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California Academy of Sciences 





]Jr. Ro))ert ('. Miller, Chairman 
Dr. G. P. Papenfiiss Dr. Edward Ij. Kessel, Editor 



No. 1. YoNGE, C. M. Observations on Hipponix antiqmitus (Lin- 
naeus) (9 text figures). Published July 15, 1953 1-24 

No. 2. Gressitt, J. LiNSLEY. The California Academy-Lingnan 
Dawn-Redwood Expedition (13 text figures). Pub- 
lished July 15, 1953 25 58 

No. 3. EssiG, E. 0. Some New and Notewortliy Aphidae from 
Western and Southern South America (Hemiptera- 
Homoptera) (56 text figures). Published July 15, 
1953 59-1 64 

No. 4. Orr, Robert T. Natural History of the Pallid Bat, An- 
trozous paUidus (LeConte) (28 text figures). Pub- 
lished January 7, 1954 165-246 

No. 5. Sasa, Manabu, and E. W. .Jameson, Jr. The Trombi- 
eulid Mites of .Ta])an (45 text figures). Published 
January 7, 1954 247-321 

No. 6. Ijoukashkin, Anatole S., and Norman Grant. Fnrther 
Studies of the Beliavior of the Pacific Sardine (/SVn-- 
(linops caerulea) in an Electrical Field (5 text fig- 
ures) . Published .luly 9, 1 954 323-337 

No. 7. TjOukashkin, Anatolk S., and Thomas C. Groody. On 
fhe Pacific Sardine {Surdinops caerulea Girard) in 
Acjuaria : Trans])ortation, Handling, jVIaintenance, 
and Survival (3 text figui'es). Published -June 30, 
1955 339-353 

No. 8. Slevin, Joseph R. Notes on Australian Amphibians (28 

text figures). Published June 30, 1955 355-392 

No. 9. Fuller, Varden. Population Growth in the West and 
its Impact on Natural Resources. Published Feb- 
ruary 17, 1956 393-404 

No. 10. lIuBERTY, Martin R, Water Resources of the West, To- 
day and Tomorrow. Published February 17, 1956. . . 405-414 

No. 11. Wyckoff, Stephen N. Forests of the Future. Pub- 
lished February 17, 1956 415-423 

No. 12. Van Cleve, Richard. The Conservation and Future De- 
velopment of West Coast IVIarine Resources. Pub- 
lished February 17, 1956 425-439 

No. 13. Fennah, E. G. Fulgoroidea from Southern China (26 

text figures) . Published February 21, 1956 441-527 

No. 14. Slevin, Joseph R., and Alan E. Leviton. Holotype 
Specimens of Reptiles and Amphibians in the Collec- 
tion of the California Academy of Sciences. Published 
September 21, 1956 529-560 

No. 15. Green, John Wagener. Revision of the Nearctic Spe- 
cies of P/foh'ims (Lampyridae: Coleoptera) (19 text 
figures). Published November 19, 1956 561-613 

No. 16. Miller, Loye. A Collection of Bird RemaiiLS from the 
Pliocene of San Diego, California (1 text figure). 
Published November 26, 1956 615-621 

Index to Volume XXVIII 623 

Errata 656 




Fourth Series 
Vol. XXVIII, No. 1 July 15, 1953 





University of Glasgow 


One of the most interesting of the commoner gastropods fonnd on the 
shores of California is the small Hipponix antiquatus. It is a member of 
one of the many groups of limpets which have independently been evolved, 
but it is of particular interest owing to its sedentary habits. While it must 
move about in early life, at some stage, at present unknown, //. antiquatus 
settles permanently. It then proceeds to lay down calcareous matter on 
the surface of the underlying rock. This cemented "ventral valve" may 
eventually attain much the same size and thickness as the true shell. 
Hippo7iix may be described as a univalve which converts itself into a bi- 
valve, but it should be noted that the valves are dorsal and ventral, resem- 
bling those of a brachiopod, not those of a lamellibranch. 

Despite their interesting ha])its little seems to liave been written about 
these animals. The only paper found which deals with habits is one on 
H. austraUs. This species lives usually on the shells of species of Turbo 
and was studied at New Caledonia by Risbec (1935). The opportunity 
was taken while visiting professor in the University of California in 1949 
to examine, first at Berkeley and later at Pacific Grove, living specimens 

'Being "Studies on Pacific Coast Mollusks, Vll" (nos. I-VI, published in University of California Publica- 
tions in Zoology, vol. SS, pp. 395-454). 



of //. (tntiquatus wliicli were o])tained from Moss Beach, a little south of 
San Francisco, and from the shores of the Monterey Peninsula. In view 
of previous work on feeding in the allied Capulus ungaricus (Yonge, 
19:^8 ) and on the ])allial organs in other limpets (Yonge, 1947), special 
attention was i)aid to the mode of feeding and to the nature and mode 
of functioning of the organs in the mantle cavity. Some specimens w^ere 
fixed in I^ouin's fluid and were later sectioned, or mounted entire, in 
the DeparfnuMit of Zoology, University of (}lasgow\ But this paper deals 
l)rimarily with observations on the living animal and no attempt has 
been made to give a detailed description of structure. 

This work was made possible by the kind cooperation of colleagues in 
the Department of Zoology, University of California, Berkeley, and 
of Dr. R. L. Bolin and members of the staflf of the Hopkins Marine Sta- 
tion, Pacific Grove. At Glasgow, Dr. H. F. Steedman gave great help by 
preparing whole mounts and cutting sections. 

Appearance and Habits 

Hipponix antiquatus is the conunonest of several species of its genus 
recorded from the coast of California (Keep and Bailey, 1935; Smith and 
Gordon, 1948). It ranges from south of the equator to about 42° N. lati- 
tude (Keen, 1987). It has a rounded, much flattened shell. As shown in 
figure lA, the apex is not central but lies well to the posterior side, and 
the outer surface is grooved. The transverse diameter of the shell is usually 
a little greatei' than the antero-posterior diameter, the specimen shown 
in figure 1 being 1.7 cm. by 1.6 cm. No larger specimen was found and 
it was also unusually regular in siiape, many shells being distorted owing 
to the confined space in which the animal had originally settled. 

The shell is smooth internally and the impression of the horseshoe- 
shaped shell muscle is clearly marked on the surface of both the shell and 
of the "ventral valve" ])elow (fig. IB and C). The latter is formed by 
the e]nthelium of the under surface of the foot and in the same way as 
the shell, i.e., the pedal margins increase it in extent while it is contiiuiaIl>' 
being thickened by secretion from the general surface of the foot. Even- 
tually it comes to have much the same maximum thickness as the shell, 
about 1 nun. It is internally concave and to about the same de])th, in 
this case some 4 mm., as the shell (cf. fig. IB and C). The only difference 
between the two is that the greatest de])th in the "ventral valve" is cen- 
tral and not ])osterior as it is in the shell. 

Smith and Gordon (1948) state that this si)ecies occurs "In colonies 
under rocks at low tide; common." It was sought with greatest success 
in narrow, often overhung, crannies in the rock from mid-tidal levels and 


Figure 1. Hiijponix antigiuilus. photographs of shell. XIVl-- A, shell in situ 
on rock; B, interior of "ventral valve" secreted by ventral surface of foot, scars of 
shell muscle shown: C, interior of "dorsal valve," i.e. true shell secreted by mantle, 
scars of shell muscle shown. 

below. It is there protected from the full force of the Pacific surf while 
the movenieuts of these waters continually brini>' in new supplies of organic 
debris and detritus of all kinds. The animals always appear to occupy 
rounded depressions in the rock surface. These have frequently, if not 
always, been made initially by rock-boring bivalves, which are extremely 
numerous in this rock. It is possible, however, that the depression may 
be further excavated during growth, presumably l^y means of the shell 
margins. But this process must cease as soon as calcareous nuiterial be- 
gins to be secreted ventrally. 

Ventral Aspect 

The appearance of an animal after careful removal from the rock 
by cutting through the attachments to the cemented "ventral valve" is 
shown in figure 2, the specimen being viewed to some extent from the 
anterior end so as to obtain a better view of the head and the more an- 
terior organs in the mantle cavity. Like the majority of limpets, e.g., 
both the archaeogastropod Patellaeea (Docogiossa) and the pulmonate 
Siphonaria (Yonge, 1947, 1952), the shell muscles (SM) unite pos- 
teriorly although in Hipponix the connexion is very narrow. Laterally, 
however, the muscles are very broad and they provide extensive attach- 
ment between the upper and lower "valves." Contraction pulls the shell 
firmly against the "ventral valve" and, as this has been laid down while 
subject to the constraining influence of the shell, it follows that the 
margins of the two "valves" make perfect contact. This ensures protec- 



Figure 2. Hipponix antiquatus, appearance in life, viewed from antero-ventral 
aspect. X5. A, anus; C, ctenidium; DD, digestive diverticula; E, exlialant current; 
HG, hypobranchial gland; I, inhalant current; 0. osphradium; P, proboscis; SM, 
shell muscle; m, p, anterior margins of mantle and of foot, forming dorsal and 
ventral surfaces of mantle cavity. Feathered arrows indicate direction of cleansing 

tion against enemies and also against the force of the sea and the danger 
of desiccation. But it has only been achieved at the expense of mobility. 
The various "homing" limpets, e.g., species of such very widely separated 
genera as Patella and Siphonaria, have retained the power of locomotion 
and yet achieved an equal measure of protection because the margins of 
the shell make perfect contact with the rock surface of the "home." 

The mantle cavity is bounded dorsally by the mantle (w) but ventrally 
by the membranous extent of the foot {p) which stretches forward be- 
tween the shell muscles which form the lateral walls of the cavity (see 
also fig. 5). Tlie head may project out of the mantle cavity, as it is 
shown doing in figure 2, but can be withdrawn into this by contraction 
of asymmetrical retractors (figs. 4 and 7; LR, RR). The head extends 
into a conspicuous proboscis (P) which is flanked by a pair of stout ten- 
tacles each with a small eye on the dorsal surface near to the base. 


The characteristically sessile habit of the adult of Hipponix is asso- 
ciated with major modifications of the foot. While, as in all limpets, 
the shell muscles are very large, the central region of the foot, which 
in other limpets forms the greater part of the creeping sole, is here de- 
void of nuiscle. It consists solely of a thin membrane much of which 
forms the floor of the mantle cavity. This is best indicated by the sections 


shown ill figures 5 and 7. The ventral surface of the foot extends peri- 
pherally so that it covers the same area ventrally as the mantle does dor- 
sally and its epithelium has a similar capacity for secreting shell. It is 
as though the animal had two mantle lobes, dorsal and ventral, but no- 
w^here connected. Secretion of calcareous matter by the pedal epithelium 
may possibly represent some elaboration of the original powers of mucous 
production. The "ventral valve" so formed is cemented firmly to the 
underlying rock. As noted above, the shell muscles serve as adductors 
drawing the free ''dorsal valve" tightly against the attached "ventral 
valve " 

At the base of the proboscis ventrally there is a small flap of tissue 
which projects forward from the mesopodial tissue below. This is the pro- 
podium (figs. 4, 5 and 7A; PR); it is terminally notched and a small 
gland opens on its dorsal surface. It appears to ])e concerned solely with 
attachment of the egg capsules in which connexion it will lie mentioned 
again later. 

Figure 3. Hipponix antiquatus. dorsal view after removed from shell, based 
on examination of living animals and of dissections; course of alimentary canal 
indicated by broken lines. X5. C — C. extent of ctenidium (dotted where viewed 
through the mantle wall); M. mouth; MG. mid-gut; OD. opening of oviduct; OE. 
oesophagus; PC. pericardium; R. rectum; ^'.'s'. style-sac; ST. stomach. Other let- 
tering as before. 


[Puoc'. 4th Sku. 

Mantle Cavity 

Aftoi' removal from the sliell, the animal may be viewed from the dor- 
sal aspect, as shown in fi<>iire 3. Owing to the forward extension of the 
shell muscles and the conse(iuent constriction of the opening of the mantle 
cavity, the anns (A) has lieen dis])laced dorsally, 0])ening sligiitly to the 
right of the mid-line. The anterior end of the ctenidinm (C) is similarly 
carried round dorsallv. terminating just to the left of the anus. 

y fj 

Figure 4. Hipponix antiqnatus, dorsal view of organs in the mantle cavity 
after opening along the right side. X5. A', kidney; LR. left retractor muscle of 
head; PL. plate where egg capsules (stalks only shown) are attached; PR. propo- 
dium; RP. renal pore; RR, right retractor muscle of head, x — x. cut ends of mantle 
edge; y — y. cut surfaces, posterior wall of mantle; z — z, cut ends of mid-gut. Other 
lettering as before. Plain arrows indicate respiratory current between ctenidial 
filaments, feathered arrows cleansing currents. 

The mantle cavity can best be examined after cutting along the edge 
of the shell muscle on the right side and then turning the roof of the 
mantle cavity over to the left, giving the appearance shown in figure 4. 
The point of immediate interest is the asymmetry due to dorso-ventral 
compression in Hipponix. In arehaeogastropod limpets, such as Acmaea 
or Patella (Yonge, 1947), and also in the mesogastropod Ca})nUi.^, whicli 
is closely related to Ilipponir, the head occupies the center of the numtle 
cavity. This is a consequence of secondary symmetry associated witii loss 
of coiling in t!ie shell and visceral mass. Tn all of these limi)ets height is 
seldom less than breadth, it is often greater. Tn Hipponix, on the other 
hand, height is always less than half the breadth. As a result of this and 


of the reduction of the pedal tissues (with which this coiiiprcssion is 
also associated), the mantle cavity extends relatively far back, as shown 
in longitudinal section in figure 5. But it also widens out internally and 
is very shallosv. Hence the head, although it projects forward out of 
the middle of the opening of the mantle cavity (figs. 2 and 3), has basally 
been pushed over to the left (figs. 3 and 7) where the oesophagus (OE) 
runs into the visceral mass. This accounts for the much greater size of 
the right as compared with the left retractor muscle of the lu'ad (figs. 4 
and 7B; KR, LR). 

A])art from this flattening and extension jiosteriorly, the mantle 
cavity is that of a typical pectinibranch prosobranch, having the same 
general disposition of the pallial organs as, for instance, in Biiccinum 
(Yonge, 1938). The pericardium, as revealed by dissection and in sec- 
tions, is situated far to the left at the base of the ctenidium (figs. 3 and 
70; PC). The large kidney (fig. 4, K) covers much of the posterior wall 
of the mantle cavity, the renal pore (RP) opening to the left of the 
mid-liiu'. Internally it is unusually capacious, as shown in figure 5. It 
communicates wnth the pericardium by way of a long reno-pericardial 
canal (figs. 5 and 70; RC). On the right side of the cavity extends the 
elongated genital aperture which, in all the specimens that were dis- 
sected, was oviducal (figs. 3 and 4 01)). The rectum (R) meanders along 
the right side of the roof of the cavity to open, near the margin of this, 
at the anus ''A). The large pectinibranch ctenidium (C) occupies the 

Figure 5. Hipponix antiquatus. longitudinal section. Xll. B. blood sinus; D. 
duct into digestive diverticula; GS. gastric shield; A', internal cavity of kidney; 
MC. mantle cavity; ODT. odontophore cartilage; RC. reno-pericardial canal; S'/S', 
style-sac with contained style. Other lettering as before. 

left side of the mantle cavity twisting over to the dorsal surface an- 
teriorly as mentioned above. Parallel to its axis on the left and so facing 
the inhalant current (fig. 3, I) extends the linear osphradium (figs. 3; 4; 


ami 7A, 0). Between the ctenidiuni and the rectum, tlie roof of the cavity 
is covered by tlic extensive tissues of the liyjiobranchial iiland (fi^s. 4; 5; 
and 7, IIG)." 

CiLiARv Currents 

The hiteral cilia on the ctenidial filaments create an inhalant cur- 
rent (I) wliicli enters the cavity on the left side, impinging first on the 
osphradium. The exludant current, as always, leaves the cavity on the 
right. The ctenidium is concerned solely with creating this current and 
with respiratory exchange. As shown in figure 4, the filaments are broad, 
like those of a typical pectinibranch. Where the filaments are modified 
in connexion with ciliary feeding, they are invariably elongated, e.g. in 
Vcnnctus novac-hoUandiae, Capulus ungaricus and, to a striking extent, 
Crepidula foniicata (Orton, 1912, Yonge, 1938). This elongation in- 
creases both lateral and frontal surfaces and so the extent of the lateral 
cilia, which create a greater inhalant current, and of the frontal cilia 
which are here concerned with food collection. In Hipponix there is only 
a moderate inhalant current, adequate for the limited respiratory needs 
of the animal, while the frontal cilia retain their primitive function 
of cleansing. 

In ciliary feeding species, moreover, collected particles are conveyed 
fo the mouth along the tips of the elongated filaments, and by way of 
special food grooves, to the mouth. This has been described in various of 
the Vermetidae (Yonge, 1932, 1938; Morton 1951&), in TurritcUa com- 
munis (Graliam, 1938), in the Struthiolariidae (^Morton, 1951rt) — al- 
though not in the related Aporrhais which has the same habit of burrowing 
in mud (Yonge, 1937) — in Crepidula and other members of the Calyp- 
traeidae (Orton, 1912, Yonge, 1938), in Capulus ungaricus (Yonge, 1938), 
and in the freshwater Viviparus viviparus (Cook, 1949). 

Careful observation in Hipponix showed that there is no passage of 
particles along the tips of the filaments or within a food groove to the 
mouth. The circulation of water and the disposal of waste particles in 
Ihe mantle cavity is essentially as in typical pectinibranchs. As de- 
scribed elsewhere (Yonge, 1938), there are three currents concerned 
with rejection of sediment, (A) those carrying heavier particles to the 
inhalant opening; {B) those carrying medium particles across the jloor 
of the mantle cavity; (C) those carrying the finest particles over and 
between the ctenidial filaments for later consolidation dorsalljf in the 
mucus from Ihe hypol)ranchial gland. Material in currents B and C is 
passed out through the exhalant aperture. The feeding currents in 
ciliary feeding Prosobranchia represent modifications of some or all of 
these currents (Yonge, 1938). 


In Hipponix the only modification is due to the constriction of the 
opening of the mantle cavity which ha.s had the effect of carrying the 
greater part of the ctenidinm on to the dorsal surface so that, as .shown 
in figures 2 and 7, the filaments hang down above the head. The inhalant 
current (I) created by the lateral cilia enters on the left side, passes 
through the ctenidium and leaves as an exhalant current (E) on the 
right (fig. 3). The heaviest particles drop out of suspension on the left 
and are removed by cilia of current ^-i on the fioor of the inhalant aper- 
ture (fig. 2). The current then impinges on the os])hradium (0) and 
pa.sses between the filaments. Larger particles arc carried to the tip of 
the filaments by the frontal cilia and are rejected by cilia on tiie surface 
of the head, on the sides of the tentacles and on the fioor of the mantle 
cavity to the right, that is current B (fig. -l). The finest ])articles are carried 
between the filaments and are then consolidated in the mucus from the hypo- 
])ranchial gland. Cilia of current C then carry the mucus-laden masses 
to the exterior. The large size of the hypobranchial gland indicates the 
amount of material carried normally in suspension. Observations in life 
revealed great quantities of mucus in the mantle cavity. Posterior to the 
line of the right retractor of the head, the floor of the mantle cavity 
is not ciliated. No doubt any material which may accumulate here is 
forced out when the shell muscles contract. 

It w^as initially most surprising to find no trace of ciliary feeding 
in this sedentary animal. The Vermetidae, which are also cemented, feed 
either by ciliary currents or by mucus strings (Yonge, 1932, 1938 : Yonge 
and lies, 1939; Morton, 19516) while Capuhis ungaricus, which is closely 
related to Hipponix and still potentially, if seldom actually, mobile is a 
ciliary feeder (Yonge, 1938). In that species particles collected by the 
ctenidia are carried in a ciliary tract to the upper surface of the propo- 
dium (much larger than that of Hipponix) where the proboscis collects 
the mucus-laden masses by means of the radula. Orton (1949) has pointed 
out that CapvJus may also live on the shells of lamellibranchs, such as 
Modiolus and ifonia, and probably takes some of the food of tlu^se ani- 
mals by inserting the proboscis into the mantle cavity. But in Hipponix 
there is no doubt that ctenidium and ciliary rejection currents are in no 
Avay modified for feeding. 

The presence of a large osphradium is interesting. If this organ be 
solely chemo-receptive then its persistence in a sedentary animal is sur- 
prising. On the other hand if, as suggested elsewhere (Yonge. 1947), it 
is, at least primarily, a tactile organ concerned with estimating the 
amount of sediment carried in with the inhalant current, then its reten- 
tion would be expected. A sedentary animal is particularly susceptible to 
danger from accumulation of sediment within the mantle cavity. 



Alimentary Canal 

The mouth consists of a vertical slit at the end of the proboscis and at 
the base of the two lateral tla])S in which this terminates. There is a 
small, and certainly functional, radula and small salivary glands. The 
buccal cavity leads into what is termed by Graham (1939) the anterior 
oesophagus and wliich possesses in llipponix a dorsal ciliated food-channel 
of unusual width. As in other style-bearing mesogastropods, such as Capu- 
lus and members of the Calyptraeidae, the lateral glandtilar pouches 
primitively associated with the mid-oesophagus (Graham, 1939) are absent 
in Hipponix. As already noted, the oesophagus passes to the left to enter 
the visceral mass; there it enters the exceptionally large stomach (ST) 
which, as shown in figures 3 and 5, occupies the greater part of the pos- 
terior region of the visceral mass. 

The general appearance of this organ when opened mid-dorsally is shown 
in figure 6. The oesophagus (OE) enters ventrally on the left side; the 

Figure 6. Hipponix antiquatus, stomach opened along dorsal surface, showing 
appearance in life. XIO. D, opening of duct into digestive diverticula (ventral); 
MG, opening into mid-gut (dorsal) separated by typhlosole (TY ) from style-sac 
(SS); OE, opening of oesophagus (ventral), q, q, material accumulated by ciliary 
currents at side of gastric shield {GS) where normally caught up by the substance 
of the revolving head of the style. 

mid-gut (]\IG) which is separated by a conspicuous typhlosole (TY) 
from an associated style-sac (SS), leaves on the same side but dorsally 
(fig. 3). There is a common opening, on the floor of the stomach, for all 
the ducts of the digestive diverticula (figs. 5 and 6; D). This opening lies 
near to the gastric shield (GS) against which the short style bears. All 
of these structures lie on the left side of the stomach together with the 
usual sorting mechanisms of ridges and grooves, all richly ciliated. But 
some three quarters of the stomach consists of a capacious caeciun which 


extends to the right. The walls of this appear corrugated when the stomach 
is opened, ciliation is poorly developed but there is evidence of muscular 
contraction, sections revealing the presence of some strands of muscle 
around this region of the stomach. 

The digestive diverticula form a compact mass on the ventral side of 
the stomach as shown in figures 2 and 5 (DD). The tubules contain 
many dark spherules (seen in the section shown in fig. 5) which are 
probably of an excretory nature. Owing to their presence the diverticula 
form a black mass when viewed from the ventral side (fig. 2). Among 
the tubules and aiound the gut generally there is abundance of a yellow, 
vescicular connective tissue. The mid-gut and rectum extend forward in 
the roof of the mantle cavity where they form a series of loops (fig. 3, 
MG and R). In an animal with a shell diameter of 12 nun. these ter- 
minal regions of the gut pulled out to a length of some 22 nun. 

Food and Feeding 

The only previous account of feeding in Hipponix appears to be that 
of Risbec (1935) on H. australis. This species lives on the shells of other 
gastropods, usually of species of Turho, and characteristically near to 
the exhalant aperture. So situated, it feeds on the faecal pellets of the 
"host," the terminal processes of the proboscis separating and then com- 
ing together rapidly when the relatively enormous food masses are swal- 
lowed. Ilipponix antiquatus feeds in essentially the same manner but 
on fragments of material — organic detritus, pieces of algae, etc. — that are 
carried within the very limited area in front of the shell where alone 
the proboscis can brow^se. In the absence of automatic supplies of food, 
such as those received by H. australis from the animal it lives upon, this 
type of feeding can only be carried on l)y a sedentary animal if food 
supplies are constantly being renewed by w^ater movements. 

The proboscis itself is muscular and very active. The terminal lobes, 
with the mouth which they flank, were frequently seen to o])on widely 
and grope forward in apparent search for food. The odontophore was 
then seen to protrude from the mouth opening. Somewhat similar ol)- 
servations were made on C. ungaricus after mucus-laden food nuisses had 
been carried on to the surface of the propodium (Yonge, 1938). In this 
species, however, the proboscis, which is grooved anteriorly, i'e]n-esents 
the much extended terminal lobes in Hipponix. How widely the mouth 
must dilate in H. antiquatus is indicated by the nature of the stomach 
contents. These, together with much amorphous matter, probably organic 
detritus, comprise sand grains and also fragments of calcareous coral- 
line algae up to 2 mm. long and 0.5 mm. wide. The radula must convey 


those fraiiim-nls into the ))uecal mass and anterior oesophagus the power- 
ful eiliation of wliieh, possibly aided by some muscular action, serving to 
carry tliem into the stomach. 

Witiiin tlie stomach material would appear initially to pass to the 
right, into the caecal extension, as indicated in figure 6. This serves as a 
store and ])erha])s to some extent also as a gizzard although the triturition 
which must occur before the larger particles can pass into the mid-gut 
is ])robal)ly completed by the action of the style and gastric shield. Cer- 
tainly particles are carried to the edge of the gastric shield (fig. 6, q) 
where, in the iiitact stomach, they will be caught up in the head of the 
rotating style. Fine particles only will finally be carried into the large 
duct (D) leading into the digestive diverticula; the greater part of the 
stomach contents must pass, essentially unchanged apart from digestion 
of starch and some triturition, into the mid-gut. A very similar type of 
stomach, with a large caecal extension, is described by Graham (1939) 
in Pomatias (Cyclostoma) elegans. 

Within the mid-gut the faecal material becomes firmly compacted into 
pellets of relatively enormous size. Each is oval in shape and from 700 to 
800 /x long and about 500 fi in diameter. When crushed they are found 
to consist of fine amorphous material, dark green in colour with fine 
fragments of lime and silica. These pellets may occur in multiple rows 
and in such numbers as to distend the mid-gut and rectum to as much 
as three times the normal diameter. Indeed the roof of the mantle cavity 
may be largely occupied by as many as 16-20 of these pellets within 
the coilings of the mid-gut and rectum. The anal oj^ening, normally small, 
is greatly distended at defecation. The exhalant current must aid in car- 
rying the ])ellets clear of the shell where water movements may dispose 
of them. In the quiet water of an aquarium tank the pellets were de- 
posited in large numbers just outside the margin of the shell. Under 
these conditions they might possibly be seized by the groping proboscis 
and swallowed. 

There is no evidence of any discrimination in feeding. Anything that 
is available, up to a relatively very large size, appears to be swallowed. 
The oesophagus is wide enough to permit the passage of, and the stomach 
capacious enough to store, large particles. The stomach is that typical of 
a style-bearing gastropod, as summarized by Graham (1939), permitting 
(1) mixing of food with the style substance, (2) sorting of fine ])articles 
for passage into the digestive diverticula, (3) removal of larger particles 
and waste from the digestive diverticula into the mid-gut, (4) rotation 
and ]iassage into the stomach of the style. The very great amount of 
indigestible matter in the food is accumulated into exceptionally large 
faecal pellets which cannot cause fouling of the mantle cavity. 

vol.. xxviii i yonge: observations on hipponix antiquatus 13 

Reproductive Organs 

Every specimen examined in life or dissected was female. One small 
si>ecimen of Avhich lon>iitndinal sections were made showed no sign of 
reproductive organs and was presumably immature. The related Capulus 
ungaricus is a protandrous hermaphrodite (Giese, 1915) and this must 
surely also be true for Hipponix.^' In C. ungaricus the reproductive sys- 
tem is simple, consisting, in the male phase, of a testis with associated am- 
2)ullae and a duct leading into a pouch with which a receptaculum com- 
municates by i fine duct. The male genital opening lies at the side of the 
right shell muscle and the sperm is carried by way of a ciliated groove to 
the ti]) of a simple penis which is without glands. After sex change, the 
gonad enlarges to form an ovary without ampullae, the pouch of the 
male phase novv^ becomes what Giese describes as a uterus into which the 
receptaculum continues to open. The sperm groove and the penis disap- 
])ear but the female reproductive aperture, although longer, remains in 
the original position. 

I'rcsumably the males, being younger and jirobably more active, are 
able to move on to and copulate with the larger and i)robably completely 
immobile females. Jones (1949) describes the presence of small speci- 
mens of ('. ungaricus on the shells of larger ones, and these may well 
have been males and females respectively. After copulation, sperms 
would ])resiunably be stored in the receptaculum and later fertilize the 
eggs when they entered the "utei'us." The latter is thick-walled and 
glandular and responsible for the formation of the ])i-<)tective ca])sule dis- 
cussed in the next section. 

The small specnnen of TI. a)iti(juatiis without reproductive organs was 
some t) mm. in diameter and was attached to rock. If this animal was 
indeed inunature and if all members of this species are ])rotandric her- 
maphrodites then they cannot be mobile in a subsequent male ])hase. Cer- 
tainly no small mobile individuals were observed and in the very turbu- 
lent conditions where //. antiquatus lives, permanent attachment in early 
life would seem to be almost essential. This matter cannot l)e regarded 
as settled but all available evidence, admittedly very limited, indicates 
that the species, almost certainly a protandric herma]ihrodite, is not mo- 
bile in the male phase and so copulation cannot occur. If this be so, then 
fertilization mav occur in one of two ways. 

The first ]iossibility is that the spermatazoa are liberated freely into 
the sea and that fertilization occurs after these have been carried into 
the mantle cavity and have entered the genital aperture of a female. 
Such ''current fertilization" is known to occur in Turritclla communis 

*The parasitic Thyca stellastuis. according to Koeliler and Vaney (1912), is not hermaplirodite al- 
though the male is smaller than the female. 


(Fretter, 1946) and in vai'ious iiu'iubers of the Vcrmetidae (Morton, 
19516). All of these animals are, however, eiliary feeders (Graham, 1938, 
Morton, 1951 ) and so with enlarged ctenidia which create a powerful in- 
halant current. In addition the pallial genital duet is widely open ventrally 
to i)erniit the recejition of sperm carried to it in the water. All of these 
animals, moreover, probably live in numbers together and in compara- 
tively still water, i.e. under conditions where a local concentration of 
sperm adequate to ensure fertilization could be jiroduced. In Hipponix 
the inhalant current is much weaker, the concentration of animals is less 
while the water movements seem normally far too great to allow any 
effective concentration of sperm. Further, the very short pallial oviduct 
in Hipponix does not provide the facilities for entrance of water-borne 
sperm that are present in TurriteUa and the vermetids. 

The other alternative is that sperm produced during the male phase 
are stored for subsequent fertilization of eggs produced by the same (inimal 
when it passes into the female phase. This would not l)e unprecedented 
in the Alollusca, having been shown to occur in the wood-boring lamelli- 
branch, Xylophaga dorsaUs (Purchon, 1941). This animal alternates in 
sex but the sperms produced in a male phase are stored in a receptaculum 
for use in the subsequent female phase. Xylophaga dorsalis lives largely 
isolated in drift wood and this is probably the only feasible method by 
which fertilization can be assured. Although the habitat is different, the 
problem in the case of II. antiquatus is not dissimilar and the solution 
may be the same. 

The female reproductive organs are very similar to those descril)ed 
by Giese in Capidus. They are situated on the extreme right side of the 
visceral mass and consist of an ovary and, to employ the terms used by 
Fretter (1946), a gonadial duct (incorporating renal constituents), and 
a wide pallial region (fig. 7, OD) forming a capsule gland into which 
opens, by a narrow duct posteriorly ( fig. 7B, RS ) , a rounded receptaculum 
seminis. In section sperm were seen within this organ. The eggs are 
large, containing great quantities of yolk, when they leave the ovary (fig. 
7C, OV) and there is no evidence of any albumen gland. A capsule is 
almost certainly laid down around each egg, after fertilization by sperm 
from the receptaculum, by the conspicuously thick and glandular walls 
of the capsule gland (uterus of Giese). In this state the eggs will l)e 
l)assed into the mantle cavity. 

Attachment op Egg Capsules 

Sedentary ^Mesogastropoda must either attach the egg capsules to the 
inside of the shell, as do the Vermetidae (Morton, 1951&), fasten them 





Figure 7. Hipponix antiquatvs. transverse section through the anterior (A), 
middle (B), and posterior (C) regions of the mantle cavity. X5. EC. egg cap- 
sules; N, nei've collar at base of proboscis; OF, ovary containing large, yolk-filled 
eggs; RC. reno-pericardial canal, showing opening into pericardium (PC) ; RS. duct 
from oviduct (capsule gland) into receptaculum seminis; T. tentacle; V. ventricle. 
Other lettering as before. 


to the rock <»i- j)el)l)l('.s on wliieh the female rests, as in Calyptraeidae such 
as CrepkluUi foDiicald ( Lehoiir, 1987), or else retain tlieni secured to the 
actiial body of the female. This is the case in both Capulns and Hipponix. 
In C. iinfidrivus a sinole large egg capsule or cocoon is attached to the 
large propodiuin of the female. Full references to literature are given 
by Thorson (l!)46j who summai'izes ]>revious work in the statement that 
*'Each fenude protects a single thin-walled cocoon of the form of a 
sausage, and each, cocoon contains several eggs, 200 fi across, which hatch 
as veligers through a fissure on the under side of the cocoon.'' 

Although Kisi)ec (1935) does not mention males and gives a very 
imperfect account of the female genital system in H. australis, he gives 
i\ good descrij)tion of the egg capsules in this species. He notes that it is 
impossible to observe egg-laying and the process of formation and attach- 
ment of the capsules. This is equally true for H. mitiquatus. He de- 
scribes the presence of six or more capsules each containing a number 
of eggs and attached by a slender stem to a plate which is itself imbedded 
in the tissues of the foot. He considered that these sacs received the eggs 
and also yellow nutritive material of a fatty nature. He was unable to 
ol)serve the role of the foot in the formation and attachment of these 

Conditions are essentially similar in H. antiqnatus. After the comple- 
tion of egg-laying the mantle cavity of the female is completely filled by 
from six to eight egg capsules each containing up to 50 yellowish eggs. 
The eggs measured about 350 /* in diameter, the greatest transverse diameter 
of the capsules being 840 fx. Each capsule was, as in H. australis, attached 
by an attenuated stalk to a perforated calcareous plate Avhich in turn 
was attached witliin a membranous area situated in the depression ven- 
ti'al of the ])i"oj)odium (figs. 4, 7C, and 8; PL). As shown in figure 8, the 
<'apsules usually bulge forward so that they may project a little distance 
out of the mantle cavity (at least when the animals have been detached). 
The head is ]iushed over to the left of the mantle cavity and the animal 
must have some difficulty in maintaining the necessary circulation of 
water through the mantle cavity. 

Unfortunately, it was impossible to observe the mode of formation 
4ind attachmein of the egg capsules. Owing to the sedentary habit of this 
animal, full dct;uls of this process could probably only be made out by 
<'xamining great numbers of animals during the period of egg-laying in 
hope of obtaining animals in all stages. The first egg-carrying female 
was obtaijied on April 5 ; during the summer all animals had o^^ capsules 
;i I though some of these were empty. 

Il is. however, certaiji that the eggs are large and yolky when they 
leave the ovaiy, where they were observed in sections (fig. 7C, OV). After 


Figure S. HiiJiJonix antiquatus. ventral aspect with egg capsules in the mantle 
cavity. X5. Lettering as before. 

fertilization tliey prestimably each receive a protective covering while 
in the capstile gland. It is, however, likely that the large capsules, each 
containing many eggs, are secreted by the gland that opens on the dorsal 
surface of the foot and which is certainly active, judging from sections, 
at this time. Moreover, it has been shown by Werner (1948) that, in 
Crepidula fornlcata, a stalked capsule is secreted arotmd the eggs by 
such a gland in the propodium. The calcareous plate and its investing 
membrane may be formed within the pocket ventral to the propoditim; 
sections show evidence that the epithelium has secreted this membrane. 
But it is not advisable to speculate further on a matter that can only 
be determined by observation. 

So far as could be determined, all eggs developed. There was no evi- 
dence that any of them formed food for others or that the capsules con- 
tained any additional nutritive material as suggested by Risbec. The 
embryos develop into fully shelled larvae but whether these actually crawl 
away after emergence from the capsules, as described by Risbec for JI . 
australis, was not seen. In \iew of the very specialized habitat, it is more 
than probable that, in //. foifiquatus also, the young crawl away from the 
parent to settle in due course permanently on the adjacent rock stirface. 


Gastropods -which possess the limpet form and the accompanying habit 
of life are particularly well fitted for survival on a hard substratum in 
the turbulence of the intertidal or shallow waters where such substrata 



iiornially occiu-. It is thereforo not surprising that the limpet foi-ni, with 
its secondary symmetry (see Yonge, 1947, p. 490, fig. 31) has imh'pend- 
ently been evolved on a number of occasions. The particular conditions 
that exist in liipixnii.r are most suital)ly discussed by coni]iaring them 
with those found in examples of the other chief types of marine lim]K'ts. 
In figur-e 9 compai-ison is made between four types of lim])ets, (A) 
Diodoro (FissureUa) and (B) Acmaea are both archaeogastropod lim])ets 
the former having paired, i.e. zygobranchous, aspidobranchiate ctenidia, 
the latter having a single aspidobranch ctenidium. (C) Hipjxniix is a 
mesogastropod limpet with a pectinibranch ctenidium and (D) Siijhonaria 
is a pulmonate with a secondary gill in what has been a pulmonary mantle 
cavity. A ca])-like shell and a horseshoe-shaped shell muscle (with the 

Figure 9. Diagrammatic comparison, from dorsal aspect, of four types of 
limpets, showing mantle cavity (MC) with inhalant (/), exhalant (E) and ma- 
jor cleansing currents (latter broken arrows), also shell muscles (black) and vis- 
ceral mass (stippled). A, Diodora (Archaeogastropoda, Fissurellidae) with paired 
and symmetrical ctenidia, exhalant current dorsal; B, Acmaea (Archaeogastropoda. 
Patellacea) with single aspidobranch ctenidium, exhalant current posterior: C, 
nil>l)0)ii.r (Mesogastropoda) with pectinibranch ctenidium, exhalant current an- 
ii'iior; D, Siijhoiutria (Pulmonata) secondary gill, exhalant current on right. 


opoiiiiiii' on the right in Siphonaria) arc common to all. The course of the 
respiratory and cleansing currents may be hrietly summarized. The 
anterior mantle cavity of the prosobranehs (A-C) has a generalized an- 
terior inhalant current (T) in A but, with the loss of the right ctenidium, 
this is confined to the left in B and C. In A the exhalant current (E) 
issues dorsally, through the shell aperture found in all zygobranchous 
gastropods and here apical in position, in B it is carried by way of the 
right pallial groove to the posterior end of the animal, whence also passes 
sediment collected in the mantle (nuchal) cavity and in the pallial grooves. 
From such a condition have been derived those in the more specialized 
Lottki, Patina, and Patella (Yonge, 1947) with their secondary pallial gills. 
These Patellacea (Docoglossa) represent the most successful of all limpets 
and this may well be associated with the use they, alone among limpets, 
have made of the pallial grooves. In Hipponix (C), apart from the flat- 
tening already discussed, the mantle cavity and its currents are those of 
a tyi>ical ])ectinibranch. In D the pulmonate mantle cavity has been suc- 
cessfully readapted for aquatic life by the appearance of secondary gills 
within this (Yonge, 1952) and not in the pallial grooves. The restricted 
opening of the mantle cavity on the right side is retained with the exhalant 
current issuing immediately ]K)sterior to the inhalant opening. 

It is in the matter of feeding that Hipponix, and the mesogastropod 
limpets in general, differ from these archaeogastropod and pulmonate lim- 
pets. Tlie latter all browse on encrusting vegetation, moving very slowly 
over the rock and scraping this with the broad radula. The mesogastropod 
limpets may be divided into (1) those in which the ctenidial filaments 
have been greatly elongated and which feed exclusively by ciliary cur- 
rents, i.e. Crcpidula and CaJjiptrara (Calyptraeidae) and (2) those in 
which there is a pronounced pro])oscis. In both groups the power of move- 
ment is lost, effectively even where the animal does not actually become 
attached. In the Calytraeidae movement ceases to be necessary because 
food, suspended in the inhalant current, is brought to the animal. But in 
the second group a most interesting variety of conditions prevails. 

In CapuJus, which is the least specialized, the grooved i)roboscis, 
formed by prolongation of the terminal lobes in Hipponix, may be used 
to take in food collected by the enlarged ctenidia (Yonge, 1938) but it 
may ])robably be also used to take in material similarly collected by a 
lamellibranch (Orton, 1949). In both Hipponix australis and H. anti- 
quatus the ctenidium is not concerned with feeding, the proboscis swal- 
lowing relatively large food masses. In the former species, which lives on 
a "host" animal, the habit has resemblance to that described by Orton for 
C. vnffaricus, but H. antiquafus is not dependent on any other animal. 
Although cemented to the sul)stratum, yet it uses the proboscis to collect 


food in front of tlie shell. Species of a third i>enns, Thyca, have also lost 
the power of movement but in association with a completely parasitic life. 
They are ectoparasitic on echinoderms, the foot being reduced but the 
long- proboscis penetrating deep into the tissues of the host (Schepman 
and Nierstras/, 1909; Koehler and Vaney, 1912). The disc of attachment 
is formed by the proboscis, the eolumellar muscle described by Koehler 
and Vaney apparently consisting of the retractor muscles of the head. 

In H. antiquatus the three most striking characteristics are (1) the tiat- 
tening of the shell, with its consequences on the form of the mantle cavity 
and the disposition of the head, (2) cementation to the substratum by the 
secretion of a ventral "valve" by the undersurface of the foot and with 
consequent loss of motility, and (3) feeding, while so attached, by means 
of the proboscis. All three, however, are related to one another and to 
the habitat. A much flattened limpet which was cemented to the sub- 
stratum would have survival value under the conditions where //. anti- 
quatus lives, namely within crevices among rocks exposed to the full and 
almost unvarying force of the Pacific surf. Under such conditions food 
will constantly be renewed so that exclusive dependence on the proboscis 
is possible. This is not the case in (\ ungaricus which lives. su])littorally, 
in very much quieter water. 

Again, members of both of these groups of limpets are lu'otandrous 
hermaphrodites (certainly Crepidula and Calyptraea, also Capulus and 
almost certainly Hipponix) but have the typical female genital system 
of the mesogastropods with large and internally fertilized eggs. The egg 
capsules must of necessity be attached either to the underlying sub- 
stratum or to the animal itself. Only the latter is possible in Hipponix 
because the animal is cemented. The precise period at which the sperm 
enters the receptaculum, which involves the question of whether self or 
cross fertilization occurs, remains to be determined, as does the precis© 
manner in which the egg capsules are attached to the underside of the 

There is finally the question of classification. Thiele (1931) i)laces 
Amalthea (^=^Hipponix) in the Amaltheacea but CapuUis and Thijca in 
the family Calyptraeidae. While there is certainly much still to be 
learned about these and allied genera and families, there can be no doubt 
that Hipponix is much more closely allied to CapuJus than would appear 
from Thiele 's classification, which cannot in this particular instance be 
supported. The precise position of Thyca, so greatly modified in form 
and habit and yet not a protandrous hermaphrodite, appears to need 
further scrutiny. 



Hipponix antiquatus is a mesogastropod limpet of ])ai'1i('ular interest 
owing to its sedentary lialnts. It lives cemented to the substratum in 
crevices among rocks often fully exposed to the Pacific surf. It is highly 
adapted for life in such an extreme habitat. 

The ventral surface of the foot secretes a "ventral valve"' closely 
resembling the "dorsal valve," i.e. shell, secreted by the mantle. The 
margins of the two make perfect contact when the horseshoe-shai)ed shell 
muscle contracts. 

The opening of the mantle cavity is constricted causing displacement 
dorsally of anus and ctenidium ; the shell is also much flattened with 
conse(iuent effects on the mantle cavity and displacement to the left of 
the head. In other respects the disposition of the pallial organs is that 
typical of mesogastropods. 

The ctenidia are solely concerned ^vith respiration. Feeding is by 
means of a muscular proboscis which is extruded from the mantle cavity 
and swallows relatively very large masses of organic detritus, such as 
fragments of calcareous algae. 

The alimentary canal is modified for the reception and utilization of 
such food masses. The stomach is large but otherwise that typical of a 
style-bearing mesogastropod. There is a single opening into the ventrally 
disposed digestive diverticula. Faecel pellets are exceptionally large. 

Available evidence indicates that H. antiquatus, like the allied Capu- 
lus ungaricus, is a protandric hermaphrodite. Owing to the sedentary 
habit and to the rough water in which it lives, cross fertilization may be 
impossible and it appears more ])robable that spermatozoa produced during 
the male phase are stored in the receptaculum for fertilization of eggs 
produced in the subsequent female ])hase. 

Large egg capsules, each containing a number of large, yolky eggs, 
are attached to a calcareous plate embedded ventral to the propodium. 
The mantle cavity is largely filled by these capsules during s[>ring and 
early summer. 

In comparison witli archaeogastro])od and also pulmonate marine lim- 
pets, mesogastropod limpets are sedentary, feeding by means of ciliary 
currents {Crepidula, Calytraea, Capulus) or by means of a proboscis 
either on organic detritus or faeces {Hipponix spp.) or else parasitically 
(Thyca). AVitli the exception of Thyca, all of these limpets are also pro- 
tandric hermaphrodites. 



Cook. P. M. 

1949. A ciliary feeding mechanism in Vivipai'u.s vivipani.s (L). Proceerlinfjs 
uf the Malacological Society of London, vol. 27. pp. 265-271. 

Fkhtiki:, V. 

1946. The genital ducts of Theodoxus, LameUaria and Trivia, and a discussion 
on their evolution in the prosobranchs. Journal of the Marine Bio- 
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GlESE, M. 

1915. Der Genitalapparat von Calyptraea sinensis Lin., Crepidula iinguiformis 
Lam. und Capulus ungaricus Lam. Zeitschrift fiir icissenschaftliche 
Zoologie. Bd. 114. pp. 169 231. 


1938. On a ciliary process of food-collecting in the gastropod Turritella com- 

munis Risso. Proceedings of the Zoological Society of London, ser. A. 
vol. 108, pp. 453-463. 

1939. On the structure of the alimentary canal of style-bearing pi-osobranchs. 

Proceedings of the Zoological Society of London, ser. B, vol. 109, pp. 

Jones. N. S. 

1949. Biological note on Cuijulus ungaricus. Mariyie Biological Station at Port 
Erin. Isle of Man. Annual Report for 1948 (no. 61). p. 29. 

Keen, A. M. 

1937. An abridged check list and bibliography of west North American marine 
MoUusca. Stanford University Press, California. 

Keep. J., and J. L. Baily, Jr. 

1935. West coast shells. Stanford University Press, California. 

KoEMLER. R., and C. Vaney 

1912. Nouvelles formes de Gasteropodes ectoparasites. Bulletin Scientifique 
de la France et de la Belgique. T. 46, pp. 191-217. 

LKnouR. M. V. 

1937. The eggs and larvae of the British prosobranchs with special refer- 
ence to those living in the plankton. Journal of the Marine Bioloqical 
Association of the United Kingdom, vol. 22. pp. 105-166. 

Morton. J. E. 

19510. The ecology and digestive system of the Struthiolariidae (Gastropoda). 
Quarterly Journal of Microscopic Science, vol. 92, pp. 1-25. 

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poma. Part 3: Novastoa lamellosa and its affinities. Transactions of 
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1912. The mode of feeding of Crepidula, with an account of the current- 
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Marine Biological Association of the United Kinydovi. vol. 9. pp. 444- 

1949. Note on the feeding hal)it of Caimlus ungaricns. Marine Biological Sta- 
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Pi KcnoN, R. D. 

1941. On the biology and relationships of the lamellibranch Xylophaga dorsalis 
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Kingdom, vol. 25, pp. 1-39. 


1935. Eiologie et ponte de mollusques gasteropodes Neo-Caledoniens. Bulletin 
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1909. Parasitische Prosobranchier der Siboga-Expedition. Siboga-Expeditie, no. 

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1948. The marine mollusks and brachiopods of Monterey Bay, California, and 
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TlIlELE, J. 

1931. Handbuch der systematischen Weichtierkunde, Bd. 1, pp. 1-376. 

TnoRSON, G. 

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Fourth Series 
Vol. XXVIII, No. 2 July 15, 1953 



Research Associate 
California Academy of Sciences 

The discovery in West Central China in 1945-46 (see Chaney, 1948) 
of a living species of the ancient, fossil, coniferous genus Mcfasequoia, a 
possible ancestor of the redwoods, aroused widespread interest in the tree. 
After Dr. R. W. Chaney, of the University of California, made a hurried 
trip to see the trees in the early spring of 1948 an arrangement was made 
between the California Academy of Sciences and Lingnan University for 
a joint expedition to spend considerable time in the area in the summer 
of 1948. The travel expenses were supplied by the California Academy 
of Sciences and personnel and equipment by Lingnan University. 

The trees were known from a few localities along the border of Sze- 
chuan and Ilupeh provinces, thirty to seventy-five miles south of Wan 
Hsien on the Yangtze River below Chungking. In the early reports 
just over 100 trees were mentioned, though it was estimated there were 
many more. AVe were able actually to count 1219 trees, though not all the 
young trees, and no doubt not quite all the large trees, are included in 
our count. We searched the country in several directions for added stands. 
If more exist, they are probably in a few limited pockets not far to the 
west, east of Chungking. 

