shape and extent of the pelvic and
peduncular dark marks.
E. polli, princeps and baxteri have denticles only slightly
stouter than those of abernethyi, but like spinax, these are ar-
ranged in random (at least anterior to the caudal peduncle) and
thus markedly different to the linear arrangement in abernethyi.
GARRICK : NEW ZEALAND ELASMOBRANCHII 189
The pelvic flank marks of polli and baxteri are much less elongate
than those of abernethyi, while the short interdorsal space of
polli (equal to less than the distance from snout tip to first gill-
opening in polli, but extending to the pectoral origin in aber-
nethyi) and the short stubby pectoral fins of princeps and baxteri
(failing to reach the first dorsal origin when laid back) clearly
distinguish these species from abernethyi.
The remaining three etmopterids, brachyurus, molleri and
lucifer, all agree fairly closely with abernethyi in their overall
proportions, their colour patterns (excluding the extensive mid-
dorsal pale band which seems to be characteristic of abernethyi) ,
and the nature and arrangement of their denticles. Bat brachyu-
rus and molleri are short-tailed species, the lengths of their upper
caudal margins reaching only two-thirds and three-fourths of the
distance from the rear ends of the pelvic bases to the lower caudal
origins, while in abernethyi the upper caudal margin is one and
a quarter times this distance. Moreover, in lateral view both
brachyurus and molleri are noticeably more sharp-snouted
species, the upper and lower profiles of the head tapering
smoothly to the snout tip. E. abernethyi is less sharp-snouted,
the head profiles little tapered from the eyes to the nostrils but
steepening rapidly from the nostrils anteriorly, where a distinct
change in the angles is evident. E. lucifer is intermediate be-
tween abernethyi, and brachyurus and molleri in the length of
its upper caudal margin (which is equal to the distance from
the rear ends of pelvic bases to the origin of lower caudal), but
differs from abernethyi in having a greater peduncular length
(distance from rear ends of pelvic bases to origin of lower
caudal equal to distance from snout tip to pectoral origin in
lucifer, but reaching only to first gill-opening in abernethyi) ;
a shorter snout (equal to or less than the length of the eye in
lucifer, but 1.5 times the eye-length in abernethyi) ; a different
dark pattern on the undersurf ace of the peduncle ; and the lack
of the extensive mid-dorsal pale band which is so prominent in
abernethyi.
190 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
LITEEATURE CITED
BlGELOW, H. B., W. C. SCHROEDER, AND S. SPRINGER
1953. New and little known sharks from the Atlantic and from the
Gulf of Mexico. Bull. Mus. Comp. Zool., 109 (3): 213-276, figs.
1-10.
Whitley, G.
1939. Studies in ichthyology no. 12. Bee. Australian Mus., 20 (4) :
264-277.
Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE
Vol. 116, No. 4
BIOLOGICAL INVESTIGATIONS IN THE SELVA
LACANDONA, CHIAPAS, MEXICO
Raymond A. Paynteb, Jr., Editor
With One Plate
CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM
April, 1957
Publications Issued by or in Connection
with THE
MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE
Bulletin (octavo) 1863 - - The current volume is Vol. ll(i.
Breviora (octavo) 1952 — No. 73 is current.
Memoirs (quarto) 1864-1938 — Publication was terminated with Vol. 55.
Johnsonia (quarto) 1941 — A publication of the Department of Mollusks.
Vol. 3, no. 35 is current.
Occasional Papers of the Department of Mollusks (octavo) 1945
Vol. 2, no. 21 is current.
Proceedings of the New England Zoological Club (octavo) 1899-
1948 — Published in connection with the Museum. Publication terminated
with Vol. 24.
The continuing publications are issued at irregular intervals in numbers
which may be purchased separately. Prices and lists may be obtained on
application to the Director of the Museum of Comparative Zoology,
Cambridge 38, Massachusetts.
Of the Peters "Check List of Birds of the World," volumes 1-3 are out
of print; volumes 4 and 6 may be obtained from the Harvard University
Press; volumes 5 and 7 are sold by the Museum, and future volumes will be
published under Museum auspices.
Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE
Vol. 116, No. 4
BIOLOGICAL INVESTIGATIONS IN THE SELVA
LACANDONA, CHIAPAS, MEXICO
Raymond A. Paynter, Jr., Editor
With One Plate
CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM
April, 19o7
No. 4 — Biological Investigations in the Selva Lacandona,
Chiapas, Mexico
TABLE OF CONTENTS
PAGE
I. Introduction. By Raymond A. Paynter, Jr 193
II. The Vegetation about Laguna Ocotal. By Robert L. Dressier 200
III. Land and Freshwater Mollusks of the Selva Lacandona, Chiapas,
Mexico. By Joseph C. Bequaert 204
IV. Ants from Laguna Ocotal. (Hymenoptera: Formicidae). By
William L. Brown, Jr 228
V. Fishes from Laguna Ocotal. By Robert Rush Miller 238
VI. Reptiles and Amphibians from the Selva Lacandona. By Ben-
jamin Shreve 242
VII. Birds of Laguna Ocotal. By Raymond A. Paynter, Jr 249
VIII. Design Quantities of some Chiapas Birds. By Charles H. Blake 286
IX. Mammals Collected at Laguna Ocotal. By Frances L. Burnett
and Charles P. Lyman 290
I
INTRODUCTION
By
Raymond A. Paynter, Jr.
In northeastern Chiapas, bordered on one side by the Rio
Usumacinta and on the other by the Rio Jatate, with northern
limits near Palenque and southern limits along the Guatemalan
border, there is a vast area of about 15,000 square kilometers
of almost unexplored and very sparsely inhabited virgin forest
(Map). The region is known as the Selva Lacandona, because
of the presence of scattered family groups of Lacandon Indians,
primitive and isolated remnants of the once vast Maya empire.
The Indians, who number less than 200 individuals,- have been
the subject of considerable publicity over the past decade, par-
ticularly after the discovery at Bonampak of well-preserved
murals in some ruined temples which are occasionally utilized by
the Lacandons. Several expeditions of archaeologists and an-
194 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
thropologists have been into the Selva Lacandona (see, e.g., Bloni
and Duby, 1955), principally in the vicinity of Bonampak, but
no biologists have reconnoitered the region, with the exception
of the Goodnights (1953), who studied the Phalangitis, and
Miranda (1953), who made a botanical survey. Both investiga-
tions were made at Monte Libano, on the edge of the forest, and
probably are only indications of what occurs in the interior.
The Selva Lacandona is on the northeastern slope of the central
highlands of Chiapas. There is a gradual decline in altitude
from a maximum of approximately 1400 meters, near the Rio
Jatate, to about 100 meters, at the Rio Usumacinta. Between
the two principal rivers are many lesser streams whose courses
are very poorly known, but which generally parallel one another
in a northwest-southeast direction. Scattered throughout are a
number of sizable lakes (lagunas), the largest of which are
Laguna Suspiro and Laguna Ocotal Grande (Plate, upper fig-
ure), nine and seven kilometers in length, respectively. The
natives call the latter lake simply "Laguna Ocotal," and all the
zoological material obtained there was so labeled, but on a map
prepared by Frans Blom (see Map) the longer name is used, in
contradistinction to Laguna Ocotal Chico, a much smaller lake
to the northeast.
It is unfortunate that nothing certain is known of the drainage
of these important lakes. While they may drain southeast into
the Rio Lacanja, thence into the Rio Lacantun, and finally
into the Usumacinta basin, there is no evidence that this is the
case, despite the predilections of cartographers for drawing con-
necting streams between known lakes and known rivers. From
personal observations, limited to the region about Laguna Ocotal,
I am inclined to believe that these lakes have formed within
closed-end solution valleys. The heavily karsted limestone
strongly suggests such an origin. Underground drainage to the
Usumacinta basin is possible, of course.
Miranda (1952) has prepared a generalized vegetation map of
Chiapas in which the Selva Lacandona is depicted as being cov-
ered by high evergreen forest, with areas of pines and oaks at
the lakes and along the southwestern boundary of the region.
It should be realized, however, that "high evergreen forest" is a
broad term embracing a great diversity of vegetational assem-
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO
195
196 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
blages. For example, at Monte Libano the forest is high, lux-
uriant, and has a relatively clear understory. It is what is usually
referred to as "rain forest," which in this part of the world
commonly contains, among other characteristic species, mahogany
(Swietenia humilis) and ramon (Trophis racemosa or Brosimum
alicastrum). On the other hand, at Laguna Ocotal there is a
forest which seems physiognomically quite similar but which is
markedly different in composition, lacking, in part, ramon and
mahogany, while gigantic oaks (Quercus spp.) are present. A de-
tailed description of the vegetation about Laguna Ocotal is found
in Dressier 's account (pp. 200-203).
Nothing is known of the climatology of the region. Since the
altitude and vegetation vary in the Selva Lacandona, it is reason-
able to suppose that the climate is also variable. The nearest
location from which weather records are aATailable is Tenosique,
Tabasco, a town at a somewhat lower elevation (60 m.) than the
lowest point in the Selva Lacandona. Here the mean annual
rainfall is 1697 mm., with June being the wettest month and
March the driest ; the warmest month is May, which has a mean
temperature of 30.4°C. and the coolest month is January, with a
mean of 22.9°C. (Ward and Brooks, 1936).
In 1954, with generous support from the American Academy
of Arts and Sciences and from the Chapman Memorial Fund of
the American Museum of Natural History, a party was formed
in order to make the first biological survey of the interior of the
Selva Lacandona. The group consisted of the author, as leader
and ornithologist, Robert T. Paine, 3rd, as assistant ornithologist,
Elisha F. Lee, as mammalogist, Robert L. Dressier, as botanist,
and Mrs. Ruth Oberg, also a botanist, specializing in the Orchi-
daceae.
In early July the party flew from the capital of Chiapas,
Tuxtla Gutierrez, to Ocosingo (alt. 850 m.), a village about one
hundred kilometers to the northeast. Heavy rains had raised
the level of the nearby rivers and delayed for several days the
arrival of our pack animals. Finally, on the morning of July 10
we started for Finca El Real (alt. 600 m.), which is approxi-
mately 40 kilometers to the east, and reached there in the after-
noon of the following day.
At El Real additional supplies, mules, and men were secured
and on July 15, with 18 mules and 9 arrieros, trail-cutters, etc.,
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 197
we moved 20 kilometers east to the settlement of Monte Libano
(alt. 600 m.). During the night a number of the mules strayed
and we were unable to recover them and move on until July 18.
The trail-cutters had been sent ahead to clear the way and pre-
pare bridges, but progress was slow and difficult owing to the
mud and obstructions in the trail. About seven hours after leav-
ing Monte Libano we arrived at El Censo (alt. 700 m.), an
uninhabited camping spot in magnificent rain forest. The fol-
lowing morning the trail was poor during the first hour of
travel, but then became worse due to an escarpment which rises
over 300 meters above the country to the east. Ascent was made
by steep switchbacks which were barely surmounted by the pack
animals. Beyond this point the trail improved somewhat. In the
middle of the afternoon our destination was reached and a camp
was prepared at the northwestern end of Laguna Ocotal (Plate,
lower figure), at an altitude of 950 meters. Because of insufficient
forage at the lake, most of the mules were returned to El Real, but
four men remained to assist with the collecting and maintenance
of the camp.
The forest is dense and the terrain rough around Laguna Oco-
tal, which means that it is seldom practicable to work far from a
trail. The only trails existing in the area are the one by which
we entered, and a badly obstructed path leading to Bonampak.
Consequently, Ave were not able to range as far as desired and
even after a month of intensive collecting did not reach the
opposite end of the lake or the pine-covered ridges along its
southwestern side. Future workers in the region would do well
to employ an additional man or two to cut new trails, thereby
enabling the collectors to sample a larger area.
The campsite was situated on a narrow strip of ground which
has cut off a small arm of the lake, impounding the water and
forming a swamp. Most of the mammals, and many of the
amphibians were collected here.
Although each member of the party concentrated on his field
of interest, unrelated material was collected whenever pos-
sible. For this reason the bird, mammal, and botanical1 col-
lections are probably the most nearly representative samples
from the region, while the fish, reptile, amphibian, and inverte-
i No complete report on the botanical collections has been prepared. A list of
the Orchidaceae has been compiled by Mrs. Ruth Oberg (in press).
198 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
brate collections contain, in the main, only the more conspicuous
elements of their faunas.
At the end of a month the mules were brought back from the
ranch and on August 20 the camp was abandoned. We returned
to El Real by the same route used in entering the forest and
on August 30 were flown from the ranch to Tuxtla Gutierrez.
An especial debt of gratitude is owed to the members of the
field party. Their whole-hearted cooperation is evident from
the large amount of material which was collected, although the
period was brief and conditions difficult. During preparations,
and while in the field, we received much assistance and many
courtesies from Horacio Albores of Ocosingo, Jose Tarano and
Juan Bulnes of Finca El Real, and Frans Blom of San Cristobal
de las Casas. I wish, also, to express my appreciation to Joseph
C. Bequaert, Charles H. Blake, William L. Brown, Jr., Frances
L. Burnett, Robert L. Dressier, Charles P. Lyman, Robert Rush
Miller, and Benjamin Shreve, who have submitted the following
reports based on material obtained during the expedition.
LITERATURE CITED
Blom, Frans and Gertrude Duby
1955. La Selva Lacandona. Mexico, D.F. Editorial Cultura, 448 pp.
Goodnight, Clarence J. and Marie L. Goodnight
1953. The Opilionid Fauna of Chiapas, Mexico and Adjacent Areas
(Araehnoidea, Opiliones). Amer. Mus. Novit., No. 1610, 81 pp.
Miranda, Faustino
1952. La Vegetacion de Chiapas, Parte I. Tuxtla Gutierrez, Chiapas,
Dept. de Prensa y Turisnio, 334 pp.
1953. Un Botanieo en el Borde de la Selva Lacandona. Mem. Congreso
Cient. Mex., 6: 285-303.
Oberg, Ruth
[In pressj. Orchids Collected at Laguna Ocotal Grande, Mexico.
Orchid Jour., 3.
Ward, Robert DeC. and Charles F. Brooks
1936. The Climates of North America. Pt. I. Mexico, United States,
Alaska. In Handbuch der Klimatologie. II. Amerika. Teil J
(1 Lief.), 325 pp.
IJIOI.OCK'AL IXVKSTIGATIONS IX CHIAPAS, MEXICO
1!)!)
The northern cud of Laguna Ocotal. The pine-covered ridges parallel the
southwestern shore of the lake.
Photo by Li i
A small liill covered by pines projects from a low deciduous forest (monte;
see Dressier, pp. 200) near the campsite at Laguna Ocotal. The lofty
tropical evergreen forest begins behind the hill and is not visible.
Photo by l.< e
200 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
II
THE VEGETATION ABOUT LAGUNA OCOTAL
By
Robert L. Dressler 1
Only a preliminary characterization of the plant cover can be
made at this time. Many important species were not in flower or
fruit, and the upper stories of the rich tropical evergreen forest
were sampled only through occasional windfalls.
Four main vegetation types may he recognized in the area :
(1) Pine Forest, or ocotal, which is usually at a higher level on
a given slope than is ni<>ut< or selva, but occurs down to the
lake shore near the campsite. Pine forest is said to extend south-
ward for some distance on the ridge west of the lakes. (2) Monte,
a dense transition of small, usually slender, hardwoods. This
vegetation generally occurs between the ocotal and the selva or
the lake shore. (3) Tropical Evergreen Forest (selva). This,
the "montana" of the natives, makes up the hulk of the Selva
Lacandona of eastern Chiapas and apparently surrounds the
lake area. (4) Cloud Scrub, a distinctive type limited to prom-
ontories overlooking the lake.
(1) Pine Forest. The pine stands include some splendid, large
specimens of Pinus tenuifolia Bentham (probably the only
species present i, but are not continuous or very extensive in the
area studied. The pines probably occupy only slopes and hilltops
which are loo well drained for the more mesic forest types. Open
park-like sites, which are few and quite small, have a grassy
ground cover including Cyperus, Scleria, Dichromena, and Pani-
cum. In favored sites an understory is formed by small hard-
wood trees, such as Hauya h< ij</< <ma Donnell-Smith, a reddish-
barked tree with much the aspect of madrono, Saurauia subal-
pina Donnell-Smith, and Myrica cerifera L. These trees are
often quite mossy and hear a rich epiphyte flora of many species
of orchids and ferns, most of which also occur in the upper
stories of the selva. The pines themselves hear many epiphytes,
hut these are of relatively few species, the genera Tillandsia,
l Gray Herbarium, Harvard University, Cambridge, Massachusetts.
BIOLOGICAL INVESTIGATIONS I N CHIAPAS, MEXICO 201
Catopsis, and Epidendrum being conspicuous. Chimaphila macu-
hifd (L.) Pursh was found on one slope, and a small fan-palm
(Brahea) occurs in dry sites. The greater pari of the pine forest
has a great deal of undergrowth and might he considered as a
marginal type passing into the monte.
(2) Transition Forest or Monte. This is a somewhat diverse
assemblage of vegetations which is characterized by its relatively
low height (mostly about 5-10 m.) and great density, its position
between the pine forest and the selva, and a very poor epiphyte
flora. Only a few of the trees making up this vegetation have
been identified. A pink-fruited Hoffmannia is frequent, Acaly-
pha gummifera Lundell and Cnidosculus midtilobus (Pax)
Johnston are local, and Cecropia was observed in one stand.
A slender Verbesina and Calliandra houstoniana (Miller
Standley are frequent in drier sites. Undergrowth is relatively
scant in most parts of the monte, but the shrubby Cephaelis
tomentosa (Aublet) Vahl is abundant on the selva side of the
transition, and the fern, Nephrolepis cordifolia (L.) Presl, is
locally very abundant on the ocotal side.
This vegetation may. on an abrupt slope, form a very narrow
zone, or it may be more widespread. In one or two sites near
the camp it is fairly extensive in small areas that have been
disturbed (cut over for fire wood ami poles for camp con
struct ion. burnt over, and doubtless heavily grazed by mules at
infrequent intervals). At the north end of the lake clear evi-
dence of disturbance is found in a nearly pure stand of small
"pomarosa", Eugenia (Syzygium) jambos L. (which may be
considered a local phase of the nioiih ). This Asiatic species i>
certainly introduced. There are two or three very large examples
of pomarosa and two large clumps of bamboo near the Bonampak
trail, which may date to its original introduction. On a slope
nearby, in the ocotal, there is a small area where Lantana camara
L., Trema micrantha (L.) Blume, Euphorbia hirta L., and
Psidium (probably /'. </u<tj<tr<i L.) occur together. None of these
weedy species was seen elsewhere in the region, and they surely
indicate past disturbance. Many of the elements of the monte
are probably those that would occur in secondary growth any-
where in the area (for example. Cnidosculus, Cecropia).
202 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
(3) Tropical Evergreen Forest or Selva. This is the Selva
Alta Siempre Verde of Miranda (1952). I have used "selva"
rather than the local "montana" in my notes and discussion to
avoid confusion with "monte". This fine forest is similar to
(and continuous with) that near Monte Libano, which has been
described by Miranda (1953b), though the forest about Laguna
Ocotal lacks ehicozapote, ramon, mahogany, and the fiendishly
spiny palm, Hexopetion. Ground cover in the selva is relatively
sparse and walking about is easy. Fallen fruits and flowers in-
dicate that Talauma mexicana (de Candolle) Don and Cymbo-
jx lain in penduliflorum (Dunal) Baillon are frequent, and the
fruits of Sloanea were seen at one place. Oaks (Quercus SJcinneri
Bentham and perhaps other species) occur frequently, especially
in the drier phases of the selva. These slightly more open, less
humid stands, curiously enough, are not to be found near the
drier ocotal, but at some distance from the lake, along the
Bonampak trail. The selva adjoining the ocotal (or transition)
usually shows its wetter phases. The upper stories of the selva are
rich in epiphytes of many kinds. The lower tree trunks bear
ferns, peperomias, gesneriads and a few species of orchids
(such as Chondrorhyncha lendyana Reichenbach til., Pleurothal-
lis eardiothallis Reichenbach til., Maxillaria maleolens Sehlechter,
and M. nasuta Reichenbach fil.) which are largely or quite re-
stricted to this habitat. Collinia, Eleutheropetalum, and several
species of Chamaedorea were the only palms seen in the selva;
tree ferns (Cyathea) are locally abundant.
(4) Cloud Scrub. This vegetation, probably comparable to
the "elfin woodland" of the West Indies, was found only on
cliff tops overlooking the west side of the lake (the prevailing
winds are from the east, across the water). These local and rela-
tively inaccessible sites were not visited by the expedition zool-
ogists. The limestone promontories are very well drained and,
probably for this reason, support only relatively small trees
(predominantly Clusia rosea Jacquin?), but, while the arboreus
elements are comparable in size and form to monte ha jo or
chaparral, the great atmospheric moisture supports an abundance
of epiphytes. These epiphytes are largely mosses and ferns, but
some orchids, Ericaceae and other epiphytes of the high selva
also flourish in the Clusia scrub.
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 208
These collections and notes are too scanty and our knowledge
of the Lacandon Forest, as a whole, too slight to say much about
the geographic affinities of the Laguna Ocotal region. A number
of the expected "Guatemalan" and "Honduran" species, new
to the known flora of Mexico, were found, especially in the rich
orchid flora (Oberg, in press). The pine forest itself is rela-
tively poor in distinctive elements, but this is perhaps to be
expected from its small and discontinuous area. It is possible
that this ocotal has lost ground to the monte in relatively recent
times through the occasional disturbances caused by chicleros,
anthropologists, and others who have used this campsite on the
Bonampak trail. A more long-term vegetational change may be
indicated by the pine logs found in selva near Monte Libano
(Miranda 1953b).
LITEEATUEE CITED
Miranda, Faustino
1952. La Vegetacion de Chiapas. Parte I. Tuxtla Gutierrez, Chiapas.
Dept. de Prensa y Turismo, 334 p.
1933a. La A'egetacion de Chiapas. Parte II. Tuxtla Gutierrez, Chiapas,
Dept. de Prensa y Turismo. 426 p.
L953b. Un Botanico en el Borde de la Selva Lacandona. Mem. Congreso
Cient. Mex., 6: 285-303.
Oberg, Euth
[In press.] Orchids Collected at Laguna Ocotal Grande, Mexico.
Orchid Journ., 3.
204 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
III
LAND AND FRESHWATER MOLLUSKS OF THE
SELVA LACANDONA, CHIAPAS, MEXICO
By
Joseph C. Bequaebt
To conform with the general plan of the reports on the col-
lections made by Dr. Raymond A. Paynter, Jr., and his associ-
ates in Mexico, during the summer of 1954, only the 39 species
obtained in the Selva Lacandona have been fully treated below.
However, in order to make the paper more generally useful, a
list is appended of 49 additional species definitely reported from
the State of Chiapas. In this list localities are recorded for 9
species obtained by Dr. Paynter 's party outside the Lacandona
area. In all, 88 species of inland mollusks are thus known at
present from the State of Chiapas, 73 of them terrestrial and
15 aquatic.
As Chiapas has not been visited thus far by a professional
malacclogist, the 88 species obviously represent only a fraction
of the actual molluscan fauna. A comparison with the better
known adjoining areas of Mexico (Veracruz) and Guatemala
(Peten and Alta Vera Paz), where the ecology and topographical
conditions are similar, suggests that the molluscan fauna of
Chiapas should comprise some 180 to 200 species. The known
88 species came from several distant localities and were obtained
by different collectors during the past 100 years. They may be
regarded as a random sample of at least the more common and
more conspicuous forms, sufficiently representative to determine
the dominant features of the molluscan fauna of the State, as
well as of the Selva Lacandona. It should be kept in mind, more-
over, that the mollusks of the better explored adjoining terri-
tories are as yet far from being completely known.
The 73 terrestrial species will be considered first, as they
form the bulk of the fauna and are most interesting from our
point of view. At present 26 of them, or about one-third of the
total, are known only from Chiapas or extend in a few cases
just north of the boundary to Tabasco. Although this figure
seems to point to a fairly higli degree of endemicity, it is prob-
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 205
ably deceptive. Several of these "endemic" species are poorly
known, often only from single specimens and not from one or
more populations. In fact I am personally acquainted with
only eight of these supposedly endemic forms. I suspect that
eventually many of them will he recognized as identical with,
or within the range of intraspecific variation of more widespread
species, recorded under other names from adjoining areas.
The largest group comprises 28 species (slightly over one-
third of the total) which generally extend from the western sec-
tion of Veracruz, through Tabasco and Chiapas, to Guatemala
and British Honduras. Of these, 15 are at present recorded from
Chiapas (and sometimes Tabasco) to Guatemala, 9 from Vera-
cruz, Tabasco and Chiapas, and 4 from Veracruz to Guatemala ;
but there can be little doubt that all of them actuall}7 occur
throughout the whole area. As suggested above, this group will
eventually grow by the addition of several of the species now
supposedly endemic in Chiapas. It appears to be the dominant
and characteristic molluscan assemblage for what may perhaps
be called the Chiapas-Guatemalan Subregion of Central America.
In all, 54 of the 73 terrestrial species known from Chiapas have
not been recorded outside this Subregion.
The remaining species are much more widely distributed. Most
of them reach their northern limit in Veracruz, although usually
extending southward far beyond Guatemala, in some cases even
to Costa Rica and Panama. Four of them are "followers of
man," who has spread them far and wide outside their original
home, which is now sometimes in doubt.
The very limited freshwater fauna consists of only 15 species.
As is often the case for freshwater mollusks in the tropics, they
are not particularly characteristic. Seven species are widely
distributed throughout Mexico and Central America, or some
even beyond; 5 are possibly restricted to the Veracruz-Chiapas-
Guatemala area, like the bulk of the terrestrial species; and 3
freshwater clams are at present known only from Chiapas, prob-
ably due to insufficient knowledge of the Central American naiad
fauna as a whole.
206 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
HEL1CINIDAE
Oligyea elavida (Menke)
Helicina flavida Menke, 1828, Synopsis Meth. Moll., 1st Ed., p. 79 ("Ja-
maica," [erroneous locality])- Sowerby, 1842, Thesaur. Conchyl., 1. Pt.
1, p. 9; PI. 3, figs. 117 and 134; 1866, Op. cit., 3, Pts. 24-25, p. 287;
PI. 272, figs. 233-236 (Guatemala).
Laguna Ocotal, 950 m.
A common species in southern Mexico (Veracruz, var. strebeli
Pfeiffer; Tabasco; Chiapas), Guatemala (Peten), and the Re-
public of Honduras (Copan).
Oligyea oweniana (Pfeiffer)
Helicina oweniana Pfeiffer, 1849 (April), Proc. Zool. Soc. London, (for
1848), p. 123 (Chiapas); 1850, Syst. Conch.-Cab., 1. Abt. 18, Pt. 1, p.
40; PI. 7, figs. 35-36 (type).
A common species in Chiapas, Tabasco, Guatemala and Costa
Rica. Two forms are represented in the collection from the Selva
Lacandona.
1. var. coccinostoma Morelet. Helicina coccinostoma Morelet.
1849, Test. Noviss. Ins. Cub. Amer. Centr., 1, (19 Peten).—
Laguna Ocotal, 950 m.
2. var. anozona von Martens. Helicina anozona von Martens,
1876 (April), Proc. Zool. Soc. London, (for 1875), p. 649
(Coban) ; 1876, Jahrb. deutsch. Mai. Ges., 3, p. 261 ; PI. 9, fig. 7.—
Monte Libano, 600 m. ; El Real, 600 m.
Helicina amoena Pfeiffer
Helicina amoena Pfeiffer, 1849 (April), Proc. Zool. Soc. London, (for
1848), p. 119 (Honduras); 1850, Syst. Conch.-Cab., 1. Abt. 18, Pt. 1,
p. 55; PI. 5, figs. 13-15 (type).
Laguna Ocotal, 950 m. ; El Censo to Laguna Ocotal, 1000 m. ;
Ocosingo, 850 m. ; El Real, 600 m. ; Monte Libano, 600 m.
Known from southern Mexico (Campeche and Chiapas),
Guatemala, British Honduras, the Republic of Honduras, Nic-
aragua, and Panama.
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 207
Helictna tenuis Pfeiffer
Helicina tenuis Pfeiffer, 1849 (April), Proc. Zool. Soe. London, (for 1848),
p. 124 (Yucatan) ; 1850, Syst. Conch.-Cab., 1. Abt. 18, Pt. 1, p. 40;
PI. 7, figs. 33-34 (type).
Helicina chiapensis Pfeiffer, 1856 (December), Mai. Blatt., 3. p. 237
(Chiapas); 1S57 (May), Proc. Zool. Soc. London, (for 1856), p. 380.
Sowerby, 1866, Thesaur. Conchyl., 3. Pts. 24-25, p. 288; PI. 7, figs.
255-257 (? type from Cuming Collection).
El Pveal, 600 m. ; Monte Libano, 600 m.
This species is widespread from southern Mexico (Jalisco;
Veracruz, Chiapas; Tabasco, Yucatan), to Guatemala (Peten;
Alta Vera Paz, and the Pacific slopes of the Cordillera), Nic-
aragua, Costa Rica, and Panama.
Schasicheila alata (Pfeiffer)
Ilelicina alata "Menke" Pfeiffer, 1849 (January), Zeitschr. f. Malakoz.,
5. (for June 1848), p. 87 (Mexico); 1850, Syst. Conch.-Cab., 1, Abt.
18, Pt. 1, p. 43; PI. 5, figs. 18-20 (type).
Menke 's name was given in MS and the original description
was written by Pfeiffer, to whom the name should be credited.
According to a note on page 96, dated January 10, 1849, the
June issue of the Zeitschr. f. Malakoz. for 1848 could not have
been published before January 1849.
Monte Libano, 600 m.
Known from southern Mexico only (Veracruz and Chiapas).
Schasicheila pannucea (Morelet)
Helicina •pannucea Morelet, 1849, Test. Noviss. Ins. Cub. Amer. Centr., 1,
p. 21 (San Luis, Peten).
Sehasichila pannucea Fischer and Crosse, 1892, Mission Scientif. Mexique,
Moll. Terr. Fluv., 2. Pt, 13, p. 448; PI. 54, figs. 5 and 5a (type from
Morelet).
Monte Libano, 600 m.
Known from much of Mexico (Chiapas; as var. misantlensis
Fischer and Crosse from Veracruz and Puebla; and as var.
hidalgoana Dall from Hidalgo, San Luis Potosi and Tamaulipas)
and Guatemala (Peten and Alta Vera Paz).
208 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Lucidella lirata (Pfeiffer)
Helicina lirata Pfeiffer, 1847, Zeitschr. f. Malakoz., 4, p. 150 (Yucatan) ;
1850, Syst. Conch.-Cab., 1. Abt. 18, Pt. 1, p. 14; PL 4, figs. 40-43 (type).
Laguna Oeotal, 950 m. ; Laguna Ocotal to El Censo, 1000 m.
Known from southern Mexico (Veracruz; Chiapas; Tabasco;
Yucatan; Quintana Roo), Guatemala, British Honduras, the
Republic of Honduras, Nicaragua, Costa Rica, and Panama ; and
as subsp. lamellosa Guppy from Venezuela and Trinidad.
CYCLOPHORIDAE
Neocyclotus dysoni aureus (Bartsch and Morrison)
Aperostoma (Neocyclotus) dysoni aurcum Bartsch and Morrison, 1942,
Bull. U. S. Nat. Mus., 181, p. 209; PI. 28, rigs. 13-15 (Panistlahuaca,
Oaxaca).
Xeocyclotus dysoni aureum Solem, 1956, Proc. Acad. Nat. Sci. Philadelphia,
108, p. 53 (Oaxaca: Gamboa. Guatemala: Amatitlan. El Salvador:
Lake Coatepeque).
Xeocyclotus dysoni Fischer and Crosse, 1888, Mission Scientif. Mexique,
Moll. Terr. Fluv., 2. Pt. 10, p. 164 (in part: specimens from Chiapas
only). Not typical N. dysoni (Pfeiffer, 1853).
Laguna Ocotal, 950 m. ; El Sumidero, Tuxtla-Gutierrez, 1000
m.
N. dysoni is a common widespread and variable species in
southern Mexico (Veracruz; Oaxaca; Tabasco; Chiapas; Campe-
che; and Yucatan), Guatemala, El Salvador, the Republic of
Honduras, Nicaragua, Costa Rica and Panama. The subspecies
aureus is known from Oaxaca, Chiapas, Tabasco, western Guate-
mala and El Salvador.
Amphicyclotus palenquensis (Pilsbry)
Aperostoma (Amphicyclotus) palenquense Pilsbry, 1935, Proc. Acad. Nat. Sci.
Philadelphia, 87, p. 3; PI. 1, figs. 3 and 3</-?> (district of Palenque,
Chiapas).
Megacyclotus palenquensis Bartsch and Morrison, 1942, Bull. U. S. Nat.
Mus., 181. p. 183; PI. 24, figs. 16-18 (type).
Amphicyclotus (Aniphicyclotus) palenquense Solem, 1956, Proc. Acad. Nat.
Sci. Philadelphia, 108, p. 44 (Veracruz: Motzorongo).
Laguna Ocotal, 950 m. ; Monte Libano, 600 m. ; El Censo to
RIOLOGICAI, INVESTIGATIONS IN CHIAPAS, MEXICO 209
Laguna Ocotal, 700-1000 m. ; Monte Libano to El Censo, 600-
700 m.
The 30 specimens from the Lacandona area agree well in
shape and in sculpture with the descriptions and figures of
Pilsbry and of Bartsch and Morrison. When well preserved, the
sculpture consists of microscopic, spiral, slightly wavy, densely
crowded lines and coarser diagonal, curved threads, spirally
ascending forward and crossing the irregular, vertical growth
lines. The retractive threads are often irregular or interrupted
by coarse malleations and are usually best marked at the peri-
phery, below which they are sometimes nearly horizontal.
The species was known thus far from only three specimens.
The type measured 20.7 mm. in height, 34 mm. in greater diam-
eter and 25.9 mm. in lesser diameter. The corresponding figures
for the paratype were 21.7 mm., 34.3 mm. and 26.1 mm. As will
be seen from the subjoined table, some of the 22 fully adult shells
of the Selva Lacandona approach these measurements closely;
a few are slightly smaller (down to 29.8 mm. in greater diameter
and 18 mm. high), and several are decidedly larger (up to 41.5
mm. in greater diameter and 24.7 mm. high). The larger speci-
mens seem to bridge the gap between A. palenquensis and A.
ponderosus (Pfeiffer), so far as size is concerned. To judge from
the specimens of A. ponderosus at the M.C.Z., that species has,
however, a decidedly higher spire and a less flattened, more
convex body- whorl, as well as a deeper and somewhat narrower
umbilicus, than A. palenquensis. The sculpture seems to be
about the same in both species.
The specimen of A. ponderosus figured by Bartsch and Morri-
son was of about the size of our largest A. palenquensis. Other
known specimens are, however, much larger (up to 48.5 mm. in
greater diameter and 29 mm. high in a series of 4 specimens from
northern Guatemala at M.C.Z.). It is therefore possible that
A. ponderosus, definitely recorded only from Guatemala (Alta
Vera Paz) and British Honduras, actually grows larger than A.
palenquensis. The latter is known only from northeastern Chia-
pas, the Selva Lacandona being some 70 km. south of Palenque.
In general shape, measurements, depressed spire and widely
open umbilicus A. palenquensis resembles Am phi cy clot us mega-
planus Morrison (1955, Proc. Washington Acad. Sci., 45, p. 160,
210 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
figs. 29-31), from El Ocote, some 35 km. south of Ocozocoautla,
Chiapas and some 180 km. west of the Selva Lacandona. How-
ever, the sculpture of the later whorls of megaplanus is described
as ' ' consisting of fine irregular axial vermiculate ribbing, ' ' being
similar to that of Amphieyclotus texturatus (Sowerby). Solem
suggests that megaplanus may be only a local race of texturatus.
Measurements of Adult Amphieyclotus palenquensis (in mm.)
eight
20.3
Width
Greater Lesser
38.4 29
Aperture
Height Width
19.2 17.7
Whorls
5%
Lagiuia Ocotal
20
36.2
27.4
17
15.5
5%
t 1
< t
20
35
27.3
16.8
15.5
5%
i t
i i
20
34.5
27.5
18
14.4
5%
1 1
1 1
23.7
38
30
18
16.5
5V2
Monte
Libano
20.7
37.3
29.3
19.8
16.8
sy2
1 1
t i
21.5
37
28
18.3
17
5V2
i 1
1 1
21.5
36
28
18
17
5y2
1 1
i 1
20.8
35
27
18
16
5y3
1 i
1 1
21
35
27
16.6
15.2
5V3
1 1
1 1
21
34.5
27.2
16
15.2
5 V*
i <
1 1
20
33.6
27.5
16.5
15.4
5 V*
< <
1 1
20.8
32.5
25
15.8
15
514
< 1
1 1
18
31.5
26.4
14
13.5
5^4
i <
1 1
18.3
31
23.5
15.3
13.7
5^4
1 1
1 1
19
29.8
24
14.3
13.7
5V5
1 1
< <
24.7
38.7
30.4
19.8
18.8
5y2
El Censo to L. Ocotal
21.2
37.6
27
18.4
16.2
sy2
t <
< <
22.8
37.3
27
18.6
16.7
5y2
t <
< <
20.5
38
30
18
15.7
sy2
Monte Libano to El Censo
20
36.5
27
17.2
15.2
5%
a it
22
36.4
28.5
17.3
16
51/3
t 1 it
PILIDAE (AMPULLARIIDAE)
POMACEA FLAGELLATA GHIESBRECHTII (Reeve)
Ampullaria ghiesbrechtii Reeve, 1856 (December), Conch. Icon., 10. Ampul-
laria, PI. 26, fig. 123 (Chiapas).
Ampullaria ghiesbreghti Bhmey, 1865, Land Fresh-Water Shells North
America, 3. (Smithson. Misc. Coll. No. 143), p. 7 (emendation of
ghiesbrechtii).
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 211
Ampullaria livescens Reeve, 1856 (August), Conch. Icon., 10, Ampullaria,
PL 5, fig. 21 (no locality).
Ampullaria miltocheilus Reeve, 1856 (December), Conch. Icon., 10. Ampul-
laria, PI. 25, figs. 102a-b (Chiapas).
Pomus giganteus Tristram, 1863, Proc. Zool. Soc. London, p. 414 (Lake of
Peten, Guatemala).
Ampullaria flagellata var. gigantea von Martens, 1S99, Biologia Cen.tr.-
Amer., Terr. Fluv. Moll., p. 412; PL 23, fig. 6 (cotype received from
Tristram).
Ampullaria malleata var. chiapasensis Fischer and Crosse, 1890, Mission
Scientif. Mexique, Moll. Terr. Fluv., 2. Pt. 11, p. 235; PL 48, fig. 5
(Las Playas, Chiapas; [immature shell duplicated by some specimens
from Laguna Ocotal]).
Laguna Ocotal, 950 m., many young shells. The largest meas-
ures 53 mm. in length, 50 mm. in greatest width, with the aper-
ture 40 mm. by 24 mm.
Pomacea flagellata (Say) is the common ampullariid snail in
Mexico and Central America, from Veracruz southward, as far
as Panama and northern Colombia. It is extremely variable in
shape and size (when adult), even in the same population. Some
30 names have been proposed for these variations and, as some
of these have been proposed either for unusual or freak speci-
mens or for immature or juvenile snails, it is extremely difficult
to dispose of them as synonyms of the few races or geographical
forms that may be usefully recognized.
One of the best characterized of these races, seemingly re-
stricted to Tabasco, Chiapas and northern Guatemala, is nearly
globular in shape, usually about as high as its greatest width,
sometimes slightly higher or slightly lower. Under optimum
conditions it may reach greater dimensions than any of the other
races of the species. ''Giant" specimens are particularly com-
mon in Lake Peten, whence Tristram described his P. giganteus.
The original measurements given by Tristram were : height,
95 mm.; greatest diameter, 90 mm.; lesser diameter, 85 mm.;
aperture, 66 by 39 mm. The cotype figured by von Martens is
92 mm. high, with the aperture 69.5 mm. long. The largest
specimen I have seen from Lake Peten is 102.5 mm. high, 88 mm.
in greatest width, with the aperture 74.3 by 42 mm., of about 6
whorls. The largest of several collected by Mr. F. G. Thompson
4 miles south of Villahermosa, Tabasco, is 83 mm. high and 75
mm. wide.
212 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
There can be little doubt that the type of Reeve's A. ghies-
brechtii was a slightly smaller specimen of P. giganteus with
unusually bright vermilion margins of the aperture. A brightly
colored aperture occurs sporadically in several species of Poma-
cea, but is never a reliable specific character. The color may be
more or less pronounced, and, moreover, is often more orange,
as in Fischer and Crosse's figure of A. ghiesbrechtii from the
Usamacinta River in Tabasco (Miss. Scientif. Mexique, Moll.
Terr. Fluv., 2, PI. 48, fig. 8), where it is present only over the
outer margin of the mouth. Reeve's figure of the type of
ghiesbrechtii is 85 mm. high, 73 mm. in greatest width, with the
aperture 59 mm. by 36.5 mm.
A. miltocheilus Reeve appears to be no more than a small
specimen of A. ghiesbrechtii; it was collected by the same per-
son in Chiapas also, and both may have come from the same
population. The figure is 48 mm. high, 43 mm. in greatest
width, with the aperture 36.5 mm. by 21.5 mm. The aperture
has the same bright vermilion color as the type of ghiesbrechtii.
A. livescens Reeve agrees well with some young specimens of
A. ghiesbrechtii found in Lake Peten with the giant specimens.
Although the name was published some months before ghies-
brechtii, I prefer to use the latter, because it was based on a
fully adult shell from a precise locality and dates from the same
year.
POMATIASIDAE
Chondropoma rubicundum (Morelet)
Cyclostoma rubicundum Morelet, 1849, Test. Noviss. Ins. Cub. Amer.
Centr., 1. p. 22 (Peten and Vera Paz, Guatemala).
Chondropoma rubicundum Fischer and Crosse, 1890, Mission Scientif. Mex-
ique, Moll. Terr. Fluv., 2, Pt. 11, p. 205; PI. 41, figs. 5f-h (cotype from
Vera Paz).
Laguna Ocotal, 950 m.
Known from Tabasco, Chiapas and Guatemala (Peten and
Alta Vera Paz).
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 213
BULIMIDAE (HYDROBIIDAB)
Amnicola guatemalensis Fischer and Crosse
Paludina hyalina Morelet, 1851, Test. Noviss. Ins. Cub. Amer. Centr., 2,
p. 21 (Lake Amatitlan, Guatemala). Not of Anton, 1839.
Amnicola guatemalensis Fischer and Crosse, 1891, Mission Scientif. Mexique,
Moll. Terr. Fluv., 2. Pt. 12, p. 264; PI. 50, figs. 5 and 5a-b (cotypes:
new name for hyalina Morelet).
Laguna Ocotal, 950 m. ; many dead specimens in a silt deposit
on the shore.
The species, known thus far from several localities in Guatemala
(Amatitlan; Peten; etc.), is now reported for the first time from
Chiapas. Guatemalan specimens were compared.
Cochliopa infundibulum von Martens
Cochliopa (?) infundibulum von Martens, 1899, Biologia Centr. -Amer., Terr.
Fluv. Moll., p. 429 ; PI. 23, fig. 3 (Guatemala, without precise locality ;
surmises it may be from Lake Peten).
Laguna Ocotal, 950 m. Many dead specimens in a silt deposit
on the shore.
Previously known only from Lake Peten and Laguna cle Ecki-
bix, in northern Guatemala (Goodrich and Van der Schalie, 1937,
Mus. Zool. Univ. Michigan, Misc. Publ. No. 34, p. 37) ; now
reported for the first time from Chiapas. Guatemalan specimens
were compared.
THIARIDAE (MELANI1DAE)
Pachychilus indiorum (Morelet)
Melania indiorum Morelet, 1849, Test. Noviss. Ins. Cub. Amer. Centr., I.
p. 25 (Palenque, Chiapas).
Melania indorum Petit, 1853, Jour, de Conchyliologie, 4, p. 162; PI. 5, fig.
7 (cotype received from Morelet).
Melania laevissi7na var. costato-plicata Brot, 1875, Syst. Conch.-Cab., 1.
Abt. 24, p. 35; PI. 5, fig. 5 (Palenque, Chiapas).
Pachychilus laevissimus var. varicosa Fischer and Crosse, 1892, Mission
Scientif. Mexique, Moll. Terr. Fluv., 2. Pt. 13, p. 329; PI. 53, fig. 6
(Palenque, Chiapas).
Monte Libano, 600 m. ; El Censo, 700 m. ; Ocosingo, 850 m. ; El
Real, 600 m. Also many dead shells, found with bones in a rock
214 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
shelter near Laguna Ocotal, and others from gopher diggings
nearby. No specimens were found alive either in the Laguna or
in the streams in the vicinity. The specimens were compared
with topotypes collected by Dr. L. Mazotti.
A common freshwater snail in the smaller streams and rivers
of Chiapas, Tabasco, Oaxaca, and of eastern and northern
Guatemala (Peten, Alta Vera Paz).
PLANORBIDAE
Helisoma caribaeum (d'Orbigny)
Planorbis caribaeus d'Orbigny, 18-41, in de la Sagra, Hist. Fis. Pol. Cuba,
Moll., Pt. 1, p. 103; PI. 13, figs. 17-19 (Havana, Cuba; and Veracruz,
Mexico).
Planorbis ancylostomus var. chiapasensis Fischer and Crosse, 1880, Mission
Scientif. Mexique, Moll. Terr. Fluv., 2. Pt. 8, p. 63; PI. 3-4, figs. 5 and
5a-b (Chiapas).
Laguna Ocotal, 950 m. ; Rio Amarillo at the Sumidero near
Las Casas (Mrs. L. Whitaker).
A common species throughout Central America, from Vera-
cruz to Panama, and in the Antilles, from Cuba to Barbados.
Tropicorbis obstructus (Morelet)
Planorbis obstructus Morelet, 1849, Test. Noviss. Ins. Cub. Amer. Centr., 1,
p. 17 (Island of Carmen, Campeche).
Planorbula obstructa Fischer and Crosse, 1880, Mission Scientif. Mexique,
Moll. Terr. Fluv., 2. p. 78; PI. 33, figs. 8 and Sa-d (type from More-
let) ; PI. 34, figs. 7 and 7o-c (var. berendti Tryon from Orizaba or
Veracruz).
Laguna Ocotal to El Censo ; a few dead, but fairly fresh speci-
mens.
Definitely known from southern Mexico ( Veracruz ; Campeche ;
Oaxaca; Yucatan; Chiapas), Guatemala and British Honduras:
The species possibly extends to southern Texas.
Taphius subpronus (von Martens)
Planorbis (Taphiua) subpronus von Martens, 1899, Biologia Centr.- Amer..
Terr. Fluv. Moll., p. 396; PI. 21, 4 figs. 15 (Amatitan, State of Tabasco,
Mexico).
BIOLOGICAL INVESTIGATION'S IN CHIAPAS, MEXICO 215
? Taphius subpronus F. C. Baker, 19-45, Molluscan Family Planorbidae,
p. 79; PI. 131, figs. 36-40 (Turrialba, Costa Rica; U.S.N. M. No.
162827).
Laguna Ocotal, 950 m., many dead specimens in a silt deposit
on the shore. I have also seen some fresh specimens of this re-
markable snail from Lake Coatepeque, El Salvador (N. C. Fassett
Coll. — Received through Dr. H. "W. Levi).
Von Martens commented upon the close similarity of subpronus
and Taphius pronus von Martens of Lake Valencia, Venezuela.
It is, indeed, impossible to point out a reliable difference in the
general shape, the method of coiling, the degree of deflection of
the aperture, the shape of the aperture and the relative width
and depth of the umbilicus, particularly as T. pronus varies
greatly in all these characters. There remains only the fine spiral
striation of the shell, present in fresh specimens of T. pronus.
There is no trace of this, neither on the weathered specimens
from Laguna Ocotal (where they might be worn), nor on the
very fresh specimens from Lake Coatepeque. The largest speci-
men from Laguna Ocotal is 8 mm. in greatest width and 2.8 mm.
thick. The largest of the 5 specimens from Lake Coatepeque is
4.5 mm. in greatest width and 2 mm. thick, approximating von
Martens' original measurements of 5 mm. and 2 mm.
I am not fully satisfied that F. C. Baker's figures, cited above,
actually represent true T. subpronus.
ANCYLIDAE
Ferrisia excentrica (Morelet)
Ancylus excentricus Morelet, 1851, Test. Noviss. Ins. Cub. Amer. Centr., 2.
p. 17 (Lake Itza [= Peten], Guatemala).
Ancylus (Ancylastrum) excentricus Fischer and Crosse, 1880, Mission
Scientif. Mexique, Moll. Terr. Fluv., 2, Pt. 7, p. 37; PI. 30, figs. 16-16a
(type from Morelet).
Laevapex excentricus B. Walker, 1924, The Ancylidae of South Africa, p. 10.
Ferrisia (Laevapex) excentrica Goodrich and Van der Schalie, 1937, Mus.
Zool. Univ. Michigan, Misc. Publ. No. 34, p. 34.
Laguna Ocotal, 950 m., several living specimens in floating
vegetation.
Known from southern Texas to Costa Rica.
216 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
SUBULINIDAE
Lamellaxis exiguus (von Martens)
L.eptinaria exigua von Martens, 1898, Biologia Centr.-Amer., Terr. Fluv.
Moll., p. 318; PL 18, fig. 10 (Teapa in Tabasco).
Laguna Ocotal, 950 m. ; Ocosingo, 1000 m.
As von Martens suspected, the original specimens, 5 mm. long,
were immature. The lot from Laguna Ocotal comprises three
immature shells agreeing with the original description and figure,
and one adult, 11.2 mm. long, 5.5 mm. in greatest width, the
aperture 5 mm. by 3 mm., of 6y2 whorls; the body-whorl is 7 mm.
long in front view. The spaced, costulate sculpture of the earlier
(post-nepionic) whorls changes gradually to close, finer vertical
striae on the later whorls ; the striation is replaced by exceedingly
fine vertical engraved lines below the periphery of the body-
whorl; the first 2 (nepionic) whorls are smooth. In the full-
grown shell the columella is shaped about as originally drawn
for the young, but the median fold is slightly more pronounced.
The outer columellar area is broad and its margin is spread over
the wide and deep umbilicus.
The species is known only from Tabasco and Chiapas.
Synopeas beckianum (Pfeiffer)
Bulimus beckianus Pfeiffer, 1846, Syinbolae Hist. Helic, 3, p. 82 (Opara I.
?); 1848, Monogr. Helie, Viv., 2, p. 164; 1854, Syst. Conch.-Cab., 1.
Abt. 13, Pt. 1, p. 125; PI. 30, tigs. 29-31 (type).
Opeas micro, von Martens, 1898, Biologia Centr.-Amer., Terr. Fluv. Moll.,
p. 294; PI. 17, figs. 10-11. Not of d'Orbigny, 1835.
Opens beckianum Pilsbry, 1906, Man. Conch., (2), 18, p. 189; PI. 27, figs.
42-46 and 54-55.
Ocosingo, 850 m. ; Laguna Ocotal, 950 m. ; Monte Libano to El
Censo, 600-700 m.
Widely distributed throughout tropical America, from Vera-
cruz to Sao Paulo, Brazil, and Peru, as well as throughout the
Antilles. It has possibly been spread by man. Pfeiffer 's original
locality appears to have been erroneous.
The species is evidently not a true Opeas and I have followed
H. B. Baker (1927, Occ. Papers Mus. Zool., Univ. Michigan, No.
182, p. 7) in placing it in Synopeas Jousseaume (1899). This
generic name appears to be antedated, however, by Synopeas
Foerster (1856) and a substitute may have to be proposed.
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 217
OLEACINIDAE
Spiraxis scalariopsis (Morelet)
Buiimus scalariopsis Morelet, 1851, Test. Noviss. Ins. Cub. Amer. Centr., 2.
p. 11 (Peten, Guatemala).
Spiraxis scalariopsis Fischer and Crosse, 1877, Mission Scientif. Mexique,
Moll. Terr. Fluv., 1. Pt. 6, p. 609; PI. 25, figs. 1 and la-b (type from
Morelet 's collection).
Monte Libano to El Censo, 600-700 m.
Known only from Chiapas and Guatemala (Peten).
Spiraxis similaris (Strebel)
Volutaxis similaris Strebel, 1882, Beitr. Mexikan. Land- Siisswasser-Conch.,
5. p. 122; PI. 7, fig. 11; PI. 17, fig. 18 (Veracruz: Pacho near Jalapa).
Laguna Ocotal to El Censo, 600-700 m., on Philodendron.
Known only from Veracruz and Chiapas.
Streptostyla chiapensis Pilsbry
Spiraxis parvula Pfeiffer, 1856 (December), Malakoz. Blatt., 3. p. 234
(Chiapas); 1857 (May), Proc. Zool. Soc. London, (for 1856), p. 379
(Chiapas). Not Achatina parvula Chitty, 1853, now placed rather
doubtfully in Spiraxis.
Streptostyla limnaeiformis var. parvula von Martens, 1892, Biologia Centr.-
Amer., Terr. Fluv. Moll., p. 100; PI. 5, fig. 24 (specimen from Chiapas,
in Pfeiffer 's Coll., but probably not the type).
Streptostyla limneiformis chiapensis Pilsbry, 1909, Man. Conch., (2), 20.
p, 111 (new name for Spiraxis parvula Pfeiffer, 1856).
Laguna Ocotal, 950 m.
The specimen collected at Laguna Ocotal, 6.5 mm. long, shows
the deep lines of growth mentioned by von Martens. I regard
chiapensis as specifically distinct from S. limneiformis (Shuttle-
worth, 1852). Streptostyla oblonga (Pfeiffer, 1856) differs from
chiapensis iu the smooth surface of the shell.
Known only from Chiapas.
Euglandina monilifera (Pfeiffer)
Glandina monilifera Pfeiffer, 1845 (October), Proc. Zool. Soc. London, p.
75 (Coban, "Veracruz" [error for Vera Paz, Guatemala]).
Achatina monilifera Keeve, 1849, Conch. Icon., 5, Achatina, PI. 14, fig. 50
(Coban; from Dennison Coll.).
218 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
Monte Libano, 600 m. ; Monte Libano to El Censo, 600-700 m.
Known from Veracruz, Guerrero, Chiapas, Guatemala ( Peten ;
Alta Vera Paz), and Costa Rica.
EUGLANDINA GHIESBREGHTI (Pfeiffer)
Oleacina ghiesbregliti Pfeiffer, 1856 (December), Malakoz. BlJitt., 3, p. 235
(Chiapas).
Achatina (Oleacina) ghiesbregliti Pfeiffer, 1857 (May), Proc. Zool. Soc.
London, (for 1856), p. 379 (Chiapas).
Glandinu ghiesbregliti Strebel, 1875, Beitr. Mexikan. Land- Siisswasser-
Conch., 2. p. 39; PI. 10, figs. 31 and 31a-c7 (possibly 2 paratypes?).
Lagima Ocotal, 950 m. ; El Censo to Laguna Ocotal, 700-
1000 m.
This species was known thus far from Chiapas and Ta-
basco ; but I have seen a specimen from Yepocapa, Dept. Chimal-
tenango, Guatemala (H.T. Dalmat Coll.), some 125 km. east of
Chiapas.
SAGDIDAE
Thysanophora impura (Pfeiffer)
Helix impura L'feiffer, 1866, Malakoz. Blatt., 13, p. 79 (Mirador, A'era-
cruz).
Thysanophora impura Strebel, 1880, Beitr. Mexikan. Land- Siisswasser-
Conch., 4. p. 30; PI. 4, 3 figs. 2 (Mirador, topotype; not Pfeiffer 's
holotype, which was never figured). Pilsbry, 1926, Proc. Acad. Nat.
Sc-i. Philadelphia, 78. p. 121, figs. 36A-B (Veracruz: Antigua; Pacho ;
Veracruz. Yucatan: Tekanta; Tunkas; Merida). Thiele, 1931, Handb.
Syst. Weichtierk., 2, p. 582, fig. 664.
Ocosingo, 850 m.
Known definitely at present from southeastern Mexico (Vera-
cruz, Chiapas and Yucatan). Published records from elsewhere
are open to question because the species has often been confused
with T. conspurcatella (Morelet, 1851).
Thysanophora pilsbryi H. B. Baker
Thysanophora pilsbryi H. B. Baker, 1922, Occ. Papers Mas. Zool., Univ.
Michigan, No. 106, p. 54; PI. 17, figs. 11-14 (Veracruz: La Laja near
the Hacienda de Cuatotolapan i.
Laguna Ocotal, 950 m.
Known only from Veracruz and Chiapas.
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 219
Tiiysanopiiora fuscula (C. B. Adams)
Helix fuscula C. B. Adams, 1849, Contributions to Conchology, No. 2. p. 35
(Jamaica).
Thysanophora fuscula Pilsbry, 1920, The Nautilus, 33. Pt. 3, p. 94, 2 figs. 1
(on p. 93, after a cotype; synonymizes with it T. fischeri Pilsbry,
1904).
Thysanophora fischeri Pilsbry, 1904 (January 30), Proc. Acad. Nat. Sci.
Philadelphia, (for 1903), p. 763; PI. 49, figs. 6-6a (Tamaulipas: 4
miles west of Victoria).
Laguna Ocotal, !)50 m. ; Monte Libano, 600 m.
Known from Tamaulipas, Veracruz, Chiapas and Jamaica.
ZONITIDAE
Habroconus trochulinus (Morelet)
Helix trochulinus Morelet, 1851, Test. Noviss. Ins. Cub. Amer. Centr., 2,
p. 10 ("H. non frecpiens in sylvas Petenenses, circa Sancti-Ludovici
pagum;" [=San Luis, Peten, probably the locality of that name in
about 16°15'N., 89°25'W.] ).
Guppya trodhulina von Martens, 1892, Biologia Centr. -Amer., Terr. Fluv.
Moll., p. 120; PI. 6, figs. 17 and lla-d (Morelet 's type, "the only
specimen which still exists in his collection").
Habroconus trochulinus H. B. Baker, 1930, Oce. Papers Mus. Zool., Univ.
Michigan, No. 220, p. 22; PI. 7, figs. 10-11 (Veracruz: Penuela to
Sumidero, 2625-3400 ft.; Necaxa, 2215-492.") ft.; "common at lower
altitudes on leaves of shrubs and trees, quite arboreal").
Laguna Ocotal to El Censo, 700-1000 m., on Philodendron; El
Real, 600 m.
The specimens from Chiapas agree with H. B. Baker's inter-
pretation of Morelet "s II. trochulina, which fortunately is in
accord with Morelet 's type as figured by von Martens. That
figure shows the body-whorl even more angulate at the periphery
than figured by Baker.
Von Martens concluded that Helix selenkai Pfeiffer (1866)
could not be separated from H. trochulinus, whereas H. B. Baker
regards them as distinct (although recognizing that they may
only represent two ecological forms). The material before me
is too small to decide the matter. A lot of 27 specimens, from
the T. Bland Collection, collected at Mirador, Veracruz, by
Berendt, who distributed them as //. selenkai, appear to be all
220 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
H. trochulinus; but the carina at the periphery varies from
very strongly marked (more so than drawn by von Martens)
to barely indicated, with all transitional stages between these
two extremes.
If restricted as H. B. Baker does, //. trochulinus is known
definitely only from Veracruz, Chiapas and Peten.
Omphalina bilineata (Pfeiffer)
Helix bilineata Pfeiffer, 1846 (February), Proc. Zool. Soc. London, (for
1845), p. 128 ("locality unknown"); 1852, Syst. Coneh.-Cab., 1. Abt.
12, Pt. 2, p. 96; PI. 83, figs. 23-25 (type; no locality).
Hyalina (Zonyalina) bilineata Pfeiffer, 1865, Malakoz. Blatt., 12. p. 16
(Veracruz).
Omplwlina bilineata von Martens, 1892, Biologia Centr.-Amer., Terr. Fluv.
Moll., p. 109; PI. 6, fig. 6 (several localities for the typical form and
the varieties, all in the State Veracruz).
Mesomphix {Zonyalina) bilineatus H. B. Baker, 1930, Occ. Papers Mus.
Zool., Univ. Michigan, No. 220. p. 28; PI. 9, figs. 2-4 (anatomy; 2
localities in Veracruz).
Omphalina (Zonyalina) bilineata Thiele, 1931, Handb. Syst. Weichtierk.,
2. p. 590.
Laguna Ocotal, 950 m.
Known thus far only from Veracruz and now recorded also
from Chiapas.
BULIMULIDAE
Bulimulus unicolor (Sowerby)
Bidinus unicolor SoAverby, 1833 (July 12), Conchol. Ulustr., Pt. 34, PI. of
Balinus, fig. 43 (with name in accompanying printed list; Panama);
1833 (September 26), Proc. Zool. Soc. London, p. 73 (Island of Perico,
Gulf of Panama).
Bulimulus unicolor Pilsbry, 1897, Man. Conch., (2), 11, p. 53; PI. 10, fig. 73.
Ocosingo, 1000 m.
Widespread in Central America, from Tabasco and Chiapas
to Panama.
Drymaeus moricandi (Pfeiffer)
Bulimus moricandi Pfeiffer, 1847 (January), Proc. Zool. Soc. London, (for
1846), p. 113 (Coban. [Guatemala]). Reeve, 1848, Conch. Icon., 5,
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 221
Bulimus, PI. 45, fig. 283 ("Central America;" ? type from Cuming
Coll.).
Bulimulus {Drymaeus) moricandi Fischer and Crosse, 1875, Mission Scientif.
Mexique, Moll. Terr. Fluv., 1. Pt. 5, p. 497; PI. 24, figs. 9-9a (Guate-
mala: Coban; Vera Paz; Duenas; with var. hyalino-albida, p. 498, from
Chiapas ) .
Drymaeus moricandi Pilsbry, 1899, Man. Conch., (2), 12. p. 78; PI. 4, figs.
62 (after Reeve) and (13-64 (after Fischer and Crosse).
El Censo to Laguna Ocotal, 700-1000 m.
The five mostly adult specimens collected belong to the var.
hyalino-albida Fischer and Crosse (1875), being clear whitish,
although very fresh. Apart from color, they agree not only with
the published figures of Reeve and of Fischer and Crosse, but
also with specimens from the Bland Collection (now at M.C.Z.),
labeled Guatemala.
There has been some doubt about the specific distinctness of
D. moricandi and Drymaeus sulphureus (Pfeiffer, 1857), prob-
ably because of the similarity in color (both species having a
pure white and a citron-yellow phase) and the fact that they
may occur together in Guatemala. In the series I have compared,
full-grown moricandi differs consistently in being broader at
the body-whorl, with a wider spire, in the longer aperture (which
reaches at least half of the total length of the shell), in a rela-
tively wider columellar area, and in a more open and perforate
umbilicus. Immature shells are, however, difficult to separate.
The largest shell seen from Chiapas is 29 mm. long, 14 mm. in
greatest width, the aperture 17 mm. by 9.5 mm.
D. moricandi is only known with certainty from Chiapas and
the adjoining northeastern section of Guatemala. D. sulphureus
is more widely distributed from Veracruz to Costa Rica.
Simpulopsis Simula (Morelet)
Bulimus simulus Morelet, 1851, Test. Noviss. Ins. Cub. Amer. Centr., 2.
p. 11 (Peten, Guatemala).
Simpulopsis Simula Fischer and Crosse, 1877, Mission Scientif. Mexique.
Moll. Terr. Fluv.. 1, Pt. 6, p. 578; PL 24, figs. 1313a (type from
Morelet).
Laguna Ocotal to El Censo, 700-1000 m., on Philodendron.
Known only from northern Guatemala (Peten) and Chiapas.
This appears to be the second record for the species, which is
222 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
probably overlooked because of its extreme fragility and its
habitat on leaves of epiphytic plants.
Orthalicus princeps (Broderip)
Bulinus princeps Broderip, 1833 (May 3), Conchol. Illustr., Pt. 27, PI. of
Bulinus, 2 figs. 18 (with name in printed list; "Conchagua, Central
America " [El Salvador] ) .
Oxystyla princeps Pilsbry, 1899, Man. Conch., (2), 12. p. 113; PL 16, figs.
1-9; PI. 17, figs. 10-12.
Monte Libano, 600 m. ; Monte Libano to El Censo, 600-700 m. ;
San Lorenzo, midway between Ocosingo and El Real.
Known from southern Mexico (Veracruz and Sinaloa) to
Panama.
TJROCOPTIDAE
Eucalodium mexicanum (Pfeiffer)
Cylindrella mexicana "Cuming" Pfeiffer, 1860 (February- June), Proc.
Zool. Soc. London, p. 139 ("Mexico").
Eucalodium (Eucalodium) mexicanum Pilsbry, 1902, Man. Conch., (2), 15,
p. 6; PI. 1, figs. 2-3; PI. 7, figs. 8-10.
Monte Libano, 600 m.
Known only from Chiapas and Tabasco.
Coelocentrum tomacella (Morelet)
Cylindrella tomaoella Morelet, 1849, Test. Noviss. Ins. Cub. Amer. Centr.,
l.p. 10 (Tabasco).
Coelocentrum tomacella Fischer and Crosse, 1873, Mission Scientif. Mexique,
Moll. Terr. Fluv., 1, Pt. 3, p. 342; PI. 15, fig. 11 (type from Morelet;
Tabasco and Palenque in Chiapas).
Laguna Ocotal, 950 m. ; Monte Libano to El Censo, 600-700 m.
Known from Tabasco, Chiapas and eastern Guatemala
(Coban).
Microceramus concisus (Morelet)
Cylindrella concisa Morelet, 1849, Test. Noviss. Ins. Cub. Amer. Centr., L
p. 12 (Yucatan).
Macroceramus concisus Fischer and Crosse, 1873, Mission Scientif. Mexique,
Moll. Terr. Fluv., 1. Pt. 4, p. 421; PI. 18, figs. 1 and la-b (type from
Morelet).
Microceramus concisus Pilsbry, 1903, Man. Conch., (2), 16. p. 155; PI. 25,
figs. 7-12.
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 223
Laguna Ocotal, 950 m. ; Ocosingo, 1000 m.
Known from Chiapas, Yucatan, Guatemala, Utilla I. off the
coast of Honduras, and Costa Rica. Most probably M. concisus
is not specifically distinct from M. gossei (Pfeiffer, 1846), from
Jamaica and (probably) Cuba, the Bahamas, and Hispaniola.
CEPOLIDAE
Leptarionta trigonostoma (Pfeiffer)
Helix trigonostoma Pfeiffer, 1844 (October), in Philippi, Abbild. Beschr.
Conchyl., 1. Pt. 7, p. 154 (or p. 24) ; PI. 5 [mislabeled 4] of Helix, 2
figs. 8 ("provincia Honduras Americae centralis"); 1845 (August),
Proe. Zool. Soc. London, p. 41 ("Veracruz, Province of Honduras,
Central America;" [a fictitious, truly Cumingian locality]).
llclix (Geotrochus) trigonostoma Fischer and Crosse, 1872, Mission
Scientif . Mexique, Moll. Terr. Fluv., 1. Pt. 2, p. 291 ; PI. 11, figs. 6 and
6a-d (Guatemala: Peten; Vera Paz; San Augustin; Sierra del Mico,
near Izabal).
Helix (Oxyehona) trigonostomu Pilsbry, 1889, Man. Conch., (2) 5, p. 132;
PI. 14, figs. 1-4; PI. 18, figs. 1-2. von Martens, 1892, Biologia Centr.-
Amer., Terr. Fluv. Moll., p. 154; PI. 9, figs. 1, la, 9, 9a, 11 and 12,
(additional localities in Guatemala: Senahu, N. side of Polochic
Valley, above Panzas; Vera Paz; Coban; near Gua^mala City, 5000
ft.; San Juan Eiv. ; Cerro Zunil, Pacific Slope near Quezaltenango ;
El Reposo between Retalhuleu and the Pacific ; slope of Cordillera, at
2500-4500 ft., at Hacienda San Francisco, Miramar and Helvetia, Buena-
vista).
Oxyehona trigonostoma Pilsbry, 1894, Man. Conch., (2), 9, p. 190.
Leptarionta trigonostoma Pilsbry, 1897, The Nautilus, 11, No. 8, p. 88.
Laguna Ocotal, 950 m. ; El Censo to Laguna Ocotal, 700-1000
m. ; Monte Libano to El Censo, 600-700 m.
The species is now for the first time recorded from reliable
Mexican localities, in Chiapas. The supposed occurrence in
"Veracruz" has never been confirmed and was evidently one
more of the many erroneous localities in the Cuming Collection.
The record from ' ' Honduras ' ' is likewise based on an error from
the same source. L. trigonostoma is restricted to Guatemala and
Chiapas, so far as known at present. Dr. H. T. Dalmat col-
lected specimens in Guatemala at Yepocapa, Dept. Chimalten-
ango, and at the Finca Montequina, Atitlan, Dept. Solola.
224 BULLETIN :' MUSEUM OF COMPARATIVE ZOOLOGY
The species varies somewhat, not only in the banding, but also
in the shape and slope of the spire. I agree with Pilsbry (1899)
that it is scarcely useful to distinguish these variants by names,
except as collector's items.
Averellia coactiliata (Deshayes)
Helix coactiliata "Ferussac" Deshayes, 1839, in Ferussac, Hist. Nat. Gen.
Part. Moll. Terr. Fluv., 1. p. 19; PI. 75, figs. 1-5 (Real-Llejos, Nicar-
agua; and "environs de Touspan, au Perou" [error for Tuxpan,
Veracruz, Mexico] ).
Helix (Patula) coactiliata Fischer and Crosse, 1872, Mission Scientif.
Mexique, Moll. Terr. Fluv., 1, Pt. 2, p. 234.
Helix (Patula, Discus, Trichodiscus) coactiliata Pilsbry, 1887, Man. Conch.?
(2), 3. p. 49; PI. 5, fig. 2.
Helix (Triclwdisoina) coactiliata von Martens, 1892, Pdologia Centr.-Amer.,
Terr. Fluv. Moll., p. 133.
Epiphragmophora (Trichodiscina) coactiliata Pilsbry, 1894, Man. Conch..
(2), 9. p. 199.
Averellia (Triehodwcina) coactiliata Thiele, 1931, Handb. Syst. Weichtierk.,
2. p. 698.
As neither the original description nor the figures were written
or published by Ferussac (who died in 1836), the name should be
credited to Deshayes.
Laguna Ocotal, 950 m. ; El Real, 600 m.
A widely distributed snail, perhaps transported sometimes by
man. It is known from Tamaulipas, San Luis Potosi, Michoacan,
Veracruz, Tabasco, Chiapas, Yucatan, Guatemala, British Hon-
duras, Nicaragua, Panama, Venezuela and Trinidad.
I am unable to separate Helix cordovana Pfeiffer (1857) from
A. coactiliata.
Additional Species Recorded From Chiapas
The following list of species known from Chiapas, but not
mentioned in the preceding pages, is based on published records.
as well as on specimens collected by Dr. R. A. Paynter, Jr. and
his associates, and more recently (1956) by Mrs. L. Whitaker,
outside the Lacandona area. Precise localities are mentioned
whenever available ; but some species have been recorded thus
far merely from "Chiapas." Species of doubtful occurrence or
identification have been omitted. The general distribution out-
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 225
side the State has been added. Asterisks mark the species which
I have seen from Chiapas.
HeUcina ghiesbreghti Pfeiffer, 1856. Chiapas. Known also
from Tabasco, Guanajuato and Guatemala.
Tamocyclus gealei Crosse and Fischer, 1872. Chiapas. Known
also from Guatemala.
Amphicyclotus texturatus (Sowerby, 1850). Chiapas: Chiqui-
huite, 6200 ft.; Escuintla, Sonconusco. Known also from Vera-
cruz and Guatemala.
Amphicyclotus megaplanus Morrison, 1955. Chiapas: El
Ocote, S. of Ocozocoautla.
*Choanopoma chiapasense Crosse and Fischer, 1877, Chiapas.
*Choanopoma sumiclirasti Crosse and Fischer, 1874. Chiapas:
El Sumidero near Tuxtla-Gutierrez, 1300 m. (R. A. Paynter, Jr.) .
Chondropoma vespertinum (Morelet, 1851). Chiapas: Palen-
que.
*Pachychilus chrysalis (Brot, 1872) (= Melania larvata Brot,
1877). Chiapas: San Pedro in the Cerro de la Gineta; Ixta-
comitan. Reported also from Nicaragua by von Martens.
*Pachychilus pyramidalis (Morelet, 1849). Chiapas: Meyapoc
near Ocozocoautla, 1000 m. (R. A. Paynter, Jr.) ; Rancho El
Eden, 2 miles from Ocozocoautla (Mrs. L. Whitaker) ; Palenque.
Also the var. maximus (Lea, 1851) from Lake Tzibal, about 50
miles west of Tenosique (D. W. Amram, Jr.). The species is
known also from Tabasco, Guatemala and the Republic of Hon-
duras.
*Physa berendti Strebel, 1874. Chiapas: Rio Amarillo at the
Sumidero near Las Casas (Mrs. L. Whitaker). Known also
from Puebla, Jalisco, Mexico City, Veracruz, Tabasco and
Oaxaca.
Vaginida moreleti (Crosse and Fischer, 1872). Chiapas: Pa-
lenque.
*Succinea brevis Pfeiffer, 1850. Chiapas: El Real, 600 m.
(R. A. Paynter, Jr.). Known also from Mexico City and the
State of Hidalgo.
Lamellaxis gracilis (Hutton, 1834). Chiapas. World-wide in
the Tropics ; spread by man.
Lamellaxis (?) semistriatus (Morelet, 1851). Chiapas: Palen-
que.
Pseudosubulina (?) chiapensis (Pfeiffer, 1856). Chiapas.
226 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Pseudosubulina (?) trypanodes (Pfeiffer, 1856). Chiapas.
*Spiraxis nitidus (Strebel, 1882). Chiapas: El Sumidero near
Tuxtla-Gutierrez, 1300 m. (R. A. Paynter, Jr.). Known also from
Veracruz.
Spiraxis sulciferus (Morelet, 1851). Chiapas: Palenque.
Known also from Veracruz and Guatemala.
Salasiella pulchella (Pfeiffer, 1856). Chiapas. Known also
from Costa Rica.
Streptostyla dubia (Pfeiffer, 1856). Chiapas.
^Streptostyla oblonga (Pfeiffer, 1856). Chiapas: El Sumidero
near Tuxtla-Gutierrez, 1300 mi. (R. A. Paynter, Jr.).
^Streptostyla streptostyla (Pfeiffer, 1846). Chiapas: El Su-
midero near Tuxtla-Gutierrez. 1300 m. (R. A. Paynter, Jr.).
Streptostyla irrigua var. shuttleworthi (Pfeiffer, 1857). Chia-
pas. Known also from Veracruz and Tabasco.
Streptostyla lurida (Shuttleworth, 1852) (= S. bocourti
Crosse and Fischer, 1868 ; S. lurida var. major von Martens,
1891). Chiapas. Known also from Veracruz, Tabasco, Guate-
mala and Costa Rica.
Streptostyla nebulosa Dall, 1896. Chiapas: San Cristobal.
Polygyra chiapensis (Pfeiffer, 1856). Chiapas.
*Polygyra yucatanea var. helictomphala (Pfeiffer. 1856).
Chiapas: El Real, 600 m. (R.A. Paynter, Jr.). Known also from
Guatemala.
Hawaiia minuscula (A. Binney, 1840). Chiapas: Palenque.
Nearly world-wide in the Tropics ; spread by man.
*Omphalina zonites (Pfeiffer, 1846). Chiapas: El Sumidero
near Tuxtla-Gutierrez (R. A. Paynter, Jr.).
Pseudohyalina cidariscus von Martens, 1892. Chiapas: Pa-
lenque.
Drymaeus chiapasensis (Pfeiffer, 1866) (— Otostomus chia-
pensis von Martens, 1893). Chiapas: Cumbre de Manzanilla.
Known also from Veracruz and Puebla.
Drymaeus recluzianus (Pfeiffer, 1847). Chiapas. Known also
from Costa Rica (var. martensianus Pilsbry, 1899).
* Drymaeus ghiesbreghti (Pfeiffer, 1866). Chiapas: the Sumi-
dero near Las Casas (Mrs. L. Whitaker). Known also from
Colima, Oaxaca and Guatemala.
*Eucalodium decollation var. ghiesbreghti (Pfeiffer, 1856).
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 227
Chiapas. Known also from Guatemala.
*Eucalodium walpoleanum Crosse and Fischer, 1872. Chia-
pas: Palenque. Known also from Guatemala.
Eucalodium sumichrasti Crosse and Fischer, 1878. Chiapas.
Coelocentrum attenuatum (Pfeiffer, 1856). Chiapas. Pos-
sibly only a variant of C. tomacella (Morelet).
Coelocentrum clava (Pfeiffer, 1856). Chiapas. Possibly only
a variant of C. tomacella (Morelet).
Coelocentrum nelsoni Dall, 1897. Chiapas: Tuxtla-Gutierrez.
Coelocentrum pfefferi Dall, 1897. Chiapas: Oeozocoautla,
1200 m.
Holospira oerendti (Pfeiffer, 1866). Chiapas. Known also
from Veracruz.
Epirobia oerendti (Pfeiffer, 1866). Chiapas.
Epirobia gassiei (Pfeiffer, 1867). Chiapas. Possibly not sep-
arable from E. berendti.
*Lysinoe ghiesbreghtii (Nyst, 1841). Chiapas: Zinacantan,
2000 m. (R.A. Paynter, Jr.) ; Rancho Nuevo, 8 miles from Las
Casas (Mrs. L. Whitaker) ; mountain above the Sumidero near
Las Casas (Mrs. L. Whitaker). Known also from Guatemala,
the Republic of Honduras, and El Salvador (Volcan de Santa
Ana)
*Xanthonyx chiapensis (Pfeiffer, 1856). Chiapas.
Elliptio sentigranosus (von dem Busch, 1845) (= TJnio corium
Reeve, 1864). Chiapas. Known also from Veracruz.
Elliptio (Nephronaias) calamitarum (Morelet, 1849) ; includ-
ing var. prolongata Fischer and Crosse, 1894; var. nephretica
Fischer and Crosse, 1894; and var. arcuana Fischer and Crosse,
1894. Chiapas : Baluntie River near Palenque.
Elliptio (Nephronaias) aeruginosas (Morelet, 1849). Chiapas:
Michol River near Palenque.
Anondontites banibousetarum (Morelet, 1851). Chiapas: Pa-
lenque.
228 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
IV
ANTS FROM LAGUNA OCOTAL (HYMENOPTERA:
FORMICIDAE)
By
William L. Brown, Jr.
The ants from Laguna Ocotal were collected for the most part
by Robert L. Dressier, and, unless otherwise indicated, the col-
lections were made from epiphytes, particularly bulbous-based
Tillandsia, growing in the pine forest or the adjacent tropical
evergreen forest. Among the 21 species represented in the col-
lection, 17 can be determined to species in accordance with
present-day classifications; the remainder belong to difficult
groups in need of revision, or else the sample is in some way
unsatisfactory for species determination, so that identification
is carried only to genus.
All of these species belong to the tropical American fauna,
and all are either widespread in South and Central America or
else range at least through Central America and extend north-
ward into Veracruz and neighboring Mexican states. Very few
ants have been recorded from Chiapas (see Brown, 1950, Was-
mann Jour. Biol., 8: 241-250), but with the present series we
have accumulated a sample sufficient to confirm the expected
close similarity of the Chiapas ant fauna to those of Guatemala
and Veracruz. A few of the records of ants received from E. 0.
Wilson, collected by him in Veracruz during 1953, are men-
tioned below where relevant. I also possess a small number of
Chiapas ants collected by C. J. Goodnight and L. J. Stannard
during the last five years, mostly from soil and leaf-litter ber-
lesates, including new species of dacetines and basicerotines that
will be described elsewhere. The soil and soil-cover samples are,
of course, quite different from the epigaeic-arboreal collections
reported below; on the forest floor, Wasmannia auropunctata
(Roger), small species of Pheidole and Solenopsis, Prionopelta
modesta Forel, and several of the smaller Dacetini are the com-
monest forms, present in nearly every Berlese sample taken,
while Pachycondyla harpax (Fabricius), Ponera nitidula Emery,
Ponera spp., and Brachymyrmex are rather frequent.
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 229
As is well known, some of the forms listed below have been
involved in considerable taxonomic uncertainty, due chiefly to
unrecognized synonymy. "Wherever such synonymy has become
obvious from the augmented samples now available in the Mu-
seum of Comparative Zoology, I have taken the minimum formal
steps necessary to list and justify it.
Platythyrea punctata P. Smith
From a nest in a fallen log, August 3. Winged forms were
present, the males being fully pigmented and apparently active,
while most of the females were still in the callow stage or were
not yet eclosed. Wilson found this ant foraging on tree trunks
after dark in Veracruz and Cuba; the nocturnal tree-climbing
habit seems characteristic of many members of tribe Platythy-
reini.
Typhlomyrmex rogenhoferi Mayr
Typhlomyrmex rogenhoferi Mayr, 1862, Yerh. zool.-bot. Ges. Wien. 12: 737,
worker. Type locality : Amazonas.
Typhlomyrmex rogenhoferi race robustus Emery, 1890, Bull. Soc. Ent. Ital.,
22: 40, worker. Type locality: Alajuela, Costa Rica. NEW SYN-
ONYMY.
Typhlomyrmex robustus subsp. manco Wheeler, 1925, Ark. f. Zool., 17A (8) :
2, worker. Type locality: Pablobamba, Peru. NEW SYNONYMY.
Prionopelta marthae Forel, 1909, Deutsch. ent. Zeitschr., p. 240, worker.
Synonymy by Brown, 1953, Psyche, 59: 104.
This species is very widespread in the forested regions of
tropical America, but a single female stray from a log is the
first sample so far recorded from Chiapas. Series from different
nests from many localities in the Museum of Comparative
Zoology show wide diversity in size and in allometric characters,
including relative head width, general robustness of body, and
sculpture. However, there is often considerable variation in these
characters within single nest series, and one particular series,
from San Juan Pueblo, Honduras, leg. W. M. Mann, contains
extremes of the variation as well as all intergrades ; bracketed are
the "diagnostic" characters for robustus and manco, as reviewed
for types and metatopotypes, or both, of these two variants
before me.
