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Full text of "Bulletin of the Museum of Comparative Zoology at Harvard College"

4 8^0 



HARVARD UNIVERSITY 

Library of the 

Museum of 

Comparative Zoology 



BULLETIN 



OF THE 



MUSEUM OF COMPARATIVE ZOOLOGY 



AT 



HARVARD COLLEGE, IN CAMBRIDGE 



VOL. 116 



CAMBRIDGE, MASS., U. S. A. 
1957 



TnK Cosmos Press, Inc. 
Cambkidiif,, Mass., U.S.A. 



1\< 



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 



Publications Issued by or in Connection 
with THE 



MUSEUM OF COMPARATIVE ZOOLOGY 
AT HARVARD < OLLEGE 



t 



Bulletin (octavo) 1863 - - The current volume is Vol. 115. 

Breviora (octavo) 1952 — No. 70 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. 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 ; Pt s 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, R 2 is forked and R 1? R 3 , R 4 , R g are simple, usually 
one crossvein occurring between Rj and R 2 . 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 R g and M 1 + 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 
R 5 and M x and at the place where Cu x 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, R 2 is forked, and R x , R 3 , R 4 , R 5 are simple; 
usually one crossvein between R x and R 2 . 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 Zoolog3 r . 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 brow T nish, 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 Cu x ; 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 R s (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. 

Mab j 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 R 2 is simple, no cross- 
vein between R 3 and R 2 . 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; R 2 a forked into R 2a i 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 Ro a 
of this species as compared with the forked R 2a 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. 



■ o v 



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, Ro a forked into Ro al 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, Ro a 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 R 2a vein of cavaleriei is forked 
into lUai an d 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; R s 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 Cu 2 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 R s 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 Cu 1} one at the base of R s , one at the subcostal 
erossvein (Scv) and one at the first fork of R s ; 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 R s 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 R s , one at subcostal crossvein (Scv) and one 
at the first fork of R s ; besides these, there is a very feeble spot 
at the fork of R 4 + 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 Cu 2 and 1A absent; pterostigma not 
prominent, connected with R s 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 Cu 1} one at the base of R 8 
and one at the first fork of R s ; 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 R s 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 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 R s 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 R s , one at the first fork of R s , 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 R s 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 Cu 1? one at the 
base of R s and one at the first fork of R s ; 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 R s 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 Cu 1} one at the base of R s , one 
at the first fork of R s , 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 R 5 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 R s 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 R s and 
one at the first fork of R s ; 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 R s 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 R s , one at the first fork of 
R p , and one at ending of Cu 2 ; 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 Cu 1 ; passing the 
fork of M 3+! , 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 R 2 +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 R s and one at the first fork of R s ; 
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 E s 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 j r ears 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 (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 Cay T 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, 3y 2 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 



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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. 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 2 g_2 7 , 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 1 g_^ 7 in the male of 338 mm., -fipnr ^ n ^ ne f ema -l e °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 aA T ailable 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 botanical 1 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 


5V 2 


Monte 


Libano 


20.7 


37.3 


29.3 


19.8 


16.8 


sy 2 


1 1 


t i 


21.5 


37 


28 


18.3 


17 


5V 2 


i 1 


1 1 


21.5 


36 


28 


18 


17 


5y 2 


1 1 


i 1 


20.8 


35 


27 


18 


16 


5y 3 


1 i 


1 1 


21 


35 


27 


16.6 


15.2 


5V 3 


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 


5V 5 


1 1 


< < 


24.7 


38.7 


30.4 


19.8 


18.8 


5y 2 


El Censo to L. Ocotal 


21.2 


37.6 


27 


18.4 


16.2 


sy 2 


t < 


< < 


22.8 


37.3 


27 


18.6 


16.7 


5y 2 


t < 


< < 


20.5 


38 


30 


18 


15.7 


sy 2 


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 6y 2 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 Miller 1 

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 (a m )- 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. 



o J 



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 I T niv. 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 2y 2 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 Sy 2 . 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. L T . 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 



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with THE 

MUSEUM OF COMPARATIVE ZOOLOGY 
AT HARVARD COLLEGE 



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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- 
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The continuing publications are issued at irregular intervals in numbers 
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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 teeth 1 ; 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 









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 row T 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 
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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 shagreening 1 
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 New T 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 three 2 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 



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. 

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1948 — Published in connection with the Museum. Publication terminated 
with Vol. 24. 

The continuing publications are issued at irregular intervals in numbers 
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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 39 i 