Metasequoia glyptostrohoides llu and Cheng, 1948 (fig. 1), is a large 
and impressive conifer. It grows to a height of 115 feet and a diameter 
of 8 feet. Actually there is only one tree this large, though there are 
several about 100 feet high and 5 feet in diameter (at 6 feet above the 

[25 1 



Figure 1. One of the largest Meta-sequoia trees, located in lower Suisapa Valley 
near Hsiao-ho. 



ground). The trees are straight, and not irregular like the relative, Glypto- 
strohiLs ijensilis, the "water-pine" of the Canton delta, for which the 
.si)eeies was named. The trunk is somewhat enlarged at the base and the 
l)ark is reddish but thin. On the smaller branches the bark is somewhat 
folded or curled. The lower branches slant slightly downward, and the 
upi)er branches point distinctly upward. The wood is relatively light 
and soft. The foliage is a conspicuous i)ale green. The needles are flat 
(fig. 2), on opposite sides of the stem, and closely resemble those of the 

Figure 2. Foliage of Metasequoia (/lyptostroboides. 

California coast redwood, Sequoia si'mprrvirens. Metasequoia ditt'ers from 
the latter, however, in being deciduous. The cones (fig. 3) are small, 
nearly round, and resemble rather closely those of the coast redwood 
in general appearance. 

The foliage, however, is much softer and the twigs and branches quite 
fragile. These characteristics seem to set it off rather conspicuously from 
many other conifers. In fact, after the species has grown longer in culti- 
vation, it may be judged one of the most beautiful of existing trees. 

The dawn redwood is called shui-hsa in Chinese, which means "water- 
fir" or "watev-Cunninghamia." The tree was apparently unknown out- 
side its range before its recent scientific discovery by Chinese plant col- 
lectors. Probablv its range in recent historic times has not been much 




Figure 3. Cones oi Metasequoia glyptosti-oboides (outer two) and Glyptofitrobus 
pensilis (inner two). 

more extensive than at the present time. This is suggested by tin- fact 
that the wood is not considered valuable, and is not carried out of the 
mountains as is the wood of Cunninghamia, or "Chinese fir." Never- 
theless, the species has probably been suffering reduction of numbers 
over a long period. The fact that it requires a damp habitat and grows 
along streams makes its survival precarious with the increase of i)opula- 
tion and the spreading of rice fields up into the higher valleys. Possibly 
the massacre 300 years ago of most of the people in eastern Szecluian by 
the Imperial forces for failure to pay taxes may have been an imjiortant 
factor in saving the tree from extinction in recent years. Another factor 
may be the apparent state of semicultivation under which the tree exists. 
The fact seems to be that a considerable percentage of the existing trees 
have been transplanted to their present situations. For example, many 
of the large trees are in straight rows up ravines, paralh'liiiu the small 
streams. Others are around the farmhouses. INIany young and iiicdium- 
sized trees are in straiglit I'ows along tlie edges of rice fields bordering 

The reason for the transplanting of tlie volunteer seedling-; from the 
shaded ravines to particular situations, often in rows, is jipparently 
based on local superstition. The mountain peo])le have tlie habit of ])re- 
dicting their crojis on the basis of cone develo])ment on the frees. Thus 
a heavy crop of cones on the upi)er portions of the trees is said 1o indi- 
cate a good rice harvest, and an abundance of cones on the lower branches 
signifies good results on the hill crops (corn, drugs, herbs, lacquei-, etc.). 
Thus it may actually l)e that the dawn redwood has been i)i-eserve(l li-om 
final extinction more or less by chance. It is interesting lo note that 
the water ])ine, (ilijptostrohus pensiJis, of southernmost ('hin;i. is also 


planted for geoinaiitic pui'i)ose.s, geiu'rally along old paths in the delta 

After observing the soft and Hexible nature of the foliage and 
branches, it is eas.y to understand how the tree has come to be semi- 
sacred and used for divination purposes. Particularly when observed in 
a breeze, the tree has a feathery and somewhat fairy-like aiipearance, 
and this seems to suggest its uniqueness. 

The principal purpose of our trip was to collect insects and other 
animals in the hope of finding some ancient faunal ek'ments of possible 
North American affinity which might have survived with Mctascquoia 
and the other ancient trees associated with it. The latest known fossil 
deposits of Metasequoia were laid down in Oregon about 20,000,000 years 
ago (John Day Miocene). A number of the other genera of trees asso- 
ciated in the same fossil deposits are still found growing with the liv- 
ing dawn redwoods. What is still more striking is that the j) resent 
dawn-redwood area is the only known place in the world where all of 
these particular trees, exclusive of Metasequoia, are now growing to- 
gether. Thus we hoped to find some animals which might be descended 
from species in ancient fossil deposits, possibly contemj^oraneous with 
the old deposits of Metasequoia which have been found in many places 
in Europe, North Asia, and North America. We even hoped to find some 
insects related to those in the present redwood association of California 
and Oregon. Since fossil birds are extremely few and the chance of wuld 
animals having survived very slight, hope and emiihasis were placed 
upon the insects. 

Our group went to western China in two parties. The first party left 
earlier and went ahead overland to Chungking late in June, going by train 
through Kwangtung, southern Ilunan, and Kwangsi provinces, and by bus 
through Kweichow and southern Szechuan provinces. The others of us went 
by plane from Canton to Hong Kong on July 3 and from there to Chung- 
king the next day. Even with a week's head start, the first party reached 
Chungking nine days behind us. During our wait, my assistant and I made 
two three-day trips to investigate citrus insect pests and ship their natural 
enemies to the Citrus Experiment Station of the University of California 
at Riverside. One tri]i was to Chiang-chin, a few hours up the Yangtze by 
steamboat, the other to Pe-pei. a major war-time educational and research 
center, up the Chialing River. Pe-pei is a progressive small city with ]Hiblic 
library, museum, park, zoo, and many schools, quite unusual in China, par- 
ticularly for such a small city. 

In the afternoon of July 14 we boarded a ship for Wan Hsien, but 
after midnight w^e crossed the Yangtze and then the Chialing River to 
take a smaller steamer, as the first ship had been taken over by the army. 
Leaving early the next morning, we reached Wan llsien at dusk. The 



[Proc. 4tii Ser. 

Figure 4. Dr. J. Linsley Gressitt. the author and leader of the expedition. 

next day we saw the magistrate and made our arrangements with por- 
ters to carry our baggage. The following morning we started our long 
walk south with a police officer, two guards, and four porters. We first 
went up the river about two miles in a sampan and crossed to the south 
bank. Passing the mouth of a small tributary coming from a waterfall, 
we saw a small rock-slide which peppered with rocks for a moment a 
river boat tied up at the base of the cliff. The south bank was high, level, 
and covered with rice paddies between a ])utte-like limestone mountain 
and a small gorge with a waterfall. 

Farther south the country was irregular and hilly, with generally 
oblique or subvertical shale strata. Parts of the second and third day 
took us through mountains of horizontal layers of sandstone, fifty or so 
feet thick, alternating with softer strata, partly shale, of similar total 
thickness. This gave a somewhat "Grand Canyon "-like effect, with 
alternating 50 to 60-degree-angle slopes and vertical drops. The sloping 
surfaces Avere generally planted with corn, sometimes up to the high- 
est level, there often being at least three such slopes between the alter- 
nating cliff's. Generally there were young pine trees (Finns viassoniana) 


growing on the tops of the mountains. In other areas more or h'ss natural 
vegetation grows on the steejjer, irreguhir slopes, though corn is often 
planted on slopes with as much as (J5-degree gradient. The more rolling 
mountains are often clothed only with grass, or with scattered pines or 
shrubbery, and are used for cattle and horse grazing. Goats are grazed 
in many of the canyons. None of these animals are milked, but instead 
they are raised for meat and hides, or as work animals. The horses, 
which are very small, are sold to traders or officials. 

Most of the lower slopes of the canyons are planted to tung-oil trees 
and corn is grown among them. iMelons and beans are also cultivated, 
as well as peppers and some cotton and tobacco. Rice, of course, is grown 
where the topography and water supply permit. Scattered groups of 
citrus trees, mostly oranges, were found in the low hills and along the 
Mo-tau-chi River, mostly near farmhouses. Some pomelo and tangerine 
trees were also grown. This was the typical agricultural situation in 
the valleys up to 2,000-3,000 feet in altitude. 

The second day, just past Chang-tang-chin, where we met the ]\lo-tau- 
chi River, we were forced to delay a few hours until the river level went 
down enough to allow the porters to pass along the main trail in the 
rock bank, which had been overflowed by high rain torrents. At this 
point there were a few salt mines along the edge of the river. The out- 
put is small, however, and most of the salt used in the mountains is 
carried in by coolies in large slabs of stratified salt mined and l)rought 
from the southwest. 

The third afternoon we reached Mo-tau-chi after climl)ing up a 
thousand feet from near the upper end of the Mo-tau-chi River valley to 
an altitude of nearly 4.000 feet. I walked past the town and saw my first 
dawn redwoods, the three originally discovered trees. The large type 
tree (fig. 5), 90 feet high and 5 feet in diameter (well above the ground) 
has a shrine at its base. The other tw^o are one-fourth grown. All three 
are more or less in a row along the banks of the stream, and at the edge 
of rice fields. JMy first view of the foliage reminded me vividly of the 
coast redwood, except for the softer and more fragile nature. However, 
the drooping terminal foliage and the swollen base of the tree suggested 
to me the water pine, Glyptostrohus, of Canton, which is a relative of 
the bald cypress, Taxodium, of southeastern United States. The dawn 
redwood also resembles the latter two in its thin bark, light wood, and 
preference for growing near water. The species, I later learned, had 
been named just two months earlier, by Dr. H. H. Hu of Peii)ing and 
Professor W. C. Cheng of Nanking, who at the same time erected the 
new family Metasequoiaceae to contain the dawn redwood and the extinct 
members of its genus, dating back, as they do, 100,000,000 years. 

The next day, after passing through a heavy stone gateway in a 



Figure 5. Type tree of Metasequoia glyijtostrohoides, located at Mo-tau-chi, 




small ravine guarding the border of Szechnan Province, we entered IIu- 
peh Province, and then went over a pass 5,100 feet high on the Chi-o 
Shan range (mountains of the seven humps) and down into a wide, fiat 
valley to Wang-chia-ying. Not far from this market town is the largest 
known dawn redwood, 81/2 feet in diameter and 115 feet tall, but we put 
off seeing it until our return because of our baggage and itinerary. On 
the twenty-first we crossed the final pass (5,000 feet) and entered the 
upper end of the Suisapa Valley (fig. 6), or Dawn Valley as I like to call it. 

P^igure G. View of upper pait of Suit^apa Valley. 

The valley was our principal objective and occupied most of our time for 
the rest of the summer. When the last of our ])arty left, it had become 
cpiite cool. 

The name Suisapa (Shui-sha-pa) refers to the local, and only Chinese, 
name, s\ii-sa, of the dawn redwood. Suisapa specifically is only a single 
small locality in the middle of the valley (fig. 7), but is the best known 
place, and more or less serves to identify the whole, as it seems to have no 
other name. Actually, there is another small farming community of 
the same name in a canyon off the lower end of the valley, near llsiaoho, 
the only market town in the 25-mile long valley. 

It is this valley which contains the unusual assemblage of j^lants of 



I Pk()( . 4 III Si;i{. 

Figure 7. View of the middle portion ot Suisapa Valley. 

ancient northern affinity, many of them being among tlie most familiar 
and conspicuous types of trees in Europe and North America. Among 
them are beeches, birches, poplars, willows, oaks, chestnut, maples, horn- 
beam, hop hornbeam, linden, sassafras, pine, and yew. If it were not 
for the rice paddies, farmhouses, and people, a European or American 
might believe himself near home. Forests of the extent found in this 
valley are rarely seen in China except on the steeper slopes of high 
mountains, and in precipitous canyons or temple preserves. Immediately 
on entering the valley, one cannot but sense its uniqueness, both from 
the standpoint of the unusual nature of the flora and from tlie extent of 
its preservation. Probably one reason for the slower rate at which 
the trees are being cut is that the nearest large commercial center is Wan 
Hsien, 120 miles walk to the north, and in another province, whereas 
the stream in the valley flows in the opposite direction for a short distance 
underground just outside the valley, and then a very long distance, round 
al)out, with dangerous rajiids and narrow gorges up wiiich boats may not 
be towed, before meeting the Yangtze River between Chungking and Wan 
Ilsien. Foresters Avho might dare to float rafts of logs downstream would 
have to dispose of their poles and ropes and walk back. Though trees are 
being cut locally at a seemingly alarming rate, they are used mostly for local 


purposes, particularly for houses, fuel, aud coffins. Coffins are made from 
trunks of Cunninghamia and carried out of the mountains as a business. 

The valley extends northeast and southwest and curves eastward at 
its upper end. The lower end is more or less closed by an east-west range 
of hills, the stream passing through a break in them, and continuing be- 
yond to the southeast for a few miles, partly underground. On each side 
of the valley extends a fairl.y sharp ridge, the east ridge reaching 5,500 
feet and the west ridge 5,100 feet in altitude. The east ridge has a num- 
ber of side ridges extending far into the valley and causing bends in the 
river though the valley is fairly straight. Some of these side ridges are 
very steep, narrow, or many-branched, and some broader. Tlie wxst ridge 
is rather uniformly narrow, being steep on each side, with only a few 
short side ridges. It is quite irregular, with several peaks and depres- 
sions. The east ridge is more regular along most of its skyline, decreas- 
ing in altitude towards the south. On the east side of it there is a con- 
sideral)le drop into a large, deep valley which slopes much more steeply 
to the soutli than does Suisapa Valley. On the west side of the west ridge, 
however, tlie drop is only several hundred feet except at the north. Im- 
mediately to the west is a large, flat-topped, treeless range of much differ- 
ent appearance from the others. It is the highest in the near vicinity 
except for Mt. Sun-sin-po to the south, located not far southwest of 
the lower end of Dawn Valley. 

We made our headquarters in one of the two large farmhouses on the 
east side of the stream at Suisapa. Part of this house was occupied by a 
former mayor, and the present mayor lived in the other house. Since Dr. 
Chancy 's visit four months earlier, the mayor's wife and one child had 
died and the mayor was now sick. We therefore could not live in his 
house, and had to be somewhat careful until we had made friends with 
the people, as they tended to ascribe this bad luck as caused by the 
foreigners coming and cutting down a dawn redwood for specimens. How- 
ever, the local people themselves cut the trees not infrequently. There 
is a prevalent local custom of cutting the branches oif the Metasequoia 
and (Uniniufihamia trees periodically, often almost to the very top, at 
least for the trees close to their houses. Thus most of the trees outside 
the shady ravines are apt to have an extremely slender appearance as new 
branches are growing out (see figs. 8 and 9). Sometimes the trees are thus 
killed, as had recently happened to two large metasequoias next to our 
farmhouse. For those not killed the practice reduces the potential self- 
seeding of the trees. 

In the canyons near our headquarters, which included many of the 
large trees outside of farmyards, we found quite a few young seedlings. 
This indicates the species is not senile, but is still able to reproduce itself. 
On close searching among the boulders and shrubbery we could often 



Figure 8. Two pruned Metasequoia trees. 




Figure 9. Metasequoia trees on floor of Suisapa Valley. 



Figure 10. Metasequoia trees, south-central part of Suisapa Valley. 




find at least as many very young trees as large ones. Though the trees 
are generally erect, sometimes young trees have widely spreading branches, 
making them as broad as high, like yew trees in the valley. 

The dawn redwoods are found scattered throughout the valley (fig. 10) 
and in some neighboring valleys to the immediate south. They are most 
abundant in the shady ravines branching from the central portion of 
the valley, though there are some large groups in certain spots near the 
southern end. J\Iany young ones are also planted along the nuiin stream, 
bordering rice paddies. In the upper portion of the valley they are 
scarcer. Their altitudinal range is api)roximately from 2,000 to 4,000 feet. 
There can hardly be said to be any Metosequoia forests or even moderate 
natural stands. ]\Iost of the trees are in rows following up side streams 
(fig. 11). There are rarely as many as 50 along a single branch stream. 

Figure 11. Straight row of Mrtii.stqiK/Ki trt't's along a small stream in a side 
valley, Suisapa. 

On arrival at Suisapa we dispensed with our guards and decided we 
would just forget about bandits in spite of the w^arnings about them. One 
of the guards who came over the pass with us from Wang-chia-ying, 
whence our Wan Hsien escort returned home, told me there were 2,000 
bandits across the second range to the west. A resident of the valley, who 
had been returning along the road with us, warned our guard at Wang- 
chia-3'ing that a group of bandits were trailing us, but they turned out 


to 1)0 i)lain-eIotliPs <iiiar(ls traekiiig' bandits. The wife of a bandit who 
had r()])])('d one of our guards on a previous trip had l)een apprehended 
in a \ilhige along our route and was interrogated by tlie offieer during 
oui' evening there. In s})ite of a total of six one-way trips we made l)e- 
tween AVan Hsien and Dawn Valley, as well as several side trips, we 
had no bandit encounters. 

Our actual host in this farmhouse was an old widower, who had only 
one child with him, a son of 16. We ate and worked, and two of us slept, 
in his main room. The other four of our group slept in the opposite wing. 
He and his son slept on low, single-plank benches on each side of the open 
hearth fire in their kitchen, keeping the fire burning all night. These houses 
were of wood and had no chimneys. The smoke filtered out through the 
tiles in the roof, and through the small windows. The roof beams or bamboo 
drying frameworks above the fire often dripped an oily pitch and were 
thickly encrusted with cinders. Rice was cooked in a large iron pot and 
other things in a large concave iron pan like a slice from the edge of a 

The local food consists mainly of rice, corn, white potatoes, beans, 
sciuash, i)eppers, some Chinese greens, cabbage, and eggplant. The peppers 
are very hot and are eaten unmixed as regular dishes in every meal as 
well as flavoring for most other dishes, so we found their meals unpalatable. 
Actually they eat few vegetables fresh and pickle most of them, mixed 
with peppers. The corn they often eat as corn-meal mush. We found it 
hard to obtain sufficient fresh vegetables. Also, most of them wei-e not 
harvested until oversized. The people made their own bean-curd, but 
never had any they wished to spare, so we bought it only at the market, 
where also we got our only meat. Fortunately, we were able to buy eggs 
most of the time, but could only get enough by buying several dozen 
every market day (every three or four days). Food was cheap and in- 
flation was behind time here. We bought eggs as cheaply as 200 to a 
United States dollar. Beef was purchased as cheaply as two cents U.S. a 
pound, but we were only able to obtain it twice during the summer. Pork 
cost four times as much. 

Most of our breakfasts and suppers, except the day after market day, 
consisted of rice, boiled potatoes, and soup containing eggs, a bouillon 
cube, and a little pork fat, with sometimes a green vegetable. Lunch con- 
sisted of one or two eggs and two or three small potatoes boiled and 
carried along with us. 

Though the food of these people seemed poor, they had ((uite enough 
of it for themselves, and in some respects they were rich. Though they 
had next to nothing of a "civilized" nature except their cooking utensils 
and carpentry tools, they could raise all the food they needed and .sell 


some, plus plant drugs, mushrooms, and the like on the iiiai-ket or to 
the outside world. They build their own houses and eottins and also sell 
some of the latter. Here their eoffins were very large, utilizing the basal 
portion of at least one Cunninghamia tree three or four feet in diameter 
for each. While we stayed at Suisapa three coffins were made in this 
house. About four families lived in this large house. The room next to 
ours was used for rice storage for the neighborhood. It was sealed u]), 
but ever}- few days it was opened when many people came to draw 
rice. On each side of the house was a row of animal pens, housing cattle, 
horses, and pigs. Great pits beneath the slat tloors collected all waste 
products, which were used for fertilizing the fields. Chickens lived in or 
under the house. Our two rooms had wooden floors, but dirt floors were 
more common. 

The language of these ]ieople was rather different from proper ^lan- 
darin. oi- standard Chinese. Though only a variant, and hardly a dia- 
lect, it was somewhat difficult to carry on conversations. For clothing, 
as in Szechuan Province, the people almost always wore long gowns of 
fairly thick, blue, home-made cloth reaching almost to the ground. This 
was true for men, women and children, and beneath this they usually 
wore jackets and long trousers of the same material. For their consider- 
able idle time, when they smoked their four- or five-foot long pipes, gos- 
siped or stared at us, their garments seemed suitable, but when they 
worked they ai)]X'ared too warm. In the fields, cutting wood, or collecting 
for us. they would often tie up the long gown around the waist. 

These peoi)le did not seem to be given to music as are people of many 
]xirts of China. However, one use for it was quite impressive. A type 
i)\' native communism existed here, whereby quite a bit of the agricul- 
tui'al o]H'rations were cooperative. A group of ten or more people would 
generally cultivate together, with their iron hoes. They all worked in 
unison to the beats of a cymbal and a drum. The leader sang out short 
ehants, and the whole group joined in the cliorus, to the accompaniment 
of the leader's cymbal and the drum of his assistant. The tempo of the 
work was not fast, and there were frequent pauses when all i-ested. 
Though I often heard these musical work-gangs, T only once came within 
close range of one, on a stee]) hillside cornfield. T was prevented from 
getting a picture by curiosity toward me breaking u]^ the routine, and 
by the lateness of the hour. These people rise before daybreak, doing 
much of their work in the early morning and late afternoon, and have 
supper at eight or nine in the evening. They find quite a l)it of leisure 
in the middle of the day. Though these people are Chinese, there is clear 
evidence in their faces and stature that here in these mountains they 
possess quite a bit of aboriginal blood. The aboriginees were apparently 
reduced and absorl)ed as the Chinese pressed into the mountains. The 


imprint of the former remain in rounder faces, thicker lijis, and browner 
skin than possessed by pure Chinese. 

In order to make our efforts more effective, we employed a widow to 
cook and wash clothes for us. Her trachoma (almost universal here) was 
so advanced that her eyesight was very feeble, and the first day she lost a 
small pocket knife and a pair of nndershorts while washing clothes in 
the river. We also frequently found hair, flies or other foreign objects in 
the food, tea leaves in the rice, etc., even though one of the boys super- 
vised most of the cooking. We tried not to think of germs too much. 
We also employed from two to four local boys or young men by the day to 
help us in collecting. It was hard to teach them to collect the small in- 
sects, and most of their catch consisted of the common butterflies, dragon- 
flies, and grasshoppers. They did help by carrying the plant-press, lunch, 
large cyanide jar, and other equipment, but they did not like to work 
every day. 

Our principal emphasis was on insects, particularly those in associa- 
tion with the Metasequoia and other interesting plants of the valley. The 
plants we collected principally for the purpose of identifying the host- 
plants of the insects, and the serial numbers of the plant specimens were 
assigned to the insects collected on them. To all the local people we met 
we advertised our miscellaneous needs, offering to buy all kinds of ani- 
mals. The returns from this method of acquiring specimens were less 
fruitful than I had experienced in some other parts of China, perhaps bo- 
cause these people seemed to have less use for money. This was also re- 
flected in the fact that we could not count on our hired collectors coming 
every day. We often lost time waiting for them in the morning, or per- 
suading some younger boys to go along in their stead. One of our more 
faithful helpers would not go up the main valley past a certain point, 
and we could only assume that it was for fear of being kidnapped to 
help fill the conscription quota of another community. 

Gradually, as a result of making rather high payments for speci- 
mens brought in, we acquired a certain number of snakes, lizards, frogs, 
and birds, but almost no mammals. The people stated that the summer 
was not the season for hunting, and they adhered to their custom. One 
of the reasons given, in addition to the fact that the animals are harder 
to find in summer, was that the summer was the breeding season. This. 
of course, was a commendable viewpoint. 

From questioning the inhabitants, we gathered that tigers, leopards, 
wildcats, bears, deer, mountain goats (serow or ghoral), muntjacs, foxes, 
civets, wild pigs, rabbits, squirrels, and others occurred here. Someone 
told us there was a family of leopards with small young at a certain point 
in the valley, but that was all we heard of them. One day I saw a long- 
tailed, slender, black and white animal which may have been a kind of 


civet. Once one of our boys saw a large dead eat at market, but could 
not bu3' it. The only specimens of mammals we got were moles, mice, and 

Among the common birds in the valley were sandpipers, a large gray 
heron, Chinese pond heron, green heron, egrets, a blackish heron, night 
hawk, black-eared kite, kestrel, dove (Streptopelia), water-cock, ring-neck 
pheasant, golden pheasant, woodpeckers, swallow, water-ouzel, forktail, da- 
yal bird, violet whistling thrush, plumbeous water redstart, gray tit, crow 
tits, scimitar babblers, quaker thrushes, other laughing thrushes, Chinese 
bulbul, yellow-vented bulbul, Chinese finch-billed bulbul, wagtails, bunt- 
ings, tree sparrow, munia, magpie, blue magpie, crow, collared crow, 
tufted mynah, and drongo. 

Among the reptiles here we obtained soft-shelled turtles, skinks, 
water snakes (Enhydris), green grass snakes (Eunjpholus), species of 
Elaphe and Natrix. Of poisonous snakes there were two species of Trimi- 
resurus (pit vipers), one of them a green bamboo snake. One of the latter 
I found at the summit of the highest peak on the west ridge, just after it 
had caught and swallowed a rat. Amphibians here included a large toad 
(Bufo asiaticus), many tree toads, and frogs of several species. 

In our daily collecting we generally divided into two groups, at least 
after lunch, as there were as many as nine of us collecting at one time. We 
attempted to collect along each ravine and ridge, to investigate all types 
of floral situations. We reached the east and west ridges at several points, 
and collected in the high valley to the west several times. Much of our 
collecting consisted of sweeping the vegetation, one species at a time, 
when possible, to collect the insects from each kind of plant. At other 
times we worked on dead branches, fences, logs, stones, streams, and rot- 
ting materials. When there were no trails up to the passes or peaks, we 
had rough going through very dense vegetation (figs. 12 and 13), or had 
to detour. 

When on the higher ridges or peaks we observed from several points 
what appeared to be two high, more or less flat-topped plateaus some dis- 
tance to the south. These began to fascinate me since it was among our 
objectives to find how far south the dawn redwoods extended. Also the 
possibility of other ancient plants or ancient animals having survived 
on a large, high, isolated plateau intrigued me and spurred the imagina- 
tion. Consequently, we made inquiries, and on August 2 I started out 
for the mountains with one of the boys and our host as guide and porter. 
On the same morning my assistant returned to Wan Hsien with a porter 
to ship specimens and send back equipment and food. The others re- 
mained at Suisapa and collected. 

Our trip commenced by following the valley south a few miles and 
then crossing the east ridge at a pass 4,000 feet high. We then followed 



Figure 12. Side valley connecting with the eastern portion of Suisapa Valley. 



Figure 13. Margin 
fields and forests above. 

of the middle portion of Suisapa Valley, showing terraced 


down a stream which went south and gradually became a river (Chen 
River) as it received streams from the large valley to the north. Along 
this narrow valley, which in parts had some vertical cliffs, we found a 
number of small groups of dawn redwoods, some apparently growing 
wild and others in farmyards. Almost w^ithout exception each farmhouse 
had one or more large trees beside it. In the upper part of this valley, as 
in Dawn Valley, this place was generally held by Metasequoia. In other 
places, as farther down this valley, the species was aj^t to be a large 
Cupressxis (no. 2505), ginkgo, or rarely pine (Pinus massoniana or Pinus 
no. 2536). 

The redwoods continued as far down the valley as we followed it that 
day, down to an altitude of 2,400 feet. The next day we saw what seemed 
to be one across the river at 2,200 feet (Tsung-lo). As the altitude lowered, 
pine (Pinus massoyiiana) gradually replaced Cunninghamia as the domi- 
nant conifer on the sides of the valleys, and the cupressine (no. 2505) 
also became more common, not only in the farmyards. The greater scarcity 
of cunninghamias in these lower valleys, however, is at least partly due 
to foresting and denser population. 

As we approached a turn to the south in this eastward-extending valley 
and then crossed on a bridge to the south bank and started to climb the 
ridge, I began to wonder strongly if this trip was really bringing us to the 
liigh plateaus we had observed. We had been surprised when told that the 
mountains about which we had inquired Avere only one day's walk away. 
After climbing a little more than 1,000 feet, we reached the farmhouse at 
which we were to stay somewhat past midafternoon. This was located only a 
short distance below the crest of the ridge. I inquired about the flat-topped 
mountains, for which we had been given the name ''male and female 
(peaks) on big-dike plateau." The answer was that they were just a short 
distance away to the east on this same ridge, hidden by fog. After a few 
moments the fog thinned enough to see two erect finger-like rock forma- 
tions near the eastern end of the ridge. Obviously in trying to describe 
the plateaus to the villagers our remarks about vertical sides, flat tops, 
and two very much alike, had suggested these landmarks, which though 
conspicuous, differed exceedingly in size from our goals. Instead of a 
few thousand, they were a few dozen, feet high, and instead of ten miles 
or more broad they were only several yards broad. There were some other 
similar rock structures on other ridges in the area, l)ut the otliers we 
saw were single. 

I walked up to the ridge and looked south, but could not make 
out my plateaus clearly. I followed the ridge west up to the top of the 
nearest peak, 4,000 feet high. It was covered with trees, but from part 
way up, at a cornfield, I had a view in the right direction. The mountains 
seemed more irregular and less like plateaus. They did not stand out so 


contjpieiiously and there were peaks in front of them, ileturniny along 
the ridge I went in the other direction to the foot of the male and female 
peaks. I hurried back towards the farmhouse, as it was getting dark. In 
leaving the ridge I took a path which led down through the forest to the 
wrong farmhouse. I had trouble inquiring, not knowing that our place 
contained the local school. I rushed down some further trails calling my 
boy, and was finally answered after dark. We spent the evening trying 
to dry out clothing and bedding which were wet from heavy rain dur- 
ing the day's walk. 

We had planned to spend three days on this trip, the second day 
climbing the mountain and collecting, but I decided to push on as far 
as we could go the second day and make it a four-day trip. The next 
morning we started early, leaving our guide to wait for us, and taking 
no mosquito nets or blankets. We crossed the pass and descended to Pa- 
sen-taau Ijy the river below its 90-degree bend around this ridge. We 
found that the good trail was on the other side, but could not cross to it. 
We saw a bridge submerged in the flooding river, a little below. We had 
a good deal of up and down as the path could not follow the river because 
of cliffs. At one point we forded a rushing stream which seemed to flow 
out of the side of the mountain above us. It did, in fact, do that, and 
was the river from Suisapa Valley, which sinks underground about two 
miles below Ilsiaoho and before reaching this ridge. We arrived in 
Tsung-lo, a market town, in the early afternoon and after resting pushed 
on, crossing the swift river in a tiny one-passenger boat, which the boat- 
man carried down to the bank over his shoulders. After passing some 
citrus orchards, we reached a narrowing in the valley where a tributary 
came out of a gorge and met the main river at an angle of over 200 de- 
grees and then entered another gorge. This formed a "Y" with the stalk 
turned to the right, the left arm and stalk being the merging tributaries. 
We crossed the east tributary on a high-roofed wooden bridge across the 
mouth of the gorge and climbed the steep-stepped rock path up the ridge 
between the two narrow gorges. 

On the opposite face of the entrance to the south gorge the very steep 
slope was cleared of its dense vegetation in the middle to expose thin 
strata of hard shale that were tilted almost vertically and exposed at right 
angles. This clearing was planted to young corn. How the farmers reached 
the middle of this near cliff was a puzzle to me. Both the gorges cut through 
solid limestone. 

As we climbed, from the 1,800-foot river level, a downpour tlireatened, 
so we decided to stop for tlie night at a house at the 2,750-foot altitude 
(Lung-keu-po), being out of breath from hurrying. After resting as the 
storm blew past, I went on alone to see how close we were to the plateaus. 
There was a continuing series of peaks of increasing height on the top 


of tliis range. Though 1 scaled one, 3500 feet high, passing thi-ough tall, 
wet grass, corntields, and ])ine groves with braiHl)les, the view was lunn- 
pered and it a])i)eared that we had gone too far to the east and in too 
roundabout a way to reaeh our ol),iective. Along the trail, between these 
conical )>eaks, were some of the serried, sharp limestoiu' projections repre- 
senting ])eculiarly eroded, vertically tilted limestone strata. These had 
also been cons])icu()us on the Chi-o range north of Suisapa. Here there were 
also some deep conical sinks like craters which, however, contained no 
water in spite of the heavy rains. The next market town to the south 
along' this route was Sa-chi, about 12 miles farther. 

After I returned we had a peculiar supper of rice and potatoes boiled 
together, something new to both me and my boy, and string beans. Our 
bouillon helped out. That night neither of us could sleep at all, the bed- 
bugs, fleas, and mosquitoes were so bad. We rose early and walked the 
ten miles to Tsung-lo for breakfast. For our return to the twin peaks, 
we left the river and climl)ed the ridge and followed it to the end. It 
was amazing' to see the extent of rice terracing and the numl)er of farm- 
houses and i^eople on the steep slopes of this high ridge. Some of the 
peaks and some branch ridges to the west, however, were well forested 
with Cunninghamia, birch, beech, oak, chestnut, pine, and other trees. 
Returning to our farmhouse somewhat early and leaving somewhat late 
the next morning, 1 had more opportunity to associate with the people. 
The students of tiie school (all boys) were interested in our insects. They 
all smoked, and many of the younger ones had their small bamboo pipes, 
of various tyi)es, tied to their clothing with a piece of string as our young- 
children might have a handkerchief pinned to their shoulder when going- 
to school. In conversation we learned that seven miles east, at Lung- 
shen-tai in the mountains across the river, there was a group of dawn red- 
woods including one apparently about five feet in diameter. This may 
represent the eastern limit of distribution of 3Ict<isequoia. We returned 
as we had come and completed a somewhat strenuous 100-mile trip. 

The next day we collected high along the west ridge again from Sui- 
sapa, and on the following we moved our headquarters to the Ilsiaoho 
temple, just outside the south end of the valley and a mile south of Ilsiaoho 
market. Actually, the small, rather rundown temple proved to be a nun- 
nery in the sense that the caretakers were three nuns. During the daytime 
the local school operated in a room on our side of the court. AVe had a 
small room nearly filled with three built-in beds for the five of us. We 
barely found room for an old table between the two beds near the single, 
])apered, outside window. Helow our room, side-by-side, were the ])ig])en 
and toilet, and below them the connnon receptacle for human and i)orcine 
waste ])roducts. In a dii't-floored room nearby we built two stone fire- 

Vol.. XXVIII I GRESSITT: />.l U'.V-A'/;/>U'r>r>/> ILX I'EIUTION 49 

places and above them, suspended fi-oin tlie rafters, our roj)es and baskets 
for drying plants and insects. 

That afternoon we hired two new collectors and in liie morning 
ascended the steep peak just north of the temple, which was the western 
end of a ridge extending to the north-south ridge between the twin i)eaks 
and Tsung-lo which we had traversed four days earlier. Though the alti- 
tude was just over 4,000 feet, we had a fair view to the south, as the nearer 
peaks in that direction were not high. From some villagers who had as- 
cended with us, we gathered more definite information about the country 
to our south, though we still did not learn the names of these plateaus. 
Descending the north side of the ridge, we came on some farmhouses 
where the people made bamboo paper by soaking the bamboo culms in lime 
pits under heavy stones. We bought some paper for pressing plants and 
packing insects. In the valley at the foot we reached the other comnnuiity 
by the name of Suisapa, though here there were not so many dawn red- 
woods. In one yard we found a tall, straight, whitish-barked ])ine. (I'inus 
no. 2536) with cones shaped something like those of knob-cone pine, though 
without the knobs. On my return to Wan Ilsien I noticed (juite a few 
young trees of this species just south of ^Mo-tau-chi, tho northernmost 
3Ietascquoia locality. On our way back to Ilsiaoho we were soaked by 
a downpour. 

The next two days we collected in various directions, operating in 
two groups. We found less forest in this area, but a wider variety of col- 
lecting situations. The altitude of the river here was just over 3,000 feet. 
From one of our friends in the market we obtained further information 
about the country to the south, and I planned another trip, this time tak- 
ing the youngest boy in our group, but no guide and no baggage except 
camera, altimeter, pure DDT powder, some food, extra socks, and our 
collecting gear. 

We started early on August 11 after eating some tough roast chicken 
and cold rice, some of which we took along with us. We went south, and 
even a little southwest at first, through a low valley which liad some more of 
the rows of limestone projections, as at Chi-o Shan, and Lung-keo-po. 
There were cornfields and lacquer trees in the valleys as well as on the 
hillsides, as the drainage was apparently not suitable for rice cultivation. 
After crossing a ridge, we headed south over somewhat level country into 
Szechuan Province. At AYen-tu, a market town near the border, we rested 
in a teahouse during the middle of the busy market day. Fi'om hei-e we 
headed southeast along a ridge which dropped down to a low valley. 
From this ridge we had some striking views to the south and east of 
some horizontal stratified mountains etched by gorges and some con- 
si)icuous erect remains of others rising from lower surrounding country. 
AVe descended our ridge obliquely from 3,400 feet, coming down through 


tung-oil trees and corn to the gorge of the Chen River as it narrowed 
and penetrated these massive stratified mountains at 1,500 feet. We were 
many miles down river from Tsung-k), and were close to 40 miles air- 
line south of the ridges at Suisapa, as we covered nearly forty miles walk 
on this first day. Though I did not fully realize it at the time, we were 
actually entering the nearer, and western, of our "plateaus," and the 
gorge cut by the river at this opening was a notch we had seen in the 
middle of its skyline. Instead of a plateau it was a large area of moun- 
tains of thick, horizontally stratified limestone cut by streams into buttes, 
pyramids, peaks or less, but with about the same maximum altitude; 
barely over 4,000 feet. The impression of high plateaus with altitudes of 
perhaps 7,000 feet was an illusion caused by their distance, being massed 
close together with a fairly solid northern face and abrupt eastern and 
western edges, as well as being seen over what was not recognized as hills 
of southwardly decreasing altitude from the ridges over 5,000 feet high 
to the north. I was also influenced by believing that these might be moun- 
tains mistaken by United States Air Force fliers for those having the 
dawn redwoods, and reported as being over 7,000 feet high. 

After entering the gorge for a short distance, we crossed the river in 
a sampan ferry at the mouth of a tributary from the east. The mouth of 
this tributary was peculiar in that though seen as a sizable stream farther 
up, and within view from the mouth, very little water actually passed the 
near dam of gravel just before its mouth. On the other hand, a rushing 
stream came out from a large hole in the foot of the cliff on the north 
side of the mouth, a few feet above the stream level. Continuing up this 
east tributary, we passed many small waterfalls. At one point far up on 
the north side, we noticed a gushing waterfall at the highest visible cliff 
level, seemingly near the top of the ridge, whereas at the lowest cliff 
level there was only a trickle of a waterfall. This could hardly have been 
caused by a very recent downpour on the ridge, or we would have seen 
the lower waterfall increase in size before it was lost to view. The cir- 
cumstance suggested that the water from the higher falls went into a tun- 
nel in the limestone rock and perhaps became the subterranean stream 
which opened at the mouth of this gorge. 

There were quite a number of goats raised in this canyon. The houses 
of the people who tended them were cliff dwellings, or partly so, being 
mostly in cavities formed by overhanging limestone cliffs, walled in front 
with rocks or woven bamboo. This vegetation in the ravine was almost 
semijungle, where it had not been destroyed for grazing or cornfields. 
At some points where it was being cleared, there were cave-ins, or small 

After the canyon widened out to gently terraced rice-field slopes, we 
reached Lin-sui. The elders of this small market urged us to spend the 


night there as it was too late to reach tlie next market over a higli pass, 
but we pushed on and stopped at dusk in a small farmhouse on the way 
up to the pass, near the upper limit of rice and tung-oil trees. The next 
morning- we continued up to the pass at dawn. Some of the steep slopes 
were grassy with scattered Cummighamia trees, and others bore bamboos 
or hardwood forests. We seemed to cross this ridge at nearly its highest 
point, 3,400 feet high, and gradually descended the slopes of a branch 
ridge from a pine grove, through hardwoods, down to tung trees and corn 

Descending furtlier to rice fields we reached the hamlet of Suen-wu, 
where there were a few relatively wealthy-looking homes witli stone walls. 
Here we saw two trees of an interesting conifer (Juniperus no. 2535) 
with drooping branches and long, sharp needles. (We had also seen one 
on the approach to the ridge.) W^e followed down a willow-bordered 
stream, crossing it many times on stepping stones till we reached a point 
where it had to be forded. Being hot and tired we took a swim. As we con- 
tinued, a powerful gusty wind came up in the now broad valley, but we 
reached Pa-se-kwan market just as the shower struck. After a rest and 
a late lunch of our first warm food for the day — rice and hulled corn 
boiled together, stewed melons with a suggestion of pork fat and quan- 
tities of tea — we continued southeast over a little-traveled route. Crossing 
a hill, we were confusesd over our directions, but came to where two boys 
were carrying wood across a river, and found we had to ford it. It was 
muddy and swollen from the recent shower and we had to strip and 
carry our things overhead as the swift water came to our chests and taxed 
our strength. Traversing more hills, partly covered with rice paddies, we 
passed south of a small hamlet, now traveling east. In spite of inquiring 
of each farmer we saw, the simple directions broke down as the tiny path 
curved and branched among paddies or terraced corn or beans. Seeing 
two farmers we made our way to them to inquire, but they refused to 
understand our questions, and thought it a big joke, though we only 
asked the way to the nearest market town. We continued upward towards 
the long ridge to the east, knowing we had to cross it at some point, till 
we found farmers who got us on the right path. Each one we asked quoted 
us a different distance to our destination, but finally we crossed the ridge 
at 2,400 feet and met two porters going our way as it became dark. W^e 
stumbled down the j^atli for nearly two miles after dark until we reached 
tlie market of Pa-hou-keo. We were so tired we dozed until our supper of 
green beans, dried beans, and rice was ready. 

The next morning we left in the early morning fog, and never really 
knew what Pa-hou-keo was like, except that it was a very small market in 
somewhat of a canyon with quite a bit of rice cultivation. We climbed 
again towards a ridge, passing rice paddies, tung-oil groves and some pine. 


oak, and other native vegetation. We breakfasted at a large far-ni house by 
the trail under some large cupressine (no. 2505) trees. From tiie pass, 
at 2,400 feet altitude again, we had a good view to the south. The valleys 
were deeper, and in i)art wider, but still the mountains were of the re- 
peated thiek limestone layers, and with about the same average maxi- 
mum height. One ])rominent vertical peak some distance to the south 
was ])robably the one down on our itinerary as our possible ultimate des- 
tination, though we had already realized that was much farther than we 
would have time to reach. There was by now no (luestion that we had 
been passing through our ''high plateaus," and that my dreams were 
unfounded. These canyons were apparently quite unsuited for the sup- 
l)ort of dawn redwoods, as their bottoms were too low, their sides mostly 
too ]irecipitous, and their tops too dry or too exposed. 

Descending again through forest, then tung groves and corn, we 
reached the Shanghai-Chungking hig'hw^ay in a relatively wide valley 
bottom with a small river at the market of Liang-wei-tan, at 1,100 feet 
above sea level. After a large lunch of noodles and our first eg-gs of the 
trip, we ended our southward march and headed east along the high- 
way towards Chien-chiang, to return north ])y another route. 

The highway wound along the sides of canyons, with some switch- 
backs and tunnels. Our route w'as partly by short-cuts, including a climl) 
over a ])ass. Beyond this pass, the valley leading to Chien-chiang became 
fairly broad, with some large, nearly flat areas of rice fields. The popu- 
lation l)ecame (|uite dense, and there was considerable coolie transport 
along the highway, though we saw no vehicles pass along the road. Hav- 
ing 20 iniles to cover after our near midafternoon start, we did not reach 
the city till well after dark. We were too late to get a regular supi)er, 
so again had noodles and eggs. We then found what was a])parently the 
best inn in the city, at the bus stop. While bathing with a basin of 
Avatei- in our room, and tending to some develoi)ing sores on my feet, we 
had several visits from the authorities who were somewhat put out be- 
cause 1 had left my i)assport in Wan llsien and neither the boy nor I had 
brought along from Ilsiaoho our official letters from the university. The 
only identification we had was my name card. That night we suffered 
severely from bedl)ugs, and I killed over fifty on my bed during several 
occasions when I forced myself awake enough to rise, light the lamp, and 
hunt them down. 

Chien-chiang (Kienkiang Hsien ) is a large town, the largest l)y fai- 
we had seen since Wan Hsien. It is a district city, one of the two ejistei-n- 
most of southeastern Szeehuan Province, and not only is near the Hu])eh 
boi'dei-, but is likewise not very far from both Hunan and Kweichow 
provinces. From here we ascended the south end of a range and ti'a versed 
the side of a large broad canyon with ])artly i)i'ecipitous sides, still of the 


same tliiek, liorizoiital limestone pattern. We then crossed some jtasses 
and descended into anotiier valley with a small i-ivcr wliidi we had to 
ford. Here naked tishermen were easting their wei<>hted Ihrow (umbrella) 
nets and eatehino' .small fish. They would dive down and eateh .some of 
the fish for certain before pulling' in the nets. As we walchcd after oui- 
swim one of them came np with the anterior end of a wiggling nine-inch 
fish in his month, to the annisement of the growing audience. We fol- 
lowed this river to its headwaters, and had to cross it many limes, fii-st 
removing shoes, and as it became small, taking running jumps whei-e there 
were .some partial stepi)ing stones. The frequent wetting did not hel]) the 
sores on my feet, which had originated from bedbug bites excei)t for one 
hlister on the top of my right ankle. During the afternoon we had our first 
heavy rain of the trip, and felt cool, being unprotected from it, whereas it 
had seemed intensely hot in the lower part of the valley at midday. 

We reached our stopping place for the night, a hamlet of four houses, at dark. Our host, an aged, somewhat confused man, w^as not i)articu- 
larly hospitable, as the tiny inn was already overcrowded. The food, 
again, was poor, but the boiled potatoes were welcome once more as a 
change. The inn consisted of two rooms, one the kitchen. My boy and I 
shared one bed, and other guests were mostly on doors and boards in the 
center of the room. The next morning, as usual, we started out in the 
early fog without breakfast. It rained on and off:' during the morning. 
We seemed to be crossing numerous passes between l)ranches of a stream 
system until finally we came out to a large deep valley again. At one 
small hamlet we found some men chopping up two thirds of the left side of 
a pig. and marketing it, though only a few people were present, and it 
was not a market. We bought one catty (liXj pounds), l)ut had to carry 
it for quite a distance before we found a farmhouse where we could cook 
it. Though it was practically the first meat we had seen on the trip we 
ate so many toasted cakes of pounded rice while the meat was cooking 
and the rice being heated that we had to carry nearly half the meat on 
for .supper. 

On towards the middle of the day we passed through a small market 
town, and descended to near the river in this large valley. We were some- 
what confused by being told widely varying distances to our destination. 
The river flowed northeast and then north till it met the Chen River, some 
miles above where we had crossed it four days before and equally far 
below where I had seen it disappear in a deep gorge on the previous trip 
eleven days earlier. We found when we reached the main river that we 
were on the wrong side of the tributary for the regular crossing, having 
been uncertain about our directions wdiere a branch of the trail had lead 
to a ford not long before. However, the boatman managed to ]nish up- 


stream above the branch and then crossed with lis to the northwest bank 
of the Chen, 

We now traveled nortlieast aloni>' the west bank of tlie Chen Kiver. 
The trail passed along steeply terraced rice fields, and we were constantly 
losing our path and taking the wrong paddy embankment. This route 
was apparently a very little-traveled one. Gradually the valley became nar- 
rower until it was a gorge, and we passed along a trail partly cut in the 
clitlf: till we came out to the mouth of a small tributary from the north- 
west. Here we noticed that both the river and the tributary entered 
steep limestone gorges, and between them, on the north, was a large cliflP 
in the middle of the steep slope leading up to the summit of a high ridge. 
I realized we were approaching the terminus i)oint of my previous trip, 
and I supposed we would cross the river here to the east and continue 
on past that point (Lung-keo-po) to Tsung-lo. There was a ferry here 
and a number of people crossed over to our side. We questioned them, 
and though the first few did not seem to realize that we were trying to 
sjjeak their language, we finally learned that they were returning from 
market at Sa-chi, the town we had fallen 10 miles short of reaching on 
the first trip. We also learned that it was shorter to go to Tsung-lo in 
the direction in which they were traveling. I was puzzled as to what that 
way could be, but gradually came to understand, as we climbed up the 
partly forested steep slope and began climbing obliquely up the face of 
the cliff on steps neatly cut in the limestone at a point in the cliff where 
there was an indentation, even possessing some vegetation, mostly small 
trees, shrubs, and vines. Above the cliff was a steep slope of corn, with 
forest near the top. Reaching the forested pass, only slightly below the 
crest of the ridge, we found the altitude 3,100 feet, whereas the river was 
],600. That 1,500-foot climl) after over 30 miles of rough going on sore feet 
was a bit exhausting. 

After resting and eating some roast field corn we continued and found 
that we were going north along a ridge which only gradually decreased 
in altitude. Passing a small hamlet Fu-chia-tang, just before dark we 
found five Metasequoia trees along a path between rice fields at a point 
where the side of the valley became steeper below. This is the south- 
ernmost known limit of distribution for the tree, though it did not ex- 
tend the range more than about ten miles farther south (latitude 29° 45' 
N.), than we found it before. We started down the slope, but as it got 
really dark and the moon was now behind clouds, we gave up the idea of 
reaching Tsung-lo that day, and retraced our steps to the hamlet. In 
asking for a place to stay, we were finally directed to an inn two miles north. 
It developed that it was on a trail to Hsiaoho, and that we did not need to 
go to Tsung-lo. A man guided us along the trail until there were no 
further branches to confuse us, but we found the ]iath full of rocks and 


puddles and thus the going inconvenient. We reached the small inn, a 
lone farmhouse on the trail, while supper was still being cooked, and had 
one of the best meals of the trip : rice, green beans, bean curd, eggs, and 
our pork, not to mention the peppers and cold pickled and peppered 
vegetables which we did not touch. After our 40-mile rough walk we 
were greatly annoyed to have to get up and fight bedbugs immediately 
after retiring. We got some brands from the tire, poured some vege- 
table oil on them and also burned some straw we found, in order to get 
light for searching out the bugs, as we could not find any of the vegetable 
oil lamps and everyone had retired. We just avoided burning the house 
down when w^e ignited our straw mattress. After killing several dozen of 
the bugs we threw the aged quilt to the other end of the room. In spite of 
all the time we spent in the hunt, w^e still had the bugs, and slept poorly. 