230 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Neoponera lineaticeps Mayr
A small colony of this rather uncommonly collected ant was
taken from a TUlandsia base, which is apparently a preferred
habitat. It has been taken in Veracruz and Costa Rica on several
occasions, but this is the first record from Chiapas. The specific
name derives from the peculiar coarse, regular longitudinal
striation covering the central part of the upper surface of the
head, a feature that will identify the species at a glance under
magnification.
Neoponera apicalis (Latreille)
Formica flavicornis Latreille, 1802, Hist. Nat. Fourmis, p. 202, pi. 7, figs.
42B, 43 (?), worker, female, nee Fabricius. NEW SYNONYMY.
Formica apicalis Latreille, 1802, ibid., p. 204, pi. 7, fig. 42A (?), worker.
Neoponera Latreillei Forel, 1905, Ann. Soc. Ent. Belg., 49: 161, nom. pro
N. flavicornis (Latreille). NEW SYNONYMY.
Neoponera apicalis var. verenae Forel, 1922, Eev. Suisse Zool., 30: 90,
worker. NEW SYNONYMY.
A single worker was taken foraging on an epiphyte. This
species and the closely related N. obscuricornis Emery have been
confused through the literature, and the confusion extends to
most of the ant collections rich in neotropical material to this
day. The outstanding differences between these two species as
I see them are : ( 1 ) N. apicalis has the five or six apical funicular
segments a bright, contrasting yellow, whereas in N. obscuri-
cornis, the apex of the funiculus is little or not at all lightened,
and does not form a sudden contrast with the rest of the antenna.
Faded or teneral specimens may seem to be intermediate, but
these are rare and are easily identified by the remaining charac-
ters. (2) N. apicalis has the sides of the petiolar node nearly
flat, scarcely or not at all concave or sulcate just next to the
posterolateral angles, so that these angles are blunt, whereas
in obscuricornis, the same angles are thrown into relief by a
slight but distinct sulcation extending along the posterior sides
of the node from top nearly to base. (3) Of the two species,
apicalis is slightly but distinctly larger on the average, though
there is some overlap in size between the two forms. (4) JV. api-
calis is more opaquely sculptured than is N. obscuricornis, though
both species are strongly opaque; direct comparison is really
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 231
needed to reveal the difference. A study of numerous nest
series, in addition to stray workers, convinces me that the fore-
going characters are consistently linked in one or the other
combination. No difficulty has arisen in assigning fresh worker
specimens to one or the other species, and no intergrades have
been seen, despite the fact that the two species frequently occur
in close proximity over a vast area reaching from the Amazon
Basin to southern Mexico. In the Museum of Comparative
Zoology, collections of both species at single localities have been
made as follows : Kartabo and Kamakusa, British Guiana, leg.
W. M. Wheeler; Barro Colorado I., Panama Canal Zone, leg. N.
Banks; Laguna Encantada, Veracruz, leg. Q. Jones and R. L.
Dressier ; Pueblo Nuevo, near Tetzonapa, and Las Hamacas, near
Santiago Tuxtla, both in Veracruz, leg. E. 0. Wilson. Both
species live in plant cavities in arboreal situations, but nothing
has been recorded concerning their ecological occurrence in any
detail.
While it seems clear enough that two and only two species
exist in this complex, the application of names to these entities
is still in some doubt. The earliest recognized description of a
member of the complex appeared when Latreille claimed to have
described two species at once, giving them the names flavicornis
and apicalis. The former name was supposed by Latreille to
apply to a Formica flavicornis earlier named by Fabricius, but
Fabricius' insect is apparently an attine species having nothing
to do with Neoponera; flavicornis is thereby a preoccupied name.
Nearly everything about Latreille 's characterization of flavi-
cornis and apicalis is either confused or patently in error, and
the confusion extends to the correspondence of the plate figures
with their respective descriptions. No reliable difference is
mentioned or shown by Latreille that will serve to separate the
two forms, and the description of the antennal coloration, if
nearly accurate, would indicate that both are referable to apicalis
in the present sense. This is my interpretation, made without
recourse to types, but a thorough examination of the original
references in conjunction with fairly good samples of the com-
plex shows that it is the simplest solution to a problem that
bothered Latreille, Emery, and Wheeler, among others, to the
point where the essentially simple species-to-species relationship
became lost to view. The name latreillei is an objective synonym
232 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
of flavicornis. I do not follow Emery's "Genera Insectorum"
assignment of latreillei as a variety of obscuricornis — an as-
signment which expressed his lack of confidence in latreillei as
a named entity in his characteristically mild, but in this case
totally confusing, fashion. The variety verenae was described
by Forel in his familiar "final melange" paper of 1922, in which
several other formicid variants, since synonymized by various
authors, were named on the basis of the most doubtful-appearing
evidence. Forel mentions no character that would serve to dis-
tinguish verenae from typical apicalis, and verenae comes from
the middle of the range of the species.
Neoponera unidentata (Mayr)
Pachycondyla unidentata Mayr, 1862, Yerh. zooh-bot. Ges. Wien, 12: 720,
worker, female.
Neoponera unidentata var. rugosula Emery, 1902, Rend. Accad. Sci. 1st.
Bologna, (n.s.) 6: 30, worker; variant spellings are "rugatula" of
Santschi and "rvgvlosa" of Wheeler. NEW SYNONYMY.
Neoponera unidentata, Wheeler, 1929, Zool. Anz., Wasmann-Festband, pp.
29-30, typical form, with the following varieties: var. eburneipes
Wheeler, p. 29, worker, female. NEW SYNONYMY, var. maya
Wheeler, p. 30, worker. NEW SYNONYMY, var. trinidadensis
Wheeler, p. 30, worker. NEW SYNONYMY.
Wheeler conceived the named variants above (plus also the
"subspecies" sulcatula Santschi, q.v. infra, which belongs in the
crenata, not the unidentata, complex) as "local varieties" based
on differences in color, sculpture, form of petiole, pilosity and
some lesser details. There is no doubt that variation exists in
these various features, and it is clear that both Emery and
Wheeler understood that the variation was graded from series
to series even in the limited samples examined by these authors.
It is also clear, from the present augmented sample drawn from
many parts of the species distribution, that the different charac-
ters do not vary according to the same geographic plan. The
region of the Upper Amazon Basin shows the strongest varia-
tion, especially in sculpture, and the range of the variation there
leaves little encouragement for racemakers. While it is possible
to trace some series to their general area of origin by the study
of trends in individual characters, other samples are ambiguous
or contradictory in the display of the same characters. Clearly,
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 233
a study of geographical variation by individual characters is
required before further attempts are made to classify the popula-
tions making up N. unidentata.
At the Laguna Ocotal collecting area, the species is character-
ized in general by a petiolar node a bit less thick from front to
rear, as viewed from the side, than in the average Amazonian
series. However, I am unable to separate some samples in the
Chiapas lot from some taken in the Amazon-Guianas region.
According to the locality and the characteristics of the most
extreme examples, I suppose the Chiapas series would fall under
Wheeler's concept of var. may a. This species is common in
bulbous-based Tillandsia at the Laguna, if Dressier 's collections
are a fair indication of relative abundance.
N. unidentata and N. crenata (Roger), and also N. carinulata
(Roger), range widely over tropical America. All three inhabit
plant cavities, and all are very similar in general habitus, but
the types of petiolar node formation are widely divergent.
Neoponera crenata (Roger)
Ponera crenata Roger, 1861, Berlin, eut. Zeitschr., 5: 3, female, nom. pro
Portera pallipes Fr. Smith, 1858, p. 98 nee p. 87.
The series from Laguna Ocotal (and most collected else-
where in southern Mexico) agree best with the form described
by Forel as X. stipitum, of which a cotype rests in the Museum
of Comparative Zoology. For the present, I am unable to find
any satisfactory characters to separate stipitum from crenata,
and I incline to the belief that a thorough study, with resort to
the scattered types of these and other named variants of the
complex, will see them all merged under the name crenata.
Meanwhile, we may tentatively associate various morphological
tendencies with the names attached to the several inadequate
descriptions involved.
The populations of which I have samples vary widely in size
from nest to nest, although intranidal variation is relatively re-
stricted. Weakly correlated with size is the shape of the petiolar
node ; this correlation holds best at the extremes of the size range,
but is poor in intergradient series. Larger specimens (ca. 11 to
12.5 mm. in TL, or total outstretched length of body, including
head and mandibles) have the posterior nodal face vertical
234 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
throughout, and distinct from the dorsal face, and the compound
eyes tend to be farther from the front of the sides of the head,
e.g., Kartabo, British Guiana, nos. 60, 425, leg. W. M. Wheeler.
These correspond to my idea of "typical" crenata (=pallipes
F. Smith preocc).
In smaller specimens, such as the Wheeler Kartabo series nos.
495, 708, 679, 286, 507, 671, 148 and 621, among many others,
TL averages only 6 to 8 mm. In these, the eyes may be closer
to the anterior cephalic margins and the posterior nodal face is
often (not always) more convex, with the surface curving con-
tinuously into the dorsal face; such specimens correspond to
stipitum Forel. It is worth noting that the female node, as usual
among ponerines, is higher and thinner in lateral-view profile
than in the workers from the same nest. This caste difference
appears to have caused some confusion in the complex in the
past.
A specimen from Espiritu Santo, Brazil (TL 8.8 mm.) and
some others away from the Kartabo locality appear to be transi-
tional between the large and small forms, but this would not
necessarily preclude the specific distinction between two closely
related forms where sympatric, as at Kartabo and elsewhere in
the Amazon-Orinoco Basins (perhaps a case of "character dis-
placement"). The intermediate forms seem to be the same as
moesta Mayr, the var. moesta of authors. The polynomials N.
crenata fiebrigi Forel, N. crenata confusa Santschi, N. crenata
confusa lata Santschi and N. unidentata sulcatula Santschi seem
from their descriptions to represent minor southern nest variants
in the small-to-medium size range of the crenata complex; these
names are almost sure to prove synonymous when properly
studied. In view of the insufficiency of my present sample
(though it is considerably better than exists in other collections
known to me) and the difficulty of seeing all the types involved,
I have left formal synonymy in this group to some future
worker.
ECITON BURCHELLI (WestWOOd)
Workers were taken from a raiding column near the Laguna
Ocotal camp. These specimens would undoubtedly be placed as
"race paririspinum" by Father Borgmeier; however, the head
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 235
of the largest major (soldier) in the series is dingy yellowish-
white in color.
PSEUDOMYRMEX PALLIDUS F. Smith
A few workers from a Tillandsia.
Pseudomyrmex gracilis (Fabricius)
This is the common bicolored form of the species often known
as var. or subsp. mexicanus. It may be that the bicolored form
is suppressed in northern South America where other bicolorous
species of similar size and appearance become common. The
geographical variation of this complex, while outstandingly con-
spicuous, has never been carefully and thoroughly investigated.
Pheidole punctatissima Mayr
Two colonies were taken in epiphytic plants.
Pheidole spp.
Two indeterminate species of Pheidole were taken in Tilland-
sia. One of these is in the confusing biconstricta group, and the
other is a much smaller species. Pheidole is one of the very
large (1,000-plus named forms) ant genera currently "out of
control" taxonomically. Until adequate revisions of these groups
become available, description of isolated new species only adds to
the confusion and the unrecognized synonymy. Possibly one half
of the names currently remaining unchallenged in Pheidole are
synonyms of older names, and identification of species with any
certainty is hopeless in most faunas, even where helpful pre-
liminary revisionary attempts have been published.
Procryptocerus scabriusculus Emery
A stray worker from foliage. E. 0. Wilson also took a worker
during 1953 at Las Hamacas, near Santiago Tuxtla, Veracruz.
This and the following two cephalotine species were determined
from revisionary papers on the cephalotines by W. W. Kempf.
The work of Father Kempf is refreshingly sound and useful as
236 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
compared to the majority of publications on neotropical ants
with which the would-be identifier has to grapple.
Paracryptocerus cristatus (Emery)
A stray soldier. E. 0. Wilson took a colony of this species at
Las Hamacas, Veracruz, on August 27, 1953, occupying three
internodes of a living Cecropia tree about 15 feet tall in tropical
evergreen forest.
Paracryptocerus scutulatus (F. Smith)
A soldier aud workers. The species is widespread from
southern Mexico to Venezuela.
Smithistruma epinotalis (Weber)
This little dacetine normally lives in plant cavities well above
the ground. The collection at Laguna Ocotal was made from a
Tillandsia, but collections from Veracruz, sent by N. L. H.
Krauss and by E. 0. Wilson, were taken from hollow twigs of
standing shrubs and trees.
Acromyrmex octospinosus (Reich)
Foraging workers of this fungus-growing ant stole rice from
the Laguna Ocotal camp. The species has been discussed at
length by W. M. Wheeler (1937, "Mosaics and other anomalies
among ants," Harvard Univ. Press. Cf. pp. 69-74), who detailed
the ambiguity of the characters supposed to separate it from
A. hystrix (Latreille). Wheeler suggested as a better separatory
character the presence or absence of bilateral tubercles or carinae
on the propodeal dorsum; however, Wheeler's own series of the
two forms in the Museum of Comparative Zoology appear to
grade through on this basis without a break.
The distinction of the "races" echinatior Forel, inti Wheeler,
volcanus Wheeler, ekchuah Wheeler and cubanus Wheeler seems
to me at least as precarious as the specific separation of hystrix
from octospinosus. The differences supposed to separate these
forms are weak and variable, and seem to mark mere individual
or nest varieties in some eases ; even if they prove later to follow
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 237
to some degree geographical trends, there seems little to be gained
by placing formal names upon these samples at this time. As
seems to be the case with a large number of the subspecies so
far described in the animal kingdom, these examples are based on
entirely inadequate samples from restricted localities, and in
their description scant thought seems to have been given to the
overaU trends in variation of the characters within the whole
species.
Azteca sp.
A small brownish form, represented by minor workers only.
Brachymyrmex obscurior Forel
Specimens from TiUandsia seem to agree best with descriptions
and other determined material of this species, though determina-
tions in this genus are doubtful in the absence of anything
better than Santschi's revision.
Nylanderia ?guatemalensis (Forel)
This slender yellowish form is usually placed as a subspecies
of vivid ul a, but since the taxonomy of this genus is so poorly
worked out, I feel that it is better to accord provisional spe-
cies rank to those names not obviously synonyms. The guati -
))iale7isis syntype in the Museum of Comparative Zoology is
badly damaged, rendering the comparison uncertain.
Camponotus circularis Mayr
Stray workers and a small colony or colony-fragment from
TiUandsia plants.
238 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
V
FISHES FROM LAGUNA OCOTAL
By
Robert Rush Miller1
Since no special collecting equipment was available, only a
very limited sample of the fish fauna of this lake was obtained.
Forty-seven specimens, representing four species in three fam-
ilies, were secured by dipnet, bent pin, and rifle (using .22 caliber
dust shot) between July 21 and August 14, 1954, by various
members of the expedition. Many of the examples are twisted,
with broken fins and missing scales, thus rendering a careful
study difficult or impossible. One viviparous species cannot be
identified with certainty to genus since no males were obtained.
Although two new species appear to be represented, no suitable
type material is available and further well-preserved series, in-
cluding both sexes, are needed to clarify their status. The fol-
lowing report, therefore, is necessarily of a preliminary nature.
The specimens listed below have been divided between the
Museum of Comparative Zoology and the Museum of Zoology
of the University of Michigan.
Despite the small size of the collection, a most interesting
fish fauna is indicated for Laguna Ocotal. Isolation no doubt has
been a potent factor in the differentiation of endemic species in
this remote area.
CHARACIDAE
Characins
Astyanax fasciatus (Cuvier). Banded tetra.
This is one of the widest ranging freshwater fishes of the
Americas. It is known from Argentina northward on the At-
lantic slope to western Texas and adjacent parts of New Mexico,
and on the Pacific slope from Colombia to western Mexico (Rio
Armeria basin of Colima and Jalisco).
i Museum of Zoology of the University of Michigan, Ann Arbor, Michigan.
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 239
There are 10 adults, 59 to 107 mm. in standard length. The
number of anal fin-rays varies as follows: 22(4), 23(4), and
24(2). Recognition of subspecies of A. fasciatus is currently
made largely on the basis of the anal-ray count. The above
specimens could be assigned to A. f. aeneus (Giinther) or they
might be interpreted as intergrades between that lowland form
and the highland subspecies, A. f. mexicanus (de Filippi). I
prefer to postpone assignment until a good series is available
from Laguna Ocotal, especially since these specimens have a
more oblique mandible and more posterior dorsal fin than speci-
mens of A. f. aeneus from the Usumacinta basin in Guatemala.
The following color notes were made in Ann Arbor on October
29, 1954. The fins of the three largest fish (98, 105 and 107 mm.
S.L.) are bright yellow-orange to deep red-orange as follows:
over seven-eighths of the pelvic fins (tips of rays clear), the
anteriormost 9 rays to all of the rays of the anal fin, the outer
one-half of the caudal rays (except 3 to 4 rays of each lobe, which
are colored their entire lengths), and the median part of the
pectoral rays (weakly colored). The dorsal fin is clear in one,
faintly yellowish on the distal half in another, and yellow-orange
on the same rays in the third. The seven smaller fish show weaker
color on these fins or none at all.
POECILIIDAE
Livebearers
PSEUDOXIPHOPHORUS BIMACULATUS (Heckel)
This species is known along the Atlantic slope of Middle
America from Veracruz, Mexico, to Miranda, Nicaragua ; its alti-
tudinal distribution is from near sea level to about 3,500 ft.
Twenty-three young to adult specimens, 16 to 49 mm. long, in-
cluding 3 mature males, were obtained. The scale crescents are
conspicuously blackened. Dorsal-ray counts are as follows: 13(3),
14(12), and 15(8). According to current practice, this sample
is assignable to P. b. taeniatus Regan (see Hubbs, 1935, Univ.
Mich. Misc. Publ. No. 28: 9-10, and references cited therein), a
subspecies known to range from Mexico to Honduras.
240 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Undetermined genus and species.
There are 2 adult females of a species of poeciliid which I
cannot identify with any known member of the family. In the
absence of males, T am uncertain to which genus the species
belongs. Reference to the Poeciliidae (rather than to some other
cyprinodont family) is assured from the nature of the first 3 rays
of the anal fin (unbranched) and of the neuromasts or pit organs
on the scale rows (see Hubbs and Miller, 1954, Zoologica, 39
(1):2).
The general body form is similar to that of Lucania parva
(Baird and Girard), a species of the oviparous family Cyprino-
dontidae, although it is more angulate anteriorly, both in dorsal
and ventral profiles. The dorsal fin is long, containing 12 rays
in each specimen; the anal fin has 9 rays. The origin of the
dorsal fin is midway between the base of the caudal fin and the
tip of the snout. There is a faint, dark line along the midside
of the body that fades out anteriorly, and the skin beneath the
anterior half of the exposed part of each scale on the back and
sides shows a concentration of coarse chromatophores. There are
no distinctive markings.
The mandible is short and nearly vertical, indicating a surface-
feeding habit. The teeth are distinctive : those in the outer row
of each jaw are large and asymmetrical (shaped like the tip of a
tableknife blade), and are tilted obliquely away from the center
towards the outer margin of the jaws; they are most strongly
oblique nearest the midpoint of each jaw, becoming almost erect
at the margins. There is a toothless gap at the midpoint in the
lower jaw. Inside of each outer row of teeth is an irregular
series of small, conical teeth.
The body shape, long dorsal fin, oblique mouth, and distinctive
dentition comprise characters which set this species apart from
any poeciliid known to me from Middle America. The jaw denti-
tion of Xiphophorus helleri (Heckel) is very similar and the
species in question may pertain to that genus ; it does not repre-
sent that species, however, which differs in coloration, mouth
width, a less oblique lower jaw, the more robust body, etc. The
dentition of the outer jaws is similar also in specimens of Phalli -
chthys pittieri (Meek), but the teeth of that species are more
loosely attached and more numerous and other marked differences
indicate no intimate relationship.
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 241
CICHLIDAE
Mo j arras
Cichlasoma species
Twelve specimens, 51 to 96 mm. long, represent a species of
(he "Section" (or subgenus) Parapetenia of Regan (1906, Biol-
ogia Centrali-Americana, Pisces, 8 : 26). However, I cannot place
the present form with any of those treated by Regan or by sub-
sequent authors. What remains of the color pattern is suggestive
of both C. salvini and C. iirophthalmus, but the Ocotal specimens
otherwise differ in many ways from both of those species. There
are rather definite to indistinct vertical to oblique bands along
the sides, numbering not more than 10 or 11, the anteriormost
2 to 3 extending obliquely forward across the nape (as in C.
nigrofasciatum) . An irregular, usually disrupted, lateral band
extends from the upper angle of the opercle, reaching backward
not farther than to below the middle of the soft dorsal fin. Some
specimens have a prominent black spot at the base of the upper
half of the caudal fin (and lying entirely above the posterior
extension of the lateral line) which is surrounded by a light
area, as in C. urophthalmus (see Giinther, 1868, Trans. Zool. Soc.
London, 6: PI. 72, fig. 1) ; this spot is indistinct or obsolete in
other specimens. The two largest fish are entirely black (a mel-
anistic phase?) and have a shorter pectoral fin (not reaching
origin of anal), but otherwise agree with the ten smaller fish.
In the latter, the pectoral fin extends to above or slightly beyond
the origin of the first anal spine. It is possible that the two black
specimens represent a different species, but this point cannot be
determined satisfactorily on the basis of the present material.
The spinous dorsal is long and low, comprising 18 spines in
9 and 19 in 3 specimens; the soft dorsal has 9 rays in all; the
anal spines are numerous : 8 in 7 and 9 in 5 ; and the soft rays of
the anal fin vary as follows: 6(1), 7(7), 8(3), and 9(1). A
narrow but definite frenum is present in each specimen and the
gillrakers (total number, including rudiments) number 9(1),
10(9), and 11(2). The premaxillary spines extend posteriorly
from about the front to the middle of the orbit.
The large number of dorsal and anal spines (with correspond-
ingly few soft rays), the body form, and the coloration appear
to be among the distinguishing characters of this species.
242 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
VI
REPTILES AND AMPHIBIANS FROM
THE SELVA LACANDONA
By
Benjamin Shreve
As this section of the report deals only with the reptiles and
amphibians collected during 1954 in the Selva Lacandona, it
is perhaps fitting to mention that Raymond A. Paynter, Jr. and
Robert L. Dressier collected reptiles and amphibians in other
parts of Chiapas and elsewhere in Mexico. Although not included
in this study, these are much appreciated.
Of those collected in Chiapas, a specimen of Stenorhina f.
freminvillii Dumeril, Bibron and Dumeril from Ocosingo, should
be mentioned as representing the first record of this race for the
state. It will be seen that of those species discussed in the
report, several are new to Mexico or to Chiapas.
It seems advisable to mention that the wholesale restriction of
type localities by Smith and Taylor (1950, Univ. Kansas Sci.
Bull., 33, pp. 313-380) is not followed here because of the numer-
ous instances of error and poor judgment, aside from being con-
sidered unnecessary. See comments of Dunn and Stuart (1951,
Copeia, p. 55; and 1951, Science, 113, p. 677).
Crocodylus moreletii Dumeril and Dumeril
Crooodylus Moreletii Dumeril and Dumeril, 1851, Cat. Meth. Kept., p. 28;
Lake Peten, Guatemala.
imm. $ (M.C.Z. 53860) Laguna Oeotal, Aug. 12.
cranium (M.C.Z. 53903) Laguna Oeotal, July-Aug.
The remains of what appears to be an immature Pseudemys
script a ornata (Gray) were found in the stomach of M.C.Z.
53860, now a made up skin.
Kinosternon leucostomum Dumeril and Dumeril
Cinosternon Leucostomum Dumeril and Dumeril, 1851, Cat. Meth. Eept., j>.
17; Mexico, etc.
1 (M.C.Z. 53861) Laguna Oeotal, July-Aug.
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 243
Anolis tropidonotus spilorhipis Alvarez del Toro and Smith
Anolis tropidonotus spilorhipis Alvarez del Toro and Smith, 1956, Herpetol-
ogica, 12. p. 9: Cerro Ombligo, 1280 m., Chiapas, Mexico.
6 (M.C.Z. 53855-7) Monte Libano, July 16-18.
4 (M.C.Z. 53858-9) Laguna Oeotal, July- Aug.
11 (M.C.Z. 53887-91) Laguna Oeotal to El Censo, Aug. 20.
8 (M.C.Z. 53894-7) El Censo to Monte Libano, Aug. 21.
On comparing this series with one from near El Potrero,
Veracruz, identified as tropidonotus, I find that the former has
a differently colored dewlap, smaller ventrals as compared with
the dorsals, and possibly larger head scales. These are mentioned
by the two authors of this new form as differences between their
new race and the typical form.
Anolis limifrons rodriguezii Bocourt
Anolis rodriguezii Bocourt, 1873, Miss. Sci. Mex., Eept., p. 62, pi. 13, fig.
1 : Panzos, Alta Vera Paz, Guatemala.
3 (M.C.Z. 53862-4) Laguna Oeotal, July-Aug.
I am doubtful about the validity of microlepis Alvarez del
Toro and Smith (1956, Herpetologiea, 12, p. 4) as a race.
Anolis capito Peters
Anolis (Draconura) capito Peters, 1863, Monatsb. Akad. "Wiss. Berlin, p.
142 : Costa Kica.
2 (M.C.Z. 53865-6) Laguna Oeotal, July-Aug.
1 (M.C.Z. 53893) El Censo to Monte Libano, Aug. 21.
This is the first record of capito from the state of Chiapas.
Basiliscus vittatus Wiegmann
Basilicus vittatus Wiegmann, 1828, Isis von Oken, 21. p. 373: Mexico.
1 (M.C.Z. 53850) Monte Libano, July 16-18.
7 (M.C.Z. 53867-71) Laguna Oeotal, July-Aug.
1 (M.C.Z. 53898) El Censo to Monte Libano, Aug. 21.
Corythophanes hernandezii (Wiegmann)
Cltamaeleopsis Hernandesii (sic) Wiegmann, 1831, Isis von Oken, 3. p. 298:
Mexico.
5 (M.C.Z. 53872-6) Laguna Oeotal, July-Aug.
244 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Lygosoma assatum cherriei (Cope)
Mocoa cherriei Cope, 1893, Proc. Amer. Philos. Soc, 31. p. 340: Palmar,
Costa Rica.
3 (M.C.Z. 53877-8) Laguna Ocotal, July- Aug.
1 (M.C.Z. 53892) Laguna Ocotal to El Censo, Aug. 20.
Recently Mittleman (1950, Herpetologica, 6, p. 19) proposed
Scincella for all New World members of Lygosoma. However,
Schmidt (1953, Check List N. Amer. Amph. Rept., p. 147) treats
Scincella as a subgenus of Lygosoma, which seems a better course.
Rhadinaea decorata decorata (Gunther)
Coronella decorata Gunther, 1858, Cat. Snakes Brit. Mus., p. 35: Mexico.
$ (M.C.Z. 53899) El Censo to Monte Libano, Aug. 21.
Midbody scale rows 17; ventrals 118; subeaudals 73+.
Lampropeltis doliata polyzona Cope
Lampropeltis polyzona Cope, 1860, Proc. Acad. Nat. Sci. Philadelphia, 12.
p. 258 : Cuatupe, near Jalapa, Veracruz, Mexico.
9 (M.C.Z. 53849) Monte Libano, July 16-18.
9 (M.C.Z. 53879) Laguna Ocotal, July-Aug.
Midbody scale rows 21-23 ; ventrals 233-238 ; subeaudals 54+
-56. This form was previously unrecorded from Chiapas.
Sibon dimidiatus (Gunther)
Leptognathus dimidiata Gunther, 1872, Ann. Mag. Nat. Hist. (4) 9. p.
31 : Mexico.
9 (M.C.Z. 53882) Laguna Ocotal, July-Aug.
Midbody scale rows 15; ventrals 182; subeaudals, with some
doubt, 109. James A. Peters, who is revising the Dipsas group,
tells me this species belongs in the genus Sibon, and that if a
subspecies is recognizable this Ocotal snake belongs to the typi-
cal form.
This specimen provides the first definite locality record for
Mexico, although dimidiatus is known from Piedras Negras,
Peten, just over the Guatemalan border.
Imantodes cenchoa leucomelas Cope
Himantodes leuoomelas Cope, 1861, Proc. Acad. Xat. Sci. Philadelphia, 13.
p. 296 : Mirador, Veracruz, Mexico.
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 245
$ (M.C.Z. 53881) Laguna Ocotal, July- Aug.
Midbody scale rows 17; verftrals 249; subcaudals 161.
Coniophanes fissidens fissidens (Giinther)
Coronella fissidens Giinther, 1858, Cat. Snakes Brit. Mus., p. 36: Mexico.
$ (M.C.Z. 53880) Laguna Ocotal, July-Aug.
Midbody scale rows 21. Neither ventral nor subcaudal counts
can be supplied as the specimen is decayed anteriorly and much
of the tail is missing. The snake was found dead.
Though this is the first Chiapas record of typical fissidens, as
defined by Smith and Taylor (1945, Bull. U. S. Nat. Mus., 187,
p. 39), it may not have much significance as the races of this
species still appear in need of revision.
Micrurus afpinis apiatus (Jan)
Flaps apiatus Jan, 1858, Eev. Mag. Zool., p. 522, col. pi. 1 ; Vera Paz,
Guatemala.
$ , 9 , imm. (M.C.Z. 53883-5) Laguna Ocotal, July- Aug.
Midbody scale rows 15 ; ventrals 205 ( $ ) , 217 ( 9 ) , 226 (imm.) ;
subcaudals 50 ( S ), 41 ( 9 ), 37 (imm.). These specimens show
evidence of intergradation with alienus, but are nearer apiatus.
In one snake the white blotch on the end of the snout is reduced ;
in another it is absent. In one the number of black body annuli
is reduced to 29, which is one in excess of the maximum given
for alienus. and one higher than the minimum for apiatus.
Micrurus elegaxs elegaxs sg veraepacis
Elaps elegans Jan, 1858, Rev. Mag. Zool. p. 524, col. pi. 2: Mexico.
Micrurus elegans veraepacis Schmidt, 1933, Zool. Ser. Field Mus. Nat. Hist.,
20, p. 32; Campur, Alta Vera Paz, Guatemala.
9 (M.C.Z. 53900) El Censo to Monte Libano, Aug. 21.
Midbody scale rows 15 ; ventrals 217 ; subcaudals 32. The
ventral and caudal counts are intermediate between those given
by Schmidt (loc. cit.) for the two races mentioned above.
Bothrops atrox (Linnaeus)
Coluber atrox Linnaeus, 1758, Syst. Nat. ed. 10, 1. p. 222: "Asia" (in
error; restricted to Surinam by Schmidt and Walker, 1943).
246 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
6 (M.C.Z. 53886) Laguna Ocotal, July- Aug.
Midbody scale rows 25 ; ventrals 212 ; subcaudals 70. Smith
and Taylor (1945, Bull. U. S. Nat. Mus., 187. p. 180) regard
asper (sic) as a race of air ox, but it seems better to use the
binomial pending a revision of the species. As Bothrops is of
feminine gender, aspera is the proper rendering for this adjecti-
val name when used with that genus.
Bufo marinus (Linnaeus)
Rana marina Linnaeus, 1758, Syst. Nat. ed. 10, 1. p. 211: America.
1 (M.C.Z. 28212) Monte Libano, July 16-18.
On comparing this specimen with toads from Veracruz, Nuevo
Leon, and Guerrero, I failed to find the differences mentioned
by Taylor and Smith (1945, Proc. U. S. Nat. Mus., 95, p. 551) as
distinguishing their Chiapas material from the rest of their
Mexican toads, although they referred all to Bufo horribilis
Wiegmann. These authors fail to state how horribilis differs from
the several other forms that undoubtedly have been included in
what is conventionally regarded as marinus, whose type locality
was probably Surinam. Head and body length 150 mm.
Bufo valliceps Wiegmann
Bufo valliceps Wiegmann, 1833, Isis von Oken, 26, p. 657: Mexico.
4 (M.C.Z. 28213-6) Monte Libano, July 16-18.
20 tadpoles (M.C.Z. 28239) Laguna Ocotal, July-Aug.
13 (M.C.Z. 28240-4) Laguna Ocotal to El Censo, Aug. 20.
4 (M.C.Z. 28251-4) El Censo, Aug. 20.
6 (M.C.Z. 28259-63) El Censo to Monte Libano, Aug. 21.
The tadpoles are only tentatively referred to this species.
Leptodactylus mystaceus labialis (Cope)
Cystignathus labialis Cope, 1877, Proc. Amer. Philos. Soc, 17. p. 90: Prob-
ably Mexico.
1 (M.C.Z. 28255) El Censo, Aug. 20.
This form seems to differ from typical mystaceus only in size,
while no differences at all could be detected between mystaceus
and the West Indian albilabris. Possibly some color differences
might be found with living material.
Eleutherodactylus alfredi conspicuus Taylor
Eleutherodactylus conspicuus Taylor, 1945, Proc. U. S. Nat. Mus., 95. p. 567:
Piedras Negras, Peten, Guatemala, near Mexico-Guatemalan border.
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 247
1 (M.C.Z. 28224) Laguna Oeotal, July-Aug.
Trinomials are used as conspicuus is probably just the southern
representative of alf recti. The differences between the two forms
appear to be very slight. Although the type locality of con-
spicuus is in nearby Peten, this is the first time that this sub-
species has been recorded from Mexico.
Eleutherodactylus laticeps (Dumeril)
Hylodes laticeps Dumeril, 1853, Ann. Sei. Nat. Paris (3), zool., 19. p. 178:
Yucatan, Mexico.
1 (M.C.Z. 28220) Monte Libano, July 16-18.
7 (M.C.Z. 28225-9) Laguna Oeotal, July-Aug.
2 (M.C.Z. 28245-6) Laguna Oeotal to El Censo, Aug. 20.
These specimens agree closely with Kellogg 's description of
laticeps (1932, Bull. U. S. Nat. Mus., 160, pp. 93, 106), which
was taken from the unique type. Kellogg does not mention the
length of the tarsal fold, which in our material extends from
about one-half to two-thirds the length of the tarsus. The entire
underside of the lower jaw and throat, not just the sides of the
throat as in the type, is stippled Avith brown. In one specimen
(M.C.Z. 28246) the usual black side of the upper jaw is reduced
to a spot under the eye.
The largest example (M.C.Z. 28227) has a head and body
length of 78 mm. The much larger size of laticeps and its some-
what different coloring seem to be the chief characters separating
it from the closely related Central American E. gollmeri
(Peters). But in color gollmeri sometimes shows the same varia-
tion as is described above for M.C.Z. 28246.
Eleutherodactylus rugulosus (Cope)
Liyla rugulosa Cope, 1869, Proc. Amer. Philos. Soc, 11. p. 160: Pacific region
of the Isthmus of Tehuantepec, Mexico.
1 (M.C.Z. 28221) Monte Libano, July 16-18.
2 (M.C.Z. 28222-3) Laguna Oeotal, July-Aug.
4 (M.C.Z. 28247-50) Laguna Oeotal to El Censo, Aug. 20.
1 (M.C.Z. 28258) El Censo to Monte Libano, Aug. 21.
All are subadult so that some are referred to rugulosus with
considerable doubt.
248 BULLETIN : MUSET7M OF COMPARATIVE ZOOLOGY
Hyla loquax Gaige and Stuart
Hyla loquax Gaige and Stuart, 1934, Occ. Pap. Mus. Zool. Univ. Michigan,
no. 281, p. 1: Ixpuc Aguada, north of La Libertad, Peten, Guatemala.
$ (M.C.Z. 28238) Laguna Ocotal, July-Aug.
The color pattern differs somewhat from that of our two
paratypes and the original description. A blackish suffusion on
the dorsum extends forwards about as far as the insertion of
the forelimbs, the anterior border being nearly straight; head
and body length 41 mm. Although known from Piedras Negras,
Peten, just across the Guatemalan frontier, this is the first record
from Chiapas.
Hyla phaeota cyanosticta Smith
Hyla phaeota cyanosticta Smith, 1953, Ilerpetologica, 8, p. 150: Piedras
Negras, Peten, Guatemala.
3 (M.C.Z. 28217-9) Monte Libano, July 16-18.
The largest of these unquestionably belongs to this race. The
two smaller examples (M.C.Z. 28218-9) are less certain. They do
not show the reticulation or spotting on the limbs and sides dis-
played by the large specimen.
This constitutes the first Mexican record for both the species
and the race, as Smith's material came from the Guatemalan
side of the Chiapas-Guatemala line (see type locality above).
PtANA PALMIPES Spix
Bana palmipes Spix, 1824, Nov. Spec. Test. Pan., p. 29, pi. 5, fig. 1 :
Amazon River, Brazil.
28 (M.C.Z. 28234-7) Laguna Ocotal, July-Aug.
2 (M.C.Z. 28270) Laguna Ocotal to El Censo, Aug. 20.
1 (M.C.Z. 28256) El Censo, Aug. 20.
1 (M.C.Z. 28257) El Ceuso to Monte Libano, Aug. 21.
It is likely that these Mexican frogs are subspecifically distinct
from topotypic Brazilian material. A revision is needed.
Raxa pipiens Schreber
liana pipiens Schreber, 1782, Der Naturforscher, Halle, 18, p. 185, pi. 4:
Raccoon, Gloucester County, New Jersey (restricted to White Plains,
New York, by Schmidt, 1953).
4 (M.C.Z. 28230-3) Laguna Ocotal, July-Aug.
Without a revision of the species, it is not known to what
race the above material should be referred.
BIOLOGICAL INVESTIGATIONS IN CHIAPAS. MEXICO 249
VII
BIRDS OF LAGUNA OCOTAL
By
Raymond A. Paynter, Jr.
INTRODUCTION
An investigation of the avifauna was one of the primary ob-
jectives of the Museum of Comparative Zoology expedition to the
Selva Lacandona, Chiapas. Between July 21 and August 19,
1954, while at Laguna Ocotal (alt. 950 m.), 490 birds were pre-
pared as skins. While these specimens are a good sample of
the resident avifauna, there can be little doubt that additional
species occur in the region but were not observed. In a heavily
forested region collecting is difficult even under the most favor-
able conditions. We were at work at the end of the breeding
season when most birds are silent and secretive - obviously the
most difficult collecting period.
The following list is based on the specimens collected, as
well as on unequivocal field observations. The specimens were
weighed on a double-pan balance. The means of the measure-
ments are accompanied by their standard errors (am)- The
Hippoboscidae (bird-flies) were identified by Joseph C. Bequaert.
ANNOTATED LIST
Tinamus major robustus Sclater and Salvin
1 $ , Aug. 7.
The species was heard on a few occasions and seen twice. The
bird weighed 1090.5 grams.
Crypturellus boucardi boucardi (Sclater)
1 9 , July 22 ; 1$, Aug. 10 ; 1$, Aug. 11.
Although by no means common, this is the more abundant
tinamou. Occasionally it was found in the tropical evergreen
forest but it occurred principally in the monte. The males
weighed 291.4 and 403.3 grams; the female 351.0 grams.
250 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Phalacrocorax brasilianus mexicanus (Brandt)
1 S , July 30.
On an undisturbed lake one would expect to find water birds
in abundance, but the cormorant was the only exclusively
aquatic form and even they were in small flocks. The weight of
the bird was 1165.0 grams.
Butorides viRESCENS virescens (Linnaeus)
1$, Aug. 11.
The specimen is immature and weighed 93.1 grams. Green
Herons were uncommon.
Sarcoramphus papa (Linnaeus)
One was observed, within a flock of Turkey Vultures, on
August 3.
Cathartes aura subsp.
A few vultures came to the camp at irregular intervals.
Elano'ides forficatus subsp.
Two kites were noted on Julv 26 and three on August 1.
i& ■
Accipiter striatus subsp.
A single individual flew back and forth over the lake on
August 16. It would seem an early date for a migrant. Prob-
ably the bird was a resident, possibly A. s. chionogaster.
Buteogallus urubitinga ridgwayi (Gurney)
1 £ , Aug. 15.
This is the only example of the species which was seen. It
harbored three species of Hippoboscidae : Omithociona erythro-
cephala, Lynchia angustifrons, and L. wolcotti.
I concur with Amadon (1949), and Amadon and Eckelberry
(1955), that the genera Hypomnrphnus and Buteogallus should
lie united.
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 251
Spizaetus tyrannus subsp.
Paine, on August 17, saw a single Black Eagle-Hawk in the
selva.
IIerpetotiieres cachinnans subsp.
Laughing Falcons were heard several times.
MlCRASTTJR SEMITORQUATUS NASO (LeSSOn)
1$, Aug. 17.
Although the dense, undisturbed, forests surrounding the
lake appeared ideally suited for certain of the birds of prey,
such as those of the genera Micrastur, Spizastur, and Spizaetus,
hawks were excessively rare. Only one example of M. semitor-
quatus was seen. It was host to the bird-flies Omithoctona
erythrocephala and Lynchia wolcotti, and weighed 749.8 grams.
Micrastur ruficollis guerilla Cassin
1 (J , Aug. 14.
The specimen was taken in the low forest near the edge of
the lake. On several occasions others were found in the thickest
parts of the evergreen forest.
Falco rufigularis subsp.
On August 10 a pair of Bat Falcons flew near the (.'amp.
where they could be seen distinctly.
Crax rubra rubra Linnaeus
U, 19, Aug. 8; 1$, Aug. 15.
Curassows were fairly abundant but not so numerous as
guans. The specimen collected on August 15 is about one-third
grown.
Penelope purpurascens purpurascens Wagler
1 $ I, July 24.
Guans were common, ranging through all types of forest, but
were most abundant where the pine and broadleaf forests met. In
252 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
the early morning they were often perched in the pines where
apparently they had spent the night. Ornithoctona erythro-
cephala was taken from the specimen.
Ortalis vetula vetula (Wagler)
1 $ , July 26 ; 19, Aug. 1 ; 1 o , Aug. 6; 19, Aug. 7 ; 1 9 , Aug. 8 ;
2$, 1$, Aug. 10; 1$, Aug. 14; 2 9, Aug. 15; 1 <$ , 3 9 , Aug. 16.
Chachalacas were very abundant in the transition zone between
the broadleaf forest and the pines, and in the monte at the end
of the lake. Specimens of Ornithoctona erythrocephala were
obtained from four birds and Lynchia plaumanni from one. Six
males ranged in weight from 319.1 to 448.5 grams, with a mean
of 402.30±16.82; nine females from 309.6 to 394.8, with a mean
of 356.21±10.31 grams.
ODONTOrHORUS GUTTATUS (Gould)
2 $ , July 25 ; 19, July 30 ; 19, Aug. 12.
Wherever the floor of the forest is dark these birds were seen
with fair regularity. The two males weighed 284.0 and 286.9
grams; the two females 314.1 and 316.3 grams.
Aram us guarauxa dolosus Peters
19, Aug. 1; 19, Aug. 11.
There seems to be no specific record of the species in Chiapas,
although Alvarez del Toro (1952) mentions that it is abundant
in the state. The shores of the lake are strewn with the empty
shells of the snail Pomacea flagellata, the preferred food of
limpkins, suggesting the presence of many of these birds. How-
ever, none was heard and just a few lone individuals were seen.
Aramides cajanea subsp.
A wood rail called in the evening of July 30. This is another
species which had been expected to occur in large numbers but
which was inexplicably rare.
Laterallus ruber (Sclater and Salvin)
16,19, July 21.
Iii the marshes near the camp, Ruddy Rails abounded, although
they were seldom seen. Their call is a rattle, similar to that of a
BIOLOGICAL INVESTIGATIONS TN CHIAPAS, MEXICO 253
small kingfisher, but somewhat slower. The male is an adult and
weighed 49.0 grams. The female retains a portion of the im-
mature plumage and weighed 40.5 grams. Elsewhere (Paynter,
1955) I have given my reasons for recognizing no subspecies of
L. ruber-.
Columba nicrirostris Sclater
1 5 , Aug. 4 ; 1 6 . J $ , Aug. 7 ; 1 $ , Aug. 11 ; 1 $ , Aug. 12 ; 19, Aug. 15.
Short-billed Pigeons were heard frequently, and sometimes
could be seen in the tallest, trees of the broadleaf forest. The
specimens, however, were taken principally in the low forest and
edges where they came to feed in fruiting trees.
The males weighed 154.3, 154.4. 159.0, and 166.1 grams; the
females 132.5 and 148.2 grams.
Leptotila cassinii ceryiniventris Sclater and Salvin
1 <5 , July 22 ; 15. July 2.5 ; 1 $ , July 30 ; 16, July 31.
Although the species has been recorded in Mexico only from
Santa Kosa, Comitan, Chiapas (Berlioz, 1939) and from two
localities on the Rio Usumacinta in Tabasco (Brodkorb, 1943),
it was reasonably abundant in the deep forest at Laguna Ocotal.
The apparent absence of Leptotila verreauxi and L. plumbeiceps,
species which are widely distributed in southern Mexico, was
surprising. However, most forms of Leptotila are secretive and
difficult to collect and I would feel certain of the absence of
these species only if more time had been spent in the field.
Two adult males and a female weighed 167.2, 176.5, and 152.1
gram, respectively; a female in juvenal plumage 138.0 grams.
Ara macao (Linnaeus)
Small flocks of Scarlet Macaws flew over quite regularly in
the morning and evening. They did not seem to feed in the
vicinity of the lake and, as a consequence, alighted rarely, and
then only in the tops of tall pines or on conspicuous dead trees
in the selva.
254 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
PlONUS SENILIS SENILIS (Spix)
1$, Aug. 14; 1$, Aug. 16.
Although parrots passed over each day at dawn and dusk,
they were nearly absent from the forest surrounding Laguna
Ocotal. Late in our stay a small flock of Pionus senilis began
to roost for the night in the pines at the edge of the broad-
leaf forest on the eastern side of the lake. It is from this
group that the specimens were secured. The birds weighed 220.4
and 221.5 grams.
The genus Amazona was represented in the flocks of parrots
seen in the air. A. ochrocephala was tentatively identified, but
another species seemed to be present also ; there may have been
still others.
PlAYA CAYANA subsp.
On August 12 there was a Squirrel Cuckoo in the top of a
pine standing in small clearing. No other was observed.
Nyctidromus albicollis yucatanensis Nelson
1 $ , July 29 ; IS, Aug. 10 ; 1 $ , Aug. 17.
Common within the burned area in the pines. The males
weighed 65.5 and 68.5 grams; the female 74.5 grams.
Phaethornis superciliosus longirostris (DeLattre)
1 9 , July 31; 1 8 , Aug. 9.
No approach to P. s. veraecrucis is exhibited by the specimens,
although birds from Palenque, which is about 100 kilometers to
the north, either were referred to that race (Brodkorb, 1943,
Tashian, 1952) or said to be intermediate (Friedmann, Griscom,
and Moore, 1950). The weight of the male was 6.4 grams; that
of the female 5.9 grams. Dressier reported seeing one of these
hummingbirds feeding at a dwarf Heliconia {H. tortuosa
Griggs), a plant with red bracts and long, curved, yellow
flowers, for which the bird's bill is well suited.
Abeillia abeillei abeillei (Lesson and DeLattre)
1$, July 27; 15,1 ?, Aug. 8.
The female weighed 3.5 grams ; the male, which is immature,
3.0 grams. No adult males were seen.
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 255
Amazilia Candida Candida (Bourcier and Mulsant)
1$, Juy 24; 1 $ , Aug. 11.
The male and female weighed 3.8 and 3.6 grams, respectively.
This species and the females and immature males of Abeillia
abeillei were impossible to differentiate in the field ; their relative
abundance is not known. Among the least common humming-
birds, they occurred in the high broadleaf forest in localities
where the sun reaches the ground and encourages the growth
of flowering plants and shrubs.
Amazilia beryllina devillei (Bourcier and Mulsant)
IS, Aug. 11.
The specimen displays none of the characters ascribed to
A. b. lichtensteini or to the nominate race and is, therefore,
the first record of A. b. devillei from the Atlantic slope of
Chiapas. It was taken in the tropical evergreen forest and was
the only example seen. It weighed 4.6 grams.
Eupherusa eximia eximia (DeLattre)
14, July 24; IS, 19, July 25; 1$, July 26; ] 9, July 27; 1$,
July 29; 2$, July 30; 13, 19, Aug. 1; 2$, Aug. 2; 29, Aug. 3:
1 S , Aug. 4, 1 $ , Aug. 6 ; 1 $ , Aug. 7 ; 2 S , Aug. 8 ; 1 9 , Aug. 9 :
2$, Aug. 10; IS, Aug. 13; 19, Aug. 14; 1$, Aug. 16;
19, Aug. 17; IS, Aug. 19.
Where there were plants flowering in the tall broadleaf forest
this hummingbird was almost always present. It was by far the
most common Trochilid. Twenty males ranged from 3.5 to 4.7
grams, with a mean of 4.18±.06; seven females from 3.0 to
4.0 grams, with a mean of 3.65±.14.
Lampornis viridi-pallens viridi-pallens (Bourcier and
Mulsant)
1$, July 23; 2$, 19, July 27; 19, July 31; IS, Aug. 1; 19,
Aug. 5; 19, Aug. 6; 1 $ , Aug. 9; 2$, Aug. 11.
These hummingbirds occurred in the same biotope as Eupher-
usa eximia and were almost as numerous.
I have examined 15 specimens of the species from and near
256 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
Mt. Ovando, Chiapas, the type locality of L. v. ovandensis, and
conclude that, although recognizahle, it is an extremely weak
race. The coloration of the dorsal surface is variable and no
consistent difference between the two races is distinguishable.
The bill length is also inconstant; there appears to be a com-
plete overlap between the forms in this character. The only dif-
ferentiating features I can appreciate are the faintly lighter
abdomen and slightly greater area of white on the lower throat
of L. v. ovandensis.
Seven males ranged in weight from 5.3 to 6.5 grams, with
a mean of 5.80±0.14 grams. Four females weighed 4.7, 4.7, 4.8,
and 5.2 grams.
Trogon massena subsp.
A single bird was seen on August 15 in a fruiting tree at the
edge of the pines.
Trogon collaris puella Gould
1 6 , July 21 ; 1 9 , Aug. 5; 16, Aug. 12.
Within the heavy forest this species was noted with regularity,
but it was uncommon.
The males weighed 63.7 and 64.5 grams ; the females 63.4
grams. It is of interest to compare these weights with those
obtained on the Yucatan Peninsula (Paynter, 1955). There two
males weighed 47.6 and 53.5 grams, and two females 41.1 and
53.9 grams. This suggests that the Peninsular birds are smaller
in mass, although no differences in linear measurements are
evident. A larger series is needed to confirm the proposal.
Trogon violaceus braccatus (Cabanis and Heine)
1 9 , Aug. 6.
The specimen, the only one of the species seen, was collected
in the transition forest. It weighed 57.1 grams.
CHLOROCERYLE AMERICANA SEPTENTRIONALIS (Sharpe)
1 <$ , July 23 ; 19, Aug. 3 ; 1 9 , Aug. 3 ; 13, Aug. 9.
Two adult females and a male weighed 40.7, 43.1, and 39.7
grams, respectively; an immature male 37.3 grams.
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 257
Chloroceryle aenea stictoptera (Eidgway)
1 9 . Aug. 10; 1 $ , Aug. 11 ; 1 $ , Aug. 15.
Both species of kingfisher were common. The two males
weighed 15.5 and 16.8 grams, the female 20.8 grams.
Htlomanes momotula momotula Lichtenstein
1 <J , Aug. 2; 1 $ , 1 9 , Aug. 3 ; 1 9 f, Aug. 7.
These small motmots were seldom seen, but because of their
silent, sluggish behavior may have been more numerous than it
appeared.
The two males weighed 32.5 and 32.7 grams; the female 29.7
and the bird of doubtful sex 31.0 grams. None possessed more
than a trace of fat. Van Tyne (1935) gave the weight of five
males from Peten as ranging from 27.0 to 28.5 grams, and
noted that the heaviest bird was fatty ; two females weighed 26.5
and 27.5 grams. The apparent difference in weight between the
two populations should be investigated when more data are
available.
Momotus momota lessonii Lesson
1$, Aug. 6.
No more than ten of these birds were seen during the period
spent at the lake. The specimen weighed 131.6 grams, which is
considerably heavier than the previously reported maximum
of 119.5 grams (Van Tyne, 1935).
Tashian (1952) listed M. m. goldmani from Palenque, but
gave no reason for resurrecting the race which Van Tyne (1935),
Wetmore (1943), and Berlioz (1952), have shown to be synony-
mous with HI. m. lessonii. I, too, am unable to appreciate the sup-
posed characters of 31. m. goldmani.
Pteroglossus torquatus torquatus (Gmelin)
15, Aug. 3; 1$, Aug. 11.
The birds weighed 209.5 and 221.0 grams, further strengthen-
ing Van Tyne's (1935) assertion that the nominate race is
markedly heavier than P. t. erythrozonus. The maximum and
minimum weights recorded for P. t. erythrozonus are: males,
258 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
177.0 (Van Tyne, 1935) and 169.2 grams (Paynter, 1955) ;
females, 181.1 (Paynter, 1955) and 147.0 grams (Van Tyne,
1935). Those for the nominate form are: males 240.0 and 209.5
grams (Van Tyne, 1935) ; female (only record) 230.0 grams
(Van Tyne, 1935).
Aracaris were fairly common.
Ramphastos sulfuratus sulfuratus Lesson
1 S , July 30 ; 1$, Aug. 18.
This large toucan was more abundant than Pteroglossus tor-
quatus.
Van Tyne (1935) found, in Peten, the weights of males of the
race range from 362.0 to 449.5 grams. These specimens weighed
483.5 and 496.4 grams and are apparently the heaviest yet re-
corded.
PlCULUS RUBIGINOSUS YUCATANENSIS (Cabot)
1 6 , July 23 ; 1 $ , Aug. 15.
The specimens collected were the only ones seen. They ex-
hibited no approach toward P. r. maximus of the mountains of
.Chiapas and Guatemala. The male weighed 83.0 and the female
75.4 grams.
Centurus pucherani perileucus (Todd)
1 9 , July 29 ; 1 <$ , Aug. IS ; 19, Aug. 19.
Within the pines this species was moderately numerous, but
it usually ranged in the tops of the trees, out of gunshot. One
of the specimens was collected in the pines and the other two
in the zone of transition between the pine and broadleaf forests.
It was rare, however, in the latter locality. The weight of the
male was 53.0 grams; that of the females 48.9 and 49.4 grams.
Amadon and Eckelberry (1955), in commenting on the failure
of many authors to unite Centurus with Melanerpes, as was
done by Peters (1948), state that "... the barred (sic) imma-
ture plumage of the Red-headed Woodpecker (M. erythrocepha-
lus) and the color pattern of M. portoricensis, do tie the two
groups together. ' ' I fail to appreciate this. The plumage of the
immature M. erythrocephalus is streaked, not barred, and notably
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 259
similar to that of the adult Acorn Woodpecker {M. formici-
vorus). No species of Centurus approximates such a condition.
M. portoricensis, an insular species presumably at the end
of an evolutionary line, rather than a link, bears no resemblance
to the Centurus group, with the exception of its red abdomen and
brown sides. Dorsally it is similar to M. formicivorus with a
glossy black back, white rump, and white forehead ; it lacks
only the red head of that species (and of Centurus) . Its behavior
is said to be like that of the Acorn Woodpecker (Wetmore,
1927). The coloration of the throat and upper chest of M.
portoricensis is similar to that of M. erythroceplialus, even to
the remnant of a black pectoral band. The extension of red to
the abdomen, when the chest is red, is a small evolutionary step
and is certainly a logical explanation for the existence of this
character in M. portoricensis.
Thus, the only real similarity between this species and the
Centurus group is the brown coloration of the sides. It seems
most likely that this character was acquired independently by
an M. forinicivorus-liiie progenitor, which was isolated in the
Antilles. Alternative explanations are that M. portoricensis is
a primitive species exhibiting the first indications of the diver-
gence of Centurus from Melanerpes, or, that Centurus is the
older group and M. portoricensis, retaining only its brown
sides, betrays the origin of Melanerpes. These are obviously
spurious hypotheses.
It is unexpected that Amadou and Eckelberry (1955) should
accept the unification of Centurus and Melanerpes, but main-
tain Tripsurus, a genus also reduced to the synonymy of Mela-
nerpes by Peters (1918). It is true that the species usually
placed within Tripsurus are separated from Centurus (sensu
stricto) by gaps greater than those existing between most species
of Centurus, but I believe the group is not of generic rank and
is best considered a subgenus of Centuries.
'B'
Veniliornis fumigatus sanguinolentus (Sclater)
1 6 , Aug. 4 ; 1$, Aug. 6 ; 1 $ , Aug. 12 ; 1$ , Aug. 15 ; 19, Aug. 17.
Where trees have fallen in the broadleaf forest, and on the
edges of other clearings, there is usually a stand of shrubs and
young trees. It is in such a habitat that this woodpecker is found.
It is, therefore, localized and uncommon.
260 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
The female weighed 29.3 grams and the males 32.0, 34.1, 35.0,
and 39.3 grams.
Phloeoceastes guatemalensis guatemalensis (Hartlaub)
1 £ , 2 9 , July 31 ; 1 9 , Aug. 2; 1 $ , Aug. 8.
An abundant resident in the high forest. The males weighed
234.0 and 237.0 grams; the females 222.5, 223.0, and 226.4 grams.
Dendrocincla anabatina anabatina Sclater
1 $ , July 23 ; 1 <$ , Aug. 7; 1 $ , Aug. 11 ; 1 <$ ?, Aug. 17.
There appears to be no specific record of this species in the
state. Alvarez del Toro (1952), however, has included it in his
book. The weight of three males was 34.2, 37.3, and 38.3 grams.
Dendrocincla homociiroa iiomochroa (Sclater)
1 ?, July 23 ; 19, July 25 ; 1 $ , Aug. 9.
Two males weighed 33.8 and 35.5 grams. Both species of
Dendrocincla were present in moderate numbers and seemed
about as common as I have found them in the rain forest of
the Yucatan Peninsula.
Sittasomus griseicapillus sylvioides Lafresnaye
19, July 23; 1 ?, July 24; 1 <$ , 1$, July 2.5; 1 £ , Aug. 4; 1 f,
Aug. 5; 1$, Aug. 8; 1$, Aug. 12; 19, Aug. 13; 1 6 , 1 ?, Aug.
14; 19, Aug. 16; 29, Aug. 17; 1 ?, Aug. 18; 1$, Aug. 19.
The linear dimensions of 8. g. sylvioides and 8. g. gracileus
have been shown to be markedly different (Paynter, 1955). It
is not unexpected that a substantial difference in the average
weight of the races also can be demonstrated. The weight of
five males of 8. g. sylvioides from Laguna Ocotal ranged from
12.8 to 14.2 grams, with a mean of 13.52±0.21; that of six
females from 11.1 to 13.8 grams, with a mean of 12.53±0.12.
On the Yucatan Peninsula, where 8. g. gracileus is found, seven
males ranged from 9.0 to 12.1 grams, with a mean of 11.01±0.35;
four females from 8.6 to 10 grams, with a mean of 9.25±0.34
(Paynter, 1955).
Within the selva this was the most abundant of the Dendro-
colaptidae.
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 261
Dendrocolaptes certhia sancti-thomae (Lafresnaye)
1 9 , July 26 ; 1 9 , Aug. 9; 1 $ , Aug. 13.
A bird of the high evergreen forest, but at times seen on the
trunks of pines when they were adjacent to its preferred habitat.
No woodhewer was more shy.
When more birds have been weighed there may be evident a
difference in mass between this subspecies and D. c. legtersi
of the Yucatan Peninsula. The male and the two females
weighed, respectively, 67.3, 66.0, and 68.3 grams. Tashian
(1952) found a female to weigh 63.0 grams. Paynter (1955)
recorded two males of D. c. legtersi as weighing 52.7 and 60.8
grams, and three females of D. c. sancti-thomae, which exhibited
an approach toward the Yucatan Peninsula endemic, as 54.3,
55.9, and 61.3 grams.
XlPHORHYNCHUS ERYTHROPYGIUS PARVUS GrisCOm
19,1 ?, July 27; 29, July 31, 2 9,1 ?, Aug. 1; 19, Aug. 3;
IS, Aug. 4; 1$, Aug. 5; 19, Aug. 6; 19, Aug. 7; 1$, Aug.
16; 19, Aug. 17; 1 ?, Aug. 18.
Xine females ranged in weight from 39.7 to 43.7 grams, and
had a mean of 41. 78 ±0.43 grams. Three males weighed 43.0,
44.0, 44.9 grams. These were abundant birds in the broadleaf
forest and occasionally ranged to the edge of the pines.
Lepidocolaptes souleyetii insignis (Nelson)
2 9 , July 26 ; 19, Aug. 16 ; 1$, Aug. 18.
L. of/in is and the present species occurred sympatrically in
the pines. They are morphologically so similar it was not possible
to distinguish them in the field ; their relative abundance is un-
known. As an aggregate, however, they were not common.
An adult male and two females weighed 30.8. 29.8, 30.9 grams,
respectively ; a juvenal female 30.7 grams.
Lepidocolaptes affinis affinis (Lafresnaye)
1 9 , Aug. 5.
It is regrettable that it was not possible to study the species
carefully while in the field. L. souleyetii is a lowland form which
usually occurs in rain forest, while L. affinis is found in pines or
262 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
other types of forest characteristic of the highlands. The fact
that at Laguna Ocotal L. souleyetii abandoned the broadleaf
forest for the pines suggests that it may have found a more
easily exploitable niche within the conifers. Neither species was
common and probably they were able to exist sympatrically
without competing.
The specimen weighed 27.6 grams.
Anabacerthia striaticollis variegaticeps (Sclater)
1 $ , Aug. 2.
The specimen, which was the only one seen, weighed 23.6
grams.
Automolus ochrolaemus cervinigularis (Sclater)
1 $ , July 31 ; 1$, Aug. 2; 1 3 , Aug. 3 ; 1 $ , Aug. 5 ; 1 $ , Aug. 6 ;
2 9 , Aug. 7 ; 1 9 , Aug. 10; 2 5,2$, Aug. 17.
None of the Furnariidae was more abundant than this species,
which was a conspicuous element of the heaviest selva.
Seven males ranged in weight from 44.7 to 50.4 grams, with a
mean of 47.34±0.87; five females from 37.5 to 44.0 grams, with
a mean of 40.48±1.12.
Xenops minutus mexicanus Sclater
1 9 , Aug. 5 j 19, Aug. 16.
Very few Plain Xenops were present. The birds weighed 10.5
and 11.8 grams.
Sclerurus mexicanus mexicanus Sclater
1 6 , July 23 ; 1 ?, 1 9 , July 24 ; 1 9 , July 27 ; 19, Aug. 17.
These birds were seen on the average of about once a day in
the most dense forest. The weight of the male and three females
was 28.0, 25.0, 26.6, and 30.0 grams, respectively.
Thamnophilus doliatus intermedius Ridgway
1 9 , July 29; 1 $ , Aug. 4; 1 $ , Aug. 18.
Antshrikes were restricted to the monte and thickets at the
end of the lake. The male taken on August 4 had enlarged
testes and was one of the few birds collected which showed indica-
tions of reproductive activity. The males weighed 28.4 and 28.8
grams; the female 30.0 grams.
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 268
Dysithamnus mentalis septentrionalis Ridgway
2 (} , 1 9 , July 24 ■ 16, July 31 ; 1 <$ , Aug. 3 ; 19, Aug. 6.
Tray lor (1941) collected two examples of this species in south-
ern Campeche, adding- the species to the known Mexican avi-
fauna. At Laguna Ocotal, it was common, and often associated
with Myrmotherula schisticalor, in the shrubs bordering trails
through the thickest parts of the deciduous forest.
Three of the males are immature and weighed 11.6, 12.7, and
13.5 grams. An adult male and two females weighed 13.6, 12.9,
and 13.8 grams, respectively.
Myrmotherula schisticolor schisticolor (Lawrence)
1(J, July 24; 1$, Aug. 6; 1 S , Aug. 14; 19, Aug. 16; 1&, 19.
Aug. 17.
Berlioz (1939) recorded seven specimens from Santa Rosa
in the district of Comitan ; no other record from Mexico is known.
It seemed to be localized in the forest, but often occurred in loose
flocks. The respective weights of three males and three females
were 7.3, 9.2, 9.2, 8.5, 8.8, and 9.6 grams.
PORMICARIUS ANALIS MONILIGER Sclater
IS, Aug. 18.
Antthrushes were noted only four times but undoubtedly
were more abundant than it seemed. On the Yucatan Peninsula
I found them exceedingly difficult to collect during the wet sea-
son, but when the forest was dry they were heard moving about
in the leaves, much like tinamous, and could be taken with little
difficulty. The specimen from Laguna Ocotal was snared in a
trammel net in the monte, although previously the species had
been seen only in the high forest. It weighed 51.3 grams.
Grallaria guatimalensis guatimalensis Prevost and Des Murs
1 $ Aug. 1.
Only this bird was seen. Its weight was 99.0 grams.
Attila spadiceus flammulatus Lafresnaye
1 9 , July 24.
This specimen came to a fruiting tree (Clusia sp.) at the
camp. It was the only one noted during the month at the lake.
It weighed 44.2 grams.
264 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Platypsaris aglaiae sumichrasti Nelson
1 $ , July 27.
The bird is almost in full adult plumage and weighed 31.5
grams. It was taken in the monte. No others were seen.
Tityra semifasciata personata Jardine and Selby
2 $ , Aug. 9.
Masked Tityras were observed on a few occasions in the tops
of pines or broadleaf trees. The birds weighed 77.9 and 86.3
grams.
Pipra mentalis mentalis Sclater
16, July 26; 1$, Aug. 2; 12, Aug. 3; 1$, Aug. 5; 29, Aug.
6; 16, Aug. 15; 19, Aug. 16; 1$ , Aug. 18.
One of the females collected on August 16 is a fledgling barely
able to fly. This species is among the latest breeders found in the
region and also was one of the most common birds wherever there
were bushes and small trees in the broadleaf forest. Adult males
made up about one quarter of the birds seen.
Adult males weighed 17.8 17.9, and 19.1 grams; an immature
male 15.8 grams; adult females 1 6.6, 17.1, 17.2, and 17.4 grams; a
juvenal female 17.4 grams.
Schifforxis turdinus verae-pacis (Sclater and Salvin)
1 9 , Aug. 2 ; 19, Aug. 8; 1 $ , Aug. 17.
Few of these manakins were in the area. The male and the
two females weighed 32.1, 32.3, and 32.7 grams, respectively.
Tyraxxus melaxciiolicus chloroxotus Berlepsch
1 6 , 1 f, Aug. 4.
The paucity of clearings made this a very uncommon species.
The few birds seen were in the pine burn or at the edge of the
lake. The male weighed 41.4 grams; the bird whose sex could
not be determined 36.1 grams.
Megarhynchus pitangua mexicaxus (Lafresnaye)
1 9, Aug. 3; 1 6, Aug. 13.
As uncommon a bird as Tyrannus melancholicus and appar-
ently for the same reasons. The weights of the male and female
wore (io. 4 and 66.3 grams, respectively.
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 265
Myiozetetes similis texensis (Giraud)
19, Aug. 6; 19, Aug. 13.
Slightly more abundant than the preceding two species. The
specimens weighed 28.0 and 33.9 grams.
Myiarchus tuberculifeb connectens Miller and Griscom
U, July 20; 1 9 , Aug. 5 ; 1 $ , Aug. 10; 2 ?, Aug. 12; 1 $ , Aug. 18.
The specimens from Laguna Ocotal are referable to M. t.
connectens, although they are rather small and fall within the
upper size range of a series of 24 specimens of M. t. platyrhyn-
chus, the smallest of the races, from the Yucatan Peninsula.
They are, however, dark dorsally and have varying amounts of
rufous on the underside of the rectrices, two characters dis-
tinguishing M. t. connectens from M. t. platyrhynchus. The race
seems not to have been reported from Chiapas, although it is to
be expected along the northern and northeastern Guatemalan
border.
Recently (1955) I noted that one character distinguishing M.
tuberculifer from M. yucatanensis is its horn-colored, rather than
black, bill. It was not realized at the time that the lighter colored
bill is not always found in M. tuberculifer. M. t. connectens and
M. t. lawrenceii occasionally have horned-colored bills, but in the
vast majority it is black. On the other hand, within the remain-
ing races horn-colored bills are frequent and black bills are some-
what of an exception; only M. t. platyrhynchus, and apparently
M. t. nigricapillus, seem to have consistently light bills.
Enough data have accumulated so that a slight difference in
weight between several races seems to be evident. Six males
of M. t. platyrhynchus ranged from 14.7 to 19.7, with a mean
of 17.15±0.74 grams; females of the race weighed 15.0, 16.0,
17.3, and 17.9 grams (Paynter, 1955). 31. t. connectens has larger
linear dimensions and the specimens from Laguna Ocotal sug-
gest that heavier weight may be an added character. Three males
weighed 19.0, 19.9, and 19.2 grams, a female 19.2 grams, and
two birds of undertermined sex 18.0 and 19.1 grams. M. t.
lawrenceii, the race with the greatest linear dimensions of the
three, may also weigh the most. Paynter (1955) recorded males
from Ocozocoautla, Chiapas as weighing 21.8 and 22.5 grams;
Martin, Robins, and Heed (1954) found males in Tamaulipas
266 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
to weigh 21 and 22 grams. Females from Chiapas (Paynter,
1955) weighed 20.0 and 22.0 grams.
Contopus pertinax pertinax Cabanis and Heine
19, Aug. 5.
Greater Pewees were restricted to the ocotal where they were
uncommon and much more shy than I have found them else-
where. The bird was in heavy molt and weighed 22.0 grams.
Empidonax minimus (Baird and Baird)
1 6 , Aug. 10.
Although this is an early date for the species, Tashian (1952)
has recorded it at Palenque, Chiapas on August 6. The specimen
weighed 10.6 grams.
Empidonax flavescens dwighti van Rossem
1 $, July 31; 1$, Aug. 13.
One bird was taken in the pines, where it was to be expected,
but the other was in the dense broadleaf forest. No more were
seen. The weight of the male was 12.4 and that of the female
12.2 grams.
Myiobius barbatus sulphureipygius (Sclater)
] $ , July 29; 1$, 1 ?, July 31 ; 1$, Aug. -t; 29, Aug. S; 16, 1
.', Au-. K); IS, Aug. 13; 1$, Aug. 14; 1 ?, Aug. 1(5.
The species was a conspicuous constituent of the selva. Five
males ranged in weight from 12.5 to 13.3 grams, with a mean
of 12.92±0.15 grams. Three females weighed 9.1, 9.9, and 10.4
grams.
Onychoriiynchus coronatus mexicanus (Sclater)
1 6 , Aug. 17.
The specimen was the only individual observed. It weighed
21.4 grams.
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 267
Platyrinchus mystaceus cancrominus Sclater and Salvin
IS, July 21; 1$, July 24; 2$, 19, 1 f, July 27; 1$, 19. 1
.'. .July 31; 1$, Aug. 9; 19, Aug. 13.
Some of the birds show a reduction in the streaking on the
abdomen, indicating an approach toward P. m. timothei, but
they are very richly colored, have breast bands, etc. — characters
of P. m. cancrominus. They frequented the underbrush of the
broadleaf forest in considerable numbers.
The weights of the males ranged from 10.0 to 12.0 grams,
with a mean of 11.16 + 0. 30. Females weighed 8.7, 9.7, 9.9 and
10.2 grams; two of indeterminate sex 9.3 and 13.0 grams.
Rhynchocyclus brevirostris brevirostris (Cabanis)
1 6 , .July 26 ; 2 6 . July 31; 1 $ Aug. 5 ; 19, Aug. 8.
Rather an uncommon resident of the broadleaf forest. The
males weighed 22.3. 23.1. 23. S, and 24.1 grams; a female 23.8
grams.
Oncostoma cinereigulare cinereigulare (Sclater)
1 6 . J uly 25 ; 1 9 , Aug. 8 ; 1$, Aug. 12.
These birds are slightly heavier than a series from the Yuca-
tan Peninsula. The males weighed 7.6 and 7.8 grams; the female
7.4 grams. Peninsular males ranged from 5.2 to 6.7 grams, with
a mean of 5.96±.18 grams and two females 5.0 and 5.6 grams.
The species was scarce.
Leptopogon amaurocephalus pileatcs Cabanis
1 ?, July 24.
This bird, the only example of the species which was seen, was
in the heavy forest. It weighed 12.7 grams. Tashian (1952)
recorded a female weighing 15.2 grams and Van Tyne (1935) a
male weighing 10.8 grams.
Pipromorpha oleaginea assimilis (Sclater)
19, July 22; 19, July 23; 1 .', July 24; 1$, July 27; 29, July
28; 16 .July 29; 1 <J , Aug. 18.
The most abundant of the Tyrannidae, and possibly of any
268 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
family. It was ubiquitous iu the broadleaf forest aud was found
in lesser numbers in the monte. Although lacking in the pines,
it was a regular visitor to an isolated fruiting tree (Clusia sp.)
at the camp. Males weghed 12.7, 13.7, and 14.1 grams; females
12.2, 12.8, 14.8, and 15.2 grams.
Xanthoura yncas vivida Ridgway
1$, July 21; IS, Aug. 2; 1 <$ , 1$, Aug. 5; 19, Aug. 9; 19,
Aug. 12; 19. Aug. 17.
These specimens are intermediate between X. y. centralis and
X. y. vivida. They are nearer to the latter in that ventrally all
are well washed with green, but few are as green as typical
X. y. vivida. In size they fall within, or above, the upper half
of the range for X. y. centralis, as given by Paynter (1955). All
the specimens are worn or in molt; it is possible that in fresh
plumage there would be less overlap with X. y. centralis.
Two males weighed 72.1 and 87.1 grams ; five females from
84.0 to 88.0 grams, with a mean of 86.08±0.68.
Jays occurred commonly in the selva. They were present, but
less abundant, in the pines and monte.
Theyothorus maculipectus umbrinus Ridgway
1$, 19, July 22; 19, July 28; 19, July 29; 1$, July 30; 13,
Aug. 3 ; 1 $ , Aug. 4 ; 1 6 , 1 9 . Aug. 8 ; 1 <? , Aug. 11 ; 19, Aug. 13 ;
1 9 , Aug. 15 ; 1 9 , Aug. 18.
It is probable that T. m. umbrinus and T. m. canobrunneus
may be distinguishable by a difference in weight. At Laguna
Ocotal, six males ranged from 16.0 to 18.7 grams, with a mean
of 16.76 ±0.47 ; five females from 13.4 to 15.4 grams, with a mean
of 14.44±0.46. On the Yucatan Peninsula (Paynter, 1955),
seven males ranged from 13.3 to 15.0 grams, with a mean of
14.36±0.77; 11 females from 11.9 to 12.8 grams, with a mean of
11.90±0.95.
These wrens ranged through the low forest in abundance.
Occasionally they were found in the high broadleaf forest and
in the brush at the edge of the pine zone.
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 269
Troglodytes musculus ixtermedius Cabanis
1 6 . July 28 ; 1 $ , Aug. 2 ; 19. Aug. 3 ; 1 $ , Aug. 17.
In the scrubby pine burn the species was common, but it was
one of the most difficult birds to collect because of its excessively
shy nature.
At Ocosingo, where one specimen (male ?) was collected
on July 8, and at El Real, where a female and male were taken
on July 13 and 14, the species lived in the vicinity of houses
and behaved like its northern counterpart, T. aedon.
The weights of the males were 11.1, 11.2, 11.4, and 12.0 grams;
those of the females 10.6 and 10.6 grams.
Hexicorhixa leucosticta prostheleuca (Sclater)
1 ?, July 22; 1$, July 23; 19 », July 24; 16, 1 ?, July 25; 1 <3 ,
July 27; 1$, July 30; 19, Aug. 1; 16, 19, Aug. 2; 19, Aug.
5; 1$, Aug. 6; 1$, Aug. 8; 1$, Aug. 14; 16, Aug. 18.
A very abundant resident of the undergrowth in the broad-
leaf forest.
Seven adult males weighed between 14.4 and 17.3 grams,
with a mean of 15.81±.32; five females from 14.1 to 16.7 "rams,
with a mean of 15.40±.41.
Hexicorhixa leucopiirys castaxea Ridgway
1 9, July 2.->; 1 .$, Aug. 2.
These specimens apparently represent the first record of H. I.
castanea from Mexico, although Hellmayr (1934) had predicted
that it would be found there on the Atlantic slope. The respective
weights of the male and female were 16.6 and 16.1 grams.
H. Jeucophnjs and //. leucosticta were found in what seemed
to be exactly the same habitat. No behavioral differences were
noted. The darker breast of H. leucophrys could not -be recog-
nized in the dark undergrowth, making it impossible to dis-
tinguish between the two species. They were, therefore, collected
at random. Presumably the ratio between the species in the
collection also represents the true ratio at Laguna Ocotal.
270 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
MlCROCERCULUS MARGINATUS PHILOMELA (Salvin)
1 ?, July 26.; 1 $ , Aug. 9.
There is no doubt that this wren was uncommon but it was
by no means rare, as the dearth of specimens would seem to
indicate. Its call is distinctive and was heard about once a day
while collecting in the broadleaf forest. It was seldom seen
because of its preference for the darkest areas of the forest
floor, where it blended ideally with the background.
The male weighed 18.1 grams ; the unsexed bird 18.4 grams.
TURDUS ALBICOLLIS LEUCAUCIIEN Sclater
19, July 28; 1$, Aug. 14.
The male retains the juvenal plumage on its throat, upper
breast, wing coverts, neck, and pileum. The remainder of the
plumage is slaty with a faint wash of olive. The other bird,
an adult female, is decidedly olivaceous dorsally ; the throat
markings are brownish black. It agrees with specimens of T. a.
leucauchen from Guatemala.
The weight of the male was 66.7 grams ; that of the female
70.3 grams.
Myadestes unicolor pallens Miller and Griscom
1$, July 31; 16, Aug. 1; 19, Aug. 7; 29, Aug. 11; 19, Aug.
12; IS, Aug. 15; 1$, Aug. 18.
The type of M. u. veraepacis, 56 specimens of M. u. pallens
from Honduras and Nicaragua, one specimen from Veracruz and
three from "Mexico" of M. u. unicolor, and the present series
from Chiapas have been examined. It is concluded that M. u.
pallens is barely distinguishable from the nominate form, on
the basis of its paler ventral color, and that 31. u. veraepacis,
which was described as an intermediate form, is referable to
M. u. pallens. The supposed differences in size between the forms
cannot be confirmed with the present material.
Because this is a montane species, it is presumed that the
population north of the Isthmus of Tehuantepec has no contact
with that which occurs from Chiapas southward. It is, there-
fore, not surprising that the series from Laguna Ocotal is
referred to the more southern population, M. u. pallens. It
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 271
is strange, however, that the species does not subspeciate more
markedly, since it is a member of a genus whose species are
rather plastic.
Adult males weighed 34.1 and 38.2 grams; an adult female
36.1 grams; two males and a female which were in almost com-
plete adult plumage 39.5, 40.7, and 36.3 grams, respectively.
Catharus mexicanus cantator Griscom
16, 19, July 2o; li, July 30; 1$, Aug. 5; 1$, Aug. 12;
19, Aug. 15.
Berlioz (1939) recorded the species from Chiapas for the
first time, but lacking comparative material was unable to assign
his series to a race.
The beautiful song of this thrush was often heard in the
late afternoon and sometimes in the morning. It must have been
fairly abundant, but it was very difficult to approach. Had it
not been for the song, it would have been assumed to be a rare
species. It was collected in the darkest parts of the broadleaf
forest.
A young, spotted female weighed 29.0 grams; two females
which were not quite adult 31.9 and 32.6 grams ; two adult males
and an adult female 33.1, 37\5, and 32.6 grams, respectively.
Smaragdolantus pulchellus pulchellus (Sclater and Salvin)
1 9 , Aug. 3 ; 1 $ , Aug. 12 ; 1 $ , Aug. 13.
Alvarez del Toro (1952) has recorded this species in Chiapas,
apparently for the first time. Blake (1953) also lists the bird
from there but has informed me (in lift.), that his citation
of the race 8. p. verticalis from Chiapas is a lapsus.
Being a species which ranges in the tops of trees it is difficult to
judge its abundance. It seemed to be uncommon. The male
weighed 25.3 grams; the females 24.3 and 26.2 grams.
Hylophilus ochraceiceps ochraceiceps Sclater
15,19, July 25 ; 1$, July 27 ; 1$, Aug. 1 ; 1 9 , Aug. 12.
Prior to Alvarez del Toro's book (1952) the species does not
seem to have been noted from Chiapas. It was common at Laguna
Ocotal in the broadleaf forest.
272 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Males weighed 11.1, 11.4, and 11.5 grams; females 10.5 and
10.8 grams.
Mniotilta varia (Linnaeus)
19, Aug. 12; 19, Aug. 15.
Black and White Warblers were first seen on August 11. The
birds weighed 10.7 and 10.8 grams.
PARULA AMERICANA INORNATA Baird
19, Aug. 12.
Parula "pitiayumi" appears to be only a morphologically pro-
nounced subspecies group of P. americana. Unless sympatry can
be shown to exist, the logical course seems to be to treat the
groups as conspecific.
A pair of the warblers was in a flowering tree in the montc
on August 12. No others were seen while at the lake.
The specimen weighed 6.9 grams, as did. a female collected at
Ocosingo on July 7.
Dendroica graciae decora Ridgway
1 9 , July 22 ; 1$ , July 30 ; 1 ?, Aug. ]8 ; 2 <5 , Aug. 19.
The species was abundant in the tops of the pines. It was
noted in the broadleaf forest a few times.
The specimens exhibit no approach toward D. g. ornata, a
distinctive form, which has been found in western Cbiapas (e.g..
Edwards and Lea, 1955) but for which there seems to be no
published report in eastern Chiapas. In the Museum of Com-
parative Zoology there are, however, specimens referable to this
race from Santa Rosa (Escuintla) and Nuevo Amatenango,
localities near the Guatemalan border.
The respective weights of three males and a female were 7.3,
8.5, 8.7 and 7.6 grams. One bird was host to the hippoboscid
Ornithoctona fusciventris.