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; R x to R 5 stalked; Mi from upper angle of the middle 
cell; M 2 and M 3 shortly stalked, connate or slightly separate, 
from the lower angle of the middle cell; Cuj from well before 
the angle of the middle cell ; Cu 2 from more or less behind the 
middle point of the middle cell ; A 2 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; M 2 and Cii! connate 
or shortly stalked, from the lower angle of the middle cell ; Cu 2 
from cell near three fourths, remote from Cuj ; A 2 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 (m 2 ) is in the middle cell. A more or less long 
spot (nil + m 3 °f the Amata wing pattern scheme) is below 
the middle cell, in the interspace between it and the vein A 2 ; 
an ovate or more or less elongate spot (m 4 ) is in the basal part 
of the interspace between the veins R 5 and M x ; two spots (m 5 
and m 6 ) are in the basal parts of the interspaces between the 
veins M 2 and Chi].. These last two spots are separated from one 
another by the vein M 3 . Some smaller extra spots are often 
present ; they occur near the basal parts of the interspaces over 
the vein R 5 and below the vein M 1; also at the outer angle 
formed by the vein Cu 2 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 Cu 2 and A 2 . This spot- 
usually crosses over the vein A 2 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 Cvl x and Cu 2 
and the middle cell. This spot usually crosses over the vein 
Cuj and reaches to the vein M 2 . 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 Cu x and Cu 2 . A dark ray 
along the vein M 2 usually connects the forewing borders with 
the discal patch. In the hindwing these borders are mostly 
dentate at the veins Cu! and Cu 2 . 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. (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, w r edge-shaped spot (m 2 ) in the 
middle cell; a long spot (m 14 ., ) below it which extends much 
farther outward than the distal edge of the spot in the middle 
cell; a long spot (m 4 ) between the veins R 5 and M t , with 
two longish extra streaks over these veins ; tw r o much broader, 
egg-shaped spots (m 5 and m 6 ) between the veins M 2 and 
Ciij separated from each other by the black vein M 3 ; 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 Cu 2 , out- 
ward from the long basal spot (m 1+;; ). 

The hindwdng with a large basal spot which extends from the 
middle cell to the vein A 2 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 (m 1+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 (m 2 ) 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 (m 4 ) between the veins R 3 and M 1? with 
two much smaller, elongate extra spots over these veins ; two 
rather short, egg-shaped spots (m 5 and m 6 ) between the veins 
M 2 and Cu x separated from each other by the black vein M 3 ; 
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 m 4 and m 5 contiguous with both; a little extra spot 
above the base of the vein Cu 2 ; 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 (m 2 ) at 
the end of the middle cell; an elongate spot (m 1+3 ) below it 
reaches about to a point on a level with the middle of the middle 
cell spot; a rather narrow, elongate spot (m 4 ) between the 
veins R 5 and M x accompanied by two short, hyaline streaks 
above and beneath; two shorter but broader, egg-shaped spots 
(m 5 and m 6 ) between the veins M 2 and Cui separated from 



406 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY 

each other by the black vein M 3 ; 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 (m 2 ) 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 (m 4 ) above the 
base of the vein M 1; accompanied by a small, slightly elongate 
extra spot below this vein, and sometimes also by a little dot 
above the base of the vein R 5 ; two spots (m 5 and m 6 ) in the 
interspaces of the veins M 2 and Cu ls egg-shaped, dilated toward 
outside, the lower of them slightly broader and longer, separated 
from each other by the black vein M 3 ; sometimes a little, ovate, 
extra spot above the base of the vein Cu 2 . 

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 A 2 . 

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 m 4 . Most of 
the specimens examined from Kuatun are like the type, others 
have extra spots above and beneath the spot m 4 . Sometimes 
another extra spot, on the outer side of the f orewing spot mu 3 
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 M 2 from the discal spot to the black wing borders ; 
these latter with a truncate tooth between the veins Cuj and 
Cu 2 ; 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 3 r ellow 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 M 2 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 Cu 2 ; 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 



BULLETIN : MUSEUM OP COMPARATIVE ZOOLOGY 



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 M 2 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 Cu 2 . 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 
M 2 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) 



416 



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 Cu 2 ; a broad, black discal spot sending a 
ray along the vein M 2 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, w T ith 
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 tw T o 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 M 2 and between the veins Cuj 
and Cu 2 ; discocellulars with a more or less broad, black spot ; 
along the vein M 2 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 (m 1+3 ) below the middle cell extends nearly 
from the wing base to the tornus; a wedge-shaped spot (m 2 ) 
in the middle cell ; a more or less large extra spot above the base 
of the vein Cu 2 ; three larger, elongate spots (m 4 to m 6 ), between 
the veins R 5 and M t and M 3 and Cu x form (together with two 
smaller and narrower extra spots above and beneath the spot 
m 4 ) 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 Cu 2 ; 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 Cu 2 . 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\i 1 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, $ l 4 Ao, 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 Cu 2 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 
(m 1+3 ) below the middle cell; a short, wedge-shaped, some- 
times diffusely blackish scaled spot (m 2 ) in the middle cell; an 
exterior series of spots consisting of more or less short rec- 
tangular spot (m 4 ) between the veins M x and R 5 accompanied 
by an extra spot above and a little, inconstant extra spot below; 
two inward pointed spots (m 5 and m 6 ) between the veins M 2 
and Cu 1} and an extra spot between the veins Cu x and Cu 2 
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 M 2 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 M 2 connecting this 
spot with the wing borders ; a border tooth along the vein Cu 2 ; 
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 
(m 2 ) in the middle cell, usually as broad as this latter; a long 
spot (m 1+3 ) below it, reaching from the wing base almost to the 
tornus, or the basal part of this spot (mj) absent; a spot (m 4 ) 
in the area between the veins R 5 and M x mostly shorter than that 
(m 6 ) between the veins M 3 and Cuj ; a spot (m 5 ) between the 
veins Mo and M 3 often the smallest in the exterior spots series ; 
sometimes more or less developed extra spots above the vein 
Cu 2 and at both sides of the spot m 4 . 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. 

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 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." 
During 1 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 w T as 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