Our remaining 13 miles or so to Hsiaoho was done before breakfast (ex- 
cept for an egg apiece), arriving at ten o'clock. AVe passed through 
country similar to that south of Tsung-lo on the other side of the river: 
fairly high, somewhat level country with small hills here and there as 
well as dry sink holes, and the same peculiar rows of erect smoothed 
limestone projections like long rows of sharp canine teeth. This con- 
tinued, through cornfields, practically all the way after getting out of 
the valley in wiiicli we had spent the night, and until we reached the 
trail down which we had started the trip, a few miles south of Ilsiaoho. 
Thus ended our somewhat disappointing 165-mile, 5i/^-day trip. 

Knowing that the porter would have returned before us, from Wan 
Hsien, with equipment and food sent back by my assistant, I was all antici- 
pation for mail, not having received any for a month. Though the porter 
liad returned several days before, to my keen disappointment there was 
no mail, except a note explaining that some mail of ours had been for- 
warded previously to Mo-tau-chi from Wan Hsien, and that the post 
office had been asked to send that mail on to us. AYe never received 
that batch of letters. That evening I received two letters forwarded from 
Chungking, the first we received at Hsiaoho, but neither- was from home. 

The next day, August 17, we moved back to Suisapa, and reoccupied 
our former quarters. We took with us two collectors w^e had trained at 
Hsiaoho, in order to have at least two regular ones to count on. But 
even these generally went back to Hsiaho on market days, so we took 
advantage of that in having them carry back food for us. On the nine- 
teenth part of our group left for Canton, with our accumulated collections 
and three dawn-redwood seedlings. On this day and the following two I 
tried to collect as usual each morning, but found it impossible, as my feet 
had become greatly swollen and the sores very large and painful even when 
I wore straw sandals. I sent the three boys out daily, hiring as much local 
liol]! in addition as ]^ossible, and liad moi-e of the people bring me in 


specimens of any sort they eanie across. By taking care of the i)lant 
drying myself and preparing- labels and other materials I was able to 
keep the boys out in the field lojiger, but still my ti-ouble was greatly 

During this period a Chinese expedition arrived and took up quarters 
in the other wing of the house. The group, led by Professor W. C. Cheng, 
codescriber of the Metasequoia, included Professor K. L. Chu, plant ecolo- 
gist from Shanghai, and Mr. C. T. Ilwa, who had made the first trip to 
collect seeds in 1947. 

On the twenty-eighth my assistant returned. Again I received no mail, 
as the letters that had arrived had been forwarded promptly. The next 
evening, market day, these did arrive, and I had my first mail from home 
in six and a half weeks. The following morning, still unable to walk, I 
left in a sedan-chair that was made from bamboo on the spot that morning. 
Our same porter carried my baggage, including our new collections and 
more Metasequoia seedlings. My sedan-chair was carried by three men : two 
in front and one behind. It was crudely made, without a backrest, but 
actually I found it quite convenient, as far as my feet permitted me to be 
comfortable, since I sat upright on my folded blanket, my feet hanging, and 
could see in all directions. There was a crude framework for a covering, 
but as I had nothing to cover it with, I kept my raincoat handy. The 
framework helped to keep the vines and thorns out of my face, particu- 
larly when I might doze. Constant sitting became monotonous, except for 
the scenery, and I would occasionally doze, even in some of the i>arts along 
small precipices. Once I nearly upset tiie chair. Since we aimed to reach 
Wan Hsien in four days, and since the going was difficult for the men 
on steep trails, we had to travel long hours with few rests. The first night, 
reaching Wang-chia-ying, we had to travel some distance after dark. The 
second night was spent at the market beyond Mo-tau-chi. 

On the afternoon of the third day, just as wo were going to cross 
to the east side of the Mo-tau-chi River after passing through part of 
a heavy downpour, the boatman yelled alarm and told everyone to quickly 
climb the bank. He had seen a crest of water coming in on the tributary 
joining the river 100 yards above just as he was about to pole off. I was 
a bit skeptical, and thought he should have crossed anyway, which he 
could have done, but his return might have been another matter. AVithin 
six minutes the water level below us rose two feet, and was still rising 
when we gave up and went back into the village to find an inn. A])])arently 
in the other valley the downpour had been much heavier, as indicated 
by the color of the sky and the thunder, and we had just experienced 
the edge of it. The rain resumed and continued all night. The next day 
the water was not down enough until early afternoon, when we con- 
tinued onlv about three miles, to find that the water here was still too 


swift for the reerossing to tlie west side. We spent the night in a very 
crowded inn and in the morning were able to move on. However, we did 
not reach the southeast bank of the Yangtze until after dark, and so did 
not arrive in Wan llsien until the forenoon of the sixth day. 

I spent three days waiting for a ship to Hankow, staying at the home 
(jf the Reverend and Mrs. J. B. Matson. With the welcome, good food, and 
comfortable rest, after the long treatment with sulfathiazol. my sores 
began to heal and the swelling to subside. By the time I left to board 
the freighter, a former "LST," I could wear a shoe on one foot, and 
when Hankow was reached three days later, both shoes were worn. After 
24 hours in Hankow, tlu- express train of the Canton-Hankow Railroad 
was boarded, and Canton reached two days later. 

Some of my dawn redwoods were promptly taken down to Hong Kong 
to be planted on the peak under the care of the Hong Kong Forestry 
Department, since Canton was thought likely to prove too warm a 
climate for the survival of the trees. Those sent earlier to Canton, how- 
ever, had already sent out new leaves, so the climatic tolerance appeared 
actually not to be so restricted. These and other trees brought by our 
main group, as well as the seeds planted, all grew and were thriving when 
I left Canton two and one-half years later. 

During the three winters that I remained at Canton it was interest- 
ing to watch the reaction of the young trees to the much warmer winters 
than those to which the tree is exposed in its natural habitat. In Canton 
(latitude 23° 06' X.), the temperature rarely drops as low as 36°F., and 
often is 70° or higher during December or early January, though cooler 
in February and whenever the wind blows down from North China for 
a period. Instead of shedding the needles in the autumn, the needles 
browned slowly and partially as the temperature dropped in winter. 
This happened in an irregular fashion, with apparently different reac- 
tions in different seedlings, some becoming brown when others were still 
almost entirely green. We frequently suspected certain plants were 
dying, only to be proven wrong later. The needles rarely actually dropped 
until the coldest winds in February, after which they were very soon 
replaced by new needles a]ipearing as soon as the temperature began to 
rise. Actually we had only about five of some thirty seedlings die. The 
percentage of germination of the seeds planted was rather low. 

Some of our group remained at Suisapa for three more weeks and 
then moved to Wang-chia-ying for five days. En route back to Wan Hsieii 
they stopped one day near the top of the Chi-o Shan (Seven Humps), 
three days at Mo-tau-chi, and two at Lung-chu-pa for general collecting. 
They spent a few weeks near Wan Hsien. Then they made a trip all the 
way back to Suisapa to collect Metasequoia seeds and to take more pho- 
tographs, with a camera I sent by airmail from Canton to Wan Hsien. 


They experienced snow while returning over the Chi-o Shan ridge, and 
returned to Canton in mid-November, after an absence of nearly five 

Our collecting resulted in the bringing back of tens of thousands 
of insects in addition to the plants and miscellaneous animals. The col- 
lections are at the California Academy of Sciences. As to the scientific 
results, it will be some time before any conclusions can be drawn as to 
possible relationships of the insect fauna of the dawn-redwood flora 
with that of western North America. It may prove more closely related 
to that of southeastern North America, as is the case with most of the 
plant genera in common between eastern Asia and North America. In 
the meantime specimens of different groups must be studied by many 
specialists. The few groups studied to date have not demonstrated any 
significant Nearctic relationships. Some of the material has been reported 
upon in some of the references cited below. 


Chanev, R. W. 

1948. Redwoods around the Pacific basin. Pacific Discovery, vol. 1, no. 5, pp. 

1951. A revision of fossil Sequoia and Taxoclium In western North America 

based on the recent discovery of Metasequoia. Transactions of the 
American Philosopliical Society, new series, vol. 40, pt. 3, pp. 171- 
263, pis. 1-12. 

Florin, Rudolf 

1952. On Metasequoia, living and fossil. F,otaniska Notiser, hafte 1. pp. 1-30. 

Gressitt. J. L. 

1950. The hispine beetles of China (Coleoptera: Chrysomelidae). Liniinan 

Science Journal, vol. 23, pp. 53-142. pis. 4-8. 

1951. The longicorn beetles of China. Longicornia, vol. 2, pp. 1-667, 22 pis., 

1 map. Paul Lechevalier, Paris. 

1952. The tortoise beetles of China (Chrysomelidae: Cassidinae). Proceedings 

of the California Academy of Sciences, vol. 27, pp. 433-592, pis. 27-36, 
1 text flg., 4 maps. 

Merrill, E. D. 

1948. Metasequoia, another "living fossil." Arnoldia, vol. 8, pp. 1-8, 2 pis., 
1 map. 

Stebbins, G. L., Jr., 

1948. The chromosomes and relationships of Metasequoia and Sequoia. Science, 
vol. 108, pp. 95-99. 

Wym.\n, D. 

1951. Metasequoia brought up to date. Arnoldia, vol. 11, pp. 25-28. 




Vol. XXVIII, No. 3 July 15, 1953 







Department ef Entomology and Parasitology 
University of California, Berkeley 

During the months of December, 1950, to and including April, 1951, 
Dr. A. E. Michelbacher and his wife, Martha, and Dr. E. S. Ross and his 
wife, Wilda, conducted an intensive insect-collecting expedition in Chile, 
in particular, and less extensively also in Argentina, Bolivia, and Peru 
with some minor collecting while en route by boat along the coast of Cen- 
tral America and Colombia. This expedition was a part of the activities 
of the California Academy of Sciences which institution arranged for the 
ocean transportation and supplied a truck for travel on land. On October 
17, 1950, the party sailed from San Francisco on the Grace Line steam- 
.ship Santa Juana — destination: Valparaiso, Chile. Their ship stopped 
at various ports on the voyage and small collections were made at Buena- 
ventura, Colombia; Callao, Peru; and other ports. Upon arrival at Val- 
paraiso the party was cordially received by the veteran Chilean ento- 
mologist, Dr. Edwyn P. Reed, and by Dr. Raul Cortes of the Chilean Min- 
istry of Agriculture, who rendered the Americans invaluable hospitality, 
information, and assistance during their stay in Chile. 

Although largo collections were made of all kinds of insects, of which 
thousands were taken. Dr. and Mrs. Michelbacher also gave special atten- 
tion to collecting as many species of aphids as time and opportunity per- 
mitted. By much hard work they procured a sizable assemblage of species 



which were preserved in alcohol and turned over to the writer for mount- 
ing and study, and for the publication of desirable information concerning 

Little previous investigational work has been done on the aphids of 
Chile, in which countr.y by far the greatest numbers of individuals and 
species were taken. 

To the present time there have been relatively few technical papers 
on the aphids of western South America. In Chile, where the greater 
part of the insect collections were made, there has been, so far as I know, 
only a single new species. Aphis citricola van der Goot (1912), described 
and this has been found to be the already described Aphis citricidus Kir- 
kaldy, 1907 {Myzus). 

In Argentina and Brazil a number of able aphidologists have pub- 
lished many important contributions on the systematics and economics 
of the Aphidae in those countries as the elaborate bibliography included 
herein testifies. It is to be expected that interest in this field of ento- 
mological endeavor will rapidly expand in the future. 

Because of the very convenient method of travel, by means of a spe- 
cially fitted Ford panel truck, the party penetrated areas rarely visited 
by foreign insect collectors and most of the specimens were taken in the 
wilds of the countries traversed. Thus their undivided time and attention 
were given to their objectives which proved very fruitful in the way of 
securing splendid insect collections. They remained in Chile from No- 
vember 25, 1950, until February 4, 1951, when they started on the trip 
over the Andes via Paso de Bermajo en route to Mendoza, Argentina, and 
thence eastward and northward through the cities of Rio Cuarto, Cordoba, 
Tucuman, Salta, Jujuy and La Quiaca, Argentina ; through Camargo, Po- 
tosi, Oruro, and La Paz, Bolivia ; via Lake Titicaca, Puno, Tinta, Abancay, 
Ayacucho, Huancayo, Oroyo and Lima, Peru. They arrived at Lima, March 
4, 1951. During the remainder of their stay they travelled about 550 miles 
in the coastal area north of Lima through Huacho, Trujillo, Chiclayo, 
and to a point north of Olmos. Dr. and Mrs. Michelbacher and Mrs. Ross 
sailed homeward from Lima on the S.S. Santa Flavia on April 6 and 
Dr. Ross followed about a month later. 

For collecting Aphidae and other small insects Drs. INIichelbacher 
and Ross had a large supply of small glass vials. Specimens of aphids 
and often portions of the host plants, and labels bearing the date and 
locality, as nearly as the latter could be ascertained, were preserved 
together with each lot. These specimens have since been mounted and 
determined as to species as far as possible. 

In cases where species collected have also ])een recorded by ento- 
mologists in the South American countries visited: Colombia, Peru, Chile, 


Argentina, and Bolivia — these have been noted in so far as available litera- 
ture permitted. 

I wish to express my thanks and great appreciations to Dr. and ^Irs. 
A. E. Michelbacher, and to Dr. and Mrs. K. S. Ross for the o])portunity 
to mount and study this fine collection of South American ai)hids. The 
large series enabled me to prepare an excellent collection of types, para- 
types, and other specimens for the department of entomology of the 
California Academy of Sciences and duplicate specimens for my own 
collection. I am grateful to Dr. E. L. Kessel for making important sugges- 
tions for the preparation of the manuscript and to Dr. Robert C. Miller, 
director of the Academy, for the publication of this paper, and especiallj^ to 
my wife, Marie, for typing, correcting, and editing it. 

I also wish to thank L. A. Bahamondes, Director General de Investi- 
gaciones Agricolas, Ministerio de Agricultura y Ganaderia, Republica Ar- 
gentina, for furnishing many specimens of Aphidae collected in Mendoza 

The total collections of aphids amounted to 25 genera ; 55 species, in- 
cluding 14 new species; and 3,395 specimens mounted on 781 slides. 




Superfamily APHIDOIDEA 



Tuberolachnus saligna (Gmelin) 

Giant Willow Aphid 
(Figure 1) 

Aphis saligna Gmelin, 1788; A. polaris Curtis, 1828; A. viminalis B. d. Fonsco- 
LOiiBE, 1840; Lachnus dentatus Le Baron, 1872; L. fuliginosus Buckton, 1891; 
Tuberolachnus viminalis (B. de Fonscolombe) Mordvilko, 1908; Pterochlorus 
saligna (Gmelin) Theobald, 1929. 

This large aphid may be readily recognized by the conspicuous pointed 
tubercle near the middle dorsum of the abdomen. It is widely distributed 
throughout the temperate regions wherever species of Salix occur and it 
has long been known as a pest of basket willow (Salix viminalis) in Europe. 
From the records it appears that this species occurs throughout the gen- 
eral habitat of Salix spp. ("Sauce") (SALICACEAE). Collections were 
made as follows : 



Figure 1. Giant willow aphid. Tuberolachnus saligna (Gmelin). Antennal seg- 
ment III of alate (top) and aptera, tip of rostrum, cornicle, cauda, and anal plate 
and abdominal tubercle all enlarged. 

Chile : 

On Salix sp. at Angol, Province of Bio-Bio. January 1, 1951; apterae 
and alatae. 

Argentina : 

On Salix sp., Uspallata, Province of Mendoza, Februai-y 6, 1951. ^NFany 
specimens of apterae and alatae. Other records by Blanchard 
1926, pp. 327-29; 1939, 871-72; 1944, 16) are: 

On SuUx sp., Tigre, Province of Buenos Aires; also from San Juan 
and Zapala. 


On English walnut (nogal or noguera) Juglans rcfjio Liiniaeus at San 
Rafael, Province of Tncnman. 

i'ERU : 

On Salix sp., Barranca, March 15, 1951 ; all a])terae. 
All collected by Dr. A. E. Michelbacher. 


Subfamily SIPHINAE 

Sipha flava Forbes 

Yellow Sugarcane Aphid 

This is a subtropical yellowish species occurring on (Jramineae in 
warm, temperate and tropical North and South America. Although not 
taken by Dr. ]\Iichelbacher the species has been reported in two of the 
South American countries where sugarcane is grown. 

Argentina : 

Blanchard (1944:17) records this species on sugarcane (caiia de 
azucar) in Tucuman. The writer has also received a good series 
from L. A. Bahamondes collected on Holcus halepensis (Lin- 
naeus) {Sorghum sp.) (sorgo, zahina), at Mendoza, April 9, 1951. 



Chileaphis P^ssig, new genus 

(Figure 2) 

AJatc oviparous female: Head with straight front; antennae 6- 
segmented ; imbricated ; hairs few and short ; segment III as long as IV 
and V ; unguis very short ; primary sensoria circular, rimmed with hairs : 
secondary sensoria linear or semicircular. Compound eyes well developed 
and with posterior ocellar tubercles. Wing venation aphis-like with radial 
sector only slightly curved and free at base. Legs unusual in that the 
alate oviparous females have numerous sensoria on the tibiae of all three 
pairs. Cornicles truncate cones and the diameter at the base exceeds 
the length ; with few hairs. Cauda appears like a chitinized lobe which 
may be recurved. Anal plate with deep median constriction. Ovarian eggs 
are globular and oval. 

In the alate male and apterae the cauda is almost globular with a 
narrow neck and wide base. Antennae of the male have both transverse 
and circular secondary sensoria on segments III, IV, V, and VI. Dorsal 
wax glands are present on all forms. Genotype: Chileaphis ynichelhacheri 



Figure 2. Chilean cypress aphid, Chileaphis michelhacheri Essig, new species. 
a, alate sexual female; b, apterous parthenogenetic female; c alate male. Anten- 
nae, cornicles, anal plate and cauda, and male sexual organs. 


Chileaphis michelbacheri Essig, new species 

Chilean Cypress Aphid 
(Figure 2) 

Type : Alate oviparous female : Pale, with the head, thorax, transverse 
glandular areas on abdomen, cornicles, and cauda, black or dusky. In 
life the body is probably partially covered with white powdery wax. An- 
tennae imbricated and with few short hairs. Secondary sensoria on seg- 
ment III linear and a few may be semicircular, scattered over the full 
length, 25-25 in number. Primary sensoria circular or oval, fringed with 
short setae. Apical sensoria on VI are compound with at least 2 small 
marginal ones. Lengths of segments: III, 0.63 mm.; IV, 0.35 mm.; V. 
0.25 mm. ; VI, 0.21 mm. Rostrum with the apical segments short and 
blunt; extends to second coxae. Sexual sensoria numerous on the tibiae 
of all legs. Wings well developed and with aphis-like venation. Radial 
sector only slightly curved. Second fork of media arises half-way be- 
tween first fork and wing margin. Abdomen with six distinct pigmented 
glandular transverse areas. Cornicles dusky, as illustrated : with few 
short hairs. Cauda unusually large and peculiar, dusky, chitinized and 
with only a few short hairs. Anal plate bilobed, with many short setae. 
Genital plate as drawn. Length of body, 2.45 mm. ; forewing, 2.8 mm. 

Aj)terous parthcnogenetic female: Whitish-gray; in living condition 
ai^parently covered with powdery wax ; widely oval with short antennae 
and legs as delineated. Lengths of antennal segments: III, 0.28 mm.; IV, 
0.09 mm.; V, 0.10 mm.; VI, 0.12 mm.; unguis nail-like. Cornicles truncate 
cones, the base about twice the diameter of the opening, with few short 
scattered hairs. Cauda large, gloliular, with slender stalk and wide base; 
with few short hairs. Anal plate deeply cleft into two lobes; with few 
short hairs. 

Alate male : Similar in color and general appearance to the alate fe- 
male but much smaller, being 1.71 mm. in length. The antennae are as 
illustrated. The sensoria are numerous, smaller and more oval and linear 
than those of the alate sexual female and occur on all the segments ex- 
cepting I and II. The globular cauda is illustrated in the turned-up 
natural ])osition and the sexual organs and cornicles are as drawn. 

Host plant and locality: This interesting new genus and species was 
collected by Dr. A. E. ]\Iiehelbacher by beating the available limbs and 
foliage of the Chilean cypress, Pilgerodenclron uviferum (Don) Florin. 
The synonymy of this host plant is given by Record and Hess (1913) as 
follows: (Juniperus uvifera Bon = Thuja tetraejona Rookev— Lib ocedrus 
tetrafjona | Hooker] Endl. = L. ciipressoides Sargent = L. uvifera [Don] 



Locality. Los Muermos, Province of Llanquihue, Chile. It was col- 
lected only in this single locality by Dr. Michelbacher for whom the spe- 
cies is named. 

The total number of individuals examined consisted of 50 apterous 
parthenogenetic females, 38 alate oviparous females, 34 immature apterous 
and alate forms, and 2 alate males, all mounted on 25 slides. 

This species differs from other known related species in having sen- 
soria on the tibiae of all legs of the alate oviparous female and the pres- 
ence of both circular and transverse sensoria on the antennae of the alate 

Neuquenaphis' michelbacheri Essig. new species 

Nothofagus Aphid 
(Figure 3) 

id.i_l^i^j^: s:5igznrg::= z^a- ^ :' ' .'' :-! ii' u simriZisS!2UZSj^ 

Figure 3. Nothofagus aphid, KeuquetiaiJhis niivhelbacheri Essig. new species, 
a, alate parthenogenetic female together with head and antenna, rostrum, some 
tubercles and spines on dorsum of abdominal segments I-II. anal plate and cauda. 
and cornicle; b, young; c, apex of one of the body tubercles. 

1. From the Province of N'euqiien, Argentina. 


TvPE: Alate parthenogenetic female: Color uncertain, the cleared alco- 
liolic specimen indicates that the head, antennal segments I and II, and 
thorax are black and the legs, cornicles, tubercle bases, cauda, anal plate, 
and abdominal glandular areas are dusky. It is probable Ihal in life ;i 
fine powdery wax may cover portions of the body. The illustration, to- 
gether with the description of the genus, will give a good idea of the im- 
portant characteristics. [It is well to point out that the aj^terous form 
on the lower right is probably a second or third instar young, but its pe- 
culiar tubercles will nevertheless assist in recognizing the genus and 
species.] Of special interest are the tuberculate first segments of the an- 
tennae; the arrangement of the oblong sensoria — 11-15 near the middle of 
segment III ; the very long flagellum ; the large, distinct tooth-like structure 
on the inner margin near the apex of the fore femora; the faintly clouded 
wing veins which do not show in the drawing; the arrangement of the 
spined body tubercles, as figured; the short reticulated cornicles and the 
rather unusual type of cauda should make the species easily recognizable. 
In other alate paratypes the number of sensoria may vary from 8-lfi. 

Host plant and locality : The species was collected by beating the 
branches of Nothofagus domhei Blume. It was collected along with Spica- 
phis niichelhacheri and Neiupienaphis chilrnsis on the Niagara Rancho 
near Temuco, Province of Cautin, Chile. Thirteen apterous sjieeimens 
and 3 alates were also collected in San Andres, Paranque, Chile. Collector, 
A. E. Michelbacher. 

The specimen described was selected from 16 adult alate i>arthenogentic 
females and has been designated as the type ; all others as paratypes. There 
were also 3 immature specimens in the lot. They were mounted on 13 slides. 

Xeuquenaphis michclhacheri differs from N. edwardsi (Laing) {Myzo- 
callis) in the following respects: 

1. Head and thorax much narrower. 

2. Presence of small setae-bearing tubercles on the pronotum. 

3. Much shorter tubercles on dorsum of abdominal segments I and 

4. The presence of a prominent tuberculate seta or spine on the 
outer surface of the cornicle. 

5. Somewhat more elongated cauda. 

Neuquenaphis chilensis Essig, new si>ecies 

Dombei Aphid 
(Figure 4) 

Type: Ahife parthenogenetic female: Color uncertain because speci- 



Figure 4. Dombei aphid, NeuquenaiJhis chilensls Essig, new species, a, tip of 
antenna; b, body tubercle; c, frontal tubercles; d, antenna sliowing secondary 
sensoria; c, rostrum; f, cornicle; g, cauda and anal lobes. 

mens were preserved in alcohol and may have faded somewhat. Mounted 
ones are brownish and black and quite similar to the alates of N euqueriaphis 
michelhacheri. All of the body, excepting the abdomen, is dusky or brown. 
The cornicles, anal plate, cauda, and body tubercles are also brown. The 
most outstanding characteristic of the species is the large number of blunt, 
fleshy, cylindrical tubercles each of which is terminated with one or two 
short, slender setae. They vary considerably in size and are arranged 
somewhat as illustrated. The antennae are long and slender — extending 
almost to the tips of the wings ; imbricated ; 6-segmented ; III much longer 
than either IV or V, but not as long as both together ; IV is longer than V ; 
the unguis of VI is li/i times the length of the base. Segments III have 
6-6 large elongate sensoria which extend to the full width and are located 
near the middle. (In three paratypes the ratio is 5-5, 6-6 and 6-7.) The 
rostrum is fairly broad and extends to the second coxae. The legs are 
sparsely clothed with short, fine hairs. The wings are aphis-like and the 
veins are cloudy. The abdomen is beset with numerous tubercles wliich 
are variable in size and with small grouped dark wax glands ; also there are 
five bands of small setae. The cornicles are short and somewhat truncate, 
with a single lateral marginal spine and somewhat imbricated; the base 
is twice the diameter of the apex. The cauda is conspicuous; knobbed. 


being- widest at the base, thickly beset witli small piniple-like areas and 
large recurved spines supported by raised conical bases. The anal plate is 
shallow-divided; the two lobes are spined much like the cauda. Length of 
body 2.20; forewing 2.60 mm.; antennae 1.30 mm.; cornicle 0.10 mm.; 
Cauda 0.15 mm. 

This species differs from Neuquenaphis michelhachfri and .V. nJirardsi 
in having tubercles on the head and thorax; capitate cauda; very much 
shorter antennal unguis; and fewer antennal sensoria. 

Host pJcmt and locality. This species was collected along with the N. 
michfUmcheri by beating the brandies of Nofhofagus domhei at Hacienda 
San Andres, near Purranque, Province of Osomo, Chile, January 15, 
1951. by Dr. A. E. Michelbacher. 

The type was selected from three alate specimens, all that were 


Spicaphis Essig, new genus 
(Figure 5) 

Apterous viviparous female : Robust, the dorsum of the entire body 
beset wdth thick, rugose, glandular-like spicules or tubercules arranged 
for the most part in longitudinal and transverse rows ; each spicule has a 
terminal glandular spine or seta. Similar spines also occur on the head, 
two basal antennal segments and cornicles. Antennae 6-segmented; eyes 
compound with terate tubercles. Cornicles cone-shaped and with a single 
tuberculate spine. Cauda knobbed. Anal plate with shallow median con- 
striction. Genotype: iSpicaphis michelbacheri Essig. 

Spicaphis michelbacheri Essig, new species 

Miehell^acher Aphid 
(Figure 5) 

Type : Apterous female : Alcoholic specimens grayish with pigmented 
areas as shown in the illustration. The most important feature is the pres- 
ence of numerous stout, somewhat curved, rugose tubercles arranged in 
what appears to be a definite pattern on the dorsum. Each of these ter- 
minates in a glandular seta. There are approximately 100 of these present. 
Between the body segments there are a series of small black glandular ( ?) 
areas. On the front of the head there are a pair of rather blunt tubercles, 
each bearing two glandular setae. The antennae are 6-segmented ; the two 
basal segments with glandular setae and a few short spines on the other 
segments. The permanent sensoria are circular, fringed, and the apical 




Figure 5. Michelbacher aphid, Spicaphis iinchelhucheri Essig. new species. 
Apterous parthenogenetic female with enlargements of head and antenna, rostrum, 
head tubercle, anal plate and cauda, body tubercles, and cornicle. 

one on segment V is compound. Segment III is longest; IV, V and VI are 
siibequal ; the base and unguis of VI are also equal in length. The eyes 
are compound with well-developed terate tubercles. The legs are rela- 
tively short with very short hairs. The rostrum extends to the third 
coxae. The cornicles, cauda, and anal plate are as illustrated. Length 
of body, 1.92 mm. ; antennae 1.26 mm. 

Described from a single apterous specimen which is designated as 
the type. 

Host plant and locality : The specimen was obtained ])y ])eating the 
limbs and foliage of Nothofagus domhei Blume, a tree native to southern 
Chile. It was taken along with Ncixini uaphis michelhaeheri described 
elsewhere. The collection was made at Los Muermos in the Province of 
Llanquihue, Chile, on January 15, 1951, by Dr. A. E. Michelbacher for 
whom the species is named. 

Type : A single apterous female so designated. 

It is barely possible that this may prove to be the apterous form of 
Neuquenaphis chilensis; the chief similarities being in the antennae, cor- 
nicles, and cauda. It differs in having a much wider body and niore nu- 
merous bod.y tubercles. The true apterous forms of N. chilensis are 




Subfamily APHIDINAE 

Tribe Rhopalosiphonini 

Hyalopterus arundinis (Fabiieius) 

Mealy Plum Aphid 

(Figure 6) 

AiJhi.s ijniHi Gkoffkov. 1762; A. arundinis Fahkku s. 1775; Hydlo/jterus pnini. 

Figure 6. Mealy plum aphid, Hyalopterus arundinis (Fabricius). Alate and 
apterous females with enlarged antennae, rostrum, cornicles, and caudas. 

This aphid is very common and often injurious to prune trees in CHILE 
according to the observations of Dr. Michelbacher, who, because of its 
abundance, did not make any collections of it. 

It has been reported in Europe, Asia, Africa, Australia, North America, 
and now in South America. It overwinters in the egg- stage on plums. 



prunes, apricots, apples, peaches, and related plants, and usually spends 
the summer on reed grass, Phragmites phragmites (L.) and cat-tail (Typha 
spp.). The eggs are laid on trees of the genus Prunus and the spring gen- 
erations occur on them. It is often injurious to the hosts and excretes 
quantities of honeydew over fruit and foliage. 

Rhopalosiphum' maidis (Fitch) 

Corn Leaf Aphid 

(Figure 7) 

Figure 7. Corn-leaf aphid, Rhopalosiplium maidis (Fitch). Alate and apterous 
parthenogenetic females with enlarged antennae, cornicles, and caudas. 

2. The ending "on" is the correct form for the Latin siphon. 


This aphid is widely distributed throughout North and South America 
and in many of the South Pacific Islands. It is frequently associated with 
cultivated corn but also feeds on sedges {Carex spp.) and Scirpus spp., 
Typha spp., many grasses, Zea mays L., and other plants. Collections were 
made as follows : 


On Johnson grass, Sorghum halepense, Callao, November l-t, 1950 — 
very abundant. 

On cultivated corn (maiz), Callao, November 17, 1950 — 1 alate and 
5 apterae. 

On cultivated corn, Cusco, Province of Cusco, March 4, 1951 — many 
apterae and alatae. 

On cultivated corn, Chiclayo, Province of Lima, March 21, 1951, 
apterae only. 

Argentina : 

On corn, Zea Mays L., Arena spp., Hordeum sp., Saccharum spp. 
In Argentina it is a vector of the mosaic of sugarcane according to 
Blanchard (1939:907-98; 1944:19). 

Rhopalosiphum nymphaeae (Linnaeus) 

Waterlily Aphid 
(Figure 8) 

Aphis nymphaeae Linnaeus, 1761. 

This is a Holarctic species widely distributed in Europe and North 
America. It overwinters on members of the genus Prunus and spends the 
summer on many aquatic and other host plants. It was collected only once. 


On Canna sp. in the Botanical Gardens, Lima, November 15, 1950 — 
many alatae. 

Rhopalosiphum prunifoliae (Fitch) 

Apple-Grain Aphid 
(Figures 9-10) 

Aphis pru7iifoliae Fitch, 1855; Rhopalosiphum pseudoavenae Patch, 1917. 

This aphid is a widely distributed economic species in North America 





Figure 8. Waterlily aphid. Rhopalosiphum nymphaeae (Linn.). Alate and ap- 
terous parthenogenetic females with enlarged antennae, cornicles. Cauda, and reticu- 
lated area on the dorsum of the aptera. (After Zimmerman. 1948.) 

where it infcst.s grains, corn, and gra.sses. In the eokler areas it may 
overwinter on apple and other deciduous fruit trees. 


Sweeping', Los ^Muerinos, Province of Llanquihue, January 20, 1951 
— 3 ahites. 

Peru : 

Beating plants at Andahuaylas, Province of Apurimac, February 7, 

1951—1 alate. 
On cultivated corn, Zea mays L., Cusco, March 4, 1951 — apterae only. 
It is a serious pest of sweet corn in California. 



Figure 9. Apple-grain aphid. Rhopalosiijhuvi prunifoliae (Fitch). Colonies on 
an ear of corn. (Photo hy Oscar Bacon, August 31, 1951.) 

Argentina : 

It is possible that some of the records of Biwpcdosiphum pseudoavence 
(Patch) on rye (centeno), maize (maiz), wheat (trigo), and Bromus sp., 



in Argentina, may refer to tliis species. Blanchard, 1939:908-1)11; 

Figure 10. Apple-grain aphid, Rhopalosiphum prunifoliae (Fitcla). Alate and 
apterous parthenogenetic females and enlarged antennae, caudas, cornicles, and 

Rhopalosiphum pseudobrassicae (Davis) 

Turnip Aphid 
(Figure 11) 

Aphis pseiidohrassicae Davis, 1916. 

This may l^e an Asiatic species which has been widely distributed by 
commerce. The writer has specimens from China (Nanking, Pehpei, Hang- 
chow), Territory of Hawaii, Iraq, Egypt, Uganda, and North America. 

Bolivia : 

On a native Nasturtimn sp. growing in a garden at Potosi, Province 
of Potosi, February 22, 1951 — -many apterae and 1 alate. 

Argentina : 

On Brassica nigra Koch and B. rapa L. in the Province of Buenos 

Aires. (Blanchard, 1939.) 
On Lepidium sp., (alheli), stocks of gilliflower (Mathiola hederacea) 

and (rabanito) or radish {liaphanus sativus L.). 




On cultivated stocks, Methiola sp., at Callao, J'rovince of Lima, No- 
vember 4, 1950 — many apterae and alatae. 

Figure 11. Turnip aphid, HhoiJalosiphum pseudohrassicae (Davis). Alate and 
apterous partlienogenetic females with enlarged antennae, cornicles, genital plate, 
anal plate and cauda, and rostrum. 

Rhopalosiphum splendens (Theobald) 

Subterranean Aphid 
(Figure 12) 

Siphonocoryne splendens Theobald, 1914; Rhopalosiplium subterraneum Mason, 

This ai)hid is chiefly a root-infesting species occurring in many parts 
of North America, in Hawaii, in Africa, and now collected for the first 
time in South America, and so far only in Peru. 

Peru : 

Feeding on roots of grasses (?) under stones in the bed of the Red 




Figure 12. Subterranean aphid, Rliopalosiphum splendens (Theobald). Alate 
and apterous parthenogenetic females with enlarged antennae, cornicles, rostrum, 
genital and anal plate, and caudas. 

River, Callao, November 16, 1950 — 8 alatae and 3 apterae. Col- 
lected by Dr. E. S. Ross. 
Beating nightshade (SoJanum sp.) at Andahuaylas, Province of Apu- 
rinac, February 7, 1951 — a single alate specimen. 

Tribe Aphidini 

Aphis alstroemeriae Essig, new species 

Alstroemeria Aphid 
(Figure 13) 

Type: Alate j)arthenogenetic female: ^Fostly black; the abdomen paler 
with black setal patches and areas around spiracles and two transverse 
bands posterior to the cornicles. Antennae with short spines and sen- 

Vol. XXVIII 1 




1» Se- e^c' .' Sf .• ,, 

Figure 13. Alstroemeria aphid, Aphis alstroemeriae Essig, new species. Anten- 
na of alate; rostrum; cornicles and caudas of alate and aptera; aptera showing dor- 
sal reticulated pattern. 

soria as drawn; unguis nearly three times the length of the base; circular 
secondary sensoria on segment III of different sizes; 9 to 11 in number; 
IV with to 2. (In paratypes the variation is III, 5 to 11 ; IV, to 2.) 
Rostrum slender and short, extending to the third coxae. Prothoracic 
tubercle present. Wings normal, second fork of media arises near the 
middle of the distance from first fork to margin of wings. Legs with 
scattered spines the length of which is less than the diameter of the 
tibiae ; hind tibiae darker at apices ; tarsi noticeably long. Cornicles nearly 
cylindrical, being broader at base, imbricated, and with slightly flanged 
opening. Anal plate rounded, spiculate, and spined. Cauda with wide 
base, the apical portion almost parallel-sided. Length of body 1.75 mm. ; 
hind tibiae 0.10 mm. ; cornicle 0.17 mm. ; cauda 0.14 mm. ; antennae 1.29 mm. 

Apterous partJienogenetic female : Robust in form and generally dark 
in color. The dorsum has a mosaic pattern of lines and dark and lighter 
usually 5-sided tile-like areas which are most pronounced on the dorsum, 
especially posteriorly. Antennae dark excepting III, IV and base of V. 
Legs dusky or black with the basal three fourths paler. Rostrum, cornicles, 
Cauda and anal plate black. Lateral tubercles present on some of the 
abdominal segments. Length 1.57 mm.; hind tibiae 0.91 mm.; cauda 0.17 
mm. ; antennae 1.55 mm. 

Xumher of specimens : The series consists of 34 mature apterous and 
23 alate females. A single alate specimen on a slide with 3 other alates 
and 6 apterae has been designated as the type. 

Host plant and locality : The specimens were collected on a native species 
of the so-called Chilean lily, Alstroemeria sp., 40 kilometers east of Los 
Andes, Province of Aconcagua, Chile, February 4, 1951, by Dr. A. E. 

This species differs from Ai)his medicaginis (Koch) in having secondary 
sensoria on antennal segment IV and from A. rumicis Linn, in having- 
fewer hairs on the cauda. 



Aphis citricidus (Kiikaldy) 

Tropical C'itrus Aphid 
(Figure 14) 
„ nmcM,. K,»K.u,„v. 1M7; AvMs t.varesi D.l Oeboo. WOS; AvKis citri- 
cola Van dkr Goot. 1912. 

Figure 14. Tropical citrus aphid, Avhis cUricidus (Kirkaldy). 

This is a medium-sized black aphid which is often very abundant on 
and injurious to the tender apical shoots of its many host plants, it 


chiefly occurs in the tropical and warin-tempcrate regions where it 
feeds mostly upon Camellia, Citrus, Hibiscus, Mangifcra, and other plants 
in the more tropical areas of Japan, China, Pacific Islands, Australia, 
Africa, Southern Brazil, and Argentina, and in greenhouses throughout 
the world. 

Chile : 

In Chile it was first reported and described as Aj)his citricola by P. 
Van der Goot in 1912. Locality and host i^lant were not given. 

Argenthsta : 

On Citrus spp., at Jujuy, Province of Jujuy, February 15, 1951, by 

Dr. A. E. Michelbaeher. It was abundant on the foliage. Three 

apterae and 3 alates were taken. 

It appears to the author that Blanchard's Paratoxoptera argentinensis, 

collected in orange groves or "naranjales en la Republica Argentina en 

Santa Ana, Missiones y en Yapeyu, Corrientes," is the tropical citrus 

aphid as designated above. (Blanchard 1944, pp. 20-22, fig. 1.) 

The writer recently received specimens from L. A. Bahamondes col- 
lected on Salix hahylonica L., Guaymallen, Province of ^lendoza, Novem- 
ber 28, 1950 ; 8 apterae. 

Peru : 

On Citrus spp. at Chicla, Province of Lima, March 21, 1951 ; 5 apterae. 

Aphis coreopsidis Thomas 

Coreopsis Aphid 
(Figure 15) 

Alate parthenogenetic female : A small species with black head, thorax, 
antennae, most of the legs, and cornicles; abdomen and cauda pale. An- 
tennae relatively short, with few short hairs ; roughly imbricated. Seg- 
ments III and IV somewhat swollen. III with 8 to 10 large circular and 
oval secondary sensoria ; IV with 4 to 7 and V with 1 to 3 similar sen- 
soria. Rostrum extending nearly to third coxae. Wing veins slightly 
dusky; two hamuli on each hind wing. Cauda pale with 5 or 6 hairs. 
Length of body 1.60 mm. ; antennae 1.50 mm. ; forewings 2.20 mm. ; cor- 
nicles 0.25 mm. ; cauda 0.15 mm. 

Apterous parthenogenetic female : Cleared specimens almost entirely 
pale with apical two thirds of antennal segment III, all of segments IV-VI, 
and the cornicles black ; body with dark areas around spiracular openings. 
Antennal segment III without secondary sensoria. Rostrum slender, ex- 
tending to second coxae ; with few hairs. Cornicles imbricated, somewhat 



V^/^J ^@T$jij_^^^^ 

n xj. ■ ,1 1 1 1 J f ,i 1 , f ) ij ; t . i w 1 1 r ) I ji I , u i 1 1 1 it roag^ 

Figure 15. Coreopsis apliid, Aphis coreo2)sidis Tliomas. Alate female showing 
antennae; cornicles, cauda, wing, and rostrum. 

curved outwardly; cauda rather thick and with 5 or 6 hairs. Length of 
body 1.60 mm.; antennae 1.60 mm.; cornicles 0.60 mm.; cauda 0.30 mm. 

Colombia : 

On a flower of the family Compositae, Buenaventura, November 4. 
1950. A single alate, one mature apterous female and two imma- 
ture specimens were taken. They are mounted on a single slide. 

Argentina -. 

Blanchard (1939. pp. 911, 912-914, fig. 13), reports this species on 
Bidens megapofamica (Speng.) at Missiones, in 1937. 
This aphid was described by Cyrus Tliomas in 1878 and has l)oen 
reported on a numl)er of Compositae in many jiarts of the United States 
from Connecticut to California. 

Aphis g-ossypii ((ilover) 1855 (1854) 

Cotton Aphid 
(Figures 16-17) 

This is a widely distributed species, especially in the warmer tem- 
perate and tropical regions. It is probably the commonest species in the 







Figure 16. Cotton aphid, Aphis gossyini Glover. A, apterous female; a, anal 
plate and cauda ; b. cornicle; c, rostrum; d, antenna; B, first instar young; e, 
antenna of second instar; f, antenna of first instar; g, cornicle, h, rostrum. 

Pacific tropical islands. It also occurs out of doors and in greenhouses 
in the temperate areas. It is well represented in South America as indi- 
cated by the reports of the entomologists of that continent. It was collected 
as follows: 

Chile : ' 

On squash, calabaza (Cucurhifo maxima), Botanical Garden, Lima, 

November 14, 1950. Alatae and apterae. 
On Bigno7iia sp. (?) at Anthofagasta, November 21, 1950. ^lany 

On Jacaranda sp. at Ovalle, Province of Coquimbo, December 1, 1950. 

A few specimens. 

Argentina : 

On myrtle-like tree, elevation 4,000 feet, Tucuman, Province of Tu- 

cuman, February 11, 1951. Many specimens — apterae and alatae. 
On thorny tree, Jujuy, Province of Jujuy, February 15. 1951. A 

few specimens of apterae and alatae. 




Figure 17. Cotton aphid, Aphis gossypii Glover. Alate female and rostrum, 
tarsi, cornicle, cauda, antennae, cauda. 

On Asclepias sp., Jiijuy, Province of Jujuy, February 16, 1951. Many 
specimens of apterae and alatae, 

E. E. Blanchard (1944, p. 23; 1939, pp. 917-918) lists this species on 
alfalfa, Clienopodium sp., Jacaranda ovalifolia R. Br., Guava (guaya), 
Liliu7n sp., (azucena), Lepidmm sp.. Hibiscus sp., Solanum sisymhriif olium 
Lam., Begonia sp., Citrullus vulgaris Schrad., Cycloma spp., Dyckia flori- 
hunda Girs., Lavatera arhorea, Mandevillea suaveolens Lindl., Morrenia 
odorata (Hook et Arn.), Oleaceae, Orthostemon sellowianus Berg., Per sea 
americana Mill., Pyrus malus L., Sechium edule Sw., Solanum lycopersi- 
cum L., Vernonia sp., Vitis vinifera L., and Zea mays L. 

The writer has recently received specimens from L. A. Bahamondes 
as follows: 

On senna (sen-sen). Cassia aphila, Mendoza Capital, October 25, 1948 

— many specimens. 
On pear (pera), Pyrus communis L., Mendoza Capital, October 15, 
1948 — many specimens. 


On Crataegus sj)., Mendoza Capital, December 12, 1950— many 

On Parthenium hysterophorus, Mendu/.a Capital, October IG, 1950— 
many apterae and alatae. 

Colombia : 

On AtripJex sp. (?), Buenaventura, November 4, 1950. Many alatae 

and apterae. 
On Baccharis sp. (?), November 4, 1950— a few alatae and apterae. 
On ComposiUe (t), November 4, 1950—1 alate and 3 apterae. 
On a shrnb, November 4, 1950— many apterae and a few alatae. 
Peru : 

On Casuarina in Botanical Garden, Lima, November 14, 1950 — many 
apterae and alatae. 

On Pomegranate, Pnnica granatum L., Callou, November 14, 1950 — 
a few aptera. 

On squash, Cucurhita maxima L., Botanical Garden, Lima, November 

14, 1950 — 9 apterae and 6 alatae. 

On cultivated potato, Solanum tuheromm L., Sicuani, Province of 
Cusco, March 2, 1951 — 1 specimen. 

On cultivated cotton, Gossypium s]x, Chanca, Province of Lima, March 

15, 1951 — 4 apterae. 

Aphis illinoisensis Shimer 

Grapevine Aphid 
(Figure 18) 

Aphis anipelophila Blanchard, 1913. 

This is a large, black North American species which commonly feeds 
on grapes and related plants. 

Chile : 

On a wild Berheris sp., 2 km. west of Porto Veras, Province of Llan- 
quihue, January 17, 1951. Only a few alates and apterous forms. 

Argentina : 

This species is reported by Blanchard (1923, pp. 33-35; 1931, p. 1002) 
on grape, Vitis sp. 
It was also described from specimens taken in Argentina by Del Guercio 
(1913, p. 159) as Aphis ampelophiJa which is a synonym of A. illinoisensis. 




Figure 18. Grapevine aphid, Aphis illinoisensis Shimer. A, antenna of alate; 
B, cornicle of alate; C, cornicle of aptera; D, rostrum of alate; E, cauda and anal 
plate of alate; F, cauda of aptera; G, prothoracic tubercle; H. forewing. 

Aphis marthae Essig, new species 

Cabildo Aphid 
(Figure 19) 

Type : Alate parthenogenetic female : A large species, black and pale 
in various patterns, one of which is figaired. Antennae black, shorter than 

Figure 19. Cabildo aphid. Ajjliis marthae Essig, new species. A, alate; B. aptera. 
a, Cauda and b, cornicle of alate; c, reticulations on dorsum of aptera; d, cornicle 
and e, antennae of alate. 


the body; unguis about twice the length of the base; segment III with a 
variable number of small and large, circular sensoria, numbering 11-16 
and distributed over much of the length. (Paratypes show the following 
pattern: 11-14, 11-15, 12-15, 12-16, 13-14, 13-16, 15-15, 15-18, 16-18, 
16-19, 17-19, 17-21, 18-18, etc.) ; IV with 1-2 sensoria. (Paratypes with 
the following numbers: 0-1, 0-2, 0-3, 1-2, 1-3, 2-3, 2-4, 5-X) ; with few 
short hairs. Rostrum quite wide with two apical segments about equal 
in length ; with hairs as shown. Primary wings large with second branch 
of media one-third distance from first fork. (This character is variable 
in paratypes. The branch may be near or even forward of the middle 
and in one instance the media had but a single branch.) Length 3.3 mm. 
Cornicles short, almost cylindrical, imbricated; 0.25 mm. long. The area 
around the base is pale. Cauda slightly shorter than the cornicle, with 
11-12 hairs. Length of body 2.5 mm. 

Apterous pa rtheno genetic female : Of the same general shape, size, 
and color of Aphis niedicaginis Koch which it also resembles in having 
the mosaic pattern on the dorsum of the abdomen. Shining black with por- 
tions of the legs and body paler. The pale areas surrounding the bases of 
the cornicles are specially noticeable. The pigmented color pattern on 
the dorsum varies considerably and may depend upon the development 
and age of the individual. Nearly full grown individuals have little dark 
markings. Length of body 2.50 mm. ; antennae 1.60 mm. ; cornicles 0.30 
mm. ; Cauda 0.30 mm. 

Host plant and locality: This species was collected on an undetermined 
shrub or tree at Cabildo, Province of Aconcagua, Chile, November 28, 
1950, by Dr. A. E. ]\Iichelbacher. A large number of apterae and alatae 
were taken. 

Type and paratypes: A singe alate individual has been designated as 
the type and the remaining 5 alatae and 20 apterae are designated as 

This species most resembles Aphis rumicis Linnaeus, but differs in the 
following respects: The bod.v is much more heavily pigmented; there are 
fewer sensoria on antenal segment III in the alatae; the unguis of the 
antennae is much shorter ; the rostrum is more robust and the apical 
segments longer, the cauda is more tapering and with few^er hairs. 

It diifers from Aphis niedicaginis Koch in having many more sensoria 
on antenal segment III and in also having a few sensoria on segment IV: 
and in having the apical segments of the rostrum broader. 

It differs from Aphis hazzi Blanchard in being more heavily pigmented ; 
with sliorter antenal unguis; in having fewer sensoria on antennal seg- 
ment IV: and in being much larger in size: 2.50 mm. as compared with 
1.80 mm. 


It was named for Mrs. Martha Michelbacher, one of its collectors, in 
recognition for her contributions to the South American apliid survey. 