Seiurus motacilla (Vieillot)
1 $ , July 25 ; 1 9 , Aug. 2 ; 1 $ , Aug. 13.
After the first of August, Louisiana Waterthrushes were seen
BIOLOGICAL [INVESTIGATIONS IN CHIAPAS, MEXICO 273
at the rate of about one per clay. The specimen collected on
July 25 is a very early arrival, but at Palenque in 1949 Tashian
(1952) observed the species on July 12.
The male weighed 18.4 grams; the females 16.7 and 18.9
grams.
Granatellus sallaei sallaei (Bonaparte)
19 ?, Aug. 10; 1&, Aug. 13.
A rare inhabitant of the low forest at the end of the lake.
The male weighed 14.0 grams, suggesting that this race is heavier
than G. s. boucardi, six males of which are known to have
ranged from 9.2 to 10.6 grams, with a mean of 10.00±.22 (Payn-
ter, 1955). The bird whose sex could not be determined with
certainty weighed 11.6 grams.
I have examined the type of 6?. s. griscomi van Rossem, a
male, and find, as the describer claimed (1934), that dorsally
it is less slaty and gray than most specimens of the species. The
supposed differences in the distribution and shade of the red of
the underparts, and of the gray on the throat, cannot be recog-
nized by me.
The bird was received, as a mount, by the Museum of Com-
parative Zoology in 1880. It is presumed to have been collected
at least a few years earlier. It was, therefore, well over fifty
years old, at a minimum, when named as the type. It is soiled
and has the oily texture that is often noticed in specimens which
have been mounted and on display for many years. The charac-
ters ascribed to the race are without doubt functions of age and
dirt. This belief is strengthened when old and fresh specimens
of G. s. boucardi, or of G. s. sallaei, are compared. The older
specimens are often noticeably darker.
Myioborus miniatus intermedius (Hartlaub)
1 & , July 26; 1 $ , July 27 ; 1 6 , July 31 ; 1 $ , Aug. 3 ; 1 $ , Aug. 13 ;
IS, Aug. 14; 1$, Aug. 16; IS, 29, Aug. 17; IS, Aug. 18;
2 S , Aug. 19.
The redstart was abundant in the selva. Two females weighed
9.0 and 9.1 grams; ten males ranged from 8.2 to 9.6 grams, with
a mean of 8.86±0.14.
274 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Basileuterus culicivorus culicivorus (Lichtenstein)
13, 19. July 22; 19, July 23; 1$, 1 ?, July 24; 1$, July 25;
1$, July 26.
These were among the most abundant birds at the lake. They
occurred in all types of habitat, with the exception of the pines,
although they were most often found in the higher broadleaf
forest.
Females weighed 8.8, 10.0, 10.3, and 10.4 grams ; two males
9.8 grams each.
Basileuterus rufifrons salvini (Cherrie)
1 ?, July 14; 19, July 20; IS, July 23; IS, 19, Aug. 12.
This species replaced B. culicivorus in the pines, where it was
common but usually too high to collect. Two males weighed 11.4
and 11.8 grams; two females 10.3 and 10.8 grams.
Todd (1929) and Griscom (1932) to the contrary, it appears
that Ridgway (1902) was correct in treating delattrii, salvini,
and rufifrons as conspecific. Griscom (1932) chose to regard
each as a distinct species, claiming that all three forms are sym-
patric in the western cordillera of Guatemala, and that salvini
and rufifrons are sympatric in Vera Paz. However, it is sig-
nificant to note that one or another of these forms has been
collected at approximately twenty localities in Guatemala (vide
Griscom, 1932, and Todd, 1929, for lists), but at no given place
has more than a single form been taken. Even when two col-
lectors' stations are adjacent, there seems always to be a dif-
ference in their altitudes. Thus sympatry does not appear to
exist. No intergradation between B. r. delattrii and either B. r.
rufifrons or B. r. salvini is known but this may be of little
significance since even in the comparatively well-studied region
of Veracruz integration between B. r. rufifrons and B. r. scdvini
was undetected until 1943 (Wetmore).
COEREBA FLAVEOLA MEXICANA (Sclater)
1 S , July 28 ; 1 9 , Aug. 11 ; 1 $ , Aug. 16.
Bananquits were rare and found only in the monte.
A mature male and female weighed 10.0 and 8.7 grams, re-
spectively. An immature male 10.7 grams. The adult male had
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 275
fully enlarged testes. It was one of the few species exhibiting
sexual activity at this season.
Amblycercus holosericeus holosericeus (W. Deppe)
1 $ , Aug. 11.
This bird, which weighed 67.0 grams, was found in a dense
tangle of vines near the shore of the lake. No more were seen.
»—
Icterus mesomelas mesomelas (Wagler)
Id, July 27; 1 9 , Aug. 12.
The male weighed 42.7 grams; the female 35.0 grams. Orni-
thoctona fusciventris was found on the latter.
The dearth of clearings meant that habitats for most of the
Icteridae were lacking. The almost total absence of orioles was
one of the impressive ornithological features of the Laguna
Ocotal region.
An oropendola was seen in the forest by one of the party, but
whether it was Zarhynchus wagleri or Gymnostinops montezuma
is unknown.
Tanagra lauta lauta Bangs and Penard
1 d . Aug. 2 ; 1 9 , Aug. 4 ; 1 $, Aug. 17.
One bird was taken in the pines and the others in the low
forest. The species was rather uncommon.
The male collected August 2 had slightly enlarged gonads;
that taken August 17 retained about half of its juvenal plumage
but had fully enlarged testes. Breeding in transitional plumage
has been reported before (e.g., Skutch, 1954).
The first male weighed 14.8 and the second 16.4 grams; the
female 17.6 grams.
Tanagra gouldi gouldi (Sclater)
1$, July 21; Id, July 23; 19, July 26; 16, 19, July 31; 19,
Aug. 4; 16, Aug. 5; 19, Aug. 6; Id, Aug. 14; Id, Aug. 16;
1 d , Aug. 18.
These were the most abundant of the tanagers, ranging through
the broadleaf forest to the edge of the pines.
Six males had a mean weight of 13.73±0.11 grams, with a
276 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
range from 12.7 to 14.5. The mean of five females Mas 14.00:
0.47 grams and their range 12.9 to 15.3.
to'
T ANGARA NIGROCINCTA LARVATA (Dll BllS )
1 6 , 1 ?, July 28 ; 19, Aug. 6 ; 1 $ , Aug. 12; 1 $ , Aug. 13 ; 1 $ . Aug. 18.
The specimens were taken in either the montc or the Clusia
tree at our camp.
Males weighed 18.6 and 19.7 grams; females 20.2, 20.3, and
21.9 (rrams.
oJ
Thraupis abbas (W. Deppe)
1 9 , July 25 ; 1 9 , July 28 ; 9 , Aug. 1 ; 1 9 , Aug. 9 ; 1 $ , Aug. 13.
The species was seldom noted in the tropical evergreen forest,
liut was very abundant in the pines, frequently moving through
the tops of the trees in flocks of about ten individuals.
A male weighed 46.2 grams; females 40.3, 46.2, 47.4, and
48.4 grams.
PlILOGOTIIRAUPIS SANGUINOLEXTA SANGUINOLENTA (LeSSOn)
1 6 , July 25 ; 1 $ , July 2(5 ; 1 $ , Aug. 12 ; 1 ?, Aug. 13.
The weights of the males were 38.8, 40.0, and 44.2 grams.
The species was moderately common but more shy than most
tanagers. It came to exposed areas at times but generally was
present in the heavier selva.
PlRANGA LEUCOPTERA LEUCOPTERA (Trildeau)
1 ?, July 22 ; 1 <J , Aug. 4; 1 $ , Aug. 3 1 ; 1 9 ?, Aug. 16; 2 $ , Aug. 18.
Usually these birds were found in small flocks along the edges
of trails and in the monte.
The mean weight of five males was 16.68±.32. They ranged
from 15.5 to 17.4 grams.
One specimen harbored the bird-fly Ornithoctona fusciventris.
Habia rubica rubicoides (Lafresnaye)
19, July 22; 1$, July 23; 19, July 24; 19, July 30; 16, 29,
July 31; 1$, Aug. 8; 19, Aug. 11; 1$, Aug. 16; 1$, Aug. 17;
3$, Aug. 18.
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 277
This form was more common than H. gutturalis. Both species
occurred throughout the broadleaf forest, with occasional ap-
pearances in the monte. There was no noticeable difference in
habitat preference between the two forms.
These specimens lend credence to the suggestion (Paynter.
1955) that II. r. nelsoni may weigh less than H. r. rubicoides,
although this still cannot be proved. Six adult male H. r. rubi-
coides ranged from 35.8 to 40.5 grams, with a mean of 38.00±.71 ;
five adult females from 27.8 to 33.6 grams, with a mean of
31.04±1.07. Adult males of typical H. r. nelsoni were reported
i Paynter, 1955) to weigh 27.7, 30.4, 31.5, and 32.1 grams; a
female 27.5 grams.
Habia gutturalis salvini (Berlepsch)
1 .', July 23 ; 1$, July 24 ; 1$, July 27 ; 1$, July 30.
The unsexed specimen weighed 42.9 grams; the remaining
birds all of which are immature, 42.7, 43.6, and 43.7 grams.
This series is unsatisfactory for subspecific determination. It
seems, however, to be more similar to immature specimens from
the range of H. g. salvini than to the limited material I have
seen of immature H. g. litioralis, a weak or possibly invalid, race.
Chlorospingus ophthalmicus dwighti Underdown
1;, 1$, July 21; IS, 1 ?, July 22; 26, 19, July 23; IS,
1$, July 24; IS, Aug. 1; 1$, Aug. 2; 1?, Aug. 7; 1 ?, Aug. 10.
Along the edges of trails and within the broadleaf forest,
wherever there was low vegetation, these birds swarmed.
Adult males weighed 18.6, 18.7, and 19.7 grams; females 17.2
and 17.4 grams. Juvenal males weighed 18.6, 18.6, 18.7, and
18.8 grams; females 16.0 and 17.3 grams.
Chlorophanes spiza guatemalensis Sclater
1 9 , July 81.
Only this bird was seen. It weighed 22.1 grams and was fatty.
Cyanerpes cyaneus carneipes (Sclater)
16, July 20; IS, July 24; IS, July 25; 1$, July 30; 1$, Aug. 18.
The species was most often seen in the Clusia tree at the camp.
278 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Adult males weighed 13.1 and 13.3 grams; an immature male
12.2 grams; females 12.3 and 13.4 grams.
Saltator atriceps atriceps (Lesson)
1 <S , Aug. 9 ; 1 9 , Aug. 10.
The gonads of the male were very enlarged and that of the
female moderately enlarged.
None of the saltators was common, but this was the most
frequently observed of the three. All the forms frequented the
brush where the broadleaf and pine forests joined.
The weights of the male and female were 77.3 and 85.5 grams,
respectively.
Saltator maximus gigantoides Cabanis
1 9 , Aug. 3 ; 1 $ , Aug. 12.
These specimens have only a trace of green on the pelium,
leaving no doubt that they are referable to this race rather than
to S. m. magnoides, which has been recorded from near Comitan
(Berlioz, 1939).
The male, which had enlarged testes, weighed 52.3 grams.
The female, whose ovary was slightly enlarged, weighed 48.4
grams.
Saltator coerulescens yucatenensis Berlepsch
1 6 , July 27.
To find this race at Laguna Ocotal was unexpected, but the
specimen is pale, has only a faint wash of buffy on the breast,
and has a light crissum, all characters differentiating S. c.
yucatenensis from 8. c. grandis. Although it may be an aber-
rant bird, additional specimens from the region will probably
confirm the identification, inasmuch as Brodkorb (1943) found
a series from eastern Tabasco to be closer to S. c. yucatenensis
than to 8. c. grandis.
The bird, which was reproduetively active, weighed 54.2 grams.
Caryothraustes poliogaster poliogaster (Du Bus)
1$, 19, Aug. 10; 19, Aug. 18.
Flocks of these birds were seen regularly in one area of transi-
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 270
tion between the selva and ocotal, where the vegetation was
fairly high but not dense. They were seldom seen elsewhere.
The male weighed 48.6 grams ; the females 49.3 and 50.7 grams.
Cyanocompsa cyanoides concreta (Du Bus)
1 $ , July 27 ; 1 $ , July 31 ; 1$, Aug. 14; 1$, Aug. 19.
The male taken July 27 had enlarged testes. That collected
on August 14, an immature bird, was host to Ornithoctona
fusciventris.
Adult males weighed 30.5 and 30.9 grams; an immature
male 27.6 grams; an adult female 30.4 grams.
Sporophila torqueola morelleti (Bonaparte)
1 $ , July 26 ; 1 $ , Aug. 8 ; 1 $ , Aug. 13.
Iii the vicinity of the marsh near the campsite, seedeaters
were seen and heard daily. A few were also found in the pine
burn, but none elsewhere.
The bird taken on August 13, which is immature, weighed
8.7 grams. The remaining birds were in full breeding condition
and weighed 8.9 and 9.6 grams.
Oryzoborus funereus Sclater
13,19, Aug. 2.
The birds, both of which had fully enlarged gonads, were
collected in the pine burn. The area was searched for additional
birds, but this pair was the only one found. The species was
reported from Chiapas once before (Brodkorb, 1943).
The weights of the male and female were 14.3 and 14.7 grams,
respectively.
Atlapetes albinucha (Lafresnaye and d'Orbigny)
1$, July 30; 1 $ , Aug. 3.
The August 3 specimen is a fledgling, which was barely able
to fly ; the other a juvenal. Both birds were taken in the under-
brush of the pine burn. Only one other was seen during the
month at the site.
The fledgling weighed 32.5 and the juvenal 33.2 grams.
2H0 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
DISCUSSION
One hundred and twenty-two species of birds were collected
or observed during the period spent at Laguna Ocotal. Of this
number, 11!) are presumed to breed in the vicinity and three are
known to be visitors (Empidonax minimus, Mniotilta varia, and
Seiurus motacilla).
The avifauna of the lowlands to the north and to the east
of the Selva Lacandona is fairly well known, affording a useful
basis for comparison with that which was found at Laguna
Ocotal. Brodkorb (1943) has surveyed the birds of Tabasco and
adjacent areas in Veracruz, Campeche, and Chiapas; Tashian's
report (1952) on the birds of Palenque, Chiapas, is an informa-
tive supplement. To the east, the district of Peten, Guatemala,
has been investigated by Van Tyne (1935) and Taibel (1955).
The localities encompassed by these papers are situated at alti-
tudes below 250 meters, with a few insignificant exceptions. A
large part of the region is covered by "high evergreen forest,"
to use the terminology of Miranda (1952). However, the com-
position of the forest differs from that at Laguna Ocotal and
is the type commonly called "rain forest." The distinction
between the high evergreen forest at Laguna Ocotal and that
which is found at lower altitudes is considered in the paper in-
troducing this series pp. 193-199).
The birds in the highlands contiguous with the Selva Lacan-
dona are not so well known. Berlioz's (1939) report on a collec-
tion from the district of Comitan is the nearest approach to a
detailed study, but it seems that the collector (Mario del Toro
Aviles) was selective and did not attempt to obtain repre-
sentatives of all the species in the region. The paucity of rap-
torial and aquatic species is particularly noticeable. The col-
lector spent a few days, in April and October, at Comitan (alt.
1650 m.), a sizable town on a plain, with oak and coniferous
forests a short distance away. He collected for 20 days in August
and for four days in October at Juncana (alt. 1(345 m.), a village
situated in the oaks and pines, 40 kilometers southeast of Comi-
tan. From May to August he worked at Santa Rosa, which,
Prof. Brodkorb {in litt.) informs me, is a finca, at an altitude
of about 1200 meters, near the Guatemala border and east of
Laguna Montebello, roughly seventy kilometers southeast of
BIOLOGICAL INVESTIGATIONS IX CHIAPAS, MEXICO 281
Coniitan. According to Prof. Brodkorb, this region is character-
ized by high rainfall and by the presence of the Sweet Gum
(Liquidambar styraciflua). Miranda (1952) maps the area
within the zone of "temperate deciduous woods," one of whose
key species is Liquidambar.
When the resident birds at Laguna Ocotal are compared with
those known from adjacent Veracruz, Tabasco, Chiapas, Cam-
peche, and Peten (Brodkorb, 1943; Tashian, 1952; Van Tyne,
1935; Taibel, 1955), it is seen (Table 1, p. 284) that 24 of the 119
species (22 per cent) have not been found in the lowlands en-
compassed by these studies. Of these 24 birds, four (Spizaetus
tyr annus, Micrastur semitorquatus, 31. ruficollis, and Chloro-
phanes spiza) are uncommon species which have been collected
at low altitudes nearby, e.g., on the Yucatan Peninsula (Payn-
ter, 1955), and in time probably will be found in the regions
under consideration. The 20 species remaining are characteristic
of higher elevations, with none having been found below roughly
300 meters, at least in northern Middle America.
With the exception of the pines, the vegetation at Laguna
Ocotal is pbysiognomically similar to that of the lowlands.
Therefore, the presence of the conifers might logically seem
to be the cause of the difference in composition of the two
avifaunas. However, this is not the case. Only four species
are confined to the pines at Laguna Ocotal. These are Lepidoco-
laptes souleyetii, L. affinis, Contopus pertinax, and Basileuterus
rufifrons; Dendroica graciae was abundant in the pines but was
seen in the broadleaf forest on a few occasions. Of these five
species, only Lepidocolaptes affinis and Dendroica graciae do not
occur in the adjacent lowlands. More subtle differences in the
environments must be responsible for the discrepancy between
the avifaunas of the two regions.
If the resident birds of Laguna Ocotal are compared with those
listed by Berlioz (1939) for the District of Comitan, Chiapas
(2100 to 1650 m.), it is seen (Table 2, p. 285) that 54 species (45
per cent) have not been recorded at the higher elevations. Owing
to what certainly was discriminating collecting in the highlands
(e.g., Cathartes aura is not recorded), the difference between
the two avifaunas is doubtless exaggerated, but still it must be
substantial. Most of the species in Table 2 are forms which
frequent lower altitudes and would be unexpected in the District
282 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
of Comitan. It is interesting to note that of the 24 birds listed
in Table 1 as not occurring in the nearby lowlands, 18 are re-
corded in the highlands (Berlioz, 1939), including the so-called
"lowland forms" Micrastur ruficollis and Chlorophanes spiza.
From this it may be concluded that the composition of the
avifauna of Laguna Ocotal is intermediate between that of the
adjacent lowlands and highlands, with a stronger element from
the lowlands. The presence of pines at Laguna Ocotal, the one
strikingly marked vegetational difference between the lowlands
and the lake district, has almost no effect on the composition of
the avifauna.
LITERATUEE CITED
Alvarez del Toro, Miguel
1952. Los Animales Silvestres de Chiapas, Tuxtla -Gutierrez, Chiapas,
Dept. de Prensa y Turismo, 247 pp.
Amadon, Dean
19-49. Notes on Harpyhaliaetus. Auk, 66: 53-56.
and Don B. Eckelberry
1955. Observations on Mexican Birds. Condor. 57: 65-80.
Berlioz, J.
1939. Etude d'une Collection d'Oiseaux du Chiapas (Mexique). Bull.
Mus. Nat. Hist. Natur. (Paris), (2), 11: 360-377.
1952. Etude Critique des Formes de Momotus momota (L.). Oiseau,
22: 20-33.
Blake, E. E.
1953. Birds of Mexico; a Guide for Field Identification. Chicago.
Univ. Chicago Press, xxix -j- 644 pp.
Brodkorb, Pierce
1943. Birds from the Gulf Lowlands of Southern Mexico. Univ. Mich.,
Mus. Zool., Misc. Pub., No. 55, 88 pp.
Howards, E. P. and E. B. Lea
1955. Birds of the Monserrate Area. Chiapas, Mexico. Condor, 57:
31-54.
BIOLOGICAL. INVESTIGATIONS IN CHIAPAS, MEXICO 288
Friedmann, Herbert, Ludlow Griscom, and R. T. Moore
1950. Distributional Cheek-list of the Birds of Mexico, Part I. Berke-
ley, Calif. Cooper Ornithological Club, Pacific Coast Avifauna,
No. 29. 202 pp.
Griscom, Ludlow
1932. The Distribution of Bird-life in Guatemala. Bull. Am. Mus. Nat.
Hist.. 64. vi + 439 pp.
Hellmayr, C. E.
1934. Catalogue of Birds of the Americas and Adjacent Islands.
Field Mus. Nat. Hist., Pub. 330, Zool. Ser., 13 (7), vi 4- 531 pp.
Martin, P. 8., C. R. Robins, and W. B. Heed
1954. Birds and Biogeography of the Sierra de Tamaulipas, an Isolated
Pine-Oak Habitat. Wilson Bull., 66: 38-57.
Miranda, Faustino
1952. La Vegetacion de Chiapas, Parte 1. Tuxtla-Gutierrez, Chiapas,
Dept. de Prensa y Turismo. 334 pp.
Paynter, R. A., Jr.
1955. The Ornithogeography of the Yucatan Peninsula. Bull. Peabody
Mus. Nat. Hist., Yale Univ., 9: 1 347.
Peters, J. L.
1948. Check-list of Birds of the World. Yol. t5. Cambridge, Mass.
Harvard ITniv. Press, xi -4- 259 pp.
Ridgway, Robert
1902. Birds of North and Middle America. Bull. U. S. Nat. Mus..
SO (2), xx + 834 pp.
Skutch, A. F.
1954. Life Histories of Central American Birds, Berkeley, Calif.
Cooper Ornithological Hoc. Pacific Coast Avifauna, No. 31.
448 pp.
Taibel, A. M.
1955. Uccelli del Guatemala con Speciale Riguardo alia Regione de!
Peten Raccolti dal Maggio al Settembre 1932. Atti Soc. Ttal.
Sei. Nat.. 94: 15-84.
284
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
TASHIAN, E. E.
1952. Some Birds from the Palenque Eegion of Northeastern Chiapas,
Mexico. Auk, 69:. 60-66.
Todd, W. E. C.
1929. A Revision of the Wood-warbler Genus Basileuterus and its
Allies. Proc. U. S. Nat. Mus., 74: 1-95.
Traylor, M. A., Jr.
1941. Birds from the Yucatan Peninsula, Field Mus. Nat. Hist., Pub.
483, Zool. Ser., 24 (19) : 195-225.
VAN ROSSEM, A. J.
1934. Critical Notes on Middle American Birds. Bull. Mus. Comp.
Zool., 77: 387-490.
van Tyne, Josselyn
1935. The Birds of Northern Peten, Guatemala. Univ. Mich., Mus.
Zool., Misc. Pubs. No. 27, 46 pp.
Wetmore, Alexander
1927. The Birds of Porto Rico and the Virgin Islands. New York
Acad. ScL, Sei. Surv. Porto Rico and Virgin Ids., 9: 245-596.
1943. The Birds of Southern Veracruz, Mexico. Proc. U. S. Nat. Mus.,
93: 215 340.
Table 1
Species of birds found at Laguna Oeotal but not recorded from adjacent
lowlands.
Accipiter striatum
Spisaetus tyrann us
Micrastur semitorquat us
Miorastur ruficollis*
Abeillia abeillei'
Amazilia beryllma
Eupherusa eximia*
Lampornis viridi- pattens*
Xiphorhynchus erythropygius*
Lepidocolaptes affinis*
Anabacerthia striaticollis*
Sclerurus mexieanus*
M yr mo t h er u la schist i color*
E mpidonax flavescens*
.Uicrocerculus mar g hiatus*
Myadestes imicolor*
( 'atharus mexieanus*
Sinargdolan ins piilchellus
Parula amcricana
Dendroica graciae'~
Myioborus miniatvs*
Clilorospingus ophthalmicus*
Clilorophaoies spisa*
A t lapet.es alb inu oha *
•Recorded from Comitan District (Berlioz, 1939).
BIOLOGICAL INVESTIGATIONS IN CHIAPAS. MEXICO
285
Species of birds found at L
highlands.
Phalacrocorax brasilian us
B 1/ tor ides v-irescens
Sarcoramphus papa
Cathartes aura
ElanoMes forficatus
Accipiter striatus
B u t eoga U us u ru b itinga
Spizaetus tyrannus
Hcrpetothercs cacliinnans
Micrastur sem itorquatus
Crax rubra
I't nelope purpurascens
Ortalis vetula
Aramus guarauna
Aramides cajanea
Laterallus ruber
Colvmba n igrirostris
Ara macao
Amasilia Candida
Amasilia beryllina
Trogon massena
Trogon violaceus
Chloroceryle aenea
Pteroglossus torquatus
Haitiphastos sulfuratus
<\ litmus pucherani
I'cuiliontis fumigatus
Table 2
aguna Ocotal but not recorded from adjacent
U( ndrocincla amabatiiui
Dendrocinela homochroa
Dendrocolaptes certhia
Lepidocolaptes souleyetii
Xenops minutus
Dysitliamn us mentalis
J'hitypsaris aglauie
Tyrannus melancholieus
Megarhynchus pitang ua
Myioset.etes similis
Myiobius barbatus
Onychorhynchus eoronat-us
Lcptopogcm amaurocephalus
Xanthoura yncas
Heni-corhina Icucosticta
Smaragdolanim pulchellvs
ILylophilus ochraceiceps
Parula americana
Granatellus sallaei
Icterus mesomeias
Tanagra lauta
Tanagra gouldi
Tangara nigrocincta
Habia gutturalis
Saltatar coerulescens
( 'yanocompsa oyanoides
Oryzoborus funereus
286 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
VIII
DESIGN QUANTITIES OF SOME CHIAPAS BIRDS
By
Charles H. Blake
What an airplane or a bird is capable of in the way of flight
is largely deducible from a few rather simple ratios. Obviously,
ability to perform as well as the design permits depends on an
adequate power plant and a sufficiently strong and responsive
control system. We do not yet know how to assess these latter
properties in a bird, but the design quality of the wings can be
discussed in general from the quantities here presented.
Three quantities are considered here : wing loading, span load-
ing, and aspect ratio. These are derived from three measure-
ments : weight, wing area, and wing span. English units are used
to render comparison with airplanes easier.
The area is that of the slightly flattened wings. Ideally it
should be the projected area of the wings fully outstretched in
flight. This can only be closely approximated.
The span is twice the length of the wing beyond the edge of
the body in the attitude in which the area is measured. This is
less than the tip to tip distance or wing expanse.
Wing loading is expressed as pounds of weight per square foot
of wing area. Similarly span loading is in pounds per foot of
span.
The aspect ratio is dimensionless and most conveniently cal-
culated as the square of the span divided by the wing area.
The wing loading is an indicator of the relative power required
for acceleration. Takeoff and climbing are more difficult with
higher wing loading. On the other hand, stability varies in the
same sense as wing loading. Span loading is a measure of
maneuverability. Birds with high span loadings will have diffi-
culty in turning rapidly unless the aspect ratio is very low. The
aspect ratio is related to lift. The higher the aspect ratio the
greater the lift in proportion to the wing area. In general, birds
with high aspect ratios glide and soar readily or remain in the
air for long periods or both. It is also generally true that a high
aspect ratio improves stability at low speeds. Similar effects
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 287
can be obtained by high wing camber or an appropriate dihedral
angle between the wings.
I am greatly indebted to Raymond A. Paynter, Jr. for making
(in the field) the outlines of the wings of 21 birds comprising 18
species, shown in the table, and recording the weights of the
individuals drawn. The series is important because it gives us
our first information on almost all the included families, and
it is the first sizable series of sedentary species. I also thank
the Geological Survey Department, Jamaica, B. W. I., for the
use of a planimeter.
There is, of course, great risk in speaking definitely about the
flight of species one has never seen alive. Some general remarks
may be made on the basis of the quantities themselves. The wing
loadings are very diverse. Tinamus has a loading that is high
even for a bird weighing nearly 2y2 pounds. By contrast, the
specimen of Ortalis has less than one-third the weight but more
than two-thirds the wing area of the Tinamus. One would readily
believe that Tinamus would take off with difficulty and even re-
luctantly. It might, however, fly well on a straightaway after it
was up to speed. This is a matter of motor rather than wings.
If the motor is only sufficiently powerful to maintain cruising
speed, then takeoff becomes virtually impossible. This is probably
not quite true of any bird. With a high wing loading consid-
erable excess power must be available. These remarks also
apply, with less force, to the toucan, Pteroglossus, and the parrot,
Pionus. If the figures for other parrots are similar to those for
Pionus, their rather labored, although rapid, flight is under-
standable. At the other extreme are a hummingbird, Abeillia,
and a woodhewer, Sittasomus, whose wing loadings are close to
the lowest on record. Their flight should be about equally un-
stable but in other respects entirely different.
Turning to the span loadings, Tinamus is again unusual in its
high loading. No other bird in the list exceeds 0.5 and span
loadings above this latter figure are very rare. The two hum-
mingbirds show the lowest loadings. It should be noted that
Eupherusa has about twice the wing loading as well as about
twice the span loading of Abeillia. It would be expected, a
priori, that the flight of these two birds would be quite different.
Abeillia ought to be the more maneuverable. It is possible that
288 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
the curious flitting flight occasionally shown by some of the larger
hummingbirds, notably Trochilus and Anthracothorax, is con-
nected with the very low loadings and high aspect ratios.
The aspect ratio shows clearly that only the hummingbirds
would be likely to fly continuously for any length of time. But
two other species in the list even attain a ratio of 5.0 and one
of these is the parrot. These low ratios may well be characteristic
of nearly non-migratory species. Generally the aspect ratio of
North American passerines seems to be from 5.5 to 7. I have
recently measured a specimen of Coccyzus americanus, migratory
but not appearing to be a strong flyer, and found an aspect ratio
of 5.9.
In this connection it is of more than incidental interest that
one of Otto Lilienthal's later gliders had an aspect ratio of
about Sy2. Such a machine would have very low stability at low
speeds and his control system was slow and cumbersome. It
is no wonder that he met his death in a crash. On the other
hand, a small bird has such low terminal velocity that the legs
can take up the shock of a fall from almost any height. The
bird gets into trouble by running into some barrier beak first
in full flight or attempting to land while its forward speed is
much in excess of stall speed.
In two cases, as shown, two individuals of the same sex were
available. The derived ratios agree within 10 per cent; quite a
reasonable agreement.
With a few exceptions, the general picture is one of low speed,
short and unstable flight. It is probable that most of these birds
do not venture out in the open where they might encounter
both wind and turbulence.
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 289
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290 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
IX
MAMMALS COLLECTED AT LAGUNA OCOTAL
By
Prances L. Burnett and Charles P. Lyman
This collection was made at Laguna Ocotal, Chiapas, Mexico,
between July 22 and August 19, 1954, by K. A. Paynter, Jr.,
and Elisha P. Lee. The specimens were all collected in the
vicinity of the camp, and consist of skins and skulls, unless other-
wise noted.
The mammals are for the most part lowland forms, with a
few subspecies which are so widespread that they are found in
the highlands as well. In subspecies with restricted ranges, the
races from Laguna Ocotal are in general similar to those col-
lected by Kuns and Tashian (1954) from Palenque to the
northwest, while Murie's (1935) collection from Uaxactun,
Peten, Guatemala, to the northeast contains more species endemic
to the Yucatan Peninsula. Of the mammals which tend to be
restricted to the Yucatan Peninsula, the fruit bat Artibeus
jamaicensis yucatanicus is the only race represented in the col-
lection from Laguna Ocotal.
We wish to thank Dr. Charles 0. Handley, Jr., of the United
States National Museum for comparing our spider monkey with
material in the United States National Museum. Also we grate-
fully acknowledge the loan of comparative material from the
following institutions: Museum of Zoology, Ann Arbor, Michi-
gan; American Museum of Natural History, New York City;
Pish and Wildlife Service, United States Department of the
Interior, Washington, D.C.; Chicago Museum of Natural His-
tory, Chicago, Illinois; and Museum of Natural History, Law-
rence, Kansas. In particular, we thank Miss Barbara Lawrence
for her help in preparation of the manuscript.
Didelphis marsupialis ?subsp.
15,299 (M.C.Z. 47274-47276)
These young opossums are relatively too large to be yuca-
tanensis and could be assigned to tabascensis on geographic
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 291
grounds. One difference between the latter subspecies and
mesamericana (—marsupialis Allen, 1901) was said to be the
greater length of tail compared to head and body. Using Allen's
tables it is found that the average of the individual ratios of tail
length to head and body length for 42 mesamericana is 89.8
(expressed in per cent) with a standard deviation of ±7.5 while
the ratio for 21 tdbascensis is 91.9 with a standard deviation of
±11.5. Obviously this difference has no statistical value. The
Laguna Ocotal specimens have ratios of tail length to head and
body length of 84, 88 and 91.
Long nasals, terminating posteriorly in a sharp point, were
also used to characterize tdbascensis. However, in four specimens
in the M.C.Z. collection, all taken within a few days in the
vicinity of Cordoba, Veracruz, one (M.C.Z. 39772) has nasals
which come to a sharp point in the midline, while the posterior
portion of the nasals of another (M.C.Z. 39770) are rounded in
outline. The two others (M.C.Z. 39771 and 39767) have nasals
which are intermediate between the extremes. There is no indi-
cation that the shape of the nasals is dependent on the age or
the sex of the animals. As the length of the nasals depends
partly on the shape of their posterior border, measurements of
nasal length on skulls of the same size would then vary according
to the outline.
Our evidence supports Hershkovitz's (1951) opinion that there
is only one widespread form of Did el phis marsupialis in Mexico
and in Central America north of the highlands of Nicaragua.
Artibeus jamaicensis yucatanicus J. A. Allen
4$ 6 ,32 9 (M.C.Z. -47278-47284)
The measurements of these fruit bats correspond with those
that Andersen (1908) gives for yucatanicus rather than those
for the larger race, jamaicensis. Andersen did not group his
results according to sex, but forearm length and various skull
measurements of the Laguna Ocotal specimens indicate that the
females tend to be smaller than the males. Our Chiapas skulls
are smaller than jamaicensis skulls of the same sex from Vera-
cruz, British Honduras, and Honduras.
Other scattered records indicate that this subspecies ranges
into northern Guatemala (Murie, 1935) and northeastern Chiapas
(Kuns and Tashian, 1954).
292 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Alouatta palliata pigra Lawrence
■2 6 6,19 (M.C.Z. 4726G-47268) Skins and skeletons
This series of howler monkeys was all collected in August from
a band of five individuals. The skin of the female is typically
black and in this resembles a male topotype from British Hon-
duras collected in April. The males, on the other hand, have an
extensive region between the shoulders, fading to a point in the
middle of the back, where the bases of the hairs are pale. In
addition, similarly colored hairs are interspersed among the
darker hairs under the forearms of one of these skins. All have
longer, denser hair than has the single topotype. The skulls of
the males are even more massive than the topotype while that of
the female is, as would be expected, considerably smaller.
The collection of these specimens extends the range of pigra
125 miles to the west and south of the type locality of Uaxactun,
Guatemala. Kuns and Tashian (1954) report a specimen from
Palenque, Chiapas as mexicana.
Ateles geoffroyi vellerosus Gray
1 9 (M.C.Z. 47269) Skin and skeleton
Dr. Charles 0. Handley, Jr. very kindly identified this speci-
men of spider monkey as vellerosus through comparison with
material in the United States National Museum. He writes that
the Laguna Ocotal skin is very similar to one from Palenque
in Chiapas, but somewhat darker and more richly colored than
specimens from Oaxaca and Veracruz. No comparison was
needed with pan from Guatemala, which is very much blacker,
nor yucatanensis which is smaller and paler. This specimen was
collected well within the range of this widespread subspecies
(Kellogg and Goldman, 1944).
Paynter reports that there were fewer spider monkeys than
howler monkeys at the lake. Small bands of three or four Ateles
were seen every few days, while bands of howler monkeys could
be heard around the camp almost every evening.
Sciurus deppei deppei Peters
5 $ $ , 3 $ 9 (M.C.Z. 47289-47296)
These small tree squirrels are as dark as typical deppei from
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 293
central Veracruz but are a little smaller when compared to the
measurements given by Nelson (1899). Apparently the north-
easterly extent of the range of this subspecies runs approxi-
mately from the western side of Lago de Izabal (Goodwin, 1984 I
north west through Laguna Ocotal to Palenque (Kuns and Tash-
ian, 1954). To the northeast Murie (1935) found the brighter
colored vivax. At Laguna Ocotal the habitat of this squirrel was
restricted to the tropical evergreen forest.
SCIUBUS AUREOGASTER AUREOGASTER F. Clivier
16,299 (M.C.Z. 47286-47288)
The collection of fire-bellied squirrels at Laguna Ocotal extends
the limits of their range more to the south and closer to the
Guatemalan border, though Dr. Remington Kellogg and Dr.
Charles 0. Handley, Jr. write us that no specimen of aureogaster
has yet been taken in Guatemala proper. In comparison with
more northern specimens no noticeable differences in the skulls
could be found, but these Chiapas squirrels are not as dark in
color. However, as Kelson (1952) has pointed out, aureogaster
varies considerably in the degree of melanism in various areas
of its range.
Paynter found these squirrels on the forest edges, amid thick
tangles of small trees and vines, in contrast to the deep forest
habitat of deppei.
HETEROMYS DESJIARESTIANIS DESMARESTIANUS Gray
1 : . 1 9 skins and skeletons, 1 9 (M.C.Z. 47298-47300;
Laguna Ocotal is near the central point of the range of this
widespread subspecies, and the specimens taken match Goldman 's
1M11 ) description.
Oryzomys colesi couesi (Alston)
30 & i . S.\ 9 9 , 1 .' sex (M.C.Z. 47331-47354, 47356-4738.3;
Fifty-one skins and skulls, as well as three skulls only, were
collected of this larger rice rat. Most of these animals were quite
young, ranging from sub-adults with the upper third molars
just erupting to adults with slightly worn molars. The pelage
of these animals is darker than that of specimens from Yaruca,
294 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Honduras, which Goldman (1918) considered typical couesi. This
darker cast is due to a more plumbeous color in the bases of the
dorsal hairs, with the guard hairs being black instead of brown-
ish. A detailed comparison of the skulls of these specimens with
over 90 couesi of similar ages (as indicated by tooth wear) from
other localities revealed no significant differences in the Laguna
Ocotal specimens. Topotypical adults of pinicola from Pine
Ridge, British Honduras, were smaller than mature but younger
animals from Laguna Ocotal.
0. c. couesi appears to be a very widespread subspecies, and,
except for the somewhat darker color, our specimens are typical
of the race.
Oryzomys alfaroi palatinus Merriam
9 6 $, 7 9 9 (M.C.Z. 47303-47305, 47319-47330, 47355)
Although some of these smaller rice rats have a dorsal pelage
dark enough to fit the description of saturatior, others are almost
as pale as Panamanian specimens of alfaroi, which palatinus re-
sembles in coloration. All of the Laguna Ocotal animals have
whitish underparts, rather than the buffy ones ascribed to satura-
tior, and are hence referred to palatinus. As Goldman points out
(1918), palatinus and saturatior inhabit the same mountain
range, and intergradation between the two races is probable.
Ototylomys phyllotis guatemalae Thomas
:'>i S (M.C.Z. 47301, 47302, 47386)
Two skins and skulls, and an additional single skull, were
taken of this smaller genus of tree climbing rats. The diagnostic
characters given by early authors (Merriam, 1901, Thomas, 1909,
and Sanborn, 1935) do not hold when applied to the present
specimens and the descriptions by Laurie (1953) were not
sufficiently definitive to clarify the situation. Furthermore, the
published records of the distribution of this group are confused.
Although these animals are generally rare in collections, it
was possible to borrow seventy specimens, including one series
from Esmeralda, Quintana Poo, Mexico, and another from Uax-
actun, Peten, Guatemala. Examination of these specimens
showed that Ototylomys could be divided into two groups. One
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 295
group consists of a smaller animal with an upper molar tooth
row measuring 5.9 to 6.5 mm., from the Yucatan Peninsula south
to Peten, and the other group consists of a larger animal with
an upper tooth row of 6.9 to 7.5 mm., from Chiapas and Alta
Vera Paz. Our specimens belong to the group of larger mammals
which also include connectens and guatemalae. Sanborn states
that connectens is the only race in which the belly hairs have
slaty bases. However, in eight specimens within the stated range
of connectens the bases of the hair of the belly vary from slaty
gray (Finca Chama and Chimoxan, Guatemala) to almost pure
white (Concepcion). Furthermore, two specimens of guatemalae
from Palenque, Chiapas, show a slaty cast of the belly fur,
particularly in the midline.
Our specimens from Laguna Ocotal also have slaty based fur on
the belly, otherwise they are closest to Thomas' description of
guatemalae. They are too small to be referred to Sanborn's
connectens and hence are considered to be the former race. It
seems apparent, however, that Ototylomys shows considerable
variation within a relatively small geographic area, and it is
suspected that some of the characters which have been used to
separate races will be found not to hold when more specimens
are available for comparison.
Peromyscus mexicanus teapensis Osgood
8 $ $ , 3 9 5 (M.C.Z. 47308-47318)
According to Osgood (1909) the various races of mexicanus
are only slightly differentiated. The Laguna Ocotal specimens
are referred to teapensis on the basis of color as well as on
geographic grounds. Although the pelage is very much darker,
the skulls closely resemble a series of mexicanus from Veracruz,
in that they lack the broader nasals, heavier rostrum, and more
massive molars ascribed to teapensis. Kims and Tashian (1954),
while identifying their specimens from Palenque as teapensis,
also noted that the skulls showed no evidence of the thickened
rostrum.
296 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Sigmodon hispidus saturatus V. Bailey
29 9 (M.C.Z. 47306, 47307)
One adult cotton rat and one subadult were collected. Both
animals are very similar to specimens in the M.C.Z. collection
from British Honduras, except that the tips of the belly hairs of
the adult are generally more whitish than yellowish. Laguna
Ocotal lies well within the range of this widespread subspecies.
Nasua narica narica (Linnaeus)
19 (M.C.Z. 47277)
One very young coati with the third molars still unerupted
resembles very closely older specimens of narica in color of
pelage.
Odocoileus virginianus subsp.
Robert Dressier found fragments of two tibiae, two femora, one
scapula and one vertebra (M.C.Z. 47476-47481) of white-tailed
deer in an Indian rock shelter near the camp at Laguna Ocotal.
Paynter found no evidence that this deer occurs naturally in
the area, and it appears probable that the remains may have been
carried there by travelling Indians. Associated with the bones
were shells of varieties of snails found in the nearb}^ lake and
more distant streams.
Mazama sp.
Bones of these small deer were also found in the rock shelter
by Dressier, along with those of the larger white-tail. Mazama
fragments consisted of a ramus, a scapula, and a vertebra (M.C.Z."
47482-47484).
Brocket deer were seen by Paynter several times in the "burn
near the camp at Laguna Ocotal. One particular spot was
noted where single animals were found bedded down on a num-
ber of occasions.
Dasypus novemcinctus subsp.
The shell of one animal was found near camp, and one live
armadillo was seen in the area.
BIOLOGICAL INVESTIGATIONS IN CHIAPAS, MEXICO 297
TAPIRELLA BAIRDII (Gill)
One tapir was seen, but not collected, at El Censo. Footprints
were seen around the shore of Laguna Ocotal. Natives reported
that they were very numerous around the lake during the dry
season.
LIST OF REFERENCES
Allen, J. A.
1901. A preliminary study of the North American opossums of the
genua Didelphis. Bull. Amer. Mus. Nat. Hist., 14:149-188.
1902. A preliminary study of the South American opossums of the
genus Didelphis. Bull. Amer. Mus. Nat. Hist., 16:249-279.
Andersen, Knud
L908. A monograph of the ehiropteran genera Uroderma, Emchisthenes,
and Artibeus. Proc. Zool. Soc. London, 1908, pp. 204-319.
Bailey, Vernon
1902. Synopsis of the North American species of Sigmodon. Proc.
Biol. Soc. Washington, 15:101-116.
Goldman, Edward A.
1911. Revision of the spiny pocket mice (genera Heteromys and
Liomys). U. S. Dept. Agric, North Amer. Fauna no. 34, 70 pp.
1918. The rice rats of North America (genus Orysomys) . U. S. Dept.
Agric, North Amer. Fauna no. 43, 100 pp.
Goodwin, George G.
1934. Mammals collected by A. W. Anthony in Guatemala, 1924-1928.
Bull. Amer. Mus. Nat. Hist., 68(1) : 1-60.
Hershkovitz, Philip
1951. Mammals from British Honduras, Mexico, Jamaica and Haiti.
Chicago Nat. Hist. Mus., Fieldiana: Zoology, 31 (47) : 547-569.
Kellogg, Remington, and E. A. Goldman
1944. Review of the spider monkeys. Proc. U.S. Nat. Mus., 96(3186) :
1-45.
Kelson, Keith R.
19o2. The subspecies of the Mexican red-bellied squirrel, Sciurus
aureogaster. Univ. Kansas Publ., Mus. Nat. Hist., 5(17) :243-
250.
298 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Kuns, Merle L., and Richard E. Tashian
1954. Notes on mammals from northern Chiapas, Mexico. Jour.
Mamm., 35(1): 100-103.
Laurie, Eleanor M. O.
1953. Rodents from British Honduras, Mexico, Trinidad, Haiti and
Jamaica collected by Mr. I. T. Sanderson. Ann. Mag. Nat. Hist.,
(12)6:382-394.
Lawrence, Barbara
1933. Howler monkeys of the palliata group. Bull. Mus. Comp. Zool.,
75(8):315-354.
Merriam, C. Hart
1901. Seven new mammals from Mexico, including a new genus of
rodents. Proc. Washington Acad. Sci., 3:559-563.
Murie, Adolph
1935. Mammals from Guatemala and British Honduras. Univ. Michi-
gan Mus. Zool., Misc. Publ. no. 26, 30 pp.
Nelson, E. W.
1899. Revision of the squirrels of Mexico and Central America.
Proc. Washington Acad. Sci., 1:15-106.
Osgood, Wilfred H.
1909. Revision of the mice of the American genus Peromyscus. LT. S.
Dept. Agric, North Amer. Fauna no. 28, 285 pp.
Sanborn, Colin Campbell
1935. New mammals from Guatemala and Honduras. Field Mus. Nat.
Hist., Zool. Ser., 20(11) :81-85.
Thomas, Oldfield
1909. A new rat from Guatemala. Proc. Zool. Soc. London, 1909. pp.
669-670.
Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE
Vol. 116, No. 5
THE GENUS TETRAGNATHA (ARANEAE,
ARGIOPIDAE) IN PANAMA
By Arthur M. Chickering
Albion College, Albion, Michigan
CAMBRIDGE, MASS.. U. S. A.
PRINTED FOR THE MUSEUM
May, 1957
Publications Issued by or in Connection
with THE
MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE
Bulletin (octavo) 1863 — The current volume is Vol. 116.
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Memoirs (quarto) 1864-1938 — Publication was terminated with Vol. 55.
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Vol. 3, no. 35 is current.
Occasional Papers of the Department of Mollusks (octavo) 1945 —
Vol. 2, no. 21 is current.
Proceedings of the New England Zoological Club (octavo) 1899-
1948 — Published in connection with the Museum. Publication terminated
with Vol. 24.
The continuing publications are issued at irregular intervals in numbers
which may be purchased separately. Prices and lists may be obtained on
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Cambridge 38, Massachusetts.
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Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE
Vol. 116, No. 5
THE GENUS TETB AGNATE A (ARANEAE,
ARGIOPIDAE) IN PANAMA
By Arthur M. Chickering
Albion College, Albion, Michigan
CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM
May 1957
No. 5 — The Genus Tetragnatha (Araneae, Argiopidae) in
Panama
By Arthur M. Checkering
Albion College, Albion, Michigan
As a result of several visits to the Canal Zone and the Republic
of Panama for the purpose of collecting and studying spiders I
have accumulated a rather large number of specimens belonging
to the interesting genus Tetragnatha Latreille, 1804. The present
study of the genus is specifically concerned with its occurrence
in Panama where it appears to have found exceptionally favor-
able conditions.
Araneologists who have interested themselves in the genus
Tetragnatha have emphasized such characters as the following:
size of the body ; shape of the abdomen ; relative position of the
eyes ; several features of the chelicerae and cheliceral teeth ; color ;
relative lengths of the different segments of the male palp ; specific
characteristics of the male palpal tarsus; presence or absence of
spines on legs and their length. F. P. Cambridge (1903) paid
close attention to the characters and relationships of the con-
ductor and embolus in male palps. Petrunkevitch (1930) also
did this and, in addition, gave careful attention to the appear-
ance of the genital fold in females which are often difficult to
place with certainty because of the absence of an epigynum and
other marked characteristics. My experience with the genus
seems to show that close attention must ahvays be given to the
specific shape of the conductor and embolus as well as to their
relationships. These features appear to be the least variable
among all of those used by taxonomists and, therefore, the most
reliable for purposes of accurate determination. In females the
characters of the genital fold are often very helpful. Size ; color ;
number, relative size, and placement of the cheliceral teeth are
all subject to a rather wide range of variation. These facts seem
to explain the numerous errors in identification which can be
found in almost every collection. In making. this study I have
tried to take into consideration all of these items in making my
identifications and in drawing conclusions regarding synonymy.
Acknowledgments arc again <lue and p'ratefullv extended to the
302 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
following' persons for their interest and cooperation : Dr. A. S.
Romer, Director, and Dr. P. J. Darlington, Jr., Curator of In-
sects, respectively, in the Museum of Comparative Zoology at
Harvard College where much of the work of preparing this paper
was carried out; Dr. W. J. Gertsch, American Museum of Nat-
ural History, for the loan of specimens from Panama ; Dr. G.
Owen Evans, Department of Zoology, British Museum (Natural
History), for the loan of almost indispensable specimens studied
by the two Cambridges ; Dr. R. V. Chamberlin, University of
Utah, for the loan of types from Panama. It is also a pleasure
again to acknowledge my indebtedness to the donors of the Pen-
rose Fund of the American Philosophical Society, the Society of
Sigma Xi, and the Trustees of the Horace H. and Mary A.
Rackham Fund whose grants have made possible six extended
periods of collecting and study of spiders in various parts of
Panama during the past thirty years.
Genus TETRAGXATHA Latrcille, 1804
The Cambridges (1889-1903) reported the following species of
this genus in Panama: T. alba F. P. Cambridge; T. pallida 0. P.
Cambridge; T. tenuis 0. P. Cambridge; T. tenuissima 0. P.
Cambridge. Petrunkevitch (1925) added T. antillana Simon to
the known list from Panama, and Banks (1929) recorded T.
mexicana Keyserling. Chamberlin and Ivie (1936) described
four new species from my first collection of Panamanian spiders
made in 1928.
T. alba F. P. Cambridge has proven to be a synonym for T.
laboriosa Hentz. T. amplidens Chamberlin and Ivie and T. siduo
Chamberlin and Ivie are synonyms for T. tropica 0. P. Cam-
bridge. The male of T. apheles Chamberlin and Ivie belongs
with T. mexicana Keyserling but the female is a T. antillana
Simon. T. ethodon Chamberlin and Ivie appears to be a valid
species. To the best of my knowledge, therefore, the total list
of known species of Tetragnatha from Panama prior to this
study may be stated as follows: T. antillana Simon; T. ethodon
Chamberlin and Ivie; T. laboriosa Hentz; T. mexicana Keyserl-
ing; T. pallida 0. P. Cambridge; T. tenuis 0. P. Cambridge;
T. tenuissima 0. P. Cambridge; T. tropica 0. P. Cambridge.
< HICKERING: TETRAGNATHA IN PANAMA 303
As a result of my studies on this genus I have definitely recorded
the following additional known species to the list : T. caudata
Emerton; T. cognata 0. P. Cambridge; T. confraterna Banks;
T. guatemalensis O. P. Cambridge; T. pallescens F. P. Cam-
bridge; T. vermiformis Emerton. In addition to these I have
been compelled to recognize and describe T. fragilis sp. nov. ;
T. gertschi sp. nov. ; T. mdbelae sp. nov. ; T. sinuosa sp. nov.
Thus the total number of species of Tetragnatha now known to
occur in Panama amounts to eighteen.
No attempt is made in this paper to give complete bibliographic
references. Only those which are regarded as particularly per-
tinent are included. Those who desire more extensive bibliog-
raphies are referred to Iioewer (1942) .
Types will be deposited in the Museum of Comparative Zoology
at Harvard College.
Key to the Species of Tetragnatha in Panama
Males
1. ALE distinctly further from PLE than AME are from PME (caudata,
fragilis, mexicana, pallescens, vermiformis) 2
1. ALE not distinctly further from PLE than AME are from PME
(antillana, cognata, confraterna, ethodon, gertschi, guatemalensis,
laboriosa, mabelae, pallida, sinuosa, tenuis, tenuissima, tropica) . .6
2. With abdomen considerably extended posterior to spinnerets
T. caudata, p. 308
2. With abdomen not extended any appreciable distance posterior to
spinnerets 3
:i. With palpal tibia distinctly longer than palpal patella (mexicana,
pallescens) 4
3. With palpal tibia not much longer than palpal patella (fragilis,
vermiformis) 5
4. Promargin of fang groove with the "large tooth"1 well developed
T. pallescens, p. 336
i. Promargin of fang groove without any "large tooth"
T. mexicana, p. 333
5. A smaller species (about 5-7 mm.) ; paraeymbium distinctly bifid
distally; with no spines on legs T. fragilis, p. 317
i The term "large tooth" is being used in the sense in which F. P. Cambridge
employed it. In certain species there is an enlarged proraarginal tooth at about
thi- "apical third or fourth."
.'504: BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
5. A larger species (7-9 mm.); paracymbium not bifid distally; legs with
spines T. vermiformis, p. 349
6. With no spines on legs; only a fine coating of hair
T. t.cnuissima, p. 344
6. With spines on all legs as well as a supply of hair (antillana, cognata,
confratcrna, ethodon, gertsclri, guatemalensis, laboriosa, mdbelae,
pallida, sinuosa, tenuis, tropica) 7
7. With palpal tibia much longer than palpal patella (antillana, guate-
malensis, pallida, tenuis, tropica) 8
7. With palpal tibia not much longer than palpal patella (cognata, con-
fraterna, ethodon, gertschi, laboriosa, mabelae, sinuosa 12
8. Paracymbium bifid distally T. antillana, p. 30(i
8. Paracymbium not bifid distally (guatemalensis, pallida, tenuis,
tropica ) 9
9. Paracymbium extended distally into a vermiform termination
T. guatemalensis, p. 326
9. Paracymbium not distally vermiform (pallida, tenuis, tropica) ....10
10. The "large tooth" proper well developed (see note, p. 303) .
T. tropica, p. 347
10. The "large tooth" proper not present (pallida, tenuis) 11
11. The conductor, embolus, and cymbium all long, slender, nearly straight
(Fig. 78) T. pallida, p. 338
11. The conductor, embolus, and cymbium all at least somewhat sinuous
(See F. P. Cambridge's Figs. 1 and 1A, Plate 41) . . T. tenuis, p. 342
12. With abdomen extended a short distance posterior to spinnerets
T. confraterna, p. 312
12. With abdomen not extended posterior to spinnerets (cognata, ethodon,
gertschi, laboriosa, mabelae, sinuosa) 13
13. The promargin of fang groove with a very large, massive tooth at base
of fang (Fig. 85) T. sinuosa, p. 340
13. The promargin of fang groove without any very large, massive tooth
at base of fang (cognata, ethodon, gertsclti, laboriosa, mabelae). .14
14. With the paracymbium somewhat serrated distally (Fig. 64)
T. tnabc lac, p. 330
14. With paracymbium not serrated distally (cognata, ethodon, gertschi,
laboriosa) 15
15. Bulb of palpal tarsus distinctly inflated (Fig. 39)
T. gertschi, p. 321
15. Bulb of palpal tarsus not unusually inflated (cognata, ethodon, lab-
oriosa) 16
16. Paracymbium distinctly notched distally T. ethodon, p. 316
16. Paracymbium without distal notch (cognata, laboriosa) 17
CHICKERING: TETRAGNATHA IN PANAMA 305
17. Conductor with a cap-like distal termination (Fig. 56)
T. laboriosa, p. 329
17. Conductor without a cap like distal termination; wdth a bluntly rounded
termination ... T. cognata, p. 310
Females
Because of the lack of distinctive features in females it is difficult to
provide readily workable keys. It is hoped, however, that the following
key will aid the student of the group in separating the females of the dif-
ferent species.
1. ALE distinctly further from PLE than AME are from PME (caudata,
fragilis, mexicana, pallescens, vermiformis) 2
1. ALE not distinctly further from PLE than AME are from PME
(antillana, cognata, confraterna, gertschi, guatemalensis, laboriosa,
pallida, tenuis, tenuissima, tropica) 6
2. Abdomen considerably extended posterior to spinnerets
T. caudata, p. 308
2. Abdomen not noticeably extended posterior to spinnerets (fragilis,
mexicana, pallescens, vermiformis) 3
3. Spines completely lacking from all legs . . . T. fragilis, p. 317
3. Spines present on all legs (mexicana, pallescens, vermiformis) 4
4. Genital fold considerably longer than wide ....T. vermiformis, p. 349
4. Genital fold wider than long (mexicana, pallescens) 5
.">. Genital fold very gently procurved (Fig. 70) T. mexicana, p. 333
5. Genital fold strongly procurved, fully twice as wide as long (Fig. 75)
T. pallescens, p. 33(5
ii. Spines completely lacking from all legs T. tenuissima, p. 344
6. Spines present on all legs (antillana, cognata, confraterna, gertschi,
giiatemalensis, laboriosa, pallida, tenuis, tropica ) 7
7. Betromargin of fang groove with a much enlarged tooth at base of fang
directed forward (Fig. 6) T. antillana, p. 306
7. Retromargin of fang groove without a greatly enlarged tooth at base
of fang directed forward (cognata, confraterna, gertschi, guate-
malensis, laboriosa, pallida, tenuis, tropica) 8
8. Genital fold fully twice as long as wide (Fig. 26)
T. confraterna, p. 312
8. Genital fold much less than twice as long as wide (cognata, gertschi,
guatemalensis, laboriosa, pallida, tenuis, tropica) 9
9. Basal segment of chelicera nearly as long as carapace ; basal segment
of chelicera with an unusual dorsal blunt tooth near base of fang
(Fig. 100) . T. tropica, p. 347
306 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
9. Basal segment of chelicera much shorter than carapace; basal segment
of chelicera without any dorsal distal tooth near base of fang
(cognata, gertschi, guatemalensis, laboriosa, pallida, tenuis) ....10
10. ALE definitely closer to PLE than AME are to PME {cognata, gert-
schi, guatemalensis) 11
10. ALE not closer to PLE than AME are to PME (laboriosa, pallida,
tenuis) 13
11. A larger species, 9-11 mm.; fang with a dorsal basal cusp
T. guatemalensis, p. 326
11. Smaller species, from 5-8 mm.; fang without a dorsal basal cusp
(cognata, gertschi) 12
12. Abdomen deeply notched at base dorsally T. cognata, p. 310
12. Abdomen unnotched at base dorsally T. gertschi, p. 321
13. Abdomen conspicuously silvery on dorsal and dorsolateral sides, with a
dark median ventral stripe and a silvery stripe on each side of the
dark stripe T. laboriosa, p. 329
13. Abdomen not conspicuously silvery and without the ventral stripes as
in laboriosa (pallida, tenuis) 14
14. ALE about as far from PLE as AME are from PME; with long robust
leg spines T. pallida, p. 338
15. ALE slightly further from PLE than AME are from PME; with rela-
tively weak leg spines T. tenuis, p. 342
Tetragnatha antillana Simon, 1897
(Figures 1-6)
T. antillana Banks, 1901
T. antillana F. P. Cambridge, 1903
T. antillana Petrunkevitch, 1911
T. eremita Chamberlin, 1924
T. antillana Seeley, 1928
T. antillana Petrunkevitch, 1930
T. apheles Chamberlin and Ivie, 193G (female only)
T. festina Bryant, 1945 (male only)
T. haitiensis Bryant, 1945 (females)
T. antillana Kraus, 1955
T. antillana Simon is well known throughout Mexico, Central
America, most of South America, and the West Indies. The vial
in the collection of the Museum of Comparative Zoology labelled
the type of T. eremita Chamberlin now contains only a single
male palp but this is very definitely from T. antillana Simon.
The female about which Dr. Chamberlin had some doubts also
CHICKERING : TETRAGNATHA IN PANAMA
307
clearty belongs here. Moreover, the female of T. apheles Cham-
berlin and I vie is quite clearly a T. antillana Simon.
Male : Lateral eyes somewhat closer to one another than AME
are to PME ; legs well supplied with short spines ; palpal patella
about two-thirds as long as palpal tibia; the conductor and
-SwBgi
External Anatomy of Tetragnatlia
Figures 1-6, T. antillana
Fig. 1. Eye group from in front.
Fig. 2. Distal ends of conductor, embolus, and cymbium.
Fig. 3. Distal end of paracymbium.
Fig. 4. Chelicera and teeth of male.
Fig. 5. Genital fold and genital area, female.
Fig. 6. Chelicera and teeth of female.
308 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
embolus as shown in Figure 2; the paracymbiuni is distally
bifurcate (Fig. 3) ; the fang is moderately sinuous and is without
a basal dorsal cusp but there is a suppressed tubercle on the
inner margin opposite the third promarginal tooth ; there is no
"large tooth" on the promargin but the spur together with the
two contiguous teeth forms a conspicuous group of three (Fig. 4).
Female : The genital fold is as shown in Figure 5 ; the retro-
margin of the fang groove has a large distinctive tooth directed
forward with a small tooth near it (Fig. 6) ; the promargin has
a similar pair of distal but smaller teeth ; the fang is moderately
sinuous and there may be a suppressed dorsolateral cusp near
its base ; the abdomen is not extended posterior to the spinnerets
in either sex.
Collection records : The male and female hypotypes are from
Boquete, R. P., August, 1950 and July, 1939, respectively.
Numerous examples of both sexes from Boquete, R. P., July,
1939, August, 19,50; El Valle, R. P., July, 1936; El Volcan,
R. P., February- April, 1936 (W. J. Gertsch) ; one female from
Madden Dam Forest, C. Z., July, 1950.
Tetragnatha caudata Emerton, 1884
(Figures 7-11)
Eucta lacerta Petruukevitdi, 1911
T. caudata Seeley, 1928
T. caudata Bryant, 1940
T. lacerta Eoewer, 1942
T. caudata Kaston, 194S
For some time this species was considered as new and was
scheduled to be described as such. After careful comparison with
T. caudata from the northern regions the close similarities seemed
to warrant regarding it as a southern variant of this species.
Further knowledge regarding it may compel araneologists to
regard it as a separate species. T. caudata Emerton has been
recorded from Maine to Florida along the Atlantic coast, from
Canada, through several middle Western states, and in the south
as far west as Mississippi. It is interesting to find it now in
Panama and not where it would be expected to come in with
goods shipped from the north. The most distinctive features of
the species are given below.
CHICKERING : TETRAGNATHA IN PANAMA
309
Male: ALE considerably further from PLE than AME are
from PME (Fig. 7); palpal patella only a little shorter than
palpal tibia ; the conductor terminates in a broad distal piece
(differing considerably from that in northern forms) (Fig. 8) ;
the paraeymbium is bluntly rounded distally; the prolateral
spur is indistinctly bifid; the "large tooth" is present and the
other eheliceral teeth are as shown in Figure 9; the fang has no
cusps; the abdomen is considerably extended beyond the spin-
nerets; leg spines are few and weak.
vsr
* J>
a Q
J%> ffo
7
11
External Anatomy of Tetragnatha
Figures 7-11, T. caudata
Fig. 7. Eye group from in front.
Fig. 8. Apex of conductor and embolus of male.
Fig. 9. Male chelicera and eheliceral teeth.
Fig. 10. Female eheliceral teeth from below.
Fig. 11. Genital fold of female.
Female: Cheliceral teeth as shown in Figure 10; the genital
fold as shown in Figure 11 ; abdomen extended posterior to spin-
nerets for about one-fifth of the total length of the organ; male
hypotype fi.05 mm. long ; female hypotype 9.36 mm. long.
310 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Collection records : The hypotypes are from Boquete, Chiriqui,
R. P., August, 1950. Two other females are in the collection and
taken at the same place and time, together with a single female
from this locality taken in August, 1954.
Tetragnatiia cognata 0. P. Cambridge, 1889
(Figures 12-18)
T. cognata F. P. Cambridge, 1903
T. cognata Petrunkevitch, 1911
T. cognata Koewer, 1942
Apparently this species has not been studied since the original
work done by the Cambridges and, hence, it seems desirable to
give a condensed description of hypotypes as follows :
Male hypotype. Total length 4.160 mm. exclusive of chelicerae ;
including chelicerae 4.875 mm. Abdomen widest about two-
fifths from base ; does not extend posterior to spinnerets. Legs
comparatively robust ; with both spines and hairs. Ratio of eyes
AME : ALE : PME : PLE = 3.5 : 2.5 : 3.5 : 3.5. Lateral eyes
separated from one another by the diameter of ALE. AME sep-
arated from PME by nearly twice as far. Carapace longer than
basal segment of chelicerae in ratio of about 3 : 2. Chelicerae :
relative^ short and robust ; the fang has a cusp on its inner
margin near the base ; the prolateral spur is a simple and fairlj-
robust spine ; the promargin of the fang groove has six teeth of
which the second is considerably enlarged and could well be con-
sidered the "large tooth" in the usual sense but F. P. Cambridge
did not so regard it ; the retromargin has six teeth with the first
two close together (Fig. 12). Legs with both spines and hair.
Palp : the tibia is only slightly longer than the patella ; the
paracymbium is relatively short and broad except terminally
where it narrows to a blunt apex (Fig. 13) ; the conductor is
broad throughout its length (Fig. 14). The hypotype male is
colored essentially like the hypotype female but is lighter
throughout.
Hypotype female. Total length exclusive of the chelicerae
5.265 mm. ; inclusive of the chelicerae 5.525 mm. Abdomen very
»il>bous about the middle (Fig. 15) ; gibbosity lacking in some
individuals which may not be mature ; not continued posterior to
spinnerets. Chelicerae : short, robust ; fang without inner or
CHICKERING : TETRAGNATHA IN PANAMA
311
outer cusps; promargin of fang groove with seven teeth1; retro-
margin with six (Fig. 16). Some variation in respect to number
and placement of teeth has been noted in both sexes. When the
female is fully mature the genital fold appears as shown in
Figure 17. The eyes are essentially as thev appear in the male
(Fig. 18).
12
1 7:-:V-
►A*
>v/
1 6
o o O o
° o o °
18
External Anatomy of Tetragnatha
Figures 12-18, T. cognata
Fig. 12. Left male cheliceral teeth.
Fig. 13. Male paraeymbium.
Fig. 14. Male conductor, embolus, and distal end of cymbium.
Fig. 15. Lateral view of female abdomen.
Fig. 16. Left female cheliceral teeth. (See footnote)
Fig. 17. Genital fold of female.
Fig. 18. Eye group of female from in front.
1 In Figure 16 there should be an additional minute tooth a short distance
proximal to the sixth tooth on the promargin (upper side).
312 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Collection records : The hypotypes are from Boquete, Chiriqui,
K. P., August, 1954. Numerous examples of both sexes from
Boquete, R. P., July, 1939, August, 1950, 1954; El Volcan,
Chiriqui, R. P., August, 1950 ; Canal Zone Experiment Gardens,
C. Z., August, 1954; Summit, C. Z., August, 1950.
Tetragnatha conkraterna Banks, 1909
(Figures 19-26)
T. eonfraterna Petrunkevitch, 1911
T. eonfraterna, Eoewer, 1942
This species was described by its author from Costa Rica with
great brevity and accompanied by a very simple figure of the
male chelicera. The species has received no attention since 1909
and would seem to deserve a detailed description which is given
below in accord with my usual formula.
Male hypotype. Total length exclusive of the chelicerae 8.775
mm. ; inclusive of the chelicerae about 10.92 mm. Carapace 3.12
mm. long, 1.722 mm. wide opposite posterior border of second
coxae where it is widest ; cephalic part only slightly raised ; with
other features as usual in the genus.
Eyes. Ocular tubercle bearing AME rather prominent ; viewed
from above, both rows moderately recurved; viewed from in
front, anterior row slightly recurved, posterior row nearly
straight, both measured by centers ; central ocular quadrangle
wider behind than in front in ratio of 6 : 5, wider behind than
long in ratio of 6 : 5. Ratio of eyes AME : ALE : PME : PLE
= 6:3: 4.25 : 4. AME separated from one another by slightly
more than their diameter, from ALE by slightly more than 1.3
times their diameter. PME separated from one another by
slightly less than 2.5 times their diameter, from PLE by twice
their diameter. Laterals separated from one another by the
diameter of AME. AME separated from PME by slightly more
than the distance between the laterals (Fig. 19). Height of
clypeus equal to 1.5 times the diameter of AME.
Chelicerae. Strongly developed, quite porrect, and moderately
divergent; basal segment 2.73 mm. long and, therefore, some-
what shorter than carapace ; fang long, slender, only slightly
sinuous, without cusps; the prolateral spur is simple, without
< IIK'KLJRING : TETRAGNATHA IN PANAMA
313
t)
22
External Anatomy of Tetragnatha
Figures 19-26, T. confraterna
Fig. 19. Eye group of male from in front.
Fig. 20. Left chelicera and cheliceral teeth of male.
Fig. 21. Distal end of male conductor and embolus.
Fig. 22. Paracymbium of male.
Fig. 23. Left chelicera and cheliceral teeth of female.
Fig. 21. Fang of female.
Fig. 25. Lateral side of female abdomen.
Fig. 26. Genital fold of female.
314 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
bifurcation or a tooth; the promargiu of the fang groove bears
nine teeth with the first robust and the second small; the retro-
margin has twelve teeth ; the teeth are spaced essentially as shown
in Figure 20. There is no "large tooth" on the promargin of
the fang groove.
Maxillae. Longer than lip in ratio of 45 : 19. Otherwise essen-
tially as usual in the genus.
Lip. Nearly as long as wide at base. Sternal suture gently
procurved ; with sternal tubercles as usual at ends of sternal
suture.
Sternum. Longer than wide in ratio of about 4:3; posterior
coxae separated by slightly more than one-third of their width.
Otherwise essentially as usual in the genus.
Legs. 1243. Width of first patella at "knee" .330 mm., tibial
index of first leg 4. "Width of fourth patella at "knee" .308 mm.,
tibial index of fourth leg 6.
Femora Patellae Tibiae Metatarsi Tarsi Totals
(All measurements in millimeters)
1. 7.865 1.175 8.060 8.710 1.755 27.565
2.
5.590
.975
4.745
4.875
1.150
17.355
3.
2.600
.552
1.625
2.080
.645
7.502
4.
0.045
.715
4.485
4.810
.950
17.005
Palp
1.826
.660
.704
1.144
4.334
Legs are provided with both spines and hairs.
Palp. Tibia only slightly longer than patella; the paracym-
bium is notched distally with the chitinous knob somewhat closer
to the base than to the apex ; the conductor is rather slender and
terminates characteristically at its distal end (Figs. 21-22).
Abdomen. Definitely extended a short distance posterior to
spinnerets ; only slightly enlarged near base ; without any dorsal
basal notch ; only overlaps carapace slightly ; 6.305 mm. long.
Color in alcohol. Legs and chelicerae yellowish with some ir-
regular grayish markings at joints. Maxillae yellowish along
median borders, grayish elsewhere. Lip and sternum grayish.
Carapace with a broad dusky gray median stripe and grayish
bands radiating from the median pit; also with a broad dusky
stripe along the border. Abdomen : the dorsum is light colored
with many yellowish silvery spangles and dusky areas; lateral
CHICKERING : TETRAGNATHA IN PANAMA 315
sides with narrow, irregular, grayish lines ; venter light
yellowish.
Female hypotype. Total length exclusive of the chelicerae 13
mm. ; including the chelicerae 15.60 mm. Carapace 3.055 mm.
long, 2.21 mm. wide opposite second coxae where it is widest.
Eyes. Essentially as in male.
Chelicerae. Basal segment 2.925 mm. long and, therefore,
slightly shorter than carapace; fang robust, markedly sinuous
and with a robust dorsolateral cusp near base and a smaller inner
cusp about one-fourth from base ; promargin of fang groove with
ten teeth; retromargin with twelve teeth the first of which is
the largest (Figs. 23-24) ; all spaced essentially as shown in
figures.
Maxillae, Lip, and Sternum. All essentially as in male.
Legs. 1243. Width of first patella at ''knee" .525 mm., tibial
index of first leg 5. Width of fourth patella at "knee" .352 mm.,
tibial index of fourth leg 6.
Femora Patellae Tibiae Metatarsi Tarsi Totals
(All measurements in millimeters)
1.
8.645
1.430
8.840
9.750
1.950
30.615
2.
5.785
1.170
5.200
5.700
1.235
19.110
3.
3.055
.660
1.820
2.405
.780
8.720
4.
6.370
.910
5.005
5.070
.810
18.165
All legs with spines and hair as in male.
Abdomen. Definitely extends a short distance posterior to
spinnerets ; considerably swollen in anterior third and quite con-
cave dorsally (concavity varies considerably among available
specimens) (Fig. 25); slightly notched at base above; 10.075
mm. long; genital fold essentially as shown in Figure 26 but
lateral margins indistinct.
Color in alcohol. Essentially as in male but in general is con-
siderably lighter with dark abdominal markings practically
absent.
Type locality. Both hypotypes from Barro Colorado Island,
C. Z., June, 1939. Numerous examples of both sexes from Barro
Colorado Island, June, 1934; February, 1936 (W. J. Gertsch) ;
June, 1936; June and August, 1939; July, 1950.
316
BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Tetragnatiia ethodon Charaberlin and Ivie, 1936
(Figures 27-31)
T. ethodon Eoewer, 1942
As I have pointed out elsewhere, the females regarded by the
authors of this species as the allotype and paratypes actually
belong" to T. tenuissima 0. P. Cambridge. The absence of spines
on the legs, the cheliceral teeth, and the genital fold as well as
other less marked characteristics make this identification certain.
External Anatomy of Tetragnatha
Figures 27-31, T. ethodon
Fig. 27. Eye group of male from in front.
Fig. 28. Left male chelicera and cheliceral teeth from below.
Fig. 29-30. Two views of distal parts of male conductor, embolus, and
cymbium.
Fig. 31. Distal end of male paracymbium.
The holotype male appears to represent a valid species and is
so regarded in this paper. It seems strange, however, that only
one specimen has been taken in view of the repeated extensive
CHICKERING: TETRAGNATHA IN PANAMA 317
collecting practiced during the past twenty-eight years. The
male holotype is rather badly mutilated and, hence, it is impos-
sible to describe it as carefully as desired. The following items
may be useful additions to the very brief description given by
the authors of the species.
Eyes. Viewed from above, both rows moderately recurved;
viewed from in front, anterior row nearly straight and posterior
row gently procurved, both measured by centers. Central ocular
quadrangle slightly wider behind than in front, slightly wider
behind than long. Ratio of eyes AME : ALE : PME : PLE =
12 : 8 : 10 : 9. AME separated from one another by their
diameter, from ALE by about five-fourths of their diameter.
PME separated from one another by about 1.7 times their di-
ameter, from PLE by slightly less than this. Laterals separated
from one another by five-fourth's of the diameter of ALE. Lat-
erals, therefore, slightly closer to one another than AME are
to PME. Height of clypeus equal to about five-fourths of the
diameter of AME.
Chelicerae. The "large tooth" is present; the prolateral spur
is bifid ; the chelieeral teeth along the fang groove are as shown
in Figure 28.
Palp. Essentially as shown in figures provided by the authors
of the species; some details relating to conductor, embolus, and
paracymbium are shown in Figures 29-31.
Type locality. The holotype is from Barro Colorado Island,
C. Z., July-August, 1928 (Chickering). No other specimens have
come to light during several collecting periods since the finding
of the holotype.
Tetragnatha fragilis sp. now
(Figures 32-38)
Male holotype. Total length including chelicerae 5.85 mm.,
exclusive of the chelicerae 5.395 mm. Carapace 1.527 mm. long,
.780 mm. wide opposite second coxae where it is widest ; cephalic
part nearly parallel sided ; other features as usual in the genus.
Eyes. Eight in two rows as usual, all dark ; viewed from above,
both rows definitely recurved ; viewed from in front, both rows
also moderately recurved, measured by centers. Central ocular
quadrangle wider behind than in front in ratio of 13 : 11 ; wider
318 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
behind than long in nearly the same ratio. Ratio of eyes AME
: ALE : PME : PLE = 7.5 : 5 : 6.5 : 6. AME separated from
one another by about six-fifths of their diameter, from ALE
by four-thirds of their diameter. PME separated from one an-
other by slightly more than twice their diameter, from PLE by
a little more than 1.5 times their diameter. Laterals separated
from one another by a little more than twice the diameter of
PLE. AME separated from PME by slightly more than 1.5
times the diameter of PME and, therefore, closer together than
ALE are to PLE. Height of clypeus equal to about the diameter
of AME.
Chelicerae. Well developed; strongly divergent; basal seg-
ment .910 mm. long; with a well-developed prolateral spur hav-
ing a pair of blunt terminal tubercles ; fang only slightly sinuous
but conspicuously bowed near the middle ; the promargin of the
fang groove has the "large tooth" about one-third from the
distal end, a small tooth distal to this and a series of five teeth
proximal to it diminishing in size to very minute at the last
(this series appears to be very variable among the paratypes) ;
the retromargin has a relatively large hook-like tooth near the
base of the fang and a series of five additional more proximal
(only four of these on the right) (Figs. 32-33) ; paratypes fre-
quently show only three proximal to the hook-like tooth on the
retromargin.
Maxillae. Nearly parallel, somewhat broadened distally ; some-
what concave along lateral surface in distal quarter ; longer than
lip in ratio of 23 : 10 ; somewhat more than three times as long
as wide at narrowest level.
Lip. Longer than wide at base in ratio of about 9:8; sternal
suture gently procurved ; with the usual sternal tubercles at ends
of sternal suture.
Sternum. Generally scutif orm ; moderately convex; longer
than wide in ratio of about 37 : 25 ; moderately scalloped opposite
each coxa and extended between all coxae, the second and third
being relatively widely separated ; continued as a narrow sclerite
between fourth coxae which are separated by only one-eleventh
of their width.
Legs. 1243. Width of first patella at "knee" .198 mm., tibial
index of first leg 4. Width of fourth patella at "knee" .137 mm.,
tibial index of fourth leg 5.
(IlICKEBING: TETRAGNATHA IN PANAMA
319
Femora
1.
4.745
2,
3.120
o
o.
1.625
4.
3.575
Palp
.748
Patellae Tibiae Metatarsi Tarsi
(All measurements in millimeters)
.585 5.085 5.525 1.397
.390 2.730 2.925 .900
.242 .902 1.170 .520
.292 2.275 2.470 .748
.264
.590
Totals
17.337
10.065
4.459
9.360
1.827
True spines are completely lacking from all legs.
Palp. The tibia is slightly longer than patella but both are
short ; the paracymbium is distinctly notched or bifid at its apex ;
the embolus describes nearly a circle on the bulb and then ex-
tends nearly straight to terminate at a point slightly beyond
35
32
34
External Anatomy of Tetragnatfta
Figures 32-38, T. fragilis
Fig. 32. Left ehelieera and eheliceral teeth of male from below.
Fig. 33. Prolateral spur of male.
Fig. 34. Distal parte of male cymhium, embolus, and conductor.
Fig. 35. Paracymbium of male.
Fig. 36. Eye group of female from in front.
Fig. 37. Cheliceral teeth of female.
Fig. 38. Genital fold of female.
320 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
the tip of the conductor which has a very characteristic form
(Figs. 34-35) . Both conductor and embolus are closely associated
throughout.
Abdomen. Not continued posterior to spinnerets; only slightly
extended over carapace ; 3.802 mm. long ; without any anterior
dorsal notch at base ; very slender and of nearly uniform di-
ameter throughout ; a little more than seven times as long as
wide near base ; genital fold only a transverse ridge.
Color in alcohol. Legs yellowish with extensive fine dusky gray
dotting. Palps, chelicerae, and maxillae yellow with a variety of
shading. Lip and sternum yellowish with dusky dotting. Cara-
pace yellowish with a broad irregular dusky median stripe from
posterior border nearly to PLE. Abdomen : dorsum yellowish
white with a small median black spot just above the anal tubercle
and four pairs of similar small black dorsolateral spots in the
posterior three fourths (with some irregularities) ; there is a
narrow dark gray lateral stripe on each side; the venter is a
dusky gray.
Female allotype. Total length including chelicerae 7.67 mm.
Carapace 1.82 mm. long, 1.25 mm. wide opposite second coxae
where it is widest ; otherwise essentially as in male.
Eyes. Essentially as in male (Fig. 36).
Chelicerae. Moderately robust ; moderately divergent ; scarcely
porrect ; basal segment .845 mm. long ; fang without particularly
conspicuous features ; promargin of fang groove with a mod-
erately large tooth near distal end and, after a long space, a series
of four teeth diminishing in size toward the proximal end ;
retromargin with six teeth arranged and spaced essentially as
shown in Figure 37. As usual there are variations in number and
placement of cheliceral teeth among the paratypes; one para-
type exhibits the same number and placement on the promargin
but has seven teeth on the retromargin spaced somewhat differ-
ently also from those of the allotype.
Maxillae, Lip, and Sternum. Except for minor details, essen-
tially as in male.
Legs. 12-43. Width of first patella at "knee" .220 mm., tibial
index of first leg 4. Width of fourth patella at "knee" .154 mm.,
tibial index of fourth leg 5.
CHICKERING : TETRAGNATHA IN PANAMA 321
Femora
Patellae
(All measm
Tibiae
ements in
Metatarsi
millimeters)
Tarsi
Totals
1.
5.200
.660
5.265
5.265
1.300
17.690
2.
3.250
.520
2.795
2.860
.780
10.205
3.
1.820
.292
.910
1.170
.560
4.752
4.
3.510
.455
2.860
2.600
.780
10.205
Legs without true spines as in male.
Abdomen. Long and slender as in male ; 5.58 mm. long ; genital
fold as shown in Figure 38.
Color in alcohol. Essentially as in male except that dusky
areas are much less conspicuous ; the single median dorsal spot
just above anal tubercle as in male but the paired dorsolateral
spots are only faintly indicated ; the dorsal and dorsolateral
areas are covered by many yellowish silvery spangles.
Type locality. Male holotype and female allotype from Barro
Colorado Island, C. Z., August, 1936. Numerous paratypes of
both sexes from Barro Colorado Island : June-August, 1936.
July, 1950 ; July, 1954 ; Ft. Davis, C. Z., August, 1936 ; Balboa,
C. Z., August, 1936; Pedro Miguel, C. Z., July, 1950; Madden
Dam Forest, C. Z., August, 1939; C. Z. Forest Reserve, July-
August, 1939 and July, 1950; Gamboa, C. Z., July, 1954; Sum-
mit, C. Z., July-August, 1950; C. Z. Experiment Gardens, C. Z.,
July-August, 1954. Arraijan, R. P., August, 1936; Porto Bello,
R. P., August, 1936.
Tetragnatha gertschi, sp. nov.
(Figures 39-45)
Male holotype. Total length including chelicerae 5.395 mm.
Carapace 2.015 mm. long; 1.365 mm. wide opposite second coxae
where it is widest ; with the usual form of the genus ; .390 mm.
tall and, therefore, about .29 as tall as wide ; with cephalic region
somewhat raised ; with the usual shallow median pit with apex
directed forward.
Eyes. Eight in two rows, all dark; ocular tubercle bearing
LE quite prominent ; viewed from above, both rows moderately
recurved ; viewed from in front, anterior row gently recurved
and posterior row gently procurved, both measured by centers;
central ocular quadrangle wider behind than in front in ratio
of 4 : 3, wider behind than long in ratio of 6 : 5. Ratio of eyes
322
BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY
AME : ALE : PME : PLE = 4 : 3.25 : 4.75 : 4. AME separated
from one another by seven-fourths of their diameter, from ALE
by eleven-fourths of their diameter. PME separated from one
another by nearly twice their diameter, from PLE by nearly the
same distance. Laterals separated from one another by about
two-thirds of the diameter of ALE and, therefore, much closer
to one another than AME are to PME. Height of clypeus nearly
equal to the diameter of AME.
39
External Anatomy of Tetraynatha
Figures 39-40, T. gertschi
Fig. 39. Male palpal tibia and tarsus; showing dilated bulb and course of
embolus and conductor.
Fig. 40. Male palpal patella, tibia, and tarsus; showing features of cym-
bium and paracymbium.
Chelicerae. Well developed and moderately divergent in distal
halves; basal segment 1.20 mm. long; with a well developed pro-
lateral bluntly pointed spur on each; fang regularly curved,
CHICKERING : TETRAGNATHA IN PANAMA 323
not sinuous ; fang groove well marked ; promargin of fang groove
with seven teeth, retromargin with seven and with both sets
spaced essentially as shown in Figure 42. There is no so-called
"large tooth" on the promargin and the two most distal retro-
marginal teeth are close together. Some variation in number and
placement of teeth has been noted among the paratypes.
Maxillae. Nearly parallel in general but slightly divergent in
distal halves; with rounded distal truncatures; slightly concave
along lateral surfaces ; longer than lip in ratio of about 32 : 15 ;
about three times as long as wide at narrowest level.
Lip. Broader at base than long in ratio of about 4:3; sternal
suture gently procurved ; with pronounced sternal tubercles at
ends of suture.
Sternum. Generally scutiform; moderately convex; longer
than wide in ratio of about 5:4; moderately scalloped opposite
each coxa and produced between all coxae ; continued as a narrow
sclerite between fourth coxae which are separated by slightly
more than one-third of their width.
Legs. 1243. Width of first patella at ''knee" .264 mm., tibial
index of first leg 5. Width of fourth patella at "knee" .198 mm.,
tibial index of fourth leg 7.
Femora
Patellae
Tibiae
Metatarsi
Tarsi
Totals
(All measurements in
millimeters)
1.
4.615
.910
4.205
4.745
1.170
15.695
o
2.990
.715
2.470
2.665
.748
9.588
3.
1.365
.500
.910
1.040
.520
4.335
4.
2.795
.585 •
2.145
2.340
.585
8.450
Palp
.943
.325
.286
1.040
2.594
All legs with spines and hair. Trichobothria have been observed
on all femora.
Palp. The patella is slightly longer than the tibia which has
the usual distal ehitinized rim. The cymbium is long, slender,
constricted in the middle third, and much broader at base. The
paracymbium is also slender, club-shaped, with a long ehitinized
border together with a ehitinized knob essentially as shown in
Figures 30-40. Both conductor and embolus are somewhat spira-
loid, closely associated throughout their length, and with the
tips extended as a terminal hook (Fig. 41). The bulb is more
inflated than usual in the genus.
324
BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
Abdomen. Not continued posterior to spinnerets; extended
slightly over the carapace ; 3.445 mm. long ; slightly notched
dorsall}" at base; about 1.235 mm. wide at widest place and,
therefore, a little less than three times as long as wide ; genital
fold essentially as in female.
Color in alcohol. Legs and palps with varying shades of
yellowish. Chelicerae brownish. Carapace yellowish with ir-
regular
inter-communicating grayish lines; just in front of
44
A
k
v d
45 f
42
\
>0
43
External Anatomy of Tetragnatha
Figures 41-45, X. gertschi
Fig. 41. Distal end of male palpal tarsus.
Figs. 42-43. Male and female cheliceral teeth, respectively.
Fig. 44. Lateral view of female abdomen.
Fig. 45. Genital fold of female.
thoracic pit there is a small gray elongated spot. Sternum, lip,
and maxillae are brownish with fine gray dots. Abdomen :
dorsum yellowish with numerous small subchitinous irregular
silvery flecks ; lateral sides irregularly black with larger silvery
(MUCKERING: TETRAGNATHA IN PANAMA 325
spots; venter with a central grayish stripe and a yellowish stripe
on each side with numerous silvery flecks.
Female allotype. Total length including chelicerae 5.525 mm.
Carapace 1.625 mm. long; 1.235 mm. wide opposite second coxae
where it is widest ; otherwise essentially as in male.
Eyes. Essentially as in male.
Chelicerae. Moderately robust; moderately divergent; basal
segment .845 mm. long. Fang without conspicuous features.
Fang groove well marked ; with five promarginal teeth and seven
retromarginal teeth, spaced as shown in Figure 43.
Maxillae. Essentially parallel; broadened distally and less
rounded there than in males. Otherwise essentially as in male.
Lip. Broader at base than long in ratio of 19 : 13. Otherwise
essentially as in male.
Sternum. Longer than wide in ratio of 15 : 11. Otherwise
essentially as in male.
Legs. 1243. Width of first patella at "knee" .242 mm., tibial
index of first leg 6. Width of fourth patella at "knee" .187 mm.,
tibial index of fourth leg 9.
Femora
Patellae
Tibiae
-Metatarsi
Tarsi
Totals
(All measurements in
millimeters)
1.
3.770
.780
3.510
3.640
.910
12.610
.>
2.470
.650
1.820
2.080
.660
7.680
3.
1.100
.430
.650
.910
.380
3.470
4.
2.405
.43.5
1.625
1.885
.575
6.945
Spines and hairs on legs as in male.
Abdoint it. Conspicuously gibbous just in front of middle
(Fig. 44) ; 3.9 mm. long; genital fold as shown in Figure 45.
Color in alcohol. Essentially as in male except that the ab-
domen is more conspicuously provided with the silvery flecks
and lias the dark lateral irregular spots much reduced; there is
also a clear middorsal stripe with narrow lateral oblique lines
extending ventrally as in so many species of this genus.
Type locality. Male holotype and female allotype from El
Volcan, Chiriqui, 11. P.. August, 1950. Several paratypes of
both sexes from El Volcan, August. 1950 and Boquete, Chiriqui,
R. P., July, 1939 and 1954.
326 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Tetragnatha guatemalensis 0. P. Cambridge, 1889
(Figures 46-53)
T. guatemalensis F. P. Cambridge, 1903
T. guatemalensis Banks, 1909
T. guatemalensis Petiunkevitch, 1911
I. seneca Seeley, 1928
T. banlcsi Levi and Field, 1954
T. guatemalensis Roewer, 1942
T. guatemalensis Kraus, 1955
Male hypotype. Total length including chelieerae 13 mm.;
exclusive of the chelieerae 8.10 mm. Carapace 2.925 mm. long;
1.95 mm. wide opposite second coxae where it is widest.
Eyes. Viewed from above, both rows recurved, anterior row
rather strongly, posterior row moderately. Viewed from in front,
anterior row moderately recurved, posterior row slightly pro-
curved (Fig. 53), both measured by centers. Central ocular
quadrangle wider behind than in front in ratio of 23 : 18; wider
behind than long in ratio of 23 : 39. Ratio of eyes AME :
ALE : PME : PLE = 6 : 4.5 : 5 : 5. AME separated from
one another by their diameter, from ALE by about 2.33 times
their diameter. PME separated from one another by twelve-
fifths of their diameter, from PLE by 2.7 times their diameter.
Laterals separated from one another by about the diameter of
PLE. AME separated from PME by about 1.33 times the di-
ameter of AME, hence further from one another than ALE are
from PLE. Height of elypeus equal to 1.5 times the diameter
of AME.
Chelieerae. Very divergent; basal segment slightly shorter
than carapace. The following features appear to be particularly
worthy of mention : there is a basal dorsal cusp on the fang but
it appears to be variable in the degree to which it is developed in
different individuals ; the prolateral spur is robust and shows
two tubercles below the apex; along the promargin of the fang
groove there is a small tooth fairly close to the spur, a hook-like
tooth behind that, then the "large tooth", and this is followed
by a series of nine teeth with the last four being very minute
(this is not in agreement with statement by F. P. Cambridge) ;
the retromargin has three teeth near the apex, then a small tooth
(not present on the right side), and this is followed by seven
CHICKERING : TETRAGNATHA IN PANAMA
327
teeth (Fig. 46). The spur, the three most apical promarginal
teeth, and the three most apical retromarginal teeth appear to
be fairly consistent in their appearance in different individuals
but a considerable range of variation has been noted with respect
to the remaining teeth.
Palp. Tibia longer than patella in ratio of 3 : 2; conductor
geniculate at beginning of apical third, somewhat spoon-shape
at apex ; paracymbium terminates in a unique slender curved
process (Figs. 49-51).
Female hypotype. Total length including chelicerae 10.40 mm.
Abdomen considerably swollen in anterior half; genital fold as
shown in Figure 52.
External Anatomy of Tetragnatha
Figures 46-48, T. guatemalensis
Figs. 46-47. Male chelicera and cheliceral teeth, and female cheliceral teeth,
respectively.
Fig. 48. Male paracymbium.
Chelicerae. The fang has a dorsal basal cusp; the promargin
of the fang groove has a large apical tooth, a somewhat smaller
tooth considerably separated from the first, and then after a
considerable space there is a series of five teeth diminishing in
size toward the proximal end of the segment; the retromargin
has a small apical tooth, a large tooth close to it followed by a
series of eight teeth with the second and third of this series
328
BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
A considerable variation in respect to
their placement has been noted among
coalescent (Fig. 47).
number of teeth and
different individuals.
Color in alcohol. The color is difficult to describe adequately.
The carapace has a pair of broad dusky dorsal stripes reaching
from the median pit to the posterior eyes ; the lateral sides of
52
49
m
O O
53
50
External Anatomy of Tetragnatha
Figures 49-53, T. guatemalensis
Figs. 49-51. Three views of distal end of male palpal tarsus; Fig. 49 drawn
at a smaller scale.
Fig. 52. Genital fold of female.
Fig. 53. Eye group of male from in front.
the carapace are also dusky. The abdomen is covered b}" many
small irregular whitish spots ; laterally there are many narrow
black irregular stripes alternating with narrow yellowish stripes ;
the venter has the usual dark median stripe with lighter spangled
areas on each side.
Type locality. The male and female hypotypes are from Barro
CHICKERING : TETRAGNATIIA IN PANAMA 329
Colorado Island, C. Z., August, 1954. Numerous specimens of
both sexes from : Barro Colorado Island, June, July, 1934 ; July,
1936; June, 1939; June, 1950; August, 1954; Gamboa, C. Z.,
July, 1954.
Tetragnatiia laboriosa Hentz 1850
(Figures 54-59)
T. illinoisensis Keyserling, 1879
T. alba, F. P. Cambridge, 1903
T. alba Banks, 1909
T. alba Petrunkevitch, 1911
T. laboriosa Petrunkevitch, 1911
T. laboriosa Seeley, 1928
T. alba Eoewer, 1942
T. laboriosa Eoewer, 1942
T. laboriosa Kraus, 1955
Notes from Dr. W. J. Gertsch suggested the synonymy given
above. Comparisons of many specimens of both T. laboriosa and
T. alba from different localities have convinced me that this is
correct. Since the species has been described many times and is
one of the best known in the genus only those features believed
to be most distinctive will be emphasized here.
Distinctive features: Male. ALE and PLE about as far from
one another as AME are from PME ; palpal patella slightly
shorter than palpal tibia ; the conductor and embolus as shown in
Figures 55-56 ; the paracymbium is not divided distally ; the
prolateral spur is robust and bifid distally; the "large tooth"
is present on the promargin of the fang groove with other teeth
as shown in Figure 57 ; the abdomen is conspicuously silvery
with alternating dark and silvery stripes; of medium size, vary-
ing from 5 mm. to about 6.5 mm.
Female : size varies from about 6 mm. to about 8 mm. in
length ; the genital fold as shown in Figure 59 ; the cheliceral
teeth as shown in Figure 58 with considerable variation noted
among the many individuals examined; F. P. Cambridge noted
a small dorsal basal cusp on the fang but I have not found this.
Collection records: Numerous specimens of both sexes from
El Volcan, Chiriqui, R. P., February 28, 1936 (W. J. Gertsch),
and August, 1950; Boquete, Chiriqui, R. P., August, 1954;
330
BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
Chilibre, C. Z., July, 1950 ; apparently also from Barro Colorado
Island, C, Z., June, 1934 and 1936 ; August, 1939 ; July, 1954.
r
r
V
"r\
<3 Q? O 4D
54
58
55
57
59
External Anatomy of Tetragnatha
Figures 54-59, T. laboriosa
Fig. 54. Eye group of male from in front.
Fig. 55. Distal end of male tarsus.
Fig. 56. Distal end of male conductor, from a different view, to show dis-
tinctive apex.
Fig. 57. Male cheliceral teeth.
Fig. 58. Female cheliceral teeth.
Fig. 59. Genital fold of female.
Tetragnatha mabelae sp. no v.
(Figures 60-64)
Male holotype. Total length exclusive of the chelicerae 6.305
min. ; including the chelicerae 7.085 mm. Carapace 1.885 mm.
long, 1.105 mm. wide opposite second coxae where it is widest;
neither eye row occupies the full width of the carapace at its
level.
Eyes. Eight in two rows as usual ; viewed from above, anterior
CHICKERING : TETRAQNATHA IN PANAMA
331
row strongly and posterior row moderately recurved; viewed
from in front, anterior row gently recurved, posterior row
slightly recurved, all measured by centers. Central ocular quad-
rangle wider behind than in front in ratio of 28 : 25; wider
behind than long in ratio of about 14 : 11. Ratio of eyes AME
: ALE : PME : PLE = 4.5 : 2.5 : 3.75 : 3. AME separated
from one another by ten-ninths of their diameter, from ALE by
a little less than twice their diameter. PME separated from
one another by a little less than twice their diameter, from PLE
by nearly the same distance. Laterals separated from one an-
o o O °
° O O 6°o
63
62
64
External Anatomy of Tetragnatha
Figures 60-64, T. mabelae
Fig. 60. Eye group of male from in front.
Fig. 61. Male chelicera and eheliceral teeth.
Fig. 62. Male palpal patella, tibia, and tarsus.
Fig. 63. Distal ends of cymbium, conductor, and embolus of male palp.
Fig. 64. Male paracymbium.
other by twice the diameter of ALE. AME separated from PME
by a little more than their diameter (Fig. 60). Height of clypeus
equal to five-fourths of the diameter of AME.
332 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Chelicerae. Well developed; very divergent; basal segment
1.235 mm. long and, therefore, about two-thirds as long as cara-
pace ; with a moderately well developed prolateral spur terminat-
ing in a pair of minute lobules; fang moderately slender, slightly
sinuous and with a small cusp on inner margin about one-fourth
from base; promargin with the "large tooth" and five others
diminishing in size toward the base of the segment; retromargin
with six teeth as shown in Figure 61.
Maxillae. Essentially parallel; with normal concavities on
outer margins toward the tip ; longer than lip in ratio of 5 : 2 ;
about four times as long as wide at narrowest place.
Lip. Broader at base than long in ratio of 4 : 3 ; sternal suture
straight ; sternal tubercles at ends of sternal suture pronounced.
Sternum. Moderately convex; generally scutiform; longer
than wide in ratio of about 4:3; continued between fourth
coxae which are separated by about one-fourth of their width.
Legs. 1243. Width of first patella at "knee" .252 mm., tibial
index of first leg 4. Width of fourth patella at "knee" .187 mm.,
tibial index of fourth leg 5.
Femora Patellae Tibiae Metatarsi Tarsi Totals
(All measurements in milimeters)
1.
5.785
.945
5.687
5.840
1.430
19.687
2.
3.770
.780
3.120
3.250
.975
11.895
3.
2.080
.465
1.170
1.495
.650
5.860
4.
4.160
.650
3.120
3.025
.780
11.735
Palp
.975
.340
.375
.845
2.535
All legs with both hairs and spines.
Palp. The tibia is only slightly longer than the patella and
both are short ; the paracymbium is short, broad, and has a
somewhat serrate distal margin ; the embolus forms a normal
loop on the bulb and then extends with the rather broad con-
ductor to its termination close to the apex of the cymbium (Figs.
G2-64).
Abdomen. Not continued posterior to spinnerets; extended
only slightly over the carapace ; only slightly notched dorsally
at base ; nearly uniform in width throughout ; 4.355 mm. long ;
.780 mm. wide near base and, therefore, nearly six times as long
as wide.
Color in alcohol. Legs, mouth parts, and sternum all with
CHICKERING : TETBAGNATHA IN PANAMA 333
varying shades of yellowish. Carapace yellowish with a dusky
stripe extending from posterior border to the median pit and
then continuing forward as a pair of somewhat diverging stripes.
Abdomen : dorsally and dorsolateral^ covered by many yellow-
ish silvery spangles; ventrally there are fewer of the spangles
with the median area free of these and somewhat transparent.
Type locality. Male holotype from Barro Colorado Island,
C. Z., July, 1954. Two paratype males from Madden Dam For-
est, C. Z., July, 1950. Females unknown.
Tetbagnatha Mexicans Keyserling, 1865
(Figures 65-70)
T. longa O. P. Cambridge, 1889
T. mexicana F. P. Cambridge, 1903
T. apheles Cliamberlin and Ivie, 1936 (male only)
T. mexicana Eoewer, 1942.
F. P. Cambridge had Keyserling 's holotype female for study
and regarded it as an immature specimen. He was certain that
T. longa 0. P. Cambridge was the same species and he had both
sexes for study. The female reported by Banks (1929) is im-
mature and its identification is uncertain. I have had specimens
from the British Museum for comparison with mine and with
those described by Chaniberlin and Ivie as T. apheles. It is quite
dear that the female of T. apheles is a recently moulted T. an-
fillana Simon. The male appears to be a T. mexicana Keyserling
and is so regarded here. There may be a single female in my
collection which belongs to this species but I consider this un-
certain. In view of the confusion relating to this species the
decision has been made to give a rather complete description of
a male hypotype in accord with my usual procedure.
Male hypotype. Total length including chelicerae 13.455 mm. ;
exclusive of the chelicerae 11.505 mm. long. Carapace 2.925 mm.
long; 1.625 mm. wide opposite second coxae where it is widest;
generally slender; with lateral ocular tubercles well separated;
with cephalic region only slightly raised ; median pit irregularly
circular; with a rather marked transverse groove between AME
and PME.
Eyes. Viewed from above, both rows rather strongly recurved ;
viewed from in front, anterior row gently recurved, posterior
334
BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
row more strongly recurved, both measured by centers. Central
ocular quadrangle wider behind than in front in ratio of 20 :
17 ; wider behind than long in ratio of 10 : 9. Ratio of eyes AME
: ALE : PME : PLE = 5.5 : 3.75 : 4.25 : 4. AME separated
from one another by about five-fourths of their diameter, from
ALE by nearly 1.6 times their diameter. PME separated from
one another by about 2.8 times their diameter, from PLE by
nearly 2.25 times their diameter. Laterals separated from one
O
70
o o
O
66
° o o °
65
68
ft
69
Fig-. 65.
Fig. 66.
Fig. 67.
Fig. 68.
Fig. 69.
Fig. 70.
External Anatomy of Tetragnatha
Figures 65-70, T. mexicana
Eye group of male from in front.
Male chelieera and chelieeral teeth.
Distal end of male palpal tarsus.
Male paracymbium.
Female chelieeral teeth from a British Museum specimen.
Genital fold of female from a British Museum specimen.
another by 2.8 times the diameter of ALE. AME separated
from PME by nearly 1.5 times the diameter of the former and,
hence, closer to the latter than the laterals are to one another.
Height of clypeus equal to a little more than twice the diameter
CHICKERING : TETRAGNATHA IN PANAMA 335
of AME. The relative distances between the different types of
eyes as they are considered here appear to be somewhat different
from those observed in specimens from the British Museum
(Natural History). These differences, however, are not con-
sidered to be particularly significant.
Chelicerae. Divergent; moderately slender; basal segment
2.08 mm. long and, therefore, about two-thirds as long as cara-
pace ; prolateral spur a simple, slender hook, not bifid terminally ;
fang slender, somewhat sinuous; fang groove with seven pro-
marginal teeth and five retromarginal teeth spaced essentially
as shown in Figure 66. The two chelicerae do not agree in the
number of teeth present along the fang groove. In the hypotype
the teeth on the left are as shown in the figure while on the right
there are only six promarginal teeth but there are seven retro-
marginal teeth. This again emphasizes the unreliability of teeth
as a certain character for identification.
Maxillae. Essentially parallel but quite concave along outer
margin at the beginning of the last third ; considerably broadened
at distal border.
Lip. Chitinized lip wider at base than long in ratio of 6 : 5 ;
sternal suture gently procurved; with well-developed sternal
tubercles at ends of sternal suture.
Sternum. Generally scutiform; longer than wide in ratio of
7:4; scalloped opposite all coxae and continued between all of
these including the fourth which are separated by a little more
than one-fifth of their width ; moderately convex ; covered by
both relatively short and long bristles.
Legs. 1423. Width of first patella at "knee" .418 mm., tibial
index of first leg 4. Width of fourth patella at "knee" .330 mm.,
tibial index of fourth leg 5.
Femora
Patellae
Tibiae
Metatarsi
Tarsi
Totals
(All
measurements
in millimeters
0
1. 9.945
1.300
10.595
10.725
2.145
34.710
2. 6.500
1.050
5.850
6.045
1.235
20.680
3. 3.185
.585
1.820
2.600
.845
9.035
4. 7.540
.780
6.400
5.785
1.235
21.740
Palp 1.650
.440
.660
.924
3.674
All legs with both spines and hairs.
Palp. The tibia is
longer than
the patella
in
ratio
Of 3 : 2;
336 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
the tibia has the usual ehitiuous rim at distal end; the para-
cymbium is long, rather slender, bluntly rounded at its distal
end ; the conductor shows three thin spiraloid plates near its
middle and appears to be somewhat spiraloid at its distal end
(Figs. 67-68). One specimen shows the distal end of the con-
ductor almost exactly as drawn by F. P. Cambridge but the
hypotype appears somewhat different. These differences are re-
garded as well within the normal variation of the species.
Abdomen. Long, slender; slightly the widest near base and
gradually tapered to posterior end which is not extended beyond
spinnerets ; 8.385 mm. long ; .975 mm. wide near base and,
therefore, more than eight times as long as wide. Genital fold
(Fig. 70) nearly transverse.
Color in alcohol. Legs yellowish with small grayish spots;
femora one and two with an obscure prolateral grayish stripe.
Chelicerae yellowish white with fang and teeth darker. Maxillae
yellowish with grayish streaks. Lip yellowish with grayish mark-
ings on the strongly chitinized portions. Sternum with various
shades of dusty gray. Abdomen : very light colored in general ;
dorsum with many irregular silvery spangles and a double series
of small black dorsolateral spots; the venter has the usual median
darker stripe.
Two females appear to go with the males but they are probably
immature. The general characteristics of females have been
studied from specimens loaned by the British Museum (Natural
History). From these specimens Figures 69 and 70 have been
drawn.
Type locality. The male hypotype is from Barro Colorado
Island, C. Z., August, 1939. Three other males and probably two
immature females are in the collection from the same locality,
dune and August, 1939.
'»■
Tetragnatha pallescens F. P. Cambridge, 1903
(Figures 71-75)
T. pallida Banks, 1892
Eugnatha pallida Banks, 1898
T. pallescens F. P. Cambridge, 1903
T. bidens F. P. Cambridge, 1903
T. pallescens Seeley, 1928
CHICKERING : TETRAGXATHA IN PANAMA
337
T. pallescens Petrunkevitch, 1930
T. pallescens Bryant, 1940
T. pallesci tis Bryant, 1945
This species is well known and has been quite adequately de-
scribed by several authors in fairly recent years (Seeley, 1928;
Petrunkevitch, 1930; Kaston, 1948). For this reason only dis-
tinctive characteristics and a few figures will be given in this
paper. Petrunkevitch (1930) was the first, so far as I know, to
point out that T. bid ens F. P. Cambridge was the same as T.
pallesci ns.
&
ii
£
\ A* i
£
71
73
I.
Fig.
Fig.
Fig.
Fig. 74
Fig. 75
73.
72
External Anatomy of Tetragnatha
Figures 71-75, T. pallescens
Eye group of male from in front.
Male ehelieera and cheliceral teeth.
Distal end of male conductor to show distinctive apex.
Female cheliceral teeth.
Genital fold of female.
Distinctive features. Male: The lateral eyes are considerably
farther from one another than AME are from PME (Fig. 71) ;
the palpal patella is about three-fourths as long as the palpal
tibia; the conductor is curiously sickle-shaped distally (Fig.
73) ; the paracymbium is long, slender, and rounded at the apex;
the chelicerae are nearly as long as the carapace ; the prolateral
spur on the basal segment of the ehelieera is unequally bifid;
the "large tooth" is absent from the promargin of the fang
338 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
groove ; the cheliceral teeth are essentially as shown in Figure
72; in some specimens the abdomen appears to be very slightly
prolonged posterior to the spinnerets ; spines of moderate length
are on all legs.
Female. The slightly extended abdomen is somewhat more
evident in this sex than in males ; the abdomen is somewhat gib-
bous anteriorly and is notched dorsally at its base ; the cheliceral
teeth are essentially as shown in Figure 74; the genital fold is
essentially as shown in Figure 75.
Collection records. Both sexes have been collected in the fol-
lowing localities : Barro Colorado Island, C. Z., June- July, 1934 ;
February-March, 1936 (W. J. Gertsch) ; June, 1936; El Volcan,
Chiriqui, Pv. P., February, 1936 (W. J. Gertsch).
Tetragnatha pallida 0. P. Cambridge, 1889
(Figures 76-80)
T. pallida F. P. Cambridge, 1903
T. pallida Banks, 1929
T. pallida Eoewer, 1942
This species was described from Bugaba, Panama, and has
only been reported once since that time (Banks, 1929) so far as
I know.
Male hypotype. Total length exclusive of the chelicerae 6.83
mm. ; including the chelicerae length is 8.455 mm. Carapace 1.755
mm. long. The whole body is long and slender. The legs are long,
slender, and bear stout spines and hairs. Ratio of eyes AME :
ALE : PME : PLE = 10 : 4 : 5.5 : 5.5. AME separated from
one another by their diameter, from ALE by a little more than
1.5 times their diameter. PME separated from one another by
slightly more than twice their diameter, from PLE by a little
less than three times their diameter. Laterals separated from
one another by a little less than twice the diameter of PLE
(Fig. 76). AME separated from PME about as far as ALE is
separated from PLE. The central ocular quadrangle is nearly
square, only slightly longer than wide. Chelicerae : basal seg-
ment 1.722 mm. long and, therefore, almost as long as the cara-
pace ; the fang has no cusps ; the promargin of the fang groove
has six teeth and the retromargin seven, all spaced essentially
as shown in Figure 77; there is no true "large tooth" in the
CHICKERING : TETRAGNATHA IN PANAMA
339
usual sense ; the prolateral spur is simple without distal bifurca-
tion. Palp: the tibia is longer than patella in ratio of about
3:2; the bulb is relatively short and the cymbium, conductor,
and embolus are all long and slender (Fig. 78). The color in
both sexes is very pale throughout with many yellowish silvery
spangles on the abdomen. A few of the females in the collection
have bright red spots at the bases of the spines and a bright red
narrow dorsolateral stripe on each side of the abdomen.
O
o
o o
O O
77
78
Fig.
76.
Fig.
77.
Fig.
78.
Fig.
79.
Fig.
80.
External Anatomy of Tetragnathn
Figures 76-80, T. pallida
Eye group of male from in front.
Male chelicera and eheliceral teeth.
Distal ends of male cymbium, conductor, and embolus.
Female chelicera and eheliceral teeth.
Genital fold of female.
340 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Female hypotypc. Total length exclusive of the chelicerae
10.01 mm.; including chelicerae 30.985 mm. Abdomen not ex-
tended posterior to spinnerets. The genital fold is shown in
Figure 80. The chelicerae are relatively short and robust with
seven teeth on each margin of the fang groove as shown in Fig-
ure 79.
Type locality. Male and female hypotypes from Canal Zone
Forest Reserve, July, 1934. Other specimens of both sexes from
Barro Colorado Island, C. Z., June-July, 1934; June-July, 1936;
June-August, 1950; July- August, 1954.
Tetragnatha sinuosa sp. now
(Figures 81-86)
Male kolotype. Total length exclusive of chelicerae 4.095 mm. ;
including chelicerae 4.355 mm. Carapace 1.27 mm. long, .877
mm. wide opposite intervals between second and third coxae
where it is widest; anterior row of eyes occupy the full width
of carapace at their level.
Eyes. Eight in two rows as usual ; ocular tubercles only moder-
ately prominent ; viewed from above, both rows recurved, an-
terior row strongly so and posterior row moderately ; viewed
from in front, anterior row gently recurved, posterior row gently
procurved, all measured by centers ; central ocular quadrangle
wider behind than in front in ratio of 14 : 11, wider behind than
long in ratio of 14 : 12. Ratio of eyes AME : ALE : PME :
PLE = 4 : '2.75 : 4 : 3. AME separated from one another and
from ALE by nearly five-fourths of their diameter. PME sep-
arated from one another by a little more than five-fourths of
their diameter, from PLE by about the same distance. Laterals
separated from one another by the diameter of PLE. AME sep-
arated from PME by slightly more than their diameter, hence
(Fig. 81) somewhat farther apart than laterals are separated
from one another.
Cliclicerae. Short, robust, only slightly porrect and divergent;
the fang is strongly sinuous and has a low cusp on the inner sur-
face and a low tubercle at its base on the dorsal side; the pro-
lateral spur has moved into a nearly dorsal position; the pro-
margin of the fang groove has five teeth with the first a massive
growth not seen in any other species; the retromargin has seven
CHICKERING : TETRAGNATHA IN PANAMA
341
teeth (Figs. 85-86). There is no "large tooth" in the usual
sense.
Maxillae. Nearly parallel with much less concavity along lat-
eral borders than usual in the genus; longer than lip in ratio
of 9 : 4 ; not quite three times as long as wide in the middle.
Lip. Wider at base than long in ratio of about 3:2; sternal
suture gently proeurved ; sternal tubercles at ends of sternal
suture shorter and blunter than usual in the genus.
Sternum. Rather strongly convex; generally scutiform; longer
than wide in ratio of 7 : 5; less strongly scalloped opposite each
coxa than usual in the genus; continued between fourth coxae
which are separated by one-third of their width.
rf£y
6
OO
©
81
84
85
86
Fig.
81.
Fig.
82
Pig.
83.
Fig.
81
Fig.
85
Fig.
8(5
External Anatomy of Tetragnatka
Figures 81-86, T. sinuona
Eye group of male hole-type from in front.
Male palp ; tibia and tarsus.
Male palpal tarsus; a different view.
Paraeymbium of male.
Male ehelicera and cheliceral teeth from below.
Male ehelicera and cheliceral teeth; prc-lateral view (from a
paratype).
342 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Legs. 1423. Width of first patella at "knee" .176 mm., tibial
index of first leg 4. Width of fourth patella at "knee" .132 mm.,
tibial index of fourth leg 7.
Femora
Patellae
(All measm
Tibiae
•ements in
Metatarsi
millimeters)
Tarsi
Totals
1.
3.835
.585
3.770
3.640
1.040
12.870
2.
2.340
.520
1.950
2.080
.780
7.670
3.
1.210
.264
.704
.836
.445
3.459
4.
3.770
.330
1.650
1.950
.650
8.350
Palp
.694
.198
.210
.638
1.740
All legs with both spines and hair.
Palp. The tibia is slightly longer than the patella and is very
broad ; both tibia and patella are short ; both cymbium and
paracymbium appear to be somewhat distinctively shaped (Figs.
82-84) ; the embolus and conductor appear to be completely sep-
arated much of their lengths, the former with a conspicuous
loop at the distal end of the bulb and terminating in a long
lash-like apex (Figs. 82-83).
Abdomen. Not continued posterior to spinnerets; long, slender
and nearly uniform in width ; 3.90 mm. long, .715 mm. wide about
one-third from base ; not notched dorsally at base.
Color in alcohol. Generally a light yellowish throughout ; the
carapace is dusky gray in the cephalic portion, behind the median
thoracic pit and along the lateral margins. Abdomen : the usual
characteristic silvery spangles are few in number in the mid-
dorsal region but are concentrated more or less as a dorsolateral
stripe on each side ; there is also a row of five or six darker spots
above the dorsolateral silvery stripes on each side and a small
median blackish spot just above the anal tubercle ; the venter is
almost uniformly yellowish with few yellowish silvery spangles.
Type locality. Male holotype from Summit, C. Z., August,
1950. Four male paratypes from the same locality, August, 1950,
and a single male from the C. Z. Forest Reserve, July, 1954
complete the known list of specimens. The female is unknown.
Tetragnatita tenuis O. P. Cambridge, 1889
(Figures 87-89)
T. tenuis F. P. Cambridge, 1903
T. tenuis Roewer, 1942
-J
CHICKERING : TETRAGNATHA IN PANAMA 343
This seems to be a rare species in collections. The Cambridges
had it from Guatemala and Panama. The British Museum
(Natural History) was unable to loan me specimens and this
indicated, I suppose, that only the original types are in that
institution. There is one female in the M. C. Z. collection from
San Domingo (Dominican Republic) doubtfully assigned to this
species. As I have already pointed out, females are often difficult
to identify with certainty but I seem to have a few specimens
which must be assigned to this species.
Female hypotype. Total length including chelicerae 10.985
mm. ; exclusive of chelicerae 9.75 mm. Carapace 2.73 mm. long,
1.625 mm. wide opposite second coxae where it is widest. The
t
.89 (\ 87
'8 8'
External Anatomy of Tetragnatha
Figures 87-89, T. tenuis
Fig. 87. Eye group of female from in front.
Fig. 88. Cheliceral teeth of female.
Fig. 89. Genital fold of female.
whole body is long and slender with the abdomen only slightly
swollen at base. Legs long and slender with both hairs and spines.
Ratio of eyes AME : ALB : PME : PLE = 5 : 3.25 : 4.25 : 4.
AME separated from one another by 1.3 times their diameter,
from ALE by twice their diameter. PME separated from one
another by about 2.33 times their diameter, from PLE by about
2.1 times their diameter. Laterals separated from one another
by slightly less than twice the diameter of ALE. AME separated
from PME by 1.2 times their diameter, hence about as far apart
us ALE are from PLE (Fig. 87). Height of clypeus equal to
1.6 times the diameter of AME. The central ocular quadrangle
is wider behind than in front in ratio of 17 : 14, wider behind
(fOOf)
344 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
than long in ratio of 17 : 15. Chelicerae : robust, moderately
porreet, slightly divergent; basal segment 1.55 mm. long and,
therefore, only a little more than one-half as long as the carapace ;
fang without cusps; promargin of fang groove with seven teeth;
retromargin with eight on right and seven on left (Fig. 88) ;
teeth observed here not quite in agreement with statements made
by F. P. Cambridge. Color : legs and mouth parts, except lip,
with varying shades of yellowish ; lip reddish brown ; sternum
yellowish ; the carapace has a few dusky streaks behind the
thoracic pit and in front of it there are two faint diverging dusky
stripes passing toward the posterior eyes ; on the abdomen dor-
sally and laterally there are many light yellowish silvery spangles
surrounded by light semitransparent reticulations ; the spangles
diminish toward the venter and disappear medially leaving a
rather narrow yellowish stripe throughout. The genital fold is
shown in Figure 89.
Type locality. The female hypotype is from Barro Colorado
Island, C. Z., Sept. 1939. Several other females are in my col-
lection from the same locality, June, 1936 ; June and August,
1939.
Tetragnatha tenuissima 0. P. Cambridge, 1889
(Figures 90-96)
T. tenuissima F. P. Cambridge, 1903
T. tenuissima Petrunkevitch, 1925
T. tenuissima Banks, 1929
T. tenuissima Petrunkevitch, 1930
T. ethodon Chamber lin and Ivie, 1936 (Females only)
T. tenuissima Bryant, 1940
T. tenuissima Boewer, 1942
T. tenuissima Bryant, 1945
This species is now known to be widely distributed throughout
Mexico, Central America, northern part of South America, and
most of the Caribbean islands. Petrunkevitch (1930) described
the most important features of the females. The most essential
characters of both sexes will be summarized here. The species
occurs in my collections most frequently of all species known
from Panama.
Male hypotype. Total length including chelicerae 8.45 mm.;
excluding chelicerae 7.455 mm. Carapace 2.265 mm. long, 1.04
CHICKERIXG : TETRAG.YATIIA IN PANAMA
345
mm. wide opposite second coxae where it is widest. Very slender
throughout whole body. Head rather sharply set off from
thoracic part and with nearly parallel lateral borders. Viewed
from above, both rows of eyes recurved, first row strongly so ;
viewed from in front, anterior row moderately recurved, poste-
rior row slightly recurved. Ratio of eyes AME : ALE : PME :
PLE = 5 : 3 : 4.5 : 4. AME separated from one another by a
little less than their diameter, from ALE by 1.3 times their
96
External Anatomy of Tetragnaiha
Figures 91-93, 96, T. tenuissima
Chelicera and chelieeral teeth of male.
Two different views of distal end of male palpal tarsus.
Chelicera and chelieeral teeth of female.
Fig. 91
Figs. 92-93.
Fig. 96
diameter. PME separated from one another by a little less than
twice their diameter, from PLE by about the same distance.
Lateral eyes separated from one another by the diameter of
PLE. AME separated from PME by a little more than the
diameter of the former and, hence, farther from one another than
ALE are from PLE. The central ocular quadrangle is wider
346
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
behind than in front in ratio of about 8 : 7, slightly wider be-
hind than long. Chelieerae : very porrect and divergent ; gen-
erally long and slender ; basal segment 2.21 mm. long and,
therefore, almost as long as the carapace ; the fang is moderately
sinuous, long and slender and has no cusps; the prolateral spur
is long, curved, with a very small dorsal apical tubercle and a
robust ventral apical tubercle ; the promargin of the fang groove
has eleven teeth including the "large tooth" but the last is mi-
nute and easily overlooked ; the retromargin has eleven teeth the
first of which is also very small and easily overlooked (teeth as
seen in the hypotype differ from those shown by F. P. Cam-
bridge but these differences are well within the normal range of
iA'
Mi
%r$* JZ&F- '%■■•
&
90
95
94
External Anatomy of Tetragnatha
Figures 90, 94-95, T. tenulssima
Fig. 90. Eye group of male from in front.
Fig. 94. Male paracymbium.
Fig. 95. Genital fold of female.
variation). Palp: the tibia is a little shorter than the patella;
the paracymbium is short, bluntly pointed at its apex and has a
chitinized knob near the distal end ; the conductor and embolus
are finely attenuated at their tips (Figs. 92-94). One of the
most distinctive features in this species is the complete absence
of spines from the legs. The hair is coarse, however, and there
are many bristles.
Female hypotype. Total length including chelieerae 10.725
mm. ; exclusive of the chelieerae 8.775 mm. Abdomen not ex-
tended posterior to spinnerets. Also long and slender like the
male but somewhat gibbous in anterior fifth of the abdomen. The
CHICKEBING: TETBAGNATHA IN" PANAMA 347
genital fold appears essentially as shown in Figure 95. Cheli-
eerae : There is a very conspicuous basal dorsal cusp on the
fang; the fang has a deep indentation on the inner side near
the middle and is moderately sinuous; the promargin of the fang
groove has seven teeth and the retromargin has eleven spaced
essentially as shown in Figure 96 (a considerable variation in
the teeth in different individuals has been noted) ; the basal seg-
ment is not quite as long as the carapace.
Type locality. The male hypotype is from Barro Colorado
Island, C. Z., July, 1954 and the female is from the same lo-
cality, August, 1954. Many specimens of both sexes are in my
collection from : Barro Colorado Island, C. Z., June-July, 1934 ;
June-July, 1936 ; June-August, 1939 ; June-August, 1950 ; July-
August, 1954; El Valle, R. P., July, 1936; Madden Dam, C. Z.,
August, 1939 ; C. Z. Experiment Gardens, July, July- August,
1950 and 1954; Boquete, Chiriqui, R. P., August, 1950 and 1954.
Tetragnatiia tropica 0. P. Cambridge, 1889
(Figures 97-102)
T. tropica F. P. Cambridge, 1903
T. tropica Banks, 1909
T. tropica Petrunkevitch, 1911
T. siduo Chamberlin and Ivie, 1936
T. amplidens Chamberlin and Ivie, 1936
T. tropica Roewer, 1942
Male hypotype. Total length including chelicerae 10.92 mm. ;
exclusive of the chelicerae 8.775 mm. Carapace 3.055 mm. long,
1.755 mm. wide opposite second coxae where it is widest. First
row of eyes project slightly beyond the sides of the carapace;
the second row occupies nearly the full width of the carapace
at their level. Viewed from above, both rows of eyes recurved,
posterior row slightly, anterior row moderately. Viewed from
in front, anterior row moderately recurved, posterior row mod-
erately procurved, both measured by centers (Fig. 97). Ratio
of eyes AME : ALE : PME : PLE = 13 : 9 : 11 : 10. AME
separated from one another by about their diameter, from ALE
by about 1.5 times their diameter. PME separated from one
another by nearly twice their diameter, from PLE by twice their
diameter. Laterals separated from one another by slightly more
348
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
than the diameter of PLB. AME separated from PMB by about
1.3 times their diameter, hence they are farther from one another
than the laterals are from each other. The central ocular quad-
rangle is wider behind than in front in ratio of about 21 : 19,
almost exactly as long as wide behind. Chelicerae : moderately
porrect ; strongly divergent ; generally long and moderately
slender ; basal segment 2.86 mm. long and, therefore, only slightly
o
o o O
o
O O
o
97
External Anatomy of Tetragnaiha
Figures 97-98, 100, T. tropica
Fig. 97. Eye group of male from in front.
Fig. 98. Cheliceral teeth of male from below.
Fig. 100. Cheliceral teeth of female from below.
shorter than the carapace ; the fang is slightly sinuous and has
a low tubercle on the inner surface about one third of its length
from the base; the prolateral spur is moderately robust and
distinctly bifid apically; the promargin of the fang groove has
the "large tooth" with two small teeth distal to it and a series
of ten small teeth proximal to it ; the retromargin has a series of
fifteen teeth shaped and spaced essentially as shown in Figure
98. The last five or six teeth along the fang groove are irregular
MUCKERING: TETRAGNATIIA IN PANAMA 349
and, apparently, quite variable among the numerous specimens
available for study.
Palp. The tibia is almost twice as long as the patella; the
paracymbium is bluntly pointed distally ; the distal end of the
conductor is subaculeate and slightly hooked (Fig. 99).
The legs have both spines and hairs some of which are quite
erect.
Female hypotype. Total length exclusive of the chelicerae
10.53 mm. ; inclusive of the chelicerae 13.13 mm. Abdomen not
extended posterior to the spinnerets; conspicuously gibbous in
anterior third and concave along dorsal surface. The genital
fold is essentially as shown in Figure 102. Chelicerae : there is
a dorsal basal cusp on the fang; there is also an unusual dorsal
distal tooth on the basal segment of the chelicera at the base of
the fang; the promargin of the fang groove has a small and a
much larger tooth at the distal end and, after a considerable
space, a series of seven teeth ; the retromargin has a total of
thirteen teeth spaced essentially as shown in Figures 100-101.
The exact number of cheliceral teeth seems to be quite variable
among the numerous specimens available for study. I regard
Chamberlin and Ivie's T. amplidens as one of these with a
larger number of teeth than heretofore recognized in this species.
Basal segment of the chelicerae not quite as long as the carapace.
Type locality. Both hypotypes are from Barro Colorado Is-
land, C. Z., June, 1939. Numerous specimens of both sexes are in
my collection from the following localities : Barro Colorado
Island, C. Z., June-July, 1934; June, 1936; June-August, 1939;
June-August, 1950; July, 1954; Madden Dam Forest, C. Z.,
July, 1950.
Tetragnatha vermiformis Emerton, 1884
(Figures 103-108)
Eucta vermiformis Petrunkevitch, 1911
T. vermiformis Seeley, 1928
T. vermiformis Eoewer, 1942.
The specimens now definitely assigned to this species have for
some time been considered to represent a new and undescribed
species. I have now carefully examined specimens from several
parts of the United States and, together with the types in the
350
BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
Museum of Comparative Zoology, compared them with my speci-
mens from Panama. This study has convinced me that I have
been dealing with the same species. There are what I consider
minor differences in the cheliceral dentition, possibly the eyes
and other features but I think these are all well within the normal
range of variations of a widely dispersed species. It seems in-
teresting to find this species so far south in Central America
when it has previously been reported only from east of the Rocky
Mountains in the United States and not yet from the West
Indies.
101
External Anatomy of Tetragnatha
Figures 99, 101-102, T. tropica
Pig. 99. Distal ends of eymbium, embolus, and conductor of male.
Fig. 101. Cheliceral teeth of a second female from below.
Fig. 102. Genital fold of female.
Male hy polype. Total length including the chelicerae 9.205
mm. ; exclusive of the chelicerae 7.67 mm. Carapace 2.86 mm.
long, 1.755 mm. wide ; with anterior row of eyes occupying the
full width of the carapace; with posterior row considerably
shorter. Logs with both spines and hairs. Ocular tubercle bear-
ing ALE quite prominent. Viewed from above, anterior row
of eyes gently recurved, posterior rowT strongly recurved; viewed
from in front, anterior row definitely procurved, posterior row
CHECKERING: TETRAGNATHA IX PANAMA 35]
straight or slightly recurved. Ratio of eyes AME : ALE : PME
: PLE = 5.5 : 3 : 3.5 : 3.5. AME separated from one another
by about their diameter, from ALE by slightly more than twice
their diameter. PME separated from one another by nearly three
times their diameter, from PLE by a little more than twice
their diameter. Laterals separated from one another by nearly
five times the diameter of ALE. AME separated from PME by
nearly twice the diameter of PME, hence much closer to one
another than ALE are to PLE. The central ocular quadrangle
is wider behind than in front in ratio of 17 : 15, wider behind
than long in nearly the same ratio. Height of clypeus equal to
twice the diameter of AME. Chelicerae : basal segment 2.47
mm. long and, therefore, about six-sevenths as long as the cara-
pace ; the fang has an inner cusp and a series of fine serrations
nearer the base (Figs. 103-104) ; the promargin of the fang
groove has ten teeth the third of which would probably be
called the "large tooth" by F. P. Cambridge ; the prolateral spur
is slender and not apically bifid ; the retromargin of the fang
groove has eight teeth. Palp : the tibia, including the chitinous
extension, is only slightly longer than the patella; the embolus
extends in a very loose spiral through the somewhat twisted
conductor to terminate in a characteristic apical form (Fig. 105) ;
the paracymbium is broad at the base where it is strongly chitin-
ized but it is very slender in the distal half and not notched at
its apex. Color : legs, cephalothorax, and mouth parts are all
yellowish ; the abdomen is covered by many yellowish silvery
spangles and grayish reticulations ; the cardiac area has a trans-
parent stripe ; the venter is also covered by the yellowish silvery
spangles and is without a median stripe which is so frequently
present in the genus.
Female Ixxjpotxjpe. Total length including the chelicerae 12.675
mm. ; exclusive of the chelicerae 10.725 mm. Carapace 2.925 mm.
long. Eyes essentially as in male (Fig. 106). Chelicerae: basal
segment 1.95 mm. long, about two-thirds as long as carapace;
the fang has no cusps; the promargin of the fang groove has
seven teeth, the first of which is relatively large ; the retromargin
also has seven teeth, the first of which is small (Fig. 107) ; the
cheliceral teeth are not in close agreement with description given
by Seeley (1928). The genital fold is essentially as shown in
352
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Figure 108. Abdomen : somewhat swollen at base but uniformly
tapered to a blunt point at posterior end ; about 4.5 times as
long as wide near base; slightly extended posterior to spinnerets.
Type locality. The male hypotype and one other male are from
Barro Colorado Island, C. Z., June, 1936 ; the female hypotype
is from the same locality, August, 1939. Four females have been
assigned to this species from El Volcan, Chiriqui, R. P., Febru-
ary, 1936 (Gertsch).
O
o *
O O
104
106
/)
108
Fig.
103.
Fig.
104.
Fig.
105.
Fig.
106.
Fig.
107.
Fig.
108.
107' 105
External Anatomy of Tetragnatha
Figures 103-108, T. vermiformis
Male chelicera and cheliceral teeth.
Lateral view of fang from male.
Distal ends of cymbium, embolus, and conductor in male.
Eye group of female from in front.
Female cheliceral teeth.
Genital fold in female.
CHICKERING : TETRAGNATHA IN PANAMA 353
BIBLIOGRAPHY
Banks, Nathan
1909. Arachnida from Costa Rica. Proc. Acad. Nat. Sci. Philadelphia,
61: 194-234, pis. 5, 6.
1929. Spiders from Panama. Bull. Mus. Comp. Zool. at Harvard Col-
lege, 69: 53-96, 4 pis.
Bryant, Elizabeth B.
1940. Cuban spiders in the Museum of Comparative Zoology. Ibid., 86
(7): 249-532, 22 pis.
1945. The Argiopidae of Hispaniola. Ibid., 95 (4) : 359-418, 4 pis.
Cambridge, O. P. and F. P. Cambridge
1889- Arachnida-Araneida. "Vols. III. In: Biologia Centrali-Ameri-
1905. cana. Dulau & Co., London.
Chamberlin, R. V. and Wilton Ivie
1936. New spiders from Mexico and Panama. Bull. Univ. Utah, 27
(5), Biol. Ser. 3 (5) : 1-103, 17 pis.
Chickering, A. M.
1957. The Genus Tetraynatha (Araneae, Argiopidae) in Jamaica,
B. W. I. and other neighboring islands. Breviora, Mus. Comp.
Zool., 68: 1-15.
Emerton, J. H.
1884. New England spiders of the family Epeiridae. Trans. Conn.
Acad., 6: 295-341.
Kaston, B. J.
1948. Spiders of Connecticut. Bull. Connecticut State Geol. Nat. Hist,
Survey. 70: 1-874, 144 pis.
Kraus, Otto
1955. Spinnen aus El Salvador (Arachnoidea, Araneae). Abh. Sen-
ckenberg. Naturf. Gesell., 493: 1-112, 12 pis.
Petrunkevitch, Alexander
1911. A synonymic index-catalogue of spiders of North, Central, and
South America, etc., Bull. Amer. Mus. Nat. Hist., 29: 1-809.
354 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGV
1 i> 2 5 . Araehnida from Panama. Trans. Connecticut Acad. Arts and
Sei., 27: 51-248, 157 figs.
1930. The spiders of Porto Eico. Pt. 2. Ibid., 30: 159-355, 240 figs.
ROEWKR, C. FR.
1942. Katalog der Araneae. Vol. 1, 1040 pp., Bremen.
SKF.LEY, B. M.
1928. Eevision of the spider genus Telragnatha. Bull. New York State
Mus., 278: 99-150.
Simon, Eugene
1892- Histoire Naturelle des Araignees. Deuxieme Edition. 2 Vols.
1903. Librairie Encyclopedique de Eoret, Paris.
Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE
Vol. 116, No 6
THE TENUIS AND SELENOPHORA GROUPS OF THE
ANT GENUS PON ERA (HYMENOPTERA : FORMICIDAE)
By Edward 0. Wilson
Biologiml Laboratories, Harvard University
CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM
May, 1957
Publications Issued by or in Connection
with THE
MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE
Bulletin (octavo) 1863 - - The current volume is Vol. ll(i.
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Memoirs (quarto) 1864-1938 — Publication was terminated with Vol. 55
Johnsonia (quarto) 1941 — A publication of the Department of Mollusks.
Vol. 3, no. 35 is current.
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Vol. 2, no. 21 is current.
Proceedings of the New England Zoological Club (octavo) 1899-
1948 — Published in connection with the Museum. Publication terminated
with Vol. 24.
The continuing publications are issued at irregular intervals in numbers
which may be purchased separately. Prices and lists may be obtained on
application to the Director of the Museum of Comparative Zoology,
Cambridge 38, Massachusetts.
Of the Peters "Check List of Birds of the World," volumes 1-3 are out
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Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE
Vol. 116, No 6
THE TENUIS AND SELENOPHORA GROUPS OF THE
ANT GENUS PON ERA ( IIYMENOPTERA : FORMICTDAE)
By Edward ( ). Wilson
Biological Laboratories, Harvard University
CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM
May 1957
No. 6 — The Tenuis and Selenophora Groups of the Ant Genu*
Ponera (Rymenoptera: Formicidae)
By Edward 0. Wilson
When W. M. Wheeler created the new genus Pseudocryptopone
(generitype: Cryptopone tenuis Emery) in 1933, it was with the
free acknowledgment that this entity could not be clearly sep-
arated from Ponera. "Indeed, I confess my inability to draw a
sharp line of demarcation between the two genera. One of the
species, incerta, new species, which I have assigned to Pseudo-
cry ptopone, might, with equal propriety, be placed in Ponera."
The purpose of this exceptional procedure was to begin a pre-
liminary, orderly reduction of Ponera. ll Ponera is now a large
and very difficult genus in great need of careful revision. The
monographer who undertakes this task will very probably divide
it into several subgenera or even genera and his definition of
these will automatically determine their relations to Pseudo-
cryptopone and therefore its true status and affinities." In a
similar fashion, Wheeler withdrew two Papuan species of Ponera
{selenophora and clavicornis) and combined them with a new
Philippine species (oreas) to form a second new genus, Seleno-
pone.
Although Wheeler's aim to partition Ponera and thereby sim-
plify its classification was an admirable one, the formal naming
of new genera on such feeble evidence as he proposed was not well
justified. A more recent examination of Pseudocryptopone and
Selenopone, along with many of the related Indo-Australian
species of Ponera, has convinced the present writer that
Wheeler's genera cannot be maintained on the basis of even the
most liberal criteria. To begin, Pseudocryptopone is linked
to Ponera by the intermediate species Ponera mocsaryi Emery,
which shows a combination of Ponera and Pseudocryptopone
characters. These two genera are further linked by several species
more closely allied to the Pseudocryptopone generitype, but
which tend strongly toward the more typical Ponera type, e.g.,
Ponera incerta (Wheeler), P. ratardorum Wilson, and P. huonica
Wilson. Similarly, Selenopone is linked to Ponera by the inter-
mediate Ponera syscena Wilson, and is closely approached within
356 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
the ranks of the "typical" Ponera by the species P. papuana
Emery. Finally, Pscudocryplopone and Selenopone are linked
to one another by several more or less intermediate species,
including Ponera clavicornis Emery, P. tenuis (Emery), and
P. huonica Wilson.
In short, there does not appear at present to be any basis for
a generic split along the lines proposed by Wheeler. The
following synonymy is accordingly proposed:
Genus PONERA Latreille
Ponera Latreille, 1804, Nouv. Diet. Hist. Nat., 24: 178-179. Generitype:
Formica eoarctata Latreille {—Formica contracta Latreille), by subse-
quent selection.
Pseudocryptopone Wheeler, 1933, Amer. Mus. Nov., no. 672: 12-13. Generi-
type: Cryptopone tenuis Emery, original designation. NEW SYN-
ONYMY.
Selenopone Wheeler, 1933, ibid., p. 19. Generitype: Ponera selenophora
Emery, original designation. NEW SYNONYMY.
Pseudocryptopone and Selenopone are of course available as
subgeneric names if any reason is found to make formal sub-
generic divisions in future revisions of Ponera. In the present
study, however, the entities considered are the two species
groups having affinities with P. tenuis and P. selenophora re-
spectively. Both groups are herein much enlarged by the addition
of a total of ten new species, most of which were collected by
the author during a recent research tour in Melanesia. To the
selenophora group have been added two older species (scabra,
sinensis) which were apparently overlooked by Wheeler in his
1933 revision.
Together the tenuis and selenophora groups comprise a large
and important section (approximately 50 per cent) of the
Papuan species of Ponera, but the present evidence indicates that
they diminish rapidly outside this area. In the tenuis group, a
single species is known from Java and one each from the Caroline
Islands, New Caledonia, and southeastern Australia. In the
selenophora group, one species each is known from the Philip-
pines, Hongkong, and southern Japan. It is possible that other
described species from outside the Papuan region may be placed
in these two groups when Ponera is more exhaustively studied.
WILSON : TENUIS AND SELENOPHORA GROUPS OF PONERA 357
At the moment the greatest concentration of species for both
groups appears to exist at intermediate elevations (500-1600
meters) in the mountains of New Guinea. No less than five
species, comprising 25 per cent of the total known, have been
collected in a limited area around the headwaters of the Mongi
River, Huon Peninsula. Further collecting in similar areas in
other parts of New Guinea will probably yield a large proportion
of the still undiscovered species.
The present contribution has been prepared as a preliminary
part of a review of the ants of Melanesia. The remainder of the
species of Ponera will be treated in a later part. Most of the type
and other material used in this study is deposited in the Museum
of Comparative Zoology at Harvard University. Other source
collections have been the Emery Collection in Genoa; Dr. E. S.
Ross' collection of New Guinea ants, deposited with the Cali-
fornia Academy of Sciences; Dr. J. L. Gressitt's collection of
Melanesian ants, deposited in the B. P. Bishop Museum, Hono-
lulu ; and Miss L. E. Cheesman's collection of New Hebridean
ants, deposited in the British Museum of Natural History.
Measurements
In the taxonomy of a genus such as Ponera, where species dif-
ferences are for the most part minute and subtle, exact measure-
ments are necessary for accurate species diagnoses. In the pres-
ent study an ocular micrometer was used, and estimations were
made to the nearest two-tenths of a unit of 0.0293 mm, or to
0.006 mm. Thus the calculated maximum error is ± 0.006 mm,
but in practice, of course, the actual maximum error varies
around this figure according to the specific measurement involved.
Head width, as defined below, is probably the "safest" measure-
ment ; repetitive measurements have shown that the actual maxi-
mum error is no more than ± 0.006 mm. Scape length and
petiolar node length are the least reliable measurements, but
even here the actual maximum error probably does not exceed
four-tenths of a micrometer unit either way, or ± 0.012 mm.
Head width (HW). Worker and queen: the maximum width
of the head held in perfect full face and excluding the eyes.
If the eyes extend beyond the lateral borders of the head in this
position, the measurement is taken across whatever parts of the
358 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
lateral borders are left exposed. Male: the maximum width of
the head across and including the eyes.
Head length (HL). The length of the head, held in perfect
full face, measured from the level of the anterionnost point of
the anterior clypeal border to the midpoint of the occipital
border.
Scape length (8L). The maximum length of the scape exclu-
sive of the basal "neck."
Cephalic index (CI). Head width X 100/head length.
Scape index (SI). Scape length X 100/head width.
Pronotal width (PW). The maximum width of the pronotum
measured from directly above and at a right angle to the long
axis of the alitrunk.
Petiole height. The height of the entire petiole, measured
from the level of the crest of the petiolar node to the level of the
lowermost point of the subpetiolar process.
Petiolar node length. When the petiole is held in exact side
view, the distance from the midpoint of the curve where the
anterior face of the node meets the anterior peduncle to the
midpoint of the curve where the posterior face of the node
meets the posterior peduncle.
Dorsal petiole width. The width of the petiolar node measured
from directly a,bove the node and at right angles to the long
axis of the body.
Characterization of the Ponera tenuis group
Worker. Small species, worker head width never exceeding
0.43 mm, head subrectangular, elongate, cephalic index not ex-
ceeding 86 ; antennal scapes short, the scape index never greater
than 90 ; antennal club massive, 4- or 5-jointed. Mandibles with
three well formed teeth occupying the apical two-fifths to one-
half of the masticatory border, the remainder of the border oc-
cupied by two smaller teeth plus a number of minute intercalary
denticles (P. huonica Wilson only) or by denticles only (other
species). Eyes minute, consisting of a single ommatidium, or
altogether absent ; when present, located 0.7 to 0.8 the distance
from the occipital border to the midpoint of the anterior clypeal
border. Junction of lateral and posterior faces of the propodeum
rounded, not marginate. Petiolar node relatively thick, seen
WILSON: TENUIS AND SELENOPHORA GROUPS OF PONERA 359
from the side, subrectangular, usually tapering very slightly
dorsally ; seen from directly above, its anterior face is more
or less semicircular or arcuate, and its posterior face is straight
to weakly concave. Subpetiolar process angular or snbangnlar,
and projecting posteriorly.
Key to the species of the Ponera tenuis group,
based on the worker
1 . Eyes absent 2
Eyes present, although represented only by a single ommatidium and
often very inconspicuous 3
2. Larger species, head width at least 0.44 mm; cephalic index at least
81 ; erect hairs numerous on scape, dorsum of alitrunk and entire
surfaces of first two gastric tergites ; antennal club indistinctly
5-jointed; body color clear yellow zwaluiuenburgi (Wheeler)
Smaller species, head width no more than 0.30 mm; cephalic index not
more than 78 ; erect hairs absent from scapes, alitrunk dorsum and
from all but the posterior strips of the first two gastric tergites ;
antennal club distinctly 5-jointed; body color brownish yellow ....
sweseyi (Wheeler )
3. Very small species, head width not exceeding 0.31 mm; (dorsal surface
of petiolar node seen from directly above, so that the posterior face
is level with the line of vision, forming in its entirety distinctly more
than a half -circle, its width 0.15 mm or less; body color light yellow-
ish brown) ; (New Guinea) .szabol Wilson, nom. nov.
Larger species, head width never less than 0.32 mm and often as
much as 0.38 mm or more; (dorsal surface of petiolar node seen
from above varying among species, from distinctly more than
semicircular to distinctly less, its width never less than 0.18 mm;
color variable between species, from light yellowish brown to very
dark brown) 4
4. Antennal club distinctly or indistinctly 5-jointed (see Fig. 1); (dorsal
surface of petiolar node seen from directly above forming distinctly
less than a half -circle) 5
Antennal club distinctly 4-jointed; (dorsal surface of petiolar node seen
from above forming in various species from distinctly less than a
half-circle to distinctly more) 6
5. Body color a uniform yellowish brown; posterior border of petiolar node
seen from directly above distinctly concave; (Java)
incerta (Wheeler)
Body color a uniform dark brown; posterior border of petiolar node
360 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
seen from directly above almost perfectly straight; (New Britain
to New Hebrides and Carolines) ratardorum Wilson, n. sp.
6. Smaller species, head width 0.34 mm or less; (body color clear yellow
to yellowish brown) 7
Larger species, head width 0.38 mm or more; (body color varying
among species, from light yellowish brown to dark brown) 8
RATARDORUM SZENTIVANYI
Fig. 1. Middle funicular segments in antennae of workers of two species
of the Ponera tenuis group, showing the principal character used to divide
i-ouplet 4 of the key. Dorsal view, semidiagrammatic.
7. Lateral surfaces of alitrunk very feebly shagreened to smooth, and
shining ; petiolar node relatively low, its height in the unique type
only 0.25 mm, or about the same as the pronotal width
petila Wilson, n. sp.
Lateral surfaces of alitrunk all moderately shagreened, and opaque;
petiolar node proportionately higher, its height in the single specimen
measurable 0.29 mm, or slightly more than the pronotal width, which
is 0.27 mm szentivanyi Wilson, n. sp.
8. Cephalic index 80 to 86 ; width of petiolar node not greater than 0.22
mm ; medium or dark brown species from New Guinea 9
Cephalic index 71 to 76; width of petiolar node not less than 0.22 mm;
light yellowish brown or light reddish brown species from New
Caledonia and Australia 10
9. Dorsal surface of petiolar node seen from directly above forming dis-
tinctly more than a half -circle (see Fig. 2) ; posterior apex of sub-
petiolar process sharply truncated; slightly smaller species, head
WILSON : TENUIS AND SELENOPHORA GROUPS OF PONERA 361
width 0.40-0.41 mm ; head dark brown, remainder of body medium
brown huonica Wilson, n. sp.
Dorsal surface of petiolar node seen from directly above forming almost
an exact half -circle or very slightly less; posterior apex of subpetiolar
process not truncated, but forming a full right-angle or an acute
angle; slightly larger species, head width 0.42-0.44 mm; entire body
uniformly dark brown tenuis (Emery)
10. Slightly smaller species, pronotal width 0.29-0.30 mm; scape index
80-86 ; entire petiolar node when viewed from directly above forming
almost an exact half-circle (see Fig. 2) ; (New Caledonia)
caledonica Wilson, n. sp.
(Based on the unique worker type). Slightly larger species, pronotal
width 0.32 mm ; scape index 90 ; entire petiolar node when viewed
from directly above forming distinctly more than a half -circle ;
(Victoria) exedra Wilson, n. sp.
PONERA CALEDONICA WilsOll, 11. Sp.
Holotype worker. HW 0.40 mm, HL 0.52 mm, SL 0.32 mm,
CI 77, SI 80, PW 0.30 mm, petiole height 0.31 mm, petiolar node
length 0.18 mm, dorsal petiole width 0.25 mm. Mandibles with
three well developed teeth occupying approximately the apical
two-fifths of the masticatory border ; the remainder of the border
occupied by an indeterminate number of minute denticles. Eyes
minute, consisting of a single ommatidium. Antennal club dis-
tinctly 4- jointed, considerably longer than the entire remainder
of the funiculus. Head seen in full-face view with nearly straight
sides, feebly concave occipital border. Petiolar node in side
view massive, subrectangular, tapering only very slightly dor-
sally ; seen from directly above, its dorsal surface forming an
almost exact half-circle, the posterior face concave. Subpetiolar
process somewhat reduced, its apex right-angular.
Dorsum of head densely punctate and subopaque to opaque ;
sides of head also densely punctate, but the punctures relatively
shallow and the surface feebly shining. Entire dorsal and lateral
surfaces of the alitrunk covered by puncturation or shagreening1
of variable density but everywhere shallow and feeble, so that the
i Shagreening as most rigorously defined means "covered with small, close-set
tubercles, suggesting shagreen leather," or "with a pebbled surface like shagreen
leather" (Webster's International Dictionary, unabridged, second edition). In
the present descriptions I have employed a somewhat broader definition of com-
mon usage in entomology, using this term to cover in addition to minute tubercu-
lation and pebbling any dense, irregular, minute sculpturing which cannot more
precisely be described as puncturation, striolation, vermiculation, or reticulation.
362
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
HUONICA
TENUIS
RATARDORUM
PETILA
CALEDONICA
SZABOI
Fig. 2. Lateral and dorsal view of the worker petioles of selected species
of the Panera tenuis group. Top: left, P. huonica Wilson, holotype;
middle, P. tenuis (Emery), worker from Ebabaang, New Guinea; right,
P. ratardorum Wilson, holotype. Bottom : left, P. petila Wilson, holotype ;
middle, P. eal.edonica Wilson, holotype; right, P. szaboi Wilson, worker from
the lower Busu River, Xew Guinea. Drawn approximately to scale.
WILSON : TENUIS AND SELENOPHORA GROUPS OF PONERA 363
surface varies from feebly to strongly shining. The dorsal petiolar
surface and gastric tergites are also feebly sculptured and their
surfaces overall feebly shining.
Body pilosity sparse, being limited almost entirely to a few
hairs on the anterior elypeal border, posterodorsal border of the
petiolar node, and entire surfaces of the gastric tergites. Ap-
pendages mostly bare, except for terminal surfaces of tibiae and
tarsal segments. Body and appendage pubescence everywhere
dense, very short and predominantly oblique to appressed.
Body concolorous yellowish ferruginous; appendages light
brownish yellow to clear yellow.
Worker paratype variation. HW 0.38-0.40 mm, HL 0.51-0.52
mm, SL 0,32-0.33 mm, CI 73-76, SI 83-86, PW 0.29-0.30 mm.
Queen paratype. HW 0.41 mm, HL 0.54 mm, SL 0.36 mm,
CI 76, SI 87, maximum eye length 0.09 mm, dorsal petiole width
0.26 mm. Distinguished from the worker by the usual queen-
worker caste differences. As in the queens of many other species
of Ponera, the petiolar node is much thinner than in the worker
caste, forming distinctly less than a half-circle when viewed
from directly above.
Relationships. P. caledonica forms with P. exedra Wilson of
Australia a discrete subgroup of the tenuis group, characterized
in the worker caste by relatively large size, elongated head, thick
petiolar node, and light coloration. The closest affinities of the
caledonica subgroup are evidently with the szentivanyi subgroup
{szentivanyi and petila). P. caledonica, can be distinguished from
P. exedra by its smaller size, thinner node, and lighter color.
Material examined. NEW CALEDONIA : Ciu, near Mt. Can-
ala, 300 m. (type locality), January 3, 1955, berlese sample of
12 workers and 1 dealate queen (E. O. Wilson) ; Mt. Mou, 180
m., December 11, 1954, 2 workers (Wilson, ace. no. 128), and
berlese samples of Dec. 12 and 27, 1954, 3 workers; Chapeau
Gendarme (Yahoue), Dec. 7, 1954, berlese sample of a single
worker (Wilson).
Ecological note. This is apparently a rather scarce cryptobiotic
species in New Caledonia. Despite rather intensive hand collect-
ing by the author in the localities cited above, only once (ace.
no. 128) was it encountered directly. In this case two workers
were found with a small amount of brood in a small cavity in the
364 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
undersurface of a rock set deeply in the soil. These individuals
were rather sluggish, and when prodded with the tip of a pair
of forceps, rolled up and feigned deatli for a short while, a
behavioral response common in other species of Ponera. All of the
other collections of calcdonica were made by filtering the ants
from masses of leaf litter and soil in a Berlese funnel. The col-
lections at Mt. Mou and Chapeau Gendarme were made in rela-
tively dry, semi-deciduous, valley-pocket forests, while that at
Ciu was in moister tropical evergreen forest.
Ponera exedra Wilson, n. sp.
Holotype worker. HW 0.38 mm, HL 0.54 mm, SL 0.34 mm,
CI 71, SI 90, PW 0.32 mm, petiole height 0.30 mm, petiolar node
length 0.20 mm, dorsal petiole width 0.24 mm. This species is
very close to P. calcdonica Wilson, and is distinguished by its
slightly larger size, more elongate head, and longer scapes, as
indicated in the measurements cited above. It also has a distinctly
thicker petiolar node ; when viewed from directly above, the
entire surface of the node forms slightly but distinctly more than
a half-circle. In addition, the subpetiolar process is somewhat
more reduced, and its apical angle is obtuse.
Queen paratype. (Tentative determination). HW 0.40 mm,
IIL 0.54 mm, SL 0.35 mm, CI 74, SI 88, petiolar node length
0.21 mm, dorsal petiole width 0.25 mm. Distinguished from the
worker by the usual queen-worker caste differences. Maximum
eye length 0.11 mm. Petiolar node thinner and more sharply
tapering than in worker; seen from directly above, the dorsal
surface of the node alone forms distinctly less than a half-circle,
but the entire node forms distinctly more. Head (except mandi-
bles) and first three gastric tergites medium brown; alitrunk
and petiole somewhat lighter yellowish brown; mandibles and
appendages brownish yellow to clear yellow.
Relationships. See under P. calcdonica.
Material examined. VICTORIA: Arthurs Seat (mountain)
at McCrae, 100-300 m. (type locality) ; April 28, 1951; a single
worker (W. L. Brown). NEW SOUTH WALES: Pymble;
October 23, 1950; a single dealate queen (Brown).
Ecological notes. Dr. Brown has supplied me with the follow-
ing information relative to the Arthurs Seat worker. This speci-
WILSON : TENUIS AND SELENOPHORA GROUPS OF PONERA 365
men was found under a rock in granitic soil in a medium
rainfall forest of Eucalyptis viminalis, E. radiata and Banksia
sp. Brown notes that the ant fauna of Arthurs Seat is unusual
for this part of Victoria, containing a number of distinctly north-
ern elements, e. g. Mayriella abstinens Forel and Camponotus
intrepiclus (Kirby). Thus the discovery of the Ponera exedra
queen at Pymble, N. S. W., hundreds of miles to the north, is not
too surprising. This latter specimen was collected from beneath
a rock in medium, dry sclerophyll forest on sandstone.
The close affinities of P. exedra to P. caledonica are of con-
siderable interest, insomuch as they represent another of a grow-
ing series of known links at the species-group level between the
ant faunas of NewT Caledonia and eastern Australia.
Ponera huonica Wilson, n. sp.
Holotype worker. HW 0.41 mm, HL 0.49 mm, SL 0.33 mm.
CI 84, SI 81, petiole height 0.33 mm, petiolar node length 0.18
mm, dorsal petiole width 0.22 mm. Right mandible with three
teeth occupying the apical half of the masticatory border, a
smaller tooth situated approximately midway between the basal-
most of the apical teeth and the basal angle, and an even smaller,
barely distinguishable tooth on the basal angle. There are no
intercalary denticles evident at magnifications up to 100X ;
higher magnifications were not used. The left mandible is
similar, but the median tooth described above is smaller and
rudimentary. Eye minute, consisting of a single ommatidium.
Antennal club massive, distinctly 4-jointed, slightly longer than
the remainder of the funiculus. Head shape about as described
in P. szaboi "Wilson. Petiolar node seen from the side relatively
thick, subrectangular, tapering very little dorsally; seen from
above its dorsal surface forms slightly more than a half -circle.
Subpetiolar process well-developed, its apex sharply truncated.
Mandibles smooth and shining ; clypeus somewhat less smooth,
and feebly shining; remainder of head roughly shagreened and
opaque. Pro- and mesonotum shagreened to contiguously punc-
tate, and subopaque ; episternum and various propodeal surfaces
variably punctate and feebly shining to subopaque. Petiolar node
mostly covered with scattered shallow punctures, feebly to
.366 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
strongly shining. C4astric tergites contiguously punctate and sub-
opaque.
Pilosity as in P. calcdonica but in addition with a few short
erect hairs on the occipital border, anterior pronotal angle,
and anterior gastric tergital surfaces. Pubescence dense, short,
predominantly oblique to appressed.
Head (except mandibles) medium brown; alitrunk, petiole,
and gaster light brown, the gaster a shade darker than the
rest; mandibles and appendages light brown.
Worker paratype variation. HW 0.40-0.41 mm, HL 0.48-0.50
mm, SL 0.33-0.35 mm, CI 80-83, SI 81-87, PW 0.29-0.32 mm.
The unusual mandibular dentition described for the holotype
occurs in most of the paratype workers. In several it is modified
by the addition of one or two smaller intercalary teeth or denti-
cles on the basal half of the masticatory border. In several others
the two principal teeth of the basal half are reduced to the size
of normal (for the tenuis group) denticles.
Queen paratypes. HW 0.43-0.45 mm, HL 0.51 mm, SL 0.35-
0.36 mm, CI 84-86, SI 80-82, dorsal petiole width 0.23-0.24 mm.
Differing from the worker caste by the usual queen-worker
differences. Compound eyes well developed, a,t least 0.29 mm
in maximum length. Petiolar node notably more slender in
side view ; seen from above its dorsal surface forms distinctly
less than a half -circle. Coloration similar to that of worker.
Male paratype. HW 0.43 mm, HL 0.45 mm, maximum eye
length 0.21 mm, dorsal petiole width 0.14 mm. Not differing
fundamentally in morphology from known Ponera males outside
the tenuis group. Antennae 13-jointed. Mandibles much re-
duced, only about 0.06 mm in length, edentate, with rounded
apices. Petiolar node seen from the side forming roughly an
isosceles triangle, with slightly concave anterior and posterior
faces and rounded dorsal crest ; seen from above, circular in out-
line. Genitalia exserted. Parameres small, 0.14 mm in length
(measured from distalmost edge of basiparamere to tip of para-
mere), tapering distally to a pointed apex. Penis valves large,
prominent, extending nearly 0.1 mm beyond the dorsal margin
of the parameres, their dorsal borders strongly convex, almost
semicircular.
Entire body covered with abundant, relatively short (length
WILSON: TENUIS AND SELENOPHORA GROUPS OF PONERA 367
never exceeding 0.06 mm) oblique to erect hairs, which merge
into the equally abundant underlying oblique pubescence.
Appendages almost entirely lacking pilosity, supplied instead
with dense, predominantly oblique pubescence.
Body uniformly dark brown ; appendages light to medium
brown; wings lightly and uniformly infumated.
Relationships. This species is closely allied to P. tenuis
(Emery) and P. ratardorum Wilson (q.v.).
Material examined. P. huonica has thus far been collected
only in a limited area in the mountainous region around the
headwaters of the Mongi River, Huon Peninsula, northeast
New Guinea. N-E NEW GUINEA: Ebabaang (type locality),
1300-1400 meters, April 16-18, 1955, 3 workers (E. O. Wilson,
ace. no. 826) and nest series with 2 workers, 2 alate queens,
and a male (ace. no. 827) ; Gemeheng, 1300-1500 m., April 11-13,
1955, worker, alate queen, dealate queen (Wilson, ace. no.
791) ; Joangeng, 1500 m., April 7-8, 1955, stray dealate queen
(Wilson, ace. no. 746).
Ecological notes. Colonies taken at Ebabaang and Gemeheng
were both small, containing probably less than 30 workers. The
one at Ebabaang was found under the moss layer covering the
upper surface of a large, soft, "rich-red" log. The Gemeheng
colony was in a small log in the same stage of decomposition.
At all localities the species was found in partly open areas at
the edge of native trails running through dense, wet midmoun-
tain rainforest.
Ponera incerta (Wheeler)
Pseudocryptopone bncerta Wheeler, 1933, Amer. Mus. Nov., no. 672: 18-19,
fig. 7, worker, queen. Type locality: Depok, Java.
The head of the unique worker type is now unfortunately
missing, so that exact measurements of cephalic and antennal
proportions could not be taken. In the present diagnosis cephalic
characters are taken from Wheeler's original description.
PW 0.29 mm, petiole height 0.29 mm, petiole node length
0.15 mm, dorsal petiole width 0.22 mm. Mandibles with three
apical teeth, behind which the masticatory border is "finely
and indistinctly crenulate" (denticulate?). Antennal club
5-jointed. Petiolar node seen from directly above forming dis-
368 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
tinctly less than a half -circle, with a shallowly concave posterior
border. Subpetiolar process well developed, its apical angle
acute.
Body sculpturing approximately as in P. huonica Wilson
(q.v.)
Body pilosity pattern as in P. caledonica Wilson. Pubescence
dense, short, predominantly appressed.
Body color uniformly yellowish ferruginous; legs clear yellow.
Relationships. This species is probably most closely related to
P. ratardorum Wilson, from which it can be distinguished by
slight differences in the antennal club composition and in petiolar
node shape, and by a strong color difference. It bears a close
habitus resemblance to P. szaboi, but can be easily distinguished
from that species by its 5-jointed antennal club, thinner petiolar
node, and somewhat more strongly shagreened alitruncal dorsum.
Material examined. JAVA : Depok, worker holotype.
PONERA PETILA Wilson, 11. Sp.
Holotype worker. HW 0.32 mm, HL 0.41 mm, SL 0.28 mm,
CI 78, SI 88, PW 0.25 mm, petiolar height 0.25 mm, petiolar node
length 0.13 mm, dorsal petiole width 0.18 mm. Very similar to
P. szentivanyi Wilson, differing slightly in body and appendage
proportions as given in the measurements cited above, and in the
much feebler body sculpturing, which can be described as fol-
lows. Sides of head densely but shallowly punctate, and feebly
shining. Entire dorsal and lateral surfaces of the alitrunk with
puncturation or shagreening of variable density but everywhere
shallow and feeble, so that the surface is feebly to strongly shin-
ing. The gastric tergites are also more feebly sculptured than
in szentivanyi and their surfaces overall feebly shining.
Relationships. Closely resembling P. szentivanyi Wilson, as
detailed in the comparative description of that species to follow.
Material examined. N-E. NEW GUINEA : lower Busu River,
near Lae; May 10, 1955; a single worker (Wilson, ace. no. 999).
Ecological note. The single worker was collected as a stray
in the superficial layers of soil beneath a rotting log on the
ground in primary lowland rainforest.
WILSON : TENUIS AND SELENOPHORA GROUPS OP PONERA 369
PONERA RATARDORUM Wilson, II. Sp.
Holotype worker. HW 0.37 mm, HL 0.47 mm, SL 0.31 mm,
CI 79, SI 84, PW 0.29 mm, petiole height 0.28 mm, petiolar node
length 0.15 mm, dorsal petiole width 0,20 mm. Mandibles with
three well developed teeth occupying about the apical two-
fifths of the masticatory border; the remainder of the border
occupied by a series of minute denticles. Eye minute, consisting
of a single ommatidium, located approximately 0.8 the distance
from the lateral occipital border to the midpoint of the anterior
genal border. Antennal club massive, indistinctly 5-jointed.
Petiolar node seen from side subrectangular, tapering almost
imperceptibly dorsally, the dorsal surface feebly convex; the
dorsal surface seen from directly above forms distinctly less
than a half-circle, and the posterior nodal border is almost per-
fectly straight. Subpetiolar process well developed, its apical
angle acute.