Aphis medicaginis Kochj^* 1854 

Cowpea Aphid 
(Figure 20) 

This aphid is shiny black in life, especially the apterous forms with 
tlie distal portions of the legs distinctly whitish. When cleared and 
mounted they appear as illustrated. This aphid is quite common, especially 
on species of Leguminosae, and occurs throughout much of the world and 
is widely distributed in South America as indicated by the collections 
made by Dr. Michelbacher and others. 

Chile : 

On thorny legume {Cassia sp. °?), Valparaiso, November 17, 1950; 

many apterae, 
On Cassia sp. Pedegua, Province of Aconcagua, November 28, 1950. 

Many apterae and 1 alate. 
On thorny legume, Illapel, Province of Coquimbo, November 28, 1950 

— 10 apterae. 
On Artemisia sp. ?, 90 km. east of end of tunnel, Illapel, November 

28, 1950 — 5 apterae and 1 alate. 
On Composite (Artemisia sp. ?), 90 km. east of end of tunnel, Illapel, 

November 28, 1950 — 6 apterae and 2 alatae. 
On Cereus sp.. Las Palmas, Province of Aconcagua, November 29, 

1950 — apterae and alatae. 
On Cassia sp. '?, Illapel, Province of Coquimbo, November 30, 1950— 

apterae only. 
On yellow-flowered cactus (cacto). La Serena, Province of Coquimbo, 

December 2, 1950 — apterae only. 
On legume (?), Huanta, Province of Coquimbo, December 6, 1950 — 

apterae and alatae. 
On woody shrub, Los Vilos, Province of Coquimbo, December 14, 1950 

— alates and apterae. 

3. D. Hille Ris Lambers informs me that he does not believe that the European Aphis mcdicaginis Koch 
occurs in America and that what we have been calling that species is probably Aphis craccivora Koch. 
In my collection there is a good series of this European species collected on Vicia cracca L. in Belgium at 
Vise by A. Collart, 1938, and at Weerde by E. Janmoulle in 1939. These appear much like our American 
Aphis medicaginis Koch in general appearances, but differ in having much shorter cornicles; greater number 
of caudal hairs: and a somewhat shorter filament, unguis or processus terminalis. It is quite possible that 
our species is distinct. 


Figure 20. Cowpea aphid, Aphis uierlicaginis Koch. Adults and antenna, Cauda 
and cornicle of aptera. (After Zimmerman, 1948.) 

On lejiumiiious tree, Valparaiso, December 17, 1950 — apterae and 

On Baccharis sp. ?, Chilian, Province of Niible, December 27, 1950 — 

many apterae and 1 alate. Very small specimens. 


On thorny legume, Antueo, Province of Bio-Bio, December 28, 1950 

— apterae only. 
On thorny legume, El Abanico, Province of Bio-Bio, December 30, 

1950 — alatae and apterae. 

On Sohmum sp., Los Andes, Province of Aconcagua, February 4, 1951, 

many apterae and alatae. 
On wild nasturtium (nasturcia), Portillo (3,000 feet). Province of 

Aconcagua, February 4, 1951. Many apterae and alatae. 

Argentina : 

On Asi r((gaJiLs sp. ?, La Quiaca, Province of Jujuy, February 18, 1951 
— many specimens apterae and alatae. 

In addition, Blanchard (1939, pp. 914-916; pp. 922-923) lists the 
following host plants (as Aphis fabae Scopoli and A. lahurni Kaltenbach) : 

Acacia sp., Arachis hypogaea L., Cicuta sp., Dyckia florihunda Oris., 
Eryngium spp., Euonymus sp., Eucalyptus spp., Foeniculum vul- 
gare, GJadiohis sp., Cirsium sp., Chrysanthemum sp.. Cosmos sp.. 
Dahlia sp., Phaseolus hmatus L., Fhoenix sp., Gerhera jamesoni, 
Senecio honariensis H. & A., Seshania punicea (DC), Spiraea 
chamaedrifoUa L., Yicia sp. 

The writer has recently received specimens from L. A. Bahamondes 
collected by him as follows : 

On Solanum sp., Mendoza, October 3, 1948 — many apterae. 

On a Composite, Mendoza, February 10, 1948 — many apterae. 

On Glo.rinia sp., IMendoza Capital, ]\Iarch 10, 1948 — a few apterae. 

Colombia : 

On legume, Buenaventura, Noveml^er 4, 1950. Collection consisted of 
21 apterous females. 


On l)road bean, Vicia faha, Callao, November 4, 1950 — 5 alates and 10 

On Astragalus sp., Sicuana, Province of Cusco, March 1, 1951 — many 
specimens, all apterae. 

On Cassia sp., Cusco, Province of Cusco, IMarch 2, 1951 — many speci- 
mens — all apterae. 

On thorny legume, Cusco, Province of Cusco, IMarch 2, 1951 — all 


Aphis nerii Boyer de Fonscolonibe, 1841 

Oleander Aphid 
(Figure 21) 


Figure 21. Oleander aphid. Ajihis iierii (Boyer de Fonscolonibe). Alate and 
apterous parthenogenetic females witli enlarged antenna of alate and cornicles and 
caudas of alate and apterous forms. A bright yellow and black species. 

Cerosipha nerii (Boyer de Fonscolonibe) Boknek. 1952. 

This is a widely distributed aphid occurring on Aschpias spp., Caltro- 
phis spp., Gonolohus spp., and related genera in Europe, Asia, South Pa- 
cific Islands, North and South America, and Africa. 

Chile : 

On oleander, Nerium sp. (adelfa, baladre), Antofagasta, November 22, 
1950 ; many apterae and alatae. 

On an unknown host, south slope of Bell Mountain, Province of Acon- 
cagua, December 17, 1950 — 1 alate. 

Beating (plants not ascertained) near the mouth of MauUin River, 
Province of Llanquihue, January 22, 1951 — a large number of 
apterae and alatae. 



Bolivia : 

On Asdejnas sp., Cumergo, February 20, 1951 — many a])torao and 

Argentina : 

On Nerium oleander L., Araujia scricofera Bert., Asclepias curas- 
savica L., and Clematis sp. (Ornamental) by Blanchard (1923, 
pp. 39-41; 1939, pp. 925-26). 


On a vine-like Asclepias (?), 65 miles west of Cusco, March 5, 1951 — 

3 alatae. 
On Asclepias sp., Chiclayo, Province of Lambayeque, March 19, 1951 

— all apterae. 

Aphis rumicis Linnaeus, 1758 

Bean or Dock Aphid 
(Figures 22-25) 

Figure 22. Bean or dock aphid, Aphin rumicis Linnaeus. Apterous partheno- 
genetic female and enlargements of a. anal plate and cauda; b, tarsus; c, lateral 
abdominal tubercle; d, mosaic and glandular areas; e, cornicle; f, rostrum; g, 

Vol. XXVIII 1 



Almost a cosmopolitan species which is widely distributed throughout 
most of the world. It feeds upon many different hosts. The following col- 
lections were made : 

Chile : 

On Chrysanthemum sp., Antofagasta, November 21, 1950 — apterae 

On Epiphyllum cactus, in a garden at La Serena, Province of Co- 
quimbo, December 2, 1950 — apterae and alatae. 

On a native shrub, 10 miles below Laguna Dam, 6,000-7,000 feet, De- 
cember 6, 1950. Plant heavily infested — only apterae collected. 

On Rumex sp.. Hacienda San Andres, near Purranque, Province of 
Llanquihue, January 1, 1951 — all apterae. 

On thistle {Cirsium sp. ?), Angol, Province of Malleco, January 1, 
1951 ; all apterae. 

Beating at Hacienda San Andres, near Purranque, Province of Llan- 
quihue, January 15, 1951 — a few apterae. 

Figure 23. Bean or dock aphid, Aphis rumicis Linnaeus. Alate parthenogenetic 
female and enlarged cauda, tarsus, cornicle, rostrum (venter), and antenna. 



On wild currant ? (grosellers), Hacienda San Andres, Parranque, 
Province of Llanciuihue, January 15, 1951 — apterae and alatae. 

Argentina : 

On wild potato (papa), Salta, Province of Salta, February 15, 1951 — 

1 single alate. 
On SoJanum sp., Salta, February 15, 1951 — many specimens ; many 

apterae and 1 alate. 
On thorny legume, Jujuy, Province of Ju.juy, February 15, 1951 — 

many specimens, all apterae. 


Figure 24. Bean or dock aphid, Aj)his rumicis Linnaeus. Sexual female and 
enlarged anal plate and cauda, antenna, cornicle, rostrum (venter) and hind tibia 
showing sensoria. 

On Solanuni sp., Jujuy, February 16, 1951 — many apterae and alatae. 

Blanchard (1935:373-375; 1939:929-930) records it 
On Rumex sp., at Tucuman. 
On Rumex sp., Cestrum parqui L.'Herit, Solanum Jycopersinun L., 

8. nodifioruyn Jacq., -S. capsicastrum Link., and other solanums. 

The writer has recently received specimens from L. A. Bahamondes, 
collected by him as follows : 

On Chaenomeles japonica (Thunb.) Lindl., at Mendoza, October 20, 
1950 — 3 apterae and 2 alatae. 

On TuJipa sp., Las Heras, Mendoza, October 10, 1950 — apterae and 



On Fumex sp., Mendoza, Guaymallen y Capital, November 1, 1950 — 
apterae only. 

On Eugenia sp., Mendoza Capital, May 10, 1918 — apterae. 

On Solanum sp. (beating), Ando Huylas, Province of ^Mendoza, Feb- 
ruary 15, 1951 — -1 alate. 

Figure 25. Bean or dock aphid, Aphis rumicis Linnaeus. Alate and enlarged 
tarus, cornicle, anal plate and cauda, rostrum, and antenna of same. 


On Agave sp., Rio Pampas, Province of Purimac, March 8, 1951 — 
many apterae and alatae. About 60 per cent killed by a fungus 

On Composite (?), on road 65 miles west of Cusco, ]\larcli 5, 1951. 
Only apterae. 

On Composite, Cusco, March 5, 1951 — 1 alate. 

Toxoptera aurantii (Boyer de Fonscolombe) 

Black Citrus Aphid 
(Figure 26) 

Aphis aurantii Boyer de Fonscolombe, 1841; Ajjhis camelUae Kaltenbach, 1843; 
Toxoptera aurantiae Koch, 1856. 



Figure 26. Black citrus aphid, Toxoptera aurantii (Boyer de Fonscolombe). At 
left: alate and apterous parthenogenetic females with enlarged antennae, Cauda, 
and cornicles. (After Zimmerman, 1948); at right: net-like saw-toothed reticula- 
tions on the venter beneath the bases of the cornicles, which are found so far 
only in this species and in Aphis citricidus (Kirkaldy). 

This species is widely distributed throughout the tropical and sub- 
tropical regions of the world and feeds upon such plants as those belonging 
to the genera Camellia, Cinchona, Citrus, Coccoloha, Coffea, Ficus, Gar- 
denia, Hibiscus, Ilex, Mangifera, Persea, Straussia, Theohroma, and others. 
Strangely enough only a single collection was made on this expedition in 
South America and that in Colombia. 

Argentina : 

This species is quite common throughout the Provinces of Buenos Aires, 
Entre Rios, Corrientes, Santa Fe, and Tucuman, according to Blanchard 
(1925, pp. 20-22; 1939, pp. 903-904; 1944, p. 18). 

On Citrus spp., Ilex paraguariensis Bonop., Mandevilla suaveolens 
Lindl., Phytolacca dioica L., Scutia huxifolia Reiss., and Vihur- 
num tinus h. 

Colombia : 

On mango {Persea sp.), Buenaventura, November 5, 1950 — many 

Vol. XXVIII 1 




Tribe Anuraphidini 

Subtribe Anuraphidina 

Brachycaudus helichrysi (Kaltenbach) 

Leaf-curl Plum Aphid 
(Figure 27) 

Figure 27. Leaf-curl plum aphid, Brachycaudus helichrysi (Kaltenbach). Alate 
and apterous parthenogenetic females with enlarged antennae, cornicles, and cauda. 
(After Zimmerman, 1948.) 

Aphis helichrysi Kaltenbach, 1843; A. bartsiae Walker, 1848; A. bellis Bickton, 
1879; A. verbenae Macchiati, 1883; A. leotiopedii Schoiteuen, 1903; Anura- 
phis cyani THEOBALn, 1915; A. abrotaniella Theobald, 1919; A. cantauriella 
Theobald. 1921; A. sherardiae. Theobald, 1926. 


This small, pale aphid has conspicuous dark body markings. It is 
widely distributed throughout tlie temperate regions of the world and 
attacks many kinds of plants. 


Very abundant on peach {Primus persica) causing the leaves to curl 
at Rivadava, Province of Huanta, December 4, 1950. Very many apterous 
and but one alate were collected at the time. The alatae were just beginning 
to appear. 

On the wing at Los Muermos, Province of Llanquihue, January 20, 
1951 — -1 alate and man}^ apterae. 

Argentina : 

Blanchard (1922, pp. 5-4-56) reports this species often abundant on 
peach and says it is "the chief enemy of the peach and causes much 
damage in the spring to the young foliage, which soon becomes curled and 
distorted. ' ' 

On Senecio sp., Aster sp., Prunus sp., and Compositae in the Province 
of Buenos Aires. (Blanchard 1922, p. 52, fig. 17; 1939, p. 894). 

On Cineraria sp., Pampa (Blanchard 1944, p. 57). 

Specimens have also been received from L. A. Bahamondes : 

On (ortiga) nettle, Godoy Cruz, Mendoza Capital, November 20, 1950 
2 alatae. 

Peru : 

On Shasta daisy, Chrysanthemum maximum (?), Lima, November 15, 
1950 — apterae and alatae. 

On potato, Solanum tuberosum, 3,000 meters altitude, Sicuana, Prov- 
ince of Cusco, March 2, 1951. Alatae only. 

On marigold. Calendula officinalis, Callao, November 14, 1950 — 6 ap- 
terae and 6 alatae. 

On Chrysanthemum sp.. Botanical Garden, Lima, November 13, 1950 
— 3 apterae. 

Tribe Brachycolini 

Brevicoryne brassicae (Linnaeus) 

Cabbage Aphid 
(Figures 28-29) 

Aphis brassicae Linnaeus, 1758; Brevicoryne brassicae (Linnaeus) Van der Goot. 



Figure 28. Cabbage aphid, Brevicoryne brassicae (Linnaeus). Alate and ap- 
terous parthenogenetic females and enlarged genital plates, caudas, cornicles, ros- 
trum, and antennae. 

This very common aphid appears to be a Holarctic species, although it 
may well have been early introduced into the Americas from Europe. It 
has become widely distributed throughout much of the world by commerce. 



Figure 29. Cabbage aphid, Brevicoryne brassicae (Linnaeus). Alate male and 
apterous female with enlarged antennae, rostrum, cornicles, caudas, male genitalia, 
and female hind femur with sexual sensoria. 

Chile : 

On cabbage, Brassica oleracea var. capitata, at Purranque, Province 
of Osorno, January 17, 1951 — many apterae and alatae. 

Argentina : 

"Common everywhere in Argentina on Brassica spp., Baphanus spp., 
and Spinacia oleracea.' ' (Blanchard, 1925, pp. 12-14.) 

Peru : 

On mustard (Brassica sp.), Callao, November 14, 1950 — many apterae 

and alatae. 
On Caper-bush, Capparis spmosa. Botanical Garden, Lima, November 
14, 1950. 


Hyadaphis conii (Davidson) 

Honeysuckle Aphid 

(Figure 30) 


Figure 30. Honeysuckle aphid, Hyadavhis conii (Davidson). Enlarged drawings 
of antennae, cornicles, rostrum, caudas, of aptera and alate and wing of alate. 

Siphocoryne conii Davidson, 1909; Aphis xylostei Schrank, 1801; Hyadaphis co- 
niella Theobald, 1925; H. sii Bokner, 1931-32; Rhopalosiphum jnelliferum 
HOTTES, 1930. 

This species is widely distributed in temperate regions and is common 
in Europe and North America and also occurs in Africa and South America. 
It particulai-ly infests Lonicera spp. and members of the large family 


A single collection of many apterae and alatae was made on Carum 
sp. (?), 60 kilometers east of Los Andes, Province of Aconcagua, 
February 4, 1951. 

Tribe Liosomaphidini 

Cavariella aegopodii Scopoli 

Parsnip and Willow Aphid 
(Figure 31) 

Aphis aegopodii Scopoli, 1763; Cavariella capreae (Fabricius), 1775 (Many 

This is probably a Holarctic species commonly occurring in P^urope 
and North America. It has been introduced into many parts of the world. 




Figure 31. Parsnip and willow aphid, CavarieUa aegopodii (Scopoli). Alate 
and apterous parthenogenetic females and enlarged antennae, cornicles, caudas, 
and caudal tubercle of female (lateral aspect). (After Zimmerman, 1948.) 


In Chile it appears to be widely distributed on ninbelliferous plants 
and was collected as follows: 

On an umbelliferous plant, at La Ligura, Province of Aconcagua, 
December 14, 1950 — apterae and alatae present in large numbers. 

On anise or dill, Ancthum graveolens, Angol, Province of Bio-Bio, 
January 1, 1951 — many apterae and alatae. 

On a wild host and by beating various plants at Hacienda San An- 
dres, Purranque, Province of Osorno, January 15, 1951 — apterae 
and alatae. 


Argentina : 

Under the above specific name and also as C. caprcae (Fabricius) this 
species has been recorded : 

On Ainuni sp., Daucus sp., Carum sp., Pastinaca sp., Pimpinella ani- 
sutn h., and Salix sp. (Blanchard 1925, pp. 14-16; 1944, pp. 

Specimens have been received from L. A. Bahamondes, collected on 
Salix hahylonica L., Guaymallen, Province of Mendoza, Novem- 
ber 28, 1951 ; many apterae and alatae. 

Tribe Myzini 

Subtribe Myzina 

Myzus circumflexus (Buckton) 

Lily Aphid 
(Figure 32) 

Siphonophora circumflexa Buckton. 1876; 8. callae Heixrich, 1909: Myzus vincae 
Gillette. 1908. 

This is a European species now widely distributed on lilaeeous plants. 
The insect infests both the bulbs and foliage. The pale green or yellowish 
apterae have conspicuous dark markings on the dorsum of the abdomen. 

Chile : 

A single apterous female was taken by beating at Purranque, Province 
of Osorno, January 15, 1951. 

Argentina : 

This species has also been collected upon a number of hosts in the 
Province of Buenos Aires, by Blanchard (1922, pp. 213-4, fig. 15; 1939, 
p. 976; 1944, p. 43) as follows: 

On Citrus sp.. Cyclamen sp.. TuJipa sp., and other Lilaceae, Orchi- 
daceae, and Vinca major L. 

Myzus omatus Laing, 1932 

Ornate Aphid 
(Figure 33) 

This species appears to have been introduced into England, where it 
was first noted on violets by Laing (1932) and described. Since then the 
writer has received specimens collected upon many host plants (Essig, 
1938, 1939, 1947). It appears that this insect may be a native of South 



^''"'- c2Si2#=^ 

A ant Cu^]ij];^xrEm 

Figure 32. Lily aphid, Myzus circumflexus (Buckton). A, adult winged female; 
a, section of the costal vein showing fenestralike areas; d, cornicle; f, cauda; k 
and 1, fenestras near the base of the subcostal vein; W, ant., antennae. B, adult 
apterous female: A, ant., antenna; b and c, setas on segments I and III of an- 
tenna; e, cornicle; g, cauda; h, rostrum; i, basal margin of cornicle; j, tip of 
cornicle. All greatly enlarged. A yellow and black species. 




America where it is associated with plants of the genera Fuchsia and Be- 
gonia and perhaps other indigenous plants. 

Chile : 

On Cereus sp., at Las Palmas, Province of Aconcagua, November 29, 

1950, along with some other aphids. 
On Bignonia sp., Antofagasta, November 21, 1950 — 3 apterae. 

Figure 33. Ornate aphid, Myzus ornatus Laing. Alate and apterous partlieno- 
genetic females with enlarged antennae, caudas, cornicles, and rostrum. 

Myzus persicae (Sulzer) 

Green Peach Aphid 
(Figure 34) 

Aphis persicae Sulzer, 1776; Aphis dianthi Schrank, 1801; A. persicwcola Bois- 
DUVAL, 1867; Myzus malvae Oestlund, 1886; Phorodon cynoglossum Williams, 
1910 ; Rhopalosiphum betae Theobald, 1913. 



Figure 34. Green peach aphid, Myzus persicae (Sulzer). A, alate partheno- 
genetic female; B, apterous female. Enlarged rostrum; cornicles, caudas, and an- 
tennae. A yellow, green and black species. 

This is one of the most widely distributed aphids and occurs on all the 
continents. It has an enormous number of host plants and is the most 
important aphid vector of plant virus diseases (Essig 1948b). 


On Datura sp., Antofagasta, November 21, 1950 — 10 apterae. 

On Cereus sp., Las Palmas, Province of Aconcagua, November 29, 

1950 — a few specimens, all apterae. 
On wild morning glory. Convolvulus sp., Angol, Province of Bio-Bio, 

January 1, 1951 — 1 alate and 3 apterae. 

Argentina : 

This aphid is widely distributed throughout this country and has been 
reported upon many host plants including: 

Ahutilon sp., Antirrhinum sp., Bellis sp., Beta sp., Brassica sp., Can- 
nabis sativa L., Capsicum sp., Chenopodiutn sp.. Cineraria sp., 
Cosmos sp., Cuminum cyminum L., Cynara sp., Descurainia ap- 
pendiculata (Gris.), Dianthus sp., Loriicera sp., Malva sp., Pim- 
pinella anisum L., Prunus spp., Pyrus sp., Senecio sp., Solanum 


spp., TuUpa sp., Vinca sp. (Blanchard 1922, pp. 211-213, %. 13; 
1939, pp. 980-981; 1944, p. 43). 

The writer has also received specimens from L. A. Bahamondes, col- 
lected by him as follows : 

On alheli, stock or gilliflower, Matiola incana L., Mendoza Capital, 
October 3, 1948—4 alatae. 

On Begonia, Mendoza Capital, ]\Iarch 7, 1949 — many specimens. 

On nettle, ortiga, rrtica urens, Godoy Cruz, Mendoza Capital, Novem- 
ber 20, 1950. 

On Tulipa sp., at Mendoza, October 13, 1950—1 aptera and 1 alate. 

On Bumex sp., Mendoza, Guaymallen y Capital, November 1, 1950^ 
sev'eral specimens 

On Barnadesia odorato (?) Mendoza Capital, November 1, 1950 — many 

Bolivia : 

On cultivated potato (patata, papa), at Camargo, February 22, 1951 
— 2 apterae. 


On caper-bush, Capporis spinosa, Botanical Garden, Lima, November 
14, 1950 — 2 apterae and 1 alate. ' 

On Phlox sp., Lima, November 15, 1950 — many apterae. 

On tobacco, Nicotiana sp.. Botanical Garden, Lima, November 14, 1950 
— 1 alate. 

On Solanum sp., Callao, November 14, 1950 — 2 alates. 

On potato, Solanum tuberosum, Callao, November 16, 1950 — 1 alate 
and 3 apterae. 

On Datura sp., Botanical Garden, Lima, November 14, 1950 — 2 alatae 
and 4 apterae. 

B}^ beating nightshade, Solarium sp., Andahuaylas, Province of Apuri- 
mac, March 7, 1951^ — 6 apterae and 1 alate. 

On cultivated potatoes, Sicuani, Province of Cusco, March 2, 1951 — ■ 
4 alates. 

On thorny tree by beating, Abaneay, Province of Apurimac, i\Iarch 6, 
1951 — 1 aptera. 

By beating, Rio Pampas, Province of Ayaeucho, March 8, 1951 — 3 


Tribe Cryptomyzini 

Capitophorus braggi (Clillette) 

Artichoke Aphid 
(Figure 35) 

Myzus braggii Gillettk, 1908. 

This pale yellow and greenish aphid often appears in overwhelming 
numbers on artichoke and other hosts. 

Chile : 

On artichoke (alcaucil) Cynara scoJymus, San Carlos, Province of 
Nuble, December 23, 1950. Many apterae and a few alatae. 

No host record ; 50 km. east of San Carlos, Province of Nuble, Decem- 
ber 25, 1950—4 alates. Collected by Dr. E. S. Ross. 

Argentina : 

On artichoke (alcaucil), Cynara scoiymus, in the provinces of Buenos 
Aires, Santa Fe, San Juan and other localities. (Blanchard, 1935, 
366-7; 1939, pp. 945-7; 1944, p. 34). 

' Tribe Nasonoviini 

Idiopterus nephrelepidis Davis 

Fern Aphid 
(Figure 36) 

A small black and white-marked aphid with clouded wings. It feeds 
upon various species of ferns {helecho, polipoclio) (Polypodiaceae) in 
tropical and sul)tropical areas. 


A single alate specimen was taken by sweeping 40 km. east of San 
Carlos, Nuble Province, December 24, 1950, by Dr. A. E. :\Iichel- 

Argentina : 

Blanchard (1939, pp. 41-42) records this species on (Acrostichuni), 
Adiantiwi, NephroJepis, and other cultivated ferns in the Province 
of Buenos Aires. 



Figure 35. Artichoke aphid. Cai)itoi>hoius liraggi (Gillette). Alate and apterous 
parthenogenetic females with enlarged antennae, caudas, and cornicles and capi- 
tate hairs or setae on the front of the head of the aptera. Alates green and black; 
aptera pale green. (After Zimmerman, 1948.) 




Figure 36. Fern aphid, IdiopteruH nephrelepides Davis. Alate and apterous 
females; antennae and cornicles of each; cauda of alate; and body tubercle of ap- 
terous female. (After Zimmerman, 1948.) 


Tribe Aulacorthini 

Subtribe Microlophiina 

Acyrthosiphon onobrychis (Boyer de Fonscolombe)' 

Pea Aphid 
(Figure 37) 

4. The nomenclature of this species has been the object of some concern of aphidologists. Although 
the above designation is now generally accepted, the eminent aphidologist D. Hille Ris Lambers (1947, p. 
247), uses Acyrthosiphon pisum (Harris). Moses Harris (1776, pp. 66-67, pi. XIII, fig. 5), gives a brief 
description and a colored illustration of what is without doubt this insect, but there may be some question 
as to whether these constitute a valid description of a new species. This name was listed in Sherborn's 
"Index Animalium" (1902, p. 756), but I know of no other references to it — not even in the extensive 
bibliography listed by Lambers. 



Figure 37. Pea aphid. AcyrthosiiJJwn onohrachis (Fouscolombe ) [Macrosiphu7n 
pisi (Kaltenbach) ]. Alate partlienogenetic female and enlarged antenna, rostrum, 
Cauda and cornicle at top; apterous female and cauda and cornicle at bottom. 
Apex of cornicle is not reticulated. Green. 


Aphis pisunt M. Harris, 1776; A. onobrychis Boyer de Fonscolombe, 1841; A. lathyri 
MosLEY, 1841; A. pisi Kai.tenbacii, 1843; Siphonophora ononis Kocir, 1855; 
Nectarophora destructor Johnson. 1^00\Macrosiphum trifolii Perganue, 1904; 
Macrosiphum pisi Patch. 1911. 

This large green aphid has a cosmopolitan distribution and is often a 
very serious pest of plants belonging to the Leguminosae. However, it does 
not yet appear to be widely distributed in western South America. Only 
one specimen was taken. 


Sweeping legumes at Mollendo, Province of Arequipa, November 19, 
] 950 — a single apterous female. 

Su])tiihe Metopolophiina 
Aulacorthum pelargonii (Kaltenbach) 
Geranium Apliid 
Aphis pelargonii Kaltenbach, 1943; A. malvae Walker, 1848-9. 

This species was collected only twice as follows : 


On geranium, at Lima, November 15, 1950 — -4 apterae. 
On Solanum sp., Callao, November 14, 1950 — 1 alate. 

Aulacorthum pseudosolani (Theobald) 

Solanum or Foxglove Aphid 
(Figure 38) 

Ajihis solani Kaltenbach. 1843; Macrosijjhum solani Theobald, 1913, Myzus ve- 
ronica Theobald, 1913; M. hydroctylei Theobald, 1925; M. pseudosolani Theo- 
bald, 1926; M. veronicellus Theobald, 1926. 

This aphid is widely distributed and lias many host plants, but there 
were only two collections : 

Chile : 

On a wild uinl)elliferous plant and by beating various plants at Pur- 
ranque. Province of Osorno, January 15, 1951 — 1 alate and 7 

On wild potato at the same phice on Janiuuy 26, 1951 — 1 alate and 
8 apterae. 



Figure 38. Solanum or foxglove aphid, AuJacorthuhi pseudosolani (Theobald) 
:= (Myzus solani (Kaltenbach) ). Apterous and winged parthenogenetic females 
with enlarged antennal segment III, cornicles and cauda. A, refers to apterous and 
W, for winged. 

Tiihe Macrosiphonini 

Sul)tribe Macrosiphonina 

Macrosiphum'^ ambrosiae (Thoma.s) 

Ambrosia A]ihi(l 
(Figure 39) 
Siplionophora amhrosiae Thom.vs. 1877. 

S. The spelling of the sjeneric name Macrosip/ium Passerini, 1860, was corrected to Macros! p/ron to 
conform to the Latin word sip/wn by Del Guercio in 1913. (Redia IX, p. 187, 1913.) However, this change 
does not conform to the present rules and regulations of the International Committee on Zoological Nomen- 




Figure 39. Ambrosia aphid, Macrosiphum ambrosiae (Tliomas). Alate and ap- 
terous parthenogenetic females with enlarged antennae, caudas, cornicles and ros- 
trum. Dark red and black. 


Tliis species is common and abundant in North America. It seems also 
to be widely distributed in South America. It is possible to confuse it with 
closely related species all of which form a complex that needs much in- 
vestigational work. 

Chile : 

On Artemisia sp. "!, Papudo, Province of Aconcagua, November 28, 
1950 — 5 apterae. 

On Baccharis sp. ?, Rivadavia, Province of Coquimbo, December 4, 
1950 — many apterae. 

On Baccharis sp. ?, Junta, Province of Coquimbo, December 6, 1950 
^many apterae. 

On Baccharis sp. ?, Concon, Province of Valparaiso, December 16, 1950 
- — 5 apterae. 

On a grindelia-like plant, 40 km. east of San Carlos, Province of Nuble, 
December 23, 1950 — 11 apterae. 

On Baccharis sp. ?, Tucapel, Province of Bio-Bio, December 28, 1950 
— 22 apterae. 

On Baccharis sp. ?, Hacienda San Andres, Purranque, Province of 
Llanquihue, January 15, 1951 — many apterae and alatae. 

On Baccharis sp. ?, Los Muermos, Province of Llanquihue, January 
18, 1951 — many alates and apterae. 

Beating along the mouth of the ]\Iaullin River, Province of Llanqui- 
hue, January' 22, 1951 — 2 alates. 

Argentina : 

On Artemisia sp. ?, Salta, Province of Salta, February 14, 1951— 

many apterae. 
On acacia-like tree (finely divided compound leaves), Salta, February 

14, 1951, many s]iecimens, all apterae. 
On Artemisia sp. ?, Salta, Fel)ruary 14, 1951 — 6 apterae. 

Peru : 

On flirijsanthemum sp., Botanical Garden, Lima, November 13, 1950 

— many apterae. 
On E7icelia sp., Botanical Garden, Lima, November 13, 1950 — many 

alatae and apterae. 
On tobacco, Nicotiana sp.. Botanical Garden, Lima, November 14, 

1950—2 alatae. 


On Gerhera janiesonii, Botanical Garden, Lima, November 14, 1950 — 

2 alatae and 12 apterae. 
On Picris sp. ?, Botanical Garden, Lima, November 13, 1950 — many 


Macrosiphum bonariensis Blanehard 
Composite Aphid 

This fine species was collected on Tanacetum vulgare L. at Flores and 
Lomas de Zamora, Argentina, by Blanehard (1912, pp. 205-6, fig. 10). It 
was raised to species statns by the same anthor (1932, pp. 19-21, fig. 10) 
and more com]iletely described by him from additional specimens collected 
on cultivated Pyrethrum, Buenos Aires; and again on Tanacetum vulgare 
L., from the Province of Buenos Aires y Cordoba, Argentina (1939, pp. 


A fine series of this species was collected by the iMichelbachers on a 
member of the Family Compositce near San Carlos, Province of Nuble, 
December 23, 1950. It appears to fall in the genus Pharalis Leach accord- 
ing to the classification by Lambers (1929). 

Macrosiphum chilensis Essig, new species 

Chilean Baccharis Aphid 
(Figure 40) 

Type: Alate parthenogenetic female: Color not indicated — possibly 
yellowish or greenish. Cleared specimens show the head, rostrum, and an- 
tennae; apical portions of the femora and tibiae; the cornicles, excepting 
the base ; and portions of the cauda and caudal plates, dusky or blackish. 
The frontal tubercles are small. Antennae slightly longer than the body; 
segment I somewhat gil)bous and much larger than II; III a little more 
than half as long as VII, with few short spine-like hairs and with 14—17 
variable circular secondary sensoria distributed irregularly or almost in a 
straight line over the basal three-fourths of the segment ; VII with short base 
and long, slender unguis. (In paratypes the number of sensoria on an- 
tenal segment III may vary from 13 to 20.) Rostrum extends to the sec- 
ond coxae. Wings as drawn. Legs rather slender. Abdomen with many 
very fine and a few short, stout spines, the bases of the latter surrounded 
by prominent dark patches somewhat like in Macrosiphum amhrosiae 
(Thomas). Cornicles cylindrical, slightly tapering; bases about twice as 
wide as the opening, slightly swollen anteriorly ; the apical reticulated area 
may be slightly, but not usually, constricted ; reticulations pronounced 



Figure 40. Chilean baccharis aphid, Mucrosii)hitni chilensis Essig, new species. 
A, alate and B, apterous parthenogenetic genetic females; a, antennae of alate; b, 
antenna of aptera; cauda, cornicle, rostrum and wing of alate. Very dark. 

and similar to those of M. solanifolii (Ashmead), Init the constriction is 
not as noticeable ; dusky except the basal one fourth. The anal plate is 
verj^ small. Cauda 0.8 mm. in length and tapering as shown. Length of 
body 3.00 mm. ; antennae 0.33 mm. ; f orewing 4 mm. ; cornicles 0.8 mm. ; 
Cauda 0.5 mm. 

Ai)terous parthenogenetic female: Of a uniform color and with dark- 
ened antennae, legs, rostrum, cornicles, cauda, and caudal plate and small 
areas at bases of thickened spines much as in the alate form. Antennal 
segment III has from 8-15 circular sensoria of various sizes and limited 
to the basal third or half of the segment — the average number is about 
11 to 12. It is rarely that the same number occurs on both antennae of 
a given individual. Length of body 3.5 min. ; width 1.6 mm. ; antennae 
3.5 mm.; cornicles 0.8 mm.; cauda 0.6 mm. 

This species somewhat resembles M. soJanifoJii (Ashmead) but differs 
in having the secondary sensoria more scattered on III in the alatae and 
many more on III in the apterae; and in having much darker or black 
antennae and cornicles. 

Host plant and locality : This species was collected on Baccharis sp. ( ?), 
at La Serena, Province of Coquimbo, Chile, December 8, 1950, by Dr. 
A. E. JMichelbaeher. 



In all, 4 alates, 25 apterae, and 5 immature individuals as well as one 
cast skin were taken. These are mounted on 5 aluminum microscope 
slides. The alate type is mounted on a slide with 4 mature apterous 

Macrosiphum cordobensis Blanchard 

Cordobens Aphid 
(Figure 41) 

Figure 41. Cordobens aphid, Macrosiphum cordobensis Elancliard. Upper — A, 
antenna, cornicle and cauda of alate parthenogenetic female ; B, antenna, cornicle 
Cauda, and rostrum of aptera. 

Argentina . 

On Baccharis sp. ( ?), at 4,000 feet on a ridge west of Tucuman, Feb- 
ruary 11, 1951 — many apterae and alatae. 

On Erigeron sp., Sierras de Cordobe, February 18, 1931. Original de- 
scription. (Blanchard, 1932, pp. 24-36, fig. 4; 1939, pp. 963-4.) 

Peru : 

On potato (papa) (casual?), Sicuani, Province of Cusco, 3,000 feet^ 
1 apterous. 

Macrosiphum edrossi Essig, new species 

Ross Aphid 
(Figure 42) 

Type: Alate parthenogenetic female: A yellowish or greenish species 
with the head, thorax, and all appendages brown or black. The cleared 
abdomen is void of any pigmented markings but has many fine hairs. The 
antennae are 4 mm. in length and longer than the ])ody, slender, with short 
knobbed hairs, and segment III wdth 15-17 large and small circular secon- 
dary sensoria arranged in a row along the underside throughout tlie full 




Figure 42. Ross aphid, Macrosiphum. edrossi Essig, new species. A, antenna, 
Cauda, rostrum and three drawings of cornicle and outline of the apterous par- 
thenogenetic female; B, antenna, cauda, cornicle and wing of alate. 

length. (In paratypes the number frequently varies for each individual 
as 15-17, 16-18, 15-16, etc.) The unguis of segment III is very long. The 
rostrum is slender, with few hairs and extends to or slightly beyond the 
second coxae. The legs are long and slender. The forewings are normal 
with venations as drawn : 4 mm. in length. The stigma is relatively lon;i'. 
The cornicles are black apically, long and slender, slightly curved out- 
wardly, somewhat constricted near tips and reticulated for a very short 
distance at the apex where it is considerably widened. In this respect 
it dilfers from most members of the genus. The apical flaring is some- 
what after the pattern of the pea aphid, Acyrthosiphon onohrachis (Boyer 
de Fonscolombe) (Macrosiphum), but in the pea aphid the reticulations 
are absent ; length 0.7 mm. The cauda is rather robust, iiaring at base 
and then slightly constricted ; 0.35 mm. in length, with six hairs. The anal 
plate is very small and crescent-shaped. Length of body, 3 mm. 

Apterous parthenogenetic female: Pale with dusky appendages and 
without any distinctive markings and with numerous small colorless hairs. 
The general characteristics are similar to those of the alate form. Anten- 
nal segment III has a variable number of basal secondary circular sen- 
soria. This variation may best be expressed by the following examples 
per individual, 2-3, 3-3, 3-5, 4-2, 4-4, 5-5, 6-9, 8-5, 8-6, 8-8, etc. The 
cauda is more rolnist than that of the alate. Length of body, 2.70 mm. ; 
antennae 2.70 mm.; cornicles 0.85 mm.; cauda 0.35 mm. 

Host plant and locality. Collected on W'hat appears to be a nettle 
(ortiga), Urtica sp., at Rio Pampas, Peru, March 8, 1951, by Dr. Michel- 

Type : An alate form mounted on a slide with 4 apterous females. The 



remaining 6 alates and all the 2-1 apterous forms are mounted on 6 other 
slides and are designated as paratypes. 

This species diifers from others in having the apices of the cornicles 
reticulated and flaring; an extremely long antennal filament; and only 
2 or 3 pairs of hairs on the cauda. 

Macrosiphum griersoni Blanchard 
Grierson Aphid 

Argentina : 

This species was described by Blanchard (1932, pp. 27-29, fig. 5) from 
the artichoke (alcachofa, arcacil, aguaturma), Cynara scoJymus L., Jujuy 
and Buenos Aires, in 1924, and on Vernonia sp.. Sierra de Cordoba. 


A single alate specimen of what appears to be this species was col- 
lected by the Michelbachers on a composite, San Carlos, Province of Nuble, 
December 23, 1950. 

An adult alate collected by sweeping at Angol, Province of ^lalleco, 
January 29, 1951. 

Macrosiphum huantana Esslg, new species 

Huanta Aphid 
(Figure 43) 

Figure 43. Huanta aphid, Macrofiiphum huantana Essig, new species. A, an- 
tenna, cornicles, and cauda of the apterous parthenogenetic female; B, antenna, 
rostrum and wing of alate. 

Type: Alate parthenogenetic female: A dark species remaining brown- 
ish when cleared and mounted; without definite pigmented areas, but 
with the appendages dark brown. Antennae slender with few blunt or 


knobbed hairs : secondary scnsoria on segment III varia])le in size and scat- 
tered over the basal half or two thirds; varying in number from 17 to 19. 
(In paratypes they vary in number after this pattern : 18-13, 17-19, 17-19, 
in the three alates available.) They are longer than the body, 4.20 mm. 
Rostrum long and slender with many small hairs or spines; extending to 
the third coxae. Wings 4 nun. long; clear; venation as drawn. Legs slen- 
der. Cornicles dark, long and cylindrical; widening basally and apically; 
apical one-fourth reticulated; length 0.75 mm. Cauda brownish, long, 
slender, gradually tapering posteriorly ; 12-14 hairs present ; 0.40 mm. 
long. Anal plate rounded, twice the width of the base of the cauda, with 
many long spines. 

Ai?terous parthenogcnctic female: Dark in color with all appendages 
brownish in cleared specimens ; without definite pigmented areas. Secon- 
dary sensoria on antennal segment III variable in size and number; usually 
limited to the basal half of the segment; ranging in number from 8 to 16 
after the pattern: 7-8, 7-9, 10-10, 12-12, 12-15, 16-16. Length of body 
3.50 mm. ; antennae 4.50 mm. ; cornicle 0.80 mm. 

Host plant and locality : Collected on Baccharis sp. along the Rio Turbio, 
branch of the Rio Elqui five miles south of Huanta fCuai.ta), Province 
of Coquimbo, Chile, December 7. 1950, by Dr. A. E. ]\Iic]ielbacher. 

This species is close to M. littoreilis Blanchard, but has more sensoria 
and hairs on rostrum and cauda and cornicles flaring at apices. 

This species also differs from other related species in having the ex- 
tremely long cauda and the long, slender rostrum. 

Altogether 30 specimens were taken : 8 alatae and 27 apterae. The 
type si)ecimen is an alate mounted on a slide along with another alate 
and three adult apterae paratypes. All other specimens are also desig- 
nated as paratypes. 

Macrosiphum li^erianum Blanchard 
Lizer Aphid 

A single apterous female of what appears to be this a]ihid was taken 
by sweeping. 

Chile : 

At Zapallar, Province of Aconcagua, November 27, 1950. 

Argentina : 

Blanchard (1922, pp. 185-7, fig. 1) collected this aphid on Sonchus 
sp., Cosmos sp., Aster sp., Wedelia gJauca, and species of Com- 
posite at Punta Chica, San Isidro, Flores and Canuelas, Province 
of Buenos Aires, and at Potrerillos and Cachueta, Pi-ovince of 
]Mendoza. He states it is close to M. soUdoginis (Fabricius). 



Macrosiphum macolai Blanchard 

Macola Aphid 
(Figure 44) 

Figure 44. Macaloi aphid, Macrosiphum mavuloi Blanchard. Apterous partheno- 
genetic female showing small knobbed setae arising from dark areas on the dorsum 
of the abdomen; cauda; cornicle; rostrum; and antennal segment III. 

Argentina : 

On Baccharis salicifolia Mendoza, June 10, 1926. (Blanchard 1936, 

pp. 29-30, fig. 6. Original description: 1939, 966-7.) 
On Erigeron sp. (chilca), Province of ]\Iendoza, by Blanchard (194-lr, 

p. 42). 


On the common dandelion, Taraxicuni vulgare, at El Abanico, Province 

of Bio-Bio, December 30, 1950 — 1 aptera. 
On Baccharis sp. ?, at Negrete, Province of l^io-Bio, January 30, 1951 

— 27 apterae. 

Macrosiphum muermosa Essig, new species 

Muermos Aphid 
(Figure 45) 

Vol.. XXVIII] 



Figure 45. Muermos aphid. Marrosiphum muermoHa Essig, new species. Ap- 
terous parthenogenetic female showing typical color pattern; a, wing, cauda, 
cornicle, and antennal segment III of alate; b. rostrum and antennal segment III 
of apterous female. 

TvpE: Alate parthenogenetic female: Probably deep red, brown, and 
black — the alcoholic preserving Huid material was stained a very deep red- 
dish-brown color. Cleared specimens appear black, dark brown, and trans- 
parently colorless. The spines are short and knobbed. The antennae are 
curved; api)roximately the length of the body and extending to the tip of 
the Cauda; dark to black in color; secondary sensoria only on segment III, 
circular to oval, rather small, ([uite variable in size, 43-44 in number 
and distributed throughout the full length, except the base and extremity; 
.spines, short, curved, slightly knobbed. (Paratypes show a variation of 
from 39-49 sensoria on antennal segment III.) Rostrum slender with 
narrow apical segments; extends to or nearly to the third coxae; with few 
incons]ucuous hairs. Legs slender ; iiind tibiae strongly curved ; tarsi very 
small. Cornicles black; wider at base, nearly cylindrical, slightly curved 
outwardly; apical one fourth plainly reticulated; 1 mm. in length. Cauda 


black, long, and well developed; with many hairs; 0.5 to 0.7 mm. long. 
Anal plate small and crescent shaped; with few rather long hairs. 

Length of body 4 mm. ; antennae 4.2 mm. ; forewing 4 mm. ; cornicles 
0.75 mm. ; cauda 0.50 mm. 

Apterous parthe^iogenetic females -. Dark reddish and black with the 
abdomen adorned with black, irregular markings. The antennae are slen- 
der, slightly longer than the body and curved. The spines are short, some- 
what curved or straight and terminate in a knob or slightly enlarged. The 
sensoria on antennal segment III are quite variable in size, extend over 
much of the length as drawn, and range in numbers after the following 
pattern: 22-25, 25-30, 26-33, 29-29, 32-34, 36-36, etc. Other charac- 
ters are similar to those in the alate. Length of body 4.5 mm. ; antennae 
4.2 mm. ; cornicles 1 mm. ; cauda 0.75 mm. 

This species belongs to the group of dark-colored aphids which have 
short, stiff, spine-like hairs; long, slender, curving antennae; slender ros- 
tra; many variable secondary sensoria only on antennal segment III of 
both alate and apterous forms ; and black pigmentation on the interseg- 
mented, marginal, antesiphunucular, and other areas of the abdomen 
such as occur in Macrosiphum amhrosiae (Thomas), Dactynotus picridis 
(Fabricius), D. taraxaci (Kaltenbach) and many other related species. 

Host plant and locality: Taken on a "yellow composite with long lance- 
like leaves, light pubescence on the undersurfaces giving silver color, ' ' at 
Los Muermos, Province of Llanquihue, Chile, by Dr. A. E. Michelbacher. 
Many specimens of l:>oth alate and apterous forms are mounted on 13 slides. 

A single alate specimen, on a slide with two apterous females, has 
been designated as the type; all others as paratypes. 

This species resembles somewhat M. lizerianum Blanchard, but differs 
in being larger ; darker ; with many more sensoria on antennal segment III 
in both alate and apterous forms, with a wide distribution of sensoria in 
the aptera ; the slender cauda ; and long, slender rostrum. 

Macrosiphum nuble Essig, new species 

Nuble Aphid 
(Figure 46) 

Type : Ajiterous pariheyiogenetic female : Color generally pale green 
(?); head, rostrum antennae, most of the legs, cornicles, excepting the 
basal portions, cauda, dark or brownish. There are no body markings 
on the apterous form described but it is l)eset with rows of short spines 
arising from basal cones and curved (drooping) and knobbed. The an- 
tennae are long and slender with 13 to 16 small circular secondary sensoria 
irregularly arranged on the ventral basal fourth or third of the segment. 
(In paratypes the ratio on 4 specimens is 7-9, 9-13, 12-15, 13-15.) The 

Vol. XXVIII 1 



Figure 46. Nuble aphid, Macroslijhum ntible Essig, new species. Apterous par- 
thenogenetic female and enlarged cauda, cornicle, rostrum and antennal segment 
III showing sensoria. 

rostrum is short, reaching to the third coxae. The proportions of the various 
segments and the spination are illustrated. The cornicles are unusually- 
long, 0.80 mm.; curved outwardly- cylindrical or slightly swollen, and 
reticulated, but not constricted apically. The cauda is relatively short, 
0.30 mm., rather thick and with many spines. The anal plate is very 
small and semioval. Length of body 3.30 mm. 

Host plant and locality: The s])eeies was collected on a composite plant 
in the Andes Mountains near Fabian, Province of Nuble, Chile, Decem- 
ber 15, 1950, by Dr. A. E. :\lichelbacher. 

The specimens consist of seven apterous females which are mounted on 
three slides. One individual is indicated as the type. All the others aro 
designated as paratypes. No alates were collected. 


This species resembles Macrosiphum solanifolii (Ashmead) in having 
the short rostrum and the apically reticulated cornicles, but differs in 
having a much larger number of secondary sensoria in the apterae which 
are irregularly arranged on the base of antennal segment III ; by the much 
longer cornicles; by the more attenuated rostral apex; and the shorter, 
thicker cauda. 

Macrosiphum rosae (Linnaeus) 

Rose Aphid 
(Figure 47) 

Aphis rosae Linnakis, 1758; A. scaMosae Scopoli, 1763; A. dii)saci Schrank, 1801; 
Hiphonophora rosaecola Passerini. 1871; Nectarophora valerianiae Clarke, 
1903; Macrosiphum centranthi Theobald, 1915; Macrosiphon rosae (Lin- 
naeus) BORNER. 1952. 

The rose aphid is almost a cosmopolitan species being widely dis- 
tributed through commerce. It occurs usually on wild and cultivated 
roses and temporarily infests such other plants as CamelUa japonica, Dip- 
sacus fullonum, D. sylvestris, Rudheckia hirta, and Sonchus spp. It is 
now well distributed in parts of South America. 

Chile : 

On cultivated rose, Valparaiso, November 26, 1950 — 1 alate and 6 

On Sonchus, sp., Valparaiso, November 26, 1950—1 alate and 17 

On cultivated rose, Vicuna, Province of Coquimbo, December 4, 1950 

— apterae and alatae. 
On beans, Fhaseohis sp., San Carlos, Province of Nuble, December 23, 

1950 — 1 alate — casual ?. 
On rose, Temuco, Province of Cautin, January 6, 1951 — apterae and 

Sweeping peafield, Lautaro, Province of Cautin, January 6, 1951 — 1 

apterae and 1 alate. 
On Sonchus sp., Rio Bueno, Province of Valdivia, January 14, 1951 

— 1 apterous. 