Sculpturing very similar to that described for P. huonica.
Pilosity and pubescence as described for P. caledonica. Body
medium brown, the head and gaster a shade darker than the
alitrunk and petiole. Appendages clear yellow.
Worker paratype variation. New Britain and New Hebrides:
HW 0.36-0.38 mm, HL 0.46-0.48 mm, SL 0.30-0.32 mm, CI 78-80,
SI 83-86, PW 0.27-0.30 mm. Carolines: HW 0.38 mm, HL
0.49 mm, SL 0.33 mm, CI 88, SI 87, PW 0.29 mm.
Relationships. This species most closely resembles P. incerta
(Wheeler) of Java, differing in its darker color, distinctly
5-jointed antennal club (versus indistinctly 5-jointed in incerta),
and straight posterior face of petiolar node. Superficially P.
ratardorum resembles P. huonica Wilson and P. tenuis (Emery)
but can be readily distinguished from these two species by its
5-jointed antennal club and much thinner petiolar node.
Material examined. NEW BRITAIN: St. Paul's, Baining
Mts., Gazelle Peninsula, 350 m. (type locality); Sept. 5, 1955;
holotype and three paratype workers (J. L. Gressitt). NEW
HEBRIDES: Ratard Plantation, 8 km. southwest of Luganville,
Espiritu Santo; Jan. 7-13, 1955; two paratype workers (Wilson,
ace. no. 348). CAROLINES : Yap I., one paratype worker (R. J.
Goss) . The holotype and two paratypes have been returned to the
Bishop Museum, Honolulu; the four remaining paratypes have
870 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
been deposited in the Museum of Comparative Zoology and
IT. S. National Museum.
This species is named in honor of Aubert and Suzanne Rataxd,
of Noumea and Luganville, the writer 's gracious hosts during his
brief stay in the New Hebrides.
Ecological notes. Dr. Gressitt's New Britain specimens were
taken from a rainforest humus berlesate. The present writer's
New Hebrides specimens were found foraging during the day
in leaf litter on the floor of primary coastal rainforest.
Ponera swezeyi (Wheeler)
Pseudocryptopone swezeyi Wheeler, 1933, Amer. Mus. Nov., no. 672: 16-17,
fig. 6, worker, queen. Type locality: vicinity of Honolulu, Hawaii.
The following measurements and descriptive notes are based
on three worker syntypes in the Museum of Comparative Zoology.
HW 0.29-0.30 mm, HL 0.38-0.41 mm, SL 0.25-0.26 mm, CI
72-78, SI 84-85, PW 0.21-0.24 mm, petiole height (single meas-
urement) 0.24 mm, dorsal petiole width 0.18 mm. Mandible with
three distinct apical teeth occupying slightly less than half the
masticatory border; the remainder of the border bearing an in-
determinate number of minute denticles. Antennal club dis-
tinctly 5-jointed. Petiolar node seen from directly above forming
slightly more than a half -circle. Subpetiolar process well de-
veloped, its apex right-angular.
Sculpturing about as in P. caledonica except that on the ali-
trunk only the declivitous faces of the propodeum are smooth
and shining, the remainder of the alitruneal surfaces being
lightly shagreened and only feebly shining.
Body concolorous light brownish yellow ; appendages clear
yellow.
Relationships. This distinctive little species does not appear to
be closely related to any of the other known members of the
tenuis group.
Material examined. HAWAII : vicinity of Honolulu, 3 syntype
workers (R. H. Van Zwaluwenburg) ; Herring Valley, Honolulu
(F. X. Williams).
Note on distribution. This species is known only from material
collected in the vicinity of Honolulu. The habitat of the type
series, "soil of cultivated and fallow sugar-cane fields," suggests
WILSON: TENUIS AND SELENOPHORA GROUPS OP PONERA 3 7 1
that it may have been introduced by man into the Hawaiian
Islands. Future collecting may show that its native range lies
somewhere in the "source areas" of Melanesia or the East Indies.
Ponera szaboi Wilson, noni. now
Crypt opone mocsaryi Szabo, 1910, Rovartani Lap., 17: 186, fig. 1, worker.
Secondary homonymy by present assignment to Ponera (nee Ponera
mocsaryi Emery, 1900). Type locality: Friedrich-Wilhelmshafen
(=Madang), N-E. New Guinea.
Pseudoeryptopone mocsaryi, Wheeler, 1933, Araer. Mus. Nov., no. 672:14.
The description offered below is based on two workers collected
by myself in the vicinity of the lower Busu River, N-E. New-
Guinea. These correspond well to Szabo 's description and figure,
differing only in having somewhat more elongate heads than
shown by Szabo.
HW 0.30-0.31 mm ; HL 0.40 mm ; SL 0.25 mm ; CI 76-78 ; SI
79-83; P\V 0.23 mm; petiole height (single measurement) 0.24
mm; petiolar node length 0.15 mm; dorsal petiole width (single
measurement) 0.15 mm. Mandible linear-subtriangular. The
apical half of the masticatory border occupied by three distinct,
acute teeth ; the basal half occupied by an indeterminate number
of minute denticles. Eyes minute, consisting of a single omma-
tidium. Antennal club massive, distinctly 4-jointed, consider-
ably longer than the entire remainder of the funiculus. Head
in full-face view subrectangular, with very feebly convex sides
and feebly concave posterior border. Petiolar node seen from
the side relatively thick, tapering slightly dorsally, with a feebly
convex dorsal border ; seen from directly above, with the posterior
face aligned with the plane of vision, the node forms distinctly
more than a half-circle, and the posterior border appears almost
perfectly straight. Subpetiolar process well-developed, its apical
angle obtuse.
Mandibles smooth and shining ; clypeus smooth and feebly
shining; remainder of head finely and evenly shagreened and
subopaque. All of alitruncal surfaces finely shagreened and
subopaque, except the episterna and declivitous faces of the
propodeum, which bear only scattered fine punctures and are
relatively smooth and more or less shining. Various surfaces of
the petiolar node bearing variably dense but fine and separated
372 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
punctures, and otherwise smooth and more or less shining.
Gastric tergital surfaces shagreened and subopaque, except for
the anterior declivity of the first gastric tergite, which is smoother
and feebly shining.
Pilosity and pubescence as described for P. caledonica Wilson.
Alitrunk and petiole yellowish brown ; head and gaster some-
what darker, approaching medium brown ; appendages nearly
clear yellow.
Relationships. In its distinctive combination of characters
in size, petiole form, and body color, szaboi stands well apart
from all the other known species of the tenuis group.
Material examined. N-E. NEW GUINEA : lower Busu River,
near Lae, 2 workers (Wilson, ace. nos. 963, 1024).
Ecological note. Both of the Busu River specimens were taken
as strays on the floor of primary lowland rainforest.
PONERA SZENTIVANYI Wilson, n. sp.
Holotype worker. IIW 0.34 mm; HL 0.45 mm, SL 0.32 mm,
CI 76, SI 94, PW 0.28 mm (petiole height not measured; see
paratype), petiolar node length 0.16 mm, dorsal petiole width
0.24 mm. Three well developed teeth occupying the apical two-
fifths of the masticatory border, followed basally by an indeter-
minate number of minute denticles. Eye minute, consisting of a
single ommatidium, located about 0.8 the distance from the lateral
occipital border to the midpoint of the anterior genal border.
Antennal club distinctly 4-jointed, considerably longer than the
remainder of the funiculus. Head elongate (CI 76) with very
feebly convex sides, and feebly but distinctly concave occipital
border. Petiolar node seen from side relatively thin, elongate-
trapezoidal; seen from directly above, so that the posterior face
is exactly parallel with the line of vision, the node as a whole
forms slightly more than a half-circle, but the dorsal surface
alone forms much less than a half-circle ; seen from the preceding
position the posterior face is feebly but distinctly concave.
Subpetiolar process somewhat reduced, its apex right-angular.
Body sculpturing approximately as described for P. huonica
Wilson.
Pilosity and pubescence as in P. caledonica Wilson.
WILSON : TENUIS AND SELENOPHORA GROUPS OF PONERA 373
Body uniformly light brownish yellow; appendages clear
yellow.
Paratype worker. A single callow worker taken with the
holotype has the integument of the head somewhat crumpled
and distorted through drying, so that regular cephalic measure-
ments could not be made. PW 0.28 mm, petiole height 0.29 mm,
dorsal petiole width 0.21 mm. Body color clear, pale yellow.
Relationships. This species most closely resembles P. petila
Wilson, as indicated under the comparative description of that
species. Together szentivanyi and petila form a subgroup of their
own within the tenuis group, characterized in the worker caste
by intermediate size, slender body form with elongate head, thin
petiolar node, and brownish yellow body color. They are closest
to the subgroup formed by P. caledonica Wilson and P. exedra
Wilson, from which they can be distinguished by their smaller
size and thinner petiolar node.
Material examined. PAPUA : Karema, near the Brown River,
about 30 miles north of Port Moresby; March 8-11, 1955; holo-
type and single paratype worker (Wilson, ace. no. 563). This
species is named in honor of Dr. J. H. Szent-Ivany, the expert
resident entomologist of the Territory of Papua-New Guinea,
whose friendly assistance greatly aided the author's field work
in this area.
Ecological note. The two type workers were taken close to-
gether on the floor of primary lowland rainforest.
Ponera tenuis (Emery)
Cryptopone tenuis Emery, 1900, Termeszetr. Fiiz., 23: 321-322, pi. 8, figs.
21, 22, worker. Original localities: Lemien, near Berlinhafen
(=Aitape), and Tamara I., N-E. New Guinea.
Pseiulocryptopone tenuis, Wheeler, 1933, Amer. Mus. Nov., no. 672: 13-14.
Lectotype worker.1 HW 0.44 mm, HL 0.52 mm, SL 0.35 mm,
CI 85, SI 80, PW 0.32 mm, petiole height 0.32 mm, petiolar node
length 0.17 mm, dorsal petiole width 0.22 mm. Right mandible
with three2 rather worn, indistinct teeth occupying the apical
half of the masticatory border; followed by an indeterminate
number of minute, blunt denticles occupying the basal half of
i By present selection, a syntype worker in the Emery Collection, kindly loaned
to the author by Dott. Delta Guiglia.
2 Not four as stated by Emery in the original description.
374 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
the border. Eyes minute, consisting of a single ommatidium.
Antennal club massive, distinctly 4-jointed, considerably longer
than the remainder of the funiculus. Petiolar node seen from
the side subtrapezoidal, tapering slightly dorsally, with a con-
vex dorsal margin, its dorsal surface seen from directly above
forming almost exactly a half-circle or very slightly less. An-
terior half of subpetiolar process perforated by a small, median
hole ; the posterior apex of the process forming an acute angle.
Sculpturing as described for P. huo7iica Wilson.
Pilosity and pubescence as described for P. huonica, except
that erect hairs are lacking from the occiput and pronotum;
these missing hairs may well have been rubbed off in this
specimen, because they are present in more recently collected
material determined as tenuis.
Body uniformly yellowish brown, appendages clear yellow
(specimen possibly faded; see below).
Variation in other worker series. The following measurements
are based on three workers from a single nest series collected at
Ebabaang: HW 0.42-0.43 mm, HL 0.49-0.52 mm, SL 0.33-0.35
mm, CI 82-86, SI 78-83, PW 0.32 mm. In these specimens the
anterior half of the subpetiolar process is not perforated as in
the lectotype, and the posterior angle of the process forms
a right angle instead of an acute angle. The body color
is uniformly blackish brown, and the appendage color is light
yellowish brown; there is an excellent possibility that the con-
siderably lighter color of the lectotype is due to fading.
Queen (tentative determination). IIW 0.50 mm, HL 0.58 mm,
SL 0.40 mm, CI 86, SI 80, dorsal petiole width 0.27 mm. Dif-
fering from the worker by the usual formicid queen-worker
differences. Maximum eye length 0.15 mm. Petiolar node much
thinner than in worker, seen from directly above forming much
less than a half-circle. Distinguished from the queen of P.
huonica by its larger size, darker body color (uniformly blackish
brown as opposed to predominantly medium brown in huonica),
and somewhat thinner petiolar node.
Relationships. P. tenuis most closely resembles P. huonica
Wilson, from which it differs principally in its more conventional
mandibular dentition, thinner petiolar node, differently shaped
subpetiolar process, and darker body color. Together these two
species bear a close habitus resemblance to P. clavicornis Emery,
WILSON : TENUIS AND SELENOPHORA GROUPS OF PONERA 375
and may in fact provide a link between the tenuis and seleno-
phora species groups.
Material examined. N-E. NEW GUINEA: (Lemien or Ta-
mara I.), lectotype worker; Ebabaang, Mongi River Watershed,
1300-1400 m., 3 workers (Wilson, ace. no. 828) ; Joangeng, near
Ebabaang, 1500 m., a stray dealate queen (Wilson, ace. no. 746).
Ecological note. The Ebabaang workers were found foraging
during the day in leaf litter on the floor of midmountain rain-
forest.
Ponera zwaluwenburgi (Wheeler)
Pseudocryptoponc zwaluwenburgi Wheeler, 1933, Amer. Mus. Nov., no. 672:
14-16, fig. 5, worker. Type locality: Oahu Island, Hawaii (by present
selection).
The following measurements and descriptive notes are based
on four worker syntypes in the Museum of Comparative Zoology.
HW 0.44-0.47 mm, HL 0.53-0.59 mm, SL 0.38-0.42 mm, CI
81-84, SI 87-88, PW 0.32-0.35 mm, petiole height (single meas-
urement) 0.28 mm, dorsal petiole width 0.20-0.23 mm. Mandibles
with three well developed teeth occupying less than half the
masticatory border, the remainder being occupied by an inde-
terminate number of minute denticles. Antennal club indistinctly
5-jointed. Petiolar node seen from directly above forming dis-
tinctly more than a half-circle. Subpetiolar process reduced to
a mere convexity.
Entire body finely and densely shagreened and subopaque.
except the mandibles, posterior face of the propodeum, and
posterior face of the petiola,r node, which are relatively smooth
and shining.
Short erect hairs numerous on anterior scape surface, entire
dorsum of alitrunk, petiolar dorsum, and entire surfaces of ex-
posed gastric tergites. Pubescence everywhere abundant, almost
entirely appressed.
Body and appendages concolorous clear yellow.
Relationships. This species, marked by its combination of
large size, lack of eyes, abundant erect pilosity, and pale color,
does not appear to stand close to any of the other species of the
tenuis group.
Material examined. HAWAII : Oahu, four syntype workers.
376 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
This species is known only from the type material, which was
collected at several localities on Oahu and on Maui. As in P.
sivezeyi, the collections were all made from the soil of cultivated
and fallow sugar-cane fields, a circumstance which suggests that
the species may have been introduced into Hawaii by man.
Characterization of the Ponera selenophora group
Worker. Medium-sized species, worker head width ranging
between 0.43 and 0.68 mm. Mandibular dentition variable, in
most cases consisting of three distinct teeth occupying the apical
two-fifths to one-half of the masticatory border, followed by a
series of minute denticles; in one case (selenophora) the basal
half of the border bears two distinct teeth in addition to the
denticles. Eyes usually small, with 3-5 indistinct facets; in one
case (elegantula) there are 11-12 facets. Junction of posterior
and lateral faces of propodeum marginate, forming an angle
of 80° to slightly less than 90°. Petiolar node massive and ex-
ceptionally broad, its dorsal width never less than 0.77 X the
pronotal width and usually much more ; arcuate or crescentic,
with the junction of the anterior and posterior faces usually
marginate. Subpetiolar process very variable in shape, from well
developed and angular or subangular to rudimentary and
rounded; when well developed, its apex generally projects pos-
teriorly.
Key to the species of the Ponera
selenophora group, based on the worker
1. Small species, head width 0.43-0.47 mm; dorsum of alitrunk completely
devoid of standing hairs; (Xew Guinea to New Hebrides)
clavieornis Emery
Larger species, head width 0.50 mm or greater; dorsum of alitrunk
either covered by abundant standing hairs (usually) or else com-
pletely devoid of standing hairs (the latter condition only in one
species from New Guinea) 2
2. Entire lateral surface of the propodeum completely sculptured and
opaque; (a larger species, head width 0.62-0.64 mm, with unusually
long scapes, scape index 90-93) ; (Japan) scdbra Wheeler
A large part of the lateral faces of the propodeum smooth and shining ;
(species variable in size, with shorter scapes, scape index 89 or less)
3
WILSON : TENUIS AND SELENOPHORA GROUPS OF PONERA 377
3. (Based on unique type). Head more elongate (cephalic index 80), with
relatively large eyes containing 11 or 12 distinct facets; alitrunk
completely devoid of standing hairs; (mountains of northeastern
New Guinea) elegantula Wilson, n. sp.
Head proportionately shorter (cephalic index 86 or greater), with
smaller eyes containing only 3-5 indistinct facets; alitrunk covered
with abundant standing hairs 4
4. Smaller species (head width 0.50 mm) with proportionately short head
(cephalic index 92-94); antennal club indistinctly 4-jointed; (Philip-
pines) oreas (Wheeler)
Either slightly larger species (head width 0.52-0.54 mm) with much
longer head (cephalic index 85-87), or much larger species (head
width 0.59 mm or greater) with head equally long to much longer
(cephalic index 86-92) ; antennal club either 5-jointed or completely
undifferentiated 5
5. Posterior face of petiolar node feebly but distinctly convex; a relatively
small species (head width of unique type 0.52 mm) from the moun-
tains of northeastern New Guinea syscena Wilson, n. sp.
Posterior face of petiolar node flat or feebly concave 6
6. Smaller species (head width of unique type 0.54 mm) ; anterior surface
of scape with abundant erect hairs; antennal club indistinctly
5-jointed ; (Hongkong) sinensis Wheeler
Larger species (head width 0.59 mm or greater) ; erect hairs scarce
to absent on anterior surface of scape; antennal club undifferenti-
ated 7
7. Smaller species (head width 0.59-0.63 mm) ; basal half of masticatory
border bearing two distinct teeth which are nearly as large as the
three teeth of the apical half; posterior border of petiolar node, seen
from directly above, distinctly concave ; (lowland rainforests of
Papua and northeastern New Guinea) selenophora Emery
Larger species (head width 0.65-0.68 mm) ; basal half of masticatory
border bearing only minute denticles which do not approach in size
the three apical teeth; posterior border of petiolar node, seen from
directly above, straight; (mountains of northeastern New Guinea)
xenagos Wilson, n. sp.
Ponera clavicornis Emery
Ponera clavicornis Emery, 1900, Termeszetr. Fiiz., 23: 317, pi. 8, figs. 7, 8,
worker. Type locality: Friedrich-Wilhelrnshafen (=Madang), N-E.
New Guinea.
Ponera clavicornis, Mann, 1919, Bull. Mus. Comp. Zool., 63: 296.
Selenopone clavicornis, Wheeler, 1933, Amer. Mus. Nov., no. 672 : 22.
378 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Worker. HW 0.43-0.47 mm, HL 0.52-0.59 mm, SL 0.35-0.42
mm, CI 81-85, SI 80-89, PW 0.32-0.37 mm, dorsal petiole width
0.27-0.32 mm. Mandibles with three well developed teeth occupy-
ing about the apical half of the masticatory border; the basal
half occupied by an indeterminate number of minute denticles.
Eye as described for P. selenophora. Antennal club relatively
slender, 5-jointed. Posterolateral margins of propodeum rela-
tively poorly defined, seen from directly above forming an angle
of only slightly less than 90°. Posterior face of petiolar node
seen from directly above almost perfectly straight. Subpetiolar
process variable in shape, ranging from a rudimentary convexity
to a strong right-angular projection.
Mandibles smooth and shining ; clypeus feebly shagreened and
shining over most of its surface ; entire remainder of the head
densely, finely, and evenly punctate (the punctures mostly under
0.01 mm in diameter) and completely opaque. Entire dorsal and
lateral alitruncal surfaces similarly punctate and opaque, except
for the ventral margins of the sides of the pronotum, a limited
central longitudinal strip on the sides of the propodeum, and
the lower half of the posterior propodeal face, which surfaces
are more or less smooth and shining. Dorsal and lateral surfaces
of petiolar node somewhat less densely punctate than head and
alitrunk, subopaque; anterior and posterior faces more or less
smooth and shining. First several gastric tergites also somewhat
less densely punctate, subopaque to feebly shining.
Pilosity completely lacking on head and alitrunk except for
a few erect hairs on the mandibles, clypeus, and frontal lobe
area. Petiolar node and first two gastric tergites bare to sparsely
pilose; terminal gastric tergites and all gastric sternites abund-
antly pilose.
Body (except mandibles and apical gastric segments) piceous
brown, approaching jet black. Mandibles, apical gastric seg-
ments, and appendages yellowish brown.
Geographic variation. The series from Espiritu Santo, New
Hebrides, have somewhat longer scapes than those from New
Guinea (SI 86-89 as opposed to 80-84).
The series from Bisianumu, Papua, have the first two gastric
tergites pilose ; in side view 5-10 standing hairs are visible along
the profile of the first tergite. The series from Tumnang, N-E.
New Guinea, and from the New Hebrides have the first two
WILSON : TENUIS AND SELENOPHORA GROUPS OF PONERA 379
gastric tergites bare of pilosity. The series from Bubia, N-E.
New Guinea, a geographically intermediate locality, has the
tergites intermediately pilose: 1-3 standing hairs are visible
along the profile of the first tergite.
Relationships. This is a very distinct species, easily separated
in the worker caste from other members of the selenophora group
by the combination of smaller size, distinctive sculpturing, and
sparse body pilosity.
Material examined, PAPUA : Bisianumu, 500 m., March 15-20,
1955 (Wilson, ace. nos. 608, 626, and 648, the last with winged
queens). N-E. NEW GUINEA: Madang, syntype worker
(Emery Coll.); Bubia, 13 km. northwest of Lae, March 26,
1955 (Wilson ace. no. 680) ; lower Busu River, May 1955 (Wilson
ace. no. 1006) ; Tumnang, Mongi River Watershed, 1500 m..
April 14-15, 1955 (Wilson ace. no. 798). SOLOMONS: Santa
Isabel (Mann, 1919). NEW HEBRIDES: A. Ratard Planta-
tion, 8 km. southwest of Luganville, Espiritu Santo, January
7-13, 1955 (Wilson ace. no. 348) ; Malua Bay, Malekula (L. E.
Cheesman). My accessions no. 608 and no. 798 were compared
directly with a worker syntype in the Emery Collection.
Ecological notes. This is an exceptionally adaptable and
widespread species. It has been collected from primary lowland
rainforest (Espiritu Santo), second-growth lowland rainforest
(Bubia), foothills forest (Bisianumu), and true midmountain
forest (Tumnang), under a variety of local ecological conditions.
My accession no. 608 (Bisianumu) was a small colony found
nesting under the bark of a large ' ' passalid-stage " log on the
ground ; larvae at various stages of development and cocoons
were present. The other two Bisianumu accessions and the
Bubia accession consisted of stray workers, also from large
passalid-stage logs. The Tumnang and New Hebrides specimens
were taken as strays in leaf litter on the forest floor.
PONERA ELEGANTULA WilsOll, n. Sp.
Holotype worker. HW 0.56 mm, HL 0.70 mm, SL 0.49 mm,
CI 80, SI 87, PW 0.43 mm, petiolar node length 0.38 mm, dorsal
petiole width 0.38 mm. Apical half of masticatory border of
(left) mandible occupied by three well-developed teeth; posterior
half occupied by six irregular denticles. Eyes relatively large,
380 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
maximum length 0.06 mm, with eleven or twelve distinct facets.
No distinct antennal club differentiated ; funicular segments from
the third outward gradually increasing in length and width.
Head subrectangular, its sides feebly convex, its posterior
border feebly concave. Posterolateral margins of propodeum dis-
tinct but rounded, seen from above forming only slightly less
than a 90° angle. Petiolar node in side view considerably thin-
ner than in any other selenophora group member, although ex-
hibiting the form and exceptional transverse width typical for
the group. Seen from directly above, the dorsal surface of the
node is thin and arc-shaped.
Mandibles smooth and feebly shining; clypeus for the most
part smooth and feebly shining. Kemainder of head covered
by punctures which are about 0.006 mm in diameter and sep-
arated from one another by about the same distance ; its surface
feebly shining. Dorsal surface of alitrunk covered by similar
punctures somewhat more widely spaced, its surface feebly shin-
ing; punctures on sides of pronotum sparser, finer, the surface
moderately shining; lower halves of episterna finely and very
sparsely punctate, their surfaces strongly shining, upper halves
finely shagreened and subopaque ; lower halves of metapleura
and of the sides of the propodeum shagreened and subopaque,
upper halves finely and sparsely punctate and shining ; posterior
face of propodeum smooth and shining. Petiolar surfaces very
sparsely punctate to smooth, moderately to strongly shining.
Gastric tergites sculptured similarly to alitruncal dorsum.
Body pilosity very sparse, limited to anterior region of head,
posterior strips of first two gastric tergites, entire surfaces of
apical gastric tergites, and entire surfaces of all gastric sternites.
Pubescence everywhere abundant and appressed.
Body color, excluding mandibles and apical gastric tergites,
jet black. Mandibles, apical gastric tergites, and appendages
yellowish brown.
Paratype queen. HW 0.62 mm, II L 0.76 mm, SL 0.53 mm, CI
82, SI 86. Distinguished from the worker by the usual queen-
worker caste differences. Maximum eye length 0.16 mm. Unlike
the queens of other members of the tenuis and selenophora
groups, the queen of elegant ula does not have a proportionately
more slender petiolar node than the worker.
Relationships. This species, with its distinctively large eye
WILSON : TENUIS AND SELENOPHORA GROUPS OP PONERA 381
size and relatively slender petiolar node, does not appear to be
closely related to any of the other members of the selenophora
group.
Material examined. N-E. NEW GUINEA : Tumnang, Mongi
River Watershed, Huon Peninsula, 1500 m. ; April 14-15, 1955 ;
one worker and one dealate queen (Wilson, ace. no. 799).
Ecological note. The two type specimens were taken together
under the bark of a rotting log in midmountain rainforest.
Ponera oreas (Wheeler)
Selenopone oreas Wheeler, 1933, Amer. Mus. Nov., no. 672: 20-21, fig. 8,
worker. Type locality: Cuernos Mrs., 1300 m., near Dumaguete, Negros
Oriental, Philippines.
}Yorker. HW 0.50 mm, HL 0.54 mm, SL 0.39 mm, CI 93, SI 78,
PW 0.36-0.38 mm, dorsal petiole width 0.29-0.31 mm. Closely
related to the members of the selenophora "subgroup" (see
under selenophora) and distinguished principally by the follow-
ing characters :
(1) Somewhat smaller size.
(2) The antennal club is f our- jointed ; the fifth funicular
segment from the apex is slightly larger than the succeeding basal
segments, but still not large enough to be considered part of the
club, as is the case in P. sinensis.
(3) The head is proportionately shorter than in any other
member of the selenophora group.
Relationships. See comparative description above.
Material examined. PHILIPPINES : Cuernos Mts., four syn-
type workers.
Ponera scabra Wheeler
Ponera scabra Wheeler, 1928, Boll. Lab. Zool. Portici, 21: 99-100, worker,
queen. Type locality: Mt. Maya, Japan (present selection).
Worker. HW 0.62-0.64 mm, HL 0.77-0.80 mm, SL 0.56-0.59
mm, CI 78-82, SI 90-93, PW 0.46-0.50 mm, dorsal petiole width
0.39-0.42 mm. This species falls close to the selenophora "sub-
group" (see under selenophora) , and can easily be distinguished
from it by the following two characters:
(1) The head is proportionately longer (maximum CI is 82
as opposed to a minimum of 85 in the selenophora subgroup).
382 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
(2) The entire alitrunk, except the posterior surface of the
propodeum, is coarsely and irregularly shagreened and sub-
opaque to opaque.
Relationships. See comparative description above.
Material examined. JAPAN: Mt. Maya, Honshu, six syntype
workers.
Ponera selenophora Emery
I'uncra selenophora Emery, 1900, Termeszetr. Fiiz., 23: 317, pi. 8, figs. 4, 6,
worker. Type locality: Lemien, near Berliuhafen (Aitape), N-E. New
(iuinea.
Selenopone selenophora, Wheeler, 1933, Amer. Mus. Nov., no. 672: 21.
Worker. HW 0.59-0.63 mm, HL 0.66-0.69 mm, SL 0.52 mm,
CI 88-92, SI 82-89, PW 0.46-0.49 mm, dorsal petiolar width
0.40-0.42 mm. Mandibles with three relatively large, well-de-
veloped teeth occupying the apical half of the masticatory
border; the basal half occupied by two smaller teeth, one located
midway between the basalmost of the apical teeth and the basal
angle, and the other on the basal angle. In addition, there are
several irregular denticles in the interdentary spaces of the
basal half of the border. Eyes minute, consisting of two or three
small, ill-defined ommatidia, located approximately 0.8 the dis-
tance from the lateral occipital border to the midpoint of the
anterior genal border. The antenna lacks a well-defined club,
the funicular segments merely increasing gradually in length
and width from the fourth outward. Posterolateral margins of
propodeum (line of juncture of posterior and lateral faces) well
marked, seen from directly above forming an angle of about
80°. Posterior border of petiole when viewed from directly above
distinctly concave. Subpetiolar process well developed, approxi-
mately right-angular.
Mandibles and most of clypeus smooth and shining. Entire
rest of head covered by contiguous punctures about 0.01 mm or
slightly less in diameter, completely opaque. Entire dorsum
of alitrunk covered by punctures about 0.006 mm in diameter,
separated by spaces of about the same width as the diameter of
the punctures, the surface feebly shining. Lateral thoracic-
surface covered by punctures of variable size, most with diameter
under 0.01 mm, the majority contiguous; the surface subopaque.
WILSON: TENUIS AND SELENOPHOKA GROUPS OF PONERA 383
The lateral and posterior propodeal faces bear only a few peri-
pherally distributed punctures and are mostly smooth and
shining. Petiolar node with sparse scattered punctures, its sur-
face entirely smooth and shining.
Short, erect hairs present on mandibles, clypeus, frontal lobe
area, entire dorsal alitruncal surface, posterolateral propodeal
margins, dorsal petiolar surface, and entire surfaces of first two
gastric segments. Apical gastric segments covered by more
abundant, much longer hairs. Pubescence almost everywhere
abundant, predominantly oblique to appressed.
Fig. 3. Lateral and dorsal views of the worker petiole of Ponera
selenophora Emery. Based on a worker from Karema, Papua, which had
been compared with a syntype in the Emery Collection.
Entire body jet black, except mandibles and apical gastric
segments, which are brownish yellow. Appendages variably
brownish yellow.
Relationships. Inside the selenophora group, P. selenophora
falls within the closely knit subgroup which also includes P.
xenagos Wilson, P. syscena Wilson and P. sinensis Wheeler. Dis-
tinguishing characters are supplied in the respective comparative
descriptions of these latter species.
Material examined. N-E. NEW GUINEA: Lemien, near
Berlinhafen (=Aitape), syntype worker; lower Busu River, near
384 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Lae, April 28, 1955 (E. 0. Wilson, aec. no. 564). PAPUA:
Karema, near Brown River, March 8-11, 1955 (Wilson, ace. no.
564). NETH. NEW GUINEA: Maffin Bay, June, 1944, a single
dealate queen (E. S. Ross). The Karema specimens have been
compared directly with a worker type in the Emery Collection.
Ecological note. Both of the author's collections were made
on the floor of primary lowland rainforest.
Ponera sinensis Wheeler
Ponera sinensis Wheeler, 1928, Boll. Lab. Zool. Portici, 22: 6-7, worker.
Type locality : Hongkong.
Holotype worker. HW 0.54 mm, HL 0.62 mm, SL 0.45 mm,
CI 87, SI 83, PW 0.41 mm, dorsal petiole width 0.35 mm. Very
close to P. selenophora and P. syscena, differing primarily by
the following combination of characters :
(1) Intermediate size.
(2) Apical five segments of antenna differentiated as a club.
(3) Posterior face of petiolar node feebly but distinctly con-
cave, approximately intermediate between selenophora and xena-
gos.
(4) Pilosity and pubescence approximately as described for
syscena.
(5) Propodeal margination as described for xenagos.
(6) Basal half of masticatory border of mandible bearing only
denticles.
Relationships. See comparative description above.
Material examined. Hongkong, holotype worker.
Ponera syscena Wilson, n. sp.
Holotype worker. HW 0.52 mm, HL 0.61 mm, SL 0.45 mm,
CI 85, SI 87, PW 0.40 mm, petiolar height 0.39 mm, petiolar node
length 0.26 mm, dorsal petiole width 0.31 mm. Closely related
to P. selenophora Emery and P. sinensis Wheeler, differing pri-
marily by the following combination of characters :
(1) Small size, distinctly smaller than the probably sympatric
P. selenophora but scarcely smaller than P. sinensis.
(2) Dorsal petiole width only 0.78X the pronotal width, as
opposed to at least 0.82X in selenophora and sinensis. Posterior
WILSON : TENUIS AND SELENOPHORA GROUPS OF PONERA 385
face of petiolar node feebly convex (feebly concave in seleno-
phora and sinensis).
(3) Propodeal margination as described for P. xenagos.
(4) Body and appendages with considerably more abundant
pilosity and pubescence than in selenophora. Thirteen to seven-
teen outstanding erect, hairs can be counted along the outer sur-
faces of the scapes in the syscena type, whereas there are no
more than five or six in selenophora. P. sinensis is close to P.
syscena in this character.
(5) Dentition of basal half of masticatory border of mandible
bearing only denticles.
Relationships. See comparative description above. Although
this species closely resembles P. selenophora in most characters,
it has a petiolar node form (q. v.) which is intermediate between
the distinctive selenophora type and the more generalized type
characterizing most of the species of Ponera.
Material examined. N-E. NEW GUINEA: native trail be-
tween Yunzain and Joangeng, Mongi Watershed, Huon Penin-
sula,, 1300 m.j April 7, 1955; a single worker (Wilson).
Ecological note. The unique type was taken as a stray on the
floor of midmountain rainforest.
Ponera xenagos Wilson, n. sp.
Holotype worker. HW 0.67 mm, HL 0.77 mm, SL 0.59 mm,
CI 87, SI 88, PW 0.52 mm, petiole height 0.53 mm, petiolar node
length 0.27 mm, dorsal petiole width 0.42 mm. Very similar to
P. selenophora Emery, differing by the following characters :
(1) Larger size.
(2) The three apical mandibular teeth occupy less than half
the masticatory border, and distinct teeth are not developed on
the basal half of the border as described for selenophora.
(3) The posterolateral margins of the propodeum are less
pronounced; viewed from directly above they form an angle of
only a little less than 90°.
(4) When viewed from directly above, the posterior margin
of the petiolar node is almost perfectly straight, as opposed to
the distinctly concave margin of selenophora.
(5) Pubescence is generally sparser. The anterior face of
the petiolar node has pubescence only over its upper quarter,
386 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
and there it is relatively sparse, whereas in selenophora it is
abundant over the entire upper half.
Paratype variation. IIW 0.65-0.68 mm, HL 0.75-0.80 mm,
SL 0.57-0.60 mm, CI 86-90, SI 83-89, PW 0.52-0.54 mm, dorsal
petiole width 0.40-0.44 mm.
Relationships. P. xenagos is the largest of the known species
of the selenophora group. Within the group, it is most closely
allied to selenophora itself, as indicated in the above comparative
description.
Material examined. N-E. NEW GUINEA: Tumnang, 1500 m.
(type locality), April 14-15, 1955, holotype and eight paratype
workers (Wilson, ace. no. 801) ; Ebabaang, 1300-1400 m., April
16-18, 1955, three paratype workers (Wilson, ace. no. 819).
Both of the above localities are in the Mongi River Watershed
of the Huon Peninsula.
Ecological notes. The Tumnang colony was found nesting
under the loose bark of a rotting stump. The Ebabaang colony
was under the loose bark of the upper surface of a large rotting
log, in the immediate vicinity of a colony of Amblyopone australis
Erichson. Both nest sites were in partial clearings at the side
of native trails running through dense midmountain rainforest.
Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE
Vol. 116, No. 7
THE CHINESE CAENERESSA SPECIES
(LEPIDOPTERA, CTENUCHIDAE)
By Nicholas S. Obraztsov
With Four Plates
CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM
June, 1957
Publications Issued by or in Connection
with THE
MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE
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Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE
Vol. 116, No. 7
THE CHINESE CAENERESSA SPECIES
(LEP1D0PTERA, CTENUCHTDAE)
By Nicholas S. Obraztsov
With Foin Plates
CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM
June, 1957
No. 7 — The Chinese Caeneressa Species (Lepidoptem ,
Ctenuchidae)
By Nicholas S. Obraztsov
TABLE OP CONTENTS
Page
LNTKODUCTION 389
ABBREVIATIONS 39i
SYSTEMATIC DESCRIPTIONS
Caeneressa n. gen 392
Pattern of the Body and Wings 393
Male Genitalia 39,5
Female Genitalia 396
Systematic Position 390
Range 39S
Key to the Species, Based on External Characters 398
Key to Male Genitalia 400
1. Caeneressa proxima n. sp 400
2. Caeneressa klapperichi n. sp 403
3. Caeneressa pratti (Leech) 405
4. Caeneressa obsoleta (Leech) 406
•">. Caeneressa swinhoei (Leech) 409
6. Caeneressa hoenei n. sp 411
7. Caeneressa dispar n. sp 413
8. Caeneressa zernyi n. sp 41,5
9. Caeneressa ningyuena n. sp 410
10. Caeneressa oenone (Butl.) 417
11. Caeneressa diaphana (Koll.) 418
12. Caeneressa graduata (Hmps.) 42S
13. Caeneressa tienmushana n. sp 430
14. Caeneressa rubrozonata (Pouj.) 432
INDEX 437
INTRODUCTION
ISome years ago, while the author was doing research on Ctenu-
chidae and other Lepidoptera families at the Zoological Collection
of the Bavarian State in Munich, he had an opportunity to study
390 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
not only the materials of that museum but also a very extensive
collection from China gathered in the course of many years by
the indefatigable investigator of the Lepidoptera of this country,
Dr. H. Hone. This collection, now one of the most important
components of the Institute of Zoological Research and Museum
Alexander Koenig in Bonn, gave the author a basis for revision
of several Chinese Ctenuchidae species in which he was especially
interested. Further materials for this revision were found in
collections of the Museum of Comparative Zoology in Cambridge,
the American Museum of Natural History in New York, and the
U. S. National Museum in Washington. The species of the related
Oriental fauna were studied from the collections of the three
last-mentioned museums and the State Museum of Natural His-
tory in Leiden. The necessary knowledge about the type speci-
mens of Ctenuchidae from China, previously described by earlier
authors, was received by the author from the British Museum
(Natural History).
The new genus treated in this paper represents a small group
of species ranging through China and the Oriental Region. These
species have up to the present time been considered as belonging
to the genus Amata F. from which they differ both in the struc-
ture of hind tibiae and in the male genitalia. The female geni-
talia could not be closely studied because the preparation of
these parts involved the complete destruction of the markings
of the abdomen which are very important for taxonomy of the
Caeneressa species.
The present paper may be considered as a complete revision
of the Chinese Caeneressa species, so far as they are known.
Concerning some species ranging also beyond China, it was pos-
sible to add information about their distribution and geographi-
cal variation in other countries.
The author wishes to express his gratitude for the friendly
cooperation of the Direction of the Zoological Collection of the
Bavarian State in Munich (Germany) in the person of Prof.
H. Krieg and the Curator of its Department of Entomology, Dr.
W. Forster, and thus for the author's opportunity to devote most
of his working-hours to research work. Hearty thanks also go
to Dr. H. Hone of the Institute for Zoological Research and
Museum Alexander Koenig in Bonn (Germany) for putting his
rich collection at the author's disposal; Dr. P. J. Darlington, Jr.,
OBRAZTSOV: CHINESE CAENEBESSA 391
of the Museum of Comparative Zoology in Cambridge, Mass., and
Dr. F. H. Rindge of the American Museum of Natural History
in New York for the opportunity to study materials in these
museums; Mr. J. F. Gates Clarke and Mr. W. D. Field of the
U. S. National Museum in Washington, D. C, for a similar op-
portunity with respect to the materials of that museum ; Dr. A.
N. Diakonoff of the State Museum of Natural History in Leiden
(The Netherlands) for sending moths for study; Mr. W. H. T.
Tams of the British Museum (Natural History) in London for
sending photographs of the type specimens of Caeneressa species
in this museum and their genitalia, and for a great deal of work
connected with this important aid; Mr. S. G. Kiriakoff of the
Zoological Laboratories of the University of Ghent (Belgium)
for some information about the above types ; Mr. F. Daniel of
the Zoological Collection of the Bavarian State for his continual
assistance in the interpretation of labels in Dr. Hone 's collection ;
Mrs. F. Tandler in Arlington, Va., for her kind assistance in the
preparation of the English text of the present paper. The author
acknowledges with thanks the support of his work on this paper
by a research grant (1952) of the Research Program on the
U.S.S.B. (East European Fund, Inc.) in New York; this grant
gave him the opportunity to study the materials and the litera-
ture in the museums of the United States.
ABBREVIATION'S
The following abbreviations of the names of collections are used in the
paper :
A.M.N.H., American Museum of Natural History, New York.
B.M., British Museum (Natural History), London.
M.C.Z., Museum of Comparative Zoology at Harvard College, Cambridge,
Mass.
M.K., Institute for Zoological Research and Museum Alexander Koenig
("Zoologisches Forschungsinstitut und Museum Alexander Koenig,
Reichsinstitut"), Bonn, Germany.
M.L., State Museum of Natural History (" Rijksmuseum van Natuurlijke
Historie"), Leiden, The Netherlands.
U.S.N.M., U.S. National Museum, Washington.
Z.C.M., Zoological Collection of the Bavarian State ("Zoologische Samm-
lung des Bayerischen Staates"), Munich, Germany.
392 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
SYSTEMATIC DESCRIPTIONS
CAENEEESSA, new genus
Syntomis (part.) Kollar, 1848, Hiigel's Kaschmir, 4 (2), p. 460; Walker,
1854, List Spec. Lep. Ins. B. M., 1. p. 117; Herrich-Schaffer, 1858,
Samml. neuer oder wenig bekannt. aussereurop. Schmett,, p. 72 ; Felder,
1862, Wien. Ent, Mschr., 6. p. 37; Moore, 1871, Proc. Zool. Soc. London,
p. 244; Butler, 1876, J. Linn. Soc. London, Zool., 12. p. 344; 1877,
Illustr. Het. B. M., 1. p. 17; Moore, 1878, Proc. Zool. Soc. London, p.
845; 1878, Anderson's Ees. W. Yunnan, p. 296; Poujade, 1886, BulL
Soc. Ent. France (6), 6. p. CXVII; Swinhoe (and Cotes), 1887, Cat.
Moths Ind., p. 45; Leech, 1889, Trans. Ent. Soc. London, p. 123;
Hampson, 1892, Fauna Brit. India, Moths, 1. p. 212 ; Swinhoe, 1895,
Trans. Ent. Soc. London, p. 30; Hampson, 1897, J. Bombay N. H. Soc,
11. p. 284; Leech, 1898, The Entom., 31. p. 152; 1898, Trans. Ent. Soc.
London, p. 319; Hampson, 1898, Cat. Lep. Phal., 1. p. 59; 1900, J.
Bombay N. H. Soc, 13, p. 46; Piepers en Snellen, 1904, Tijdschr. v. Ent.,
47, p. 51; Seitz, 1909, Gross-Schm. Erde, 2. p. 38; Matsumura, 1911,
Thousand Ins. Jap., Suppl., 3. p. 69; Zerny, 1912, Wagner's Lep. Cat.,
7. p. 12; Seitz, 1912, Gross-Schm. Erde, 10, p. 67; Van Eecke, 1925,
Zool. Meded. Rijksmus. Leiden, 8, p. 208; Draeseke, 1926, Iris, 40, p.
46; Wileman, 1929, Trans. Ent. Soc. London, 76. p. 420; Matsumura,
1931, 6000 Illustr. Ins. Jap., p. 995; Kawada, 1934, Cat. Ins. Jap., 5.
Lep. Syntomidae, p. 1; Wu, 1938, Cat. Ins. Sin., 4. p. 629; Sonan, 1941,
Trans. N. II. Soc Formosa, 31. p. 95.
Hydrusa (part.) Swinhoe, 1891, Trans. Ent. Soc. London, p. 473; 1892, Cat.
East, and Austral. Lep. Het., 1. p. 50; Kirby, 1892, Synon. Cat. Lep.
Het., 1. p. 902; Hampson, 1892, Fauna Brit. India, Moths, 1. p. 220;
1898, Cat, Lep. Phal., 1. p. 66; Kiriakoff, 1954, Ann. Mus. Congo
Tervuren, in 4°, Zool., 1. p. 431.
Zygaena (part.) Kirby, 1892, Synon. Cat. Lep. Het., 1, p. 89.
Eressa (part.) Hampson, 1892, Fauna Brit. India, Moths, 1. p. 221; Swin-
hoe, 1895, Trans. Ent. Soc. London, p. 32.
Amata (part.) Rothschild, 1910, Novit. Zool., 17. p. 433; 1912, ibid., 19, p.
375; Hampson, 1915, Cat. Lep. Phal., Suppl., 1, (1914), p. 13; Fletcher,
1925, Cat. Ind. Ins., 8. p. 6; Matsumura, 1927, J. Coll. Agr. Hokkaido
Univ., 19. p. 74; Candeze, 1927, Enc Ent. (B), Lepidoptera, 2. p. 74;
Joannis, 1928, Ann. Soc Ent. France, 97, p. 245.
Head rather roughly scaled, the frons usually a little more
smooth. Antennae bipectinate or serrate in the male, serrate or
simple in the female, sometimes simple in both sexes, always
ciliate. Palpi labiales porrect, rather short, roughly scaled,
OBRAZTSOV: CHINESE CAENERESSA 393
with a subacute terminal joint. Proboscis moderately long, weak.
Legs smooth, only the coxae somewhat rougher scaled from the
exterior side ; hind tibiae without middle spurs, with a pair of
terminal ones only. Abdomen smoothly scaled.
Forewing moderately broad, dilated outward; dorsum nearly
two thirds as long as the costa ; termen straight or slightly convex ;
12 veins; Rx to R5 stalked; Mi from upper angle of the middle
cell; M2 and M3 shortly stalked, connate or slightly separate,
from the lower angle of the middle cell; Cuj from well before
the angle of the middle cell ; Cu2 from more or less behind the
middle point of the middle cell ; A2 more or less arched, extends
to the tornus.
Hindwing subovate, shorter than the forewing dorsum ; 5 veins ;
Sc coincident with R and Mi, to the costa; M2 and Cii! connate
or shortly stalked, from the lower angle of the middle cell ; Cu2
from cell near three fourths, remote from Cuj ; A2 to the tornus.
Pattern of the Body and Wings. In their pattern the Caener-
essa species are similar to most other Ctenuchidae genera of the
Eastern Hemisphere. The predominant scaling of the head and
body is black or dark brown, often with a blue, greenish or
violet, silk or metallic reflection. The markings are formed by
white, yellow, orange, or red scaling on the dark ground ; some-
times the dark scaling is completely replaced by these colored
scales or pushed into the background. On the head the colorous
markings may be represented as a patch on the frons, also as
streaks on the cheeks; exceptionally the dark ground of the
head is completely taken over by the colorous scaling. On the
patagia the dark ground is often similarly replaced, or they are
patched with color. The tegulae are usually more or less widely
patched with color, sometimes without any black. On the thorax
colorous markings (streaks and patches) are often present; the
pectus usually with lateral patches. The interior side of the
coxae and some other parts of the legs are often colored, es-
pecially in the males. The pattern of the abdomen is formed of
variously developed transverse segmental bands and girdles, com-
plete or interrupted on the dorsal or ventral surface; longi-
tudinal lines are also sometimes present. The scaling of the body
and its parts is never more than trichromatic, usually it is
bichromatic.
394 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
The wing pattern of Caeneressa is similar to that of Amata F.
and formed by hyaline spots on a dark ground. In this way, the
wing pattern scheme of this latter genus (cf. Obraztsov, 1935,
Ent. Anz., 15, p. 262; 1941, Univ. Kijev., Acta Mus. Zool., 1,
(1939), p. 114) can be used also in Caeneressa. In the forewing
a triangular spot (m2) is in the middle cell. A more or less long
spot (nil + m3 °f the Amata wing pattern scheme) is below
the middle cell, in the interspace between it and the vein A2 ;
an ovate or more or less elongate spot (m4) is in the basal part
of the interspace between the veins R5 and Mx ; two spots (m5
and m6) are in the basal parts of the interspaces between the
veins M2 and Chi].. These last two spots are separated from one
another by the vein M3. Some smaller extra spots are often
present ; they occur near the basal parts of the interspaces over
the vein R5 and below the vein M1; also at the outer angle
formed by the vein Cu2 and the middle cell. The subcostal area
is sometimes hyaline, the supradorsal area pale scaled.
In the hindwing a basal spot is present. It occupies the inter-
space between the middle cell, vein Cu2 and A2. This spot-
usually crosses over the vein A2 and reaches almost to the wing
dorsum. Not infrequently it also crosses over the lower vein
of the middle cell which latter is in this case at least partly
hyaline. The second hindwing spot is a distal one. It occupies
the basal part of the interspace between the veins Cvlx and Cu2
and the middle cell. This spot usually crosses over the vein
Cuj and reaches to the vein M2. In case both spots of the
hindwing are enlarged and confluent, they occupy most of the
surface of the wing, and the hindwing becomes hyaline with
dark borders.
Frequently all wing spots are very enlarged, and the wings
become predominantly hyaline. In this instance, the dark ground
of the forewing is reduced to a discal patch and borders along the
wing margins. These borders are usually dilated at the wing
apex, often also between the veins Cux and Cu2. A dark ray
along the vein M2 usually connects the forewing borders with
the discal patch. In the hindwing these borders are mostly
dentate at the veins Cu! and Cu2. The veins of both wings are
always more or less dark.
OBRAZTSOV : CHINESE CAENERESSA
395
The wing pattern is bifacial and the under surface matches the
upper one. Exceptionally the dark interspaces of the under
wing surface are lightened by yellow, whitish or other scales.
Male Genitalia (Pig. 1). Uncus long, more or less curved;
tegumen simple or with lateral appendages; saccus variously
111%
v mm-,* ■(' 'i^:,
1 m
Fig. 1. Male genitalia of Caeneressa diaphana (Koll.) ; preparation no. 2
(M.C.Z.).
a, lateral view ; b, ventral view.
896 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
long, at least moderate. Valvae elongate, variously shaped, some-
times slightly asymmetrical, in this instance the left valva is
shorter; the upper edge of the valva (costa) mostly indifferenti-
ate ; sacculus more or less thickened, distally rounded, without
a free apical point ; interior side of the valvae connected with
the vallum penis by more or less long processus basales. Vallum
penis membranous, only the fultura inferior sclerotized, vari-
ously shaped. Aedoeagus moderate or long, more or less straight
or slightly curved ; coecum penis moderate or underdeveloped ;
vesica with a cuneus of numerous, little, chitinous spines, or with
well developed elongate cornuti, sometimes with both.
Female Genitalia (Fig. 2). The seventh abdominal segment
with a broad, strongly sclerotized tergite and a narrow, less
sclerotized sternite ; it forms caudad a wide, roundish opening
into which the papillae anales are retracted in the position of
rest. The postsegmental edge of the seventh sternite with a wide
indentation displaced to the left which borders with the ostium
bursae. The bottom of the sinus vaginalis membranous, with a
narrow, arched sclerite cephalad from the ostium bursae, between
it and the postsegmental edge of the seventh sternite. The eighth
segment in form of a narrow ring, with a tergite more sclerotized ;
I he eighth sternite semimembranous and forms a kind of medial
ventral plate slightly widened at the middle. The narrow, lateral
commissurae of both eighth tergite and sternite joined into
rather short apophyses anteriores. The papillae anales soft,
hairy, broad coniform, the apophyses posterioi'es nearly three
limes as long as the apophyses anteriores. Two short, narrow
papillae genitales between the papillae anales.
Corpus bursae round, membranous, with a large, dented, sclero-
tized signum dilated cephalad and constricted and pointed
eaudad. Cervix bursae wide, with a broad lateral appendix
joined to the ductus seminalis. Ductus bursae rather narrower
than the cervix, constricted and being stronger near the wide
ostium bursae.
The above description of the female genitalia was made from
Caeneressa diaphana (Koll.) only, and it is not safe to say that
it may relate to all species of the genus.
Systematic Position. The new genus Caeneressa is closely
related to Eressa Wkr., Trichaeta Swinh. and Amata F., and in
OBRAZTSOV: CHINESE CAENERESSA
397
the system has to be ranked among this generic group of the
Ctenuchidae. From Ercssa it differs chiefly in the veins Mo
and Ciij connate or stalked on the hind-wing; the male genitalia
Kiy. J. Female genitalia of Caeneressa diaphana (Koll.) ; preparation no.
Ot. 0 (M.L.).
398 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
of Caeneressa are similar to those of the multigutta group of
Eressa. The new genus resembles especially Trichaeta, but dif-
fers from it in the absence of the lateral hair-tufts on the abdo-
men, and in genitalia. The features distinguishing Caeneressa
from Amata consist of the absence of the middle spurs of the
hind tibiae and of the male genitalia. The latter in Caeneressa
do not have any developed upper angle of the valva, and the
cornuti are never numerous and ranged in a longitudinal row,
both of which features are so typical for Amata.
Range. Information about the geographical distribution of the
genus Caeneressa is currently very insufficient, in so far as the
systematic position of many non-Palearctic species ranked under
Amata P. is not proved. About the non-Palearctic species of the
new genus it is only known at present that albifrons Moore, actea
Swinh., oenone Btlr., era Swinh. and serrata Hmps. belong to
Caeneressa. In the Palearctic region the genus is represented by
thirteen species, seven of them new.
The range of the genus Caeneressa is restricted to China (with
the north extremity of distribution reaching the southern part
of the province of Shensi), North India, Burma, Indo-China,
Chusan, Formosa, and the Great Sunda Islands. The most widely
distributed species of the genus is diaphana Koll. found in almost
all parts of this range, while the distribution of other species is
very restricted. Except for diaphana, all species known to be
from China are endemics of this fauna. They have been found
in the provinces of Shensi, Szechwan, Kweichow, Yunnan, Anh-
wei, Hunan, Kiangsu, Chekiang, Kiangsi, Fukien, and Kwang-
tung. It is very probable that they may be present also in other
provinces which have been studied only to a very limited extent.
There is evidence that rubrozonata Pouj. and diaphana Koll. are
the most widely distributed Caeneressa species in China, although
it would be premature to deny wide distribution with respect
to the rest of the species of this genus.
Key to the Species,
Based on External Characters
1. Antennae bipeetinate in the male, serrate in the female 2
Antennae serrate in the male, simple in the female, sometimes simple
in both sexes 12
OBRAZTSOV: CHINESE CAENERESSA 399
2. Abdomen with only transverse, yellow or red bands seldom joined
together at the middle line 3
Abdomen with transverse bands and in addition with longitudinal
lateral lines 13
3. No red sealing on any part of the head and body 4
Head, patagia, tegulae, thorax, and abdomen with red sealing on a
blaek ground ningyuena, n. sp.
•4. Frons white or greyish 5
Frons yellow or black 6
5. Patagia blaek; the elongate spot below the middle cell of the forewing
extends farther outward than the spot in the middle cell
proximo,, n. sp.
Patagia yellow; the elongate spot below the middle cell of the forewing
extends no farther outward than the spot in the middle cell
pratti Leech
6. Patagia black 7
Patagia yellow, at least laterally 10
7. Hyaline areas between the forewing veins M2 and Cut extend to the
middle cell 8
Hyaline areas between the above-mentioned veins formed as separate
spots which do not reach to the middle cell 9
8. Thorax with a posterior yellow patch; first abdominal tergite with
lateral yellow patches hoenei, n. sp.
Thorax without a posterior yellow patch; first abdominal tergite
broadly yellow zernyi, n. sp.
9. Frons black ; tegulae yellow with blaek end-hair ; hindwing with a broad
hyaline area obsoleta Leech
Frons diffusely yellow scaled ; tegulae yellow only on shoulders ; hind-
wing with two separate hyaline spots Jdapperichi, n. sp.
10. Some of the yellow abdominal bands narrower at the middle
oenone Btlr.
The yellow abdominal bands not narrower at the middle 11
11. Head yellow; subcostal area of the forewing hyaline . . . .dispar, n. sp.
Head black; subcostal area of the forewing black sivinlioei Leech
12. Abdomen with transverse bands and also with yellow or red longitudinal
lines 13
Abdomen with yellow transverse bands only diapluina Koll.
13. Tegulae yellow or red, at least on shoulders 14
Tegulae blaek graduates, Hmps.
14. Abdomen besides two dorso-lateral yellow lines with a medio-dorsal,
longitudinal, yellow line; tegulae entirely yellow . .tienmushana, n. sp.
Abdomen with two dorso-lateral, longitudinal, yellow or red lines only;
tegulae with some black, at least in the end-hairs . . .rubrozonata Pouj.
400 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Key to Male Genitalia
1. Processus basales of the valvae reach no farther than to the vallum
penis 2
Processus basales of the valvae extend far over the vallum penis
zernyi, n. sp.
2. Uncus dilated distally, shaped like a turkish broadsword . . 3
Uncus not dilated distally 7
.'i. Aedoeagus with a cuneus of numerous, little spines . .4
Aedoeagus with one or two cornuti 5
4. Uncus with a short, pointed tip diaphana Koll.
Uncus with a long, pointed tip oenone Btlr.
5. One cornutus 6
Two cornuti rubrozonata Pouj.
6. The left valva shorter; saccus rather long tienmushana, n. sp.
Both valvae almost equally long; saccus short. graduata Hmps.
7. Saccus broad, rather short 8
Saccus rather narrow, more or less long 10
8. Cornuti large, strong 9
Instead of cornuti, two pairs of longitudinal rows of numerous short
cones proximo,, n. sp.
9. Tips of both valvae curved inward; the distal cornutus very broad at
the base pratti Leech
Tips of valvae straight ; the distal cornutus an almost regular cone . . .
Mappericlti, n. sp.
1 0. Distal part of the valva with two angles 11
Distal part of the valva with an acute point 12
11. The left valva distinctly shorter than the right one; its lower angle
acute hoenei, n. sp.
Both valvae almost equally long; the lower distal angle of the left
valva broad, stout swvnhoei Leech
12. Tegumen simple; the upper edge of the valva with an acute point; one
cornutus obsoleta Leech
Tegumen with lateral appendages ; the upper edge of the valva equally
arched ; three cornuti dispar, n. sp.
1. Caeneressa proxima, new species
PI. 1, figs. 1-3
Male. Antennae bipectinate, black, the apical part of the shaft
white. Head black ; f rons white. Patagia black ; tegulae orange-
yellow (at least on the shoulders), black bordered. Thorax black
1 No material available for ningyuena, new species, described from a female.
OBRAZTSOV: CHINESE CAENERESSA 401
with a narrow, orange-yellow, posterior edge; pectus with two
yellow patches on each side. Legs concolorous with the body or
slightly paler; the interior side of the fore coxae white; fore
tibiae sometimes with Avliite, longitudinal streaks ; tarsi more or
less long whitish at the base. Abdomen black-brown, shot with
greenish or violet ; first tergite orange-yellow ; second to sixth
tergites (inch) usually with incomplete, orange, postsegmental
bands, the fifth segment mostly with such a complete girdle; the
corresponding sternites with complete, orange-yellow bands.
Wings brownish black, with a dull, violet gloss ; spots white-hya-
line. Length of the forewing : 25-29 mm.
In the forewing a rather long, wredge-shaped spot (m2) in the
middle cell; a long spot (m14., ) below it which extends much
farther outward than the distal edge of the spot in the middle
cell; a long spot (m4) between the veins R5 and Mt, with
two longish extra streaks over these veins ; twro much broader,
egg-shaped spots (m5 and m6) between the veins M2 and
Ciij separated from each other by the black vein M3 ; the up-
per of these spots slightly shorter than the lower one ; a more
or less developed, oval extra spot at the base of the vein Cu2, out-
ward from the long basal spot (m1+;;).
The hindwdng with a large basal spot which extends from the
middle cell to the vein A2 and is accompanied by a streak behind
this vein; a distal spot, almost equal in size to the basal one,
more or less separated from it, divided by the black vein Cuj
into two unequal parts.
Female. Similar to the male. Antennae serrate. Fore coxae
entirely black. The orange-yellow band on the sixth abdominal
segment mostly absent. The subcostal area of the forewing whit-
ish hyaline, the supradorsal area sometimes whitish.
Male Genitalia (Fig. 3) . Tegumen elongate, moderately arched ;
uncus long, slightly curved downward ; saccus broad and large.
Valvae almost symmetrical, with a strong, thickened sacculus;
distal edge of the valva dentate; the whole valva equally nar-
rowed toward the rounded tip ; processus basales curved, extend-
ing to the upper part of the vallum penis. Fultura inferior bottle-
shaped. Aedoeagus rather thick, moderately curved downward,
funnel-shaped at the tip; coecum penis rudimentary, broad;
cuneus of numerous short cones forming two pairs of longitudinal
rows.
402
BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
Types. Holotype, male, Lienping, Province Kwangtung, May,
1922, H. Hone (genitalia preparation no. S.050 ; M.K.) ; allotype,
female, Hoengshan, Province Hunan, May 29, 1933, H. Hone
(M.K.) ; paratypes, two males, Lienping, Province Kwangtung,
May, 1922, II. Hone (M.K. and Z.C.M.).
Additional material examined. Two females, Yenping, Prov-
ince Fukien, June 13, 1917 (A.M.N.H.) ; one female, Nanking,
Province Kiangsu, June 15, 1933, H. Hone (Z.C.M.).
Range. Chinese provinces Kwangtung, Fukien, Hunan, and
Kiangsu.
Fig. 3. Male genitalia of Caeneressa proxima, new species ; preparation no.
S.050 (M.K.).
a, lateral view; b, ventral view; c, aedoeagus.
Remarks. Superficially very similar to pratti Leech and klap-
perichi, n. sp., except for a much longer forewing spot (m1+3)
below the middle cell. Moreover, proxima can be distinguished
from pratti by the black color of the patagia and dissimilar color
of the fore coxae. The markings of the abdomen of proxima are
unlike those of klapperichi; also the frons scaling is white, not
yellowish as in this species.
OBRAZTSOV: CHINESE CAENERESSA 403
2. Caeneressa klapperichi, new species
PL 1, figs. 7, 8
Male. Antennae bipectinate, black, the apical three-fourths
of their shafts white. Head black; frons diffusely pale-yellow
scaled. Patagia and tegulae black, the latter with yellow shoulders.
Thorax violet-brown, with a narrow, yellow, posterior edge ; pec-
tus with two yellow patches on each side. Legs concolorous with
the body ; the interior side of the fore coxae whitish yellow ; tarsi
whitish scaled. Abdomen violet-brown; first tergite orange-yel-
low; second to seventh segments (incl.) with complete, orange-
yellow, postsegmental girdles. Wings brownish black with a
coppery gloss; spots white-hyaline. Length of the forewing:
21 mm.
In the forewing a rather short, wedge-shaped spot (m2) in
the middle cell ; an elongate spot (n^ +3) below it extends farther
outward than to the middle of the above spot ; an almost equally
broad, elongate spot (m4) between the veins R3 and M1? with
two much smaller, elongate extra spots over these veins ; two
rather short, egg-shaped spots (m5 and m6) between the veins
M2 and Cux separated from each other by the black vein M3 ;
the upper of these spots slightly shorter than the lower one.
The hindwing with a rather large, roundish basal spot which
extends from the middle cell almost to the dorsum ; a separate
distal spot divided by the black vein Cui into two unequal parts ;
the middle cell whitish scaled.
Female. Similar to the male. Antennae serrate. Yellow of
the tegulae only slightly developed. Abdomen with the first
tergite orange-yellow ; similarly colored, incomplete, postseg-
mental bands on the fourth and fifth segments. Forewing broader
than in the male, all spots larger ; the extra spot located between
the spots m4 and m5 contiguous with both; a little extra spot
above the base of the vein Cu2 ; the subcostal area hyaline. The
middle cell of the hindwing partly hyaline. Length of the fore-
wing : 26 mm.
Male Genitalia (Fig. 4). Tegumen strongly arched; uncus
long, dilated at the middle, strongly curved downward ; saccus
short. Valvae almost symmetrical, with a narrow, slightly thick-
ened sacculus ; distal edge of the valva dentate ; the whole valva
404
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
equally narrowed toward the truncate tip ; processus basales
angularly curved, hardly reaching with their tips to the vallum
penis. Fultura inferior subquadrate. Aedoeagus rather short
and thick, with a coecum penis directed downward ; a large, thick,
claw-shaped cornutus on the tip of the vesica, and a much
shorter one at its middle ; some fine, sclerotized dotting at the
bases of these cornuti and above the second of them.
Types. Holotype, male, Kuatun, Province Fukien, 2300 m.alt.,
June 16, 1938, J. Klapperich (genitalia preparation no. S.046 ;
M.K.) ; allotype, female, the same locality, June 20, 1938, J.
Klapperich (M.K.).
Fig. -i. Male genitalia of Caeneressa Jclapperichi, new species ; preparation
no. S.046 (M.K.).
a, lateral view; b, ventral view; c, aedoeagus.
Range. The species is known from the above locality only.
Remarks. The appearance of this new species is very like that
of pratti Leech and proxima, n. sp. ; klapperichi is especially
closely related to the first of these species. It is likely that
klapperichi may be a geographical subspecies of pratti, but
in view of our scanty knowledge of both at present, it is better
to consider them provisionally as two independent species. Super-
ficially both are distinctive in the coloring of the frons and
patagia, also in a dissimilar pattern of the abdomen. The dis-
tinguishing features in the male genitalia are given in the pratti
OBRAZTSOV : CHINESE CAENERESSA 405
description. As to the distinction between klapperichi and
proximo,, refer to the remarks on this latter species.
3. Caeneressa pratti (Leech, 1889), new combination
PI. 1, figs. 4, 5
Syntomis pratti Leech, 1889, Trans. Ent. Soc. London, p. 123, pi. 9, fig.
3; 1898, ibid., p. 325; Hampson, 1898, Cat. Lep. Phal., 1. p. 64; Seitz,
1909, Gross-Sehm. Erde, 2. p. 40; Zerny, 1912, Wagner's Lep. Cat., 7.
p. 25; Wu, 1938, Cat. Ins. Sin., 4. p. 632. — ORIGINAL DESCRIP
TION: "Allied to Syntomis muirheadii, Feld., to which species it bears
a strong superficial resemblance, but is separated therefrom by having
only two hyaline spots towards base of primaries, and blackish margins
to abdominal fold of secondaries. There is no yellow patch on the
posterior edge of thorax, but one is situated band-like on first segment
of abdomen, and this is followed by five yellow belts in the male and
four in female. These last are interrupted on the back of the female
by a stripe of the blackish ground colour. Antennae strongly pectinated
in the male, a character which at once distinguishes it from male
S. muirheadii. Expanse, $ 47 mm., $ 56 mm." (Leech, 1889).
Zygaena pratti Kirby, 1892, Synon. Cat. Lep. Het., L p. 95.
Male. Antennae bipectinate, black, the apical half of the shaft
white. Head black ; f rons white. Patagia and tegulae yellow, the
latter bordered with black. Thorax black ; pectus with two yellow
patches on each side. Legs concolorous with the body or slightly
paler ; the interior side of the fore coxae yellowish white, the base
of the tarsi whitish. Abdomen brownish black; the entire first
tergite orange-yellow ; second segment with orange-yellow, lateral
patches on the tergite and a similarly colored, complete, postseg-
mental band on the sternite; third to sixth segments (incl.) with
complete, orange-yellow, postsegmental girdles, paler on the
ventral surface. Wings brownish black, spots white-hyaline.
Length of the fore wing : 23 mm.
In the forewing a rather short, wedge-shaped spot (m2) at
the end of the middle cell; an elongate spot (m1+3) below it
reaches about to a point on a level with the middle of the middle
cell spot; a rather narrow, elongate spot (m4) between the
veins R5 and Mx accompanied by two short, hyaline streaks
above and beneath; two shorter but broader, egg-shaped spots
(m5 and m6) between the veins M2 and Cui separated from
406 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
each other by the black vein M3 ; the upper of these spots is
slightly longer than the lower one.
The hindwing with a large basal spot which begins in the
middle cell and extends almost to the dorsum; a smaller distal
spot separated from the basal spot by a transverse black bar and
divided by the black vein Cut into two unequal parts.
Female. Similar to the male. Antennae serrate. Frons greyish
white. Patagia black, yellowish at sides. First tergite of the
abdomen orange-yellow, second to fifth tergites (incl.) with
similarly colored, broad lateral patches; the corresponding ster-
nites with paler yellow postsegmental bands. The subcostal
area of the forewing whitish hyaline. Length of the forewing:
26-28 mm.
Male Genitalia (PI. 1, fig. 6). Like those of klapperichi, n. sp.,
but the valvae tips strongly curved inward. Fultura inferior more
elongate. The distal cornutus narrower but more dilated at the
base ; the proximal cornutus slightly longer than in klapperichi.
Types. Holotype, male, and allotype, female, Kiukiang, Prov-
ince Kiangsi, June, 1887 A. E. Pratt (genitalia preparation of
the holotype no. 221; B.M.).
Additional material examined. One female. Province Kiangsi,
June 15 (A.M.N.H.).
Range. Chinese province of Kiangsi.
Remarks. Very similar to both preceding species whose dis-
tinguishing features are discussed above. The resemblance of
pratti to diaphana Koll. ssp. muirheadii Fldr. with which this
species has been compared by Leech (1889) is very remote and
neither can be mistaken for the other. The hyaline wing spots
in muirheadii occupy a larger surface, the supplementary ele-
ments of the markings are more developed, the abdominal
girdles more numerous. In addition to these differences and
those in the male genitalia, the antennae of muirheadii are serrate
in the male and simple in the female.
4. Caeneressa obsoleta (Leech, 1898), new status
and combination
PI. 2, figs. 7-9
Syntomis swinhoei ab. obsoleta Leech, 1898, The Entoni., 31. p. 152. — ORIG-
INAL DESCRIPTION: "In this form the upper hyaline spot of the
OBRAZTSOV: CHINESE CAENERESSA 407
subapical trio is absent, and also the spot between the interno-niedian
bar and the two submarginal spots; the border of secondaries is
broader. Expanse, 34 millim." (Leech, 1898).
Syntomis aotca ab. obsolcta Zerny, 1912, Wagner's Lep. Cat., 7, p. 19;
Seitz, 1913, Gross-Schm. Erde, 10. p. 74.
Male. Antennae bipectinate, black, one-fourth white-tipped.
Head and patagia entirely black ; tegulae yellow with black end-
hairs. Thorax black with a large, yellow, posterior patch ; pectus
with a faint-yellow patch on each side. Legs black, the interior
surface of the coxae yellow. Abdomen violet-black; first tergite
yellow, at the middle broadly interrupted by black ; yellow, post-
segmental bands on second to seventh segments (incl.), enlarged
medio-dorsally and ventro-laterally, sometimes absent on fifth
and sixth sternites. Wings black, spots white-hyaline. Length of
the forewing: 17-19 mm.
In the forewing a long, wedge-shaped spot (m2) in the middle
cell; below it a long, more or less broad, slightly arched spot
(nijls) which extends from near the wing base to about three-
fourths of the dorsum; an elongate-ovate spot (m4) above the
base of the vein M1; accompanied by a small, slightly elongate
extra spot below this vein, and sometimes also by a little dot
above the base of the vein R5 ; two spots (m5 and m6) in the
interspaces of the veins M2 and Culs egg-shaped, dilated toward
outside, the lower of them slightly broader and longer, separated
from each other by the black vein M3 ; sometimes a little, ovate,
extra spot above the base of the vein Cu2 .
The hindwing spots form a common hyaline area bordered by
black ; these borders are broad at the costa, dilated at the apex,
with an obtuse tooth at the vein A2.
Female. Similar to the male but with antennae serrate. The
eyes circumciliated with yellow. The interior surface of the
coxae black. First abdominal tergite with two yellow, lateral
patches and a similarly colored streak at the middle ; second to
sixth segments (incl.) with yellow bands dilated medio-dorsally
and ventro-laterally, the two posterior ones sometimes reduced
or absent.
Male Genitalia (Fig. 5). Tegumen moderately arched; uncus
long, undulate, with a short tip curved downward; saccus long,
narrow. Valvae symmetrical; sacculus large, thickened, with a
round tip ; upper edge of the valva almost straight ; an acute
408
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
projection between it and the irregularly concave distal edge;
valva tip truncate, with a prolonged, acute, distal angle ; proces-
sus basales extend to the vallum penis, their tips dilated. Fultura
inferior elongate. Aedoeagus slender and long, slightly curved
downward at coecum penis and upward at the tip; a single,
grain-shaped cornutus.
Fig. 5. Male genitalia of Caeneressa obsoleta (Leech) ; preparation no.
S.049 (Z.C.M.).
a, lateral view; l>, dorsal view; c, ventral view; d, aedoeagus.
Type. Holotype, female, Ningpo, Province Chekiang, July,
1886 (B.M.).
OBRAZTSOV: CHINESE CAENERESSA 409
Additional material examined. Two males and four females,
Kuatun, Province Fukien, 2300 m.alt., May 19 till June 11, 1938,
J. Klapperich (M.K.) ; one male and one female, the same data
(preparation of the male genitalia no. S.049 ; Z.C.M.).
Range. Chinese provinces of Chekiang and Fukien ; Chusan
Islands.
Variation. The nominate form of the species was described
as having a single extra-spot below the f orewing spot m4 . Most of
the specimens examined from Kuatun are like the type, others
have extra spots above and beneath the spot m4. Sometimes
another extra spot, on the outer side of the f orewing spot mu3
is present. A male specimen from Kuatun (May 20, 1938) lacks
extra spots completely.
Remarks. This species was established by Leech as an aberra-
tion of sivinhoei Leech and considered by later authors as belong-
ing to actea Swinh. but it has nothing to do with either of these
species. It bears rather some likeness to proxima, n. sp., but is
much smaller and with no spot on the frons, has dissimilar
markings of the abdomen and very distinct genitalia.
5. Caeneressa swinhoei (Leech, 1898), renewed status
and new combination
PI. 2, figs. 4, 5
Syntomis sivinhoei Leech, 1898, The Entom., 31, p. 152; 1898, Trans. Ent.
Soc. London, p. 322. — ORIGINAL DESCRIPTION: "Allied to S.
actea, Swinh., but the frons and head are black; the fronts of the
tegulae and the metathorax are marked with yellow. The abdomen of
male has seven yellow bands, and that of the female six. On the
primaries the black along fifth vein between the diseal bar and marginal
border is narrower, as also is the marginal border of secondaries.
Expanse, $ 35 millim., $ 36 millim." (Leech, 1898).
Syntomis actea ssp. 1 Hampson, 1898, Cat. Lep. Phal., 1» p. 64.
Syntomis actea ssp. sivinhoei Hampson, 1898, op. cit., p. 537; Seitz, 1909,
Gross-Schm. Erde, 2. p. 40; 1913, op. cit., 10. p. 74; Zerny, 1912, Wag-
ner's Lep. Cat., 7. p. 19; Wu, 1938, Cat. Ins. Sin., 4, p. 629.
Amata actea swinhoei Fletcher, 1925, Cat. Ind. Ins., 8, p. 7.
Male. Antennae bipectinate, black, presumably white tipped
(in the holotype the antennae tips are broken). Head entirely
black. Patagia black, yellow laterally ; tegulae yellow with black
end-hairs. Thorax black with a large, yellow, posterior patch;
410 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
pectus with yellow, lateral patches. Abdomen black ; first tergite
with yellow lateral patches; second to seventh segments (incl.)
with yellow postsegmental bands. Wings hyaline with veins and
borders black, the latter enlarged at the apex. Forewing, more-
over, with a black, subquadrate discal spot and a black ray
along the vein M2 from the discal spot to the black wing borders ;
these latter with a truncate tooth between the veins Cuj and
Cu2 ; subcostal and supradorsal areas black. Hindwing with
the subcostal area and the middle cell black. Length of the fore-
wing : 16 mm.
Female. Similar to the male but antennae serrate, black, white
tipped. Abdomen with the first tergite patched not only laterally
but also with presegmental and postsegmental 3rellow patches at
the middle; all yellow abdominal bands dilated at the middle;
seventh segment without any band. Length of the forewing:
17 mm.
Male Genitalia (PI. 2, fig. 6). Like those of hocnei (cf. below),
but differing from them as follows : Tegumen broader ; uncus
with a more curved tip. Both valvae almost equally long ; the
lower distal angle of the left valva stout, the upper angle directed
more upward ; in the right valva the upper distal angle more
acute.
Types. Holotype, male, Mupin, Province Szechwan, June,
Kricheldorff (preparation of genitalia no. 223; B.M.) ; allotype,
female, Chiatingfu, Province Szechwan, July, A. E. Pratt (B.M.).
Range. Chinese province Szechwan.
Remarks. The acquaintance of the author with this species
is based on photographs of the type specimens in the British
Museum and the male genitalia of the holotype ; certain charac-
ters of the markings of those specimens were verified by Mr.
S. G. Kiriakoff at the author's request.
This species was considered by Hampson (1898) as a sub-
species of actea Swinh., while in point of fact swinhoei differs
from Caeneressa actea (Swinh.) (PI. 2, figs. 1-3) both super-
ficially and in the male genitalia. The frons of actea is yellow
in the female ; the eighth abdominal segment of the male (seventh
of the female) is yellow; in swinhoei it is black. The male geni-
talia of actea are very typical : uncus deeply undulate, gibbous
before a rather narrow and long tip; saccus rather short; the
right valva with the lower distal angle pointed and the upper
OBRAZTSOV: CHINESE CAENERESSA 411
angle broadly rounded, underdeveloped; the left valva much
narrowed distally, with an almost straight, sharply pointed lower
distal angle. Aedoeagus of actea is shorter and thicker than in
swinhoei.
For a discussion of distinguishing features of very similar
hoenei, dispar and zernyi, refer below to the descriptions of these
species.
6. Caeneressa hoenei, new species
PI. 1, figs. 9, 10
Male. Antennae bipeetinate, black, the apical part of the shaft
white. Head black ; f rons yellow. Patagia black ; tegulae yellow
with black end-hairs. Thorax black with a posterior yellow patch ;
pectus with two yellow patches on each side. Legs concolorous
with the body; the interior side of the coxae entirely yellow.
Abdomen black ; first tergite with lateral yellow patches ; yellow
postsegmental bands (dilated on the ventral side) on the follow-
ing six segments ; the tip of the abdomen black with bluish-violet
gloss.
Wings hyaline with black veins and narrow (at the apex
dilated) black borders. In the forewing a black, subrectangular
discal patch ; the vein M2 connecting this patch with the wing
border black scaled ; a broad, truncate tooth on the interior side
of the black wing border between the veins Cu! and Cu2 ; sub-
costal and supradorsal areas of the forewing black. Hindwing
with very narrow black borders slightly dilated at the apex.
Reverse of both wings with a strong yellowish scaling along the
costa and the dorsum, on interior edges of the black wing borders,
partly also along the veins. Length of the forewing : 15-19 mm.
Female. Similar to the male. Antennae serrate. Head en-
tirely black. Coxae of all legs yellow streaked. Yellow bands,
narrower on the ventral side, on second to sixth abdominal seg-
ments (incl.).
Male Genitalia ( Fig. 6 ) . Tegumen elongate, moderately arched ;
uncus long, equally curved; saccus long and rather narrow.
Valvae asymmetrical, the right one longer; upper and distal
edges not differentiated from one another; the distal part of
the valva subrectangular, with a truncate-concave edge and
412
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upper and lower angles both acute and longer on the right valva ;
sacculus long and wide ; processus basales narrow, dilated at
tips. Vallum penis with two lateral dentate plates joined to the
tips of processus basales. Fultura inferior rounded, inversely
heart-shaped. Aedoeagus slender and very long, slightly curved
downward at the coecum penis and upward at the tip ; a single,
thorn-shaped cornutus.
Types. Holotype, male, Tapaishan in Tsinling, Province Shensi,
1700 m.alt., July 7, 1936 ; allotype, female, of the same locality
and date; three male paratypes taken July 7 to 10, 1936, H.
Fig. 6. Male genitalia of Caeneressa hoenei, new species; preparation no.
S.047 (Z.C.M.).
a, lateral view; b, ventral view; c, aedoeagus.
Hone (M.K.). A further male paratype from the same locality
(preparation of genitalia no. S.047; Z.C.M.).
Range. The species is known from the above locality only.
Remarks. From swinhoei Leech to which the new species is
closely related, it differs in having black patagia and a yellow
irons in the male. C. hoenei also resembles actea Swinh., dispar
n. sp. and zernyi n. sp. but in actea the frons is black in the
male, yellow in the female, and the patagia and the anal abdom-
inal segments are yellow. Both sexes of dispar have entirely yel-
low head and patagia, and the yellow abdominal bands are almost
OBRAZTSOV : CHINESE CAENERESSA 413
joined together; zernyi, in which the female is unknown, has an
entirely black head. The genitalia of all these species are unlike
those of hoenei.
7. Caeneressa dispar, new species
PI. 4, figs. 1, 2
Male. Antennae bipectinate, dark brown, the two apical thirds
of their shafts yellowish. Head, patagia, and tegulae yellow.
Thorax brownish black with a posterior yellow patch ; pectus with
two yellow patches on each side. Legs brownish, diffusely yellow
scaled. Abdomen black ; first tergite with a yellow rectangle,
black patched in the middle ; second to seventh segments ( incl. )
with postsegmental yellow bands joined at the middle line, each
of the tergites consequently with two dorsolateral black patches ;
eighth tergite black postsegmentally with yellow hairs; sternites
whose scaling is very damaged may presumably be entirely yel-
low.
Wings hyaline, veins and narrow borders (dilated at the apex)
brownish black. Forewing, moreover, with a brownish-black
discal patch; vein M2 rather more blackish scaled, supradorsal
area black ; subcostal area with longitudinal hyaline streak. Costa
of the hindwing broad black ; upper part of the middle cell
hyaline. Length of the forewing : 16 mm.
Female. Similar to the male from which it differs as follows :
Antennae serrate, with short, yellowish-white tips. Head black;
frons, cheeks and vertex diffusely yellow scaled. Legs black.
Abdomen with bands as in the male but on first to sixth tergites
(incl.) only; the joining of bands not so clear; anal segments
and all sternites black. The black pigmentation of the body
more intensive and the yellow markings more orange. Black
markings more dilated in both wings, and the black forewing
borders with a distinct, broad tooth at the vein Cu2. Length of
the forewing : 18-19 mm.
Male Genitalia (Fig. 7). Tegumen with two lateral appendages
curved upward; uncus rather long, moderately curved; saccus
rather long. Valvae short, almost symmetrical ; sacculus narrow ;
the terminal part of the valva much narrower than the basal
part; the upper distal angle slightly acute; processus basales
414
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
straight, extending to the vallum penis. Fultura inferior elongate-
ovate, with a concave upper edge. Aedoeagus comparatively
large, slightly curved ; three thorn-like cornuti.
Types. Holotype, male, Kuatun (27°40' N. and 117 °40' E.),
Province Fukien, 2300 m. alt., May 19, 1938, J. Klapperich (geni-
talia preparation no. S.048; M.K.) ; allotype, female, the same
locality, May 6, 1938, J. Klapperich (M.K.) ; paratype, female,
April 25, 1938, the same locality and collector (Z.C.M.).
Fig. 7. Male genitalia of Caeneressa dispar, new species ; preparation no.
S.048 (M.K.).
a, lateral view; b, ventral view; c, aedoeagns.
Additional material examined. Female, Shanghai, Province
Chekiang, H. Hone (M.K.).
Range. Chinese provinces Fukien and Chekiang.
Remarks. The similarity of both sexes in certain characters and
the fact that the moths were found in the same locality within a
comparatively short time period, argues in favor of considering
them conspecific. If this is not the case, the female may be con-
sidered as a new species because it is unlike any other known
species.
The female specimen from Shanghai has an abdominal pattern
like that in the male holotype; in the rest of its characters it
OBRAZTSOV: CHINESE CAENERESSA 415
does not differ from the Kuatun females except that the black
wing markings are rather more developed and some of the hyaline
areas are more spot-like.
Some similarity exists between dispar and actea Swinh., swin-
hoei Leech, and zernyi n. sp., but all these species have their
abdominal bands free, not joined. The male genitalia and some
other characters in the above species are unlike those of dispar.
The male of dispar recalls slightly the Formosan Amata kara-
pinensis (Strd.) but the latter is not a Caeneressa species.
8. Caeneressa zernyi, new species
PL 4, fig. 3
Male. Antennae bipectinate, black, the apical third of their
shafts white. Head, patagia, and thorax black; tegulae yellow
with black end-hairs. Legs brown. Abdomen black with the first
tergite orange patched (the scaling of the remaining abdominal
segments is damaged, and only some orange scales indicate thai
bands were originally present ) .
Wings hyaline with veins and borders black. Forewing with
a subquadrate, black discal patch ; a black ray along the vein
M2 joins the discal patch with the dilated apical border; a
truncate interior tooth of the black wing border between the
veins Cuj and Clio ; subcostal and supradorsal areas black. Hind-
wing with black borders dilated at the apex; costa and the middle
cell black. Length of the forewing: 16 mm.
Male Genitalia (Fig. 8). Tegumen scarcely developed; uncus
very long, curved, dilated at the base and before the narrow,
rounded tip; (the saecus is missing). Valvae symmetrical; sac-
culus well developed, broad, rather Hat ; the upper edge of the
valva almost straight to the distal angle ; the apical part of the
valva narrow and elongate ; processus basales much longer than
the vallum penis, curved, arranged caudad. Fultura inferior
irregularly shaped, narrow in the upper part and dilated in the
lower. Aedoeagus slender, moderately long, slightly curved, with
a moderate coecum penis; a single, thorn-shaped cornutus.
Type. Monotype, male, Shinchow near Canton, Province
Ivwangtung (genitalia preparation no. S.008; Z.C.M.).
Remarks. This species was identified by Dr. H. Zerny (Vienna)
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BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
as new but was not described. It is superficially similar to actea
Swinh., swinhoei Leech and dispar n. sp. but' differs from them
in the black patagia. From hoenei n. sp. it differs in having an
entirely black head. The peculiar processus basales and the
uncus of zernyi are unlike those of the remaining known Caener-
essa species.
Fig. 8. Male genitalia of Caeneressa zernyi, new species ; preparation no.
S.008 (Z.C.M.).
a, dorsal view; b, ventral view; c, lateral view; d, aedoeagus.
9. Caeneressa ningyuena, new species
PI. 4, fig. 4
Female. Antennae deeply serrate, black, one-fourth white
tipped. Head red ; a narrow streak between the eyes and the
mouth parts, black. Patagia entirely red ; tegulae red with
OBRAZTSOV : CHINESE CAENERESSA 417
brownish-black end-hairs. Thorax black with a posterior red
patch ; pectus with two red patches on each side. Legs entirely
black. Abdomen black with six broad, red, postsegmental bands
on first to sixth tergites (incl.).
Wings hyaline with black veins and borders. Forewing with
the borders broadly dilated at apex and forming a broad, in-
terior tooth at the vein Cu2 ; a broad, black discal spot sending a
ray along the vein M2 to the border; subcostal and supradorsal
areas rather diffusely black scaled. Hindwing borders narrow,
dilated only at the apex ; costa and the greatest part of the middle
cell black. Length of the forewing 15 mm.
Type. Monotype, female, mountains near Ningyuenfu, Prov-
ince Szechwan (Z.C.M.).
Remarks. Because of its red pigmentation, ningyuena may
be compared with rubrozonata Pouj. to which it has no other
similarity. The female of rubrozonata has simple antennae ; only
its frons, not the whole head, is red ; the red pigmentation on
the tegulae and the pectus is less developed, the thorax is en-
tirely black. Also the postsegmental edge of the seventh abdom-
inal sternite is distinct in both species (Pig. 12). From graduata
Hmps., also red-pigmented, ningyuena can be distinguished by
its broader forewing shape, less developed black wing markings,
and absence of the longitudinal red lines on the abdomen. The
antennae of graduata are simple in the female.
10. Caexeressa oenone (Butl.), new combination
and renewed status
Syntomis diaphana var. ? Walker, 1854, List. Spec. Lep. Ins. B. M., 1. p
126. — ORIGINAL DESCRIPTION: ' « Nigro-viridis, flavo varia; an-
tennae nigrae serrate, apice albae; alae hyaline subluridae, purpureo-
fusco marginatae, anticae purpureo-fusco faseiatae. Blackish green.
Head pale yellow; vertex black. Proboscis tawny. Antennae black,
serrated along the whole length, white above towards the tips. Thorax
with an interrupted yellow band in front, and with a large subtriangular
yellow mark on each side; scutellum and pectus mostly yellow. Wings
hyaline, with a slight lurid tinge; borders, band on the tip of the
discal areolet, and an opposite mark on the hind border purplish brown.
Abdominal segments with more or less interrupted yellow bands. Legs
blackish brown. Length of the body 6 lines: of wings 14 lines."
(Walker, 1854).
418 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Syntomis oenone Butler, 1876, J. Linn. Soc. London, Zool., 12, p. 344 ;
Swinhoe (and Cotes), 1887, Cat. Moths Ind., p. 49. — OKIGINAL DE-
SCRIPTION: "S. diaphana, var. ?, Walker (nee Kollar) is a distinct
species, and may be named S. oenone." (Butler, 1876).
Zygaena vitreata (part.) Kirby, 1892, Synon. Cat. Lep. Het., 1. p. 95.
Syntomis diaphana (part.) Hampson, 1892, Fauna Brit. India, Moths, I.
p. 216; 1898, Cat. Lep. Phal., 1. p. 67; Seitz, 1909, Gross-Schm. Erde,
2, p. 40; Zerny, 1912, Wagner's Lep. Cat,, 7. p. 20.
Antennae shortly bipeetinate in the male, serrate in the female.
In other respects, this species cannot be distinguished from C.
diaphana (Koll.).
Male Genitalia. Uncus rather longer than in diaphana, with
a more prolonged and acute tip ; saccus larger ; valvae somewhat
broader. Fultura inferior almost regular ovate.
Type. Holotype, male, North India (B.M.).
Additional material examined. Five males and five females,
Kooloo, Himalaya, Carleton (preparation of male genitalia no.
1, Obr.; M.C.Z.).
Range. Western Himalaya.
Remarks. Although this species has been not found within
the limits of China, its presence in the Western Himalaya and
a great similarity to diaphana Koll. are reasons for including it
in the present revision. Walker (1854) described the antennae
of oenone as "serrated along the whole length." This is an
evidence that he correctly distinguished between oenone and
diaphana. The later authors neglected this feature and ranked
in none to diaphana. As matter of fact, the antennae of oenone
are shortly bipeetinate in the male, but without a good magnifier
seem serrate ; those of diaphana seem, under the same conditions,
simple, and Hampson (1898) therefore placed this moth among
the species with antennae simple in both sexes.
The series of the oenone moths examined at the Museum of
Comparative Zoology, represent specimens with markings like
those of the nominotypical subspecies of diaphana. It is very
significant that one diaphana female specimen was also caught
in Kooloo.
11. Caeneressa diaphana (Koll.), new combination
The synonymy is given under subspecies.
Male. Antennae serrate, black, apical part of the shaft white
OBRAZTSOV: CHINESE CAENERESSA 419
above. Head black; frons and narrow streaks behind the eyes
white, cream-white, yellow or orange. The ground of the whole
body black, often with a greenish, violet, or bronze reflection.
Patagia from cream-whitish to orange, in the middle usually
divided by black; tegulae concolorous with the patagia, wTith
black end-hairs. Thorax with a broad transversal, whitish, yellow
or orange, posterior patch ; this patch is often divided into two
patches ; the middle part of the thorax sometimes with two longi-
tudinal, concolorous lateral streaks ; pectus with twTo similar
patches on each side. Legs black or dark brown, often with a
bronze or greenish reflection; sometimes the femora and tibiae
with whitish or yellow longitudinal streaks, and the inner
surface of the coxae is of the same color; first joint of the tarsi
sometimes whitish. Abdomen with whitish, yellow or orange
bands on seven segments; at least some of these bands dorsally
narrowed or interrupted at the middle; some of the anterior
bands often paler than the remaining bands ; on the ventral side
the bands are usually somewhat paler; the anal segment black
or yellow.
Wings predominantly hyaline with black veins and borders. In
the forewing these borders are always dilated at the apex, usually
more or less dilated at the vein M2 and between the veins Cuj
and Cu2 ; discocellulars with a more or less broad, black spot ;
along the vein M2 usually a black ray joining the discal spot
with the wing borders. All these black markings are variously
developed, and the hyaline area is sometimes reduced to separate
spots: a long spot (m1+3) below the middle cell extends nearly
from the wing base to the tornus; a wedge-shaped spot (m2)
in the middle cell ; a more or less large extra spot above the base
of the vein Cu2 ; three larger, elongate spots (m4 to m6), between
the veins R5 and Mt and M3 and Cux form (together with two
smaller and narrower extra spots above and beneath the spot
m4) an exterior row of forewing spots. The basal parts of the
forewing veins often yellow. In the hindwing the black borders
dilated at the apex and slightly indented at the vein Cu2 ; the
middle cell locked by a black discal spot. A stronger development
of the black markings may make two spots of the whole hyaline
area of the hindwing ; they are almost separated from each other by
the vein Cu2. Middle cell and dorsum of the hindwing often
whitish or yellowish scaled. Length of the forewing : 15-25 mm.
420 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
Female. Similar to the male. Antennae simple. Abdomen with
six whitish or orange bands. Length of the f orewing : 17-30 mm.
Male Genitalia (Pig. 1). Tegumen elongate, arched; uncus
long, curved, dilated toward the tip and then pointed; in the
dorsal view the uncus is equally narrowed from the base to the
tip; saccus broad, short. Valvae almost symmetrical, or the
left valva is somewhat shorter, both leaf-shaped; sacculus mod-
erately thickened ; distal edge of the valva more or less dentate ;
processus basales extend to the vallum penis. Fultura inferior
rounded. Aedoeagus moderately thickened, slightly curved at the
middle; coecum penis moderate, broadly rounded; cuneus com-
posed of numerous, diffusely arranged, small, sclerotized cones
and a plate on the vesica.
Female Genitalia (Fig. 2). Discussed in the description of the
genus.
Range. From Kashmir and North India through most of China
and Indo-China to the Great Sunda Islands ; Formosa.
Remarks. In the limits of its range, this species is found in
three subspecies. They have no difference in the genitalia and
are linked together by intermediate forms.
11a. Caeneressa diaphana diaphana (Koll.) , new status
PL 3, figs. 1-4
Syntomis diaphana Kollar, 1848, Hiigel's Kaschmir, 4. part 2, p. 460, pi.
19, fig. 7; Walker, 1854, List. Spec. Lep. Ins. B. M., 1. p. 126; Herrich-
Schaffer, 1838, Sarnml. neuer oder wenig bek. aussereurop. Schmett.,
p. 72; Swinhoe (and Cotes), 1887, Cat, Moths Ind., p. 47; Hampson,
1892, Fauna Brit. India, Moths, 1. p. 216; Swinhoe, 1895, Trans. Ent.
Soc. London, p. 31; Hampson, 1898, Cat. Lep. Phal., 1. p. 67; Snellen
en Piepers, 1904, Tijdschr. v. Ent., 47, p. 51, 53; Seitz, 1909, Gross-
Schm. Erde, 2, p. 40; Zerny, 1912, Wagner's Lep. Cat., 7. p. 20; Seitz,
1913, op. tit., 10. p. 74, pi. 9g [fig. 4]. — ORIGINAL DESCRIP-
TION : ' ' Alis diaphanis, marginibus, macula in anticis costali nervisque
nigris; fronte, maculis humeralibus, metathoracis cingulisque abdomi-
nis, medio interruptis, flavis. Expans, alar. 1", 8"' (mas.) — 2", Vfa'"
(femin.)." "Die grosste mir bekannte, sehr ausgezeichnete Art.
Die Pliigel alle glashell, durchsichtig, nur ihre Rander und die Adern
schwarz. Auf den Vorderfliigeln erstreckt sich an der Spitze die
sehwarze Farbung am weitesten naeh innen, dann verbindet beilaufig
in der Mitte ein schwarzer Fleck die beiden Hauptaste der Fliigeladern,
OBRAZTSOV: CHINESE CAENERESSA 421
und entsendet einen sehmalen Streifen nach der Spitze hin; auch vom
Aussenrande, nahe am hinteren Winkel wird der schwarze Saum
breiter. Die Adern sind verhaltnismassig dick. Die schwarze Em-
saumung der Hinterfliigel ist ziemlich gleichformig und das durchsich-
tige Feld nur von drei feinen Adern durchzogen. Die Stirne, beiderseits
ein Schulterfleck, ein in der Mitte unterbrochener Querstreifen am
Hinterriicken gelb, auch der Hinterleib erscheint mit sieben in der
Mitte unterbrochenen, beim Manne ockergelben, beim Weibchen mehr
lichtgelben Eingen. Die Fiihler sind schwarz, gegen die Spitze weiss
bestaubt." (Kollar, 1848).
Syntomis vitreata Herrieh-Schaffer, 1855, Samml. neuer oder wenig bekannt.
aussereurop. Schmett., pi. 50, fig. 267. There exists no description of
vitreata, only a figure has been published. In the text accompanying
the plates, Herrieh-Schaffer considered this name as synonymous ■with
diaphana Koll. to which the figure of vitreata has an undoubted
similarity.
Hydrusa oaiaea Swinhoe, 1891, Trans. Ent. Soc. London, p. 473, pi. 19,
fig. 10; Kirby, 1892, Synon. Cat. Lep. Het., 1, p. 902; Hampson.
1892, Fauna Brit. India, Moths, 1. p. 222; Swinhoe, 1895, Trans. Ent.
Soc. London, p. 32. — ORIGINAL DESCRIPTION: "S2. Palpi
and antennae black, antennae white above towards the tips ; f rons,
head, and body bright ochreous; space between the antennae, a thin
band behind, three longitudinal stripes on thorax, which meet in a band
before and behind, segmental bands on abdomen, and extreme tip, deep
black. Wings mostly hyaline, with black veins and borders. Fore
wings with the costal line black, the band on disco -cellular broadly
black, the black colour on the lower discoidal veinlet and on the first
and second median veinlets thickening towards the irregular marginal
band, some ochreous colour on the veins towards the base and on the
space below the submedian vein. Hind wings with the costa broadly
black, and with a marginal band somewhat as on fore wings. Under
side as above; legs black, streaked with ochreous grey; tarsi for the
greater part whitish. Expanse of wings, l%o in." (Swinhoe, 1891).
NEW SYNONYM.
Hydrusa diaphana Swinhoe, 1892, Cat. East, and Austral. Lep. Het., 1.
p. 51.
Zygaena diaphana Kirby, 1892, Synon. Cat. Lep. Het., 1, p. 95.
Zygaena vitreata Kirby, 1892, op. cit., p. 95.
Syntomis oaiaea Hampson, 1897, J. Bombay N. H. Soc, 11, p. 284; 1898,
Cat, Lep. Phal., 1. p. 67; 1900, J. Bombay N. H. Soc, 13. p. 47; Seitz,
1913, Gross-Schm. Erde, 10. p. 74, pi. 9f [fig. 9].
Syntomis muirhcadi (non Fldr.) Hampson, 1898, Cat. Lep. Phal., 1. pi. 3,
fig. 13; Sonan, 1941, Trans. N. H. Soc Formosa, 31. p. 96.
Syntomis horishana Matsumura, 1911, Thousand Ins. Jap., Suppl., 3, p. 69,
422 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
pi. 35, fig. 19; Wileman, 1929, Trans. Ent. Soc. London, 76. p. 429;
Matsumura, 1931, 6000 Illustr. Ins. Jap., p. 995, fig. — OEIGINAL
DESCRIPTION: "9. Fore wing yellowish, hyaline in certain lights
reflecting blue; costa, outer and hind margin, a spot upon the cross
vein, a longitudinal stripe each upon the veins III and V, as well
as the total veins dark brown; at the hind margin with a yellow stripe.
Hind wing just like the fore wing, veins except II yellowish, costa and
outer margin as well as the vein II dark brown. Body dark brown,
irons, collar, tegulae, mesonotum in the middle, a broad band to each
segment of the abdomen and pygidium yellow. Legs dark brown, tarsi
somewhat paler. Length: 16 mm.; exp. 48 mm." (Matsumura, 1911).
NEW SYNONYM.
Syntomis hoppo Matsumura, 1911, Thousand Ins. Jap., Suppl., 3, p. 70, pi.
35, fig. 20; Wileman, 1929 Trans. Ent. Soc. London, 76. p. 431; Mat-
sumura, 1931, 6000 Illustr. Ins. Jap., p. 995, fig. — ORIGINAL DE-
SCRIPTION: "It differs from S. horishana m. as follows: $. 1.
Hyaline spot of the cell lb somewhat narrower. 2. Veins of the hind
wing dark brown. 3. Frons, collar, tegulae orange yellow. 4. Abdomen
orange yellow, to each segment with a spindle shaped black band, 2
last segments black, shot with blue. Length: 16 mm.; exp. 48 mm."
(Matsumura, 1911). NEW SYNONYM.
Syntomis uajaca Zerny, 1912, Wagner's Lep. Cat., 7, p. 19.
Amata oaiaea Fletcher, 1925, Cat. Ind. Ins., 8. p. 8.
Amata diaphana Fletcher, 1925, op. cit., p. 11; Candeze, 1927, Enc. Ent.,
ser. B, Lepidoptera, 2, p. 74; Joannis, 1928, Ann. Soc. Ent. France,
97, p. 245.
Syntomis muirheadi ab. horidhana Kawada, 1934, Cat. Ins. Jap., 5, Lep.
Syntomidae, p. 2.
Syntomis muirfieadi ab. hoppo Kawada, 1934, loc. cit.
Wings mostly hyaline with the black only on their borders,
discocellulars, and other veins. The hyaline areas separated by
veins and merely by a black ray along the forewing vein Mo.
The interior tooth of the forewing terminal border between the
veins C\i1 and C112 never meets the middle-sized discocellular
patch and just along these veins reaches sometimes to the middle
cell. Patagia yellow ; thorax with or without longitudinal yellow
streaks; yellow abdominal bauds interrupted at the middle, at
least on two basal tergites.
Types. Syntomis diaphana: Holotype, male, and allotype, fe-
male, Masuri, N. W. Himalaya (location of types unknown) ;
Ilydrusa baiaea: holotype, male, Khasia Hills, Assam (B.M.) ;
S. horishana: monotype, female, Horisha, Formosa (Hokkaido
OBRAZTSOV: CHINESE CAENERESSA 423
Imperial University, Sapporo); 8. hoppo: monotype, female,
Hoppo, Formosa (the same collection).
Additional material examined. One female, Kooloo, Himalaya,
Carleton (M.C.Z.) ; one female, Morendro Doonai, Shillong,
Assam, 1936 (M.L.) ; two males (genitalia preparation no. S.002 ;
Z.C.M.) and one female (Zoological Museum of the Kiev State
University), Ningyuenfu, Province Szechwan; one female, Kiu-
huashan, Province Anhwei, September, 1932, G. Liu (M.C.Z.) ;
two females, Chiengmai, Siam, October 26-28, 1920 (A.M.N.H.) ;
three males (A.M.N.H.) and one female (genitalia preparation
no. Ct. 9; M.L.), Java; one male and one female, Tjibodas, Java.
1400-1800 m. alt., November 1-20, December, 1927, H. Burgeff
(Z.C.M.) ; two males and two females, the same locality, April
1-10, 1907 (male genitalia preparation no. 2, Obr. ; M.C.Z.) ; two
males, Mt. Gede, Tjibodas, Java, April, 1909, Bryan and Palmer
(genitalia preparation no. 4519 W.D.F. ; U.S.N.M.) ; one male
and one female, Gedeh, W. Java, 1350 m. alt., 1893 ; 1600 m. alt.,
1887 (M.L.) ; one male, Sindinglaya, W. Java, 1885 (M.L.) ; one
female, Preanger, W. Java, 5000 ft. alt., Sythoff (M.L.) ; one
male, "Java Sea" (M.L.) ; two males (genitalia preparation no.
Ct. 3 ; M.L.) and one female (M.C.Z.), without data.
Range. Kashmir; N. W. Himalaya; Chinese province Szech-
wan ; S. China ; from N. India to Burma ; Indochinese Peninsula ;
Great Sunda Islands; Formosa.
Variation. The chief characters of the subspecies are more or
less stable except for the width of the black wing borders and the
yellow body pigmentation which may vary from specimen to
specimen. All intermediate forms between the more yellow pig-
mented baiaea and the less yellow pigmented diaphana were
found, and there is no necessity to separate these forms under
special names. The same should be said of horishana and hoppo
which are only synonyms for diaphana. The extreme individual
forms are the following two.
ab. melas Wkr., new status (Plate 3, fig. 4)
Syirtomis melas Walker, 1854, List. Spec. Lep. Ins. B. M., 1, p. 133;
Butler, 1877, Illustr. Het. B. M., 1. p. 17, pi. 6, fig. 10; Swinhoe (and
Cotes), 1887, Cat. Moths Ind., p. 49; Seitz, 1913, Gross-Schm. Erde,
10, p. 73, pi. 9g [fig. 5] ; Wileman, 1929, Trans. Ent. Soc. London, 76.
424 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
p. 430, pi. 20, fig. 12. — ORIGINAL DESCRIPTION: "Purpureo-
f usca, albido varia ; proboscide fulva ; palpis antennisque nigris, his
apiee albis ; abdomen albido subinterrupte fasciatum ; alae hyaline,
longae, subluridae, marginalibus venisque infuscatis. Purplish brown.
Head whitish in front, on each side and beneath. Proboscis tawny.
Palpi and antennae black, the latter white above towards the tips.
An interrupted band on the prothorax, four broad stripes on the
mesothorax and scutellum whitish. Wings hyaline, long, with a very
slight lurid tinge, bordered with brown round the margin and along
the veins, especially at the tips and across the tip of the discal areolet
and along the opposite space of the hind border; a whitish streak
along the hind border of the fore wings, and another along the fore
border of the hind wings. Abdomen nearly linear; segments from the
first to the sixth with whitish bands which are partly interrupted
above. Length of the body 10 lines; of the wings 28 lines." (Walker,
1854).
Zygaena melas Kirby, 1892, Synon. Cat. Lep. Het., 1. p. 94.
Syntomis melaena (nom. emend.) Hampson, 1892, Fauna Brit. India.
Moths, 1. p. 216; Swinhoe, 1895, Trans. Ent. Soc. London, p. 31;
Hampson, 1898, Cat. Lep. Phal., 1. p. 96; 1900, J. Bombay N. H. Soc,
13. p. 48; Zerny, 1912, Wagner's Lep. Cat., 7, p. 23; Wu, 1938, Cat. Ins.
Sinens., 4, p. 630. •
Amata melas Fletcher, 1925, Cat. Ind. Ins., 8, p. 17; Joannis, 1928, Ann.
Soc. Ent. France, 97, p. 245.
Amata melaena Candeze, 1927, Enc. Ent., series B, Lepidoptera, 2, p. 75.
Syntomis owstoni subsp. melas (ex err.) Wu, 1938, Cat. Ins. Sinens., 4, p.
631.
Female specimens with the body pigmentation partly whitish
instead of yellow, especially on the frons, patagia, tegulae, on the
interior side of the fore coxae, and on the abdomen. Some of
the abdominal bands are sometimes nevertheless yellow.
Type. Holotype, female, Nepal (B.M.).
ab. andersoni Moore, new status (Plate 3, fig. 3)
Syntomis andersoni Moore, 1871, Proc. Zool. Soc. London, p. 244, pi. 18,
fig. 1; 1878, ibid., p. 845, 857; 1878, Anderson's Res. W. Yunnan, p.
296, pi. 81, fig. 4; Swinhoe (and Cotes), 1887, Cat. Moths Ind., p. 45.
— ORIGINAL DESCRIPTION: "Male and female. Wings hyaline,
veins bluish black ; body black, with orange-yellow bands : fore wing
with the costa and exterior and posterior margins black; space be-
tween the submedian vein and posterior margin pale yellow; a broad
transverse discicellular black quadrate spot, which is recurved out-
OBRAZTSOV: CHINESE CAENERESSA 425
wards: hind wing with the anterior border pale yellow, and having
a small discoidal black spot; apex and exterior margin black;
posterior margin tinged with yellow. Spot on front of head, coxae,
legs above, and band on each segment of abdomen beneath white. Collar
round thorax, tegulae, spots on thorax, and band on each segment of
abdomen above orange-yellow; tip of abdomen in male purplish black,
in female yellowish grey. Proboscis, palpi, antennae, and legs beneath
black, the antennae tipped with white. Expanse, $ l4Ao, 9 1% inch."
(Moore, 1871).
Zygaena andersoni Kirby, 1892, Synon. Cat. Lep. Het., 1. p. 96.
Syntomis melaena Hampson, 1900, J. Bombay N. H. Soc, 13. p. 50.
Amata flavolavata Rothschild, 1910, Novit. Zool., 17. p. 434; 1912, ibid.,
19. p. 375, pi. 3, fig. 24; Hampson, 1915, Cat. Lep. Phal., Suppl., 1.
(1914), p. 33; Fletcher, 1925, Cat. Ind. Ins., 8. p. 14. — ORIGINAL
DESCRIPTION: "9. Nearest A. melaena Wlk., but distinguishable at
once by the last abdominal segment being orange, and not blue-black as
in melaena and melaena andersoni. Frons orange; tegulae and patagia
orange; thorax black, orange at hind edge; antennae entirely black;
abdomen bright orange with five black rings. Forewing hyaline
orange-yellow, costal area between costal and subcostal nervures with
basal three-fifths orange-yellow, area between vein 1 and inner margin
orange, a black patch on discocellulars, apex and outer margin nar-
rowly black, nervures black, veins 4 and 5 stalked. Hindwing hyaline
orange-yellow, outer margin and nervures black. Length of forewing:
•J 7 mm." (Rothschild, 1910). NEW SYNONYM.
Syntomis flavolavata Zerny, 1912, Wagner's Lep. Cat., 7. p. 21; Seitz, 1913,
Gross-Schm. Erde, 10, p. 73.
Black wing markings reduced. The costal hindwing margin
yellowish. Abdominal bands wide and not interrupted at the
middle.
Types. S. andersoni: Holotype, female, Yunnan (B.M.) ; A.
flavolavata: monotype, female, Khasia Hills, Assam (B.M.)
Remarks. There is no doubt that diaphana and baiaea are
conspecific. The author had at his disposal both these forms and
could not find any constant features which would distinguish one
form from the other. The specimens like horishana and hoppo
were found among the populations of ssp. diaphana from the
continental part of China, and Kawada (1934, loc. cit.) was right
in considering both Formosan "species" as forms of diaphana,
which he erroneously called muirheadi. The ab. melas was found
in almost all parts of the ssp. diaphana range; similar female
specimens with the yellow abdominal markings replaced by
426 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
white, the author studied from Ningyuenfu, Morendro Doouai
and Chiengmai, also from Java. In the literature melas was
mentioned also from Himalaya, Nepal (type), Sikkhim, Burma,
and Indochina. This form like ab. andersoni has no geographical
adaptation. A separation of flavolavata from this latter form
was unreasonable: Rothschild (1910) was wrong in describing
the last abdominal segment of andersoni female as blue black.
The remaining characters of flavolavata and andersoni, the
yellow tinge of the hyaline wing membrane included, are com-
mon and may be observed in any population of ssp. diqphana.
lib. Caeneressa diaphana muirheadii (Fldr.), new status
PI. 3, figs. 5-9
Syntomis muirheadii Felder, 1862, Wien. Ent. Mschr., 6, p. 37; Leech, 1889,
Trans. Ent. Soc. London, p. 123. — ORIGINAL DESCRIPTION:
"Alis anticis nigricantibus, maculis tribus vittaeformibus aliisque
quatuor pone diseum hyalinis, posticis, margine costali et externo
exceptis, hyalinis, fronte scapulisque luteis, cingulis abdominalibus
ochraceis. <$$." "Regiones montanas circa Ning-po incolit ista,
S. diaphanae Kollar affinis. Mas maculas hyalinas exteriores alarum
anticarum aliter habet ordinatas quam feniina. In eo macula prima
inter venam subcostalem et discoidalem seperiorem jaceat, in femina
inter ramos ultimos subcostales. Macula supra rami mediani primi
basin minuta est, in femina autem vittaef ormis. " (Felder, 1862).
Zygaena muirheadii Kirby, 1892, Synon. Cat. Lep. Het., 1. p. 95.
Syntomis muitheadi Hampson, 1898, Cat. Lep. Phal., 1. p. 95; Leech, 1898.
Trans. Ent. Soc. London, p. 322; Seitz, 1909, Gross-Schm. Erde, 2.
p. 40, pi. 9g [fig. 3]; Zerny, 1912, Wagner's Lep. Cat., 7. p. 24; Seitz.,
1913, op. cit., 10. p. 70; Draeseke, 1926, Iris, 40. p. 46; Wileman, 1929,
Trans. Ent. Soc. London, 76. p. 421, 429-431, pi. 20, fig. 11; Wu.
1938, Cat. Ins. Sinens., 4. p. 630.
Syntomis muirheadi (ssp. or ab.) aucta (non Leech) Hampson, 1898, Cat.
Lep. Phal., 1. p. 95; Seitz, 1909, Gross-Schm. Erde, 2. p. 40; 1913, op.
cit., 10. p. 70; Wu, 1938, Cat. Ins. Sinens., 4. p. 630.
Amata muirheadi Fletcher, 1925, Cat. Ind. Ins., 8. p. 17.
Wings with black markings more distributed and hyaline areas
forming spots on the black ground. The spot edges, the forewing
subcostal area, and the middle cell of the hindwing, often with
a taint of yellow scales. Orange-yellow abdominal bands mostly
interrupted or considerably narrowed dorsally. Thorax usually
with orange-yellow longitudinal streaks well developed.
OBRAZTSOV: CHINESE CAENERESSA 427
Types. Allotype, female, Ningpo, Province Chekiang (Tring
Museum; ef. Wileman, 1929); holotype, male, is apparently
missing.
Additional material examined. Three males and three females,
Suifu, Province Szechwan, April 8, 1922, November, D. C.
Graham (U.S.N.M.) ; four males, Shinkaisi, Omeishan, Province
Szechwan, 4400 ft. alt., August, D. C. Graham (U.S.N.M.) ; one
female, Chengfu, Province Szechwan, June, D. C. Graham
(U.S.N.M.) ; one female, Chungking, Province Szechwan, Sep-
tember, 1941 (M.K.) ; one female, Tungjen, Province Kweiehow,
September 8, 1928, C. B. Wahl (A.M.N.H.) ; one male, Nang-
king, Province Kiangsu, June 15, 1933, H. Hone (M.K.) ; one
male, Lungtan near Nangking, Province Kiangsu, June 3, 1933,
II. Hone (M.K.) ; one male, East Tienmushan, Province Cheki-
ang, 1500 m. alt., June 13, 1931, H. Hone (M.K.) ; one male and
one female, the same locality, May 25, 1931, H. Hone (Z.C.M.) ;
three males and one female, West Tienmushan, Province Cheki-
ang, 1600 m. alt., May 29, September 2-3, 1932, H. Hone (M.K.) ;
one male, Mokanshan, Province Chekiang, May 31, 1931, H. Hone
(M.K.) ; four females, Yenping, Province Fukien, June 28, Au-
gust 8, and September 9, 1917 (A.M.N.H.) ; one female, Foochow,
Province Fukien (U.S.N.M.) ; one male and two females, Shaowu,
Province Fukien, 500 m. alt., May 9 till 24, August, 1937, J. Klap-
perich (M.K.) ; one male, the same locality (genitalia prepara-
tion no. S.036 ; Z.C.M.) ; one male and three females, Kwangtseh,
Province Fukien, August 28 till September 7, 1937, J. Klapper-
ich (M.K.) ; one male, the same locality (Z.C.M.).
Range. Chinese provinces Szechwan, Kweiehow, Kiangsu,
Chekiang, and Fukien.
Variation. This subspecies varies chiefly in size. The wing
pattern is more or less constant; only the varied shape and size
of the forewing extra spot above the vein Cu2 affects the width
of the black wing border. The yellow abdominal girdles are
mostly well developed; only in one female from Chungking is
there no girdle on the second tergite.
Remarks. Hampson (1898) and some other authors ranked
Syntomis ancta Leech (The Entom., 31, 1898, p. 153) to muir-
headii, but by mistake. The present author had an opportunity
to study the male genitalia of the type of aucta on the basis of
a photograph received from the British Museum. This examina-
428 BULLETIN :. MUSEUM OF COMPARATIVE ZOOLOGY
tion showed that aucta was an independent species belonging to
the genus Amata F.
lie. Caeneressa diaphana hunanensis, new subspecies
PI. 3, figs. 10, 11
From the preceding subspecies this differs by a slight develop-
ment of the yellow pigmentation of the patagia and thorax.
Patagia black with some yellowish scales at the anterior edge ;
longitudinal yellow streaks of the thorax obsolescent.
Types. Holotype, male (May 30), allotype, female (June 6),
and two paratypes (M.K.) ; two further paratypes, one male
and one female (Z.C.M.). The series originates from Hoeng-
shan, Province Hunan, 900 m. alt., May 28 till June 6, September
4, 1933, H. Hone.
Range. Known from the above locality only.
Remarks. Although the distinction between the new sub-
species and ssp. muirheadii seems minimal, it is constant in all
the specimens examined from Hoengshan. On the other hand,
not any specimen of muirheadii, of the large series examined, has
the yellow of the patagia so much reduced as in hunanensis.
This fact gives the ground for considering the Hunan specimens
a separate geographical form.
12. Caeneressa graduata (Hmps.), new combination
PI. 4, figs. 16, 17
Syntomis graduata Hampson, 1898, Cat. Lep. Phal., 1. p. 67, pi. 2, fig. 28 ;
Seitz, 1909, Gross-Schm. Erde, 2. p. 40, pi. 9f [fig. 4]; Zerny, 1912,
Wagner's Lep. Cat., 7. p. 22; Wu, 1938, Cat. Ins. Sinens., 4, p. 629.
-OBIGINAL DESCEIPTION: "$. Head, thorax, and abdomen
black; tegulae and a patch on metathorax crimson; abdomen with
subdorsal and lateral crimson streaks, the subdorsal streaks conjoined
by segmental lines. Fore wing with hyaline streak above vein 1 from
near base to near termen, and a series of spots between veins 2 and 5
and 6 and 8, diminishing in size towards costa. Hind wing with
hyaline patches below the cell and above vein 2, and spot between
veins 3 and 5, which are stalked." " $ with broad dorsal crimson
fascia on abdomen." (Hampson, 1898).
Male. Antennae biserrate, black, short white tipped. Head
OBRAZTSOV: CHINESE CAENEKKSSA
429
black. Patagia red ; tegulae black. Thorax black with a posterior
red patch ; pectus with two red patches on each side. Legs
brownish, slightly paler than the body. Abdomen blackish
brown ; postsegmental, crimson bands on first to seventh tergites
(inch) ; a crimson mediodorsal and two lateral, longitudinal
lines; sternites entirely black. Length of the forewing: 14-15
mm.
Wings black brownish, rather diffusely scaled, with hyaline
spots. Forewing with a three-fourths long, slightly arcuate spot
(m1+3) below the middle cell; a short, wedge-shaped, some-
times diffusely blackish scaled spot (m2) in the middle cell; an
exterior series of spots consisting of more or less short rec-
tangular spot (m4) between the veins Mx and R5 accompanied
by an extra spot above and a little, inconstant extra spot below;
two inward pointed spots (m5 and m6) between the veins M2
and Cu1} and an extra spot between the veins Cux and Cu2
bordering on the lower edge of the middle cell ; these exterior
spots are separated from each other by veins only. Hindwing
black bordered, with an elongate hyaline area below the middle
cell.
Fig. 9. Male genitalia of Caeneressa graduata (Hmps.) ; preparation no.
S.064 (M.K.).
a, lateral view ; b, ventral view ; c, aedoeagus.
Female. Similar to the male. Antennae simple. Abdomen
with a broad, dorsal, longitudinal, crimson fascia.
Male Genitalia ( Fig. 9 ) . Tegumen rather narrow ; uncus mod-
erately long, dilated and pointed distally; saccus broad, rather
short, with a narrow tip. Yarvae symmetrical; sacculus short;
430 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
costa slightly arched between the upper and distal edges; tip of
the valva acute, directed slightly upward ; processus basales
extending only to the vallum penis. Fultura inferior ovate.
Aedoeagus moderately sized, curved upward ; one single, long
cornutus, thickened at the base, very narrowed distally.
Type. Ilolotype, male, Kiangnan (B.M.).
Additional material examined. One male, Lungtan near Nang-
king, Province Kiangsu, May 8, 1933, H. Hone (genitalia prepa-
ration no. S.064; M.K.).
Range. Chinese province Kiangsu.
Remarks. In the abdominal pattern, this species is similar to
C. tienmushana, n. sp., but it has crimson bands instead of the
yellow ones in the latter species, and otherwise shaped geni-
talia. From all other Caeneressa species, gr adnata differs in its
peculiar venation having the hindwing veins M2 and CUi
stalked.
13. Caeneressa tienmushana, new species
PI. 4, fig. 5
Male. Antennae slightly serrate, black, short yellowish tipped.
Head black; frons and palpi yellow. Patagia yellow, at the
middle blackish ; tegulae entirely yellow. Thorax blackish brown ;
pectus with two yellow patches on each side. Legs brownish,
diffusely yellowish scaled, especially on the tarsi ; interior side
of the coxae yellow. Abdomen brownish black ; first to seventh
tergites (inch) with narrow, postsegmental, yellow bands; a
yellow mediodorsal and two longitudinal, lateral lines; the
corresponding sternites in their greatest part yellow ; tip brown-
ish black with sparse, postsegmental, yellow hairs. Length of
the forewing: 14 mm.
Wings hyaline with brownish black veins and borders dilated
at the apex. Forewing moreover with a broad, brownish black,
discal spot and a black ray along the vein M2 connecting this
spot with the wing borders ; a border tooth along the vein Cu2 ;
all the hyaline areas more or less spot-shaped ; subcostal area
yellow. Hindwing narrower, brownish black bordered; dorsum
yellow scaled ; middle cell and costa brownish black, the latter
slightly tinged with yellow. The reverse of both wings with a
diffuse, yellow sealing along the costa and dorsum.
OBRAZTSOV : CHINESE CAENERESSA
431
Male Genitalia (Fig. 10). Teguinen narrow, moderately
curved, dilated distally and with an acute tip ; saccus moder-
ately long, almost straight. Valvae leaf-shaped, the left valva
considerably shorter ; tip pointed ; sacculus rather narrow,
slightly thickened, with an obtuse tip; processus basales short,
slightly curved at the tips, extending only to the vallum penis.
Fultura inferior elongate-ovate. Aedoeagus rather long, slender.
Fig. 10. Male genitalia of Caeneressa tienmushana, new species; prepara-
tion no. S.051 (M.K.).
a, lateral view; b, dorsal view; c, ventral view; d, aedoeagus.
slightly curved upward; a single, thorn-like cornutus, and a
fine, distal cuneus of numerous, little, chitinous cones.
Type. Monotype, male, West Tienmushan, Province Chekiang,
June 9, 1935, II. Hone (genitalia preparation no. S.051; M.K.).
Range. Known from the above locality only.
432
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Remarks. Allied to rubrozonata Pouj., especially to its ab.
leucoma Leech, from which the new species differs in its anten-
nae, head color, tegulae, legs, and abdomen, in more enlarged
hyaline wing areas, and in genitalia. The wing pattern is some-
what similar to that of ob sol eta Leech but the forewing spots
are less far apart.
14. Caeneressa rubrozonata (Pouj.), new combination
The synonymy is given under subspecies.
Male. Antennae slightly serrate or simple, black, short white
tipped. Head black ; f rons white. Patagia black, sometimes shot
with reddish or yellowish, especially at sides, seldom entirely red
or yellow ; tegulae red or yellow with black hair-tufts on the
Fig. 11. Male genitalia of Caeneressa rubrozonata (Pouj.) ; preparation
no. S.052 (Z.C.M.).
a, lateral view; b, ventral view; c, aedoeagus.
tips, or black with colored shoulders. Thorax black ; pectus uni-
colorous, or with a pale reddish or yellowish patch on each side.
Legs black; the interior side of the coxae white, sometimes also
the femora and tibiae white or whitish streaked. Abdomen black ;
first tergite and the further six segments with postsegmental,
red or yellow girdles; they are more or less broad, complete or
interrupted ventrally, jointed between themselves laterally;
those on the second and third tergites often narrower or absent,
that on the seventh tergite also absent sometimes. Wings black
OBRAZTSOV: CHINESE CAENEBESSA
433
with white, hyaline spots. Length of the forewing: 11-16 mm.
Forewing with five ground spots : an elongate-cuneiform spot
(m2) in the middle cell, usually as broad as this latter; a long
spot (m1+3) below it, reaching from the wing base almost to the
tornus, or the basal part of this spot (mj) absent; a spot (m4)
in the area between the veins R5 and Mx mostly shorter than that
(m6) between the veins M3 and Cuj ; a spot (m5) between the
veins Mo and M3 often the smallest in the exterior spots series ;
sometimes more or less developed extra spots above the vein
Cu2 and at both sides of the spot m4. Hindwing with a more
or less large spot placed right at the dorsum and separated from
it by a narrow black border ; occasionally this spot reaches
across the vein Cui.
Female. Similar to the male. Antennae simple. Frons red-
dish, yellowish, grey or black. Legs entirely black. Girdle on
the seventh tergite always absent.
Fig. 12. Postsegmental edge of the seventh abdominal sternite of two
Caeneressa species; a, C. ningyuena, new species; b, c, C. rubrozonata
(Pouj.).
Male Genitalia (Fig. 11). Like those in graduata Hmps., with
symmetrical valvae. Aedoeagus equally broad, slightly curved
upward, with a small coecum penis; two elongate, sharp cornuti.
Female Genitalia (Fig. 12). The postsegmental edge of the
seventh abdominal sternite more or less concave.
434 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
Range. Chinese provinces Szeehwan, Chekiang, and Fukien.
Variation. Individual variation of the wing and abdomen
markings, and of the body color and head scaling. Two geo-
graphical subspecies.
Remarks. Hampson (1898) wrote on this species : "both wings
with veins 4 and 5 shortly stalked." As a matter of fact, this
character is inconstant, and the two veins mentioned are often
only coincident.
14a. Caeneressa rubrozonata rubrozonata (Pouj.), new status
PI. 4, figs. 6-9
Syntomis rubrosonata Poujade, 1886, Bull. Soc. Ent. France, ser. 6, 6. p.
CXVII; Leech, 1898, Trans. Ent. Soc. London, p. 324; Hampson,
1898, Cat. Lep. Phal., 1. p. 85; Seitz, 1909, Gross-Schm. Erde, 2. p.
40; Zerny, 1912, Wagner's Lep. Cat., 7, p. 25; Draeseke, 1926, Iris..
40. p. 46. — ORIGINAL DESCRIPTION: "Envergure: $, 30 mill.;
9, 32 mill. Ailes hyalines, avec les taches, les nervures et les bords
noirs. Les superieures sont tres allongees et assez pointues; la bordure
externe s'elargit a l'apex environ du quart de l'aile, oil elle est reliee
par une bande etroite a une tache presque carree qui limite la cellule
et touche au bord costal; cette bordure s'elargit encore en une tache
presque carree, plus large que la prec£dente, entre les deux derniers
rameaux de la nervure mediane. Ailes inferieures tres petites, lanceo-
lees, n'egalant pas en longueur la moitie des superieures, ayant l'apex
jusqu'au tiers de l'aile environ et le bord costal noirs. TSte, thorax
et abdomen noirs, se dernier ayant les arceaux superieures bordes de
rouge vermilion ; antennes noires, blanches a 1 'extremite. Le male seul
a le front blanc ainsi que la poitrine, le devant des hanches et des
lignes sur les euisses. " (Poujade, 1886).
Zygaena rubrizonata Kirby, 1892, Synon. Cat. Lep. Het., 1, p. 93.
The black of the wings considerably reduced ; the hyaline
spots of the forewing exterior spots series placed near the middle
cell. In the male the abdomen not girdled ventrally.
Types. Mupin, Province Szeehwan (probably in the Musee
d'Histoire Naturelle, Paris).
Additional material examined. Four males and one female,
West of Yachow, June, 2000 to 7000 ft. alt., D. C. Graham
(U.S.N.M.) ; one male, Shinkaishi, Omeishan, Province Szeeh-
wan, 4400 ft, alt, July, D. C. Graham (U.S.N.M.).
Range. Chinese province Szeehwan.
OBRAZTSOV: CHINESE CAENERESSA 485
Variation. The nominotypical form has red body markings.
Tliat with yellow markings was described as an aberration.
ab. leucoma Leech (Plate 4, figs. 6, 8, 9)
Syntomis consequa Leech, 1898, The Entom., 31. p. 153; 1898, Trans. Ent.
Soc. London, p. 324; Hampson, 1898, Cat. Lep. Phal., I. p. 96, pi. 5,
fig. 2 ; Seitz, 1909, Gross-Schm. Erde, 2. p. 40, pi. 9e [fig. 1] ; Zerny,
1912, Wagner's Lep. Cat., 7, p. 20. — ORIGINAL DESCRIPTION:
' ' Female. Wings almost exactly identical with those of S. rubrozonata,
but the frous is greyish, the collar is yellow, and there are six yellow
bands on abdomen, the first of which is broad. Expanse, 28 millim."
(Leech, 1898). NEW SYNONYM.
Syntomis leucoma Leech, 1898, The Entom., 31, p. 154; 1898, Trans. Ent.
Soc, Loudon, p. 324. — ORIGINAL DESCRIPTION: "Frons, tegulae,
and fore tibiae white; thorax and abdomen black, the latter with
seven yellow bands, the last two of which are almost confluent. Prim-
aries hyaline, venation black; there is a black spot at outer extremity
of cell, and this is united by a bar with the broad apical portion of
the black outer marginal border; the latter is toothed at veins 2 and 3;
there is a curved black streak along inner margin. Secondaries hya-
line, with black outer border which is broadest at apex. Expanse, 28
millim." (Leech, 1898).
Syntomis rubrosonata (part.) Hampson, 1898, Cat. Lep. Phal., 1, pi. 3, fig. 7 ;
Seitz, 1909, Gross-Schm. Erde, 2, pi. 9f [fig. 5].
Syntomis rubrozonata ab. leucoma Hampson, 1898, Cat. Lep. Phal., 1, p. 85;
Seitz, 1909, Gross-Schm. Erde, 2, p. 40; Zerny, 1912, Wagner's Lep.
Cat., 7, p. 25.
With yellow body markings instead of red ones.
Types. Syntomis leucoma: Monotype, male, Omeishan, Prov-
ince Szechwan, 3620 ft. alt., May and June, 1890 (B.M.) ; 8. con-
sequa: monotype, female. Mupin, Province Szechwan, June,
Kricheldorff (B.M.).
Remarks. There is no doubt that leucoma and consequa are
different sexes of the same form although consequa has been
erroneously considered by most authors as an independent
species. I retain for this aberration of rubrozonata the first of
the two simultaneously-established names because it has already
been used in this sense. This aberration is known to me in one
male specimen from the above-mentioned series from Yachow.
436 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
14b. Caeneressa rubrozonata eurymelaena, new subspecies
PI. 4, figs. 10-15
The black of the wings distributed; the exterior series of the
hyaline forewing spots remote from the middle cell. The abdo-
men of the male usually girdled also ventrally.
Types. Holotype, male, allotype, female, eight males and
seven females, paratypes, Kuatun, Province Fukien, 2300 m. alt..
May 11 till June 6, July 23 till August 8, 1938, J. Klapperich
(M.K.) ; further paratypes, two males and one female, the same
locality (Z.C.M.).
Additional material examined. One male and two females,
Mokanshan near Hangchow, Province Chekiang, June 9 till 16,
August 7 till 19, 1930, H. Hone (M.K.) ; one male and two
females, the same locality (male genitalia preparation no. S.053 ;
Z.C.M.) ; one male, West Tienmushan, Province Chekiang, 1600
m. alt., June 28, 1932, II. Hone (genitalia preparation no. S.052;
Z.C.M.) ; one male and one female, East Tienmushan, Province
Chekiang, 1500 m. alt., June 7 till 20, 1931, H. Hone (M.K.).
Range. Chinese provinces Chekiang and Fukien.
Variation. As in the preceding subspecies, ab. leucoma is
known here also (one male from West Tienmushan and one
male and one female from East Tienmushan). In six female
specimens (one from Mokanshan and five from Kuatun) the
patagia are entirely red. Sometimes there are no girdles on the
ventral surface (two females from Mokanshan, three males and
nine females from Kuatun). Most of the specimens have extra
spots in the forewings; they are absent in only four specimens
(one male and three females from Mokanshan).
INDEX
actea auct., 409
actea Swinh., 398, 410. 412, 415, 416
albifrons Moore, 398
Amata (part.) auct., 392
Amata P., 394, 396, 398
andersoni Moore, 424, 426
aucta (non Leech) Hmps., 426
aucta Leech (Amata), 427
baiaea Swinh., 421, 425
bajara Zerny, 422
Caeneressa, new genus, 392
consequa Leech, 435
diaphana (part.) auct., 417, 418
diaphana Koll. (species), 395, 396,
397, 398, 399, 400, 418
diaphana Koll. (subspecies), 420
dispar, new species, 399, 400, 411, 412,
413. 416
era Swinh., 398
Eressa (part.) auct., 392
Eressa Wkr., 396
eurymelaena, new subspecies, 436
flavolavata Eothsch., 425, 426
graduata Hmps., 399, 400, 417, 428
433
hoenei, new species, 399, 400, 411. 416
hoppo Mats., 422, 423, 425
horishana Mats., 421, 423, 425
hunanensis, new subspecies, 428
Hydrusa (part.) auct., 392
karapinensis Strd. (Amata), 415
klapperichi, new species, 399, 400, 402,
403. 406
leucoma Leech, 432, 435. 436
melaena Hmps., 424, 425
melas Wkr., 423. 425
muirheadi auct., 421, 425, 426
muirheadii Fldr., 406, 426. 428
multigutta Blanch. (Eressa), 398
ningyuena, new species, 399, 400, 416.
433
obsoleta Leech, 399, 400, 406
oenone Btlr., 398, 399, 400, 417
owstoni (non Eothsch.) Wu, 424
pratti Leech, 399, 400, 402, 404, 405
proxima,new species, 399, 400. 404, 409
rubrizonata Kirby, 434
rubrozonata (non Pouj.) Hmps., 435
rubrozonata Pouj. (species), 398, 399.
400, 417, 432
rubrozonata Pouj. (subspecies), 434
serrata Hmps., 398
swinhoei Leech, 399, 400, 409. 412,
415, 416
Syntomis (part.) auct., 392
tienmushana, new species, 399, 400, 430
Trichaeta Swinh., 396, 398
vitreata HS., 421
vitreata (part.) Kirby, 418
zernyi, new species, 399, 400, 411, 412,
415
Zygaena (part.) Kirby, 392
i Synonyms are italicized,
basic discussion is given.
Figures in bold-face refer to pages on which the
Plate 1
Fig. 1. Caeneressa proximo, new species, holotype, male, Lienping,
Province Kwangtung, May, 1922, H. Hone (M.K.).
Fig. 2. Idem, paratype, male, the same data.
Fig. 3. Idem, allotype, female, Hoengshan, Province Hunan, 900 m. alt..
May 29, 1933, II. Hone (M.K.).
Fig. -1. C. pratti (Leech), holotype, male, Kiukiang, Province Kiangsi,
June, 1887, A. E. Pratt (B.M.).
Fig. 5. Idem, allotype, female, the same data (B.M.).
Fig. 6. Idem, male genitalia of the holotype (B.M.).
Fig. 7. C. Mapperichi, new species, holotype, male, Kuatun, Province
Fukien, 2300 m. alt., June 16, 1938, J. Klapperich (M.K.).
Fig. 8. Idem, allotype, female, the same locality, June 20, 1938, J.
Klapperich (M.K.).
Fig. 9. C. hoenei, new species, holotype, male, Tapaishan in Tsinling,
Province Shensi, 1700 m. alt., July 7, 1936, H. Hone (M.K.).
Fig. 10. Idem, allotype, female, the same data (M.K.).
Published with the permission of the corresponding museums.
Plate 2
P^ig. 1. Caeneressa actea (Swinh.)i iiolotype, male, Khasia Hills, Assam,
Hamilton (B.M.).
Fig. 2. Idem, allotype, female, the same data (B.M.).
Fig. 3. Idem, male genitalia of the Iiolotype (B.M.).
Pig. 4. C. swinhoei (Leech), Iiolotype, male, Mupin. Province Szechwan,
June, Kricheldorff (B.M.).
Fig. 5. Idem, allotype, female, Chiatingfu, Province Szechwan, July,
A. E. Pratt (B.M.).
Fig. li. Idem, male genitalia of the Iiolotype (B.M.).
Fig. 7. ('. obsolcta (Leech), Iiolotype, female, Xingpo, Province Chekiang,
July, 188(i (B.M.).
Fig. 8. Llcm, female, Kuatun, Province Fukien. May 26, 1938, J. Klap-
perich (M.K.).
Fig. 9. hli id, female, the same locality, June 11, 1938, J. Klapperich
(M.K.).
Published with the permission of the corresponding museums.
PLATE 2
Plate 3
Fig. 1. Caeneressa diaplvana diaphana (Koll.), male, Ningyuenfu, Prov-
ince Szechwan (Z.C.M.).
Fig. 2. Idem, female, Tjibodas, Java, 1400-1800 m. alt., November 1-20,
1927, H. Burgeff (Z.C.M.).
Fig. 3. Idem, all. andersoni Moore, holotype, female, Province Yunnan
(B.M.).
Fig. 4. Idem, ah. melas YVkr., holotype, female, Nepal, India (B.M.).
Fig. 5. C. diaphana muirheadii (FldrJ, male, East Tienmushan, Province
Ohekiang, May 25, 1931, H. Hone (M.K.).
Fig. (i. Tdem, male, Nangking, Province Kiangsu, June 15, 1933, H. Hone
(M.K.).
Fig. 7. Idem, male, West Tienmushan, Province Chekiang, September
2, 1932, II. Hone (M.K.).
Fig. 8. Idem, male, the same data (M.K.).
Fig. 9. Idem, female. East Tienmushan, Province Chekiang, May 25,
1931, 11. Hone (Z.C.M.).
Fig. 10. C. diaphana hunan.ensis, new subspecies, holotype, male, Hoeng-
shan, Province Hunan, 900 m. alt., May 30, 1933, H. Hone (M.K.).
Fig. 11. I/I/ in, allotype, female, the same locality, June 1, 1933 (M.K.).
Published with the permission of the corresponding museums.
Plate 4
Fig. 1. Caeneressa dispar, new species, holotype, male, Kuatun, Province
Fukien, 2300 m. alt., May 19, 1938, J. Klapperich (M.K.).
Fig. i'. Idem, allotype, female, the same locality. May 6, 1938, J.
Klapperich (M. K.).
Fig. 3. C. zernyi, new species, monotype, male, Shinchow near Canton.
Province Kwangtung (Z.C.M.).
Fig. 4. ni/ngyuena, new species, monotype, female, mountains near
Ningyuenfu, Province Szechwan (Z.C.M.).
Fig. 5. C. tit nmushana, new species, monotype, male, West Tienmushan,
Province Chekiang, June 9. 1935, H. Hone (M.K.).
Fig. 6. C. rubrozonata rubrozonata (Pouj.) ah. leucoma, Leech, male,
west of V.-ichow, June, 2(K)(l to 7000 ft. alt., D. C. Graham (U.S.N.M.).
Fig. 7. C. rubrozonata rubrozonata (Pouj.), female, the same data
(F.S.N.M.).
Fig. 8. ('. rubrozonata rubrozonta (Pouj.) ab. leucoma Leech, monotype
of Syntomis leucoma Leech, male, Omeishan, Province Szechwan. 3620 ft.
alt., .May and June, 1890 (B.M.).
Fig. 9. Idem, monotype of Syntomis conseqva Leech, female, Mupin,
Province Szechwan, June, Kricheldorff (B.M.).
Fig. 10. C. rubrozonata eurymelaena, new subspecies, ah. leucoma Leech,
male, West Tienmushan, Province Chekiang, 1600 m. alt., June 28, 1932.
II. I Line (M.K.).
Fig. 11. ('. rubrozonata eurymelaena, new subspecies, holotype, male,
Kuatun, Province Fukien, 2300 m. alt., May 19, 1938, J. Klapperich (M.K.).
Fig. 12. Idem, allotype, female, the same locality. May 23, 1938, J.
Klapperich (M.K.).
Fig. 13. till in. paratype, female, the same locality, June 3, 1938, J.
Klapperich ( M.K. ).
Fig. 14. Ill: in, female. Mokanshan, Province Chekiang, August 19, 1930,
11. Hone (M.K.).
Fig. 15. Idem, ah. leucoma Leech, female, Last Tienmushan, 1500 in. alt.,
June 20, 1931, 11. II fine (M.K. .
Fig. 10. C. graduata (Hmps.)j holotype, male, Kiangnan, Province
Kiangsu ( B.M. ).
Fig. 17. Idem, male, Lungtan, Province Kiangsu, May 8, 1933, H. Hone
( M.K. ).
Published with the permission of 1 he corresponding museums.
6
8
9
14
10 r ,.
15
a
16
r
13 | 17
PLATE 4
Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE
Vol. 116, No. 8
A COMPARATIVE MORPHOLOGICAL STUDY
OF THE PROVENTRICULUS OF ANTS
(HYMENOPTERA: FORMICIDAE)
By Thomas Eisner
Biological Laboratories, Harvard University, Cambridge, Mass.
With Twenty-five Plates
CAMBEIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM
July, 1957
Publications Issued by or in Connection
with THE
MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE
Bulletin (octavo) 1863 — The current volume is Vol. 116.
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Vol. 3, no. 35 is current.
Occasional Papers of the Department of Mollusks (octavo) 1945 —
Vol. 2, no. 21 is current.
Proceedings of the New England Zoological Club (octavo) 1899-
1948 — Published in connection with the Museum. Publication terminated
with Vol. 24.
The continuing publications are issued at irregular intervals in numbers
which may be purchased separately. Prices and lists may be obtained on
application to the Director of the Museum of Comparative Zoology,
Cambridge 38, Massachusetts.
Of the Peters "Check List of Birds of the World," volumes 1-3 are out
of print; volumes 4 and 6 may be obtained from the Harvard University
Press; volumes 5 and 7 are sold by the Museum, and future volumes will be
published under Museum auspices.
Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE
Vol. 116, No. 8
A COMPARATIVE MORPHOLOGICAL STUDY
OF THE PROVENTRICULUS OF ANTS
(IIYMBNOPTERA: FORMICIDAE)
By Thomas Eisner
Biological Laboratories, Harvard University, Cambridge, Mass.
With Twenty-five Plates
CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM
July, 1057
Xo. 8 — A Comparative Morphological Study of the
Proventriculus of Ants (Hymenoptera :
Formicidae )
By Thomas Eisner '
CONTENTS
INTRODUCTION 441
ACKNOWLEDGMENTS 445
PART I : MATERIALS AND METHODS 445
Cuticular framework preparations (p. 446) ; histological serial
sections (p. 447); method of illustration (p. 448).
I 'ART II: TYPES OF FORMICID PROVENTRICULI 449
MYRMECIOID COMPLEX 449
Subfamily Myrmeciinae 449
Myrmeeia (p. 449, fig. 4)
Subfamily Pseudomyrmecinae 4,52
Pseudomyrmex (p. 452, fig. 5)
Subfamily Aneuretinae 453
Aneuretus (p. 453, fig. 6)
Subfamily Doliehoderinae 453
HypocUnea (p. 453, fig. 7); Leptomyrmex (p. 454, fig. 9);
Liometopum (p. 455, fig. 8) ; Tapinoma (p. 456, fig. 26) ; Az-
teca (p. 458, fig. 30); Iridomyrmex (p. 459, fig. 32); Frog-
gattella (p. 460); Tumeria (p. 461); Dorymyrmex (p. 461,
fig. 42) ; Forelius (p. 462, fig. 40) ; Conomyrma (p. 463,
fig. 33); Technomyrmex (p. 464, fig. 48).
Subfamily Formicinae 40,3
Asepalous formicine proventriculus 465
Xotoncus (p. 465, fig. 50); Melopharus (p. 469, fig. 52)
Arropyga (p. 469, fig. 51); Anoplolepis (p. 469, fig. 64)
Acantholepis (p. 469, fig. 65) ; Diodontolepis (p. 469)
Myrmoteras (p. 469, fig. 66).
Sepalous formicine proventriculus 470
Camponotus (p. 470, fig. 68) ; Formica (p. 473, fig. 69)
Xotostigma (fig. 76) ; Lasius (fig. 78) ; Prenolepis (fig. 77)
Myrmecocystus (p. 474, fig. 79); Gesomyrmex (fig. 80)
Paratrechina (fig. 81); Cataglyphis (fig. 82); Brachymyrmex
i This study was supported largely by the Lalor Foundation, and in part
by a grant from the U. 8. Public Health Service. It was published by a grant
from the Wetmore Colics Fund, and a grant-in-aid from the Sigma-Xi-RESA
Research Fund.
440 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
(fig. 84); Oecophylla (fig. 83); Gigantiop.s (p. 474); Dcn-
dromyrmex (p. 474) ; Opisthopsis (p. 474) ; Polycrgus (p.
474); Polyraehis (p. 474); Pseudolasius (p. 474).
Proventriculi with incipient sepals 474
Cladomyrrna (p. 474, fig. 85) ; Myrmecorhynchus (p. 474,
lig. 86).
PONEROID COMPLEX 475
Subfamily Ponoiinae 475
Odontomaohus (p. 475, fig. 90) ; Amblyopone (p. 476, fig. 91).
Subfamily Cerapachyinae 476
Phyracaees (p. 476, fig. 92).
Subfamily Myrmicinae 477
Pogonomyrmex (p. 477, fig. 95) ; Hylomyrma (p. 478).
Subfamily Dorylinae 478
Eeiton (p. 478, fig. 93).
THE STOMODAEAL VALVE 478
IDENTITY OF THE SEKRETSCHICHT OF EMERY (1888) . . 479
PART III : DISCUSSION 481
LITERATURE CITED 487
KEY TO ABBREVIATIONS 489
EISNER: ANT PROVENTRICULUS 441
INTRODUCTION
Adult Hymenoptera are distinctly discontinuous in their
feeding habits, and subsist very largely on fluids (Bischoff '27).
The relatively large amount of liquid nutrient gathered during
one of the intensive feeding periods is not passed directly into
the midgut (text fig. 1, m. g.), but is first temporarily retained
within a sac-like dilation of the stomodaeum called the crop
(text fig. 1, cr.). At intervals, controlled amounts of liquid are
released from the crop to the midgut. Since in this way food
is passed gradually into the midgut, thereby avoiding a sudden
dilution of midgut enzymes at feeding, the digestive and ab-
sorptive processes within the midgut proceed at optimal effic-
iency. The organ that regulates the delivery of fluid from crop
to midgut is the proventriculus (text fig. 1, pv.). In adult
Hymenoptera the proventriculus, as usually developed, is no
mere strait between crop and midgut. It consists of a relatively
voluminous, strongly-muscled, bulb (text figs. 2, 3), opening
anteriorly to the crop through a cruciform portal, and poster-
iorly into the midgut through a slender, tubular, stomodaeal
valve.
From the structure of the organ it is clear that this type of
proventriculus represents a distinct adaptation to the fluid
nature of the diet. It is, in fact, nothing but a pump, with
intake and exhaust valves provided respectively by the portal
and the stomodaeal valve. Proventricular activity consists of
rhythmic contractions and expansions of the bulb by means of
which a regulated pumping is accomplished (Bailey '52, Schrei-
ner '52). Small particulate matter, such as pollen grains, present
no obstacle to proventricular pumping. We know, for instance,
that in Prosopis, Vespa, and Bombus, even relatively dense sus-
pensions of pollen grains are effectively transferred to the mid-
gut without proventricular obstruction (Bailey '54). In Apis,
the proventriculus may even, under special circumstances, con-
vey pollen grains to the midgut independently from their liquid
medium (Bailey '52, Schreiner '52) ; but this filtering ability
442 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
is probably secondary to the primary fluid-pumping function of
the organ (Bailey '54).
Among the hymenopterous insects thus far investigated, the
proventriculus is monotonously uniform in structure from fam-
ily to family. This circumstance throws into striking relief the
exceptional group : the ants. Within this single medium-sized
family, the proventriculus shows a greater diversity of structural
types than can be found in any other insect family — greater
even than in most entire orders.
According to the theory of Wheeler ('23), a fundamental
bond of social life among the ants is the exchange of nutrient
through regurgitative feeding. In adaptation to regurgitative
feeding the crop assumes a new and overwhelmingly important
function in the social community. It serves not only as a recep-
tacle for the nutrient reserves of the individual, but also as a
communal reservoir, or, as Forel aptly termed it, a "social
stomach," from which the non-foraging members of the com-
munity may derive their nourishment. Some of the higher ants,
in which regurgitative feeding achieves its highest degree of
specialization, have even developed special storage castes ("re-
pletes") with enormously distended crops and expansible gasters
(e.g. Leptomyrmex, Melophorus, Prenolepis, Myrmecocystus,
Proformica, etc.).
In ants, as in all Hymenoptera, the proventriculus serves pri-
marily as a pump. However, the development of a crop capable
of storing nutrient, not only in amounts far beyond those re-
(luired by the individual forager, but also for prolonged periods
of time, has been accompanied by modifications in the proventric-
ulus allowing that organ to assume the additional function of
effectively damming the posterior outlet of the crop. The ant
proventriculus is thus seen to serve a dual purpose, being re-
sponsible not only to the individual, for whom it regulates the
nutrient supply to the midgut, but also to the community, which
it serves by acting as a passive dam to the "social stomach."
During1 evolutionary refinement of its emergent social function as
EISNER: ANT PROVENTRICL'LUS
44:{
sal. gl
Mssjea*
Fig. 1. Digestive tract of the ant Myrmica rubra (simplified, with
some cephalic structures omitted; after Janet). Abbreviations: ant. int.,
anterior intestine; b. c, buccal cavity; cr., crop; inf. chb., infrabuccal
chamber; m. <j., midgut; m. t., malpighian tubule; oes., oesophagus; pny.,
pharynx; pv., proventrieulus; rec, rectum; sal. dct., salivary duct; sal. ffl.,
salivary gland.
Fig. 2. Proventrieulus of Apis (after Snodgrass, relabelled). Abbrevia-
tions: bl., bulb; cp., cupola; Cr., crop; .¥. G., midgut; pt., portal; qd.,
quadrant.
Fig. 3. Longitudinal section through proventrieulus of Apis (after
Snodgrass, relabelled). Abbreviations: c. m., circular muscles; I. m., longi-
tudinal muscles; 6^. vlv., stomodaeal valve; other labels as in Figure 2.
444 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
a dam, the ant proventriculus has solved its new mechanical
problems by means of a number of different morphological
changes, accounting for the extraordinary diversity in structure
by which the organ has come to express itself in the ants.
The present study has as its chief objectives the analysis of
structure, and the explanation of function, of each of the princi-
pal kinds of proventriculi known to occur among the ants. This
structural analysis draws from, and is resynthesized with a view
to augmenting, the general body of knowledge concerning for-
micid evolution. All three elements of this approach — anatomi-
cal, mechanical, and phylogenetic — have been inextricably in-
volved in each step of the study.
The ant proventriculus has been the subject of considerable
interest in the past ; yet, despite several involved descriptions
of its anatomy that have appeared, its exact function and mode
of action have never been properly understood. Due to its struc-
tural variability, the organ has been used extensively as a taxo-
nomic character in the classification of the Dolichoderinae and
Formieinae, and in phylogenetic speculations concerning these
and other groups. The first accurate anatomical study of the
dolichoderine-formicine proventriculus was that of Forel (1878).
Emery (1888) followed up Forel 's work in a classic comparative
monograph, still the basic reference on this organ. He reviewed
most of the dolichoderine and formicine tribes, as well as the
aberrant myrmicine tribe Cephalotini. Additional concise de-
scriptions of the proventriculus of the Dolichoderinae and For-
micinae were given in 1912 and 1925 in the fascicles of Wyts-
man's Genera Insectorum dealing with these groups. The most
recent work deals only with the aneuretine proventriculus (Wil-
son, Eisner et al. '56) and with the specialized camponotine
type (Eisner and Wilson '52). Wherever necessary, reference is
made to these works throughout this paper.
The suprageneric classification of the family Formicidae fol-
lowed here is that advanced by Brown ('54). The aneuretines
are given subfamilial rank in accordance with Wilson, Eisner
et al. ('56). On the advice of W. L. Brown, Jr. and E. 0.
Wilson, I am avoiding the use of tribal designations, classifica-
tion at this level being artificial and currently under revision.
EISNER: ANT PBOVENTRICULUS 445
ACKNOWLEDGMENTS
1 should like to express my gratitude to Professor P. M. Car-
penter, not only for specific aid in connection with this study,
hut for general guidance, inspiration and kindnesses throughout
my graduate training at Harvard.
.Many thanks are due Dr. W. L. Brown, Jr., who supplied and
identified much of the material, and contributed generously of
his knowledge of ant taxonomy and evolution.
I also wish to thank Dr. E. 0. Wilson for supplying some of
the formicine genera, for help in field work, and for offering
advice during the earlier phases of the study. The section de-
voted to the sepalous formicine proventriculus represents the
results of a study conducted jointly with Wilson in 1952, and
recently published (Eisner and Wilson '52). The contents of
this paper have been incorporated herein in slightly modified
form.
To each of the following persons I am indebted for contribut-
ing histologically preserved material : Dr. C. P. Haskins of the
Carnegie Institution, Washington, D. C. ; Dr. N. Kusnezov of
the Instituto Miguel Lillo, Tucuman, Argentina; and Dr. T. C.
Schneirla of the American Museum of Natural History, New
York.
The help of my wife, Maria, who inked many of the drawings,
is gratefully acknowledged.
Most of this research was completed while I was a Fellow of
the Lalor Foundation of Wilmington, Delaware. Special thanks
are due its director. Dr. C. Lalor Burdick, for his personal in-
terest.
PART I
MATERIALS AND METHODS
It is impossible fully to understand any proventriculus with-
out considering the closely interdependent structure and func-
tion of the two major organic constituents, the cuticular frame-
work and the muscular envelope. Structural data were obtained
principally from two types of microscopical preparations. The
lirst of these consisted of the isolated proventricular framework
divested of all attached cellular portions. The second type of
446 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
preparation consisted of the simple serial sections of the whole
proventricuhis. The functional and phylogenetic considerations
were derived from a careful comparative study of these prepara-
tions and such other incidental data as became available.
The following is a brief outline of the procedure used in the
making of the preparations and a brief account of the method of
illustration employed, with a consideration of its limitations.
Cuticular framework preparations
This method is outlined here in some detail, since it may be of
aid to the systematist who uses the ant proventricuhis as a
taxonomic character.
In order to remove all cellular constituents from the proven-
triculus, the whole ant is immersed in hot (ca. 90°C) 10% aque-
ous KOH for V2 to 1 hour, or in the same solution at room
temperature for 8 to 15 hours. Specimens from either fluid
preservative or dry mounts may be treated in this way.
The whole ant is then transferred through several changes of
distilled water to remove the alkali, and finally to 70% alcohol,
where the proventricuhis is dissected out. To this end, a small
window is first cut in the gaster, through which the crop can
readily be seen as a thin cuticular sac extending backwards from
the petiole; the proventricuhis itself appears as a small terminal
protuberance on the crop. At no time should the proventriculus
be severed from the crop, since the crop provides an ideal hold-
fast by which the proventriculus may be transferred through
the various solutions which follow. Furthermore, the positional
relationship and proportions of the two organs are in themselves
important things for the observer to note. The crop with the
proventriculus is seized with fine forceps, extracted from the
gaster and transferred directly into the stain.
Chlorazol black E (1% in 70% alcohol) proved to be the
most suitable of the stains used (Pantin '48). One to five min-
utes suffices for adequate staining; overstained specimens are
usually, but not always, readily destained in 70% alcohol.
Other cuticular stains such as eosin, acid fuchsin, fast green
and picro-creosote (Kennedy '49) offered no advantages over
chlorazol black E. Acid fuchsin occasionally proved useful as a
counterstain to chlorazol black ; the latter stain has a low affinity
EISNER: ANT PROVENTRICULUS 447
for the cupolar or sepalar region of the proventriculus, while
acid fuehsin has a strong affinity for these regions only, and
was therefore especially useful where it was otherwise impossible
to delimit the exact extent of the portal opening of the pro-
ventriculus.
The specimen is then passed through two changes of 100%
alcohol for about 3 minutes each, and into cedar oil, where it
may be stored indefinitely. If whole mounts are desirable, the
cedar oil is rinsed off with xylol, and the specimen mounted
under a coverslip in balsam or synthetic resin.
In cases in which valuable dried specimens, such as holotypes
or uniques, must be studied, the proventriculus can usually be
removed without materially damaging the ant. To isolate the
proventriculus from such a sample, the specimen is first relaxed
in a suitable fluid, such as Barber's fluid or a mild detergent-
water mixture, and then the crop and proventriculus are ex-
tracted together through an incision in the membrane between
the first and second gastric segments. The location of the in-
cision may, of course, vary with the kind and condition of the
ant, but a careful and commonsense approach adapted to the
individual specimen will usually yield a good proventriculus.
As Forel long ago noted, this operation can be performed so
neatly that the re-moiinted ant is only the better for its inci-
dental bath. Once isolated, the crop and proventriculus can be
treated with cold 10% KOH until free of soft tissues, and then
handled like any other proventriculus.
Histological serial sections
Unfortunately, many of the genera available for this study
were either dried, or had been stored in alcohol for many years,
which rendered them useless for histological study. There was
also available, however, an adequate series of representative
genera properly preserved by histological fixatives (Schaudinn's,
Bourn's, Carnoy's). Some of the material that had been pre-
served in alcohol for relatively short periods of time (about 5
years or less) yielded adequate and, in fact, often remarkably
good histological preparations.
Serial sections were made by routine histological technique.
The proventrieuli were embedded in Tissuemat [}/% hour), sec-
448 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
tioned at 5 to 10 micra, and stained with Ehrlich's hematoxylin
and eosin, or in some eases with Masson's trichrome, Mallory's
triple or Heidenhain's iron-hematoxylin. Both longitudinal and
transverse sections were always made.
Method of Illustration
The drawings of ant proventriculi used throughout this paper
are of two distinct types.
The first of these consists of three-dimensional semi-diagrams
of the whole cuticular framework of the proventriculus. The
camera lucida drawings on which these illustrations are based
were made as follows. With the proventriculus submerged in
cedar oil in a small dish placed under the dissecting microscope,
the fine, drawn-out end of a glass capillary needle was introduced
into the proventriculus through the point of intersection of the
slit-like anterior openings (portal) of the organ. The opposite
end of the needle was held in a universal joint by means of
which the glass needle could be turned and the proventriculus
oriented in the desired way. The hollow glass needle draws in
t lie oil by capillarity, and in so doing becomes almost transparent
and does not hide any of the structural details of the proventric-
ulus. Camera lucida drawings were made with 15x ocular and
6x objective. The histological serial sections were of aid in
filling in some of the more inconspicuous structural detail, not
seen through the dissecting microscope.
In some of those few proventriculi studied in which the cuticu-
lar framework is excessively thin and flaccid, usually losing its
shape in the absence of the supporting musculature (the pon-
erine, eerapachyine, dory line, and myrmicine representatives,
and to some extent also the aneuretines and pseudomyrmecines),
the drawings are in large part reconstructions based on the
serial sections, and are labelled as diagrams.
With the exception of the conspicuous rugae on the outer
margins of the plicae of the bulb, no effort was made in any of
these drawings to represent the finer details of cuticular sculp-
turing. Wherever possible, these details were represented in
the histological drawings discussed below.
For the purpose of clarifying some of the internal intricacies
of the proventriculi, the cuticular framework is occasionally
EISNER: ANT PROVENTRICULUS 44')
represented with certain sections cut away. The depiction of
the cutaways was based on the serial sections.
The second type of illustration consists of semi-diagrams of
selected histological cross sections and longitudinal sections
through the proventriculus, and were made with the aid of a
camera lucida.
Due to the variety of fixatives used, the material served only
for gross histological study, and lacked the uniformity necessary
for comparative cytological observations of any value. The
illustrations are therefore semi-diagrammatic, inasmuch as no
attempt was made to represent intracellular detail. In a limited
number of cases the only histological sections obtained were
oblique, rather than transverse or longitudinal, and the drawings
represent reconstructions from camera lucida outlines of the
oblique sections. Wherever such reconstruction was employed,
the drawings are labelled as diagrams.
The outstanding difficulty encountered in the interpretation
of the histological sections was raised by the outermost longi-
tudinal muscle fibers of the proventriculus (l.m. 3 or similarly
disposed fibers). These fibers usually consist of only a few
scattered strands, which during histological processing may
shrink away from their insertions on the cuticle, thereby be-
coming easily displaced, especially since they often lack the
support of other tissues and lie free in the hemocoel. Tracing
such muscle fibers through successive sections was difficult at
best and sometimes impossible.
PART II
TYPES OF FORMICID PROVENTRICULI
MYRMECIOID COMPLEX
Subfamily MYRMECIINAE
MykMECIA Fabricius
Species examined: M. regidaris* Crawley, M. piliventris F.
Smith, M. nigrocincta F. Smith, .¥. vindex* F. Smith, M. forfi-
cata* (Fabricius).
• An asterisk is used herein to denote those proventriculi which were sec-
tioned histologically. A lack of an asterisk indicates that only cuticular frame-
work preparations were made.
450 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
The proventriculus of Myrmecia is one of the least specialized
among that of ants, and in most major respects it is similar to
that of the few other aculeate hymenopterous families in which
the organ has been described.
The cuticular framework (Fig. 4) consists of a thin- walled,
flask-shaped bulb (bl.), opening anteriorly through a wide,
roughly cruciform portal (pt.). The Avails of the bulb are four
broad, sclerotized, inwardly-curved plates, joined basally and
merging into the slender, tubular stomodaeal valve (st. vlv.).
The plates are separated one from another by relatively narrow
longitudinal strips of thin, flexible cuticle, the pleats or plicae
(pi.). The plates themselves are here termed inter plicaries
(inter pi.). The interplicaries extend free beyond the plicae and
the portal to project into the lumen of the crop as four apically
rounded lobes or quadrants (qd.), collectively constituting the
cupola (cp.). Actually, the cruciform shape of the portal is de-
termined by the presence of the cupolar quadrants. The inner
sclerotized surface of the quadrants is covered with relatively
sparse short pile (Figs. 10, 12).
The bulb is surrounded by a strongly developed layer of
circular muscles (Figs. 10, 13, 14; cm.). With the exception of
the innermost fibers, which insert on the lateral margins of the
plicae, the circular muscles envelop the bulb without attachment
to the cuticle.
The longitudinal muscles are arranged in three distinct groups
(Figs. 10, 12, 13, 14; l.m.l, l.m.2, l.m.3). The most strongly
developed of these (l.m.l) extends over the interplicary cuticle
of the bulb, originating posteriorly near the base of the bulb and
inserting anteriorly on the quadrants of the cupola. A second
group (l.m.2) inserts along the posterior half of the interplicary
plates, pierces the circular muscles, and fans out to become a
part of the longitudinal musculature of the midgut. The third
group (l.m.3) consists of only a few scattered fibers that insert
on the quadrants, and extend posteriorly over the circular
muscles to become continuous with l.m.2.
It proved impossible to determine the exact fate of those
longitudinal muscles that approach the bulb from the crop,
although the preparations suggested that these fibers distribute
themselves among the various intrinsic muscles of the proventric-
ulus (see dotted lines, Fig. 10).