Peru : 

On cultivated rose, Lima, July 15, 1937. Collected by Mrs. Selma 
Gahl — 1 alate and 4 apterae. In author's collection. 
On cultivated rose, Botanical Garden, Lima, November 13, 1950 — 
abundant; many apterae and 1 alate. 




Figure 47. Rose aphid. Macrosiphum ro.sae (Linnaeus). A, alate parthenogenetic 
female; B, apterous female; C, antennal segment III of alate showing sensoria; 
D, antennal segment III of aptera ; E, cornicle of alate; F, cornicle of aptera; G, 
Cauda of alate; H, cauda of aptera. 



\ ^c^M^r^'^ 

Figure 48. Potato aphid, Macrosiphum solanifolii (Ashmead). A, apterous par- 
thenogenetic female with enlarged: a, cornicle; b, dorsum of rostrum; c, cauda; e, 
antenna; f, another antennal segment III. B, first instar young; g, tip of the an- 
tenna of same; h, sensoria in the process on antennal segment VI of embryo; i, 
enlarged antennal setae. 

On cultivated rose, Chielayo, Province of Lima, ]\Iarch 21, 1951 — many 
specimens; all apterae. 

Argentina : 

Common on cultivated roses throughout Argentina and also on loquat 
(nispero) Eriohotrya japonica. (Blanchard 1922, pp. 187-190, 
fig. 2; 1939, pp. 967-9). 

Macrosiphum solanifolii (Ashmead) 

Potato Aphid 
(Figures 48-49) 

Siphonopho7-a solanifolii Ashmead, 1882; S". euphorbiae Thomas, 1878; Macrosi- 
phum ciicurhitae del Guercio, 1913; Macrosiplium gei (Koch) Hottes and 
Frison, 1931; Macrosiphon solanifolii (Ashmead) Borner, 1952. 

This is a very widely distributed Holarctic species and appears to 




Figure 49. Potato aphid, Macrosiphum solatrifolii (Ashmead). A, alate par- 
thenogenetic female; a. hind tarsus; b, upper and lower portions of the hind tibia; 
c, Cauda; d. rostrum — ventral aspect; e, cornicle; f. antenna with two additional 
segments of segment III from other individuals to show variation ; g. tip of antenna. 

1)0 \\v\\ reprosonted in South America. The extensive collections help 
to substantiate this l)eliet'. 

Chile : 

On squash, Cucuvhitn maxima, Antofagasta, Xovcniber 21, 1950 — abun- 
dant ; apterae and alate. 

On Sonchus sp., Zapallar, Province of Aconcagua, November 27, 1950 
1 alate. 

On Cere us cactus. Las Palmas, Province of Aconcagua, November 29, 
1 950 — apterous. 

On beans, Phascolus sp., San Carlos, Province of Nuble, December 23, 
1950—1 alate. 


On artichoke, Cynara scolymus, San Carlos, Province of Nublc, De- 
cember 23, 1950—2 alatae. 

On anise, or dill, A^iethum graveolens, Angol, Province of Bio-Bio, 
January 1, 1951 — 1 alate. 

On morning glory, Convolvulus sp., Angol, January 1, 1951 — many 
apterae and alatae. 

On cultivated tomato (tomatera), Lycopevsicon esculentum, Angol, 
January 2, 1951 — 1 alate. 

On squash, Cucurhita maxima. Agricultural School, Angol, January 
4, 1951 — many apterae and alatae. 

On cultivated potato (patata, papa), Solamim tuberosum, Angol, Janu- 
ary 5, 1951 — 1 alate and 5 apterae. 

On garden pea, Lathynis sativus L., by sweeping, Lautaro, Province 
of Cautin, January 6, 1951 — 1 aptera and 1 alatae. 

Sweeping at Los Muermos, Province of Llanquihue, January 20, 1951 
1 alate. 

On wild potato, Puyehue, Province of Osorno, January 26, 1951 — 1 
apterous and 4 alatae. 

Argentina : 

Blanchard (1922, pp. 190-2, fig. 3; 1939, pp. 969-70: 1944, p. 43) lists 
the species on many host plants, including : 

Antirrhinum sp., (cardos), Carduus sp.. Citrus sp.. Cosmos sp., Cris- 
taria corchorifolia Oris., Curcurhita .spp., Ipomaa batatas (L.), 
Lactuca sp., Malva sp., Fyrus communis L., P. mahis L., Prunus 
avium L., Solanum lycopersicum L., 8. tuberosum L., Spiiiacia 
ohracea L., Vicia faba L., and Watsonia sp. 
The writer has recently received specimens from L. A. Bahamondes, 
collected by him 

On Tulipa sp. at jMendoza, October 13, 1950 — 1 aptera and 1 alate. 

Colombia : 

On Lantana sp., at the port of Buenaventura, November 3, 1950 — 
many si^ecimens of apterae and alatae. 


On bean, Phaseolus vulgaris L., at Callao, November 14, 1950 — 1 alate 

and many apterae. 
On marigold. Calendula officinalis, Callao, November 14, 1950 — 1 alate 

and many apterae. 
On marigold, Calendula officinalis, Callao, November 14, 1950 — 1 alate 

and a number of apterae. 




On pomegranate, Punica granatum L., Callao, November 14, 1950 — 
10 apterae. 

On marguerite, Chrysanthemum fnitescens, Callao, November 14, 
1950 — many apterae. 

On Datura sp., Botanical Garden, Lima, November 14, 1950 — 2 ala- 
tae and 10 apterae. 

On mallow, Malva sp., Callao, November 14, 1950 — 1 alate and many 

On tobacco, Nicotiana sp., Botanical Garden, Lima, November 14, 1950 
— 5 alates. 

On Solanum sp., Callao, November 14, 1950 — 2 alates and many 

On potato, Solanum tuberosum, Callao, November 16, 1950 — 5 alatae 

and 8 apterae. 
On cotton, Gossypium sp., Chanca, Province of Lima, ^Nlarch 15, 1951 

1 apterous. 
On Asclepias sp., and on thorny shrub (Leguminosae) at Chiclayo, 

Province of Lima, March 19, 1951 — all apterae. 

Macrosiphum tucumani Essig, new species 

Tucuman Aphid 
(Figure 50) 

Figure 50. Tucuman aphid, Macrosiphum tucumani Essig, new species. Ap- 
terous parthenogenetic female showing general color pattern; A, two antennal 
segment III of the alate; B, same for the aptera; C, cornicle of alate; D, rostrum 
of alate; E, wing; F, cauda of alate; G, cauda of aptera. 


Type: Alate parthenogenetic female: The color is apparently nearly 
black in the living forms and retains much of the same when cleared and 
mounted. A relatively small species averaging 2 mm. in length. The body 
spines or hairs are short and mostly knobbed or enlarged apically. The 
antennae are black; long and slender; longer than the body; 2:20 mm. 
The unguis or filament is about six times the length of the base. Segment 
III has the following distribution of secondary circular sensoria ar- 
ranged nearly in a row: 14-14. (In paratypes the following variation was 
noted: 13-15, 15-16 on the other six alates collected.) The rostrum is of 
even width to the pointed apical segment and extends to or just beyond 
the third coxae. The legs are mostly black, slender, and the tibiae curved 
as shown. The wings are as drawn and are 2.70 mm. in length. The second 
branch of the media is two-thirds beyond the base. The cornicles are nearly 
cylindrical, l)lack or somewhat paler basally with the apical one-fourth 
reticulated; length 0.47 mm. The base and apex are somewhat wider than 
the rest. The cauda is not quite half of the length of the cornicles or 0.22 
mm. and has 8 or 9 spine-like hairs. 

Apterous parthenogenetic female : Dark, body is covered with ver}^ 
many small blackish areas around tlie bases of the short knobbed spines as 
is a characteristic of such species as Macrosiphon amhrosiae (Thomas), 
Dactynotus jaceae Linnaeus, D. picridis (Fabricius), D. taraxaci (Kalten- 
bach), and others. Antennal segment III has circular secondary sensoria 
loosely distributed almost the full length in the following combinations: 
7-8, 7-12, 9-9, 9-10, 11-13, 12-12. The cornicles are somewhat longer 
than in the alate, being 0.50 mm. Length, 2 mm. 

Host plant and locality: On BaccJiaris sp. at Huanta (Guanta) on the 
Rio Turbia which flows into the Rio Elqui, Province of Coquimbo, Chile, 
December 4, 1950, by Dr. A. E. Michelbacher ; and on BaccJiaris sp. (?), 
on a ridge at 4,000 feet altitude, 24 kilometers west of Tucuman, Argentina, 
February 11, 1951, by Dr. Michelbacher. The collections consisted of 43 
adult and a number of immature apterous forms, and 7 alates. One of 
these, mounted on a slide with a number of alates and immature forms 
has been designated as the type. All others are labeled paratypes. 

This species is unusual in having a relatively short cauda and some- 
what bhuiter rostrum. The cornicles somewhat resemble those of M. solani- 
folii (Ashmead) in the arrangement and amount of reticulation, but num- 
ber and distribution of sensoria are quite different, especially the greater 
number on III of the apterae as figured. 

Vol. XXVIII 1 



Tribe Dactynotini 

Subtribe Dactynotina 

Amphorophora peruviana Essig, new species 

Peruvian Aphid 

(Figure 51) 

Type : AJate parthcnogeyietic female : A pale and dark species with 
dusky-bordered wing veins. The body, excepting the abdomen, is mostly 
dark; the abdomen has a few regular dark patches similar to those shown 
in the drawing of the apterous form. The antennae are slender, longer 
than the body — the unguis is about six times the length of the base. Cir- 
cular secondary sensoria of various sizes occur nearly in a row on the 
underside of segment III. The number may vary from 14 to 15. (In para- 
types there is a variation in the number of 14-19.) The rostrum is rela- 
tively short and extends onlj- to the third coxae; with many short spines 
as illustrated. The forewings are as drawn. The legs are slender. The 
cornicles are about twice as long as the cauda and are smooth, distinctly 
swollen, with two or three rings or broken reticulations at the apex, the 
flange is irregularly flaring, 0.55 mm. long. The cauda is trumgular, being 
more pointed in the alate than in the apterae; 0.30 mm. long and 0.14 mm. 
to 0.17 mm. wide at the base and with the normal number of hairs. Length 
of bodv 3.50 mm.: length of forewing 3.50 mm. 

Figure 51. Peruvian aphid, Amphoroiihora periiviana Essig, new species. A, 
aptera showing color pattern ; a, antenna! segment III of alate above and aptera 
below; b, cornicles; c, cauda of alate; d, cauda of aptera; e, rostrum upper and 
lower surfaces; f, wing. 



Figure 52. Sow-thistle aphid, Amphorophora sonchi (Oestlund). Alate and ap- 
terous forms with enlargements of antenna of alate, and antennal segment III of 
the apterous; cornicles of both, and cauda of alate. (After Zimmerman. 1948.) 


Apterous parthenogenetic female: Normal in size and form, 3 mm. in 
length. The general characteristics similar to those of the alate. Antennae 
without secondary sensoria. Cauda broader tlian in tlie alate, with few 

The type specimen was chosen from three alates (one designated as the 
type) and five mature apterae and one immature specimen. All mounted 
on four aluminum slides. 

The type is mounted singly on a slide. The remaining aUitae and all 
apterae are designated as paratypes. 

Host: Unknown; specimens were obtained by beating onto a canvas 

Locality: Rio Pampas, Peru, March 8, 1951. 

Collector: T)r. A. E. Michelbacher. 

This species somewhat resembles A. sonchi (Oestlund) but differs 
markedly in having much shorter and more triangular cauda ; thickened 
cornicular openings; hairy rostrum; and dark body markings. 

Amphorophora sonchi (Oestlund) 

Sow-thistle Ai)hid 
(Figure 52) 

Rhopalosiijhum sotirhi Okstumi, 1886; Amphorophora cosmopolitana M.vsox. 1925. 

This is a large green and black species which, in colder areas of its 
range, overwinters on liihes s])])., and passes the summer on Sonchus spp., 
and other related hosts. In the warmer areas it may pass both winter and 
summer on the latter. 

Chile : 

On Sonchus sj)., Zapallar. Province of Aconcagua, November 27, 1950 

— many ajiterae. 
On Sonchus sp., Rio Bueno, Province of Valdivia, January 14. 1951 

— a number of apterae. 

Argentina : 

On Sonchus h\)., in Province of Buenos Aires. Listed as Amphoro- 
phora lactucae (Kaltenbach) Blanchard (1922, pp. 207-9, fig. 11). 
On Lactuca spp., So7ichus spp., and Cichorium endivia in Province of 
Buenos Aires. Listed as Amphorophora cosmopolitan us Mason 
(Blanchard, 1939, pp. 951-53). 


PMgure r).!. Woolly apple aphid. Kriosoma laniyero (Hausmann), as it appears 
on apple. 




Tribe Eriosomatini 

Eriosoma lanigera (Hausmann) 

Woolly Apple Aphid 
(Figures 53-54) 


A2}his lanigera Hausmann, 1802; Coccus mali Bingley, 1803; Myzoxylus mali Blot, 
1831; Eriosoma mali Samoiklle, 1819; Scliiozoneura lanigera Hausmann, 

Figure 54. Woolly apple aphid, Eriosoma lanigera (Hausmann). Alate and 
apterous parthenogenetic females x66; antennae of alate and aptera ; cauda and 
anal plate and cornicle of alate; wax glands of aptera x 166. (After Zimmerman, 


This is a Holarctic species now eoininon throughout temperate Europe 
and North America. It has been widely distributed through commerce 
to all temperate regions where its hosts flourish. 

Chile : 

On cultivated apple (manzano), Anglo, Province of Bio-Bio, January 
1, 1951 ; only apterae were taken. The material showed about 20 
per cent parasitism by Aijhelinus mali (Ilaldeman) Avhich was 
introduced into Chile many years ago. 

Argentina : 

According to Blanchard (1926, pp. 333-35; 1939, pp. 983-84), the 
woolly a])ple aphid is common on apple wherever grown. He also states 
that the "introduction of ApheUnus mali Ilaldeman has done much to 
keep it in control." 


Tribe Pemphigini 

Pemphigus populi-transversus Riley 

Transverse Poplar Gall Aphid 
(Figure 55) 


What is believed to be this species was collected by Dr. A. E. Michel- 
bacher on the roots of plants exposed by turning stones in the bed of the 
Red River, Callao, November 16, 1950. The lot consisted of 7 apterae and 
15 alates. 

Argentina : 

Blanchard (1926, })p. 335-337: 1939, pp. 986-87) records this species as 
forming subglobular galls on the leaf petioles of Populus canadensis which 
may have been introduced into the Province of Buenos Aires. Also on 
Popidus anf/ulata Ait. 

Thecabius populi-monilis (Riley) 

liead-like Cottonwood Gall Aphid 
(Figure 56) 

Pemphigus populi-monilis Rti.p:y, 1879; Thecabius popuHvionilis (Riley) Gillette, 


This aphid makes elongated marginal galls in rows along the veins of 


Figure 55. Galls of the transverse poplar gall aphid. Pevtphujus i)oi)uU-traus- 
i-ersus Riley, on the petioles and leaf bases of Populus sp. 



the leaves of PoiJiilus spp. A single alate specimen was taken at Zapallar, 
Province of Aconcagua, November 27, 1950. 

Figure 56. Bead-like cottonwood gall aphid, Thecabius poimli-monilis (Riley). 
A, alate; B, antenna of alate enlarged. 


Abeto. Vease Abies sp. 
Abies sp. 

Cinara grossus (Kalt.), Argentina. 

Cinara hyalina (Koch). Argentina. 

Dreyfusia nordmanniana Ecks. 

Abrojo. Vease Xantliium sp. 
Abut Hon sp. 

Myzus persicae (Sulzer). Argentina. 
Acacia sp. 

Aphis fabae Scopoli. Argentina, 
medicaginis Koch. Argentina. 
Acacia mansa. Vease Sesbania punicea 

(D. C). 
Achicoria. Vease Chicorium intybus L. 
Acrostichum sp. Helecho. 

Idopterus nephrelepidis Davis. 

Adelfa. Vease Nerium spp. 
Adesmia sp. 

Aphis laburni Kalt. 
Adiantum sp. Helecho. 

Idiopterus nephrelepidis Davis. 
Agave sp. (Maguey, cabuya). 

Aphis rumicis L. Peru. 
Agraciejo. Vease Berber-is sp. 
Aguacate Vease Persea americana Mill. 
Aji. Vease Capsicum spp. 
Alamo. Vease Populus spp. 
Albaricoque. Vease Pruuus armeniaca 

Alcaucil. Vease Cynara scolymus L. 
Alfalfa. Vease Medicago satiiKi L. 
Algodonero. Vease Gossyinum spp. 
Alheli. Vease Mathiola incana. 
Almendro. Vease Prunus domestica L. 

(P. communis Huds.) 
Altamisa. Vease Descuraitiia appendi- 

culata (Gris.). 
Alstrameria sp. 

Aphis alstroemeria, n. sp. Chile. 
Amaryllis belladonna L. 

Aphis gossypii (Glover). 
Andropogon bicornis L. 

Hysteroneura ogloblini Blanchard. 

Anethum graveolens L. 

Cavariella aegopodii (Scopoli). 

Macrosiphum solanifolii (Ashm.). 
Anis. Vease Pimpinella anisum L. 
Aniirrliinum majus L. 

Macrosiphiun solanifolii (Ashm.). 

Myzus persicae (Sulzer). Argentina. 
Apio. Vease Apium graveolens L. 
Apium graviolens L. 

Cavariella aegopodii (Scop.). 

Trifidaphis phaseoli (Pass.). 
Arachis hypogaea L. (cacahue, 
Aphis laburni Kalt. Argentina. 
Aphis medicaginis Koch. Argentina. 
Araujia sericofera Bert. 

Aphis nerii B. d. Fonsc. 
Arrayan. Vease Orthostemon sellotvi- 

aiius Berg. 
Arroz. Vease Oryza sativa L . 
Artemisia spp. 

Macrosiphum ambrosias (Thomas). 

Argentina, Chile. 
Aphis medicaginis Koch. Chile. 
Arum sp. 

Pentalonia nigronervosa Coquer. 
Argentina, Brazil. 
Asclepias sp. 

Aphis gossypii Glover. Argentina, 

Aphis nerii B. d. Fonsc. Bolivia, Peru. 
Macrosiphum solanifolii (Ashm.). 
Asclejjias curassavica L. 

Aphis nerii B. d. Fonsc. Argentina. 
Asparagus officinalis L. 

Myzus persicae (Sulzer). Argentina. 
Aster spp. 

Brachycaudus helichrysi (Kalt.). 

Macrosiphum lizerianum Blanchard. 




Astralago. Vease ANtragalus 
Astragalus sp. (astragalo). 

Aphis medicaginis Koch. Argentina, 
Atriplex sp. 

Aphis gossypii Glover. Colombia. 
Avellano. Vease Corylus avellana L. 
Avena. Vease Algeria sativa L. 
Avena sativa L. 

Rhopalosiphum maidis (Fitch). 

Toxoptera graminum (Rondani). 
Azucena. Vease Amaryllis belladonna 

L. and Lilium spp. 
Baccharis spp. 

Brachycaudus helichrysi (Kalt.). 

Macrosiphum ambrosiae (Thomas). 

chilensis, n. sp. Chile, 
cordobensis Blanchard. 

huantana n. sp. Chile, 
lizerianum Blanchard. Argentina, 
macolai Blanchard. Argentina. 
tucumani n. sp. Argentina, Chile. 
Aphis gossypii Glover. Colombia, 
medicaginis Koch. Chile. 
Baccharis ))iel<tst())iiaefolia Gris. 
Brachycaudus helichrysi (Kalt.). 

(BL). Argentina. 
Aphis gossypii Glover. Argentina. 
Macrosiphum littoralis Blanchard. 
Baccharis salicifolia Pers. 

Macrosiphum macolai Blanchard. 
Baladre. Vease Nerium spp. 
Barnadesia odorata Griseb. 

Myzus persicae (Sulzer). Argentina. 
Batata. Vease Iponwea batatas (L.). 
Begonia sp. 

Aphis gossypii Glover. Argentina. 
Aulacorthum pseudorosaef olium 

Blanchard. Argentina. 
Myzus omatus Laing. General 

persicae (Sulzer). Argentina. 
Bellis perrenis L. 

Myzus persicae (Sulzer). Argentina. 

Berberis sp. (Berbero, agracejo.) 

Aphis illinoisensis Shinier. Chile. 
Berberis buxifolia Lam. 

Aphis patagonica Blanchard. 
Berbero. Vease Berberis sp. 
Beta vulgaris L. 

Myzus persicae (Sulzer). Argentina. 
Bidens megaiiotanrica Spreng. 
Aphis coreopsidis (Thomas). 
Biguonia sp. 

Aphis gossypii Glover. Chile. 
Myzus ornatus Laing. Chile. 
Bdhvieria nivea Gaud. 

Myzus persicae (Sulzer). 
Bonetero. Vease Euonymus sp. 
Brassica nigra L. (Mostaza negro.) 
Brevicoryne brassicae (L.). 

Argentina, Brazil, Chile, Peru. 
Rhopalosiphum pseudobrassicae 
( Davis). Argentina. 
Brassica oleravea L. Coliflor. 

Brevicoryne brassicae (L.). Argen- 
tina, Brazil, Chile, Peru, Trini- 
dad Island. 
Myzus persicae (Sulzer). Argentina. 
Brassi((( rapa L. 

Brevicorjme brassicae (L.). Argen- 
tina, Chile, Brazil, Peru. 
Myzus persicae (Sulzer). Argentina. 
Rhopalosiphum pseudobrassicae 
(Davis). Argentina. 
Brassica sinapistrum Boisd. 
Brevicoryne brassicae (L.). 
BroiHus sp. 

Rhopalosiphum prunifoliae (Fitch). 
Browns unioloides Kth. 

Rhopalosiphum psundoavenae 
(Patch). Argentina. 
Cabay. Vease Agave. 
Cacahuate. Vease Arachis hypogaea L. 
Cacahue. Vease Arachis hypogaea L. 
Cacao. Vease Theobroma sp. 
Cacto. Vease 

Cactus spp. 
Epiphyllum sp. 
Cafe. Vease Coffea sp. 
Calabaza. Vease Cucurbita spp. 




Calendula officinalis L. 

Brachycaudus helichrysi (Kalt.). 

Macrosiphum solanifolii (Ashm.). 
Camelia. Vease Camellia sp. 
Camellia spp. 

Aphis camellicola Del G. Argentina. 
Macrosiphum rosae (Linn.). 
Toxoptera aurantii (B. d. Fonsc). 
Cafia de aziicar. Vease Sacchai-um 

offlcinarum L. 
Caiiamo. Vease Cannabis sativa L. 
Canna sp. 

Rhopalosiphum nymphaeae Linn. 
Cannabis sativa L. (canamo). 

Myzus persicae (Sulzer). Argentina. 
Capsic u ni spp. ( A ji ) . 

Myzus persicae (Sulzer). Argentina. 
Capparis spinosa L. 

Brevicoryiie brassicae (Linn.). Peru. 
Myzus persicae (Sulzer). Peru. 
Cardencha. Vease Dipsacus fullonum L. 
Cardoncillo. Vease Eryngium panicula- 

tum Cav. 
Cardo negro. Vease Cirisium lanceola- 

tuni ( L. ) . 
Cardos. Vease Card u us sp. 
Carduus spp. 

Macrosiphum solanifolii ( Ashmead ) . 
Car ex spp. 

Rhopalosiphum maidis (Fitch). 
Car urn sp. 

Hydaphis conii (Davidson). Chile. 
Carum varri L. (Carvi). 

Cavariella aegopodii (Scopoli). 
Carvi. Vease Carum carvi L. 
Cassia sp. 

Aphis medicaginis Koch. Chile, Peru. 
Cassia (tphi/Ila Cav. 

Aphis gossypii Glover. Argentina, 
medicaginis Koch. Chile. Peru. 
Cassia bacillaris. (Taro). 

Aphis gossjrpii (Glover). Trinidad 
Castanea sativa Mill, (castafiio). 
Myzocallis castanicola Baker. 

Castaiio. Vease Castanea spp. 
Casuarina sp. 

Aphis gossypii Glover. Peru. 
Cebada. Vease Hordeum vulgare L. 
Cebadilla. Vease Brotnus unioloides 

Centeno. Vease Secale cereale L. (Rye). 
Cerens sp. (Genero de cactos). 

Macrosiphum solanifolii (Ashmead). 

Myzus persicae (Sulzer). Chile. 

ornatus Laing. Chile. 
Aphis medicaginis Koch. Chile. 
Cerraja. Vease Sonchus oleraceus L. 
Cestrum parqui L'Herit. (Duraznillo 
negro ) . 
Aphis rumicis L. Argentina. 
Chaenomeles japoniva (Thunb). Lindl. 
(Membrillo japones.) 
Aphis rumicis L. Argentina. 
Chaguar de las penas. Vease Dyckia 

poribunda Gris. 
Chayote. Vease Sechiuni edule Sw. 
Chenopodiuw, sp. 

Aphis gossypii Glover. Argentina. 
Myzus persicae (Sulzer). Argentina. 
Chilca. Vease Erigeron spp., Baccharis 

Chrysanthemum sp. (Crisantemo). 
Aphis rumicis L. Chile. 

medicaginis Koch. Argentina. 
Brachycaudus helichrysi (Kalt.). 

Capitophoraphis williamsoni 

Blanchard. Argentina. 
Macrosiphum artemisiae (Thomas). 
Chrysanthemum frutescens L. 

Macrosiphum solanifolii (Ashmead). 
Chrysanthemum indicum L. 
Aphis fabae Scopoli. Argentina. 
Brachycaudus helichrysi (Kalt.). 

Macrosiphoniella sanborni 
(Gillette), Argentina. 
Chrysanthemum ntaxiuiuni Ramond. 
Brachycaudus helichrysi (Kalt.). 



Cicer arietinum L. 

Rhopalosiphum pseudoavenae 
(Patch). Ar<;t'ntiiia. 
CicJwriiim end i via L. (Escarola). 
Amphorophora sonchi (Oestlund). 
(A. cosmopolitanus Mason.) 
Macrosiphum sonchi (L.). 
Cichormm intybus L. (Achicoria). 

Macrosiphum sonchi L. Argentina. 
Ciclamen. Vease Cyvlamen indicum L. 
Cicnta sp. 

Aphis medicaginis Koch. Argentina. 
Cicuta. Vease Coniuvi viaciihitmn L. 
Cineraria sp. 

Brachycaudus helichrysi (Kalt.). 

Myzus persicae (Sulzer). Argentina. 
Cipres. Vease Cupressus sp. 
Cirsium lanceohitum (L.). 
(cardo negro). 
Aphis fabae Scopoli. Argentina, 
rumicis L. Cliile. 
medicaginis Koch. Argentina. 
Ciruelo. Vease Prunus doniestica L. 
Citrulhis rulgaris Schrad. 

Aphis gossypii Glover. Argentina. 
Citrus spp. 

Aphis citricidus (Kirkaldy). 
Argentina, Chile, Peru, 
gossypii Glover. Argentina, 
laburni Kalt. Argentina. 
Macrosiphum solanifolii (Ashmead). 

Myzus circumflexus (Buckton). 

Paratoxoptera argentinensis 
(Blanchard). Argentina. 
Toxoptera aurantii (B. d. Fonsc). 
Argentina, Brazil, Trinidad 
Clavel. Vease Dianthus caryophylUts 

Clematide. Vease Clematis sp. 
Clematis sp. (clematide). 

Aphis nerii (B. d. Fonsc). 
Coccoloha sp. 

Toxoptera aurantii (B. d. Fonsc). 

Coffea (cafe). 

Toxoptera aurantii (B. d. Fonsc). 
Coliflor. Vease Brassica oleracea L. 
Colocasia esciilenta (L.). (Taro.) 
Aphis gossypii Glover. Trinidad 
Comida de vibora. Vease Lyciuin 

argentinum Hier. 
Comino. V6ase Cuminum vyviinum L. 

Compositae (various genera and 
Aphis coreopsidis Thomas. Colombia, 
gossypii Glover. Colombia, 
medicaginis Koch. Argentina, 
rumicis Linn. Peru. 
Macrosiphum beretica Blanchard. 

bonariensis Blanchard. Chile, 
griersoni Blanchard. Chile, 
lizerianum Blanchard. Argentina, 
muermosa, n. sp. Chile, 
nuble, n. sp. Chile. 
Conejitos. Vease Antirrhinum majus L. 
Conium maculatuvi L. 

Aphis fabae Scopoli. Argentina. 
Conrolvuliis sp. (Domperdo, Dondiego 
de dia). 
Macrosiphum solanifolii (Ashmead). 

Myzus persicae (Sulzer). Chile. 
Coqueta. Vease Bellis perennis L. 
Corona de novia. Vease Spiraea 

chamaedrifolia L. 
Coryliis arellana L. (Avellano). 

Myzocallis coryli (Goeze). 
Cosmos sp. 

Aphis medicaginis Koch. Argentina. 
Macrosiphum solanifolii (Ashmead). 

Myzus persicae (Sulzer). Argentina. 
Cosnujs hipen)iatus Cav. 

Aphis fabae Scopoli. Argentina. 
Macrosiphum lizerianum Blanchard. 

solanifolii ( Ashmead ) . 
Myzus persicae (Sulzer). Argentina. 
Crataegus sp. (Espino). 

Aphis gossypii Glover, Argentina. 




Crisantemo. Vease Chrysanthemum 

indicum L. 
Crisfdiiti corchorifoUa (iris. 

Macrosiphum solanifolii (Ashmead). 
Cucumis melo L. 

Aphis gossypii Glover. Argentina. 
Cucumis sativus L. (Coliombro o 
Aphis gossypii Glover. Trinidad 
Cucurhita maxi))ia L. (Calabaza). 
Aphis gossypii Glover. Brazil, 

Chile, Peru, Trinidad Island. 
Macrosiphum solanifolii (Ashmead). 
Argentina, Chile. 
Cioninum cyminum, L. (Comino). 

Myzus persicae (Sulzer). Argentina. 
Cupressus macrocarpa Hartw. 
Cinara fresai Blanchard. Argentina. 
CychiDirn iw^Jicum L. (Ciclamen). 
Myzus circumflexus ( Buckton ) . 
Cyvloma spp. 

Aphis gossypii Glover. Argentina. 
Cyclonia japonica Pers. (Membrillo 
Aphis fabae Scopoli. Argentina, 
gossypii Glover. Argentina. 
Cydonia vulgaris Pers. (Membrillo.) 
Aphis gossypii Glover. Argentina. 
Cynara srolymus L. (Alcaucil). 
Capitophorus braggi (Gillette). 

Argentina, Chile. 
Macrosiphum griersoni Blanchard. 

solanifolii (Ashmead). Chile. 
Myzus persicae (Sulzer). 
Cynodon dactylon (L.). 

Paraprociphilus graminis Blanchard. 
Dahlia sp. 

Aphis medicaginis Koch. Argentina. 
Dahlia rariabilis Desf. (Dalia). 

Aphis fabae Scopoli. Argentina. 
Dalia. Vease Dahlia variabilis Desf. 
Damasco. Vease Prunus armeniaca L. 

Datura sp. 

Macrosiphum solanifolii (Ashmead). 

Myzus persicae (Sulzer). Chile, 
Daurus carota L. (Zanahoi'ia). 
Cavariella aegopodii (Scopoli). 
Descurahiia appendiciilata (Gris.). 

Myzus persicae (Sulzer). Argentina. 
Dlanthus varyophyUus L. (Clavel). 
Myzus persicae (Sulzer). Argentina. 
Trifidaphis phaseoli (Passerini). 
Diente de leon. Vease Taraxacum vul- 
gar e (Lam.). 
Dipsaciis fuUonum L. (Cardencha). 
Macrosiphum rosae (Linn.). 
Dipsacus sylvestris Huds. 
Macrosiphum rosae (Linn.) 
Domperdo. Vease Convolvulus spp. 
Dondiego de dia. Vease Convolvulus 

Duraznillo negro. Vease Cestrum 

parqui L'Herit. 
Duraznero. Vease Prunus persica L. 
Dyckia floribunda Gris. (Chaguar). 
Aphis fabae Scopoli. Argentina, 
gossypii Glover. Argentina, 
medicaginis Koch. Argentina. 
Echinodorus sp. 

Rhopalosiphum nymphaeae (L.). 
Encclia sp. 

Macrosiphum ambrosiae (Thomas). 
Epiphyllum sp. 

Aphis rumicis L. Chile. 
Eragrostis pilosa ( L. ) . 

Carolinaia ogloblini (Blanchard). 
(Hysteroneura). Argentina. 
Erigero)i sp. 

Macrosiphum cordobensis 
Blanchard. Argentina. 
macolai Blanchard. Argentina. 
Eriohotrya japonica Lindl. 

Macrosiphum rosae (L.). Argentina. 



Eryngium sp. 

Aphis eryngii Blanchard. Argentina, 
medicaginis Koch. Argentina. 
Eryngium paniculatiivi Cav. 

Aphis fabae Scopoli. Argentina. 
Escarola. Vease Cichorium encliina L. 
Espadana. Vease Typha sp. 
Esparrago. Vease Asixiragiis ojjirinalis 

Espinaca. Vease Spinacia oleracea L. 
Espino. Vease Crataegus sp. 
Eucalipto. Vease Eucalyptus globulus 

Eucalyptus globulus Lab. (Eucalipto). 
Aphis fabae Scopoli. Argentina, 
medicaginis Koch. Argentina. 
Eugenia sp. (Arra>au). 

Aphis rumicis L. Argentina. 

gossypii var. malvoides Van der 
Goot. Argentina. 
Euonyinus sp. 

Aphis bazzii Elanchard. Argentina. 
Euonymus europaea L. 

Aphis fabae Scopoli. Argentina. 
Euonymus japanica Thunb. 

Aphis fabae Scopoli. Argentina, 
medicaginis Koch. Argentina. 
Eyphorlna portulacoicles Speng. 
Aphis pseudopulchella Blanchard. 
Feijoa sp. (Feijao). 

Smynthurodes betae Westwood. 
[Trifidaphis phaseoli (Pass.)]. 
Ficus sp. (Higuero). 

Toxoptera aurantii (B. d. Fonsc). 
Flox. Vease Phlox. 

Foeniculum vulgar e Gaertn. (Hinojo). 
Aphis fabae Scopoli (?). 

medicaginis Koch. Argentina. 
Cavariella aegopodii (Scopoli). 

Sappaphis apiifolia (Theobald). 
Fragaria resca L. Frutilla. 

Capitophorus frag:^efolii (Ckll.). 
Frijol. Vease Phaseolus. 
Frutilla. Vease Fragaria vesca ]j. 

Fuchsia spp. 

Myzus ornatus Laing. Peru, Chile 

Fucsia. Vease FucJisia. 

(Jarbanzo. Vease Cicer arietnum L. 

Gardenia forida L. (Jazmin). 

Aphis gossypii Glover. Argentina. 
Toxoptera aurantii (B. d. Fonsc). 
Genero de cactos. Vease Cereus sp. 
Geranio. Vease Pelargonium spp. 
Gerbera jamesoni Balus. 

Aphis fabae Scopoli. Argentina. 
medicaginis Koch. Argentina. 
Macrosiphum ambrosiae ( Thomas ) . 
Gladiolo. Vease Gladiolus communis 

Gladiolus communis L. (Gladiolo). 
Aphis fabae Scopoli. (?). Argentina, 
medicaginis Koch. Argentina. 
Gliricidia sepium (G. maculata). 

Aphis medicaginis Koch. Trinidad 
Gloxinia sp. 

Aphis medicaginis Koch. Argentina. 
Gossypium spp. ( Algodonero) . 

Aphis gossypii Glover. Argentina, 

Peru, Trinidad Island. 
Macrosiphum solanifolii (Ashmead). 
Gourliea decorticans Gill. & Hook. 

Aphis laburni Kalt. Argentina. 
Graminea- (Hierba). 

Geoica lucifuga (Zehntner). 

Ehopalosiphum maidis (Fitch). 

splendens (Theobald), (sobre 
Raices). Peru. 
Toxoptera graminum (Rondani). 
Granada. Vease Punica granatum L. 
Grosellero. Vease Ribes sp. 
Guava. Vease Psidium guajava L. 
Guayabo. Vease Psidium guajava L. 
Guayava. Vease Orthostemon seV.oiri- 

anus Eerg. 
Guayule. Vease Parthenium acetafu)n 

Guindo dulce. Vease Prunus arium L. 




Guisante. Vease Lathyrus sp. 

Haba. Vease Vicia faba L. 

Haya sudamerieana. Vease Nothofagus 

Hedera helix L. (Hiedra). 
Aphis hederae Kaltenbach. 
Helecho. Vease Acrostichum. Arlian- 

turn. 'Neplirolepis. 
Hibisco. Vease Hibiscus sp. 
Hibiscus sp. (Hibisco). 

Aphis gossypii Glover. Argentina, 

Trinidad Island. 
Toxoptera aurantii (B. d. Fonsc). 
Hiedra. Vease Hedera helix L. 
Higuera. Vease Ficus sp. 
Hinojo. Vease Foeniculum rule/are 

Holcus halepense (L. ). Vease Sorghum 

Hordcum rulgare L. (Cebada). 

Aploneura lentici Pass. Argentina. 
Rhopalosiphum maidis (Fitch). 

Sipha carrerai Blanchard. 

Toxoptera graminum (Rondani). 
Hyptis spicata Poit. 

Macrosiphum hyptidis Blanchard. 
Ilex paraguayensis Bonop. 
(Yerba mate). 
Toxoptera aurantii (B. d. Fonsco- 
lombe). Argentina. 
Iitiopaa batatas Lam. (Batata). 

Macrosiphum solanifolii (Ashmead). 
Iris Jlorcntina L. 

Yezabura tulipze (B. d. Fonsc). 
Jacaranda sp. 

Aphis gossypii Glover. Chile. 
Jacaranda ovaUfolia R. Br. 

Aphis gossypii Glover. Argentina. 
Jazmin. Vease Gardenia jiorida L. 
Jazmin de Chile. Vease Mandevillea 

suaveolens Lindl. 
Judia. Vease Phaseolus spp. 

Juglans regia L. (Nogal). 

Tuberolachnus salignus (Gmelin). 
Juniperus uvifera Don. Vease Phil- 
gerodendron uviferuin (Don) 
Lactuca spp. (Lechuga). 

Amphorophora sonchi (Oestlund) 
(A. cosmopolitanus Mason.) Ar- 
gentina, Brazil. 
Macrosiphum lizerianum Blanchard. 

Myzus persicae (Sulzer). Argentina. 
Lagenaria siceraria (Mol.) Standi. 
Aphis gossypii Glover. 
Trinidad Island. 
Lantana sp. 

Macrosiphum solanifolii (Ashmead). 
Lathyrus sp. (Guisante. Pesal.) 

Macrosiphum solanifolii (Ashmead). 
Lathyrus satirus L. (Guisante. Pesal.) 
Acyrthosiphon onobrachis (B. d. 
Fonsc.) [Macrosiphum pisi 
(Kalt.).] Cosmopolitan. 
Macrosiphum solanifolii (Ashmead). 
Laurel negro. Vease Nectajidra sp. 
Laurel rosa. Vease Nerium oleander L. 
Laurel fino. Vease Viburnum tinus L. 
Lavatera arborea L. (Malvon). 

Aphis gossypii Glover. Argentina. 
Lechuga. Vease Lactuca sativa L. 

Acyrthosiphon onobrachis (B. d. 

Fonsc). Peru. 
Aphis medicaginis (Koch). 
Chile, Colombia, Peru, 
rumicis Linn. Argentina. 
Lens esculenta Moench (Lenteja). 
Aphis labumi Kalt. Argentina. 
Lenteja. Vease Lens esculenta Moench. 
Lepidium sp. 

Aphis gossypii Glover. Argentina. 
Rhopalosiphum pseudobrassicae 
(Davis). Argentina. 
Libocedrus cupressoides Sargent. 

Vease Pilgerodendron urifera 
(Don) Florin. 



Libocedrus tetragona (Hooker). 

Vease Pilgerodendron uvifera 
(Don) Florin. 

Myzus circumflexus (Buckton). 
Lilium sp. (Azucena). 

Aphis gossypii Glover. Argentina. 
Lonicera confusa DC (Madreselva). 
Hyadaphis conii (Davidson). 

Myzus persicae (Sulzer). Argentina. 
Loto. Vease Nelnmhium sp. 
Lyciuv) argentinum Hier. 
(Comida de vibora). 
Aphis rumicis L. Argentina. 
Ly coper sicon esculentuv} L. 
(Tomate, Tomatera). 
Aphis gossypii Glover. Argentina, 
rumicis L. Argentina. Brazil. 
Macrosiphum solanifolii (Ashmead). 

Argentina, Chile. 
Myzus persicae (Sulzer). Argentina. 
Madreselva. Vease Lonicera confusa 

Maguey. Vease Agave. 
Maiz. Vease Zea mays L. 
Malva spp. 

Macrosiphum solanifolii (Ashmead). 
Argentina, Peru, 
solutum Blanchard. Argentina, 
urtica (Schr.). Argentina. 
Myzus persicae (Sulzei-). Argentina. 
Malva rosa. Vease Pelargonium 

graveolens L'Herit. 
Malvon. Vease Lavatera arborea L. 
ManderiUea snai^eolens Lindl. 
(Jazmin de Chile). 
Aphis gossypii Glover. Argentina. 
Toxoptera aurantii (B. d. Fonsc). 
Mangifera sp. (Mango). 

Toxoptera aurantii (B. d. Fonsc). 
Mani. Vease Arachis hypogaea L. 
Manzano. Vease Pyrus malus L. 
Mango. Vease Mangifera sp. 

Aphis fcenecionicoides Blanchai'd. 

Mathiola hederacea (Rabanito). 
Bhopalosiphon pseudobrassicae 

(Davis). Argentina, Peru. 
Myzus persicae (Sulzer). Argentina. 
Mathiola incanu L. 

Myzus persicae (Sulzer). Argentina. 
Medicago satira L. (Alfalfa). 

Aphis laburni Kalt. Argentina, 
gossypii (Jlover. Argentina. 
Melon. Vease Cucumis vielo L. 
Membrillo. Vease Cydonia vulgaris 

Membrillo japones. Vease Cydonia 

japonica Pers. 
i\Ienta. Vease Mentha arvensis L. 
Mentha arrensis L. 

Kaltenbachiella pallida (Haliday). 
(K. menthae Schout.). 
Mimosa sp. 

Hysteronettra ogloblini Blanchard. 
Molle. Vease Schinus dependens 

( Ortega ) . 
Morrcnia odorata (H. & Arn.). 

Aphis gossypii Glover. Argentina. 
Mostaza negra. Veasa Brassica nigra. 

Mostaza silvestre. Vease Brassica sina- 

pistrum Boise. 
Musa sp. (Banana). 

Pentalonia nigronervosa Coquer. 
Brazil, Trinidad Island. 
Mo ntezu m a speciosissim a 

Aphis gossypii Glover. Trinidad 
Nabo. Vease Brassica rapa L. 
Nasturcia. Vease. Nasturtium offici- 
nalis L. 
Nasturtium officinale R. Br. 
Aphis medicaginis Koch. Chile. 
Rhopalosiphum pseudobrassicae 
(Davis). Bolivia. 
Neciandra sp. (Laurel negro). 

Neolizerius tuberculatus Blanchard. 
XcliiDihiu)!! sp. (loto, ninfea). 

Rhopalosiphum nymphaeae (Linn.). 

Vol.. XXVIII] 



Xephrolepis spp. (Helecho). 

Idiopterus nephrelepidis (Davis). 
yerium oleander L. (adelfa, baladre). 
Aphis nerii (B. d. Fonsc). 
Argentina, Chile. 
Nieotiana sp. (tabaco). 

Macrosiphvim ambrosiae (Thomas). 

solanifolii (Ashmead). Peru. 
Myzus persicae ( Sulzer ) . Peru. 
Ninfea. Vease Neliimbium sp. 
Nispero. Vease Eriohotrya japonicu 

Nogal. Vease JugJans regia L. 
Noguera. Vease Juglans regia L. 
Xothophagus sp. 

Neuquenaphis edwardsi (Laing). 
Xot]iofagi(s domhei Blume. 

Neuquenaphis michelbacheri n. sp. 

chilensis, n. sp. Chile. 
Spicaphis michelbacheri, n. sp. Chile. 
yyri}l)h<i<t sp. 

Rhopalosiphum nymphaeae L. Brazil. 
Ocottd (irutifolia (Nees. ). 
Lizerius ocoteae Blanchaid. 

Aphis gossypii Glover. Argentina. 
Ohno. Vease UIdius spp. 
Ombu. Vease Phytolacca dioica L. 
Onohrychis satira Lam. 

Aphis laburni Kalt. Argentina. 
Orch idaceae ( Orquidea ) . 

Myzus circumflexus (Buckton). 
Orquidea. Vease Orchidaceae. 
Ortlwstemon sello^riaiuis Berg. 
Aphis gossypii Glover. Argentina. 
Ortiga. Vease Vrtica spp. 
Oryza satira L. 

Aresha setigera Blanchard. 
Palmera. Vease Phoenix sp. 
Papa. Vease Solanum tuberosum L. 
Parthenium aretatvm (Gray). 

Macrosiphum lizerianum Blanchard 

Parthenium hysfcrojjhorus L. 

Aphis gossypii (Hover. Argentina. 
Macrosiphum lizerianum Blanchard. 
Pastinaca. Vease Pastinaca satira L. 
Pastinaca satira L. (Pastanica). 
Cavariella aegopodii (Scopoli). 
Patata. Vease Solanum tuherosuDi L. 
Pelargonium sp. (Geranio). 

Aulacorthum pelargonii (Kalt.). 

Macrosiphum solutum (Blanchard). 
Pelargonium graveolens L'Herit. 
(Malva rosea). 
Macrosiphum bosqui Blanchard. 
Pelon. Vease Prunus persica var. 

nucipersica Schn. 
Peral. Vease Pyrus com munis L. 
Persea americana Mill. 

Aphis gossypii Glover. Argentina. 
Persea sp. 

Toxoptera aurantii (B. d. Fonsc). 
Pesal. Vease Lathyrus sativus L. 
Phaseolus sp. (Judia. Frijol). 

Macrosiphum solanifolii (Ashmead). 

Smynthurodes betae ( Westwood ) . 
[Triiidaphis phaseoli (Passe- 
rini)]. Argentina. 
Phaseolus lunatus L. 

Aphis laburni Kalt. Arengtina. 
medicaginis Koch. Argentina. 
Phaseolus rulguris L. (Judia, Frijol). 
Aphis gossypii Glover. Brazil. 
Macrosiphimi solanifolii ( Ashmead ) . 
Phlox sp. (Flox). 

Myzus persicae (Sulzer). Peru. 
Phabe porphyria Griseb. 

Lizerius ocoteae Blanchard. 
Phoenix sp. 

Aphis fabae Scopoli. Argentina, 
medicaginis Koch. Argentina. 
Ph rag mites ph rag mites (L.). (P. 
communis Trin.). 
Hyalopterus arundinis (Fab.). Chile. 



Phytolacca dioica L. 

Toxoptera aurantii (B. d. Fonsc). 
Picris sp. 

Macrosiphum ambrosiae (Thomas). 
Pilgerodenflron urifcrum (Don) 
Chileaphis michelbacheri n. sp. 
Pinipinella anisin)i L. 

Cavariella aegopodii (Scopoli). 

Myzus persicae (Sulzer). Argentina. 
Pino. Vease Pinus spp. 
Pinus spp. 

Pineus havrylenkoi Blanchard. 
Pinus halepensis Mill. 

Cinara pini L. (Lachnus pineti 
Koch). Argentina. 
Pinus radiata Don. 

Cinara pini L. (Lachnus pineti 
Koch). Argentina. 
Pinus pinaster Ait. 

Cinara pini L. (Lachnus pineti 
Koch). Argentina. 
Piretro. Vease Pyrethrum sp. 
Pittosporuyn sp. 

Aphis fabae Scop. var. bazzii 
Blanchard. Argentina. 
Polipodia. Vease Helecho, Ploypo- 

Polypodiaceae (Polypodia). 

Idiopterus nephrelepidis (Davis). 
Populus alba var. pyramidalis Bunge 
Pemphigus bursarius (L.). 

Phloemyzus passerinii (Signoret). 
Populus angulaia Ait. (Alamo). 

Pemphigus populi-transversus Riley. 
Argentina, Brazil. 
Populus canadensis Moench (Alamo). 
Pemphigus populi-transversus Riley. 
Populus simonii Ca)-r (Alamo). 
Pterocomma populeum (Kalt.). 

(Aphis populea Kalt.) Argentina. 

Porotilla de los sapos. Vease Vigna 
luteola (Jacq.). 

Prunvs spp. 

Aphis prunicola Kalt. Brazil. 
Appelia schwartzi (Borner). 

Brachycaudas helichrysi (Kalt.). 

Argentina, Chile. 

persicae-niger (Smith). 

Hyalopterus arundinis (Fab.). Chile. 
Myzus persicae (Sulzer). Argentina. 
Rhopalosiphum nymphasse (L.). 


Prunus armeniaea L. (Damasco). 
Brachycaudus schwartzi Boerner. 

Prunus avium L. 

Brachycaudus helichrysi (Kalt.). 

Macrosiphum solanifolii (Ashmead). 


Prunus douiestica L. (Ciruelo). 

Brachycaudus schwartzi (Borner). 
helichrysi (Kalt.). Argentina, 

persicae-niger ( Smith ) . 
Prunus persica L. (Duraznero). 
Appelia schwartzi (Borner). 

Brachycaudus helichrysi ( Kalt. ) . 
Argentina, Brazil, Chile, 
persicae-niger (Smith). 
Myzus persicae (Sulzer). Argentina. 

Psidium guajara L. (Guayabo, 
Aphis gossypii Glover. 

Punica granatuni L. (Granado). 
Aphis gOEsypii Glover. Peru. 
Macrosiphum solanifolii (Ashmead). 

Pyrethrum sp. (Piretro). 

Macrosiphum bonariensis Blanchard. 

Pyrus sp. 

Myzus persicae (Sulzer). Argentina. 




Pyrus commiini.s L. (Peral). 

Aphis gossypii Glover. Argentina. 

laburni (Kalt.). Argentina. 
Eriosoma lanigerum (Hausmann). 

Brazil (?). 
Macrosiplmm solanifolii (Ashmead). 
Pyrus mains L. (Manzano). 

Aphis gossypii Glover. Argentina. 
pseudopomi Blanchard. 
Eriosoma lanigerum (Hausmann). 

Argentina. Brazil, Chile, Peru. 
Hyalopterus arundinis (Fab.). 

Macrosiphum solanifolii (Ashmead). 
Quercus sp. 

Tuberculoides querciplatensis 
Blanchard. Argentina. 
Quercus ilex L. 

Tuberculoides grodsinskyi 
Blanchard. Argentina. 
Quercus rohur L. 

Tuberculoides elegans Blanchard. 
Rabanito. Vease Raphanus sativus L. 
Ramino. Vease Boehmeria nivea Gaud. 
Raphanus sativus L. 

Brevicoryne brassicae (L.). 

Khopalosiphum pseudobrassicae 
(Davis). Argentina. 
Remolacha. Vease Beta vulgaris L. 
Repollo. Vease Brassica oleracea L. 
Revienta caballo. Vease Holanum capsi- 
castrurn Link., iS'. sisyvibi'ifolium 
Ribes sp. 

Aphis rimiicis L. Chile. 
Roble. Vease Quercus spp. 
RoMnia pseudoacacia L. 

Aphis laburni Kalt. Argentina. 
Rosa. Vease Rosa centifolia y R. 

gallica L. 
Rosa spp. 

Aulacorthum pseudorosaefoliimi 

Blanchard. Argentina. 
Macrosiphum rosae (Linn.). 

Argentina, Brazil, Chile, Peru. 

Rosa centifolia and R. f/nlUca L. 
Aulacorthimi pseudorosaefolium 

Blanchard. Argentina. 
Macrosiphum rosae (L.). Argentina. 
Myzus rosarum (Kalt.). Argentina, 

Passerinia tetrahoda (Walker). 
Rumex spp. 

Aphis rumicis Linn. Argentina, 

Myzus persicae (Siilzer). Argentina. 
Rumex paraguayensis D. Parodi. 

Baizongiella solanophila Blanchard. 
Saccharum offlcinarum L. (caiia de 
Ehopalosiphum maidis (Fitch). 

Sipha flava Forbes. Argentina, 
Brazil, British Guiana. 
Salix spp. (Sauce). 

Aphis neosaliceti Blanchard. 

Cavariella aegopodii (Scopoli). 

Tuberolachnus saligna (Gmelin). 
Argentina, Brazil, Chile, Peru. 
Salix bahylonica L. 

Aphis citricidus (Kirkaldy). 

Cavariella aegopodii (Scopoli). 

Paratoxoptera argentinensis 
Blanchard. Argentina. 
Salix viminalis L. (Sauce "Osier"). 
Tuberolachnus saligna (Gmelin). 
Salsifi. Vease Tragopogon porrifolius 

Sambucus austi-alis Cham. & Schl. 

Aphis sambuci L. Argentina. 
Sandia. Vease Citrullus vulgaris 

Sauce. Vease Salix spp. 
Sauco. Vease Sambucus australis 

Chem. et Schl. 
Schijius dependens Ortega. 

Aphis schinifoliae Blanchard. 



Scirpus sp. 

Rhopalosiphmn maidis (Fitch). 
Scutia huxifolia Reiss. 

Toxoptera aurantii (B. d. Fonsc). 
Secale cereale L. (Centeno). 

Rhopalosiphum prunifoliae (Fitch) 
(R. pseiidoavenae Patch). 
^echiuiu edule Sw. 

Aphis gossypii Glover. Argentina. 
Senevio spp. 

Brachycaudus helichrysi (Kalt.). 

Macrosiphum capitophoroides 

Blanchard. Argentina. 
Myzus persicae (Sulzer). Argentina. 
Senivio honariensis Hook & Arn. 
Aphis fabae Scopoli. Argentina, 
medicaginis Kocli. Argentina. 
Sen-Sen. Vease Cassia aphilla. 
Seshania punicea. (DC). 

Aphis laburni Kalt. Argentina, 
medicaginis Koch. Argentina. 
Sida acuta Burm. 

Aphis gossypii Glover. Trinidad 
Sisyvihriiini Arnotiianum Gill. & Hook. 
Rhopalosiphum sisymbrii Del 
Guercio. Argentina. 
Solano. Vease Solanum spp. 

Aphis gossypii Glover. 
Solanum spp. 

Aulacorthum pelargonii (Kalt.). 

Peru. (Casual ?). 
Aphis medicaginis Koch. Argentina. 

rumicis L. Argentina. 
Macrosiphum solanifolii (Ashmead). 

Myzus persicae (Sulzer). Argentina, 

Rhopalosiphum splendens 

(Theobald). Peru. 
Smynthurodes betae (Westwood). 
(Trifidaphis phaseoli (Passe- 
Solanum capsicastrum Link. 

Aphis rumicis L. Argentina. 
Solanum lycopersicum L. (Tomate, 
tomatera). See Lycopersicon 
esculentum L. 

Solanum melongena (Berengena). 
Aphis gossypii Glover. Brazil, 
Trinidad Island. 
Solanum nodiftorum Jacq. 

Aphis rumicis Linn. Argentina, 
solanophilus Blanchard. 
Solanum sisymbriifolinm Lamb. 
Aphis gossypii Glover. Argentina, 
rumicis Linn. Argentina. 
Solanum tuberosum L. (Patata, papa). 
Aphis gossypii Glover. Peru, 
rumicis Linn. Argentina. 
Aulacorthum pseudosolani 

(Theobald). Chile. 
Macrosiphum cordobensis 

Blanchard. Argentina, Peru, 
solanifolii ( Ashmead ) . 
Argentina. Chile, Peru. 
Myzus persicae (Sulzer). Argentina, 

Bolivia, Peru. 
Smynthurodes betae (Westwood). 
[Trifidaphis phaseoli (Passe- 
rini)] Argentina. 

Solidago sp. 

Brachycaudus helichrysi (Kalt.). 
Sowlius spp. 

Amphorophora lactucae (Kalt). 

sonchi (Oestlund). Argentina, 
Brazil, Chile. 
Macrosiphum lizerianum Blanchard. 

rosae (Linn.). Chile, Peru, 
solanifolii (Ashmead). Chile. 
Sonchus olcraceus Linn. 

Amphorophora sonchi (Oestlund). 
Sorgo. Vease Sorghum halepense (L.). 
Sorghum halepense (L.) (Sorgo). 
Rhopalosiphum maidis (Fitch). 

Brazil, Chile, Peru. 
Sipha flava Forbes. Argentina. 
Sinnacio oleracea L. 

Brevicoryne brassicae (L.) 

Macrosiphum solanifolii (Ashmead). 

Vol. XXVIII 1 



Spiraea chamaedrifoHa L. (Cornoa de 
Aphis fabae Scopoli. Argentina, 
medicaginis Koch. Argentina, 
pseudopomi Blanchard. 
Straussia sp. 

Toxoptera aurantii (B. d. Fonsc). 
Sunchillo. Vease Wedelia glauca (Ort.). 
Sugarcane. See Saccharuvi officwarum 

Sorglium. See Holcus halepensis (L.). 
Suncho. Vease Baccharis salicifolia 

Tanaceto. Vease Tanacetum vulgare L. 
Tabaco. Vease Nicotiana sp. 
Tanacetum vulgare L. 

Macrosiphum bonariensis 
Blanchard. Argentina. 
cocoensis Blanchard. Argentina. 
Taraxacum vulgare (Lam.), (diente 
de leon). 
Macrosiphum macaloi Blanchard. 
Tare. Vease Colocasia esculenUivi L. 
Tasi. Vease Araujia sericifera Bert. 
Tasi fragante. Vease Morrenia odorata 

(Hook. & Arn.). 
Theobroma sp. (Cacao). 

Toxoptera aurantii (B. d. Fonsc.) 
Trinidad Island. 
Thuja occidentalis L. 

Cupresobium juniperi (DeGeer) 
(Cinara). Argentina. 
Thuja tetragona Hooker. Vease Phil- 
gerodendron uviferum (Don) 
Tomate. Vease Solan u))i lycopersicuni 

Tomatera. Vease Solatium lycopersi- 

cuni L. 
Tragopogon porrifoUus L. 

Anuraphis tragopogonis (Kalt.). 
Trigo. Vease Trificum vulgare Vill. 
Triticuni vulgare Vill. 

Pemphigus populi-transversus Riley. 

Rhopalosiphum prunifolae (Fitch). 
(R. pseudoavenae Patch). 
Schizaphis graminm (Rondani). 
(Toxoptera). Argentina. 

Tulipa sp. (Tulipan). 

Aphis rumicis L. Argentina. 
Aulacorthum eumorphum Blanchard. 

Brachycaudus tulipae (B. d. Fonsc). 

Macrosiphum solanifolii (Ashmead). 

Myzus persicae (Sulzer). Argentina, 
circumflexus Buckton. Argentina. 
Tulipan Vease Tulipa sp. 
Tuya. Vease Thuja. 
Typha sp. (espadana). 

Hyalopterus arundinis ( Fab. ) . 

Chile, Cosmopolitan. 
Rhopalosiphum maidis (Fitch). 
Ulmus sp. (01 mo). 

Eriosoma lanigerum (Hausmann). 

Aulacorthum pseudosolani (Theo- 
bald). Chile. (Myzus.) 
Urfica sp. (Ortiga). 

Brachycaudus helichrysi (Kalt.). 

Argentina. (Aphis.) 
Macrosiphum edrossi, n. sp. Peru, 
solutum Blanchard. Argentina. 
Urtica urens L. (Ortiga). 

Myzus persicae (Sulzer). Argentina. 
Yeriiovia sp. 

Aphis gossypii Glover. Argentina. 
Macrosiphum griersoni Blanchard. 
Viburnum, tinus L. (Laurel tino). 
Toxoptera aurantii (B. d. Fonsc). 
Vicia faba L. (Haba). 

Aphis fabae Scopoli. Argentina, 
medicaginis Koch. Argentina, Peru. 
Macrosiphum solanifolii (Ashmead). 
Vid. Vease Vitis vinifera L. 
Vigna luteola (Jacq.). 

Picturaphis vignaphilus Blanchard. 
Vinca major L. 

Aulacorthum eimiorphum Blanchard. 

Myzus circumflexus Buckton. 
persicae (Sulzer). Argentina. 



Viola odorata L. (Violeta). 

Idiopterus violae (Pergande). 
Violeta. Vease Viola odorata L. 
Vitis vinifera L. (Uva). 

Aphis ampelophila Blanehard. 

gossypii Glover. Argentina, 
illinoisensis Shinier. Argentina. 
Viteus vitifolii (Fitch). (Phyllox- 
era). Argentina, Brazil, Peru. 
Walnut, English. See Juglans regia L. 
Watsonia sp. 

Macrosiphum solanif olii ( Ashmead ) . 
Wedelia ghiiica (Ort.). 

Macrosiphum lizerianum Blanehard. 

Xanthiiim sp. (Abi'ojo). 

Anuraphis xanthii Del Guercio. 

Yerba mate. Vease Ilex paragitayensi.s 

Zahnia. Vease Holcus halepensis (L.). 
Zanahoria. Vease Daucus carota L. 
Zapallo. Vease Cucurbita sp. 
Zea mays L. (Maiz). 

Aphis gossypii Glover. Argentina. 
Rhopalosiphum maidis (Fitch). 
(Aphis). Argentina, Brazil, 
Peru, Trinidad Island, 
prunifolae (Fitch). [R. pseudo- 
avenae (Patch)]. Argentina, 



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1938. The cabbage aphis {Brericoryne brassicw L.). Journal Ministry of Agri- 
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Fourth Series 
Vol. XXVIII, No. 4, pp. 165-246; 28 text figs. January 7, 1954 






Curator of Ornithology and Mammalogy 

California Academy of Sciences 

Introduction and Acknowledgments 

So little is known regarding tlie natural histor}- of many of our com- 
mon species of North American bats that it is frequently difficult or 
impossible to provide satisfactory answers to seemingly simple questions 
on the subject. These voids in our knowledge do not reflect a lack of 
scientific interest in bats but rather have resulted from the technical 
difficulties involved in studying these animals. 

Almost every above-ground movement of many kinds of diurnal mam- 
mals can 1)6 noted by careful observers. The use of live traps and of 
marking has provided us with considerable information on the activi- 
ties of many nocturnal, terrestrial species. Who, however, can accur- 
ately state how far a bat flies in the evening after it emerges from its 
daytime retreat? So far as the observer is concerned the animal's iden- 
tity is usually lost in the gloom of twilight within a matter of seconds. 
The distance that it travels in search of food, whether a few hundred 

[165 1 


yards or some miles, must largely remain a matter for conjecture until 
more advanced techniques for study are developed. 

Vaguely aware of some of the difficulties involved in a project con- 
cerning chiropteran life history, the writer began such an undertaking 
in May, 1947. The pallid bat, Antrozous pallidus (LeConte), was the 
species selected for this purpose for several reasons. It is relatively 
common in central California. It can easily be kept in captivity. It is 
sufficiently large and distinctive in appearance to be readily recogniz- 
able in the field, even at night in the beam of a flashlight. The results 
obtained to date are presented in this paper. These include field obser- 
vations, information derived from studies of bats in captivity, and rele- 
vant data extracted from the literature. It is felt that there are still 
many voids in our knowledge of events concerning the natural history 
of this one species but it is to be hoped that future studies may solve 
some of these problems. 

Many persons contributed to this project. I wish to thank Dr. Robert 
C. Miller, director of the California Academy of Sciences, for the funds 
and facilities provided by that institution which made this undertaking 
possible. I am particularly indebted to ]\Iiss Mary Louise Perry for her 
painstaking care of captive animals during a period of over four years 
and for accurately recording and interpreting data obtained from labo- 
ratory studies. I am also especially grateful to iVIr. Eben McMillan for 
his enthusiastic cooperation in locating bat colonies in San Luis Obispo 
and Kern counties, California, and for his very active participation in 
much of the field work. For the identification of many ectoparasites 
and of insects used as food by bats I wish to thank Dr. Edward W. 
Baker, United States Department of Agriculture, Washington, D. C. ; 
Professor G. F. Ferris, Stanford University; Mr. Robert Iloldenried, 
United States Public Health Service, Santa Fe, New Mexico; Dr. Glen 
M. Kohls, United States Public Health Service, Hamilton, Montana; 
Mr. Frank Prince, United States Public Health Service, San Francisco, 
California; Dr. Edward S. Ross, Dr. Edward L. Kessel, and the late Dr. 
Edwin C. Van Dyke of the Department of Entomology, California Academy 
of Sciences. For permission to borrow or study specimens under their 
care I wish to thank Dr. Seth B. Benson, Museum of Vertebrate Zoology, 
University of California; Dr. William IT. Burt, Museum of Zoology, 
University of Michigan; Dr. David H. Johnson, LTnited States National 
Museum; and Dr. Colin C. Sanborn, Chicago Natural History Museum. 
Others who contril)u+ed to this study include Mr. Lionel Berryhill, Mr. 
Sterling Bunnel, IM's' L.^ura Grainger, ]\Irs. eT. Gordon Irving, Mrs. 
Berta B. Kessel, Mr. Tlu-^h B. Leech, ]\Ir. I.-^n ^McMillnn. Mrs. Dorothy 
B. Orr, Mr. Frank L. Rogei-s, Dr. Charles G. Sibley, Mr. Leon E. Salanave, 
and Mr. M. Woodbridge Williams. 



A number of summering colonies of pallid bats were known to the 
writer in or fairly close to the San Francisco Bay region. Most of the 
field work, however, was concentrated in eastern San Luis Obispo and 
western Kern counties because conditions in that region seemed highly 
favorable for bats of this species, judging from their abundance, and 
because the environmental changes resulting from human usage had 
been less there than in many parts of California. 

Thirty-one days, devoted primarily to obtaining data relevant to this 
species, were spent in the field as follows: 

May 9-11, 1947, San Luis Obispo and Kern counties. 

August 9-10, 1947, Humboldt County (M. L. Perry). 

September 19-22, 1947, San Luis Obispo and Kern counties. 

February 27-March 1, 1948, Monterey, San Luis Obispo, and 
Kern counties. 

June 5-9, 1948, San Luis Obispo and Kern counties. 

April 1-4, 1949, Monterey, San Luis Obispo, and K'ern counties. 

October 21-24, 1949, Monterey, San Luis Obispo, and Kern 

January 22, 1950, Marin County. 

April 13-16, 1951, San Luis Obispo and Kern counties. 

June 13, 1952, Santa Clara County. 
Although all of the field observations made on Antrozous pallidus 
were within the range of the race pacificus, as given by Grinnell (1933), 
occasional references are made to published reports concerning the natu- 
ral history of other geographic forms of this species. 

Pallid bats were maintained in captivity during the entire period 
of this study. IMost of the wild-taken individuals were released, with 
L^^nited States Fish and Wildlife Service bird bands attached to their fore- 
arms, from 10 to 13 months after capture. Such releases were made either 
at the original place of capture or, for experimental purposes in an attempt 
to study homing instinct, at some other locality within the same gen- 
eral region in which field studies were made. 

As a rule bats were captured at their daytime retreats, usualh^ with 
the aid of insect nets and long-handled forceps. They were transported 
to the laboratory in cages 16 inches wide, 16 inches high, and 12 inches 
deep, made of plywood and quarter-inch hardware cloth, with sliding 
doors at the back. At the California Academy of Sciences bats were 
housed in several sizes of cages for various purposes. Small retaining 
cages of the type just mentioned were used for the confinement of fe- 
males with newlv born voung to facilitate studies on behavior and 


growth, for isolating certain individuals, and in enforced dormancy ex- 
periments. A larger cage, 48 inches wide, 38 inches high, and 42 inches 
deep, made of quarter-inch hardware cloth on a wooden frame, was 
used for small colonies kept in tlie laljoratory. This was sufficiently 
large to permit limited tiight. Two large flight cages, 4 feet wide, 8 
feet high, and 10 feet deep, that had previously been used as aviaries 
(for detailed descrij^tion see Orr, 1945, p. 179) on the roof of the 
Academy's North American Hall, served to maintain small colonies of 
bats out-of-doors. 

The smaller cages were kept darkened during the day by hanging- 
burlap or paper over the screened side or by placing this side next to 
a wall. In the larger cages bricks, boards, or small boxes provided day- 
time retreats for the bats. Each occupied cage was provided with 
food and water daily, except on week-ends. The food consisted prin- 
cipally of meal worms (the larvae of the flour beetle, Tenebrio molitor) 
which were killed by momentarily scalding them before they were placed 
in the feeding dishes in the late afternoon. Since, on occasion, as many 
as 10,000 meal worms were used weekly, it was not found practical to 
raise them. Tliey were secured from the Sure-Bite Live Bait Company 
in Torrance, California. 

Toward the end of this project it was found more practical, eco- 
nomical, and satisfactory fiom the standpoint of maintaining healthy 
captive animals, to use a food mixture recommended by Mr. Ernest 
P. "Walker, assistant director of the National Zoological Park, Washing- 
ton, D. C. This consisted of equal parts of hard-boiled egg yolk, cot- 
tage cheese, ripe banana, and meal worms, with small amounts of Jecu- 
lin, wheat germ oil, and vitamin mixture added. The ingredients were 
ground into a paste. It was found expedient to prepare a fairly large 
quantity of this food at one time, dividing it into small portions which 
were wrapped in wax paper and kept frozen until needed. 


The pallid bat is readily distinguished, on the basis of external char- 
acters, from all other North American bats by its combination of large 
size, proportionately large ears and broad wings, peculiar shape of )iose, 
and color of pelage. 

Of the 24 species of bats recorded as occurring in California (Grin- 
nell, 1933; Constantine, 1946; Olson, 1947) only 4 approximately equal 
or exceed the pallid bat in general body size. These are the hoary bat 
(Lasiurus cinereus (Beauvois)), the western yellow bat [Dasypterus ega 
(Gervais)), the pocketed bat {Tadarida femorosacca (Merriam) ), and 
the mastiff bat (MoJossus perotis Schinz). The hoary bat is not often 




Figure 1. The ears and eyes of the pallid bat are proportionately large. Note 
the serrated outer edge of the tragus. Photographed at the California Academy of 
Sciences, November 12, 1952. 

found in the same region as the pallid bat, except in spring and autumn 
when the former species is migrating. Only once during this study 
were these two species noted together (Orr, 1950). The western yel- 
low l)at has been recorded only once from California (Constantine, 
1946). The pocketed bat and the mastiff bat are both members of the 
family Molossidae. The former species is very rare in California, known 
only from the extreme southern part of the state. The mastiff bat is 
more widely distributed but of rather local occurrence and cannot be 
considered common. Ftirthermore, it is readily distinguished from the 
pallid bat by its greater size, having an average wingsi)read in excess 
of 500 millimeters. 

The ears of the pallid bat are separate and large, altliough not as 
long, proportionately, as in members of the genera Covynovhinus and 
Euderma. In shape the pinna is roughly rhomboidal, obliquely attached 
at the base and rounded at the tip. When laid forward it extends con- 
siderably beyond the nose. Along the postero-lateral half of the ear 
there are usually 9 to 11 horizontal creases which permit the pinna to 
be folded back. The tragus is slender, tapering distally, rounded at the 
tip, and serrate along the outer edge (fig. 1). The ears are pale gray- 
ish tan in color. A narrow strip of hairs is present on the antero-dorsal rim 



[Proc. 4th Seb. 

of the pinna, extending from the base halfway to the tip. Two narrow 
bands of hairs, extending parallel to the long axis of the ear, are pres- 
ent inside the pinna. 

The wings are proportionately broad with the third metacarpal only 
slightly longer than the fifth. Only the extreme tip of the tail extends 
beyond the uropatagial membrane. The flight membranes, in general, 
are essentially naked except those parts immediately adjacent to the 
body. In color they are a dark slate gray with a slight vinaceous tinge. 
The calcar terminates in a small but distinct lobe jnst short of the 
middle of the free edge of the nropatagial membrane. The feet are 
proportionately large and broad, with a few sparse hairs on the backs 
of the toes. 

The end of the muzzle of the pallid bat is decidedly truncate with 
the nostrils opening forward. The rhinarium is scroll-shaped and ele- 
vated into a slight ridge above the nares. Behind the rhinarium, on either 
side of the muzzle, is a large, flattened, glandular swelling (fig. 2), The 
eyes are relatively large for vespertilionid bats. 

Table I 

Average and extreme measurements, in millimeters, of adult specimens of 
Antrozous j)allid2is paciflcus from eastern San Luis Obispo and western Kern 
counties, California. 




Ear from 































Number avei 






























Number averaged 







Pelage: The fur of the pallid bat is not dense and is of moderate 
length and medium texture. The hairs on the dorsal surface of the body 
are, for the most part, longer and more widely separated from one an- 
other than are those on the ventral surface. The area between the 



Figure 2. Large, glandular swellings are situated on either side of the muzzle 
of the pallid bat, immediately behind the scroll-shaped rhinarium. Photographed 
at the California Academy of Sciences, December 10, 1952. 

shoulders is very scantily haired. Some of the longest hairs from the 
back of an average adult specimen in fresh pelage measured 8 milli- 
meters from tip to base when taut. Owing to a slight natural kinkiness, 
however, the actual distance irom tip to base rarely exceeds 5 or 6 milli- 
meters for the dorsal hairs and 3 to 4 millimeters for hairs on the ven- 
tral parts of the l)ody. 

The hairs on the dorsal areas of the head and body are bicolored. In 
specimens of Antrozous ijaUidus pacificus Merriam from San Luis Obispo 
County, in fresh pelage, the distal third of these haii's is nearest Ridg- 
way's (1912) Olive-Brown and the proximal two-thirds is nearest Cart- 
ridge Buff. The hairs on the ventral surface of the head and body are 
generally unicolored and, in fresh pelage, vary from a creamy white 
to a very pale grayish white. Occasionally some of the tips of the ven- 
tral hairs have a fuscous tinge, especially those on the posterior part 
of the body. In w^orn pelage the distal tliird of the hairs of the upper 


parts tends to become lighter, approaching Saccardo's Umber while the 
proximal two-thirds of these hairs becomes more yellowish than in iresh 
pelage. Likewise, the hairs on the ventral parts become more buffy. A 
certain amount of this seasonal change in pelage color, particularly on 
the underside of the body, appears to be adventitious, possibly because 
of frequent contact with urine and excrement in the roosts. 

One example of albinism is known. Setzer (1950, p. 350) records 
an albinistic specimen of this species in the collection of the United 
States National Museum. It was secured by Stanley G. Jewett on Feb- 
ruary 9, 1940, 26 miles north of Las Vegas, Nevada. 

Molt: There is but one molt annually in this species. This takes place 
during the summer months, the time of occurrence varying consid- 
erably with different individuals in the same colony. Examination of 
museum specimens, captive bats, and those living in the wild failed to 
indicate any sign of molt starting before early ^lay or of not having 
begun l)y the end of August. Most signs of molting were to be observed 
during the months of June and July. Pregnancy does not appear to 
influence the molt. Examination of females in late stages of pregnancy 
on June 6, 1948, in eastern San Luis Obispo County, showed that some 
had not yet begun to molt while others in the same colony were either 
in process of molting or had already assumed new pelage. 

New pelage first makes its appearance on the relatively bare area of 
the back, between the shoulders, and, midventrally, at the base of the 
neck. The new hairs on the intershoulder area radiate from a point on 
the middorsal line while the new hairs on the underside of the neck 
are arranged in the form of a whorl. Simultaneously with the ap- 
pearance of new hair on these two parts of the body the skin of the back 
appears darkly pigmented. Following this, new hair is soon in evidence 
from the crown of the head to the lower part of the back and on the 
ventral part of the head and body. The new hairs grow more rapidly 
on the middorsal surface than elsewhere and appear last of all on the 
front of the head, sides of the neck, and rump. Examinations of study 
skins as well as living individuals have shown that the old fur on a 
particular part of the body falls out when the new fur on that area 
has grown out to about half of its full length. When captive molting 
bats were handled at this stage the old fur came out readily in clumps. 

The time required to complete the molt seemingly is not very long. 
One captive individual was observed in the very early stages of molt 
on August 26, 1949. New fur was just appearing above the surface of 
the skin between the shoulders and on the underside of the neck. When 
this bat was examined 13 days later it was completely in new pelage 
although some of the hairs had not yet quite attained full length. 



Antrozous paUidus belongs to the family Vespertilionidae and the sub- 
family Nyctophilinae. The only other genus belonging to this subfamily, 
according to Miller (1907, p. 234) and Simpson (1945, p. 60), is Nycto- 
pkilus of Australia and the East Indies. The genus Antrozous is re- 
stricted to North America where its range extends from southern British 
Columbia east to Kansas and south to south-central Mexico. It has been 
recorded' in the following states or provinces: southern British Colum- 
bia (Racey, 1933, p. 18), eastern Washington (Dalquest, 1938, p. 213), 
parts of Oregon (Bailey, 1936, pp. 390-392), parts of California (Grin- 
nell, 1933, pp. 93-94), western and southern Nevada (Hall, 1946, p. 164), 
southern and eastern Utah (Durrant, 1952, p. 60), parts of Arizona 
(Miller, 1897, p. 44; Swartli, 1929, p. 347; McKee, 1932, p. 71; Cahalane, 
1939, p. 422), parts of New Mexico (Bailey, 1931, p. 379), southern Colo- 
rado (Warren, 1910, p. 284), southern Kansas (Hibbard, 1934, p. 227), 
northwestern Oklahoma (Burt, 1945, p. 309), western Texas (Bailey, 
1905, p. 214), Baja California (Nelson, 1921, p. 128), Sonora (Burt, 
1938, p. 27), Durango (Allen, 1903, p. 612), Nuevo Leon (Davis, 1944, p. 
380), and Queretaro (Miller, 1897, p. 45). There are previously unre- 
corded specimens from Tamaulijias. Mexico, in the University of Michi- 
gan Museum of Zoology and it is likely that further collecting will dis- 
close the presence of this genus in a number of other states in northern 
and central Mexico as well as in southern Idaho. 

Within the range of the genus there are two currently recognized 
species, Antrozous paUidus and A. bunkeri Hibbard. These iwo forms 
are very closely related and likely will prove to be conspecific. So far 
as known, A. bunkeri has been recorded only from the type locality in 
southern Kansas (Hibl^ard, 1934) and from one locality in northwest- 
ern Oklahoma (Burt, 1945). Antrozous jjaUidus, whose distribution is 
much more extensive, is represented by four currently recognized geo- 
graphic races whose ranges are approximately as follows: paUidus, south- 
eastern California and northeastern Baja California east to Colorado 
and Texas and south to south-central Mexico; cantweUi, southern Brit- 
ish Columbia and eastern Washington south to northeastern California 
and northwestern Nevada; pacificus, northwestern Oregon south to north- 
western Baja California; minor, central to southern Baja California. 

Antrozous paUidus is primarily a species of the Ijower and Upper 
Sonoran life zones. In parts of Oregon and California, however, it 
ranges well up into the Transition Zone, locally, and to the south, in 
parts of jNIexico occurs in the Arid Tropical Zone. It has an altitudinal 

1. The references cited are not necessarily the first published records but, wherever possible, those 
that give the most complete distributional accounts for each state or province concerned. 


range known to extend froia -178 feet in Death Valley, California 
(Grinnell, 1933, p. 94), to at least 6700 feet in the Qninn Canyon Moun- 
tains, Nevada (Hall, 1946, p. 166). These altitudinal extremes are 
both within the range of the i-ace palUclus. In central California, how- 
ever, the writer has records of Antrozoiis p. pacificus from essentially 
sea level at Inverness, Marin County, to 5000 feet at Long Barn, 
Tuolumne County. 


No attempt is made here to give a general description of the habitat 
of the pallid bat since environmental conditions vary so much through- 
out different parts of its rather extensive range. To facilitate a better 
understanding of the requirements of memljers of this species, however, 
some of the situations in which they have been observed are described. 
These include coniferous forests of several different types, nonconiferous 
woodland, brushy terrain, rocky canyon, open farm land, and desert. 
Suitable daytime I'etreats or roosts, of course, are essential for the oc- 
currence of these bats in any region. It is possible that the presence 
of available fresh water nearby also is necessary. 

Fairly conclusive evidence was obtained indicating that pallid bats 
occur in Richardson (irove State Park, numl)oldt County, California. 
This grove of redwoods (Sequoia sempervirens) is close to the Eel River 
and there is open country nearby where bats of this species might 
forage at night. A torpid pallid bat was found at an abandoned cabin 
in a dense redwood forest in ]\Iill Valley, Marin County, California, 
on January 15, 1950. 

At various times in 1948, 1949, and 1950 Mr. :\I. Woodbridge Wil- 
liams noted pallid bats in or around his home in Inverness, Marin 
County. An individual was captured here on November 7, 1948, and 
another in August, 1950. Much of the forest growth in the vicinity 
of the Williams' home is native, consisting principally of Bishop pine 
(Pinus viuricata), coast live oak (Quercus agrifoJia), tan oak (Litho- 
carpus densiflora), and madrone (Arhutus menziesii) . 

On the evening of May 29, 1939, the writer captured a pallid bat 
at Long Barn, 5000 feet, Tuolumne County, California. This indi- 
vidual flew in through an open window of a cabin situated in a forest 
composed principally of yellow pine (Pinus ponderosa), white fir (Ahies 
concolor), and incense cedar (Libocedrus decurrens). Bailey (1936, p. 
390) records pallid bats associated with yellow pine in southwestern 

In San Luis Obispo and Kern counties, California, summering colo- 
nies of these bats were found either in buildings or in crevices in rock. 



One group of ranch bnildiniis inlial)ited hy them was situated near the 
mouth of a small canyon which opened out onto a fiat that was used 
for raising grain. The vegetation behind the buildings consisted largely 
of grasses with clumps of Eriogomim fasciculatum and Artemisia cali- 
fornica scattered about. There were some large cottonwood, pepper, 
and fig trees growing near the buildings. Permanent water, provided by 
a spring, was available close by. Mr. Eben McMillan found another 
colony of ]iallid bats living in the attic of a garage in the town of 
Shandon. There were a good many cultivated trees nearby and open 
grassland within a quarter of a mile. 

Figure 3. Rui'ky outcrops provide numerous roosting .-iles in the form of 
crevices for pallid bats. Photographed near Carneros Spring, Kern County. Cali- 
fornia. May 20, 1948, by G Dallas Hanna. 

Five colonies were found in natural rock crevices. Two of these 
colonies were located within several hundred yards of each other in 
a large outcrop of sandstone at the northwestern edge of the Carrizo 
Plain, San Luis Obispo County. Some of the rocks were 50 to 60 feet 
in height. Inuncdiately surrounding the outcrop was grassland inter- 
mingled with a good many trees and shrubs including blue oak (Quer- 
cus doughisii), islay (Prnnus ilicifolia), great-berried manzanita (Arcto- 



[Proc. 4th Ser. 

staphijlos glauca), black sage (Salvia meUifera), California sagebrush 
(Artemisia calif ornica), and Aplopappus cuneatus. During the dry sea- 
son the nearest available water was about one-half mile distant. An- 
other colony was found in a sandstone outcrop 21/2 miles northwest of 
Carneros Spring, Kern County. The surrounding terrain was extremely 
arid (fig. 3). A few low-growing shrubs, principally California sage- 
brush and Eriogonum sp., were present near the base of \he rocks. 
\Vhether or not there was any water available to bats nearer than Car- 
neros Spring was not determined. Two other colonies were found in- 
habiting rocky escarpments bordering a small valley through which 
San Juan Creek flows, 3 miles southeast of La Panza, San Luis 01)isiio 
County (fig. 4). The valley was no more than one-quarter of a mile 
in width. A few oaks were scattered about in the grassland and an occa- 
sional willow grew along tlie streamside. 


Figure 4. Favorable habitat for pallid bats on the La Panza Ranch along San 
Juan Creek, San Luis Obispo County, California. Photographed June 9, 194S. 

A number of other chiropteran species were observed in the general 
area inhabited by the pallid bats in San Luis Obispo and Kern coun- 
ties. These included Myotis yumaneyisis, .1/. thymnodes, 2L. voJans, M. 
californicus, M. suhuhitus, Pipistrellus hespcrus, Eptesicus fuscus, Cory- 
norhinus rafinesquei, Tadarida mexicana, and Molossus perotis. 


Hall (1946, p. 163) comiiu'iits that in the Carson Basin, Churchill 
County, Nevada, the pallid bat "frequents the edges of the basin, forag- 
ing there as well as roosting in the caves along the outcrops of rock at 
the foot of the hills." Burt (1934, p. 397) records this species as a 
common inhabitant of the lower desert regions of southern Nevada, 
especially around ranch houses where water and vegetation are present. 


During most of the spring, summer, and autumn months pallid bats 
are found during the daylight hours in groups or colonies generally 
consisting of no more than one hundred individuals and frequently 
composed of a considerably smaller number. The cavities or crevices 
which provide daytime roosts for these colonies may be in rocks, trees, 
or various man-made structures as has already been indicated. Irre- 
spective of their diversity in location, all of the roosting places ex- 
amined appeared to have certain features in common. All provided 
semidarkness, and jn-otection from above. The ventral surface of the 
body of a roosting bat was always against a solid surface which was 
either a part of the roost or the body of another bat. Koosting pallid 
bats were never observed hanging freely from a horizontal surface, 
such as the roof of a cave, in the daytime. Some other species of bats 
regularly hang in this latter manner. The opening from the roosting- 
crevice to the outside was, with one exception, at least several feet, 
usually much more, al)ove the substratum, thus permitting the bats 
both to take flight immediately after emerging and to fly directly into 
the opening when returning. The opening was generally beneath the 
roost although when the roost was in a building it was frequently neces- 
sary for the entrance to be on a horizontal plane. 

In the event that ])allid l)ats are disturbed at their roost they fre- 
quently will leave and go to an alternative roost which is usually lo- 
cated nearby. The permanence of such moves is not known. In at least 
two instances roosts that were so vacated, as a result of disturbance, were 
found unoccupied during the succeeding several years. There is some 
indication that the shifting of a colony from one site to another is 
sometimes done without apparent provocation. On several occasions 
roosts were found to be inhal)ited in the spring and were left undis- 
turbed by the observer. AVhen examined several months later they 
were unoccuiiied. Although it is possible that these colonies were dis- 
turbed by someone in the meantime, their locations were such as to 
make this exceedingly improbable. Ryberg (1947, pp. 74—77) comments 
on similar alternative roosts used by certain species of bats in north- 
ern Europe, and Pearson, Koford, and Pearson (1952, p. 276) com- 


ment on the alternative roosts of summering colonies of Corynorhinus. 

Since pallid hats are not primarily forest dwellers, roosts in cavi- 
ties in trees are of relatively uncommon occurrence. On August 9 and 
10, 1947, Mary Louise Perry (MS) visited Richardson Grove State 
Park, Humholdt County, California, in search of pallid hats reportedly 
inhabiting hollow redwood trees in the grove. A number of hollow 
trees were examined and two were found to have been inhabited by 
bats although no bats were observed at this time. In the base of one 
tree tlie guano indicated that the inhabitants were prol)ably all Myotis. 
About 98 per cent of the droppings in the other appeared to be those 
of Myotis but the remainder were large, about the size of those of 
Antrozous. According to Mr. Earl P. Hanson, assistant district super- 
intendent at Humboldt Redwood State Park, Antrozous palJidus and 
Myotis califoniicus were the only species of bats that had, up to that 
time, been identified from Richardson Crove. Bailey (1936, p. 390) 
records a colony found in a hollow yellow pine in Jackson County, 
Oregon. The same author (1931, pp. 379-380) mentions another in- 
stance in which pallid bats that were disturbed in a cave in the day- 
time were noted seeking refuge in a hollow tree in the Cloverdale 
Hills in southwestern New Mexico. Hall (1946, p. 163) records a small 
grou]) of these l)ats found roosting in a hole in a cottonwood tree at Ash 
Meadows, Nevada. Davis (1944, p. 380) found a colony of pallid bats 
using a cavity in a dead cypress as a daytime retreat in Nuevo Leon, 

While cavities in trees occasionally provide natural daytime re- 
treats, crevices in rocks are most frequently used for this purpose. 
Several such daytime retreats were observed in San Luis Obispo and 
Kern counties. One of these was found on May 10, 1947, beneatli a 
partly loosened slab of rock on a per]iendicular cliff of an is-olated 
sandstone outcrop "Ji/o miles northwest of Carneros Spring, Kern County 
(fig. 5). The entrance to the crevice was 7 feet above the ground and 
measured 2 feet in length and 4 inches in width. The crevice extended 
upward between the slab and the cliff for about 3 feet. The bats 
were crowded into the upper part of the cavity. The cliff below the 
opening was stained with urine and excrement and the ground be- 
neath was covered with guano. Disturbance caused the bats to leave 
and fly around to another side of the outcrop where they were thought 
to have taken refuge in another crevice about 75 feet pbove the 
ground. The distance between these two crevices was about 100 yards. 
The roost which the bats had left was found unoccupied on Se]itember 
20. 1947. although uumerou'^ dronpings in the vicinity indicated that 
a colony of pallid bats still inhabited tliis isolated outcrop. 

On .Tune 7, 1948, two more colonies were located benenth loose slabs 







Figure 5. The crevice beneath the loose slab of sandstone in the center of the 
picture served as a day roost for a colony of pallid bats. Two and a half miles 
northwest of Carneros Spring, Kern County, California, September 20, 1947. 


of sandstone at a rocky outcrop on the northwestern edge of the Car- 
rizo Plain in San Luis Obispo County. One of the colonies occupied a 
daytime roost very similar to that just described. In this instance the 
entrance to the crevice was 12 feet above the ground. The colony was 
not disturbed on this date. On October 23, 1949, pallid bats were 
again found in this crevice and in a similar crevice 30 feet away. The 
entrance to the latter was only 6 feet above the ground. These two 
roosts were considered as belonging to the same colony. The second 
colony found on June 7, 1948, was about 200 yards from the one 
just described. The retreat occupied by these bats was also behind a 
partly loosened slab of sandstone on a cliff but it was situated about 
20 feet above the ground and just above a narrow ledge of rock that 
was sufficiently wide for a person to walk on. The main entrance to 
the roost was a narrow, vertical opening, about 12 inches in length 
and 2 inches in width, at one side of the slab. The bottom of the 
opening was 12 inches above the ledge. Along the lower edge of the 
slab, where it contacted the ledge, there were two holes about a foot 
apart. Each hole measured approximately 3 inches in diameter. Ac- 
cumulations of excrement that had rolled out through the holes were 
present on the ledge. All of the bats in this colony were crowded 
tightly together at the upper end of the crevice which was about 18 
inches above the ledge. This was the only roost observed where the 
bats were unable to drop immediately after emerging. The ledge pre- 
vented them from so doing. Approximately two-thirds of the occu- 
pants were captured as they attempted to come out. The rest escaped. 
Those that escaped appeared to have no difficulty in taking flight im- 
mediately upon coming out of either the vertical slit or the two holes. 
The captured bats were released a short time later, after their sex had 
been recorded and the females palpated to determine pregnancy. The 
majority of those that either escaped or were released flew to a crevice 
in the roof of a shallow cave about 50 yards away. This alternative 
roost was known to have housed a colony of pallid bats the previous 
summer. When these roosts w^re visited on October 23, 1949, both were 
found uninhabited. It was difficult to tell whether the alternative roost 
in the roof of the cave had been used in the daytime during the sum- 
mer of 1949. There were large accumulations of guano belonging to 
both pallid bats and Mexican free-tailed bats but both species were 
known to use the roof of this cave regularly as a night roost during 
the summer. The roost above the ledge showed no sign of having been 
occupied in 1949. When it was visited again on April 14, 1951, it was 
found to be occupied by a wood rat (Neotoma lepida). This suggests 
the possibility that wood rats occasionally compete with these bats 
for home sites. 


On June 8, 1948, two other colonies of pallid bats were found on 
the La Panza Ranch along San Juan Creek, San Luis Obispo County, about 
9 miles west of Simmler. One was located in a crevice beneath a loose slab 
of rock on the face of a nearly vertical cliff (fig. 6). The entrance to 
the crevice was 10 feet above the ground and consisted of an open- 
ing 3 feet in length and about 4 inches in width. The crevice extended 
upward approximately 3 feet and the bats were in the uppermost part. 
When some of these bats were captured the following day the other 
members of the colony left. All those seen leaving flew down the can- 
yon through which the creek flowed and were out of sight as soon as 
they rounded a bend several hundred feet away. One gained the im- 
pression that they were moving to an alternative roost known to 
them although this was not found by the observers. The roost de- 
serted at this time was found occupied in the summer of 1949 and 
again in the spring of 1951. It was not examined in 1950. 

The second colony found on June 8, 1948, on the La Panza Ranch 
was about half a mile farther up the canyon in a small cave. The 
cave, situated at the base of a sandstone cliff, was dome-shaped and 
about 10 feet in diameter. The distance from the floor of the cave 
to the highest point was also about 10 feet. On one side of the dome 
there was a funnel-shaped recess which led into a short, narrow crevice 
in which the bats were observed roosting. There were two openings 
into the cave from the outside. One of these was irregular in shape. 
It extended from the ground up to a height of 31/2 feet and was about 
21/2 feet in width. Above this, about 5 feet from the ground, there 
was another opening 1 foot wide and 2 feet in height. The lower open- 
ing, judging from the presence of many droppings, was the one more 
frequently used by the bats when entering or leaving. In the center 
of the floor of the cave there was a broad, cone-shaped pile of excre- 
ment 3 to 4 feet in diameter and attaining a maximum depth of 12 
inches. The presence of observers close to this cave during the night 
caused the colony to move to another roost by dawn the following 
morning. It w^as visited in the summer of 1949 and again in the 
spring of 1951 and found unoccupied in the daytime on each occa- 
sion although fresh accumulations of guano in each instance showed 
that the dome of the cave still served as a night roost. 

There is a possibility that these bats might roost beneath rock slides 
or in talus slopes in certain localities although no such colonies have 
been located. Racey (1933, p. 18), however, records the discovery of 
a single bat of this species beneath a pile of stones in British Columbia. 

Pallid bats frequently flnd suitable roosts in attics, between walls, 
between bridge timbers, and in other similar hiding places in man- 
made structures. One such colony that was periodically under obser- 



[Proc. 4th Sek. 


Figure (J. The creviee on the sandstone eliff in the center of tlie picture housed 
a colony of pallid bats. La Panza Ranch, San Juan Creek, San Luis Obispo 
County, California, June 9, 1948. 



vat ion during tliis study inhabited a barn (fig. 7) and a shed nnmedi- 
ately adjacent to the bam on a ranch 4i/o miles northeast of Shandon, 
San Luis Obispo County. Sometimes the entire colony was in one or 
the other of the buildings and sometimes it was divided, with bats in 
both buildings. In each structure the bats occupied small spaces at 
either end between the timbers that supported the roof. Since the 
crevices were small and could not hold many bats, the colony was 
divided into a numl)er of groups. Disturbance never caused members 

Figure 7. Pallid l)ats inlnihitiut; ihis ham, 4io miles northeast of Shandon, 
San Luis Obispo County, California, were observed entering and leaving through 
the opening seen immediately below the peak of the roof. Photographed Octo- 
ber 22, 1949. 

of this colony to desert the liuildings although they might fly from 
one building to the other or move to the opposite end of the I'oof of 
the same building. 

On June 7, 1951, Sterling Bunnell examined a colony of pallid bats 
living in the attic of Encina Hall at Stanford University in Palo Alto. 
Some adult females and young were captured. The writer examined 
the attic of this building a few days later on June 13. The attic was 
found to be divided into a number of sections, each about 40 feet long 
and equally wide. Brick walls served as partitions between adjoining 
sections. Beams 2 inches by 12 inches rested on the brick partitions 
and su]:)ported the roof. No bats were found in the section where they 
had been roosting on June 7 although there were large accumula- 


tions of droppings at the bases of the hriek ))artitions at each end. Two 
partitions away, however, about 40 pallid bats were found roosting 
in spaces between the top of one of the brick partitions and the roof. 
It seemed likely that these were the same bats that had been living in 
the other part of the attic on June 7. 

Another colony discovered by Eben McMillan {fide letter of April 24, 
1949) in western San Luis Obispo County, between Paso Eobles and 
Cambria, was located in a space between the window panes and some 
boards that had been nailed across the windows of an old house. 

There are numerous records in the literature of pallid bats using 
man-made structures, a few of which are cited here. Krutzsch (1946, p. 
241) records bats of this species living in a barn, in the attic of a 
church, and in cracks betw^een the beams of a bridge. Hall (1946, p. 163) 
describes a group of these bats found near Lahontan Dam, Nevada, that 
had "ensconced themselves in the crevices, 1/2 to 1 inch in width, be- 
tween the 18-inch timbers on the underside of a bridge." Storer (1931, 
p. 244) records a summering colony of pallid bats found living in the 
walls of a residence in Berkeley, California. Bailey (1905, p. 214) men- 
tions pallid bats roosting behind a signboard at Comstock, Texas. 

At night pallid bats frequently alight, at what are generally re- 
ferred to as night roosts, to rest or to consume food that has just been 
captured. These so-called night roosts are much more accessible to the 
bats than tlie crowded recesses that constitute daytime retreats. Bats 
living in barns or attics usually use the ridgepole of the roof for such 
purposes. In rocky areas the roofs of shallow caves frequently pro- 
vide night roosts (fig. 8). Large accumulations of guano and parts of 
insects that have been discarded are present beneath such roosts (fig. 
9). Early in April, 1949, the floor of one such roost in a small cave 
on the La Panza Ranch was scraped clean. When examined six months 
later on Octolier 23 a new accumulation of guano w^as found that meas- 
ured 4 feet in diameter and 7 inches in depth in the center. 

The night roosts of all the colonies whose daytime retreats were 
known were usually within a few yards of these daytime retreats. In 
one instance the night roost of a colony was not located although no 
great effort was made to find it in the numerous rocky recesses nearby. 
Several night roosts regularly used by pallid bats were found although 
diligent search failed to reveal the whereabouts of the day roosts. In 
each instance, however, possible hiding places nearby were inacces- 
sible to the observer. 

The night roosts of these bats in buildings are frequently a source 
of considerable annoyance to the owners. Bats belonging to one colony 
under observation used, for this purpose, either the ridgepole of a barn 
or the ridgepole of an adjacent shed that served as a garage. As a re- 



Figure 8. The roof of this shallow cavern near San Juan Creek on the La Panza 
Ranch, San Luis Obispo County, California, served as a night roost for pallid 
bats and Mexican free-tailed bats. Photographed June 8, 1948. 



Figure 9. Accumulation of guano beneath the night roost shown in figure 8. 
Photographed October 23, 1949. 

suit, the bat excrement fouled hay that was stored in the barn and 
made it necessary to hang a cloth covering over the car in the garage 
to keep it clean. Members of another colony were observed using the 
beams supporting the roof of an open porch of a ranch house for a 
night roost. The house was less than one year old when examined on 
the evening of April 13, 1951, and the two-by-four beams of the porch roof 
had been freshly creosoted. This apparently was no deterrent as 6 
bats were seen hanging from a beam at one end of the ]iorch and 3 
from a beam at the opposite end at 9 :55 p.m. Fresh droppings and 
insect remains were found on the concrete beneath each small group. 


The fact that an electric light had been burning on the porch all eve- 
ning did not seem to discourage the bats from coming here any more 
than did the creosote. 