EISNER: ANT PROVENTRICULUS 451
lii the arrangement of its longitudinal muscles, the Myrmecia
proventrieulus is unique : all other ant proventrieuli examined,
with the possible exception of Pseudomyrmex, Aneuretus (see
next below) and some genera of the Poneroid complex, lack the
J.m.2 fibers.
A longitudinal section through the stomodaeal valve and associ-
ated structures (Fig. 11) shows how the extremely thin and flex-
ible cuticular intima of the valve extends far into the lumen of
the midgut and then turns inside out and back upon itself so as
to form a compound tube. A strong muscular sphincter embraces
the valve at the point of emergence from the bulb. Figure 11
is of interest also in that it shows in histological detail the nature
of the foregut-midgut junction. Attention is called to the con-
spicuous columnar cells (a) grouped as a ring around the stomo-
daeal valve at the point where the latter perforates the midgut.
The nature of these cells becomes clear from the work of
Dobrovsky ('51) who has shown that in Apis these cells form a
continuous barrier that separates the foregut from the midgut
during the course of prepupal reorganization. At. this time the
developing stomodaeal valve is held back as an invagination
within the bulb, and it is not until later that the cellular barrier
is pierced by the intrusion of the valve into the midgut. A point
of incidental interest is that the basement membrane (b.m.).
which is strongly eosinophilic and readily traced, forms a con-
tinuous sheet over the foregut and midgut epithelium.
In all other ant proventrieuli, the stomodaeal valve is essen-
tially similar to that of Myrmecia, and except for some minor
(but functionally significant) variation in the degree of develop-
ment of the muscular sphincter (see page 478), no further con-
sideration will be given to the stomodaeal valve in subsequent
proventrieuli.
My interpretation of the function of the Myrmecia proventrie-
ulus is as follows. The circular and longitudinal muscles act as
antagonists to one another in compressing and expanding the
bulb. By the combined action of the various longitudinal fibers,
the interplicary plates of the bulb first spread apart from one
another with the unfolding of the plicae, and then are flattened
out. The quadrants of the cupola separate, and crop fluid is
admitted through the widened portal into the expanded bulb. A
forward flow of liquid from the midgut is prevented by the
452 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
stomodaeal valve, the thin wall of which collapses under the
pressure of the midgut contents. By the contraction of its
circular muscles, the bulb is then compressed and its contents
forced back through the stomodaeal valve into the midgut.
Backflow to the crop is hindered by the tight closure of the
cruciform portal (approximation of the quadrants) and by a
tight folding of the membranous plicae, closed by the circular
fibers along their margins.
In addition to its function as a one-way valve, the stomodaeal
valve undoubtedly serves also as a flexible, self-adjusting link,
which by bending is capable of adjusting to peristalsis and vol-
ume changes of the crop.
The proventriculus of Myrmecia apparently is no better
adapted to serve as a dam for the crop than is that of other
Hymenoptera. It is evident from the structure of the organ that
prolonged damming is possible only if either the cruciform portal
or the stomodaeal valve is maintained tightly occluded under
sustained muscular contraction. In preserved proventriculi of
all Myrmecia species examined, the portal was always seen to be
slightly agape, suggesting that sustained occlusion is main-
tained exclusively at the level of the stomodaeal valve, which in
this genus is provided with a particularly powerful sphincter.
Subfamily PSEUDOMYRMECINAE
PSEUDOMYRMEX Lund
Species examined: P.pallidus* (F.Smith).
In its principal features, the proventriculus of Pseudomyrmex
(Fig. 5) is similar to that of Myrmecia, and it can be safely
assumed that it functions in much the same way. In Pseudomyr-
mex the quadrants of the cupola are shaped to fit together more
closely at the axis, resulting in a more narrow cruciform portal,
and a more compact cupola.
The Pseudomyrmex material available was in such a poor state
of preservation that exact delineation of the longitudinal muscu-
lature was impossible. Although the presence of l.m.l and l.m.3
fibers could be determined with certainty (Figs. 15, 16), there
remains some doubt concerning the presence of l.m.2. A few
fibers, strongly suggestive of l.m.2 were seen in cross sections to
EISNER: ANT PROVENTRICULUS 453
extend through the circular muscles at the level of the posterior
third of the bulb (see l.m.2 ( ?), Fig. 16), but final proof of the
presence of l.m.2 must await further investigation.
Subfamily ANEURETINAE
AneURETUS Emery
Species examined: A. simoni* Emery.
In all of its major features, this proventriculus (Fig. 6), like
that of Pseudomyrmex, conforms closely to that of Myrmecia.
The l.m.l and l.m.3 fibers are fully retained (Fig. 21). Al-
though I found no evidence for the presence of l.m.2 fibers, I
must admit that some uncertainty remains regarding their ab-
sence ; the small size of the proventriculus, and the rather poor
condition of the preparations available, made precise observa-
tions difficult.
In view of the overall structural similarity of this proventricu-
lus with that of Myrmecia and Pseudomyrmex, its function is
probably also much the same.
Subfamily DOLICHODERINAE
HYPOCLINEA Mayr
Species examined: H. piistulata* (Mayr).
The cuticular framework of the proventriculus of Hypoclinea
(Fig. 7), although distinctive in its ovoid bulb and flattened
dome, still conforms to the basic structural plan of Myrmecia,
Pseudomyrmex, and Aneuretus, except that the plicae have be-
come sclerotized toward the base of the bulb.
Histologically, the longitudinal musculature has undergone a
simplification by the loss of l.m.2 (Fig. 17). In Hypoclinea this
could be ascertained with certainty. Only l.m.l and l.m.3 are re-
tained (Figs. 17-20). As will be recalled, the main function of
l.m.2 in Myrmecia is to expand the bulb by pulling outward on
the interplicary plates. With the sclerotized posterior parts of
the plicae, the base of the bulb in Hypoclinea forms a cup of
uninterrupted relatively thick cuticle, capable of elastic ex-
pansion by itself when the circular muscles relax. L.m.2 has
therefore become superfluous and is lost. The l.m.l and l.m.3
4.)4 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
fibers serve mainly to disengage the cupolar quadrants so as to
open the bulbar portal during the intake phase of proventricular
activity, and from their attachment, it can be seen that they
play no major part in the expansion of the bulb itself. Except
for the fact that bulb expansion has become a passive process,
the action of the Hypoclinea proventriculus is probably similar
to that of Myrmecia.
LEPTOMYRMEX Mayr
Species examined: L. pollens* Emery, L. cnemidatus Wheeler.
The proventriculus of Leptomyrmex (Fig. 9) has become
specialized in several major respects. The cupola is character-
istically concave in its anterior aspect. The bulb is relatively
small, and occupies less than the posterior half of the proventric-
ulus. In the anterior half, the interplicary plates are thickened
and pinched inward so as to frame a narrow cruciform tract,
the occlusory tract (oec. tr.), an inward extension of the cruci-
form portal of the cupola. The entire occlusory tract and cupola
are lined with dense, long, pile (Figs. 23, 24).
The walls of the occlusory tract are still articulated laterally
by thin, flexible plicae, but as the occlusory tract opens pos-
teriorly into the bulb, the plicae lose their membranous character
and thicken to become sclerotized and hollow (C-shaped in cross
section), reinforcing the bulb at its four angles.
Histologically, Leptomyrmex resembles Hypoclinea in that
l.m.2 is absent, and only l.m.l and l.m.3 are retained (Figs. 23,
24, 25).
In its pumping action, the proventriculus of Leptomyrmex
resembles that of Hypoclinea inasmuch as the bulb, being later-
ally reinforced by sclerotized plicae, is so constituted as to ex-
pand through its own elasticity. The longitudinal muscles serve
only to widen the occlusory tract (the walls of which are allowed
to separate by the flexibility of the plicae), thereby opening the
pathway by which fluid enters the bulb.
In the acquisition of an occlusory tract we see in Leptomyrmex
the first major specialization serving to improve the damming
potential of the proventriculus. The extensive, relatively nar-
row, and densely pilose occlusory tract would seem capable by
itself of effectively containing to a large extent the posteriorly
EISNER: ANT PROVENTRICULUS 455
directed pressure exerted by the crop contents. Thus, the need
for muscular force in the maintenance of proventricular occlu-
sion has become somewhat reduced, and the efficiency of the
damming action of the proventriculus correspondingly increased.
LlOMETOPUM Mayr
Species examined: L. sp.* (near apiculatum Mayr).
Although superficially Liometopum (Fig. 8) is quite distinct
from Leptomyrmex, its general structural plan betrays its debt
to a stock common to these two genera. Like Leptomyrmex
Liometopum has an extensive occlusory tract, and the bulb itself
is considerably reduced in size. Both the occlusory tract and
cupola bear dense, long pile (Fig. 22). In contrast to that of
Leptomyrmex, the cupola of Liometopum is broad and convexly
reflexed, and is somewhat more rigid in consistency than any
previously discussed. The plicae of Liometopum are, as in Lepto-
myrmex, fully sclerotized at the level of the bulb. Unlike in
Leptomyrmex, the plicae are also somewhat thickened at the level
of the occlusory tract, although they undoubtedly still retain
sufficient flexibility to be readily unfolded when the occlusory
tract is dislodged during bulbar intake. The ready mobility
of the occlusory tract is clearly evidenced by prying with a glass
needle. The muscles (Fig. 22) are arranged as in Leptomyrmex.
In its pumping action, Liometopum cannot differ grossly from
Leptomyrmex. Some improvement might be afforded by the more
rigid reflexed cupola, which acts to assure a passively maintained
intrusion of the anterior proventriculus into the crop and
thereby provides greater freedom of access to the cruciform
portal. In the genera discussed previously, superficially similar
intrusion of proventriculus into crop is also seen, but in these
the texture of the cupola is largely membranous, and its intru-
sion seems to be maintained entirely by the tension of those
longitudinal muscles extending from the crop over the pro-
ventriculus.
In its action as a dam, the Liometopum proventriculus shows
some improvement over that of Leptomyrmex, in that the oc-
clusory tract is provided with somewhat sclerotized and there-
fore spring-like plicae, serving as adjuncts to the circular
muscles in sealing the occlusorv tract.
45(3 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
TAPINOMA Forster
Species examined: T. sessile* (Say).
The proventriculus of Tapinoma (Fig. 26) is much more
specialized in structure than any of the proventriculi discussed
so far. It is a strong, rigid, subglobose structure, consisting of
a well-developed bulb surmounted by a cupola made of thick,
strongly sclerotized cuticle. The face of the cupola is ornamented
with a characteristic, raised, densely pilose reticulum (Figs. 27,
28). This sculpturing extends inward to line the slit-like cruci-
form portal. This portal opens almost immedia'tely into the
bulb, and does not proceed farther inward to form an extensive
occlusory tract as in Leptomyrmex and Liomeiopum. The bulb
itself is characterized by the extreme rigidity and c-shaped
cross sections of the plicae (Figs. 26, 29).
The longitudinal musculature has undergone some new special-
ization (Figs. 27, 29). One group of fibers, consisting of only
a few strands, joins the margins of the cupola to the sides of
the bulb. On the basis of their position inside the circular
muscles, I consider these fibers as homologues of l.m.l of previ-
ous proventriculi ; from these they differ only in their more
peripheral insertion on the cupola. A second group of longi-
tudinal fibers inserts anteriorly on the crop intima, in such a
way as to maintain the posterior fold surrounding the proven-
triculus, and originates posteriorly on the base of the bulb near
the origin of the stomoclaeal valve. This was the first proven-
triculus studied in which the full course of these fibers, including
origin and insertion, could be traced with ease. Possibly they
are homologues of l.m.3 that have shifted their insertion outward
to the crop intima and have taken posterior origin secondarily
on the bulb. This interpretation is reasonable, in view of the
absence of any other obvious l.m.3 homologues in Tapinoma.
There exists, in fact, some evidence that in previously discussed
proventriculi, the outermost components of l.m.3 are in a similar
wise already fold-maintaining fibers, but in these cases origin
and insertion could usually not be made out, and the fibers
seemed to extend from the crop over the proventriculus with-
out attachment to the cuticle. In view of the uncertainty that
remains concerning the identity of these fibers in Tapinoma, and
EISNER: ANT PROVENTRICULUS 457
subsequent dolichoderines having similarly placed fibers, they are
labelled as Z.ro.3(?).
In its function as a dam, Tapinoma shows significant im-
provement over previous proventriculi. In the resting proven-
triculus, the quadrants of the cupola are rigidly maintained in
the adducted position and the portal orifice is held to a narrow
slit by the spring-like reinforcement of the strongly sclerotized
plicae. Here is seen the first instance in which proventricular
damming is probably accomplished without any muscular rein-
forcement whatsoever.
The structural advances in the Tapinoma proventriculus are,
of course, linked closely with functional changes in the pump-
ing action. The restriction of the portal orifice to a narrow slit
tends to handicap the ingress of fluid during bulbar intake, and
it becomes necessary to widen the portal through a disengage-
ment of the cupolar quadrants. Tapinoma lacks an effective
occlusory tract, and only a minimal dislocation of the quad-
rants suffices for the ready access of fluid into the bulb. I
believe, in fact, that a wide disengagement of the quadrants is
not only unnecessary but impossible. The thickness and rigidity
of the cupola and the full sclerotization of the bulbar plicae
(these acting as spring antagonists to the quadrant retractor
muscles (l.m.l)) speak for a very narrow range of mobility of
the quadrants. The narrowness of this range is further demon-
strated by the resistance to displacement of the portal when a
glass needle is inserted. It is interesting to note in this con-
nection that l.m.l inserts on the rim of each quadrant, thereby
achieving optimal leverage.
When the circular muscles contract, the bulb is compressed
and fluid is forced back into the midgut in the usual fashion.
Backflow into the crop is prevented by a maximal compression
of the cruciform portal and a tight engagement of its pilose
lining. During the subsequent relaxation of the circular muscles,
the bulb expands under the intrinsic elasticity of its cuticular
framework. In this expansion, the spring-like plicae play an
important role. Simultaneously, the portal orifice is forced
slightly agape under tension from l.m.l, and crop fluid rushes
into the newly emptied and now expanding bulb.
458 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
AZTECA Forel
Species examined: A. sp* (common ochraceous species from Vera
Cruz, Mexico), A. instabilis (F. Smith).
Although in general shape, constitution of the bulb and ar-
rangement of the musculature, the proventriculus of Azteca
(Fig. 30) seems close to that of Tapinoma, it has certain features
that suggest that it may stand alone as an evolutionary offshoot.
The cupola is unique in that the four quadrants are each
deeply and angularly excised, lending the cupola a characteristic
cruciform shape in frontal view. In section, the strongly scler-
otized cuticle of the cupola is seen to possess an inner zone
composed of closely approximated cuticular hairs which give it
a finely and evenly striated appearance. These hairs extend
inward along the cruciform portal to line a short occlusory tract.
In possessing this tract, Azteca further differs from Tapinoma.
In musculature (Fig. 31) the Azteca proventriculus does not
differ greatly from that of Tapinoma. Emery (1888) claimed
a complete absence of longitudinal muscles in Azteca instabilis
(F. Smith), but he probably just missed them.
The Azteca proventriculus at rest is maintained in the oc-
cluded state by its intrinsic elasticity in much the same way as in
Tapinoma, and it can therefore also serve effectively as a passive
dam to the crop. Its damming action is, in fact, improved over
that of Tapinoma, owing to the presence of the occlusory tract.
Although obvious structural similarities render it likely that
the proventriculus of Tapinoma and Azteca function in much
the same way, it is not clear to me whether, during the active
intake phase of bulb operation in Azteca, there is a full disen-
gagement of the cupolar quadrants, or whether disengagement is
damped as in Tapinoma. In Azteca, the presence of an occlusory
tract may offer sufficient resistance to the inflow of fluids to make
its opening necessary. The angular excision of each quadrant
may confer greater flexibility upon the cupolar portal arrange-
ment, thus facilitating the action of the strong "retractors"
(l.m.l) found in Azteca. Further evidence for mobility of the
occlusory tract is demonstrated by its relatively easy displace-
ment when a glass needle is introduced.
The five genera Forelius, Dorymyrmex, Turneria, Froggat-
tella, and Iridomyrmex are unmistakably related on the basis of
EISNER : ANT PROVENTRICULUS 450
proventricular structure. In this series, the proventriculus
achieves its highest degree of specialization among the Dolicho-
derinae. An understanding of the structural intricacies of these
proventriculi is perhaps best accomplished by considering first
the most complex proventriculi of the series {Iridomyrmex,
Frog gat tella, Turneria) and then their more simple evolu-
tionary relatives (Dorymyrmex, Forelius).
Iridomyrmex Mayr
Species examined: I. detect us* (F. Smith), I. viridiaeneus*
Viehmeyer, /. punctatissimus Emery, I. nitidus Mayr.
The proventriculus of Iridomyrmex (Figs. 32, 44, 45) features
a broad, strongly reflexed cupola with involuted margins, hold-
ing tightly within its hollow a greatly reduced bulb. (Caste
dimorphism is displayed in the relative development of cupola
and bulb ; in the female caste, the bulb is slightly larger, and
barely projects from beneath the cupolar shell.)
The cupola is primarily divided into four quadrants by the
cruciform portal in the usual fashion. A major new development
is the secondary division of the quadrants, each of which is split
radially by a narrow phragma (phr.).
The external cupolar cuticle is covered by dense, fine, short
pile (Figs. 34, 35), extending into the cruciform portal. A simi-
lar pilose vestiture, but slightly longer, was already seen to cover
the cupola of Azteca.
Each arm of the cruciform portal opens into a vestibular cleft
or sinus (Figs. 32, 35; sin.), which in turn communicates with
the mesial lumen of the bulb via a submedian zone of construc-
tion (Figs. 32, 35; a). The zones of construction are secondary
developments within the bulb proper that have arisen in response
to a functional need to be discussed below. The bulb is further
characterized by the subdeltoid cross section of the plicae.
Histologically, the proventriculus is seen to lack all intrinsic
longitudinal muscles (Figs. 34, 35). There remain only those
fibers (l.m.3) that maintain the posterior fold of the crop sur-
rounding the cupola.
An outstanding advance in Iridomyrmex is the complete en-
casement of the bulb within the rigid, reflexed cupola, a circum-
stance that tends to render the bulb motionbound. This handicap
460 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
has been overcome by the development of the cupolar phragmata,
which act as flexible radial hinge-lines, allowing the cupolar seg-
ments to yield inward to the contraction of the bulb. The
powerful development of the circular muscles (Figs. 34, 35)
reflects the added burden imposed on the bulb by the need for
embowing the cupolar segments during bulbar contraction. The
bulbar expansion that follows is activated not only by the intrin-
sic elasticity of the bulb itself, but also by the cupolar segments,
which, after becoming embowed during bulbar contraction,
spring back to their original position and carry along the walls
of the bulb.
The necessary development of radial hinge-lines, however,
brings with it a new difficulty. During bulbar contraction, the
lateral arms of the cruciform portal tend to be forced open by
the contraction of the circular muscles (see Fig. 35) instead
of becoming occluded, as would be the case if the cupolar quad-
rants were not pleated. It therefore has become necessary to
develop a secondary valve seal in the form of the constriction
zones within the bulbar lumen. During bulb contraction, the
four constriction zones close early in each pumping cycle, and
serve to prevent leakage from bulb to crop.
Since lridomyrmex has lost all intrinsic longitudinal muscles,
it is incapable of actively expanding the cruciform portal during
bulb expansion. The need for such expansion is avoided by the
reduction of bulb size (relative to portal orifice) and also because
resistance to inflow is minimized in the absence of an occlusory
tract.
This proventriculus is ideally suited to serve as a passive dam.
The very narrow, densely pilose portal, maintained immobile by
the rigid bracing of the bulbar core within the cupola, acts as an
effective plug guarding the outlet of the crop ; fluid can probably
be forced inward past the portal only under suction pressure
exerted by the bulb.
FeOGGATTELLA Forel
Species examined: F. kirbyi (Lowne).
EISNER: ANT PROVENTRICULUS 461
TlJRNERIA Forel
Species examined: T. sp* (near pacifica Mann, collected by
B. 0. Wilson in New Hebrides, 1954-1955).
In these two genera the proventriculus is identical in all
respects to that of Iridomyrmex. Caste dimorphism is again
manifested in the slightly longer bulb in the female.
DORYMYRMEX Mayr
Species examined: D. ensifer* Forel.
On the basis of the proventriculus, Dorymyrmex (Figs. 42,
43) may be considered close to the direct evolutionary antecedent
of Iridomyrmex.
In Dorymyrmex, unlike Iridomyrmex, the posterior third of
the bulb, constituting the most voluminous and therefore the
contractile portion of the bulb proper, is free and projects from
beneath the cupolar shell. Each cupolar quadrant is, as in
Iridomyrmex, incised by radial hinges or phragmata, but in-
cision is incomplete and stops considerably short of the cupolar
summit. As in Iridomyrmex, the cupola and portal bear short
dense pile, and the plicae of the bulb are subdeltoid in cross
section. There is no trace of intrinsic longitudinal muscles, and
there persist only those fibers (l.m.3) that sustain the circum-
cupolar fold of the crop (Fig. 37, representing a longitudinal
section through the proventriculus of Conomyrma, may be taken
also to represent Dorymyrmex) . Dorymyrmex has not acquired
the secondary valve-seal mechanism provided in the bulb of
Iridomyrmex in the form of submesial zones of constriction.
From a functional standpoint, one can readily explain the
incipient condition of quadrant incision, and the lack of a sec-
ondary valve-seal mechanism in the bulb. Both of these condi-
tions are, in fact, directly attributable to the incompleteness of
the envelopment of bulb by cupola. First of all, the fact that
most of the contractile portion of the bulb projects from be-
neath the cupola, thereby remaining largely unimpeded in its
operation, obviates the need for a complete radial incision oi
the quadrants. Only the rim of the cupola must yield to bulbar
contraction, and consequently only the rim is incised. Secondly,
since the quadrants are only incompletely pleated, it is still
462 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
possible to accomplish a valve-seal at the level of the portal
during bulbar compression, and no secondary valvular mechan-
ism had to be acquired.
In its action as a passive dam, the Dorymyrmex proventric-
ulus presents the same structural advantages as that of Iridomyr-
mex.
FORELIUS Emery
Species examined: F.foetidus* (Buckley).
In its basic plan, the proventriculus of Forelius (Figs. 40, 41)
is just about what would be expected of a precursor member of
the Dorymyrmex-Iridomyrmex line. Forelius displays in in-
cipient form the outstanding feature of this line, namely the
envelopment of the bulb by the cupola; most of the bulb is
free, and only the anterior third is encased within the cupola.
Relative to the cupola, the bulb is much larger in Forelius, than
it is in Dorymyrmex and Iridomyrmex. Forelius, like Dorymyr-
mex, shows a beginning of cupolar pleating: only the margins
of each quadrant are interrupted by phragmata. The subdeltoid
outline of the plical cross section and the short dense evenly-
distributed pile of the cupolar face and portal, characteristic of
both Dorymyrmex and Iridomyrmex, are already fully developed
in Forelius.
Histologically, the picture presented by the longitudinal
muscles is puzzling (Fig. 36). The most conspicuous fibers
(l.m.f) insert on the underside of the quadrants outside the
circular muscles, bend posteriorly around the circumeupolar fold
of the crop, and fan out anteriorly over the crop. These fibers
would seem to represent a newly captured component of the
proventricular musculature, possibly representing crop fibers
that have secondarily inserted on the cupola. A single fiber (see
t, Fig. 36) is shown as a remnant of l.m.l. A question mark has
been used to denote this fiber, since its presence is not of general
occurrence, and I have, in fact, seen it only as a single fiber in
one of the three proventriculi sectioned longitudinally (all were
workers of the same nest series). This variation in the presence
of l.m.l points to Forelius as an evolutionary stage in the
Dorymyrmex-Iridomyrmex line at which l.m.l has become
totally superfluous and is disappearing. Longitudinal fibers,
EISNER: ANT PROVENTRICULUS 463
similar in position to l.m.3 of previous proventriculi are present
in Forelius.
Although its general structural affinities to Dorymyrmex sug-
gest that the Forelius proventriculus functions in much the same
way, there remains to be found a fully satisfactory explanation
of the function of the unique longitudinal muscles (l.m.?) of
Forelius. In view of their insertion on the underside of the
quadrants, their obvious function would appear to be quadrant
disengagement and portal widening. The general rigidity of
the cupola (evidenced again in this genus by resistance to dis-
placement when a glass needle is inserted through the portal)
would, of course, preclude anything but the narrowest margin
of cupolar mobility, and the most that the muscles can accom-
plish is a minimal distention of the portal slits (in much the same
way as in Tapinoma). But why has portal distention become a
necessity in Forelius when it is obviously superfluous in the
basically similar proventriculi of Dorymyrmex and Iridomyr-
mexl The answer lies in the relatively larger size of the Forelius
bulb, the midistended portal of which offers an insufficient intake
orifice during bulbar expansion. One may ask why Forelius did
not retain a fully developed set of l.m.l to accomplish portal dis-
tention instead of acquiring a completely new set of muscles, but
careful examination of Figure 36 shows that l.m.l, due to its
adaxial position, would be ill-suited to accomplish quadrant
retraction even if strongly developed.
The following two genera {Conomyrma and Techno my rmex)
have much in common with the Forelius-Dorymyrmex-Irido-
myrmex line, but certain unique characteristics of their own
suggest their phyletic divergence from this series.
Conomyrma Forel
Species examined: C. thoracica* Santschi
Conomyrma, in common with Forelius, Dorymyrmex and
Iridomyrmex, shows a partial envelopment of the bulb by the
cupola, the degree of involvement being intermediate between that
of Forelius and Dorymyrmex (Figs. 33, 46, 47). The cupolar
cuticle bears short dense pile, and the bulbar plicae are sub-
deltoid in cross section (Figs. 37, 38, 39). L.m.l fibers are
totally lacking, and there are present only those longitudinal
464 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
fibers (l.m.3?) that maintain the circumcupolar fold of the crop.
Conomyrma differs from the Forelius-Dorymyrmex-Iridomyr-
mex series in that it has no phragmata to interrupt the strongly
convex, evenly sclerotized surfaces of the quadrants. The four
convex quadrants, separated by the recessed lines at the arms of
the portal, produce a four-lobecl or somewhat clover-leafed
pattern in frontal view (Fig. 47).
The "yielding lines" in the cupola, needed to allow bulbar
contraction, coincide with the arms of the cruciform portal.
The elastic arch formed by each quadrant confers the necessary
flexibility to allow recession at the "yielding lines" during
bulbar contraction. The placement of the "yielding lines" at
the portal slits also ensures a tight closure of the latter during
contraction of the bulb, and this closure acts as an efficient
valve opposing backflow. Thus, by means of a relatively simple
variation in the cupolar plan, the Conomyrma proventriculus
has avoided the development of phragmata and the ensuing dif-
ficulties that had to be met with in Iridomyrmex by the acquisi-
tion of a secondary valve-seal in the bulb.
TECHNOMYRMEX Mayr
Species examined: T. detorquens* (Walker) (=albipes F.
Smith).
This proventriculus (Figs. 48, 49) is so similar to that of
Conomyrma in the major features of its framework that one is
hound to assume that it functions in identical fashion.
Technomyrmex, like Conomyrma, has acquired arched quad-
rants as a means of conveying flexibility to the cupola ; the arch-
ing is more pronounced than in Conomyrma and affords a dis-
tinguishing character.
Emery (1888) and Forel (1878) have already pointed to
the unique sculpturing manifested by the Technomyrmex cupola.
In T. detorquens this sculpturing takes the form of a reticulum
of hexagonal units (Fig. 49), raised on the underside of the
cupola. My preparations suggest (although not with certainty)
that the hexagonal units delimit the areas of cuticular deposition
of the individual underlying epithelial cells. Emery interprets
the reticulum, not as a surface sculpturing, but as an intra-
cuticular labyrinth ; it is likely that he was led to an erroneous
EISNER : ANT PROVENTRICULUS 465
interpretation by the obliquity of his seetions.
Technomyrmex differs from Conomyrma, Forelius, Dorymyr-
mex, and Iridomyrmex in that the bulbar plicae are not subdel-
toid, but C-shaped in cross section (Fig. 49). This feature, the
unique type of cupolar sculpturing, and also divergent external
characters of the whole ant that have long been recognized by
taxonomists (Brown, W. L., personal communication) point up
the possibility that the form and function of the Technomyrmex
proventriculus may have developed along an independent line.
In this case, the proventriculi of Technomyrmex and Conomyrma
would have to be counted as a striking instance of convergent
evolution.
Subfamily FORMICINAE
Among the formicine genera, the proventriculus falls far short
of developing the heterogeneity of types we have seen to occur
in the Dolichoderinae. In fact, all Formicinae possess either one
or the other of two basic types of proventriculi, with relatively
slight variation, except for the very few known species, belong-
ing to only one or two genera, in which the proventriculus is of
intermediate character. The tAvo main types, w.ith the intermedi-
ates, seem to express a simple evolutionary progression from the
more generalized asepalous type to the advanced sepalous type.
From among many possible examples, the two described here in
detail as representative of asepalous (Notoncus) and sepalous
(Camponotus) types were chosen because of their relatively large
size and because adequately preserved material happened to be
available.
Asepalous Formicine Proventriculus
Notoncus Emery
Species examined: N. ectatommoides* (Forel).
The cuticular framework (Fig. 50) consists of cupola and
bulb, both characterized by extreme sclerotization and rigidity.
The cupola, unlike that of any dolichoderine proventriculus.
is not hollowed out, but consists of a thick solid cuticular cap
roofing the frontal aspect of the bulb. The face of the cupola is
466 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
flattened in Notorious (but domed in many other asepalous formi-
cine genera) and frames a narrow cruciform portal. The portal
continues inward through the cupola in the form of a narrow
occlusory tract, except for the outer extremities of the arms of
the portal, each of which is immediately adjacent to the upper
extension of its respective plicary canal (Figs. 53, 55-58). The
face of the cupola, the cruciform portal, and the occlusory tract
all bear short, densely approximated pile, similar to but longer
than that found in the advanced dolichoderine proventriculi.
The cruciform portal and occlusory tract are rigidly braced by
the massive cupolar quadrants ; it is impossible to pry open the
portal with a glass needle without cracking the bulbar frame-
work.
The bulb itself is marked externally by the four prominent
plicae, one at each corner. The plicae are heavily sclerotized and
c-shaped in cross section, so that the lumen of each one forms a
distinct plicary canal (pi. en.). Posteriorly, the bulb merges
into the stomodaeal valve in much the same way as in all other
proventriculi, by a posterior constriction and sudden thinning of
the cuticle.
The proventriculus of Notoncus, like that of probably all the
other asepalous formicine genera, is peculiar in that it does not
intrude into the crop in dolichoderine fashion, but is actually
prevented from such intrusion by a constriction of the crop
immediately in front of the cupola (precupolar constriction;
Figs. 53, 54, precp. const.).
Histologically, Notoncus has undergone some striking innova-
tions (Figs. 53, 55-58). The circular muscles are most strongly
developed around the bulb, and at this level they do not differ
significantly in their arrangement from those of previously dis-
cussed proventriculi : the innermost fibers insert along the plicae,
while the peripheral fibers envelop the bulb without cuticulav
attachments. Anteriorly, the circular muscles extend past the
cupola to form a relatively strong sphincter surrounding the
precupolar constriction of the crop. In this arrangement the
Notoncus proventriculus differs from all previously discussed
proventriculi, in which, it will be recalled, the strongly developed
circular musculature terminates abruptly at the anterior ex-
tremity of the bulb, whence it continues as a loose network of
fibers over the crop.
EISNER: ANT PROVENTRICULUS 467
The principal group of longitudinal muscles (Fig. 53) 1
consider, on the basis of their position inside the circular
muscles, as homologues of l.m.l of previous proventriculi. These
fibers originate on the interplicary cuticle of the bulb and extend
anteriorly to insert, not on the cupolar quadrants, as in previous
proventriculi, but on the crop intima of the precupolar constric-
tion, in such a way as to act as antagonists to the sphincter sur-
rounding the constriction (some fibers apparently fail to insert
on the cuticle and extend anteriorly over the crop; see Fig. 53).
Outside the circular muscles there is present, as usual, a second
rather weakly developed group of longitudinal fibers. I am
tentatively homologizing these with l.m.3 of previous proven-
triculi, although their anterior insertion on the crop intima could
not be verified with certainty in Notoncus.
Thus we see that the most important modifications of the
musculature of the asepalous formicine proventriculus are cor-
related, on the one hand, with the loss of portal mobility, and
on the other with the acquisition of the precupolar constriction.
The development of a rigid immobile portal eliminated the need
for longitudinal muscles of the kind needed for quadrant dis-
engagement. The muscles were not lost, however, as occurred in
the higher dolichoderines with similarly immobile portals ; a shift
of their anterior insertion engaged them instead with the opera-
tion of the precupolar constriction of the crop.
In its function as a dam, the asepalous formicine proventricu-
lus presents the same advantages as that of the higher dolicho-
derines, inasmuch as the portal orifice is permanently reduced
to a narrow, immobile and densely pilose slit, rigidly braced by
the sclerotized cupola and plicae. Its damming function is
actually improved over that of the advanced dolichoderine pro-
ventriculus by the presence of the occlusory tract.
It is in the pumping action of the proventriculus, and specifi-
cally in the mechanics of ingress past the cruciform portal and
occlusory tract, that we find in Notoncus a functional deviation
from previous proventriculi.
It will be recalled that almost the entire occlusory tract is
by-passed at its four corners by an anterior extension of the
plicary canals. These four tips apparently constitute the princi-
pal incurrent pathways, since they represent the points of least
resistance to inflow. In conformity with this functional plan,
468 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
the occlusory tract is densely pilose, while the canals have naked
walls. In this method of ingress, the Notorious proventriculus
differs from the dolichoderine proventriculi with similarly im-
mobile portals. In these, there are no structurally differentiated
pathways of resistance through the portal ; resistance to inflow,
and therefore rate of inflow, is generally distributed over the
entire portal.
Following is my interpretation of the pumping action of the
proventriculus. As the bulb cavity collapses under muscle con-
traction, the contents are forced back through the cylinder into
the midgut. The entire occlusory tract becomes maximally oc-
cluded through a tight engagement of its pilose lining, and thus
acts to bar backflow into the crop. Also, as the bulb starts its
contraction, its four plicary canals are sealed off by approxima-
tion of their inner edges, perhaps imperfectly in the posterior
portion, but at least completely enough to act as an auxiliary
valve to prevent most of its contents from flowing back and forc-
ing their way through the portal tips.
As the circular muscles relax and the bulb cavity expands
due to the elasticity of its cuticular framework, the original
narrow orifice of the cruciform portal is restored. Simultan-
eously the plicary canals open again into the bulb lumen and
communicate the bulbar suction to the tips of the portal. Un-
doubtedly, some liquid is sucked in also through the occlusory
tract, but this portion is so extensive and densely covered with
hairs that traffic through it must be minor as compared to that
through the lateral canal system.
What is the function of the precupolar constriction? If we
follow the basic assumption that the longitudinal muscles con-
tract alternately with the circular muscles, it is apparent that
the constriction opens and the cupola is uncovered to project
freely into the crop just at the time when bulb expansion has
initiated fluid inflow through the portal. Thus, the opening
of the constriction acts as a pump in itself, inasmuch as it
assures the transfer of crop fluid to the immediately antecupolar
lumen of the crop, and this function is probably especially im-
portant when the crop contains only small amounts of fluid. An
additional function of the precupolar constriction is suggested
below, on page 480.
EISNER: ANT PROVENTRICULUS 469
MELOPHORES Lubbock
Species examined: M. sp.* probably ludius Forel), M. bay at i
Lubbock.
The outstanding peculiarity of Melophorus is to be found in
the cupola (Fig. 52), the face of which is slightly depressed in
the center and protuberant at its four corners. Otherwise, both
in its cuticular framework and arrangement of the muscles
(Figs. 59-61), the proventriculus does not differ grossly from
Notorious.
ACROPYGA Roger
Species examined: A. myops* Forel (or species near, collected
by W. L. Brown. Blaekall Range, S. E. Queensland, Australia,
1951 ) .
ANOPLOLEPIS Santschi
Species examined: A. lonyipes (Jerdon), A. custodiens (F.
Smith).
ACANTHOLEPIS Mayr
Species examined: A. frauenfeldi (Mayr).
DlODOXTOLEPIS Wheeler
Species examined: D. hiekmani (Clark).
The proventriculi of these genera (Figs. 51, 64, 65) resemble
one another on the basis of the strongly convex shape of the
cupola. Diodontolepis, which has not been illustrated herein,
bears very close resemblance to Anoplolepis. Histological^, only
Acropyya (Figs. 62, 63) was examined, and except for minor
structural peculiarities, such as the extreme thickness of the
plical walls, there were found no specializations in the arrange-
ment of the muscularis to suggest a functional deviation from
Notoncus. All these genera have a precupolar constriction like
Notorious.
AlYRMOTEEAS Forel
Species examined: M. williamsi Wheeler.
The single dried specimen that was available of this rare
470 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
species revealed a distinctly peculiar proventriculus (Fig. 66),
unique among those of asepalous formicines in the flattened and
cruciform aspect of the cupola, and in the elongate and pos-
teriorly tapering shape of the bulh. (Histologically preserved
specimens of this genus have recently been collected by Dr. E. 0.
Wilson and should be available shortly to amplify the present
study.)
Sepalous Formicine Proventriculus
(Figure 67, a diagrammatic representation of the sepalous
proventriculus, should be used as a reference throughout this
section.)
CamPONOTTIS Mayr
Species examined: C. vicinus* Mayr, C. americanus* Mayr, C.
herculednus (Linnaeus).
In the eamponotine proventriculus (Figs. 67, 68, 70-74), the
obvious distinctive character, here met for the first time, is a
long anterior extension called the calyx, consisting of four
heavily sclerotized strap-like cuticular arms, or sepals (sep.),
extending anteriorly from the bulb to capture part of the pos-
terior wall of the crop. (Sexual dimorphism is manifested in all
sepalous proventriculi by the slightly longer calyx in the
female.)
The sepals may be thought of as the anterior drawn-out bulbar
plicae of an asepalous precursor, that have carried along in their
extension the four corners of the cupola with their enclosed
portal tips. Each sepal is seen to contain a cylindrical canal, the
sepal canal (sep. en.) (continuous posteriorly with the plicary
canal of the bulb), that communicates along its entire length
with the lumen of the calyx through a narrow and densely
pilose longitudinal cleft (pt.) representing an extended arm of
the cruciform portal. On their luminal surface, the sepals are
covered with dense short pile (Fig. 71) similar to that lining
the cupolar face of asepalous formicine proventriculi. The
sepals themselves are joined by means of membranous sheets of
inter sepalary cuticle (intersep.), continuous with and of the
same texture as the intima of the crop wall proper.
EISNER: ANT PROVENTRICULUS 471
Posteriorly, the sepals curve together and fuse medially at
the level of the occlusory tract. In the structure of the occlusory
tract and bulb, Camponotus does not differ grossly from asepal-
ous proventriculi, and the rigid c-shaped plicae are again so
constituted that the canals they enclose merge with the bulb
lumen by way of a zone of constriction that can be sealed off
during bulb compression (Fig. 74).
Only relatively minor changes have occurred in the muscularis
(Figs. 70-74). The l.m.l fibers are seen to originate on the inter-
plicary cuticle of the bulb, and extend anteriorly past the oc-
clusory tract, at which point the inner fibers become inserted on
the intersepalary cuticle, while the few remaining outer fibers
continue anteriorly without cuticular attachments past the calyx
and over the crop. Attention is called to the inward fold of the
intersepalary cuticle at the bottom of the calyx (Fig. 70, a), and
into which insert the l.m.l fibers. This fold is undoubtedly all
that remains in this proventriculus of the precupolar constriction
of an asepalous ancestor.
Outside the circular muscles, a few scattered longitudinal
fibers extend from crop to midgut (Fig. 70, ?), apparently by-
passing the proventriculus without cuticular attachments. Prob-
ably these fibers contain only the usual crop-to-midgut fibers,
although it is possible that they are joined by homologues of
l.m.3 of previous proventriculi, that have here become detached
from the cuticle. In other sepalous formicine genera (see For-
mica next below) the muscles differ in their arrangement.
In its action as a pump, this proventriculus remains essentially
the same as its asepalous antecedent. The principal difference
between the two types lies, of course, in the sepalary develop-
ment and its functional consequences. In asepalous formicine
proventriculi, the presence of an occlusory tract — although a
distinct asset to the function of the proventriculus as a dam —
considerably restricts the portal area available for bulbar intake.
Only the outermost tips of the portal, being immediately ad-
jacent to the anterior extensions of the four plicary canals that
by-pass the occlusory tract, remain as pathways for fluid ingress.
Camponotus and the other sepalous formicines have successfully
exploited a means of increasing the bulbar intake area by extend-
ing, in the form of the calyx, both the plicary canals and their
associated portal tips. Thus, the calyx is to be regarded, not as
472 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
au adjunct to the damming function of the proventriculus, but
as a further refinement of the proventriculus as a pump. The
damming potential has, however, been retained, since increase in
portal area is accomplished without widening the portal at any
point.
During bulb contraction, when fluid is being evacuated from
the bulb through the stomodaeal valve, backflow through the
occlusory tract is prevented by the tight engagement of its oppos-
ing pilose linings. Simultaneously, the potential backfloAV route
through the plicary and sepalary canals is closed (or nearly
closed ) by the meeting of the edges of the grooves through which
canals and bulbar lumen communicate. When the bulb recovers
by means of elastic expansion, the grooves are reopened, and
the lumen again communicates with the canals, initiating a new
inflow through the sepalary portal slits. As in the asepalous
formicines, it seems that a small amount of fluid must also enter
directly through the occlusory tract at the stage of expansion;
however, the length, narrowness, dense vestiture and extremely
restricted mobility of the four clefts forming the tract surely
reduce such inflow to a negligible quantity, especially when one
considers the opportunity for fluid ingress offered by the portal-
plus-canals.
Emery (1888) believed the calyx to function exclusively as a
valve that, by sustained occlusion, acts to relieve the proventricu-
lus of fluid pressure from the crop. He disregarded the possi-
bility that the principal intake system of the bulb might be
routed through the sepals. He believed that the pilose lining
of the portal clefts actually seals off the sepal canals from the
calyx cavity: "Die Kinne wird gegen das Lumen des Kelches
durch das Ineinandergreifen der die Flugel besetzenden Har-
chen verschlossen." Accordingly, he was inclined to treat the
sepals as relatively unimportant structures. Subsequent workers,
including Forbes ('38), have observed that the canals really
open into the lumen, but not until recently (Eisner and Wilson
'52) has the possibility been considered that they might serve
to conduct fluid past the occlusory tract.
Brief mention should be made at this point of an incidental,
I nit probably significant function of the calyx. It is likely that
during bulbar activity the calyx undergoes pulsations synchron-
ous with those of the bulb, being alternately constricted by its
EISNER: ANT PROVENTRICULUS 47o
circular muscles and expanded by the elastic spring-like action
of the sepals. During calyx expansion, crop fluid tends to be
drawn into the calyx lumen, thereby assuring an adequate sup-
ply of the bulb just at the time when bulb expansion initiates
fluid intake through the sepalar-portal arrangement. Thus, while
increase of intake area was the principal evolutionary justifica-
tion for the appearance of the calyx, it seems that it came to
function also as an analogue of the precupolar constriction of
asepalous proventriculi.
It is doubtful that the l.m.l fibers play any significant con-
tributory role in the expansion of the calyx, since they insert,
not on the sepals themselves, but on the loosely folded and
flaccid intersepalary cuticle. Furthermore, in other sepalous
formicines (Formica), the similar structure of which suggests
no great functional divergence from Camponotus, l.m.l has been
lost entirely. I regard l.m.l in Camponotus as a mere remnant
persisting from an asepalous antecedent in which the function
of l.m.l was, as in Notoncus, the dilation of the precupolar con-
striction.
FORMICA Linnaeus
Species examined: F. sp.* (fusca group), F. subnuda Emery
(sanguinea group).
Aside from obvious but relatively minor differences in the
shape of the bulb and calyx, Formica (Figs. 69, 75, 100-103) dif-
fers significantly from Camponotus only in the arrangement of
the longitudinal muscles (compare Figs. 70 and 75). L.m.l
is entirely absent. There is present, outside the circular muscles,
a group of longitudinal fibers that insert anteriorly on the crop
intima so as to maintain a circumsepalar fold of the crop (Fig.
75) ; I am tentatively homologizing these fibers with l.m.3 of
previous proventriculi. In Camponotus (Fig. 70) no such cir-
cumsepalar fold was found, and the scanty outermost longitud-
inal muscles of the proventriculus extend from crop to midgut
without apparent cuticular connections.
Figures 76-84 illustrate a variety of the forms of cuticular
framework found among different sepalous formicine genera, in-
cluded here chiefly because of their taxonomic interest. The
features of each type as known at this time are clear from the
474 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
figures, and the individual cases require no discussion. (In
Myrmecocystus (Fig. 79) the tips of the sepals curve outward
only in the worker; in the female they are straight.)
Proventriculi were examined for a number of additional gen-
era, but since each of these closely resembles one of the genera
figured, it will suffice here merely to list the correspondences.
Gigantiops destructor (Fabricius), Dendromyrmex chartifex (F.
Smith), Opisthopsis haddoni Emery, and Polyergus breviceps
Emery resemble Formica. Polyrachis dives (F. Smith) is similar
to Camponotus. Pseudolasius mayri Emery closely resembles
Lasius.
Although such characters as the shape of bulb and plicae
and length of the sepals may in themselves be valuable phyletic
indicators, it is hoped that the internal evolutionary relation-
ships of this series will be further clarified by future work, and
especially by a comparative histological analysis of the longi-
tudinal musculature. The possibility should not be lightly dis-
missed that a major phylogenetic divergence may be outlined
by the presence or absence of l.m.l, as exemplified in this study
by Camponotus and Formica respectively.
Proventriculi with Incipient Sepals
ClADOMYRMA Wheeler
Species examined: C. hewitti Wheeler.
Only two dried specimens, a queen and a worker, probably
lit'stmates, were available for study of this very rare genus. The
proventriculus of the queen (Fig. 85) shows an incipient but
nevertheless distinctly typical calyx, and on this basis Clado-
myrma would seem to represent a survivor of the ancestral stock
that gave rise to the higher sepalous formicines. Sexual di-
morphism is strikingly exemplified in this genus by the com-
plete absence of a calyx in the worker, the proventriculus being
endowed with a typical asepalous cupola.
MyeMECORHYNCHUS Andre
Species examined: M. emery i Andre.*
Although distinctly suggestive of an incipient sepalar condi-
tion (Fig. 86), other peculiarities of the proventriculus, as well
EISNER: ANT PROVENTBICULUS 47.")
as characteristics of the whole ant (W. L. Brown, and E. 0.
Wilson, personal communication), point to the possibility that
Myrmecorhynckus may stand apart as an evolutionary offshoot,
having acquired its sepal-like structures by convergence.
The bulb is more capacious than usual (Fig. 89), and the
occlusory tract is not reduced to a narrow impassable cleft as in
other formicines, but is seen to he held agape (Fig. 88), sug-
gesting that bulbar intake is generally distributed over the entire
portal and occlusory tract instead of just at the tips. Most prob-
ably, this widening of the occlusory tract developed in response
to the increase in bulbar capacity. The cruciform portal also
differs from other formicines in that it lacks the usual pilose
vestiture.
In its musculature, Myrmecorhynckus is like Notoncus. The
precupolar constriction and l.ni.l are well developed and prob-
ably still functional.
PONEROID COMPLEX
Only very few representatives of this complex, including one
or two genera of each of the four major subfamilies currently
recognized (Brown '54), were examined for the present study.
In part this was due to the lack of properly preserved material,
an especially unfortunate handicap here since the relative uni-
formity of the cuticular framework of the poneroid proventricu-
lus forces one to turn to the arrangement of the muscles in the
search for phyletic indicators. The aberrant subfamily Leptanil-
linae could not be studied at all ; I did not succeed in recovering
a proventriculus from the single dried specimen that was avail-
able to me.
Subfamily PONERINAE
OdONTOMACHUS Latreille
Species examined: 0. haematoda* (Linnaeus), or near.
In its principal features (Fig. 90), the proventricular frame-
work conforms to the general plan of Myrmecia. The bulb is
elongate and posteriorly tapering. Only the interplicary plates
are somewhat sclerotized; the plicae themselves are thin and
flaccid. The cupolar quadrants are globose, and diverge from
47(i BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
one another, framing a widely gaping cruciform portal. The
presence of l.m.l and l.m.3, similarly arranged as in Myrmecia,
could be determined with certainty. The apparent absence of
l.m.2 is not to be taken as conclusive, since my preparations were
inadequate.
Both in its pumping action and damming potential, the pro-
ventriculus of Odontomachus probably resembles that of Myr-
mecia.
AMBLYOPONE Erichson
Species examined: A. australis* Erichson.
This proventriculus (Fig. 91) is strikingly unique among those
of ants, and possibly all other Hymenoptera, in that it deviates
from the basic quadripartite plan. The cupolar quadrants,
portal arms, plicae, and interplicary plates are six in number,
instead of the usual four. Except for this remarkable feature,
Amblyopone does not differ greatly from Odontomachus. The
cupolar quadrants are more closely approximated, and do not
diverge apically as in Odontomachus. The plicae are mem-
branous, and the entire proventriculus is weakly sclerotized and
flaccid. The material available was poorly preserved, rendering
impossible an exact delineation of the longitudinal musculature.
The l.m.l fibers are grouped into six separate bundles instead of
the usual four, each bundle following the course of its corre-
sponding interplicary plate.
The structural peculiarities of Amblyopone do not suggest
any major functional deviations. Its function, like that of
Odontomachus, is probably similar to that of Myrmecia.
Amblyopone is generally agreed to be one of the most primitive
of all living ants. The distinct peculiarities of its proventriculus
point up the possibility that Amblyopone may actually be more
highly specialized than previously suspected.
Subfamily CERAPACHYINAE
PHYRACACES Emery
Species examined: P. dumbletoni Wilson.
Although distinctive in its barrel-shaped bulb (Fig. 92), the
EISNER : ANT PROVENTRICULUS 477
proventriculus of Phyracaces conforms to the structural plan of
Myrmecia, and presumably functions in a similar way. The
presence of l.m.l and l.m.3, arranged as in Myrmecia, was deter-
mined with certainty, but there remains some doubt about the
apparent absence of l.m.2.
Subfamily MYRMICINAE
POGONOMYEMEX Mayr
Species examined : P . occidentalism (Cresson).
The cutieular framework (Fig. 95) consists of an elongate,
extremely flaccid and funnel-shaped ''bulb," devoid of a cupola,
merging anteriorly with the crop through a gradual expansion
of its lumen. Although the Avail of the bulb is produced into
four inward folds, vaguely delimiting plicary and interplicary
regions (Fig. 96), there are no textural differences between these
regions, the cuticle being evenly membranous throughout. The
circular muscles are strongly developed as usual, but have lost
all cuticular connection. The exact course and homologies of the
longitudinal muscles could not be determined. The principal
group of fibers, extending as four bundles along the inward folds
of the bulbar cuticle, are strongly suggestive of l.m.l.
It is difficult to explain the functional implications of the
obviously degenerate condition of this proventriculus. The loss
of an anterior valve-seal, elsewhere provided by the cupola, and
the loosely-folded membranous condition of the bulbar cuticle,
speak for a radical departure from the usual fluid pumping
action. It may well be that this type of proventriculus has be-
come modified to yield to the passage of solid matter, forced
through the bulb under muscular pressure. This possibility
deserves further attention, especially since the myrmicines are
known for their unusual feeding habits, several genera commonly
partaking of seeds and other vegetable material.
The structural divergences of the Pogonomyrmex proventric-
ulus would not seem to have reduced its capacity as a dam, since
the membranous bulb is readily occlusible under muscular con-
traction.
Forel (1878) and Emery (1888) called attention to the highly
aberrant proventriculus of the myrmicine tribe Cephalotini. This
proventriculus has re-acquired a cupola, in the form of a rigid
478 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
and elaborately ornamented umbrella, that abuts a typical funnel-
shaped and membranous myrmicine "bulb." The functional sig-
nificance of this unusual cupolar development is obscure at pres-
ent.
Hylomyema Forel
Species examined: H. colwmbica* Forel, or near.
This proventriculus is identical in all major respects to Pogo-
nomyrmex.
Subfamily DORYLINAE
ECITON Latreille
Species examined: E. hamatum* (Fabricius).
This proventriculus (Fig. 93), although closely resembling
that of Pogonomyrmcx, appears to have progressed even further
in its degeneration, the folding of the bulbar cuticle being com-
pletely randomized and irregular (Fig. 94). This proventriculus
is, in fact, nothing but a stomodaeal valve, preceded by a mus-
eularly maintained constriction of the posterior wall of the crop.
The same functional speculations advanced for Pogonomyrmex
apply to Eciton, except that the latter is wholly predaceous.
THE STOMODAEAL VALVE
Except in Myrmeeia, where it was described in considerable
detail, no emphasis has been granted to the stomodaeal valve
in the various, other proventriculi. The valve is, in fact, the
most uniform constituent of the ant proventriculus, and the only
structural modifications that seem to have functional overtones
occur in the degree of development of the muscular sphincter
that surrounds the neck of the valve at the point where it
pierces the midgut.
In Myrmeeia, as well as in all other generalized proventriculi
with divergent cupolar quadrants and a mobile portal (pseudo-
myrmecines, ponerines, and cerapachyines), the sphincter is
strongly developed, and probably serves in maintaining an occlu-
sion of the neck of the valve when the resting proventriculus
prevails in its function as a dam. In all the higher dolichoderines
EISNER: ANT PROVENTRICULUS 479
and formicines, in which proventricular damming is accom-
plished passively by an elastically maintained occlusion of the
cupolar or sepalar portal arrangement, the need for sustained
occlusion at the level of the stomodaeal valve has been lost, and
the sphincter surrounding the neck of the valve has accordingly
become reduced, sometimes consisting of not more than a single
layer of fibers.
IDENTITY OF THE "SEKRETSCHICHT" OF EMERY
(1888)
Emery granted some emphasis to the fact that the cupola of
most dolichoderine proventriculi seen by him (Tapinoma, Tech-
nomyrmex, Forelius, Conomyrma (=Dorymyrmex Emery nee
Mayr), Bothriomyrmex and Iridomyrmcx) is tightly overlain by
a discrete layer or "membrane" of varying thickness. Emery
assumed that this layer, which he called in its different forms
Sekretschicht, SeJcrethautchen, or Sekretblattchen, was a deposit
secreted by the cupolar epithelium. He found no such layer pres-
ent in Azteca, Liometopum, Leptomyrmex, or tribe Dolicho-
derini. Among the formicines, Emery found the layer present in
Camponotus ligniperdus, but makes no mention of it in the two
asepalous genera he examined, Plcujiolepis and Brachymyrmex.
In the histological preparations made for this work, a dense
layer, more or less unevenly granulose in texture (Fig. 98), was
found covering the cupola or sepals of the following genera :
Doliehoderinae : Forelius, Dorymyrmex, Iridomyrmex, Turneria,
( 'onomyrma, Teeh nomyrmex.
Formicinae: Camponotus, Formica.
The layer varied considerably in thickness, even among nest-
mates. Structurally, it showed considerable differentiation from
genus to genus, particularly in texture and in staining reaction ;
in some cases the layer was weakly basophilic, in others totally
refractory to the stains used.
There wTas no trace of a layer in Myrmecia, Pseudomyrmcx,
Aneuretus, Tapinoma, and members of the poneroid complex.
In Azteca, Liometopum, and Leptomyrmex, as well as in all
genera of asepalous Formicinae that were sectioned (Notoncus.
Melophorus, and Acropyga), there was present no compact
480 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY
layer, but the cupolar hairs held some clustered particles in loose
irregular arrangement (Fig. 99).
It is evident that the degree of development of the layer is
closely correlated with the amplitude of portal mobility. With
the notable exception of asepalous formicines, a compact layer
is present only in those proventriculi endowed with an immobile
slit-like portal. Given this correlation, my interpretation of the
"Sekretschicht" is that it is most likely not a secreted structure,
but merely a filtration aggregate consisting of those small parti-
cles that, having escaped the withholding action of the infra-
buccal chamber and the filtration devices within the buccal
cavity, have massed on the cupolar or sepalar surfaces, held back
by the filtering guard hairs of the proventricular portal.
There remains to be explained the absence of the layer in the
asepalous formicines. The precupolar constriction may afford
this explanation, considering its probable action of sweeping the
cupolar face repeatedly during the pumping cycle (see under
Notoncus, p. 468).
One apparent inconsistency in this hypothesis is the difference
found between Tapinoma and Forelius, despite the similar func-
tions proposed for these genera insofar as their damped portal
opening is concerned. Tapinoma sessile lacked the filtrate layer
in my preparations, while my Forelius possessed the layer in a
well-developed form. The probability that the portal does open
would at first glance seem to render difficult the accumulation
of sufficient filtrate to form a layer, but the degree to which
this opening is damped in these two genera actually makes it
more surprising that Tapinoma lacks the layer than it does that
Forelius has it.
As noted above, Emery found the layer present in Tapinoma,
but in a different group of the genus, in T. mclanocephalum
(Fabricius). Absence of the layer in my Tapinoma could be
taken as a sign that my interpretation of proventricular func-
tion is incorrect, and that the portal really opens more widely
than seems possible in view of the cupolar rigidity. However, the
structural qualities of the Tapinoma proventriculus lead me to
believe that the lack of a filtrate layer in my limited samples is
either a histological artifact or else reflects a lack of particulate
dietary constituents in these specimens, all drawn from a uni-
nidal series.
EISNER: ANT PROVENTRICULUS 481
PART III
DISCUSSION
One of the organs long used in ant classification is the pro-
ventriculus, a structure in which major changes are largely
concordant with subfamily, and even with many tribal breaks
based primarily on other structural characters. It is this con-
cordance that renders proventricular structure such a valuable
indicator of phylogeny within the ants, and at the same time
affords a basis for checking evolutionary steps in proventricular
modification. Even at the generic level, the proventriculus so
far as known offers no serious contradiction to presently ac-
cepted classifications, and the deeper one probes into the details
of musculature and other components, the more new useful in-
formation the organ yields for the systematist.
In Figure 97, a dendrogram is based on the structural rela-
tionships of the known proventriculi as revealed by the present
study, and interpreted in the light of the most recent phyletic
se-hemes. It will be necessary to refer to this figure in following
the discussion below.
The outstanding feature of the adult hymenopterous digestive
tract is the crop, a stomodaeal food reservoir acquired in adapta-
tion to the relatively discontinuous feeding habits characteristic
of these insects (Bischoff '27). The proventriculus is essentially
a structure accessory to the crop, and in this capacity serves
both as a dam, capable of containing the pressure exerted by
the liquid crop contents, and as a pump, regulating the delivery
of fluid to the midgut. It consists of a flexible but strongly-
muscled bulb, the pump, with intake and exhaust valves provided
respectively by the cupolar portal, and the stomodaeal valve.
The stomodaeal valve is embraced by a sphincter, the sustained
contraction of which presumably effects the dam.
This simple type of proventriculus is well suited to the habits
of the solitary Hymenoptera, the crop of which stores food for
its own use, or at the most for the use of itself and its mate
(Given '54).
With the acquisition of social habits and division of labor, the
burden of securing, transporting and transmitting food falls
entirely upon a part of the adult population. The brood, re-
482 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
prod uc tives, and non-foraging workers are more or less continu-
ous consumers of food, but contribute little or nothing to the
larder. There thus arises a necessitj' for a storage system — a
necessity accentuated under unfavorable environmental condi-
tions. Honeybees, for instance, solve this storage problem by
means of the comb, a communal storehouse external to their own
bodies.
In the ants, such storage facilities as exist, at least for fluids,
are usually limited to the crops of the individual workers. The
storage of amounts of fluid far in excess of the needs of the
individual worker, and for protracted periods of time, places a
strain on the arrangements by which damming depends upon
sustained muscular contraction of the stomodaeal valve sphincter.
Any structural changes tending to relieve the load of the
sphincter are therefore at a premium, especially if these changes
can shift the load to a passive barrier.
The generalized formicid pro vent riculus
In the lower ants (Ponerinae, Cerapachyinae, Myrmeciinae,
Pseudomyrmecinae, and Aneuretinae) the proventriculus is simi-
lar in all major respects to that of the other Hymenoptera so
far studied. Proventricular damming is probably maintained
exclusively b}* the stomodaeal valve sphincter, which in these
ants is especially well produced. The bulb is a flaccid structure
with gaping portal, incapable in itself of plugging the crop.
Damming is therefore a dynamic, energy-consuming process.
This basically inefficient mechanism limits the full exploitation
of the social functions of crop storage and regurgitative feeding.
Indeed, recent work emphasizes the fact that in these primitive
ants, regurgitative feeding is still in its incipient stages (Le
Masne '53, Haskins and Whelden '54). None of these ants is
known to have ' ' replete ' ' castes.
Dolichoderinae and ITormicinae
In the advanced dolichoderines (tribe Tapinomini) and all
formicines, proventricular damming no longer depends upon the
continuous muscular pressure of the stomodaeal valve sphincter,
and the sphincter in these subfamilies is greatly reduced in size.
EISNER: ANT PROVENTRICULUS 483
The damming function has shifted to the reinforced cupola,
which now forms a rigid shield, pierced by a narrow, densely
pilose cruciform slit, and capable by itself of stemming leakage
from the crop. Damming is therefore accomplished passively,
without expenditure of muscular energy. The evolution of the
proventriculus in these subfamilies is the history of the elaborate
and varied pathways of change along which the proventriculus
adapted to its new function as a passive dam.
An early step in the evolution of the damming cupola is
exemplified in the dolichoderines by the Azteca-Tapinoma pro-
ventricular type, and in the formicines by the asepalous type.
Already these proventriculi have acquired the pilose portal slit,
strongly braced by the sclerotized cupola and bulbar plicae, and
able to hold back fluids when the inactive proventriculus pre-
vails in its function as a dam. That these remarkably similar
refinements arose convergently in the dolichoderines and formi-
cines is attested by the differences in the arrangement of the
longitudinal muscles and by other characters.
The restriction both in size of orifice and in mobility of the
cupolar portal arrangement — although a distinct asset to the
damming function of the proventriculus — tends to render the
portal orifice inadequate for bulbar intake during the pumping
cycle. This handicap is accentuated in Azteca and the asepalous
Formicinae by the interposition, between portal and bulbar
lumen, of an occlusory tract. In order to reconcile the basic
pumping action of the bulb with the new damming qualities of
the portal, Azteca and Tapinoma retain a limited range of
portal mobility, the quadrants being slightly dislocated during
bulbar intake under the action of l.m.l muscle fibers. Dislocation
is damped, with the sclerotized bulbar plicae acting as spring
antagonists to the retractor muscles. Unlike those of Azteca and
Tapinoma, the asepalous formicine proventriculus can accom-
plish its pumping action without quadrant dislocation. It has,
in fact, no muscles adapted to moving the quadrants, the l.m.l
fibers having shifted their insertion away from the cupola itself
to become concerned with the operation of the precupolar con-
striction of the crop. Bulbar intake is routed through the plicary
canals, the anterior extensions of which by-pass the occlusory
tract to join the four corners of the portal. This mechanism of
ingress, although using only a relatively small portion of the
484 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
portal, must provide sufficient flow for the active pumping action
of the organ.
The most advanced proventricular types of the dolichoderines
(Iridomyrmex, Dorymyrmex, Conomyrma, and Technomyrmex)
and formicines (Camponotus, and all other sepalous proventric-
uli), although differing radically in structure, are actually re-
markably similar in their functional achievements. They retain
the narrow slit-like portal attained by their respective evolu-
tionary antecedents, and are therefore similarly adapted to serve
as passive dams. In these proventriculi, the need for portal
disengagement is completely eliminated, and the quadrant re-
tractors (l.m.l) are entirely absent or if retained, as is the case
in some sepalous proventriculi (Camponotus) , they are function-
ally impotent as retractors. The complete loss of portal mobility
is coincident with certain structural refinements designed to
increase the portal orifice: bulbar capacity ratio needed to assure
adequate bulbar intake during the pumping cycle. In the doli-
choderines this is accomplished by a reduction in bulbar capacity
through progressive envelopment of bulb by cupola, a process
shown in its incipient stage in Forclius, and culminating in
Iridomyrmex. In the formicines, bulbar capacity remains un-
altered, there being instead an increase in the portal orifice
through anterior extension of the portal tips in the form of
sepals. Since the portal is increased in length but not in width,
its damming potential is not sacrificed. The development of
sepals is seen in its incipiency in Cladomyrma.
Two relatively primitive dolichoderine genera, Liometopum
and Leptomyrmex, having in common certain proventricular
specializations unique among the dolichoderines, apparently rep-
resent one or two isolated and probably long-standing evolution-
ary offshoots. Both proventriculi feature an extensive occlusory
tract, a distinct asset to the damming action of the proventricu-
lus. Potentially, this occlusory tract tends to obstruct bulbar
operation, but since it is articulated laterally by membranous
plicae, it is easily opened by l.m.l during bulbar intake.
The adaptive significance of the passive-dam proventriculus of
the dolichoderines and formicines is reflected by the full exploita-
tion in these ants of the social functions of crop storage and
regurgitative feeding (Le Masne '53). All ant genera known
EISNER: ANT PBOVENTRICULUS 485
to have "replete" castes are restricted to one or the other of
these subfamilies.
Myrmicinae and Dorylinae
The proventriculus of the myrmicines (with the exception of
the Cephalotini; see Emery 1888) and the dorylines has become
degenerated to the point where it consists essentially of only a
stomodaeal valve; the bulb is presumably non-functional as a
pump, since it is reduced to a membranous and more or less
irregularly folded constriction of the posterior crop. From the
structure of the organ it is clear that proventricular damming
is not a passive process, but must be dependent upon muscular
contraction, at the level of the stomodaeal valve and/or the
"bulb." Although regurgitative feeding is known to occur
among some of these ants (Crematogaster; "Wilson and Eisner
'57) nothing is known about the length of time over which the
crop can store nutrient. It is interesting to note in this connec-
tion that some myrmicines have become independent of the crop
as the exclusive communal reservoir. Instead, they have de-
veloped a storage system outside their own bodies, hoarding
seeds or other types of solid nutrient.
It may be relevant here to emphasize a major functional
limitation of the passive-dam dolichoderine and formicine pro-
ventriculi. The complete loss of portal mobility, resulting from
the acquisition of a rigid slit-like portal, would seem to preclude
completely the transfer of any but the most minute particulate
matter to the midgut. We have seen, in fact, that in the advanced
doliehoderines, even minute particles are not always passed to
the midgut, but actually accumulate over the cupolar face in
more or less discrete layers. We have also seen that in the
asepalous formicine proventriculi, such accumulation of particles
does not take place, presumably being prevented by the sweeping
action of the precupolar constriction of the crop.
It would be interesting to know whether, in all ants with a
rigid slit-like portal, there has been an increasing dependence
upon cephalic solid-withholding devices such as the infrabuccal
chambers or the ridges of the buccal cavity mentioned by
Wheeler ('10). It would even be worthwhile considering
whether the refinement of such buccal particle-traps was a pre-
486 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
adaptation necessary for the evolution of the passive-dam pro-
ventriculus. There is much need for detailed observations on the
feeding behavior of ants having such proventriculi. Just because
an ant is seen feeding on a dead insect, for instance, does not
mean that it is swallowing solid matter, since it may actually take
in only the juices. One would even expect that small particles
such as pollen grains, which are known to figure prominently
as a source of protein in the diet of adult honeybees, are of too
large diameter to be transferred through the passive-dam pro-
ventriculi.
The proventriculi with mobile portals, on the other hand, such
as those of the myrmicines, pseudomyrmecines, aneuretines, pon-
erines, and cerapachyines (and, for that matter, of all other
aculeate families in which the organ has been studied), are con-
structed in such a way that no hindrance would appear to be
offered to the passage of at least small particles. Perhaps some
lesser particles such as pollen grains actually constitute a staple
source of protein in some of these ants. From the work of Bailey
( '52) we know, for instance, that in Vespa, Bombus, and Proso-
pis, pollen suspensions do not choke the proventriculus and are
effectively pumped to the midgut. In Apis, the proventriculus
has certain structural peculiarities enabling it, at least under
some circumstances, to strain out the pollen grains from the fluid
crop contents and selectively transfer them to the midgut
(Bailey '52, Schreiner '52). This type of activity, while prob-
ably incidental to the primary fluid-handling function of the
proventriculus, may be of importance when the bee requires large
quantities of pollen protein for the synthesis of brood-food secre-
tion (Bailey '54). Whether any activity similar to the pollen-
transfer action of the Apis proventriculus occurs in any of the
ant proventriculi with mobile portals remains to be shown.
The proventricular structure of the dorylines and myrmicines
(with the single known exception of the cephalotines) presents
no obvious obstacle to the passage of even large particles, but we
do not yet know whether these ants actually swallow solids. It
would be interesting to know this, because it is not beyond possi-
bility that the "degenerate" condition of the proventriculus in
these ants is actually an adaptive response to changed feeding
habits involving greater use of injested solids.
EISNER: ANT PROVENTRICULUS 487
I am fully cognizant of the fact that the often elaborate functional
schemes advanced here for the various proventricular types are largely
speculative, and that future work may uncover fallacies in my reasoning.
However, there is only one basic assumption underlying this study and for
which I have no direct substantiating evidence, namely, that the circular
muscles and longitudinal muscles contract alternately rather than simul-
taneously. Actually, it seems impossible to conceive of any workable
proventricular mechanism based on a synchronous activity of these muscles.
The other lines of reasoning used in connection with functional interpreta-
tions, such as evaluation of the relative effectiveness of the various muscles,
are not completely speculative, since they are based in part on an estimation
of the relative mobility of the various proventricular components that these
muscles operate. Mobility of the cupolar quadrants, for instance, is readily
disclosed by prying with a glass needle. Similarly, the elasticity of the
bulb can be evaluated by observing its resilience under varying pressure
directly applied.
It would be interesting to test the functional schemes advanced here by
direct experimentation. I have not yet tried to evaluate the relative damming
efficiency of the various proventricular types by determining 1) the amount
of fluid that can be stored in the crop, and 2) the length of time over
which storage can be maintained. However, such studies, and others along
these general lines, are currently planned or already under way in this
laboratory. The use of radioactive tracers appears to be especially promis-
ing. A preliminary investigation on the relative importance of regurgitative
feeding in several representative species of ants is in press (Wilson and
Eisner '57, Eisner and Wilson, '57).
LITERATURE CITED
Bailey, L.
1952. The action of the proventriculus of the worker honeybee, Aph
mellifera L. J. Exp. Biol., 29: 310-327.
1954. The filtration of particles by the proventriculi of various
aculeate Hymenoptera. Proc. Roy. Ent. Soc. London, A29:
119-123.
BlSCHOFF, H.
1927. Biologie der Hymenopteren. Springer Verlag, Berlin.
Brown, W. L.
]9."4. Remarks on the internal phylogeny and subfamily classification
of the family Formicidne. Insectes Sociaux, 1: 21-31.
488 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
Dobrovsky, T. M.
1951. Postembryonic changes in the digestive tract of the worker
honeybee (Apis mellifera L.). Memoir, Cornell Univ. Agr. Exp.
Sta., 301: 3-45.
Eisner, T. and \V. L. Brown
1957. The evolution and social significance of the ant proventriculus.
Proc. Tenth Internat. Congr. Ent. (in press).
Eisner, T. and E. O. Wilson
1952. The morphology of the proventriculus of a formicine ant.
Psyche, 59: 47-60.
1957. Radioactive tracer studies of food transmission in ants. Proc.
Tenth Internat. Congr. Ent. (in press).
Emery, C.
1888. Uber den sogenanuten Kaumagen einiger Ameisen. Zeitschr.
wise. Zool., 46: 378-412.
1912. Formicidae, subf. Dolichoderinae. Wytsman's Genera In-
sectorum, fasc. 137: 1-50.
1925. Formicidae, subf. Formicinae. Ibid., fasc. 183: 1-302.
Forbes, J.
1938. Anatomy and histology of the worker of Camponoius hercu-
leanus pennsylvanicus De Geer (Formicidae, Hymenoptera ').
Ann. Ent. Soc. Am.. 31: 181-195.
EOREL, A.
1S7S. Etudes myrmecologiques en 1878 (premiere partie) avec l'an-
atomie du gesier des fourmis. Bull. Soc. Vaudoise Sci. Nat..
15: 337-392.
Given, B. B.
1954. Evolutionary trends in the Thynninae (Hymenoptera: Tiphii-
dae) with special reference to feeding habits of Australian
species. Trans. Roy. Ent. Soc. London, 105: 1-10.
11 vSKiNS, C. P. and R. M. Whelden
1954. Note on the exchange of ingluvial food in the genus Myrmecia.
Inseetes Sociaux, 1: 33-37.
Kennedy, C. H.
1949. Methods for the study of the internal anatomy of insects. Ohio
State Univ., Columbus, Ohio.
Le Masne, G.
1953. Observations sur les relations entre le couvain et les adultes ehez
lew fourmis. Ann. Sci. Nat. Zool., (11) 15: 1-55.
EISNER: ANT PROVENTRICULUS 489
Pantin, C. F. A.
1948. Notes on microscopical technique for zoologists. Cambridge
Univ. Press.
SCHREINEB, T.
1952. Tiber den Nahrungstransport im Darm der Honigbiene. Zeitschr.
vergl. Physiol., 34: 278-298.
Snodgrass, R. E.
1956. Anatomy of the Honey Bee. Comstock Publishing Associates
(Cornell Univ. Press).
Wheeler, W. M.
1910. Ants, their structure, development, and behavior. Columbia
Univ. Press.
1923. Social life among the insects. Harcourt, Brace and Co., New
York.
vVilson, E. 0. and T. Eisner
1957. Quantitative studies of liquid food transmission in ants. Inseetes
Sociaux (in press).
Wilson, E. O., T. Eisner, G. C. Wheeler and J. Wheeler
1956. Aneuretus simoni Emery, a major link in ant evolution. Bull.
Mus. Comp. Zool. Harvard, 115: 81-99.
KEY TO ABBREVIATIONS
The measurements given for proventricnlar length were taken
between cupolar summit, or sepal tips, to the base of the bulb
at the point of insertion of the stomodaeal valve.
All longitudinal sections show only one half of the proventric-
ulus, and are taken through the interplicary cuticle of the bulb,
midway between the plicae.
bl bulb
b.m basement membrane
cm circular muscles
cp cupola
Cr crop
interpl interplicary plate of bulb
intersep intersepalary cuticle
l.m.l longitudinal muscles no. 1
l.m.2 longitudinal muscles no. 2
l.m.3 longitudinal muscles no. 3
M. G midgut
490 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
occ. tr occlii8ory tract
phr phragma
pi. . plica
pi. en plicary canal
precp. const precupolar constriction
pt portal
pv proventriculus
qd quadrant
sep sepal
sep. en sepal canal
sin sinus
st. vlv stomodaeal valve
PLATES
PLATE 1
Fig. 4. Myrmccia regularis Crawley. Cuticular framework. (Worker;
0.40 mm.)
Fig. 5. Pseiidomyrmt.r 'pallidm (F. Smith). Cuticular framework, diag-
rammatic. (Worker; 0.14 mm.)
Fig. 6. Aneuretus simoni Emery. Cuticular framework, diagrammatic.
(Worker; 0.08 mm.)
PLATE 1
I -LATE 2
Fig. 7. Hypoclinea pustulata (Mayr). Cuticular framework. (Worker;
0.18 i .)
Fig. 8. Liometopum sp. (near apicidatum Mayr). Cuticular framework.
(Worker; 0.24 mm.)
Fig. !•. Leptomyrmex onemidatus Wheeler. Cuticular framework
Worker; 0.44 mm.)
PLATE 2
PLATE ::
Fig. 10. Myrtneda vindex F. Smith. Longitudinal section through bull)
;ill:l Cupola.
Fig. 11. M. regularis Crawley. Longitudinal section through stomodaeal
valve.
Fig. 12. M. vindex F. Smith. Cross section through cupolar quadrants.
Fig. L3. Same. Cross section through middle of bulb.
Fig. 14. Same. Cross section through posterior fifth of bulb.
PLATE 3
PLATE 4
Fi^'. 1~>. Pseudomyrmex pallidus (F. Smith). Cross section through
middle of bulb.
Fig. Hi. Same, ('mss section through base of bulb.
Pig. 17. Hypoclinea p-untulata (Mayr). Longitudinal section through
bulb and cupola ( diagrammatic i.
Fig. 18. Same. Cross section through cupolar quadrants.
Pig. 19. Same. Cross section through middle of bulb.
Pig. 20. Same. Cross section through base of bulb.
Pig. 21. .tin tin l us simoni Emery. Longitudinal section through bulb
.■mil cupola.
PLATE 4
PLATE 5
Fig. 22. Liometopum sp. (near apiculatum Mayr). Longitudina] section
through cupola, oeelusory tract, and bulb (diagrammatic).
Fig. 23. Leptomyrmex pollens Emery. Longitudinal section through
cupola, oeelusory tract, and lmlti.
Fig. 2-4. Same. Cross section through middle of oeelusory tract,
Fig. 25. Same. Cross section slightly below middle of bulb.
occ.tr.
y ' ima
l.m. 3
occ. tr.
«t.wlv.AVXf
PLATE 5
PLATE 6
Fig. li * » . Tapinoma sessilt (Say). Cuticular framework. (Worker;
0.12 mm.)
Fig. 27. Same. Longitudinal section through cupola and bulb.
Fig. 28. Same. Cross section through cupola near extremes of porta] 1i]>s.
Fig. 29. Same. Cross section through middle of bulb.
Fig. 30. Asteca sp. Cuticular framework. (Worker; 0.18 mm.)
Fig. 31. Same. Longitudinal section through cupola, occlnsory tract, and
bills ( diagrammatic I .
■j^«= — stvlv
PI.ATK6
'LATE 7
bvig. 32. Iridomyrmex detectus (F.Smith). Cuticular framework, (queen;
0.37 mni. >
Fig. '■>■':. Conomyrma thoracica Santschi. Cuticular framework. (Worker;
0.12 mm. >
PLATE 7
PLATE 8
Fig. 34. Iridomyrmex detectus (F. Smith). Longitudinal section through
cupola and bulb.
Fig. 35. Same. Cross section through widest portion of r-upola.
Fig. 30. Forelius foetidtts (Buckley). Longitudinal section through
cupola and bulb.
Fig. 37. Conomyrma thoracica Santschi. Longitudinal section through
cupola and bulb (diagrammatic).
Fig. 38. Same. Slightly oblique cross section through cupola.
Fia\ 39. Same. Cross section through middle of hull'.
fS2^ stvlv
PLATE 8
PLATE !»
Fig. 40. Fun h us fa' I nl us (Buckley). Outline of lateral view. (Worker;
0.10 linn. I
Fig. 41- Same. Frontal view.
Fig. 4:2. Dorymyrmex ensifer Forel. Outline of lateral view. (Worker;
0.12 nun.
Fig. 43. Same. Frontal view.
Fig. 44. Iridomyrmex viridiaeneus Viehmeyer. Outline of lateral view.
Worker; 0.15 mm.
Fig. 4". Same. Frontal view.
PLATE 10
Fig. 46. Conomyrma thoracica Santschi. Outline of lateral view.
( Worker : 0.12 mm.)
Fig. 47. Same. Frontal view.
Fig. 48. Tedhnomyrmex detorquens (Walker). Outline of lateral view.
( Worker ; <>.l 1 mm. I
Fig. 49. Same. Frontal view.
PLATE 11
Fig. 50. Notorious ectatommoides fForel). Cuticular framework, (queen;
(i.22 mm.)
Fig. 51. Acropyga myops Fore! (or species near)- Cuticular framework.
(Worker; 0.23 mm. J
Fig. 52. Melophorus bagoti Lubbock. Cuticular framework, (queen;
0.55 mm.
'tiC
PLATE I I
PLATE 12
Fig. 53. Notoncus ectatommoides (Forel). Longitudinal section through
cupola, occlusory tract, and bulb.
Fig. 54. Same. Proventrieulus in relation to crop and midgut. (Note
the precupolar constriction of the crop.)
Fig. 55. Same. Cross section through cupola.
Fig. 56. Same. Cross section through occlusory tract.
Fig. 57. Same. Cross section through anterior third of bulb.
Pig. 58. Same. Cross section through base of bulb.
loiSL Z_occ Ir
precp cons!
»3g2_lm 3(?)
l.m.3(?)
precp- const.
PLATE 12
PLATE 13
Fig. 59. Melophorus sp. (probably ludius Forel). Cros3 section through
cupola.
Fig. 60. Same. Cross section through middle of occlusory tract.
Fig. 61. Same. Cross section through middle of bulb.
Fig. 62. Acropyga myops Forel (or species near). Cross section through
middle of cupola.
Fig. 63. Same. Cross section slightly above middle of bulb.
pi en.
I. ml
l.m.3(?)
PLATE 13
PLATE 14
Fig. 64. Anoplolepis custodiens (F. Smith). Cuticular framework.
(Worker; 0.35 mm.)
Fig. 65. Acantholcpis frauenfeldi (Mayr). Cuticular framework. (Worker;
0.25 mm.)
Fi<>'. (><i. Myrmoteras williamsi Wheeler. Cuticular framework. (Worker;
0.1 1 mm.)
PLATK 14
VI
PLATE 15
Fig. 67. Exploded diagram of the generalized sepalous formieine pro
'ntriculus (based on Camponotus).
PLATE 15
PLATE 16
Ki or. 68. Camponotus vicintis Mayr. Cuticular framework. (Major
worker ; 1 .07 nun. I
Fig. 69, Formica sp. (fusca group). Cuticular franiewoi'k. (Worker;
0.63 mm. I
PLATE 16
'LATE 17
Fig. 70. Cavipoiiotus arnericanus Mayr. Longitudinal section through
ealyx (intersepalary cuticle i and bulb.
Fig. 71. Camponotus vicinus Mayr. Cross section through middle of
calyx.
Fig. 72. Same. Cross section through base of calyx.
Fig. 7:'.. Same. Cross section through ocelusory tract.
Fig. 74. Same. Cross section through middle of bulb.