Seasonal Behavior 

Spring and summer: By late March or early April the period of 
dormancy appears to terminate in central California and summer colony 
formation occurs. Mrs. Grinnell (1918, p. 356) records March 27 as 
the earliest known date of observation of this species in spring m this 
state. In Texas, Bailey (1905, p. 214) indicates April 18 as the earli- 
est seasonal date on which the species was noted. 

In the spring of 1949 an examination of four sandstone crevices 
in eastern San Luis Obispo County, where summering colonies had 
been observed in 1947 and 1948, failed to reveal the jn-esence of any 
pallid bats on April 1, 2, and 3. A very few fresh droj^pings beneath 
two of the colony sites indicated that one or at most a very few bats 
had recently been present. During the ensuing two weeks ]\Ir. Eben 
McMillan periodically visited one of these colony sites and a barn in 
which pallid bats had been found every summer for some years. Bats 
were found in the barn on April 10 and in the sandstone crevice on 
April 14. In each instance they appeared to have arrived not more 
than a day or so previously. No effort was made to determine the num- 
ber of individuals present on the first day bats were found in these 
colonies, but on April 15 it was estimated that about 30 individuals 
were present in the colony first occupied on April 10. Seventeen of 
these were captured, of which 15 were females and 2 were males. Five 
of the females were banded bats that had been captured by the writer 
at the same place on September 20, 1947, and kept in captivity ]n San 
Francisco until June 6, 1948, at which time they were released at the 
original site of capture. This represented a 25 per cent return since 
15 other pallid bats, taken at the same place and on the same date, 
were also released here with bands on their forearms on June 6, 1948. 
It is probable that additional banded individuals were present in the 
colony on April 15, 1949, but they were not captured. 

This same colony was visited on April 14, 1951. The owner of the 
ranch on which the barn that housed the bats was situated stated that 
the colony had made its appearance several weeks before. Difficulty 
in gaining access to the roof of the barn made it possible to capture 
only 5 females and 4 males, although many more bats were heard and 
known to be present. Two of the females and one of the males were 
banded bats. One female belonged to the group originally captured 
on September 20, 1947, and released on June 6, 1948, after being kept 


in San Francisco for some months. It was not one of those recaptured 
on April 15, 1949. The other female and the male had been banded 
at this colony on October 22, 1949, and released immediately. On 
March 20, 1953, Dr. and Mrs. Karl B. Koford visited this colony and 
found 4 females and 1 male present. An unusually mild winter and 
early spring may have accounted for the early seasonal appearance of 
bats at this summer colony in 1953. 

A second roost, known to have been inhabited the previous three sum- 
mers, was visited on April 14, 1951, and found to be occupied. The 
small quantity of fresh guano on the previous years' deposit beneath 
the roost indicated that the colony had not been established here more 
than a week or two. Although about 40 pallid bats were estimated 
to be present, only 11 were captured. Two were males and 9 were fe- 
males. In addition, 7 Mexican free-tailed bats were taken (2 males, 5 
females). All were weighed, examined for parasites, and then released. 

So far as known pallid bats are gregarious during the spring and 
summer months. The number of adult individuals comprising a colony 
was found to vary from about 30 to approximately 100. There is con- 
siderable variation in the sex ratio in different colonies and, seemingly, 
even within the same colony at different times. Luther Little (]\Iail- 
liard, MS) captured 19 adults from a fairly large colony inhal)iting 
a deserted building in Lake County, California, on April 16, 1919. 
Eighteen were males and 1 was a female. Twenty-two adults captured 
at random from a colony composed of about 40 individuals inhabiting 
the attic of Encina Hall, Stanford University, California, on June 13, 
1951, consisted of 15 males and 7 females with young. Five adult fe- 
males, however, had been taken from this same colony by another per- 
son the week before. Of 15 adults captured in western Kern County 
on May 10, 1947, from a colony estimated to consist of about 100 indi- 
viduals, 8 were males and 7 were females (either pregnant or with 
young attached). Three adult males and 4 females, pregnant or with 
newborn young, were secured from a colony of 60 bats in eastern San 
Luis Obispo County on June 9, 1948. Two days previously a random 
sample from another colony of similar size several miles away con- 
sisted of 7 adult males and 35 pregnant females. Thirty-seven pallid 
bats captvired from another colony in this region on September 20, 1947, 
consisted of 8 males and 29 females. At this time the number of indi- 
viduals in the colony was estimated to be somewhat in excess of 60. On 
June 6, 1948, 26 adults were captured at random at this same colony. 
Sixteen were males and 10 were pregnant females. As has been pre- 
viously stated, on April 14, 1949, shortly after the formation of this 
colony in the spring, a random catch produced 2 adult males and 15 
adult females. Another random catch on April 14, 1951, produced 4 



adult males and 5 adult females, and on March 20, 1953, there were 
1 adult male and 4 adult females in the colony. 

Storer (1931, p. 244) records the capture of 8 adult or nearly adult 
males and 13 females, either pregnant or with young, from a colony 
of pallid bats living between the walls of a residence in Berkeley, Cali- 
fornia, on June 24, 1919. Hall (1946, p. 163), on the other hand, re- 
ferring to this species in Nevada, states that "Pallid bats separate by 
sex in spring before the young are l)orn. Tlie females then are in colo- 
nies. About fifty females is the largest number observed in one colony." 
Dalquest (1947, p. 24) records a colony of pallid bats, consisting of 
about 60 males, found in the attic of an old winery near Angwin, Napa 
County, California, on July 8, 1945. 

Figure 10. Mexican tree-tailed bats are very treyueiitly associated with pallid 
bats in the wild. When housed in the same laboratory cage members of the two 
species stayed together. Photographed at Iho California Academy of Sciences, 
December 12, 1952. 


From the foregoing observations it can be seen that, in Califor- 
nia at least, summering colonies may be composed of adults of both 
sexes and that either sex may predominate in numbers. 

Although big brown bats (Eptesicus fuscus), several species of Myotis, 
and lump-nosed bats (Corynorhinus rafinesquei) may occasionally be found 
in the same cave, building, or even attic with pallid bats in the daytime 
during the summer, they do not seem to share the same roost with 
members of the last species. Only the Mexican free-tailed bat (Tada- 
rida mexicana) is commonly found in pallid bat roosts (fig. 10). Most 
of the pallid bat colonies examined were found to contain one or more 
of these small molossids. The greatest number observed with pallid 
bats in one colony was 17. Members of either or both sexes may be 
represented, including pregnant females in the late spring and sum- 
mer. Individuals of this species were found to be equally common in 
colonies of big brown bats. 

Krutzsch (1946, p. 241), discussing big brown bats in San Diego 
County, California, states that : "Eptesicus, Tadarida, and Antrozous have 
been found living successfully in the same building, a barn on the 
grounds of The Old People's Home, in the attic of the San Marco 
Church, and in various cracks between the "I" beams of the bridge at 
the juncture of Highways 79 and 80." Dalquest (1947) mentions find- 
ing Corynorhinus rafinesquei, Myotis yumitnensis, Myotis thysanodes, and 
Tadarida mexicana inhabiting the same buildings as pallid bats during 
the daytime. Bailey (1936, p. 392) records a mixed colony of pallid 
bats and Mexican free-tailed bats found in Carson Valley, Nevada. Hall 
(1946, p. 165) records 2 pipistrelles (PipistreUus hesperus) and 1 little 
brown bat (Myotis californicus) hanging in a mine tunnel in which 2 
pallid bats were found hibernating near Yerington, Nevada, on Decem- 
ber 27, 1939. 

FedJ: By the middle of September the adults have all completed the 
annual molt (many have done so by the middle of summer) and the 
young of the year have matured to the stage where they are indis- 
tinguishable, at least in the field, from adults. No evidence was obtained 
to indicate dissolution of the summer colonies this early. By the middle 
of Octol)er, however, the summer colonies tend to break up into smaller 
groups. At this time pallid bats may be found in situations where 
they do not occur in summer. 

On October 22 and 23, 1949, some known sunuuer colony sites in 
eastern San Luis Obispo County were examined. The first colony 
visited was in the barn, previously mentioned, where 20 banded indi- 
viduals captured on September 20, 1947, had been released on June 6, 
1948. Although over 60 bats were generally found here in summer 
there were onlv 12 on October 22. Six were males and 6 were females. 


One of the females was a banded individual, one of the group captured 
on September 20, 1947. 

Examination of a deserted cabin a])out one mile from this colony 
resulted in the discovery of a male and a female pallid bat hanging side 
by side behind a loose board in one of the rooms. Although this cabin 
had been investigated many times before in winter, spring, and sum- 
mer, the only bats previously found here had been of the species Myotis 
suhiihitus. Later that evening the night roost of another colony of pallid 
bats was visited and 8 or 10 individuals were observed hanging there 
before the approach of intruders disturbed them. 

The following day the site of another summer colony was visited. 
It contained only 7 bats, 5 of which were captured. Two of these 
were males and 3 were females. About 30 feet away another small 
group composed of 13 pallid bats was discovered. Only two of these 
were captured. They proved to be females. P^xamination of two other 
summer colony sites that day showed that one had recently been va- 
cated while the other contained 5 pallid bats and 15 Mexican free- 
tailed bats. None of the former were captured. 

Mr. Eben McMillan made some interesting observations on the 
appearance of pallid bats in the fall of 1948 and 1949 on the Pinole 
Ranch at the north end of the Carrizo Plain in eastern San Luis 
Obispo County. No pallid l^ats were seen about the ranch buildings 
in the spring or summer months of these two years. In 1948, how- 
ever, between the latter part of October and the middle of Novem- 
ber, pallid bats were found hanging each night from beams support- 
ing the grain bins. Each moi-ning dui-ing this period numerous drop- 
pings and fragments of insects, presumably discarded by the bats, 
were found on the ground beneath the beams. After the middle of 
November the bats were no longer seen. In 1949 pallid bats again 
a]ipeared here during the second week in October. On October 23 
the writer observed many fresh droppings as well as remains of Jeru- 
salem crickets on some clean sacks that had been placed beneath this 
temporary night roost the evening before. Dui'ing the next week or 
so Mr. McMillan examined the sacks each day and then cleaned 
them. After October 25 there was a gradual decrease in the number 
of di-oppings found daily until November 1. On this date there were 
only two droppings and none were found subsequently that year. 

Winter: Very little information is available regarding the where- 
abouts of pallid bats between the middle of November and the end 
of March. They are not to be found in their summer retreats nor are 
they active at night, as far as could be determined. There is no evi- 
dence to indicate that members of this species migrate in the strict 
sense of the word although unquestionably there is a local shifting 


and probably a dispersal of summer populations. Such meager infor- 
mation as was acquired seems to indicate that single individuals or 
small groups of these bats seek remote retreats where they are not 
likely to be disturbed. 

Referring to this species in Nevada, Hall (1946, p. 165) says: "The 
hibernating individuals consisted of two males found December 27, 
1939, in a mine tunnel 9 miles east and 2 miles north of Yerington, 
120 and 138 feet from the mouth of the tunnel. Each bat was in a 
crevice in the roof of the tunnel and had its ears erect and its eyes 
open. The bats were unable to fly and their movements were slow. 
The temperature inside the mine where the bats were was 60 °F. ; out- 
side the mine it M^as between 40° and 50°F. (Alcorn, MS)." Alcorn 
(1944, pp. 309-310), referring to the same cave, mentions that pallid 
bats were noted here during three winters. 

Mrs. Grinnell (1918, p. 356) records two females in the collection of 
Stanford University taken on January 1, 1895, at Carmel Mission, Mon- 
terey County, California. A female (no. 106583) in the collection of 
the Museum of Vertebrate Zoology, University of California, was taken 
at AVoodside, Santa Clara County, on January 17, 1947. It weighed 
22.3 grams. 

On January 18, 1950, a male pallid bat was received that had been 
captured three days previously in Mill Valley, Marin County, Califor- 
nia, by Thomas Leech. Although the animal had been in captivity for 
three days its weight upon receipt was 26.5 grams. A few days later, 
January 22, the writer visited the place where the bat had been found. 
It proved to be an old, deserted, one and one-half story cabin situated 
in a rather dense redwood forest on a northeast-facing slope. The bat 
liad been found hanging on the outside of the building beneath an old 
]>iece of damp canvas where it undoubtedly was dark, cool, and damp 
all winter. Further search failed to uncover any additional pallid bats, 
although a female big brown bat (Eptesicus fuscus) and a female lump- 
nosed bat (Corynorhmus rafinesquei), each dormant, were hanging 
within the cabin. 

On December 1, 1940, E. Raymond Hall, Ned Stone, and Thane 
Riney found a male and two female pallid bats in a crevice in a lime- 
stone cliff 6 miles east of Walnut Creek, 1450 feet. Contra Costa County, 
California. The surrounding territory was covered with chaparral 
(Perry, MS). 

Dr. Charles G. Sibley informed the writer that on January 28, 1940. 
he found 4 pallid bats in a semitorpid condition inside a hollow post 
holding up one end of an old lean-to on Bolinas Ridge, 1500 feet, Marin 
County, California. His attention was directed to the presence of the 
bats by faint noises, reminiscent of small mice, coming from within 


the post. Two of tlie bats were males, weighing 25.8 and 28.5 grams, 
and 2 were females, weighing 28.3 and 32.5 grams. They are now in 
the collection of the Museum of Vertebrate Zoology (nos. 90572-90575). 

On March 13, 1950, Mr. Eben McMillan found a torpid female pallid 
bat hanging beneath a loose board in a deserted cabin about 5 miles north- 
east of Shandon, 1300 feet, San Luis Obispo Count}^ California. This 
is the only possible winter record that was obtained from San Luis 
Obispo and Kern counties. Mr. McMillan and the writer had found 
a male and female of this species, in breeding condition, hanging be- 
neath this same board on October 22, 1949. Careful search for winter- 
ing bats, on other occasions, in old buildings, mine tunnels, eaves, and 
other possible retreats in parts of Monterey, San Luis Obispo, and Kern 
counties failed to reveal the presence of any pallid bats although bats 
of the following species were located : Myotis thysanodes, Myotis voJans, 
Myotis suhulatus, Eptesicus fuscus, and Corynorhinus rafinesquei. In 
this region, however, there are countless natural crevices in cliffs and 
rocky outcrops that are essentially inaccessible for human examination 
and which could serve as adequate hiding places for wintering bats. 

Effect of enforced dormancy: Efforts were made to determine the 
effect of a simulated winter environment on captive individuals. On 
September 25, 1947, 6 females and 2 males, that had been captured a 
few days previously in eastern San Luis Obispo County, were placed in 
a small retaining cage in a refrigerated room where the temperature 
ranged from 40°F. to 50°F. To maintain fairly high humidity wdthin 
the cage the screened front was covered with burlap which was damp- 
ened daily. A container with water was kept on the floor of the cage. 
No food was given the animals during the ensuing weeks. 

The 2 males died, one after 14 days and the other at 40 days. Two 
of the females also died, one at 8 days and the other at 36 days. The 
death of some of these bats was suspected to have lesulted from the 
toxic effect of aniline dyes placed on their bodies as a means of iden- 
tification. The remaining 4 females were removed to a warm labora- 
tory at the end of 43 days. During this period of enforced dormancy 
they had been weighed weekly. Occasional increases in weight were 
attributed to water that was believed to have been taken from time to 
time. The average loss in weight, per bat, at the end of 43 days 
amounted to 8.1 per cent or 0.184 per cent per day. The gradual loss 
in weight of 2 of these bats is shown in figure 11. 

During the 28 days following the period of enforced dormancy these 
two bats were kept in a small cage in the laboratory and fed daily. 
Daily records were kept of their weights. As is shown in figure 11 there 
was a very rapid increase in weight during this time. The average in- 
crease, per bat, at the end of 27 days amounted to 60.8 per cent or 2.25 


per cent per day. On the last day there was a slight decline in the 
weight of each bat. 

At the end of this 28-day period these two bats were returned to the 
refrigerated room and kept there from December 5, 1947, until April 
27, 1948, under the same conditions used in the first dormancy experi- 
ment. At the end of this period of 144 days the decline in body 
weight, per bat, averaged 27.6 per cent or 0.193 per cent per day 
(fig. 11). 

On November 15, 1950, 4 pallid bats (3 males, 1 female) that had 
been captured at Farmington, San Joaquin County, California, on 
October 27, 1950, were placed in a cage in a refrigerated room and 
kept there until March 14, 1951. They were provided with drinking 
water and the front of the cage was kept covered with damp burlap. 
During this period of 119 days the temperature of the room was kept 
at 38°F. to 40°F. The bats were weighed weekly. The average loss in 
weight, per bat, amounted to 24.96 per cent or 0.209 per cent per day 
(fig. 12). 

In each of the three dormancy experiments mentioned the bats 
hung from the screened side of the cage. The water container was 
visited by one or more of the bats about every 3 to 5 days. This was 
demonstrated by placing a thin layer of fine dry sand on the bottom 
of the cage and examining it every day or so for tracks. When tracks 
were found leading to the water dish the sand was smoothed over 
again. Several investigators have previously suggested the possibility 
that hibernating bats are periodically aroused by thirst. This may 
have been true of these captive individuals since they periodically 
visited the water container and on several occasions a temporary in- 
crease in body weight was noted. Under natural conditions most hiber- 
nating bats that have been studied are not far from water, either in 
the form of accumulated droplets in the hibernacula or in streams 
and ponds nearby. Folk (1940, p. 312) observed little brown bats 
(Myotis hicifugus), aroused from a torpid condition in winter in a 
cave in New York, lapping drops of moisture condensed on the wall. 
Furthermore, there is occasional shifting of individuals during the 
general period of dormancy which may be the result of movement to 
and from water. As is shown in figures 11 and 12 the greatest loss in 
weight occurred during either the first or second week of enforced 

The respiration rate in dormant individuals varied greatly and 

Figure 11. Graph showing changes in body weights of 2 pallid bats kept with- 
out food at 40°F. to 50°F. for 43 days, then removed to a warm laboratory room 
and fed for 28 days, following which they were again kept at 40°F. to 50°F. 
without food for 144 days. 



(JiOOr^vDift^rOcJ — O(T>e0t^(X)inM-rO<M — O(T)00 
fOrOrOrorororOrororOCvJOJCNJOoojCMcvJcocJOJ — — 

-»• "^ 

aicor^<x)if)"3-rooj — ocDcD^-t£)in^rO(M — OCT) 




[Proc. 4th Seb. 

was difficult to determine accurately. Bats cannot be said to become 
truly torpid since even the turning on of a light in the refrigerated 
room at times stimulated breathing. The noise involved in the open- 
ing of a door or the sound of footsteps had the same effect. Numerous 
respiratory counts made on the bats maintained in a refrigerated room 
showed a range of to 52 per minute. The greatest period in which 
no obvious sign of breathing was observed was 4 minutes. 

Dormant bats, when suddenly handled, reacted feebly at first. The 
mouth was generally opened wide and a weak buzzing sound pro- 
duced. Following this, if the wings were free, one or both of them 
would be slowly extended and then held stiffly in an outspread posi- 
tion. No attempt, of course, was made to disturb dormant individuals 
any more than was necessary. When two females were placed in the 
sun in an outdoor flight cage after 144 days of confinement without 
food in a refrigerated room they were capable of sustained flight 
within 15 minutes. 

Neither bats kept in the laboratory nor those in outdoor flight 
cages became dormant for any considerable period of time during the 
winter. They were, however, more sluggish and ate less at this sea- 
son. Those in outdoor cages occasionally failed to take any food on 
cold or stormy nights. 















77 84 


DAYS 7 14 21 28 35 42 49 56 63 70 

Figure 12. Graph showing gradual loss in body weights of 4 pallid 
at 38°F. to 40°F., without food, for 119 days. Solid lines indicate males, 
line a female. 

98 105 112 119 

bats kept 
the dotted 



Seasonal fluctuation in body weight: As is shown in figure 11, captive 
liallid bats are capable of increasing- body weight rapidly after a period 
of dormancy. Data on bats from a single colony, presented in table II, 
indicate that this is also probably true of these bats in the wild. Males 
captured in April, 1951, several weeks after their appearance in the 
summer colony, weighed about the same as males taken in June. Males 
captured only a few days after their arrival in the summer colony 
in April, 1949, weighed considerably less. Females continued to increase 
in weight from the time of their arrival in spring until the young were 
born. Although no opportunity was afforded to secure weights of fe- 
males from this particular colony immediately after the period of par- 
turition, the average and extreme weights of 7 females, captured June 
13, 1951, in Santa Clara County, California, that had young from about 3 
to 6 weeks old, were 26.8 (25.1-29.3) grams. There appeared to be 
an increase in the weight of bats of both sexes again in the fall. 

Table II 

Average weights, in grams, of pallid bats from a single colony 4Vi; miles 
northeast of Shandon, San Luis Obispo County, California, in spring, summer, 
and autumn. Figures in parentheses represent numbers averaged. 





April 16. 1949 

20.6 (2) 

24.3 (15) 

Females had 
recently ovulated 

April 14, 19512 

25.1 (3) 

31.7 (2) 

Females with well 
developed embryos 

June 6, 1948 

25.2 (16) 

34.7 (10) 

Females in late 

October 22, 1949 

29.4 (7) 

28.3 (7) 


Daily Behavior 

Voice: A number of different kinds of audible sounds are produced by 
]iallid bats. Each of these seems to be associated with a particular type of 

The "intimidation" note is a loud, strong, dry, insect-like buzz that 
is uttered when a bat is frightened, angered, or very annoyed. The 

2. The fact that bats secured from this colony weighed more in the middle of April, 1951, than 
those secured about the same date in April, 1949, is attributed to their earlier seasonal appear- 
ance in 1951. 


mouth is half-opened and tlie teeth bared at such times. From 1 to 7 
successive buzzes may be given with equal time intervals between. The 
rate of utterance is about 3 to 5 per second. The individual buzzy 
notes are all of approximately the same pitch, quality, intensity, and 
duration. Any disturbance of a roosting colony during the day imme- 
diately resulted in one or more of the bats uttering this .sound. Captive 
bats uttered the intimidation buzz when disturbed by persons attempt- 
ing to handle them, when disturbed by other bats in the same cage, or 
when suddenly startled. This note was frequently uttered during the 
day by a female that was attempting to discourage her nearly grown 
young from nursing. 

The "squabble" consists of a series of high-pitched, dry, rasping, 
thin, double notes of duration varying up to one second. In certain 
respects it resembles the squeaky note of the Anna hummingbird. This 
sound is produced with the mouth open and the teeth bared. Squab- 
bling appeared to be a sign of irritability and anger and was uttered 
by individuals that were being crowded too much or slightly hurt by 
other bats. It was frequently heard during the day, especially when 
it was warm and the bats were restless. On several occasions the squab- 
bling of pallid bats led to the discovery of colonies in the field. 

The "directive" call usually consists of 1 to 5 rapidly repeated notes 
with equal time intervals between. The notes are single, clear, resonant 
in quality, and of a high pitch. They might be phonetically described 
as "sit" or "sit-sit-sit, " depending on the number of notes uttered, and 
remind one of the sound made by a power line with a short circuit. 
This call is usually given as soon as a bat takes flight in the evening 
on emerging from a daytime roost. It was occasionally uttered by caiv 
tive individuals when they were released for exercise in the laboratory. 
Captive bats were also heard to utter these notes sometimes while 
crawling about the laboratory or even when hanging in the room. This 
call is frequently heard at night in areas where pallid bats are abun- 
dant. On one occasion, in the midafternoon, 42. bats that had just been 
captured and examined were released one at a time close to their 
roost. The first 5 flew about for a few moments then entered a crevice 
in a rock. They uttered the directive notes after they disappeared 
from sight. These notes were answered by other bats, that were then 
released, and seemed to serve as a means of directing the latter to 
the same crevice. On another occasion, in the laboratory, it was noted 
that the utterance of directive notes by a bat 3i/4 months old caused 3 
adults to fly to the young, hang next to it, and nuzzle it. 

The directive note of the young pallid bats may be described as a 
"chirp." This is a high-pitched, bird-like note that the young are 
capable of uttering from the very moment of birth until they are 


old enough to i)rodiice tlie adult type of directive note previously 
described. In captivity chirps were uttered as soon as a young bat 
became detached from its mother and were repeated monotonously until 
the parent rejoined the young. The frequency of utterance of these 
notes during periods of detachment increased from 2 or 3 per second 
during the first day of postnatal life to 5 per second by the tenth 
day. By the time young bats were about 12 days old this chirp was 
replaced by a directive call similar to that given by adults. In one 
instance this change in voice occurred at the age of 7 days. The chirp 
seemed to serve as a means of acquainting the female with the loca- 
tion of her young. When a female was separated from her young 
and released in another part of the room, or a young bat was taken 
from its mother and placed in another part of the room, the parent 
seemed to depend upon this sound to locate her offspring. 

A sound which may be described as "chittering" was heard on 
some occasions. Although its significance is not clearly known, it was 
suspected of being a note of contentment. Each of these notes was 
of a high pitch, although the intensity varied slightly, of about one- 
half second duration, and repeated 3 or 4 times with half-second inter- 
vals between notes. On one occasion these chittering notes were heard 
coming from a cage in which an adult female and her 6-weeks old 
young were hanging side by side. Laboratory-reared bats that were 
very tame were occasionally heard uttering these notes during the day- 
time until they were 18 months old. 

A "plaintive" note, presumably associated with pain, was uttered by 
females in labor. Such notes were high-pitched but not as high as the 
directive notes. They were loud, gutteral, harsh, double notes repeated 
3 or 4 times, each note having a duration of about 2 seconds. They 
were uttered with the mouth open, the lips drawn back, and the eyes 
partly closed. They evoked no response from other bats in the same 

On occasion as bats groom themselves they produce a non-vocal, ex- 
plosive, nasal sound. This apparently is the result of a sudden release 
of air through the nasal passages and may serve as a means of dis- 
lodging loose hairs that adhere to the nose. 

When bats were flying about in a laboratory room they could l)e 
heard making faint clicks as they passed close by the observer. These 
same clicks were often produced by tame bats held in the hand at 
times when they were restless and about to take flight. The writer 
did not have an opportunity to study ultrasonic sounds produced ])y 
these bats but was kindly supplied with some information by Dr. F. 
Alton Everest of the Moody Institute of Science in Los Angeles. 
Through the cooperation of Mr. Eben McMillan 3 adult female and 3 



[Proc. 4Tn Ser. 

nursing young pallid bats were secured by Dr. Everest at one of the 
colonies under observation by the writer on the Carrizo Plain. By 
means of an ultrasonic amplifier it was found that during flight these 
bats emit a very narrow beam of sound from the open mouth. The sig- 
nal strength of this sound was found to fall very rapidly when the 
mouth w^as not aimed directly at the microphone. The same change in 
rate of click was observed as that described by Galambos and (h-iffin 
(1942). Pallid bats appeared to have an ultrasonic energy peak of 
about 40 kilocycles per second. 

Locomotion: Pallid bats are capable of crawling quite well. When 
moving forward on a horizontal surface the weight of the body is borne 
on the wrists and feet. The knees are bent and directed upward and 
the tail is partly curled under. The fore part of the body is elevated 
more than the hind part. This position differs noticeably from that 
assumed by the Mexican free-tailed bat. Members of the latter species 
keep the body parallel to the surface on which they are crawling. 

Figui-e 13. Pallid bat photographed at the moment it was taking flight from 
the edge of a table. Photographed at the California Academy of Sciences, Octo- 
ber 26, 1952. 



Figure 14. Shortly after the beginning of the downstroke. 
Bob Lackenbach, September 1, 1949. 

Photograplied by 

Figure 15. Position of wings in downstroke slightly later than shown in figure 
14. Photographed at California Academy of Sciences, June 26, 1952. 



Figure 16. Midway in the downstroke. Photographed at the California Academy 
of Sciences, December 10, 1952. 

Figure 17. Near the end of the downstroke. Photographed by Bob Lackenbach, 
September 1, 1949. 



Figure 18. At the end of the downstroke. Photographed at the California 
Academy of Sciences, November 20, 1952. 

Pallid bats can climb readily, relying on the sharp claws of the 
hind toes and thumbs to hold on to vertical surfaces. If undisturbed 
when climl)ing they usually face the direction in which they are going. 
If they are bothered or frightened, however, they will frequently re- 
verse their position and back up. No doubt this enables them to take 
flight more rapidly than would be possible if the head were facing 
upward. Likewise it generally makes it possible to direct intimidation 
notes and gestures more effectively toward the source of annoyance. 

In flight, individuals of this .species appear to be somewhat slower 
and less agile than many smaller bats. This impression results partly 
from the large size of the pallid bats. Their wdng beats are slower and 
they require more room in which to turn. There is no question but that 
they lack the ability to maneuver in a limited space as effectively as 
smaller kinds of bats. It is questionable, however, whether they are 
actually slower flyers than most other vespertilionids. The Mexican 
free-tailed bat, on the other hand, which is a molossid with propor- 
tionately long, narrow wings, can outdistance the pallid bat very easily. 
Since these two species are frequently associated with one another, 
this was observed many times in the field as well as in the laboratory. 

Field studies on flight under natural conditions were of little value 
as pallid bats emerge from their daytime roosts rather late in the eve- 



[Pkoc. 4th Ser. 

Figure 19. Midway in the upstroke as viewed from the side. Photograplied 
at tlie California Academy of Sciences, November 20, 1952. 

Figure 20. Midway in tlie upstroke as seen from below. Photographed at 
the California Academy of Sciences, November 20, 1952. 



Figure 21. Near the end of the upstroke. 
Academy of Sciences, April 29, 1953. 

Photographed at the California 

niiig when the light intensity is very low. Stndies, however, were made 
on the flight of captive bats in laboratory rooms. Direct observations 
were supplemented by slow motion films (64 frames per second) and 
stills made with a Speed Graphic and an Exacta VX using a Dormitzer 
Synctron electronic flash with a speed of about one five-thousandth 
of a second. 

Before discussing the mechanics of flight it might be well to com- 
pare briefly a vespertilionid bat with a bird as regards the structures 
used in this form of locomotion. The chiropatagium may be considered 
as analogous to the outer half of a bird's wing. From the functional 
standpoint it serves the same purpose and acts as a i>ropeller to effect 
forward progress. The plagiopatagium is analogous to the inner half 
of a bird's wnng and like the latter serves to supply much of the neces- 
sarv lift. The movement of the plagiopatagium is considerabh' less 



tlian tliat of the chiropatagiuin in flight. The anterior part of the bat's 
wing, like the anterior part of a bird's wing, possesses the greatest 
rigidity and it is liere that the drive or pressure is exerted on the down- 
stroke. The uropataginm is analogous to the tail feathers of tlie bird 
and may perform many of the same functions. 

On extending the wing of a bat it will be seen tliat the tension of 
the membranes is sufficient to cause the fourth and fifth digits to flex 
slightly so that the ventral surface of the wing is concave and the 
dorsal surface convex. The greatest bend occurs between the fifth meta- 
carpal and the first phalanx of the fifth digit. The downward bend of 
the fourth finger is less pronounced. As will be seen in figure 16 the 
position of the hind leg and the pressure exerted by the air beneath 
the plagiopatagium serve to maintain this cam])er in flight. 

Figures 13 to 21 illustrate the action of the bat's wung. The mo- 
tion is essentially the same as that of the wing of a l)ird in forward 
flight (Aymar, 1935, pp. 136-137; Storer, 1948, pp. 31-32). The down- 
stroke is downward and forward while the upstroke is upward and back- 
ward. This has already been noted by Griffin (1946) whose studies on 
fliglit were based upon high-speed photography developed l)y Profes- 
sor Harold E. Edgerton at the ]\Iassachusetts Institute of Technology. 

In one respect the upstroke may be considered less efficient in the 
bat's wing than in the bird's wing. In the bird the flight feathers of 
the wdng can separate during this action to allow air to pass between 
them, thus minimizing resistance. No action comparable to this is pos- 
sible in the bat. However, in the bat, as in the bird, resistance is mini- 
mized by the generally convex shape of the upper surface of the wing 
as well as by a partial folding of the wing during the upstroke. Slow 
motion pictures also show that the wing action is much more rapid on 
the upstroke than on the downstroke. Furthermore, on the downstroke 
the uropatagium is depressed considerably as a result of the legs l)eing 
pushed forward and the tail curled ventrally (fig. 18). This tends to 
increase the lift in a species such as the pallid bat which has a large 
uropatagial membrane and thus partly compensates for the loss occur- 
ring with the succeeding upstroke. The uropatagium does not straigliten 
out until the wings are raised. 

The uropatagium not only aids in increasing lift during sustained 
flight but serves as a brake when a bat is slowing down to alight or to 
avoid an obstacle suddenly placed in its w^ay. Under these circum- 
stances the legs are separated more widely than in regular flight and 
thrust far forward, thus depressing the uropatagium. At such times 
the angle of the body and wings is increased so as to approach the 
.stalling point. 

While in flight the thumb is always extended, essentially at right 



Figure 22. A close view of tlie liead of a pallid bat in flight showing the posi- 
tion of the pinna, tragus, and mouth. Photographed at the California Academy 
of Sciences, November 20, 1952. 

angles to the anterior edge of the wing. The moiitli is opened slightly 
and the upper and lower lips are decidedly drawn apart anteriorly (fig. 
22). This is believed to l)e associated with the ntterance of ultrasonic 

In the laboratory, pallid bats were occasionally seen to hover mo- 
mentarily when flying close to the floor. At such times their forward 
]n'ogress was essentially stopped. Such hovering action was thought to 
be associated with the habit these bats have of capturing food on the 
ground. Occasionally pallid bats were observed to glide for short dis- 
tances with the wings fully extended and up and down motion sus- 
pended or so slight as to be hardly discernible. 

The number of wing beats per second was found to l)e subject to 
considerable variation, depending on the type of flight. Griffin (1946, p. 
119) states that "Bats make about 15 strokes of the wing per second. . . ." 
This may be true of the vespertilionid bats of eastern United States that 
he studied. The pallid bat, which is considerably larger than any of 
the cave-dwelling species of eastern North America, was found to make 
10 to 11 strokes per second in ordinary straight flight. The rate of wing 
beat was found to increase under certain circumstances, as for example 
in executing a turn. On such occasions as many as 131/2 strokes per second 


were recorded, while in a glide the number of strokes per second was 
reduced to zero for a brief period. 

Pallid bats usually alight head up on vertical surfaces, catching on 
to the landing surface by means of the claws on their thumbs and hind 
feet. If the hind claws have found satisfactory crevices to cling to, the 
thumb holds are immediately released, as is also that of one hind foot. 
This causes the body to swing down to an inverted position at which 
time the claws of the thumbs and the released hind foot again grasp 
the landing surface. This method of alighting differs from that de- 
scribed by Dalquest (1947, p. 25), for Conjnorhinus rafinesquei, a 
smaller and more agile species. It also differs decidedly from the way 
the phyllostomatid bat, Macvotus caUfornicus, alights (Hatfield, 1937). 

Odor: Pallid bats produce a very characteristic, skunk-like odor. On 
occasion one could detect this odor as much as 15 feet from an entrance 
to a roosting crevice. Disturbance of the bats results in a marked in- 
crease in the intensity of the odor. The odor is produced by the se- 
cretion of the glands on either side of the muzzle. On the surface of 
each gland are a number of pore-like depressions. A bristle is located in 
the center of each depression and each bristle is surrounded by about five 
smaller hairs (fig. 2). When a bat is disturbed, numerous tiny drop- 
lets of secretion are exuded within a few seconds and the odor becomes 
very strong. When the glands of a freshly killed bat are cut the odor 
becomes intense. It has been suggested that oil secreted by the nose 
glands of some bats is used to rub on the flight membranes to protect 
them from climatic conditions (Abdulali, 19-49, p. 423). In the pallid 
bat, however, it seems more likely that the secretion may serve as a de- 
fense mechanism. It is true that members of this species frequently 
nuzzle their wing membranes but it is also true that the muzzle glands 
seem actively to secrete an odorous exudate only wiien the bats are 
alarmed. Perhaps this odor tends to repel certain other kinds of 

Diurnal behavior: Since the body temperature of many kinds of bats, 
including the species under discussion, has been found to be close to 
that of the environment when the animals are at rest during the day 
(fig. 23), it is not surprising that the behavior of these animals differs 
on cold days from that on warm days. On cool days, or early in the 
morning before it had warmed up, members of pallid bat colonies were 
found to be silent and essentially motionless until disturbed. If an 
individual was prodded at such times it would open its mouth and 
utter the intimidation buzz. ]\Iovement was slow at first and it was only 
after several minutes of continued disturbance that the bats would 
scramble around actively or be able to fly. On approaching a colony on 
a warm day one could frequently hear bats squabbling when 100 feet 



■.SS>R%'^,. 3»i> V*> .>"^«(S 

Figure 23. Typical posture assumed by captive pallid bats when sleeping 
during the day. Photographed at the California Academy of Sciences, Decem- 
ber 12, 1952. 

away. Any attempt to reacli into a crevice resulted in numerous in- 
timidation and squal)ble notes as the bats actively backed to the inner- 
most part of the roost. They were very alert at such times, keeping 
tlieir heads raised and often their mouths open. The latter act was 
inter])reted as an intimidation gesture. If further disturbed, various in- 
dividuals would try to move rapidly to the roost entrance and take flight. 
Likewise, in the laboratory cages much more activity and quarreling 
Avere evident when it was warm than when it was cool. 

Newly captured bats usually bit viciously when grabbed. Anger at 
other bats was often expressed by biting if the intimidation buzz failed 
to discourage the annoyer. This was often preceded or accompanied by 


the squabble note. The majority of tlie bats that were kept in captivity 
for several months and handled regularly became accustomed to such 
treatment and ceased calling and biting when handled. However, they 
would usually open the mouth widely, bare the teeth, and occasionally 
move the jaws in a chewing soi-t of motion. Certain individuals never 
Ijecame tame while others became very tame. Some of the young born 
and reared in capti\'ity would come of their own accord to the person 
that regularly cared for them and seemed to prefer being held in a 
hand to hanging in a cage. Bats that were wild or were being held 
in hand against their will would usually resort finally to spasmodic 
jerking of the body if all other means failed to give them freedom. 
Such behavior was frequently exhibited when body measurements or 
vaginal smears w^ere being taken or when pelage examinations were 
being made. Such jerking ceased immediately when the animal was 

During the day, captive bats were frequently observed nuzzling their 
fur and combing it with the hind claws. Whether or not the tongue was 
used in this cleaning process could not be determined. The flight mem- 
branes were also nuzzled and considerable time was devoted to scratch- 
ing ectoparasites. 

Captive pallid bats, both in laboratory and out-of-door cages, usually 
slept together in compact groups. Only occasionally would individuals 
stay apart from others in the cage and generally such behavior was the 
result of illness or injury. Not only did they customarily stay together 
but, after living in a cage a short while, developed the habit of roost- 
ing each day in the same place. When returned to a large flight cage 
after liaving been kept elsewhere for several weeks they would fly di- 
rectly to the place where they had previously roosted. On several occasions 
roosts, consisting either of bricks or boards, were moved a few feet to 
determine the reaction of the bats upon being returned to a cage after 
an absence of two weeks. In each instance they flew to the original site 
of the roost and appeared bewildered. After a few moments of inves- 
tigating, the new site of the roost was found. It was difficult to deter- 
mine whether sight, odor, echo location, or all were involved in deter- 
mining the position of the roosts. 

On several occasions bats which had been captured in a barn in San 
Luis Obispo County were kept in San Francisco for as long as 13 
months and then liberated at the original site of capture. Almost all 
of the bats, immediately upon being released outside of the barn in 
the daytime, flew directly to a small opening beneath the peak of the 
roof, that was regularly used by members of the colony as an exit and 
entrance to the attic of the barn where the roost was located. 


Nocturnal behavior: Pallid ])ats do not eineroe from tlioir daytime 
roosts until fairly late in the evening. Observations made on June 8, 
1948, on a colony in eastern San Luis Obispo County, showed that the 
first bats emerged 47 minutes after sunset at wliioh time tlie illumina- 
tion was less than 0.1 foot candles. By way of comparison, the first 
western pipistrelles were observed 2 minutes after sunset at which 
time the illumination was very high, and the first Mexican free-tailed 
bats emerged 17 minutes after sunset when the illumination was 3.0 
foot candles. In the early morning of June 9, 1948, the last pallid bat 
was seen in flight 51 minutes before sunrise when tlie illumination was 
less than 0.1 foot candles. There is some evidence that in the fall of 
tlie year pallid bats, as well as several other species of bats observed, 
emerge earlier, relative to sunset, than in summer. On September 20, 
1947, the first pallid bats wei-e seen 19 minutes after sunset when the 
illumination was 0.15 foot candles. On that same evening western pipis- 
trelle bats were seen 6 minutes before sunset and on the following eve- 
ning Mexican free-tailed bats were seen emerging from daytime roosts 5 
miiuites before sunset. On this latter date, however, an overcast sky 
greatly reduced the amount of illumination. The earlier appearance of 
liats, in relation to sunset, in the autumn in the semiarid region where 
these observations were made may have been due to the scarcity of in- 
sect food at this season. Insects are less numerous in late September 
than in the middle of summer and less active after dark because of the 
cold. Certain other essentially crepuscular species of mammals, such as 
rabbits (Orr, 1940) are known to begin feeding activities much earlier 
in relation to sunset in winter when food is scarce than in summer when 
there is an abundance of palatable vegetation. Among diurnal birds 
Hinde (1952, p. 189) has shown that the great tit (Parus major) and 
the great spotted woodpecker (Dendrocopos major) roost later, relative to 
sunset, and emerge earlier, relative to sunrise, in midwinter than in 
autumn or spring. 

Griffin and Welsh (1937) have shown that several species of insec- 
tivorous bats studied possess a definite activity rhythm, such as occurs 
in many other kinds of animals, and suggest (p. 342) that "Perhaps 
when the environment is constant, it is this internal mechanism that 
stimulates the animal at 24-hour intervals." No doubt this accounts for 
the rather uniform time of appearance of bats of the same species in the 
same region on a given date although seasonal changes in environ- 
mental conditions relating to food, temperature, and i-elative length of 
day and night undoubtedly affect the daily activity of bats in tem- 
perate regions. 

On June 8, 1948, Mr. Eben McMillan, Mr. Ian IVIcMillan, the writer, 
and his wife camped near a pallid bat colony along San Juan Creek in 


San Luis Obispo County. From sunset that evening until after daybreak 
the following morning the entrances to the daytime roost and an adja- 
cent night roost were kept under observation constantly by one or more 
members of the party. This was the second colony found on June 8, 
1948, and has already been described on p. 181. At 7:30 p.m. a num- 
ber of Mexican free-tailed bats were seen flying in and out of the cave 
in which the roost was located. The first pallid bat emerged at 8:00 
P.M. During the next 15 minutes 4 more pallid bats emerged and 
during the next 5 minutes between 15 and 20 individuals made their 
exits. In each instance the lower opening of the cave was used. Each 
bat appeared to void urine a moment or so after emerging. On six occa- 
sions urine fell on a 10 b}^ 12-foot tarpaulin located about 10 yards 
from the entrance to the cave and directly beneath the line of flight 
taken by the bats. After emerging from the cave each bat flew directly 
to the west about 50 yards where the ground level dropped abruptly 
about 20 feet to the creek. The bats upon reaching this point also 
dropped so that they were out of the observer's line of vision. It was 
suspected that the bats went directly to the creek to secure water but 
this was not determined. By 8:30 p.m., at which time it was nearly 
dark, some pallid bats were observed entering the night roost, which 
consisted of a shallow cavern in a cliff a few yards to the south of the 
daytime roost. During the remainder of the night bats were noted en- 
tering or leaving the night roost periodically. Sometimes quite a few 
individuals seemed to come at one time, sometimes they arrived singly 
in fairly rapid succession, and occasionally none would be noted for 
some minutes. Individuals approaching the night roost regularly ut- 
tered the directive call. Pallid bats were seen to enter the cave in which 
the daytime roost was located on only a very few occasions. At 11 :00 
p.m. 2 bats were found hanging from the roof of a small cavern about 
30 feet north of the daytime retreat. At 12:30 a.m. about 8 or 10 pal- 
lid bats entered the main night roost and remained there until an ob- 
server approached with a flashlight at 1 :10 a.m. All but two tlew out. 
These remaining two were hanging from the roof of the cavern. 

Between 1 :57 a.m. and 3 :00 a.m. an observer was stationed at the 
entrance to this night roost. A flashlight was occasionally used and 
there is little doubt that the presence of the observer disturbed the 
bats. However, a record was kept of the time and number of l)ats, 
whenever it was possible to determine the latter, that entered, as follows: 
1:57 A.M., 1; 1:59 a.m., group; 2:01 a.m., 1; 2:02 a.m., 2; 2:04 a.m., 
3; 2:07 a.m., 1; 2:09 A.M., group; 2:11 a.m., 3; 2:12 a.m., 1; 2:13 
A.M., 2; 2:18 A.M., 2; 2:23 A.M., 2; 2:24 A.M., 1; 2:28 A.M., 12; 2:29 
a.m., 2; 2:31 A.M., large group; observer left for IjO minutes to ex- 
amine daytime roost and other night roost; 2:43 a.M(.,-1; 2:45 a.m., 1; 


2:46 A.M., 1; 2:50 A.M., 1; 2:52 A.M., 2; 2:53 A.M., 1; 3:00 A.M., 1. It be- 
gan to get light at 3 :00 a.m. and from then on the number of bats seen de- 
creased. The last pallid bat was seen flying near the entrance at 3 :53 a.m. 
at which time it was fairly light. Investigation of the daytime roost at 
sunrise showed that it was deserted. Apparently the proximity of human 
beings during the night so disturbed the bats that they moved to an 
alternative roost, although the latter was not found by the observers. 

The behavior pattern of the bats at this night roost conformed in 
general to that observed at other niglit roosts. Indi\'iduals came both 
singly and in groups. Frequently they would enter and seem to flutter 
momentarily against or hover near the wall of the cavern and then 
fly out. Occasionally individuals would alight. The time that they re- 
mained hanging varied from a few seconds to 40 minutes. It is prob- 
able that they often remain in a night roost for a considerably greater 
period of time than 40 minutes if undisturbed. Two other night roosts 
in natural caves were observed at night in addition to the roost ob- 
served on the night of June 8 and early morning of June 9. In most 
instances the bats observed hanging in these roosts were suspended by 
their feet from the center of the ceiling rather than hanging on the 
walls of the caves. As noted previously, however, this was not always 
true of bats using man-made structures for night roosts. Every pallid 
bat observed in a night roost was seen to be very alert and ready to 
take flight if alarmed. Nevertlieless, by approaching slowly with a flash- 
light it was often possible to come within several feet of some individuals. 


In captive males enlargement of the gonads became noticeable toward 
the latter part of August. This was accompanied by a marked increase 
in the size of the cremaster sack which had been hardly apparent dur- 
ing May, June, and July. No pallid bats were examined in the field 
in August. Enlargement of the testes was evident in both captive males 
and those examined in the field in September. By the middle of Octo- 
ber, however, there was a marked decrease in the size of the testes of 
both captive and wild individuals. Weekly examinations of four caged 
males from the middle of October, 1947, to the end of April, 1948, 
showed a steady decrease in the size of the cremaster sack. At the 
end of this time it presented tlie appearance of two darkly pigmented 

As has been noted by others who have studied the reproductive 
cycle of vespertilionid bats in the northern hemisphere, this decrease 
in the size of the cremaster sack in the fall is associated with a de- 


crease in the size of the testes rather than the epididymes. As long 
ago as 1879 Fries described the prolonged retention of sperm in the 
epididymis of certain Enropean bats of the families Vespertilionidae 
and Rhinolophidae and also noted that the accessory male reproductive 
glands were functional from fall until spring. Similar observations 
were made by Rollinat and Trouessart (1896). The first detailed his- 
tological studies of the male rei)roductive tract in the Microchiroptera 
were made by Courrier (1927) on Pipistrelhis pipistreUus. Courrier 
not only confirmed the conclusions of earlier workers but also found 
that spermatogenesis occurs during the summer months. Miller (1939), 
working on Myotis lucifugus and Myotis grisescens in this country, 
found a similar seasonal sequence in the reproductive cycles of males 
of these species as have Pearson, Koford, and Pearson (1952) in Cory- 
n orh inns rafin esqnci. 

Although the gonads of male pallid bats have attained maximum 
size by late August and September, no evidence was obtained to indi- 
cate that copulation occurs before the latter part of October, by which 
time a definite decrease in size of the gonads has occurred. Vaginal 
smears from 29 females taken from a colony containing members of both 
sexes, in eastern San Luis Obispo County, on September 20, 1947, failed 
to reveal the presence of any sperm. Since no uterine examinations were 
made of these bats this cannot be considered as conclusive evidence that 
no copulation had occurred. However, the absence of spermatozoa in 
smears from such a large number of females makes it seem unlikely 
that many, if any, had been inseminated. On October 22, 1949, vaginal 
smears were taken from 7 females captured in the same region. Spermatozoa 
were found present in 2 of the females. 

The spermatozoon of this si)ecies possesses a rather elongate, oval- 
shaped head which is truncate posteriorly. The body or connecting 
piece is of but slightly lesser diameter than the head. Its length is 
nearly half of the total length of the spermatozoon. Posteriorly it 
tapers to its junction with the tail piece. The total length of an average 
spermatozoon, measured wdth an ocular micrometer, was 36//,. 

A number of observations were made on the breeding activities of 
pallid bats in captivity. On September 25, 1947, 3 males and 3 females 
that had been captured 5 days previously were placed together in an 
indoor laboratory flight cage where the temperature ranged from 50°F. 
to 65 °F. These bats, as well as others that were later placed in this 
cage, were banded so that the activities of each individual could be 
followed. A few minutes were devoted to observation each morning 
and again in the afternoon when the bats were fed. Two of the males 
failed to show anv signs of sexual activity during the succeeding six 


months. These may have been young of the year althougli this could 
not be determined on the basis of external characters. Pearson, Ko- 
ford, and Pearson (1952) have sliown that it is improbable that the 
males of Corynorhinus rafinesquei copulate the first year. The third male 
was noted copulating with the same female on November 3, 18, 20, and 
28. He was found copulating with the second female on November 9, 
and a vaginal smear taken from the third female on November 11 
showed the presence of spermatozoa. The first female was permanently re- 
moved from the cage on December 5. The second and third females 
were removed on December 29 and replaced by two other females. On 
January 16, 1948, however, the second female was returned to the cage. 
The male was found copulating again with her on February 6, 13, and 
16. She was removed to a separate cage on February 25, and on April 
24 aborted two embryos. A vaginal smear taken on February 24 from 
one of the females placed in the breeding cage on December 29 re- 
vealed an abundance of spermatozoa, indicative of recent copulation. No 
signs of copulation were observed after this date. 