Fig. 77). Formica sp. (fused group). Longitudinal section through calyx
(intersepalary cuticle) and bulb.
PLATE 1
PLATE 19
Pig. 71*. Myrmecooystus mexicanus (Wesmael). Cuticular framework.
(Worker; 0.53 mm.)
Fig. 80. Gesomyrmeal lusonensis (Wheeler). Cuticular framework.
(Worker; 0.30 mm.)
Fig. 81. Paratrechina longicornis (Latreille). Cuticular framework.
(Worker; 0.17 mm.)
PLATE 1<>
PLATE 20
Fig. 82. CataglypMs sp. (near bicolor Falnicius). Cuticular framework.
( Worker; 0.64 mm.)
Fig. 83. Oecophylla smaragdina (Fabricius). Cuticular framework.
(Worker; 0.43 mm.)
Fig. 84. Brachymyrmex obscurior Forel. Cuticular framework. (Worker;
0.30 mm.)
PLATE. 20
PLATE 21
Fig. 85. Cladomyrma heioitti Wheeler. Cuticular framework. ( Queen :
(1.37 mm. )
Fig. 86. Myrmecorhynchus emeryi Andre. Cuticular framework. (Worker;
0.15 mm. )
Fig. 87. Same. Cross section through middle of bulb.
Fig. 88. Same. Cross section through occlusory tract.
Fig. 89. Same. Cross section through middle of bulb.
PLATE 21
PLATE 22
Fig. 90. Odontomachus haematoda (Linnaeus) or near. Cuticular frame
work (diagrammatic). (Worker; 0.25 nvni.)
Fig. 91. A in hi I/O pone dlistralis Erichson. Cuticular framework (diagram-
matic I. i Worker ; 0.20 mm.)
Fig. 92. Phyracaces dumbletoni Wilson. Cuticular framework (diagram
matic). ('Worker; 0.15 mm.
PLATE 22
PLATE 24
Pig. !»7. Dendrogram showing evolution of the formicid proventriculus.
CN
<
PLATE 25
Fig. 98. Iridomyrrrkex detectus (F. Smith). Longitudinal section through
proventriculus. Note (arrow) the more or less compact layer of filtrate
overlaying the eupola.
Fig. 99. Leptomyrmex pattens Emery. Longitudinal section through pro-
ventriculus. Note (arrow i the loose aggregate of filtrate among the
cupolar hairs.
Fig. 100. Formica sp. (fusca group). Cross section through anterior third
of calyx.
Fig. 101. Same. Cross section through base of calyx.
Fig. L02. Same. Cross section through occlusory tract.
Pig. 103. Sunie. Cross section through middle of bulb.
PLATE 25
Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE
Vol. 116, No. 9
THE IXODES RASVS GROUP OF AFRICAN TICKS WITH
DESCRIPTIONS OF FOUR NEW SPECIES
(IXODOIDEA, IXODIDAE)
By Don. R. Arthur
Department of Zoology, King's College, London,
and Consultant, Department of Medical Zoology,
U. S. Naval Medical Research Unit No. 3,
Cairo, Egypt
In Collaboration with
Colin Burrow
Harold Row Student, King 's College, London
CAMBRIDGE, MASS., U.S.A.
printed for the museum
July, 1957
Publications Issued by or in Connection
with THE
MUSEUM OF COMPARATIVE ZOOLOGY
AT HARVARD COLLEGE
Bulletin (octavo) 1863 — The current volume is Vol. 116.
Breviora (octavo) 1952 — ■ No. 78 is current.
Memoirs (quarto) 1864-1938 — Publication was terminated with Vol. 55.
Johnsonia (quarto) 1941 — A publication of the Department of Mollusks.
Vol. 3, no. 35 is current.
Occasional Papers of the Department of Mollusks (octavo) 1945 —
Vol. 2, no. 21 is current.
Proceedings of the New England ZoSlogical Club (octavo) 1899-
1948 — Published in connection with the Museum. Publication terminated
with Vol. 24.
The continuing publications are issued at irregular intervals in numbers
which may be purchased separately. Prices and lists may be obtained on
application to the Director of the Museum of Comparative Zoology,
Cambridge 38, Massachusetts.
Of the Peters "Check List of Birds of the World," volumes 1-3 are out
of print; volumes 4 and 6 may be obtained from the Harvard University
Press; volumes 5 and 7 are sold by the Museum, and future volumes will be
published under Museum auspices.
Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE
Vol. 116, No. 9
THE IXODES RASVS GROUP OF AFRICAN TICKS WITH
DESCRIPTIONS OF FOUR NEW SPECIES
(IXODOIDEA, IXODIDAE)
By Don. R. Arthur
Department of Zoology, King's College, London,
and Consultant, Department of Medical Zoology,
U. S. Naval Medical Research Unit No. 3,
Cairo, Egypt
In Collaboration with
Colin Burrow
Harold Row Student, King 's College, London
CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM
July, 1957
No. 9 — The Ixodes rasus Group of African Ticks with
Descriptions of Four New Species (Ixodoidea, Ixodidae)
By Don R. Arthur x and Colin Burrow
INTRODUCTION
Hitherto the African ticks of the genus Ixodes which possess
closed circular anal grooves have been incompletely investigated
and all such ticks have been referred to the species Ixodes rasus
Neumann 1899. This approach to the diagnoses of these ticks
was established by Nuttall, Warburton, Cooper and Robinson
(1911) and used indiscriminately until Schulze (1943) sub-
divided rasus into three subspecies (see later). Consequently,
at the present time the status of rasus and other forms as yet
undescribed presents an interesting problem to the systematist.
This uncertainty of structure, coupled with our ignorance of the
biology of the rasus group, opens up a particularly interesting
and virgin field for research, and not a few unknown allied
species undoubtedly await discovery.
Nuttall (1911) classified I. rasus in the biological group within
the genus Ixodes in which males and females coexist together
on a host that either wanders or does not travel far and in the
subgroup where the sexes are found in copula on the host. Even
so, the published reports show that the host range of the adults
of the I. rasus group varies from small insectivores (mice, ele-
phant shrews) to leopards, large antelopes, domestic dogs and
man. The picture for immature stage-host relationships is more
uncertain.
Specimens on which this report is based were obtained from Dr.
H. Hoogstraal, NAMRU-3, Cairo, Egypt; Dr. Gertrud Theiler,
Onderstepoort Veterinary Research Department; The Museum
of Comparative Zoology (through the courtesy of Dr. J. Be-
quaert) ; Musee Royal du Congo Beige (through the courtesy of
Dr. E. Dartevelle) ; Dr. Pierre Morel, Laboratoire Federal de
l'Elevage George Curasson, Dakar, Senegal; Rocky Mountain
i The opinions and statements contained herein are the private ones of the
Writers and are not to be construed as official or reflecting the views of the Navy
Department or the Naval Service at large.
494 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Laboratory, Montana (through the courtesy of Mr. Glen M.
Kohls) ; Dr. J. Mouchet, Direction des Services de Sante Pub-
lique, Yaounde, Cameroun Frangais; British Museum (Natural
History) through the courtesy of Dr. G. Owen Evans and Mr. E.
Browning; Mr. W. Grey, Mazabuka, Northern Rhodesia; Dr.
Brazzard, Laboratoire d'Histoire Naturelle, Ecole Nationale
Veterinaire, Toulouse, France.
This work was initiated when the writer (D.R.A.) was in re-
ceipt of a Leverhulme Research Award and completed on a
visit to East Africa under a Colonial Welfare and Development
Grant made available by the Colonial Office. This author wishes
to record his thanks to these organizations as well as to members
of the East African Veterinary Research Organization (particu-
larly to Miss Jane Walker) for their hospitality, and to U. S.
Naval Medical Research Unit No. 3, Cairo, Egypt for their great
assistance and many kindnesses.
The following abbreviations are used to denote the sources of
the material examined: BM, British Museum (Natural History) ;
CNHM, Chicago Natural History Museum ; EAVRO, East Afri-
can Veterinary Research Organization; GHFN, Nuttall Collec-
tion, British Museum (Natural History) ; HH, Harry Hoogstraal,
Cairo ; JM, J. Mouchet, Cameroons ; MC, Musee Royal du Congo
Beige, Tervuren (Belgique) ; MCZ, Museum of Comparative
Zoology; OP, Onderstepoort Research Station; RML, Rocky
Mountain Laboratory Collection.
SYSTEMATIC DESCRIPTIONS
Ixodes rasus Neumann 1899
(Figures 1-13)
Ixodes rasus Neumann (1899), pp. 137-39, Figs. 12-14, described from 3
females and 1 male (cf. Remarks below) from Belgian Congo (cf.
Bequaert, 1931, who refers type locality to French Equatorial Africa).
Nuttall, Warburton, Cooper and Robinson (1911) repeat Neumann's
description of the male, and describe another species of female (see
under Ixodes pseudorasus) . Schulze (1943) subdivided rasus into three
subspecies, viz: /. rasus rasus, I rasus cumulatimpunctatus and /. rasus
eidmanni. (Cf. Remarks below.)
ARTHUR: IXODES RASUS GROUP 495
Type material. Originally described by Neumann (1899, pp.
137-39) from three females and a copulating male, from Hyrax
species collected in "the Congo" in A. Mocquerys coll. 1899.
The present labelling of these specimens reads "740. Ixodes
rasus, 1 male, 2 females (-1 female) Hyrax species, A. Mocquerys
coll. 1899. G. Neumann det." An anomalous situation occurs
here as Neumann stated that there were three females, whilst
the present information states "2-1 females," and accordingly
1 propose that the remaining female specimen becomes the electo-
type. Electotype female, and allotype male deposited in the
Neumann collection, No. 740, at the Ecole Nationale Veterinaire,
Toulouse, France.
Paratype: 1 female, Viverra civetta, Congo Beige. Deposited
at the Ecole Nationale Veterinaire, Toulouse, France, No. 761.
MATERIAL EXAMINED. Total 43 females; 6 males. 1 fe-
male (no host data) Dakar, Senegal: 1 female, wild pig, Came-
roons, Fr. Berlin Zoo Museum, Dr. Schafer (GHFN— 3005) : 1
female, Felis pardus L., primary forest, Mainyu Bridge, Mamfe,
Cameroons 500 ft. alt., 12.5.33. P. Sladen Trust Expedition
(BM) : 2 females, Manis tricuspis (=Phataginus tricuspis (Ra-
fmesque) ) secondary forest, Bashamii, Mamfe, Cameroons 23.3.33.
I. T. Sanderson leg. (BM) : 3 females (originally, 6 females ac-
cording to legend in vial) , Cephalophus leucogasier Gray, Efeileu,
Bulu Country, Cameroons. 28.6.33. I. Sanderson leg. (BM) : 6
females. 1 nymph, 8(5)1 N, White mongoose (783 M) Old sec-
ondary forest, Bashan, Mamfe Division Cameroons 28.6.1933, P.
Sladen Trust Expedition, I. T. Sanderson leg. (BM) : 3 females,
Nandiniabinotata, [probably binotata binotata (Reinwardt)] high
deciduous forest, Mamfe, Cameroons, 30.4.1933. I. T. Sanderson
leg. (BM) : 2 females, "Schuppentier" (probably Phataginus) ,
Lolodorf, Africa, 29.3.1907 (GHFN coll: no number) : 1 female,
2 males, Genetta tigrina (subspecies not stated), Mongbivalu,
8/1939, Mme. Lepersonne leg. (MC 4502/4506) : 4 females (no
host data), Simba,- (MC 8259 and 8262) : 1 male (no host data)
Flandrina, 6.3.1928. R. P. Hubsbaert leg. (MC no other infor-
mation) : 1 female, Cercocebus albigena, Okongena (Lububu),
22.9.1929, A. Collart leg. (MC 42096), nymphs ?, antelope,
Masua, Lububu, 9.9.1929, A. Collart leg. (MC 43372/43386): 1
female, Aulacodus (=Thryonomys Fitzinger) swinderianus
Temminck 1827, Aruwimi, Panga, -8.1925, J. Schouteden leg.
496 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
(MC 8220 and 8222) : 1 female, Sus ? Leverville, 1927 (no other
data) (MC 46569): 1 female, Colobus badius badius (Kerr),
N'Dzida, Ivory Coast, 20.9.53. A. Villiers leg. (MCZ: 1 female,
Neotragus pygmaeus, Yapo, Ivory Coast, A. Villiers leg. (MCZ) :
1 male from Mongoose, Mt. Du Chaillu, Mt, Bijou, French Equa-
torial Africa, 8.8.1951. H. A. Beatty leg. (CNHM 73796) 2 fe-
males (no other data), OP coll. 2906 : 2 females and 1 male "man
and dog," Kumasi, Ashanti (GHFN 928) — this collection is a
bulk sample of 9 females and 1 male, i.e. we have no knowledge
of which specimens came from man or dog but the two rasus
forms are readily distinguishable from the 7 female pseudorasus
forms which are discussed later, with no intermediates : 3 females,
"Pangolin or Scaly manis, Mubango, Mabina Forest, Kyagle,
Uganda, 4000 ft, Capt, C. R. S. Pitman leg. (BM 6.19. 1-20) : 1
female, Lophuromys aquilus aquilus (True), Nyika Plateau,
Nyasaland, 9.10.1948. A. Loveridge leg. (MCZ) : 1 female, Manis
tricuspis {=Phataginus tricuspis (Rafinesque) ), Fernando Po.
(BM).
Distribution. From the records we have examined, Ixodes
rasus is to be found in many parts of West and Central Africa
and is common locally in East Africa. WEST AFRICA. Sene-
gal, Ivory Coast, Gold Coast; CENTRAL AFRICA. Fernando
Po, Cameroons, French Equatorial Africa (Bequaert 1931 re-
fers the type locality to French Equatorial Africa, not to the
Belgian Congo), Belgian Congo; EAST AFRICA. Uganda,
Nyasaland, Northern Rhodesia, Sudan. Previous records of I.
rasus from Southern Rhodesia (Nuttall 1916) refer to I. pseudo-
rasus and those of Cooley in the Rocky Mountain Laboratory,
Montana to Ixodes pilosus. The material of the records of Bed-
ford 1929 and 1932 are not available for re-examination.
Hosts. The wide range of hosts for the adults previously given
for /. rasus has now become considerably reduced (cf. Hoog-
straal, 1956). The immature stage-host relationships must re-
main in abeyance until such time as the larvae and nymphs are
bred and their diagnostic characters ascertained. The present
known hosts of the adults are: "wild pig," Felis pardus, Phata-
ginus tricuspis, Ccphalophus leucogaster, white mongoose, Nan-
dinia binotata, Genetta tigrina, Cercoeebus albigena, antelope,
Aulacodus (= Thryonomys) swinderianus, Colobus badius
ARTHUR : IXODES RASUS GROUP 497
badius, Neotragiis pygmaeus, dog, man, Lophuromys aquilus
aquilus, Hyrax (type specimen), Viverra civetta.
Biology. Unstudied.
Remarks. The Neumann collection at the ficole Veterinaire,
Toulouse, contains in addition to the electotype and the para-
type two lots of specimens identified hy Neumann as 7. rasus
and bearing the following data: (1) "Ousambara (Afr: or
Allem:) det. by G. Neumann 1900, 1 female," and (2) "7. rasus
Nn. Bismarckburg (Togo). Conradt leg. G. Neumann det. 1899.
Berlin Mus." 1 female. The first specimen is I. pseudorasus, the
second 7. oldi Nuttall 1913. Schulze (1943) indicated that cir-
cular anal grooves are characteristic of this species and in some
instances they may be drawn out or narrowed posteriorly. This
is in fact true for the electotype (see description). To what ex-
tent Schulze (1943) was justified in dividing rasus into three
subspecies is problematical. Schulze 's 7. rasus rasus undoubtedly
refers to Neumann's rasus and he adds little to Neumann's de-
scription of the male beyond directing attention to the lobes on
the ventral side of the basis capituli and the correction of the
position of the genital orifice. Similarly he gives the same
characters for 7. rasus eidmanni stating that they are "more
strongly chitinized (!) and darker," the denticles of the hy-
postome with a small apical hook (true also for 7. rasus), prox-
imity of sensory organs in the integument and absence of a
definite ' ' peripheral zone ' ' of the integument. These distinctions
would appear to me to be of doubtful value in establishing a
subspecies, particularly as the numbers examined were inade-
quate. This subspecies has been collected at Rio Muni and Span-
ish Guinea. I have no specimens of 7. rasus from either source
and the only specimens from Rio Muni constitute a distinct new
species (Ixodes muniensis) which is described later. I have
failed to see the original material of 7. cumulat imp unci atus and
have seen nothing in the extensive African tick fauna investi-
gated that is comparable with it. The occurrence of the "sichel-
haar" in the capsule of Haller's organ in 7. rasus (see Schulze
1943) is applicable to a large number of Ixodes ticks, and the
break in the chitin within the depression (trough), to which
Schulze alludes, is due to a failure to appreciate that the cuticle
498 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
A , 0. 5 MM.
ARTHUR: IXODES RASUS GROUP 499
in this region is saddle shaped in the majority (if not all) Ixodes
ticks (Arthur, 1956).
Rcdescription of the electotype FEMALE. Body well en-
gorged, dried specimen, dark red color, sclerotized parts dark
red brown.
Capitulum (Figures 1, 2). Length of basis capituli to hy-
postomal base, 0.43 mm., breadth of basis across dorsal ridge,
0.62 mm., sub-triangular, straight lateral margins slightly di-
vergent to palpal base, postero-lateral margin produced into
cornua which are broader basally than long, rounded apically;
well-defined posterior margin, straight and salient, black pig-
mentation peripherally (Figure 2). Surface gently convex with
reticulate sculpturing, lateral surface slightly curved. Porose
areas strongly depressed, sub-triangular in outline separated by
a distance equivalent to their greatest breadth. Basis capituli
broad ventrally ; auriculae as large strong blunt retrograde proc-
esses, anterior angle sharp, posteroventrally directed and stand
well out from the periphery of the basis. Distal part of ca-
pitulum in electotype broken off. Hypostome (Figure 7) 0.56
mm. in length, tapering apically, profile curved ; dentition from
base to apex, 1 row of 1/1 ; 4 rows of 2/2, 5 rows of 3/3, 4 rows
of 3/3, slight corona present, denticles long, hook-like. (Hypo-
stomal structure determined from females collected in Simba,
MC 8259 and 8262.)
Scutum. Broadly ovate (Figure 3) but widest in front of
mid-length, tapering more strongly to rounded posterior margin ;
colour, dark red-brown, surface reticulate. Cervical grooves as
wide depressions most pronounced about mid-way along; cervi-
cal field flat, surface is strongly elevated lateral to these grooves
but without indication of lateral ridge; punctations large, deep
and close together marginally, smaller and more widely sep-
arated elsewhere.
Figs. 1-7. Ixodes rasus, female: 1. Capitulum, ventral; 2. Capitulum,
dorsal; 3. Scutum; 4. Coxae and trochanters I-IV; 5. Anal plate; 5B,
diagram of anal plates in side view as drawn in Figure 5. Male: 6A. Tarsus
1; 6B. Tarsus IV; 7. Hypostome of female. (Scale A refers to Figs. 1,
12, 4-7; scale B refers to Fig. 3.)
500 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Legs. Moderately long and stout, distal segments missing
from first three legs on right side, similarly on second leg on left
side. Coxae slightly convex, reticulately patterned, coxae I, II
and III syncoxae, coxa I with distinct short bi*oad internal spur,
coxa IV with external spur, short, broad ; blunt tapering tro-
chanter spurs on legs I to IV (Figure 4) . Tarsi missing in electo-
type and paratype specimens. In specimens from Simba (MC
8259 and 8262), long, tapering with slight hump in front of
Haller 's organ on tarsus I ; similar humps on succeeding tarsi ;
length of tarsus I, 0.85 mm., metatarsus I, 0.45 mm. ; tarsus IV,
0.71 mm., metatarsus IV, 0.54 mm. (Figures 13A, B). Genital
opening, level with the third coxal interspace, genital apron
bilobed. Anal opening located far back on the body, convex anal
valves with two pairs of fine long hairs, anal grooves circular,
closed, and drawn out into a point in the electotype specimen
(circular in paratypes and other specimens examined) ; in Fig*
ure 5 the region bounded by groove is deeply sunk in front so
that the anal valves are almost vertical ; this sinking is less pro-
nounced posteriorly. Spiracular plate transversely oval, macula
central, goblets small, numerous. Body hairs fine, short, sparse.
Redescription. MALE. Elongate oval body, narrowing slightly
anteriorly, posterior extremity broadly rounded, length exclud-
ing basis, 2.56 mm., breadth 1.6 mm. Legs, scutum, dark red-
dish brown.
Capitulum (Figures 8, 9). Greatest breadth of basis capitidi
posterior to palpal insertion, 0.43 mm., much broader than long,
and converge by rectilinear and curved postero-lateral margins
to a slightly convex, salient posterior border; surface flattened,
brown in colour, bordered by darker band of pigmentation (stip-
pled in Figure 8) which does not extend to the periphery, small
scattered distinct pores. Palpi short, broad, length of article 2,
0.22 mm., article 3, 0.23 mm., greatest width of 0.19 mm. at
junctions of articles 2 and 3 ; lateral margins almost straight,
mesial profile of article 2 convex, that of article 3 straight, taper-
ing to broadly rounded apex; pronounced flanging effect ("roll
collar") along meso-dorsal edge of article 2, continued for some
distance along article 3 ; mesial surface of palp very slightly
concave ; hairs short to moderate in length, numerous, par-
ARTHUR: IXODES RASUS GROUP
501
ticularly on outer side, article 1 bears mesial spur. In ventral
view basis capituli b»oad, traversed by undulating ridge; mesial
lobe of which is more strongly convex than those on either side,
Figs. 8-12. Ixodes rasus, male: 8. Capitulum, dorsal; 9. Capitulum,
ventral; 10. Dorsum; 11. Venter; 12. Coxae and trochanters I-IV.
502 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
steep slope from ridge to hypostomal base, behind ridge surface
declivitous, broken by irregular ridges and particularly by two
sharp triangular elevations (Figure 9). (The hypostome and
chelicerae are missing, but Neumann (1899) gives the length of
the former as 0.55 mm.)
Scutum (Figure 10). Length 2.33 mm., breadth 1.33 mm.
Elongate, sides straight between the sharply angled antero-
lateral border and the broad rounded extremity ; surface convex,
dark reddish-brown colour, marginal fold white, of uniform
width, 0.175 mm. wide [Neumann (1899), repeated by Nuttall
et al. (1911) states that body fold is narrow (0.1 mm.)]. Cervi-
cal grooves faint anteriorly, leading into short, moderately deep,
wide depressions. P initiations of moderate size and depth, uni-
formly distributed, pronounced. Ventral plates (Figure 11) :
pregenital plate longer than broad, posterior and lateral borders
straight, anterior margin rounded; median plate large, diverg-
ing quite strongly to junction with adanal plates: sides and
posterior margin sinuous; adanal plates not joined behind anal
plate, latter circular but drawn out into a small but distinct
point behind, as in female; numerous punctations of similar
form to those on scutum. Abundantly supplied with short hairs,
which are shorter than those on the epimeral plates. Anus
eccentric, nearer anterior rim of anal groove.
Legs. Long, last two segments of fourth pair of legs extend
beyond posterior limits of body. Coxae (Figure 12). Large,
shiny, dark reddish brown, slightly convex and somewhat rugose,
such hairs as are present, long ; coxae 1 to III syncoxae ; coxa I
short, tapering spur on posterointernal angle, external spur short
and broad on coxae IV, coxae II and III unarmed ; distinct taper-
ing trochanter spurs on legs I to IV ; tarsi (Figures 6 A, B) much
as in female ; length of tarsus I, 0.79 mm. ; tarsus IV, 0.65 mm.
Ixodes pseudorasus new species
(Figures 14-31 j)
Ixodes rasus pro parte Nuttall, Warburton, Cooper and Robinson 1911, Pt.
II, pp. 229-30, Figures 225-226. Described females from cattle in
ARTHUR : IXODES RASUS GROUP
503
Uganda (GHFN 877d), from leopard, Obuasi, S. Ashanti (GHFN
503) and from man and dog (GHFN 928) — this last record is dis-
cussed under host list of /. rasas. It would appear that these writers
did not see the types of Neumann and state that their description
differs from that of Neumann (1899) in respect of the female.
Fig. 13. Ixodes rasus, female: A, Tarsus and metatarsus I; B, Tarsus
and metatarsus IV.
MATERIAL EXAMINED. Two females, goat, Beeba-Sharo,
Belgian Congo, 23.V.1913, F. Harker leg. (GHFN 2353) : 2 fe-
males, (no host cited) Ibembo, 6.7.50 (MC 70016) : 4 females,
dog, Port Franqui, -2.1934, Dr. Bouvier leg. (MC 38651) ; 1 fe-
male, Okapia, Epulu, 1933, R. Fr. Hutsebaut leg. (MC 46515/
46518) : 6 females, Okapia, Epulu, 1938, R. Fr. Hutsebaut leg.
(MC 46674/46662) ; 3 females, Okapia johnstoni, Epulu River,
Ituri District, Belgian Congo, P. Putnam leg. (BM) : 2 females,
Potamochoerus porcus, Ibembo, 6.7.1950, R. Fr. Hutsebaut leg.
(MC) : 1 female, Procavia ? Burunga, Kiru, -12.1925, Dr.
Schouteden leg. (MC 8178 & 8182) : 1 female, 3 nymphs, Tragel-
504
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Figs. 14-29. Ixodes pseudorasus and variant form; 14-15 female of /.
pseudorasus: 14. Capitulum, dorsal; 15. Capitulum ventral; 15A, B. Palpal
article 1, dorsal and ventral; 16, 17. Hypostome, I .pseudorasus and variant
form respectively; IS. Scutum (I. pseudorasus) ; 19, 20. Coxae and tro-
chanters, I.pseudorasus and variant form respectively; 21. Tarsus I (I.
pseudorasus) ; 22. Tarsus I (variant form) ; 23. Tarsus IV (I.pseudorasus) ;
24. Tarsus IV (variant form) ; 25. Genital apron (variant form) ; 26.
Genital apron (I.pseudorasus) ; 27, 28. Spiracular plate, I.pseudorasus and
variant form respectively; 29. Scutum (variant form). The letters A,D,P,V
refer to anterior, dorsal, posterior and ventral directions, respectively.
ARTHUR: IXODES RASUS GROUP 505
aphus scriptus, Kibombo, Belgian Congo, (MCZ) : 2 females,
Buffel, Ibembo, 2.8.1950, R. Fr. Hutsebaut leg. (MC 67307) : 1
female, cow, Costermansville, Vercommen 1950 (MC 61269) : 2
females, Potamochoerus porcus, Ibembo, 6.7.1950, R. Fr. Hutse-
bant leg. (MC 70016) : 2 males, 2 females, Cricctomys gambianus,
Mt. Selinda, Southern Rhodesia -12.55 (OP 2433 ii) : 3 females,
man, Mt. Selinda, Sth. Rhodesia, 23.9.55 (OP 2433 v) : 1 female,
5 nymphs, Chevrotain, north central Rio Muni, 23.4.1954, K. C.
Brown leg. B.22832, gift of R. Traub, (HH) : 1 nymph ? forest
antelope, north central Rio Muni, 18.5.1941, gift of R. Traub,
(HH) : 4 females, leopard, Obuasi, Ashanti, -12.1908 (GHFN
503), 7 females, man or dog? -10.1907, Dr. Graham leg. (GHFN
928) : 1 female, Gold Coast, no other data (Entomol. Research
Com. Cat. No. 762a. 14.4.1922. 8. BM) : 2 females, Cricetomys
gambianus, Bibianaha, Gold Coast, 3.12.1911. N. C. Rothschild
leg. (BM) : 6 females. Sierra Leone (no other data. BM) : 1 fe-
male, Cephaloplius, Tanga, German East Africa (BM) ; 2 fe-
males, Umboyasi River, Mgongo, British East Africa, no other
data (BM) : 1 female, cattle, which came from Bukedi to Mpumu,
Uganda; 10.9.1909, D. Bruce leg. (GHFN 877d) : 1 female, 10
nymphs, Bushbuck, Kyagwe, Uganda, N. W. Mettam leg. ( GHFN
3829a) : 1 female, giant rat, Mubango, Mambina Forest, Kyagle,
Uganda 4000 ft. alt., 1932, Capt, C. R. S. Pitman leg. (BM 6.19.
21) : 3 females, Centropus (?) superciliosus Loande, Kampala,
Uganda, -9.39. (BM) : 1 female, several nymphs, Pygmy Antelope,
Neotragus moschatus akeleyi, Mt. Kenya, 7000', Kenya : 9 females,
cattle, Kitale, A. Wiley leg. 5.4.49 (EAVRO) : 19 females,
Boocercus eurycerus, Kabolet Forest, near Kapenguria, 23.2.56,
S. F. Barnett leg. (EAVRO) : 3 females, Cricetomys, Mlange,
Nyasaland -.10.1914 (BM 13.12.30. 29-30) : 3 females, Mungos
melanurus, Zomboe, Nyasaland 1915 (BM 1915. 12. 30, 3/-33) :
1 female, 1 male "tick bird" stomach ? Tanganyika, (Vet. Lab.,
Kabete) : 1 female, Dakar, Senegal (No other data).
Distribution. The general pattern of the distribution of this
species follows closely that of I. rasus, except that as far as pres-
ent valid records are concerned there are far more records from
East Africa. WEST AFRICA. Senegal, Ivory Coast, Gold
Coast. CENTRAL AFRICA. Rio Muni, Belgian Congo. EAST
AFRICA. Uganda, Kenya (British East Africa), Tanganyika
(German East Africa), Nyasaland, Southern Rhodesia.
506 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Description. FEMALE. Body of unfed female short oval,
broadest at about the posterior third.
Capitulum (Figures 14, 15). Basis capituli sub-triangular,
posterior margin straight or slightly undulate, sides curved,
surface convex, broad, rounded, very short broad cornua ; porose
areas pear-shaped, less frequently oval, separated by a distance
less than their maximum diameter, when oval usually set
obliquely to long axis of body. Palpi long, lateral profile slightly
concave except for slight baso-lateral swelling, mesial profile of
article 2 broadly curved, that of article 3 straight, apex rounded
with inner angle acute, outer angle rounded, meso-dorsal margin
of article 2 and proximal part of article 3 flanged, palpal hairs
moderately long basally on article 2, shorter on article 3. Ven-
trally (Figure 15) basis capituli has straight lateral margins,
with postero-lateral and posterior margins broadly rounded, two
pairs of hairs in position indicated in Figure 15; surface either
flat or gently convex; auriculae form broad flat lobes with
rounded apices. Palpal article 1 (Figures 15 A, B) drawn
out into a mesodorsal flange and a ventro-lateral lobe supplied
with a long hair; article 2 with few hairs of moderate length
basally, hairs shorter and more abundant on article 3; inner
face of article 2 flat, that of 3 slightly concave. Length of capit-
ulum, 1.1 mm. ; breadth of capitulum across dorsal ridge, 0.48
mm. ; breadth of capitulum across auriculae, 0.57 mm. ; length of
palpal article 2, 0.47 mm. ; greatest breadth of palpal article 3,
0.16 mm.; length of palpal article 3, 0.37 mm. Hypostome (Fig-
ure 16) long, gently curved profile lines, rounded at the tip,
small "corona"; dentition, 2 rows of 4/4, 4 rows of 3/3, and 7
rows of 2/2 teeth.
Scutum (Figure 18). Length 1.42 mm., breadth 1.15 mm.,
widest in front of middle, curving strongly to scapular base, less
strongly posteriorly, sides almost rectilinear and terminate in a
rather narrowly curved posterior margin, scapulae short, pointed,
emargination slight; lateral carinae slightly indicated, short,
ceasing just beyond the greatest width; cervical grooves super-
ficial, shallow, not reaching to postero-lateral border. Punctations
consisting of closely set groups of small pores, uniformly dis-
tributed, short hairs.
Legs. Moderate length; coxae I, II and III syncoxae: coxa I
sub-triangular with well developed areae coxales supplied with
ARTHUR: IXODES RASUS GROUP 507
stout hairs, other hairs fine, variable in length, internal spur
lacking, the postero-internal extremity rounded, coxae II and III
longer than broad, supplied with hairs of uniform length, coxa
IV subtriangular, spurs absent; trochanters I— III with short,
broad, blunt spurs. Tarsi long, tarsus I, 0.74 mm., tarsus IV,
0.67 mm. (Figures 21, 23) tapering fairly gradually to slight
hump beyond Haller's organ, pad almost as long as claws.
Genital opening. Between coxae IV, covered with translucent
unilobed genital apron (Figure 26) genital grooves horseshoe-
shaped.
Anal grooves. Situated far back, rounded and drawn out
posteriorly into a small point.
Spiracular plate (Figure 27). Round, macula almost antero-
ventral in position.
Variants. The only consistent variant forms, which I have en-
countered are 9 females collected from "Vache, Costermansville,
Vercommen 1950, MC coll. 69733."
Description. FEMALE. These specimens are dark brown and
the alloscutum bears scattered short white hairs.
Capitulum (Figures 30, 31). Length 0.98 mm., breadth
across basis capituli just behind insertion of palpal article 1,
0.54 mm. ; surface flattened, porose areas subtriangular in out-
line, superficial, interporose interval about equal to maximum
breadth ; marginally basis capituli heavily pigmented, almost
black, lateral margins diverge anteriorly, posterior margin
straight, cornua short, broad basally and rounded apically;
palpi relatively short and broad, article 1 short, broader than
long, internal mesial spur present (Figure 30) ; article 2
swollen basally thence lateral profile concave to suture line with
article 3, that of article 3 nearly straight and terminating in a
broad rounded apex, mesially article 2 gently convex, article 3
straight and tapering to tip ; hairs long on lateral and mesial pro-
file of article 2, shorter on article 3; mesially, palp drawn out
into a broad flange and long hairs arise on its mesial face, i.e.
above and below the flange, length of article 2, 0.4 mm., article
3, 0.31 mm. Ventrally basis capituli broad, auriculae much re-
duced and form flattened projections (cf. with 7. pseudorasus) ,
508 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
surface generally flat as far back as slight posterior depression,
beyond which it rises strongly (stippled in Figure 31). Two
pairs of hairs in positions indicated in Figure 31 ; posterior
margin broadly rounded. Hypostome (Figure 17) long, rounded
apically, largest teeth about mid-length, strongly pointed, denti-
tion from base to apex, 7 rows 2/2, 4 rows 3/3, 2 rows 4/4, 2
rows 5/5 teeth, small "corona," median triangular unarmed area.
Scutum. Mean scutal dimensions are 1.3 mm. x 1.17 mm. (cf.
with I. pseudorasus, 1.4x1.1), and the scutum is broadly rounded
posteriorly (Figure 29), surface elevated between the short cervi-
cal grooves; lateral carinae ill-defined, antero-lateral margins
slope steeply, scapulae short, rounded, emargination slight;
punctations small, shallow, most abundant posteriorly but even
so well separated, less numerous in the cervical field and between
the cervical grooves and the lateral carinae.
Legs. Long and strong, coxa I with short, salient trenchant
internal spur, coxa IV with short rounded external spur (Fig-
ure 20), trochanter spurs short, broad, pointed; tarsus I (Figure
22) long, narrow, slightly tapering, strongly humped beyond
Haller's organ, tarsi II-IV with progressively weaker humps,
hairs progressively shorter and stouter from leg I-IV ; length of
tarsus I, 0.76 mm., tarsus IV (Figure 24), 0.67 mm.
Spiracular plate. Almost rounded, macula anteriorly placed,
large number of moderately large goblets (Figure 28).
Genital aperture. Between coxae IV : genital apron slightly
concave (Figure 25) (cf. with I. pseudorasus and I.rasus).
Anal grooves. Circular.
Description. MALE. From bovines, Sura, Arusha, Tangan-
yika, 13 Dec, 1955. F. W. White leg.
Body. Elongate oval, greatest width slightly in front of the
middle, surface more steeply rounded posteriorly. Colour (of
alcohol preserved specimens) alloscutum uniformly dark red-
dish brown, with many fairly long brownish white hairs es-
pecially on the periphery, hairs generally closely adherent to
surface, scutum and basis capituli deep red brown with paler
posterior and postero-lateral margins, legs and palpi less heavily
pigmented.
ARTHUR : IXODES RASUS GROUP
509
Capitulum (Figures 31 A, B). Basis capituli about 1-3 times
as broad as long, length 0.26 mm., breadth 0.34 mm., dorsal ridge
broadly convex, more sharply rounded postero-laterally and
thence to almost straight divergent margins, greatest width of
basis capituli immediately behind palpal insertions. Surface of
basis capituli reticulately patterned, elevated in mid-line behind
anterior cheliceral foramen, broad depression on each side run-
ning antero-laterally from posterior margin to palpal insertions,
Figs. 30, 31. Ixodes pseudorasus, variant form, female:
dorsal; 31. Capitulum, ventral.
30. Capitulum,
510 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
sloping away ventrally around base of latter, posterior margin
weakly elevated. Palpi broad, short, length to breadth as about
2.6:1.0, taper from base to broadly rounded apex, articles 2 and
3 of about equal length, about 0.21 mm., lateral profile of article
3 indented baso-laterally thence slightly sinuous to the suture
line between articles 2 and 3, that of article 3 almost straight;
mesial margin of article 2 as a broad arc of a circle, that of
article 3 straight and convergent to the rounded apex; greatest
breadth of 0.16 mm. just below suture line of articles 2 and 3,
hairs of moderate length and quite numerous apically. Ventrally
posterior border concave, postero-lateral angles steep, lateral
margins gently concave, diverging to palpal insertion; trans-
verse ridge strongly defined, with auriculae as faintly curved
and flange-like edges, connected by a median, broad, tongue-like
ridge ; basis capituli slightly longer than broad, less heavily pig-
mented than dorsal surface except for median tongue and
auricular elevations of the ridge and periphery; surface slopes
to hypostomal base from the transverse ridge, but short and de-
clivitous behind the latter; two triangular elevations on either
side of a depression behind, pair of erect hairs situated lateral
and posterior to hypostomal base, mesial surface of both palpal
articles 2 and 3 concave, that of 2 irregularly so. Hypostome
(Figure 31P) : broad, with indented apex, rounded on either
side of the indentation, 12 lateral teeth increasing in size from
apex to p re-basal teeth, basal tooth angular, teeth arranged as
2-3 rows of 3/3 files, 2 rows of 4/4 files, 4 rows of 5/5 files, pre-
basal and basal tooth subtended by two crenulations each ; length
0.23 mm.
Scutum (Figure 31G). Colour uniformly deep red brown,
with smaller less heavily pigmented patches antero-lateral to
scapulae. Elongate oval, tapering a little anteriorly, steeply
rounded posteriorly, greatest breadth approximately at mid-
length, length 1.93 mm., breadth 1.30 mm., distance between the
scapulae, 0.38 mm., antero-lateral margins behind scapulae un-
dulate, lateral and postero-lateral margins curved, lateral carinae
lacking, cervical grooves faint and very shallow, most apparent
in front of the pseudoscutal outline, weak behind the scapulae.
ARTHUR: IXODES RASUS GROUP
511
Figs. 31A-H. Ixodes pseudorasus male: A. Capitulum, dorsal; B. Capit-
ulum, ventral; C. Opisthosoma, ventral; D. Coxae I-IV; E. Tarsi I and IV;
F. Hypostome; G. Scutum; H. Spiracular plate.
512 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
Punctations numerous, subequal, shallow, small, more or less
evenly distributed over the surface. Scapulae broad based, short,
subtriangular with blunt apices, emargination moderate. Hairs
white, small, fairly numerous, uniformly distributed over the
surface.
Ventral plates (Figure 31C). Pregenital plate slightly con-
cave behind, convergent sides, rounded anteriorly, extends from
anterior margins of coxae III to posterior margin of coxae I ;
median plate about one and a third times as long as broad, great-
est breadth very near to posterior margin, sides curved and con-
vergent to genital opening ; adanal plates subrhomboidal with
sharp anterior angles, posterior junction between adanal plates
on either side ill-defined or absent ; anal plate transversely oval
and drawn out into a small point posteriorly.
Legs (Figure 31D). Broad, long, coxae I-III with extensive
syncoxal areas; all coxae large and broad, coxae I with strong,
conical internal spur, II and III unarmed, IV with smaller
rounded, broad external spur, several long hairs on each coxa,
with posterior margins of coxae I-III strongly trenchant, tro-
chanters I and II with large, broad-based spurs, III with a similar
but more flange-like spur, IV with smaller, more slender pointed
spur: tarsi (Figure 31E) long with slight hump, length of tarsus
I, 0.67 mm., metatarsus I, 0.32 mm., tarsus IV, 0.57mm., meta-
tarsus IV, 0.38 mm.
Spiracular plate (Figure 31H). Large, elongate oval,
broadly rounded in front, narrower behind, long axis parallel to
that of body, length 0.36 mm., breadth 0.22 mm., macula antero-
ventral.
Genital orifice. On a level with the anterior edges of coxae
III, genital apron slightly indented.
Hosts of Ixodes pseudorasus. Leopard, goat, cattle, bushbuck,
Neotragus mosckatus akeleyi, man, dog, Cricetomys gambianus,
Cephalophus, Mungos melanurus, giant rat, Okapia (which ap-
pears to be a frequent host in Belgian Congo), Centropus super-
ciliosus, Neotragus pygmaeus, Potamochoerus porcus, Tragela-
phus script us, "tick-bird," Chevrotain, forest antelope, bovines.
Biology. On larger mammals the adults occur on the ears and
thighs of hosts. Immature stages probably occur on small mam-
mals but until breeding experiments of these stages are completed
it is inadvisable to name possible hosts.
ARTHUR: IXODES RASUS GROUP 513
Related species. Confusion regarding the Ixodes rasus group
has arisen because of the acceptance of the closed anal groove as
the sole diagnostic character, which is based on the key prepared
by Nuttall, Warburton, Cooper and Robinson (1911). At pres-
ent we recognize the following five species having closed circular
grooves: I. rasas Neumann, I. pseudorasus, sp. nov., I. muniensis
sp. nov. 7. procaviae sp. nov. and I. thomasae sp. nov. The affini-
ties between rasus and pseudorasus are to be found only in the
form of the anal groove and the long palps. A comparison of
the descriptions and figures reveal that the differences in the
form of the auriculae, the spurs on coxae I and IV and in the
shape of the genital apron constitute distinctive characters. Both
species have a similar geographical range and if I. rasus and I.
pseudorasus are to be considered as variations within the species,
as suggested by Nuttall et al. (loc. cit.) then all grades of inter-
mediates would be expected. This is not so and out of 140 fe-
males examined 43 belong to I. rasus and 97 to I. pseudorasus
with no trace of intermediate forms. Whether the variants of
I. pseudorasus from Costermansville, all of which agree among
themselves, should have subspecific rank is problematical as only
nine females were available for examination, and far more col-
lecting for this tick is desirable.
Remarks. Nuttall, Warburton, Cooper and Robinson (1911)
base their description on females from Uganda (GHFN 877a),
Oubasi (GHFN 503) and from man and dog, Kumasi, Ashanti.
It would appear that these authors did not see the type and con-
cluded that their description of the female differed from that of
Neumann (1899).
Ixodes muniensis sp. nov.
(Figures 32—51)
Holotype. Female, from Cephalophus sp. "Mongele," Epulu
River, Ituri district, Belgian Congo, P. Putnam leg. Deposited
in Museum of Comparative Zoology, Harvard University.
Paratopes. Total 3 females, 9 nymphs and 8 larvae, all from
the same host as the holotype and deposited in the Museum of
Comparative Zoology, Harvard University.
MATERIAL EXAMINED. Total 13 females, 15 nymphs, 8
514 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
larvae. 1 female, 3 nymphs ex Duiker, north central Kio Muni
24 May, 1954, K. C. Brown leg., gift of Col. R. Traub to HH
No. B. 22842 ; 1 female, 1 nymph, tabby cat, north central Rio
Muni, 23 April, 1954, K. C. Brown leg., gift of Col. R. Traub to
HH No. B. 22836 ; 3 females, 2 nymphs, forest antelope, north
central Rio Muni, 10 June. 1954, K. C. Brown leg. gift of Col.
R. Traub to HH No. B22845; 1 female, Neotragus pygmaeus,
Yapo, Ivory Coast, A. Villiers leg. (MCZ) ; 1 female, Felis
pardus, primary forest, 12.5.1933, 500 ft. Mainyu Bridge, Mamfe,
Cameroons, I. T. Sanderson leg. P. Sladen Trust Expedition
(BM 1954 6.14.52) ; 2 females, tube No. 20 (no host data) ; Yoka-
douma, Cameroons, 30.5.55 (JM).
Description. FEMALE. Body, oval-oblong. Alloscutum
cream, few short curved white hairs in alcohol preserved speci-
mens, sclerotised parts medium brown, except for more heavily
pigmented basis capituli, maximum width slightly in front of
spiracle ; anal aperture near posterior border ; genital orifice
level with posterior edge of coxae IV, genital apron unilobed;
scutum reaches back beyond half the opisthosomatic length in
unfed or partly fed specimens.
Capitulum (Figures 32, 33). Basis capituli deep reddish-
brown, approaching black peripherally; posterior margin be-
tween cornua slightly convex, cornua large, sub-triangular,
rounded apices; lateral margins slightly divergent to greatest
width at palpal base; surface of basis reticulately patterned,
elevated in mid-line, slightly convex to antero-lateral margins
being more emphasized near the hypostomal base. Porose areas
indistinct, sub-triangular, shallow, inter-porose length about
equal to breadth of porose areas. Palpi long, rounded apically,
tapering most strongly distal to suture line between articles 2
and 3, lateral profile of article 2 swollen latero-basally, thence
gently concave to suture line ; mesial profile of article 2 convex
proximally, greatest breadth occurs about half-way along article
2, article I with prominent dorsally-directed flange-like pro-
jection (Figure 32), hairs long, fine, especially laterally, fewer in
number mesially. Ventrally basis capituli slightly longer than
broail, greatest breadth across auriculae, posterior border broadly
curved, lateral borders slightly constricted (Figure 33), auriculae
large, flat, broad-based, lobes, directed ventrally, pair of small
ARTHUR: IXODES RASUS GROUP
515
Figs. 32-38B. Ixodes municnsis, female: 32. Capitulum, dorsal; 33.
Oapitulum, ventral; 33A. Variation in the auricular form of the female;
34. Hypostome; 35. Scutum; 36. Coxae and trochanters I-IV; 37, Anal
plate; 38A. Tarsus and metatarsus I; 3SB. Tarsus and metatarsus IV.
516 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
hairs mesial to auriculae, similar pair posterior and lateral to
hypostomal base. Palpal article 1 produced into a prominent
pointed spur ventro-laterally and supplied with two hairs ; mesial
surface of article 2 flat, that of article 3 slightly concave. Hy-
postome (Figure 34), long, tapering, slender, apex rounded;
dentition from base to apex consists of six rows of 2/2, 6 rows
of 3/3, 2 rows of 4/4, small "corona.
> >
Measurements (in mm.) of 10 females
of I. muniensis
Length of capitulum from
Holotype
specimen
Average
Smallest
Large
dorsal ridge
0.31
0.31
0.30
0.32
Breadth along dorsal ridge
0.35
0.36
0.34
0.39
Breadth across auriculae
0.46
0.45
0.41
0.48
Length of palpal article 2
0.32
0.32
0.29
0.34
Length of palpal article 3
0.25
0.24
0.23
0.27
Scutum (Figure 35). Medium brown colour in alcohol pre-
served specimens; long, greatest width anterior of mid-length,
antero-lateral margins gently convex, postero-lateral margins
weakly concave before terminating in rounded posterior margin ;
lateral carinae as short ridged elevations not extending to the
margins, lateral field slopes gently from the ridge ; cervical
grooves shallow, becoming broadest about mid-length, not attain-
ing postero-lateral border, convergent at first then divergent
postero-laterally. Punctations equal, moderate-sized, distinct,
shallow, more numerous posteriorly than anteriorly, absent be-
yond carinae. Scapulae very short, broad, blunt, emargination
almost negligible, hairs few, scattered, longer and more numerous
anteriorly between cervical grooves.
Measurements of scutum of 10 females (in mm.)
Holotype
female Average Smallest Largest
Length of scutum 1.08 1.09 1.02 1.16
Breadth of scutum 0.76 0.80 0.72 0.83
ARTHUR: IXODES RASUS GROUP 517
Legs. Long, slender, coxae 1, IT, and III (Figure 36) with ex-
tensive syncoxal areas, internal angle of coxa I slightly trenchant,
internal spurs lacking; coxae IV, subtriangular, small postero-
external spur; hairs slender, white, prominent broad flange-like
saliences on trochanters II and III reduced salience on tro-
chanter I. Tarsi (Figures 38 A, 38B) relatively long and thin with
numerous hairs of moderate length.
Measurements of tarsi and metatarsi of 10 females (in mm.)
Holotype
female
Average
Smallest
Large;
0.62
0.59
0.55
0.62
0.31
0.30
0.29
0.32
0.53
0.53
0.51
0.55
Length of tarsus I
Length of metatarsus I
Length of tarsus IV
Length of metatarsus IV 0.35 0.34 0.33 0.37
Spiracular plate. Oval, long axis transverse to that of body,
macula slightly ventrally placed, dimensions, holotype female,
0.23 mm. x 0.18 mm., average size based on 10 females, 0.23 mm.
x 0.19 mm., smallest, 0.20 m. x 0.17 mm., largest, 0.24 mm x
0.21 mm.
Anal grooves (Figure 37). Circular, closed, ending in a slight
point behind.
Genital orifice. Level with posterior margin of coxae IV,
small, crescentic, unilobed genital apron.
Description. NYMPH. Body broadly rounded posteriorly,
narrowing anteriorly, greatest width just behind level of coxae
IV; in alcohol preserved nymphs alloscutum pale cream, with
curved white hairs of moderate length, legs, pale brown : anterior
half of scutum and palps more heavily pigmented but less so than
basis capituli, scutum reaches back to about one-third of opistho-
somatic length in unfed or partially gorged specimens.
Capitulum (Figures 39, 40A, 40B). Basis capituli twice as
broad as long, yellow-brown centrally, reddish-brown periph-
erally, dorsal ridge gently convex and extended to lateral points,
cornua lacking, antero-lateral margins sharply convergent to
palpal base ; surface of basis reticulately patterned and elevated
near dorsal ridge, gently sloping ventro-laterally, but becoming
steeper along the antero-lateral margins. Palpi relativly short,
article 2 (0.12-0.13-0.15 mm.) slightly longer than article 3
(0.095-0.11-0.12 mm.) ; tapering distinctly along length of article
518
BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
3, lateral profile strongly constricted basally, thence almost
straight to the suture between articles 2 and 3, greatest breadth
about mid-length of article 2, article 3 rounded apically ; mesially
article 1 has a small dorsally directed flange-like projection, hairs
long on lateral profile and basally on article 2, fewer in number
mesially, tuft of smaller hairs apically. Ventrally basis capituli
has straight posterior border, postero-lateral angles rounded con-
verging to well defined "waist," auriculae consist of bilobed flat-
jWgggggggjj
O.I MM.
0.5MM.
Figs. 39-45. Ixodes muniensis, nymph: 39. Capitulum, dorsal; 40a.
Capit ulum ventral; 40B. Auricula; 41. Scutum; 42. Hypostome; 43. Coxae
and trochanters I-IV; 44A. Tarsus and metatarsus I; 44B. Tarsus and
metatarsus IV; 45. Anal plate.
ARTHUR: IXODES RASUS GROUP 519
tened projections with rounded apices, directed postero-laterally,
with the more posterior lobe ventrally inclined ; a prominent
postero-laterally directed hair between the lobes, auriculae con-
nected by strong curved transverse ridge, becoming less elevated
in mid-line ; surface reticulately patterned, yellowish-brown cen-
trally, deeper reddish-brown peripherally. Surface convex behind
the ridge but flat in front, pair of moderate length hairs pos-
terior and lateral to hypostomal base. Mesial surfaces of both
palpal articles 2 and 3 flat. Hypostome. Length of hypostome
figured (Figure 42), 0.26 mm.; mean length, 0.24 mm., shortest,
0.22 mm., longest, 0.26 mm., greatest width about mid-length,
tapering to rounded apex, denticles broad, blunt, arranged in
8 rows of 2/2, 2 rows of 3/3.
Measurements of 15 nymphs (in mm.)
Average Smallest Largest
Length of eapitulum from dorsal
ridge 0.13 0.12 0.15
Breadth along dorsal ridge 0.2<i 0.26 0.31
Breadth across auriculae 0.29 0.25 0.34
Scutum. Length, 0.47-0.53-0.57 mm., breadth, 0.55-0.60-
0.63 mm., colour pale to medium brown ; transversely ovate, an-
terolateral margins converge to scapulae in slight convexity with
minor undulations, postero - lateral margins behind greatest
breadth converge more strongly to broadly rounded posterior
margin; lateral carinae as slight straight elevations fading into
general elevation of scutum anteriorly and widening postero-
laterally, lateral field slopes gently peripherally ; cervical grooves
shallow, widest about mid-length, fading posteriorly and not
attaining postero-lateral margins, divergent in front : scapulae
minute, broad, blunt, emargination weak. Punctations few in
number, small and shallow of equal size, most abundant posteri-
orly ; few short white adpressed hairs, most dense on lateral fields.
Legs. Moderate size, coxa I (Figure 43) with moderate sized
pyramidal pointed external spur and longer conical pointed
internal spur, coxae II, ITT and IV with successively decreasing
pyramidal, pointed spurs, few hairs of varying lengths on each
520 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
coxa, small flange-like saliences on trochanters I-III. Tarsi (Fig-
ures 44A, 44B) relatively short, broad, with slight hump distal
to Haller's organ on tarsus I ; length of tarsus I, 0.37 - 0.41 - 0.44
mm., metatarsus I, 0.17 - 0.18 - 0.19 mm. ; tarsus IV, 0.29 - 0.34 -
0.36 mm., metatarsus IV, 0.18 - 0.22 - 0.23 mm.
Spiraculab plate. Oval, longitudinal axis transverse to that
of body, macula ventral and slightly anterior of center, dimen-
sions : average 0.16 mm. x 0.14 mm., smallest 0.12 mm. x 0.11 mm.,
largest 0.17 mm. x 0.17 mm.
Anal grooves. Oval, long axis parallel to that of body, closed,
circular terminating in a slight point behind (Figure 45).
Description. LARVA. Fully fed larvae almost elliptical in
shape, in alcohol preserved specimens alloscutum black, few
scattered short, curved white hairs, legs and palpi pale brown,
basis capituli medium brown, scutum yellow or dark brown with
yellowish margins, scutum reaches back to less than a quarter of
opisthosomatic length in fully fed specimens, greatest width
about mid-length, anal grooves closed, circular.
Capitulum. Total length of capitulum, 0.19 mm., breadth
across dorsal ridge, 0.15 mm. Basis capituli twice as long as
broad (Figures 46, 47) dorsal ridge convex, drawn out to lateral
points, no cornua ; lateral margins short, strongly convergent to
palpal bases; surface of basis capituli reticulately patterned,
elevated into two prominent mounds mesial of palpal bases, de-
pressed in mid-line behind hypostome, gentle slope antero-lat-
erally, becoming steeper along rostral margins: Palpi short,
article 2 (0.065 mm.) a little longer than article 3 (0.058 mm.),
lateral profile of article 2 indented basally, thence gently con-
vex to apex, mesially article 2 almost straight and article 3
terminates in a fairly acute rounded apex, long hairs present
laterally, fewer mesially; mesial surface of articles 2 and 3 flat.
Ventral view (Figure 47) posterior margin of venter of basis
capituli slightly curved, posterolateral angles gently rounded,
constricted laterally ; auriculae as in nymph, each lobe flattened
ventrally, subtriangular with blunt apices, directed more or less
laterally, breadth across auriculae, 0.13 mm., strong curved
transverse ridge, auriculae elevated above posterior extension of
basis capituli: surface reticulately patterned; anterior to trans-
verse ridge, pale brown, behind ridge more heavily pigmented.
Hypostome (Figure 49) short, approximately parallel-sided,
ARTHUR: IXODES RASUS GROUP
521
rounded apex with broad "corona," succeeded by 1 row of 3/3,
6 rows 2/2, 1 row 1/1 ; length 0.13 mm.
4Gg25MM* — -47
0.25 MM.
Figs. 46-51B. Ixodes muniensis, larva ; 46. Capitulum, dorsal ; 47. Capitu-
lum, ventral; 48. Scutum; 49. Hypostome; 50. Coxae I-III and trochanters
II-III; 51A. Tarsus and metatarsus I; 51B. Tarsus and metatarsus III.
522 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
Scutum. Colour variable in alcohol preserved specimens, e.g.,
black, yellow, black with yellow margins — black with paler color-
ation anteriorly : about as broad as long, length 0.29 mm., breadth
0.29 mm., greatest breadth at about mid-length, antero-lateral
margins curve gradually to almost negligible scapulae, lateral
margins rounded, lateral carinae very slight elevations close to
antero-lateral margins; cervical grooves shallow, widest about
midway along, narrowing appreciably behind and not reaching
postero-lateral margins. Punctations very few, equal, indistinct,
small and shallow, confined to posterior and lateral portions of
scutum ; specimens examined glabrous.
Legs. Moderate length and thickness, coxa I (Figure 50) with
pointed, horn-like external spur, smaller conical internal spur,
few long hairs on coxae ; small flange-like saliences on trochanters
I and III, largest on II. Tarsi (Figures 51A, B) with several
long hairs, tarsus I broad, humped, length of tarsus I, 0.22 mm.,
length of metatarsus I, 0.1 mm. ; length of tarsus III, 0.20 mm.,
length of metatarsus III, 0.1 mm.
Anal aperture. Fairly near posterior border; anal grooves
closed, circular.
Belated species. I. mnniensis, like /. rasus and I. pseudorasus,
is not a host specific tick and appears from the limited data
available to be a West African species. This new species is
readily recognizable from other species of the rasus group by the
characters which appear in the key (p. 535). Males are unknown.
Hosts. Cephalophus, duiker, tabby cat, forest antelope, Neo-
tragus pygmacus, Felis pardus.
Ixodes procaviae new species
(Figures 52-72)
Holotype. Female, from Dendrohyrax adolfifriederici, Kisenyi,
Belgian Congo, 6 September, 1949. Deposited in the Rocky
Mountain Laboratory, Hamilton, Montana, U. S. A. Number
(25) 27839.
Allotype. Male, no host or locality data, Rocky Mountain
Laboratory, Hamilton, Montana, U. S. A. Number 26006.
Paratypes. 2 females, 1 male, 1 nymph. Data as for allotype
ARTHUR: IXODES RASUS GROUP 523
and all deposited in the Rocky Mountain Laboratory, Hamilton,
Montana, U. S. A.
OTHER MATERIAL EXAMINED. Total, 15 females, 2
males, 1 nymph. 3 females, Procavia adolfifriederici Brauer,
Burunga, Belgian Congo, H. Schouteden leg. (MC) ; 5 females,
Procavia ?, Burunga, Kini, -.12.1925, H. Schouteden leg. (MC
8178-8182) ; 1 female, Phacochoerus sp., Bururi, 1.6.1949, J.
Francois leg. (MC 68842) ; 1 female, Dendrohyrax arboreus,
Kisenyi, Ruanda, J. Deom leg. (MC 72375) ; 2 females, Rift Val-
ley Province, Kenya (HH collection).
Description. FEMALE. Elongate oval, tapering more strongly
anteriorly, maximum width just in front of spiracle ; anal aper-
ture situated far back, scutum reaches back beyond half opistho-
somatic length. Alloseutum dark grey in alcohol preserved speci-
mens, with white hairs closely applied to the body.
Capitulum. Basis capituli average length from dorsal ridge
to rostral base 0.30 mm. (holotype 0.26 mm., range 0.26-0.32
mm.), average breadth across dorsal ridge, 0.37 mm. (holotjrpe
0.37 mm., range 0.34-0.39 mm.) ; dorsal ridge straight between
cornua, latter well-developed, relatively slender, subtriangular
with blunt salient apices; lateral margins carinate, slightly di-
verging anteriorly to level of palpal insertion ; surface reticu-
lately patterned, flat except for pronounced lateroventral curva-
ture in the rostral region. Porose areas distinct, of moderate ex-
tent, piriform, moderately depressed, separated by an interval
about equal to their maximum breadth (Figure 52). Palpi, long,
slender about five times as long as the greatest width; apex
broadly rounded with more acute curvature mesially; mean
length of article 2, 0.40 mm. (holotype, 0.42 mm., range 0.39
mm. -0.42 mm.) ; article 3, 0.25 mm., (holotype, 0.26 mm., range
0.23 mm. -0.26 mm.) ; lateral profile of article 2 strongly in-
dented baso-laterally, thereafter distinctly concave to the suture
line between articles 2 and 3, article 3 with straight or slightly
convex lateral outline ; mesial profile of article 2 convex, that of
3 almost straight ; article 1 broader than long with small, pointed
dorsally directed spur, situated close to meso-dorsal margin;
bristles few, moderate to long. Basis capitidi more heavily pig-
mented ventrally than dorsally, posterior border straight with
prominent, rounded postero-lateral extensions (Figure 53), lat-
eral margins slightly constricted behind auriculae, latter short,
524 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
flattened backwardly-directed horns narrowing to their tips and
directed postero-ventrally ; palpal article 1 possessing a saddle-
shaped spur adjacent to antero-lateral margin of basis capituli,
two fine white hairs arise dorsal to this spur; mesial surface of
article 2 flattened, that of 3 slightly indented. Hypostome (Fig-
ure 54), average length 0.59 mm., (holotype, 0.60 mm., range
0.57 mm. -0.60 mm.), tapering slightly from base to apex which
is domed, denticles long, slender becoming slightly shorter
towards the mid-line, dentition from base to apex as 2 rows of
2/2 files, 3 rows of 3/3 files,, 3 rows of 4/4 files, 6 rows of 5/5
files surmounted by a small "corona."
Scutum. Colour brown, elongate oval in shape with greatest
width in front of mid-length, average length, 1.24 mm., (holo-
type, 1.25 mm., range 1.18 — 1.27 mm.), average breadth, 0.86
mm., (holotype, 0.88 mm., range 0.80—0.89 mm.). Antero-
lateral margins converge to broad, blunt, short scapulae (Figure
55) by a few minor undulations, behind greatest width conver-
gent margins either rectilinear or slightly concave and termi-
nate in broadly rounded posterior extremity. Lateral carinae
as slight, straight elevations which are most pronounced about
three quarters of their length posteriorly, lateral field slopes
strongly from the carinae. Cervical grooves strongly convergent
posteriorly before becoming divergent for the greater part of
their length, shallow and narrow at first subsequently broaden-
ing, do not attain postero-lateral borders. Punctntions small to
medium-sized, distantly spaced and more numerous posteriorly
than anteriorly. Hairs few, irregularly dispersed but most fre-
quent anteriorly between the cervical grooves.
Legs. Long. Coxae I, II and III with small syncoxal areas
(cf. Ixodes muniensis), trenchant behind, prominent sharp pos-
tero-internal spur on coxa I which may or may not reach to or
slightly overlap the anterior margin of II, postero-internal angle
of II forms a marginal salience, long tapering external spur
on coxa IV ; coxae each supplied with a variable number of long
white hairs. Each trochanter supplied with long, sharp spurs
(longer than indicated in Figure 56). Tarsi with dorsal surface
converging slightly to Haller's organ (Figure 57A), and followed
ARTHUR : IXODF.S RASUS OROFP
525
Figs. 52-58. Ixodes procaviae, female: 52. Capitulum, dorsal; 53. Capitu-
lum, ventral; 54. Hypostome; 55. Scutum; 56. Coxae and trochanters I-IV ;
57A. Tarsus and metatarsus I; 57B. Tarsus and metatarsus IV; 58. Anal
plate.
526 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
by a moderate sub-apical hump, the latter is less well defined on
tarsi II-IV, hairs becoming stronger and shorter from tarsi I to
IV; average length of tarsus I, 0.61 mm. (holotype, 0.62 mm.,
range 0.57 mm. — 0.63 mm), metatarsus I, 0.32 mm. (holotype,
0.32 mm., range 0.31 mm. — 0.32 mm.); tarsus IV, 0.51 mm.,
(missing in holotype, range 0.49 mm. — 0.52 mm.) ; metatarsus
IV, 0.34 mm. (missing in holotype, range 0.33 m. — 0.35 mm).
Spiracular plate. Slightly ovate, long axis transverse to that
of body, macula slightly anterior of center; dimensions, mean,
0.25 x 0.21 mm., holotype, 0.23 x 0.20 mm., range, 0.22 — 0.26
mm. x 0.19 — 0.23 mm.
Anal groove. Oval, closed and in some specimens terminates
in a small point, long axis parallel with corresponding axis of
body (Figure 58).
Genital orifice. On a level with posterior margin of coxae
IV; narrow, long crescentic.
Description. MALE. Elongate oval, but tapering quite strong-
ly in front with a broad posterior extremity. Length from tip of
palpi to posterior body margin 2.13 mm. ; greatest body width at
the level of the spiracles 1.25 mm., body flat. Colour of alcohol
preserved specimens, medium brown alloscutum, sclerotized parts
golden brown.
Capitulum. Overall length from dorsal ridge to palpal tips
0.49 mm., greatest breadth behind palpal insertion, 0.31 mm.,
breadth across dorsal ridge, 0.22 mm. Basis capituli (Figure 59)
broader than long, mid-region golden brown, black peripherally,
posterior margin straight, postero-lateral angles rounded, sides
divergent to base of palps; surface irregularly sculptured, con-
vex medianly behind cheliceral "foramen," otherwise flattened
except for a slight elevation along the posterior ridge, few scat-
tered small pores. Palpi, short, broad, globular, about twice as
long as the greatest breadth; dorsally article 2 slightly longer
than article 3 (as 0.15 mm. is to 0.14 mm.), with the greatest
width at the apex of article 2; apex broadly rounded; lateral
margin of article 2 swollen basally then either straight or gently
concave to suture, profile of article 3 straight, mesial margin of
articles 2 and 3 broadly convex; hairs of moderate length, fine
particularly laterally, two long strong hairs arise mesodorsally
and basally from article 2. Ventrally (Figure 60) basal margin
either straight or faintly concave, lateral margins concave, di-
ARTHUR: IXODES RASUS GROUP
527
Figs. 59-66B. Ixodes procaviae, male: 59. Capitulum, dorsal; 60. Capitu-
lum, ventral; 61. Scutum; 62. Venter; 63. Coxae and trochanters I-IV ;
64. Spiracular plate; 65. Hypostome; 66A. Tarsus and metatarsus I; 66B.
Tarsus and metatarsus IV.
528 BULLETIN: MUSEUM OP COMPARATIVE ZOOLOGY
vergent ; auriculae as small, raised lips linked by a transverse
ridge to form a broad, tongue-shaped eminence in the midline,
gentle slope from the ridge anteriorly; steeper, sharper slope
posteriorly ; one pair of small white hairs — one on each side
lateral to and behind the hypostomal base, second pair of hairs
on ridge between the auriculae and the median tongue-shaped
eminence ; mesial surfaces of articles 2 and 3 slightly concave.
Hypostome (Figure 65) short (0.22 mm.), broad, tapering, and
indented apically. Dentition consists of at least 7 rows of crenu-
lations arranged in 4/4 files.
Scutum. Colour, medium to dark brown. Scutal surface very
slightly convex centrally, gradually downcurved laterally, long
oval in shape (Figure 61) but narrowing more strongly in front
than behind, length 1.69 mm., breadth 1.05 mm. Cervical grooves
short, not extending back to the level of the faintly indicated
pseudoscutum, slightly converging and thence widely divergent,
shallow — narrow initially before broadening. Punctations nu-
merous, shallow, small, fewer in number anteriorly. Scapulae
short, broad sharp ; emargination moderate. Hairs few, short,
white more abundant posteriorly than anteriorly.
Legs. Moderate length and breadth. Coxae I, II and III with
relatively small syncoxal areas, coxa I with distinct broad taper-
ing postero-internal spur, coxa II with short internal spur
mesially and forward of the postero-internal position, coxa IV
with broad-based conical external spur; coxae I and II trenchant
behind. Several hairs of varying length on all coxae. Tro-
chanters I, II and III with prominent conical spurs, that on IV
less pronounced — these spurs are not as strong as the com-
parable ones in females. Tarsi humped, broad, with moderate to
long hairs ; length of tarsus I, 0.54 mm., metatarsus I, 0.25 mm. :
tarsus IV, 0.46 mm., metatarsus IV, 0.30 mm. (Figures 66A, B).
Spiracular plate. Elongate oval (Figure 64) long axis paral-
lel to that of body, goblets numerous, macula anteroventral ;
dimensions 0.20 mm. x 0.28 mm.
Ventral plates. Pregenital plate with straight posterior edge,
lateral margins curved and converging anteriorly; median plate
large, much wider behind than in front, lateral and posterior
ARTHUR : IXODES RASUS GROUP 529
margins curved (see Figure 62) ; adanal plates almost rhoru-
boidal, sides adjacent to anal plate incompletely concave; anal
plate almost egg-shaped with anterior edge flattened ; small trans-
verse subrectangular plate between genital orifice and pregenital
plate.
Genital orifice. Wide, located between coxae 111.
Description. NYMPH (described from a single specimen from
lot RML — -26006). Body shape as in female except that the
greatest width occurs across coxae IV; has some of the salient
characters of the female.
Capitulum. Length, 0.30 mm., breadth across auriculae 0.20
mm., breadth across dorsal ridge, 0.245 mm., oasis capituli about
twice as broad as long, dorsal ridge slightly concave, and pro-
duced to salient lateral projections, which mark the greatest
width, lateral margins very short and sharply convergent to
palpal insertions ; surface reticulately patterned, gently convex
except for a slight elevation along the dorsal ridge, and strong
slope peripherally. Palpi long, club shaped, length of article 2,
0.16 mm., length of article 3, 0.14 mm., article 3 stouter than
article 2, greatest breadth 0.05 mm. Article 2 swollen basally
for a short distance thence concave to the rounded palpal apex,
mesial profile of articles 2 and 3 broadly convex ; article 1 with
small mesodorsal spur, hairs, long, particularly laterally. Ven-
trally, posterior edge of basis capituli straight (Figure 68),
posterolateral edges sharply rounded, sides slightly constricted;
auriculae small flange-like lobes, not particularly elevated above
surface, more heavily pigmented than rest of basis ; pair of small
hairs placed posterior and lateral to hypostomal base, one long
curved white hair on each side in front of auriculae ; palpal
article 1 with small elongate saddle-shaped spur, mesial surface
of article 2 flat, that of 3 concave. Ilypostome length, 0.26 mm.,
profile curved, apex rounded, external teeth larger than internal
teeth, dentition from apex to base as 5 rows of 3/3 files, 6 rows
of 2/2 files (Figure 71).
Scutum. Colour, brown beyond cervical grooves, paler in
between latter and posteriorly. Large, nearly as broad as long ;
length 0.54 mm., breadth 0.52 mm., greatest breadth about one-
530
BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
Figs. 67-72B. Ixodes procaviae, nymph: 67. Capitulum, dorsal; 68. Capit-
ulum, ventral; 69. Coxae and trochanters I-IV; 70. Scutum; 71. Hypostome;
72A. Tarsus and metatarsus I; 72B. Tarsus and metatarsus IV.
ARTHUR: IXODES RASUS GROUP 531
third distance from scapulae, latter short, broad, blunt; antero-
lateral margins almost rectilinear, convergent to scapulae, at
greatest width sharply angled (Figure 70) thence by slightly
sinuous postero-lateral sides to broadly rounded posterior ex-
tremity ; lateral carinae as minor elevations, becoming wider and
losing height as lateral margins reached, short, straight; lateral
field slopes gently from the ridge ; cervical grooves convergent,
indistinct and shallow anteriorly, then diverging, widening but
not attaining postero-lateral margins. Punctations distinct, few,
small, shallow, most numerous behind greatest breadth; hairs
short, white, most prominent anteriorly and laterally.
Legs. Of moderate length and thickness. Coxa I with sharp
pointed, conical, internal and external spurs (Figure 69), coxae
II and III Avith moderate-sized, round-ended, pyramidal, ex-
ternal spurs, coxa IV, with broad-lobed external salience ; pos-
terior margin of coxa I and postero-external angle of coxa IV
trenchant. Trochanter spurs on all coxae, that on I being small
and conical, on II-IV broad, flange-like. Tarsi (Figures 72A,
B) taper gradually from proximal to distal end, slight pre-apical
hump on I, becoming less well defined from II to IV; length of
tarsus I, 0.36 mm., metatarsus I, 0.15 mm. ; tarsus IV, 0.28 mm.,
metatarsus IV, 0.17 mm.
Spiracular plate. 0.12 mm. x 0.10 mm., long axis transverse
to that of body, macula almost centrally placed.
Anal groove. Oval, slightly open behind, longitudinal axis
parallel to that of body.
LARVA. Unknown.
Related species. The females of procaviae and thomasac agree
in the form of their auriculae and the spurring of coxae I and
1 1 . apart from the closed anal groove. The postero-lateral exten-
sion of the venter of the basis capituli together with the long,
tapering trochanter spurs make it readily distinguishable from
other members of the rasus complex of species.
Hosts. Dendrohyrax adolfi-friederici, Dendrohyrax arboreus,
Procavia, Phacochoerus sp. (Names of hosts given here are as
on data enclosed with the specimens in the tubes.)
Remarks. At present known only from the Belgian Congo,
Uganda and Kenya.
532 BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY
Ixodes thomasae new species
(Figures 73-79)
Holotype. Female, Host : HH 4093, Arvicanthis abyssinicus
nubilans, Njoro, Rift Valley Province, Kenya, 7500 ft., 14 June,
1948, II. Hoogstraal leg. Deposited in Rocky Mountain Labora-
tory, Hamilton, Montana.
Paratype. One female, Host : HH 4106, Otomys tropicalis el-
gonis, Njoro, Rift Valley Province, Kenya, 7500 ft., 15 June,
1948, H. Hoogstraal leg. Deposited in British Museum (Natural
History).
Description. FEMALE. Body shape of unfed female elongate
oval, narrowing anteriorly, broadly rounded posteriorly, greatest
width just behind the spiracle; Colour, sclerotized parts dark
brown, alloscutum reddish brown in alcohol preserved specimens,
well covered with white adpressed hairs of moderate length :
Anal aperture far back. Scutum extends well beyond half opis-
thosomatic length.
Capitulum. Overall length from cornua to palpal tips 0.85
mm., breadth across dorsal ridge, 0.38 mm., dorsal ridge straight
between cornua, latter small, broad-based with blunt apices, lat-
eral margins slightly convex, divergent almost to level of palpal
insertion, which is the widest point (Figure 73) ; surface reticu-
lately patterned depressed around and in front of the porose
areas, inter-porose area and surface adjacent to the dorsal ridge
elevated, strong ventral curvature near antero-lateral borders.
Porose areas subtriangular, slightly depressed, inter-porose in-
terval less than the maximum breadth. Palpi long, about five
times as long as the greatest width ; lateral profile distinctly con-
cave, with mesial convexity more or less parallel to it, apex
rounded ; length of article 2, 0.39 mm., article 3, 0.26 mm. ; article
1 with small dorsally-directed flange-like spur arising adjacent
to lateral margin of basis capituli : latter nearly one and a half
times as broad as long. Ventrally (Figure 74) basis capituli
more heavily pigmented than dorsally, reddish-brown pigmenta-
tion peripherally, surface reticulately patterned, flat, except be-
hind the auriculae and the antero-lateral margins which curve
away dorsally; posterior border broadly curved, postero-lateral
edges sharply rounded, lateral margins gently and slightly con-
stricted ; greatest breadth (0.45 mm.) across auriculae, latter
ARTHUR: IXODES RASUS GROUP
533
well developed, tapering horn-like spurs which are flattened on
their ventral surfaces, horizontal and directed postero-laterally ;
Figs. 73-79. Ixodes thomasa.e, female; 73. Capitulum, dorsal; 74. Capitu-
lum, ventral; 7.1. Scutum; 76. Coxae and trochanters I-IV; 77. Hypostome;
78. Tarsus and metatarsus I. 79. Anal groove.
534 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
pair of hairs posterior and lateral to hypostomal base; palpal
article I with saddle-shaped spur from which a long white hair
projects postero-laterally, mesial surface of article 2 flat, that of
3 slightly concave. Hypostome (Figure 77) long (0.57 mm.),
narrow, tapering to a pointed apex, outer file of denticles larger
than those nearer the mid-line and arranged from base to apex
as 1 row of 1/1 files, 7 rows of 2/2 files, 6 rows of 3/3 files, sur-
mounted by a small corona.
Scutum. Colour, brown ; large, rhomboidal, longer than broad
(1.32 mm. x 0.88 mm.) with greatest breadth in front of mid-
length ; antero-lateral margins faintly undulate and convergent
to scapulae, postero-lateral margins straight, converging to pos-
terior convexity. Lateral carinae distinct, broadening postero-
laterally and reaching the margins just behind greatest width,
lateral field beyond carinae sloping strongly. Cervical grooves
weak anteriorly but on diverging become deeper and wider de-
pressions about mid-length, do not extend to the periphery.
Punctations medium size, moderate depth and uniformly dis-
tributed. Glabrous in holotype.
Legs. Small to moderate syncoxae on coxae I-III (Figure 76),
short conical internal spur on coxa I, large external spur on
coxa IV, all coxae trenchant; few hairs of varying sizes on all
coxae. Trochanters I-III bear well developed broad rounded
saliences, that on IV reduced to a smaller spur. Tarsi long, taper-
ing with a distinct hump beyond Haller's organ (Figure 78).
Spiracular plate. Oval, with long axis transverse to corres-
ponding axis of body, macula antero-ventral, dimensions 0.26 x
0.22 mm.
Anal groove. Circular and drawn out to a small point pos-
teriorly, closed (Figure 79).
Genital orifice. Between coxae IV, genital apron large, oval,
unilobed.
This species is named for Miss Dilys G. Thomas of the Ad-
ministrative Staff of King's College, London, who has exhibited
the greatest patience as well as a high degree of efficiency in deal-
ing with our continual typing requirements on acarological
matters.
Related species. See the key on page 535 and the information
on page 531.
ARTHUR: IXODES RASUS GROUP 535
Hosts. Arvicanthis abyssinicus nubilans, Otomys tropicalis
elgonis.
Remarks. Known only from Kenya.
Key to the known species of African Ticks allied to Ixodes rasus
Neumann by virtue of their possession of a closed anal groove.
(Characters mentioned are sufficient to serve as a diagnosis of
new species described herein.)
Females
1. Auriculae as sharp retrograde spurs Ixodes rasus
Auriculae otherwise 2
2. Coxa I lacking internal spurs 3
Coxa I with internal spur 4
3. Cornua about as long as basal breadth, auriculae straight edged, directed
postero-ventrally, coxa IV with slight external spur I.muniensis
Cornua shorter than basal breadth, auriculae lobed, directed laterally,
coxae IV without external spur I.pspudorasus
4. Auriculae lobed as in pseudorasus Variant of I. pseudorasus
Auriculae tapering nearly to a point, directed postero-laterally ....5
5. Posterior margin of venter of basis capituli extended laterally, tro-
chanter spurs long, pointed. Hypostomal dentition 3/3 files for nearly.
the whole length I.procaviae
Posterior margin of venter of basis capituli not extended laterally,
trochanter spurs as broad rounded saliences. Hypostomal dentition 3/3
for distal third I.thomasat
Males
(The males of I.muniensis and I.thomasae are not known.)
1. Coxae II with short internal spur mesial and forward of the postero-
internal position, basis capituli relatively narrow I.procaviae
Coxa II lacking such spurs, basis capituli broad 2
2. Syncoxal areas on coxae II and III extensive, scutum with steadily
convex margins, projections on transverse ridge not well developed ....
I .pseudorasus
Syncoxal areas on coxae II and III nut extensive, scutum with rec-
tilinear margins, projections on transverse ridge well developed and
large I. rasus
536 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY
SUMMARY
1. The male and female of Ixodes rasus of Neumann (1899)
are redescribed and the systematic status of the female of
the species bearing this name, as described by Nuttall et al
(1911), is critically reviewed. After examination of a large
number of ticks we have arrived at the conclusion that the
female of rasus, as described by Nuttall et al should be as-
signed specific rank as pseudorasus. No such great variation,
as suggested by them, occurs in the specimens we have
studied.
2. The subspecific forms of I.r.cumulatimpunctatus and I.r.eid-
manni of Schulze (1943) have not been observed, and we con-
sider that in certain respects his subspecific characters are in-
adequate for diagnostic purposes.
3. The male of pseudorasus is described and a variant of the
female of this form noted.
4. Three other new species having closed, circular, anal grooves
are described. These are I.muniensis (female, nymph, larva).
I.procaviae (female, male, nymph) and I.thomasae (female).
5. All species appear to have a wide range of potential hosts.
REFERENCES
Arthur, D. R.
1 9 .""» 6 . The morphology of the British Prostriata with particular refer-
ence to Ixodes liexagonus Leach, III. Parasitology, 46: 261-307.
Bequaert, J.
1931. Synopsis des Tiqucs du Congo Beige. Rev. Zool. Bot. Afr., 20:
209-251.
HOOGSTRAAL, H.
1956. African Ixodoidea Vol. I. The ticks of the Sudan. Dept of the
Navy. Bureau of Medicine and Surgery. Pp. 11101.
Neumann, G.
1899. Revision de la Famille des Ixodides. 3. Mem. Soc. Zool. France,
12: 107-294.
Nuttall, G. H. F.
1911. On the adaptation of ticks to the habits of their hosts. Para-
sitology, 4: 46-67.
ARTHUR: IXODES RASUS GROUP 537
Xtttall, G. H. F.
19U3. Notes on ticks, IV. Relating to the genus Ixodes and including
a description of three new species and two new varieties.
Parasitology, 8: 294-337.
Nuttall, G. H. F., C. Warburton, W. F. Cooper and L. E. Robinson
1911. The genus Ixodes. A monograph of the Ixodoidea, Pt. II.
pp. xix + 105-348. Cambridge University Press.
ScHUIiZE, P.
1943. lieber zwei bemerkenswerte afrikanische Schwesterarten von
Ixodes: I.rasus Neum. und vanidicus n.sp. Zool. Anz., 142: 121-
141.
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