Captive bats were observed to copulate both on the floor of the 
cage and while hanging upside down on the vertical screen walls. On all 
occasions the male lay over the back of the female but was never seen 
holding her with his mouth. The uropatagium of the female was pushed 
to one side at such times. All copulations were observed during daylight 
hours. No vaginal plug was ever found in the female although a jelly- 
like substance, rich in spermatozoa, was found in the vagina imme- 
diately after copulation. 

To determine what effect out-of-door temperature might have upon 
the breeding activities of pallid bats in San Francisco, 3 females and 
a male were placed in an outdoor flight cage on September 25, 1947. 
These bats, like those kept in the indoor flight cage, had been captured 
5 days earlier in eastern San Luis Obispo County. One of the females 
was observed copulating with the male on October 31, and on Novem- 
ber 4, 5, 6, 10, 13, 14, 17, 18, 19, and 24. There were no indications 
that either of the other 2 females bred. A fourth female was placed 
in the cage on November 1 and was observed copulating with the male 
on November 12. No sexual activity was observed during December 
and on the 29th of that month the 4 females were removed and 3 other 
females were placed in with the male. There was no further indication 
of breeding, however, on the part of these bats during the succeeding 
months. The air temperatures, in degrees Fahrenheit, for San Francisco 
as compiled by the United States Department of Commerce, Weather 
Bureau, in the Monthly Weather Review, for October, 1947, to February, 
1948, are shown as follows: 



























Summarizing these observations on the breeding of pallid bats in 
captivity, therefore, it would appear that those maintained in the in- 
door laboratory cage, where the extreme ranges in daily temperature 
were from 50°F. to 65°F., copulated during the months of November, 
December, January, and February. Bats kept in an outdoor flight cage 
in San Francisco bred only from the end of October to the end of No- 
vember. The mean minimum temperatures for these two months were 
56°F. and 49°F., respectively. There was no indication of breeding during 
December, January, and February when the mean minimum temperatures 
were 46°F., 48°F., and 45°F., respectively. None of the captive bats 
kept in this outside flight cage went into dormancy in the usual sense 
of the word as applied to colonial bats. They ate regularly during the 
winter and, on all except the coldest nights, appeared to be active. 

Field observations failed to show that pallid bats breed in Septem- 
ber. There is some evidence (cf. Seasonal Behavior) that the sum- 
mer colonies break up in October. During this month and up until 
the middle of November these bats were found in smaller aggregations, 
in one instance just a pair, and in places where they were not observed 
in summer. Females with spermatozoa in the vaginal passage were examined 
on October 22. On the basis of these few facts, combined with those 
obtained from captive pallid bats kept in an outdoor cage in San 
Francisco, it would appear that in central California this species breeds 
at least in the latter part of October and in November. 

Very little is known regarding the whereabouts, and nothing per- 
taining to the sexual behavior, of pallid bats in the wild during the 
months of December, January, February, and March. Certain North 
American species of colonial, vespertilionid bats that have been studied 
in winter have been found to show signs of periodic activity during 
the season of dormancy. This, as has been noted, was also true of cap- 
tive pallid bats kept without food at temperatures ranging from 38 °F. 
to 50°F. Possibly during these brief periods of activity copulation oc- 
casionally occurs. Pearson, Koford, and Pearson (1952) have shown 


that in Corynorhinus rafinesquei in California copulation occurs mostly 
during the winter months. These authors also indicate that the males 
are much more active that the females at this season and probably 
copulate with the females while the latter are torpid. 

The literature on this subject has been carefully summed up by 
Ilartman (1933), Baker and Bird (1936), Wimsatt (1945), and others. 
With most vespertilionid bats occurring in the Northern Hemisphere 
copulation takes place principally in the fall of the year prior to hiber- 
nation. There may or may not be occasional copulations by bats aroused 
from dormancy during the winter, and again in spring just prior to or 
about the time of ovulation. Miniopteriis schreihersn, which was studied 
by Courrier (1927) in France is, of course, an exception to this rule. In 
this species copulation and ovulation both occur in the fall. Fertiliza- 
tion takes place immediately and pregnancy lasts over the winter until 
spring. Miniopterus australis, studied by Baker and Bird (1936) in the 
New Hebrides, breeds in the spring at which time fertilization occurs. 
Miniopterus is considered exceptional in so far as the family Vesper- 
til ionidae is concerned. Bats of the family Rhinolophidae behave essen- 
tially like most of the Vespertilionidae in that they copulate in the fall 
of the year and ovulate in the spring. On the other hand, phyllosto- 
matid bats (Hamlett, 1935) and molossid bats (Hartman and Cuyler, 
1927; Sherman, 1937) that have been studied, copulate in the spring 
at the time of ovulation. 

During this study an attempt was made to determine whether or 
not spring or late winter insemination is necessary, both for normal 
development and for fertilization, in the pallid bat. Guthrie and Jeffers 
(1938) induced ovulation in Myotis lucifugus in December and January 
by injections of hypophyseal extract and found that normal development 
did not follow, whereas ovulations induced in February were followed 
by normal cleavage and the formation of blastodermic vesicles. These 
authors, as well as Miller (1939) in his paper on the reproductive cycle 
of male bats, concluded that this difference in the activation reaction 
was conditioned by the introduction of fresh sperm during the late 
winter copulations. Pearson, Koford, and Pearson (1952, p. 299) con- 
clude that in Corynorhinus rafinesquei "there is a refractory period dur- 
ing autumn and early winter during which ovulation does not normally 
occur. . . ." They further conclude "that ovulation is not readily in- 
duced by removing Corynorhinus from hibernation to warm rooms before 
mid-February, but that injections of pregnancy urine bring about ovu- 
lation at this time." 

On December 5, 1947, a female pallid bat that had been captured on 
September 20, and was known to have copulated repeatedly during 
November, was removed from a cool room in which she had been 


caged with a male and ])laeed alone in a cage in a heated greenhouse 
where the temperature ranged from 75 °F. to 78 °F. It was hoped that 
this would induce ovulation. On February 26, 1948, she gave birth to 
one young which was reared successfully. The total elapsed time from 
the date on which the female had been placed in the warm room until 
parturition occurred was 83 days. Although the time of ovulation 
was not known, it is probable that some days elapsed between the time 
the bat was placed in a warm environment and the time of ovulation. 
Normal development apparently occurred in this instance in midwinter. 

Wimsatt (1944) has shown that fall and early winter insemination 
can effect fertilization in the spring in Myotis lucifugus and Eptesicus 
fuscus. Eyberg (1947), apparently unaware of Wimsatt 's (1944) work, 
mentions two instances in which females of Nyctalus noctuJa were cap- 
tured in the fall after hibernation had commenced and kept cool and iso- 
lated during the winter. He states (p. 149) that "Long after they had 
awakened in the spring, they began, gradually, without any possibility 
of copulating again, to show signs of pregnancy." 

To determine whether or not late winter or spring insemination is 
necessary for fertilization in the pallid bat, a female that was known 
to have copulated on November 12, 1947, and may have copulated sub- 
sequently, was isolated from all males in an outdoor flight cage on 
December 29, 1947. Sometime between the night of May 21 and the 
morning of May 24, 1948, she gave birth to one young. The young was 
dead when found on May 24, but appeared normal as regards develop- 
ment. Presumably, as often happens in captivity, the female deserted 
the young at birth and the latter died from neglect. Birth in this 
instance occurred nearly five months after the female ^\as isolated from 
any males. As has already been indicated and will be shown subse- 
quently, this greatly exceeds the gestation period of this species. 

Vaginal smears were taken approximately once a week from 12 cap- 
tive females, between December 4, 1947, and June 4, 1948, to determine 
whether or not any epithelial changes could be observed and corre- 
lated with the reproductive cycle. Between June and December, 1948, 
smears were taken once or twice a month. Small, sterile, rubber spatu- 
las were used and each smear was diluted with a drop of Ringer's 
solution on a slide. The slides were examined with an 8 mm. objective 
and X7 ocular. Some of the bats used in the study were kept out-of- 
doors, others in a warm laboratory, and others in a refrigerated room 
during the winter and early spring of 1947-48, to see if environmental 
temperature had any effect. 

In general it Avas found that squamous epithelial cells were present 
all year around. Although occasional cornified cells were noted during 
all months of the year, they were found in abundance from the end of 


January until early in May. The greatest number of cornified cells, in 
proportion to nucleated epithelial cells, were observed in smears from 
bats kept at lower environmental temperatures. Leucocytes appeared 
in March and April, varying in time of appearance with the individual. 
There was no continuous invasion of the vagina by leucocytes during 
these months but rather a sudden rise and fall in the count. It was sus- 
pected that the appearance of leucocytes in numbers might be asso- 
ciated with ovulation, but this was not determined. An occasional leu- 
cocyte was found in early May, after which not one was seen. Guthrie 
(1933) noted large numbers of leucocytes in the vaginal passage in 
Myotis lucifugus at the time of ovulation in the spring. 

As has previously been stated, a captive female that had bred was 
removed from a moderately cool environment and placed in a warm 
room on December 5, 1947. Eighty-three days later she gave birth to 
a normal young. It was believed that the increased temperature induced 
ovulation and that the gestation period may have been about eighty 
days, possibly less, depending upon the time required for the matur- 
ing of the follicle (see Guthrie and Jeffers, 1938). In order to further 
check the gestation period, 15 female pallid bats were secured on April 
15, 1949, in eastern San Luis Obispo County. This was 5 days after 
the first appearance of bats at this colony site. They were transported 
to San Francisco and on tlie following day 4 were killed and their re- 
productive tracts fixed in Bouin's fluid. After this material was sec- 
tioned and stained it was found that one female had recently ovulated, 
possibly within 24 hours, although no tubal egg could be found. The 
ruptured follicular cavity had not as yet formed a corpus luteum. 
The three other reproductive tracts contained blastocysts in one or both 
horns of the uterus. In each instance implantation was beginning to oc- 
cur on the antimesometrial side of the uterus and the beginnings of 
chorionic villi were evident on the side of the trophoblast next to the 
inner cell mass. A thin, single-celled layer of entoderm adhered to 
the underside of the inner cell mass and the inner surface of the tro- 
phoblast of each blastocyst. In one embryo the amniotic cavity had not 
as yet made its appearance. In each of the others a small amniotic 
cavity was present. In the most advanced embryo the flattened em- 
bryonic disc was quite apparent. To state the age of these embryos is 
a difficult matter. When compared with the macaque, studied by Heuser 
and Street er (1941), which has a gestation period of 168 days, they 
are comparable to the development attained between the 9th and 12th 
days following ovulation. One would expect more rapid growth in the 
case of a mammal having a shorter gestation period. Guthrie and 
Jeffers (1938), however, found free uterine blastocysts in Myotis Juci- 
fuf/us at what was estimated to be the tenth day following ovulation 


induced by injections of hypophyseal extract. If such slow develop- 
ment occurs in vespertilionid bats under natural conditions following 
normal ovulation in the spring, the most advanced embryos found in 
Ant7'ozous females killed on April 16 might be estimated to be about 
12 days of age. This would indicate that 3 of the 4 females ovulated 
sometime during the first week of April. Two more females were killed 
on May 2. One contained a single embryo, the other contained 3 em- 
bryos, 2 in the right horn of the uterus and 1 in the left. All 4 embryos 
appeared to be at about the same stage of development, but only 1 
was removed from the foetal membranes for measurement. This in- 
dividual had a crown-rump length of 5 mm. The others were preserved 
intact in their foetal membranes for future study. On May 18, 2 
more females were killed. One contained 1, the other 2 embryos. Each 
embryo had a crown-rump measurement of 12 mm. On May 23 an- 
other female aborted 2 young. The first of these was eaten by the 
parent. The second dropped to the floor of the cage and was removed 
and preserved for later study. It measured 16 mm. from crown to 
rump. On June 1, one of the remaining 6 females showed obvious signs 
of illness and was found dead on June 6. A post-mortem examination 
indicated that she had probably died on June 4. One horn of the 
uterus contained an 18 mm. embryo. On June 7 two of the females 
gave birth to 2 normal young each. Another female produced 2 young 
between the evening of June 10 and the morning of June 13, another 
female gave birth to 1 young on June 14. The remaining female failed 
to produce young. Although she may not have been pregnant, there 
is the possibility that she aborted at some time without this fact l)eing 
observed by the writer. 

Thus, if we assume that the 4 females that gave birth to young 
between June 7 and 14 ovulated sometime between April 4 and 15, as 
was believed true of the 4 females killed on April 16, the possible 
maximum and minimum gestation periods would be 71 and 53 days 
respectively. It is reasonable to presume, however, that those that bore 
young first were the first to ovulate. This would indicate an average 
gestation period of about 9 weeks. However, Pearson, Koford, and Pear- 
son (1952) as well as Eisentraut (1937) earlier, have suggested that en- 
vironmental temperature, in so far as it affects the temperature of the 
mother, may influence the length of the gestation period in certain 
species of bats. Consequently it is possible that extended periods of 
cold weather in spring, during the time of pregnancy, may prolong tlie 
gestation period in this species. 

Wimsatt (1945) estimated the gestation period for Myotis lucifugus 
to be between 50 and 60 days. Pearson, Koford, and Pearson (1952) in- 
dicate that in Corynorhinus rafinesquei the gestation period may vary 


from 59 to 100 days, depending upon the mother's body temperature. 
Sherman's (1937) studies on Tadarida cynocephala, a molossid bat of 
the southeastern United States, led him to conclude that the gestation 
period in that species is between 11 and 12 weeks. 

Birth and Growth of Young 

Number of young per female: Records of 28 females, from San Luis 
Obispo and Kern counties, that either bore young or were pregnant (4 
captured with 2 young each, 10 gave birth to young in captivity, 14 
had embryos in utero) are as follows: 1 female had 3 embryos, 20 fe- 
males had 2 embryos or young each, 7 females had 1 embryo or young 
each. This is an average of 1.8 young per bearing female. 

Hall (1946, p. 165) indicates that uterine examinations of 13 preg- 
nant pallid bats taken in Nevada showed that 11 contained 2 embryos 
each, 1 contained 3 embryos, and 1 contained 1 embryo. Bailey (1936, p. 
392) mentions 2 females taken in a cave in Nevada, each with 2 young 
attached. Burt (1934, p. 397) records 4 females taken at Indian 
Springs, Nevada, on the nights of June 2 and 7, 1929. Three of the 
females each contained a single, well-developed embryo and the other 
female contained 2 embryos measuring 22 mm. J. Grinnell (1914, p. 
263) records a female taken along the Colorado River with 2 embryos. 
H. Grinnell (1918, pp. 352 and 356) records a female taken at San 
Bernardino, California, with 3 embryos and another taken at Painted 
Rock, San Luis Obispo County, California, with 2 embryos. Grinnell 
and Storer (1924, p. 61), on examining 19 females captured on May 
27, 1915, at Snelling, Merced County, California, found that 15 con- 
tained 2 embryos each, 3 had 1 embryo each, and one was not pregnant. 

Two young, therefore, a]^pears to be the usual number borne an- 
nually by most female pallid bats. Occasionally there may be only one 
young and rarely three. It is difficult to understand how females of 
this species could successfully rear 3 young since they only possess 2 nipples 
and the young seem to hold tightly to these for the first few days 
after birth. It is possible that in females with 3 embryos one is re- 
sorbed prior to birth, although resorption of advanced embryos has 
quite justifiably been questioned by Ryberg (1947, p. 139). 

Dates of birth: The time at which the young are born varies with 
individual females in the same colony and seemingly to some extent 
with locality and year. 

On May 10, 1947, 7 female pallid bats were captured 2i^ miles 
northwest of Carneros Spring in Kern County. Two had newly born 
young clinging to them and the remaining 5 were pregnant. The latter 
had all given birth to young in captivity by May 14. These were the 


earliest seasonal field records for young obtained during this study. 
The region where these bats were found, however, was the hottest and 
most arid part of the entire area where observations were carried on. 

On June 6, 1948, 10 pregnant females were captured at random 
from a colony 4i/2 miles northeast of Shandon, San Luis Obispo County. 
This locality was about 27 miles northwest of the site of the colony 
previously mentioned and was in a region that may be described as 
cooler and less arid in summer. Two of the females were prepared as 
study skins and each was found to contain 2 nearly full-term foetuses. 
The latter averaged 45.5 (45-46) mm. in total length and 2.55 (2.5-2.6) 
grams in weight. Four of the females were released 2 days later and 
the remaining 4 gave birth to young in captivity between June 14 and 
23. On the evening of the same day the 10 pregnant females were cap- 
tured, a lactating female pallid bat was shot near Carneros Spring in 
Kern County. It was thought that she had probably borne young sev- 
eral weeks previously. On the following day, June 7, 1948, 35 female 
pallid bats were taken from a colony on the northwestern edge of the 
Carrizo Plain, San Luis Obispo County. All were palpated before 
being released and were found to be in advanced stages of pregnancy. 

Four females captured at the colony 4I/2 miles northeast of Shan- 
don on April 15, 1949, gave birth to young in captivity between June 
7 and 14. Two pregnant females were captured at this colony on April 
14, 1951. One was killed on April 17 and found to contain 2 embryos 
with a crown-rump length of 7 mm. which indicated a much earlier 
ovulation date than was suspected for females taken from this same 
colony on April 16, 1949 (cf. table II). The second female was killed 
on May 25 and found to have 2 foetuses about ready for birth. 

From the foregoing evidence, therefore, there is some indication 
that the time of parturition in eastern San Luis Obispo and Kern coun- 
ties varies from year to year within the same colony. It is quite pos- 
sible that early warm spring weather may be conducive to early sea- 
sonal appearance and ovulation. Within any one colony in this region 
such data as were gathered indicate that, in any one year, all the fe- 
males bear their young within a period of several weeks. 

On June 13, 1951, 10 young were secured from a colony of pallid 
l)ats inhabiting an attic at Stanford University, Santa Clara County, 
California. The smallest individual was about half grown and unable 
to fly. Its age was estimated at about 3^2 weeks. Tliree others were 
about three-fourths grown but not quite able to fly. The remaining 6 
had body measurements within the range of adults and could fly well. 
The oldest was believed to be about 6 weeks of age and probably was 
born the first week in May. 

Regarding reproduction of this .species in Nevada, Ilall (1946, p. 


165) may be quoted as follows: "It was noted that the embryos taken 
on May 17 measured only 3 mm. from crown to rump. The female 
taken on June 4, 6 miles south of St. Thomas, had two embryos nearly 
ready for birth, and the embryos found on June 7 at Indian Springs 
(Burt, 1934:397) also were nearly full term. On June 15, 1939, in 
crevices in the roof of a cave 10 miles west and 5 miles south of 
Fallon, Alcorn (AIS) found about fifty females nearly every one of 
which had one or two young securely fastened to her mammae. Bailey 
(1936:392) found a colony of females wnth young in the same cave on 
June 27, 1927. The young of one female, he thought, were but one day 
old. Burt (1934:397) mentions young bats nearly full grown taken at 
Indian Springs on August 2 and 3; our specimen taken August 7, west 
of Lahontan Dam, is of similar age. It appears that young ordinarily 
are born in the first half of June." 

Sex ratio of young: Fifty young comprising 25 taken at random 
from colonies in the wild, 21 born in captivity, and 4 removed from 
the uteri of pregnant females just prior to birth, were carefully ex- 
amined to determine sex. Twenty-seven were males and 23 were fe- 
males. These figures seem to indicate a nearly equal sex ratio at birth. 

Parturition: The relatively meager literature on parturition in the 
Chiroptera has been fairly well summed up by Ramakrishna (1950). 
In general, breech presentation is characteristic of most of the species 
of Microchiroptcra in which birth has been observed, exceptions hav- 
ing been noted in the case of NyctaJus noctula by Daniel! (1834) and 
Hipposidcros speoris by Ramakrishna (1950). Observations made on a 
number of captive pallid bats giving birth to young indicate that par- 
turition in this species conforms in general to the pattern thought to 
be characteristic of the suborder. 

During the period of lal)or and birth captive females hung upright. 
The uropatagium was curled ventrally at such times to form a basket 
and thus prevent the young from falling to the floor when born. The 
period of labor vai'ied from a few minutes to nearly two hours. Fe- 
males respired rapidly when in labor and frequently uttered notes of 
pain during contractions. When females had 2 young, the time re- 
corded between the births varied from 12 to 65 minutes. All births 
observed were by breech. The rump and tail of the young, with uro- 
patagium folded over the ventral part of the body, emerged first. As 
soon as the hind feet of the young were free they seemed to be used to 
further assist emergence by pushing against the mother's body. On 
several occasions a female was seen to sever the umbilical cord by bit- 
ing it and later to eat the placenta when it was expelled. In most in- 
stances, however, the females were not under observation at times when 
this occurred. If the placenta was not found in the cage the following 



[Proc. 4th Ser. 

day it was believed that the female had eaten it. Females that did 
not consume the placentae refused to accept their young and either 
killed the latter by biting them or else let them fall to the floor of 
the cage. The refusal of some captive females to accept their young 
was thought to be an abnormal condition associated with confinement 
in an artificial environment. 

Females that behaved in what might be termed a normal manner 
licked their young frequently during the process of birth and then 

Figure 24. Female pallid bat with young 2 days old. Photographed by Lionel 
T. Berryhill, May 15, 1947. 

licked them thoroughly immediately after birth. One mother was seen 
to extend the wings of her newborn young with her mouth so that she 
could lick the membranes. Newly born bats seemed capable of climb- 
ing of their own accord to the mother's mammary glands. They used 
their thumbs, hind feet, and mouths to hold on to their mother's fur and 
pull themselves up. The female usually would assist by nuzzling the 
young. After a young bat became attached to its mother's nipple the 
female would envelop the young with her plagiopatagium so that the 
baby could no longer be seen. The young oriented itself in the same 
direction as the parent. Consequently, if she hung upside down the 
offspring did likewise. 

Since young bats hold on very tightly to their mother's nipples (fig. 


24) it is often very difficult to remove them to take body measure- 
ments. Furthermore, distur])ance of this sort occasionally results in a 
female refusing to accept her young. Consequently, few newborn young 
were measured in captivity. However, the following selected measure- 
ments were made of two seemingly healthy young males a few hours after 
birth (figures in parentheses represent percentages of corresponding 
average adult male measurements): Total length, 48, 50 (42 per cent); 
tail length, 13, 15 (37 per cent) ; hind foot, 9, 10 (71 per cent) ; ear 
from notch, 8, 9 (30 per cent); forearm, 17, 18 (31 per cent). Their 
weights were 2.9 and 3.3 grams. 

Develoijment of young: At the time of birth young pallid bats 
have their eyes closed and their ears folded down tightly against the 
sides of the head. The skin of the body and the appendages is pink 
and to the unaided eye appears naked. When examined under a low- 
power microscope a few hairs will be found on the following parts: 
on eitlier side of the head extending as a band from behind the rhina- 
rium almost to the base of the antitragus, along the margin of the 
lower jaw, on the flight membranes immediately dorsal to the humerus 
and femur, and on the underside of the distal phalanges of the toes. 

The information relating to growth of young given in the follow- 
ing paragraphs is based upon bats born and reared in captivity, unless 
otherwise stated (figs. 25 to 28). It is quite possible that under such 
circumstances the normal rate of growth and development may have 
been affected. A number of young developed malformations of the 
])ones before maturity, presumably as a result of dietary deficiencies. 
Others seemed fairly normal. 

By the fourth day of postnatal life the regions on which minute 
liairs could be detected had increased considerably so as to include an 
area extending from the tip of the tail anteriorly to tlie posterior lum- 
1)ar region, then forward on either .side of the body to the posterior 
thoracic area. Hairs also occurred on the dorsal surface of the legs and 
toes as well as on the wings from the thumb to the shoulder. These 
hairs could barely be seen by the naked eye at this time. By about the 
tenth day scanty fur was clearly evident on most of tlie upper parts of 
the body, excluding the area between the shoulders. Heaviest pelage 
was present on the lower back although the longest hairs were to be 
found on the posterior sides of the lower jaw. 

The eyes, which were closed at time of birth, opened between the 
eighth and tenth days. The ears, although capable of being raised 
slightly by the third or fourth day, were not held erect until about 
the time the eyes opened. The tragus was pressed against the main 
])ody of the pinna thus closing the external auditory canal until this 



Figure 25. Dorsal view of a 3-day-old male pallid bat. Photographed by Lionel 
T. Berryhill, May 15, 1947. 


ll ' 21 ^ 3i l 


•> H? II 12 

Figure 26. Ventral view of a 3-day-old male pallid bat. Photographed by 
Lionel T. Berryhill, May 15, 1947. 





Figure 27. Dorsal view of a female pallid bat 30 days old. Photographed by 
Lionel T. Berryhill, June 9, 1947. 




•■• ._ij;WLi'S'j»iii''iK?-«i' 

Figure 28. Ventral view of a female pallid bat 30 days old. Photographed 
by Lionel T. Berryhill, June 9, 1947. 


Captive young were observed to exhibit fear when 10 days old. 
When a hand was placed near a young bat at this age it would draw 
back and utter an intimidation buzz. 

Whether or not females in the wild carry their young with them 
when they forage at night was not determined. On several occasions 
females that were disturbed at their daytime roosts were observed fly- 
ing away with their young attached. Griffin (1940, p. 182) states that 
"For the first three or four days after birth young of Myotis lucifugus 
are usually carried by the mother as she hunts." By the time young 
captive pallid bats were two weeks old they ceased hanging onto the 
mother during the daytime. The parent, however, kept the young be- 
neath the plagiopatagium until they were too large for this arrange- 
ment and were forced to hang beside her. 

By the eighteenth day a short, scanty pelage was seen on all of the 
upper parts of the body, except for the intershoulder area, and to a 
limited extent on the ventral parts. The skin, which was pink during 
the first week, had gradually darkened on the body wherever hair was 
present. The flight membranes had become somewhat grayish with the 
darkest coloration evident along the top of the tail and dorsal parts 
of the legs. 

By the time the young were 24 days old the fur was sufficiently 
dense and long to obscure the skin except between the shoulders, a 
region relatively bare even in adults. At this age the general color of 
the dorsal pelage was much darker than that of adults. This resulted 
largely from the fact that only the darkly pigmented distal parts of 
the hairs had made their appearance and the light basal portions had 
not as yet grown out. The fur on the ventral parts of the body was a 
pale cream-buff, somewhat like that of the adult but with a more gray- 
ish east. The flight membranes appeared vinaceous. 

By the thirty-fourth day the light basal parts of the hairs of the 
dorsal parts of the body were evident. The young still appeared much 
darker than adults. Apart from the intershoulder area, which acquired 
a scanty growth of hair at 2i/^ months, pelage changes after the flrst 
month seemed only to involve increase in length of hairs. 

The earliest age that a captive-born young was observed to fly was 
7 weeks. However, wild-taken young whose measurements were ap- 
proximately similar to captive young 6 weeks old were capable of fly- 
ing quite well. It is significant perhaps in this regard to note that 
captive young began eating some mealworms by the time they were 6 
weeks old, although they continued to nurse for several more weeks 
until the female would finally discourage this by keeping away from 
them or by threatening them when they came close to her. 

The pelage of young when they have attained adult size is indis- 


tinguishable from that of adults in new pelage. Many pallid bats 
were examined in the field in September and October. It seems highly- 
probable that some of these must have been young of the year. If 
so, however, they could not be distinguished from adults at this sea- 
son on the basis of external characters. 

Tooth replacement: There is considerable variation in the deciduous 
dentition of the Chiroptera. Some bats, such as members of the family 
Rhinolophidae, possess deciduous teeth only in embryonic life. These 
teeth are reportedly absorbed prior to birth. Others retain their de- 
ciduous teeth for some weeks of postnatal life. In certain groups these 
teeth have the form of simple, straight spicules. In others they may be 
noticeably recurved. In some members of the family Vespertilionidae 
the deciduous teeth are not only recurved but may possess one or two 
accessory cusps. 

The complete number of deciduous teeth in the pallid bat is 22, 
represented by the formula: i f, c |, pm f. As pointed out by Miller 
(1907, p. 20) this represents the maximum number known to occur in 
the order and is characteristic of a number of species of vespertilionids. 
In studying the sequence of appearance of the permanent teeth in this 
species the following material was used : 14 alcoholic specimens, rang- 
ing from foetuses nearly ready for birth to young 8 days old; 7 captive- 
born young whose dental development was followed from birth until the 
permanent dentition w^as acquired; 7 skulls of young whose approximate 
ages were determined on the basis of information derived from a study 
of young bats born and reared in captivity. 

An examination of nearly full-term foetuses indicates that the first 
deciduous teeth to erupt through the gums are the upper and lower 
canines. Shortly following this the upper and lower incisors erupt, al- 
though in some specimens the first pair of upper incisors were some- 
what slower than the others in appearing. This in general is the condi- 
tion at time of birth, although Barrett-Hamilton (1910, p. 14) states 
that young bats, with the exception of members of the family Rhino- 
lophidae, are born with complete deciduous dentition. The upper and 
lower premolars in Antrozous usually do not make their appearance 
until the third to the fifth day of postnatal life. In several newborn 
young, however, these teeth were in the process of breaking through 
the gums. By the early part of the second week the deciduous teeth 
are about fully grown out. 

There is considerable individual variation in the shape of these teeth, 
especially the canines and premolars. The crowns of the upper and 
lower incisors possess a definitely trifid pattern with a main central cusp 
and two smaller accessory cusps, one on either side. The canines, both 
upper and lower, may vary from a condition in which three distinct 



cusps are present to one in which only two cusps are faintly indicated. 
The upper premolars may be trifid, bifid, or have the crown more or 
less truncate in shape. The last form is most often seen in pm -. Pm j 
is generally trifid while pm ^ varies like the upper premolars. The 
cusps, which are shaped like small booklets, are curved inward. This 
is most pronounced, however, on the upper and lower incisors. 

The deciduous dentition has long been considered to represent a 
primitive type of dentition. This was the contention of Winge (1941) 
and is a view generally held by mammalogists. However, with respect 
to the clasping type of deciduous teeth found in certain bats one must 
concur with Spillman (1927, p. 251) who describes this as a remark- 
able specialization not found in any other living or fossil groups of 
mammals. Such teeth unquestionably enable the young to hold tightly 

Table III 
Sequence of loss of deciduous teeth in Antrozous palUdus. 

Age at 

Age at 

Upper teeth 

ivhich lost 

Loioer teeth 

which lost 


5 weeks 


2-3 weeks 


5-6 weeks 


2-3 weeks 


6-8 weeks 


4 weeks 


6-8 weeks 


4 weeks 


8-10 weeks 


5-8 weeks 
8-9 weeks 

Table IV 
Order of appearance of permanent teeth in Antrozous pallidus. 

Upper teeth 

Age at time 
of eruption 

Lotver teeth 

Age at time 
of eruption 


4 weeks 


4 weeks 


4 weeks 


5 weeks 


5 weeks 


5 weeks 


3 weeks 


4 weeks 


4 weeks 


4 weeks 


4 weeks 


4 weeks 


5 weeks 


5 weeks 


to the mother's nipple when she is in flight. It is very difficult to re- 
move young from the mother during the first week or two following 
birth without injuring the mouth of the former or the mammary gland 
of the latter. 

The permanent dental formula for the pallid bat is: \ ^, c \, pm ^, 
m 1 = 28. These teeth erupt through the gums internal to the deciduous 
teeth and for the most part their appearance precedes the loss of the 
deciduous teeth (tables III and IV). Certain of the latter teeth are 
present for some time after the young are weaned. 


Since pallid bats do not emerge from their daytime retreats until 
dusk it is difficult to make observations on their foraging habits. Some 
observers report that these bats go directly to water after taking to 
the air in the evening. Bailey (1936, p. 391), referring to members 
of this species, says: "At early dusk they begin to come out and 
usually fly directly to the nearest still water wliere they drink by re- 
peatedly dipping to the surface, then begin their search for insect 
food." Nelson (1918, p. 493) makes the following statement regarding 
pallid bats: "Often when camping at desert waterholes, I have seen 
them come in just before dark to drink, scooping up water from the 
surface while in flight, and then circling back and forth over the 
damp ground at an elevation of a few yards for the capture of some 
of the insects common in such places." 

Whether or not drinking water is essential for pallid bats is not 
known. However, only one of the summering colonies located during 
the course of this study was situated more than half a mile from 
known available fresh water. In this instance there was a spring 21/0 
miles away and it is possible that there may have been sources of 
water nearer. In cajitivity bats regularly drank water. 

When foraging, members of this species generally fly fairly low and, 
as has long been known, frequently alight on the ground to capture 
food. Burt (1934, pp. 397-398) observed these bats usually flying 3 or 
4 feet above the ground when foraging. He also records them fre- 
quently alighting on a lawn to capture June beetles (Polyphylla) in 
front of a ranch house at Indian Springs in southern Nevada during 
the summer. Nelson (1918, p. 493) reported the capture of a pallid 
bat that lit on the ground in an apple orchard in northern Arizona. 
The bat had caught a Jerusalem cricket (Stenopelmatus fuscus) and 
was so engrossed in its prey that it was readily picked up by the nape 
of the neck. Huey (1936, p. 285) records 2 pallid bats caught in mouse 
traps set for desert rodents in Mono County, California, on August 2, 


1932. Since maii}^ insects, especially pinacate beetles and desert crickets, 
had been observed coming to the bait the night before it was suspected 
that the bats had come to feed on the insects. 

Nearly every pile of droppings beneath the day and night roosts 
of pallid bats examined during this study contained numerous remains 
of Jerusalem crickets which, of course, are flightless and must have been 
captured on the ground. The remains of scorpions were also frequently 
found in such places. 

Mr. Eben Mc^Iillan, who contributed so much to this study, placed 
some clean grain sacks beneath some grain tanks wliere pallid bats 
were noted hanging up at night on the Pinole Rancli on the Carrizo 
Plain in San Luis Obisjio County, on October 22, 1949. Each morning 
during the succeeding week the sacks were examined and then cleaned. 
On each of the first six mornings fresh remains of Jerusalem crickets 
were found beneath the roost. 

On October 23, 1949, the writer found numerous insect remains be- 
neath a pallid bat roost along San Juan Creek, 9 miles west of Simm- 
ler, San Luis Obispo County. Some of these were saved and later 
identified as parts of the following insects : Orthoptera : Stenopehnatus 
probably fuscus: Coleoptera : PolyphyUa probably decinilineata, Roma- 
leuni simplicicoUe, Prionus californicus, Eleodes acuticauda. 

Mrs. Grinnell (1918, p. 355) records the remains of Jerusalem 
crickets and sphinx moths as being most numerous during the summer 
on the floor of a barn loft occupied by pallid bats at (llendora, Los 
Angeles County, California. A sample of insect remains dropped by 
these bats in this same loft in September contained parts of the fol- 
lowing: Orthoptera: Stenopelmatus sp., Microcentrum sp., Gryllus sp.; 
Lepidoptera: Deilepkila [Celerio] lineata; Coleoptera: Prionus californi- 
cus, Ligyrus gihhosus. 

Hatt (1923, p. 261) lists the arthropod remains found with drop- 
pings beneath a pallid bat roost in September at jNIission San Antonio 
de Padua, Monterey County, California. Most of these were Jerusalem 
crickets (Stenopelmatus fuscus) and scorpions (Anuroctonus phaiodac- 
tylus). A few other insect fragments identified included a grasshopper 
(Schistocerca sp.) and a beetle (PolyphyUa decinilineata) . 

Borell (1942, p. 337) records the remains of grasshoppers (Mela- 
nopliis cliff erentialis and Schistocerca shoshoni), large June bugs, and 
one large ground beetle, found beneath a night roost used by pallid 
bats near Albuquerque, New ]\Iexico. 

It is not known whether bats of this species regularly prey upon 
animals other than arthropods. Engler (1943, pp. 96-97), however, 
records captive pallid bnts which were observed to eat western skinks 
(Eumeces skilto7iia7ius), a Sonoran desert gecko (Coleonyx variegatus), 


and were suspected of eating the head and neck of a Mexican free- 
tailed bat (Tadarida mexicana). It seems likely that starvation was re- 
sponsible for such deviation from an insectivorous diet, although, as the 
author suggests, it is possible that small night lizards may locally be 
preyed upon by pallid bats. 

It is practically impossible to determine the quantity of food con- 
sumed nightly by insectivorous bats in the wild. In captivity, however, 
it is a relatively simple matter to ascertain the amount of food a bat 
consumes each night, although this does not necessarily represent the 
amount that would be taken under natural conditions. Mathias and Se- 
guela (19-40, pp. 15-19) found that Myotis mystacinus and M. myotis, 
fed on fl.y pupae, consumed an amount equal to about one-third of 
their own weight daily, Eamage (1947, p. 61) found that several spe- 
cies of Myotis, kept in small flight cages, ate about one-half their own 
weight in fly pupae or termites each night while a specimen of Epte- 
sicus fuscus, kept in a small cage in which it could not fly, ate an 
amount equal to about one-third of its weight per day. As noted by 
that author the quantity of food eaten by captive bats varies greatly 
on different nights. 

From May 25 to September 23, 1947, records were kept of the 
amount of food (meal w^orms in this instance) consumed by 3 captive 
pallid bats (1 male, 2 females) each night. The food was weighed be- 
fore being placed in the cage each afternoon and the amount left in 
the morning was again weighed. From May 25 to August 13, the bats 
were kept indoors 41 per cent of the time. After this date they were 
constantly kept indoors. From ]\Iay 25 to June 1 the average amount 
of food consumed per bat per night was 3.5 grams. The average 
weight per bat was 29.2 grams on June 5. During July the average 
amount of food consumed per bat per night was 4.2 grams. The aver- 
age weight per bat was 29.9 grams on July 10. From August 1 to 
August 13, the nightly food consumption per bat was 4.5 grams. The 
average weight per bat was 28.3 grams on August 7. From August 
14 to September 23 each bat ate an average of 4.7 grams of food per 
night. During this period, however, there was a marked increase in 
the weight of the bats. Their average weights on various dates were 
as follows: August 21, 30.6 grams; August 29, 32.3 grams; September 
5, 33.5 grams; September 12, 35.1 grams; September 19, 38.6 grams. 
During this entire period of nearh' 4 months these bats were given ade- 
quate amounts of food daily as evidenced by remains found in the 
mornings. The marked increase in weight during the latter part of 
August and in September may be attributed partly to lack of sufficient 
exercise, although there is evidence that under natural conditions bats 
show an increase in weight at this time of year. 


AVJiile captive bats were principally fed meal worms during the 
early phases of this study and a prepared diet later (see p. 168) cer- 
tain other kinds of animal food were occasionally offered them. They 
accepted the pupae of the following species of Diptera : Lucilia seri- 
cata, CaUiphora erythrocephala, and Calliphora vomitoria. On one oc- 
casion an omnivorous looper moth ( Sahulodes caherata) wrh accepted 
by a bat and eaten. On another occasion two of three snails (Helix 
aspersa) were eaten by captive pallid bats. California oak moths (Phry- 
ganidia caJifornica) were refused. 

No personal field observations were obtained relating to the man- 
ner in which these l^ats eat large insects. On several occasions, how- 
ever, live Jerusalem crickets, held by forceps, were offered captive bats 
that w^ere hanging head downward in cages. In each instance the bat 
viciously bit the insect, then reversed its position in the cage so that 
its head was uppermost. The tail and uropatagium were then curled 
up ventrally so as to form a basket before eating began. The posi- 
tion of the bat at such times was similar to that assumed by females 
when giving birth to young. The basket served to prevent the food 
falling to the ground until all desirable parts were consumed. The 
head and legs of these large crickets were discarded. Borell (1942, p. 
337) observed a pallid bat in a similar position when it was eating 
a grasshopper, and Pittman (1924) described a similar posture for 
captive individuals of Myotis lucifugus when they were eating moths. 


The following ectoparasites w^ere found on pallid bats from San Luis 
Obispo and Kern counties: 

Cimex pilosellus Horvath. Western bat bedbug. 

Basilia antrozoi (Townsend). Bat fly. 


Myodopsylloides palposa (Rothschild). Flea. 


Ornitliodoros stayeri Cooley and Kohls. Tick. 

Ornithodoros sp. Tick. 

Spinturnix sp. Mite. 

Steatonyssus sp. Mite. 


Cimex pilosellus Horvath. On May 10, 1947, a western bat bedbug was 
found on a pallid bat captured at a daytime roost in western Kern 
County. On April 14, 1951, while trying to remove some pallid bats 
from a colony along San Juan Creek, 9 miles west of Simmler, San 
Luis Obispo County, a western bat bedbug was found adhering to a 
net which had been poked into the crevice housing the bats. 

Basilia antrozoi (Townsend). This species of nycteribiid w^as found 
to be fairly common on pallid bats. Nearly all bats of this species ex- 
amined during the spring and summer months had a few, usually 1 
to 3, of these parasitic flies on them. The greatest number found on 
any one individual was 8. It is possible that these parasites are less 
numerous in the fall as only 10 out of 37 pallid bats captured on Sep- 
tember 20, 1947, in San Luis Obispo County had flies on them. These 
insects showed a preference for the sides of the bodies of their hosts. 
They usually remained hidden in the fur unless disturbed. This spe- 
cies has previously been reported from Antrozous as Avell as from sev- 
eral other genera of the Family Vespertilionidae (cf. Stiles and Nolan, 
1931, p. 648). 

MyodopsyUoides pal pom (Kothschild). Fleas were found to be rela- 
tively uncommon parasites on pallid bats. They were recorded only 
three times and no more than one flea was ever found on a bat. Two 
specimens w^ere collected on September 20, 1947, and another was se- 
cured on April 14, 1951. 

Ornithodoros sp. Many pallid bats collected or examined in the field 
during the months of September and October were rather heavily in- 
fested with ticks. Those ticks collected during these months proved to 
])e larval forms of the genus Ornithodoros but could not be identified 
as to species. A count was made of the number of ticks found on a 
grou]) of 37 bats secured in San Luis Obispo County on September 
20, 1947. Eleven of the bats were free of these parasites. The other 26 
had the following numbers, respectively: 1, 1, 1, 1, 1, 1, 1, 1, 2, 2, 3, 3, 
3, 3, 4, 4, 4, 4, 6, 6, 8, 10, 12, 13, 16, 26. Ticks were most frequently 
found on the sides and back of the neck, the shoulders, and sides of 
the body. Larval ticks were not found on the flight membranes. 

On April 14, 1951, while capturing bats from a colony living be- 
neath a loose slab of rock on a cliff 9 miles west of Simmler, San Luis 
Obispo County, a large number of adult ticks were found crawling up 
the perpendicular face of the cliff beneath the roost. In probing for 
the bats the ticks had apparently been dislodged and they were at- 
tempting to return to the crevice that housed the bats. A number of 
these were saved and proved to be Ornithodoros stageri Cooley and 
Kohls. An examination of 11 pallid bats captured here at this time 
revealed the presence of larval ticks on 5 individuals. The number of 


ticks found on the bats varied from 1 to 11. A single adult tick of 
the above-mentioned species was found boring into the upper surface 
of the uropatagium of one bat. 

Spinturnix sp., and Steatonyssus sp. Mites were commonly noted 
on the flight membranes of nearly all bats examined in the field and 
those kept in the laboratory. All those identified were either adults 
or nymphs of the genera Spintuniix or Steatonyssus. Individuals of the 
latter genus were generally more al)undant than those of the former. 
The number of mites on pallid bats usually far exceeded that of all 
of the other ectoparasites together. No marked seasonal fluctuation in 
numbers was apparent. 

Although no Diptera belonging to the family Streblidae were ob- 
served by the writer on any bats of this species, Jobling (1949) 
records TrichoMus corynorMni Cockerell from Antrozous in California. 
This streblid primarily parasitizes bats of the genus Corynorhinus (cf. 
Kessel, 1952) but has been recorded on several other kinds of cave- 
inhabiting bats. Possibly the occurrences of these flies on bats other 
than those of the genus Corynorhinus is accidental. 


As was noted by the late Dr. Glover M. Allen (1939, p. 280), "Bats 
have few enemies.'' There is no indication that pallid bats are an ex- 
ception to this I'ule. It is likely that their greatest enemy among the 
vertebrates is man who dislikes having small native mammals living 
in his attics and barns. 

Pallid bats are occasionally preyed uiron l)y owls. Baker (1953, p. 253) 
records the remains of 6 pallid bats found in ]iellets, belonging to either 
horned owls {Buho virginianus) or barn owls {Tyto alha), collected in cen- 
tral Coahuila, Mexico. 

Since bats of this species emerge rather late in the evening there 
is little likelihood of their being attacked by diurnal birds of ]n-ey 
under ordinary conditions. If a colony is subjected to occasional day- 
time disturbance, however, there is some evidence that individuals 
may be captured or at least harassed by small hawks. On -June 9. 
1948. 7 banded bats that had IxH'n ca]>tured 3 days previously at a 
colony about 30 miles away, were released at 7:00 a.m. close to a cliff 
bordering San Juan Creek in central eastern San Luis Obispo County. 
There were numerous crevices and small caves in the nearby cliff and 
several of the bats immediately flew to one of the recesses. Another 
lit on a sandstone outcrop and was immediately attacked by a Brewer 
blackbird (Eupliagus cyanocephaJus). It was able to avoid the at- 
tacker bv scrambling into a narrow crevice. The other bats flew ar-ound 


the cliff and disappeared from sight. Shortly afterwards, however, a 
sparrow hawk (Faico sparverius) flew over the cliff' with an object that 
appeared to be a pallid bat in its claws. Another instance of a hawk 
attacking a bat of this species was observed about midday on October 
23, 1949, at a rocky outcrop along the northwestern edge of Carrizo 
Plain in San Luis Obispo County. On this occasion while the w^riter 
and several others were attempting to capture a group of bats, found 
beneath a loose slab of sandstone, in order to band them, 2 individuals 
escaped. One of these was suddenly attacked, as it was flying about in 
the sunlight, by a sharp-shinned hawk (Acci2nter striatus) that had 
been perched in a nearby oak. The bat evaded the hawk and dropped 
safely into a crevice out of sight. 

Allen (1939, p. 288) mentions a record, supplied him l)y B. Patter- 
son Bole, of a snake (species not known) that was found to have cap- 
tured a pallid bat in California. As suggested by Allen it is not sur- 
prising that bats of this species, which occasionally alight on the 
ground, are sometimes caught by nocturnal snakes. 

Hatt (1923, p. 261) in commenting on the remains of scorpions 
found beneath pallid bat roosts, a fact confirmed during the present 
study, makes the following comment: "It is difficult to understand how 
these bats capture the scorpions without being severely hurt or killed, 
for the venom of this scorj^ion is ]ioisonous enough to kill most mam- 
mals of this size and the ]>hysical injury of the injection alone should 
be a serious matter to so small an animal. These scorpions are able to 
bend the postabdomen to such a wide range of positions that it would 
seem impossible that the bat could avoid being stung the greater num- 
ber of times that it captured such ])rey. It is ]iossible then that this 
bat is immune to the venom of a scorpion." Nothing is knowai regard- 
ing the immunity of pallid bats to scorpion venom or their ability to 
kill such arthropods without being stung. Consequently, the scorpion 
must be considered as a potential enemy until proved otherwise. 


Pallid bats are to be found in colonies during the spring, summer, 
and early fall months. The number of individuals in established sum- 
mer colonies observed dui'ing this study was found to vary from ap- 
proximately 30 to 100. Daytime roosts may be in crevices in rocks 
or recesses in man-made structures, such as houses, barns, and bridges. 
Night roosts, where bats may rest or consume insects that they have 
captured, are usually close to the day roosts. 

In central California pallid bats arrive at the summer colonies by 
the latter part of ]\Iarch or early April. Members of both sexes may 


be present in a colony. The annual molt may occur any time be- 
tween May and Auo-ust. Summer colonies start to break up in October. 
During this month and early November the bats are found in smaller 
groups, often in situations where they are not to be seen earlier in 
the year. Little is known regarding the whereabouts of pallid bats in 
winter. They are absent from their summer retreats and have not been 
observed flying about at night. There is no evidence, however, that 
any extended migration occurs. The few records available of win- 
tering individuals indicate that single bats or small groups pass the 
winter, probably in a torpid condition, in remote recesses. A study 
was made on the effect of lowered environmental temperatures on cap- 
tive pallid bats. Individuals kept in a dark room without food for 144 
days at temperatures ranging from 40° F. to 50 °F. lost an average of 
27.6 per cent in body weight or 0.193 per cent per day per bat. Bats 
kept under similar conditions but at temperatures between 38°F. and 
40°F. for 119 days lost an average of 24.96 per cent in body weight 
or 0.209 per cent per day per bat. Bats kept for extended periods of 
time at low temperatures without food occasionally drank water and 
did not survive unless high relative humidity was maintained in their 

Seasonal fluctuations were apparent in the body weights of pallid 
bats. Lowest w^eights were recorded for bats when they first made 
their appearance in sununer colonies in the spring. Males gained weight 
for several weeks, then maintained a fairly constant level until the end 
of summer. Females continued to gain weight until they had their 
young. Members of l)oth sexes again showed an increase in weight in 
the fall. 

Several kinds of vocal utterances are produced by pallid bats, in- 
cluding an intimidation note, a squabble, a directive call, chittering, 
and a plaintive note associated with pain. The odor, produced by glands 
on either side of the nose of these bats is distinctive and it is suggested 
that it may serve as a defense mechanism to repel other animals. 

Studies were made on the flight of these bats, using slow motion 
pictures and stills made with the aid of an electronic flash. The mo- 
tion of a bat's wing is essentially the same as that of a bird's wing in 
forward flight. The downstroke is dow^nward and forward and the up- 
stroke is upward and backward. The upstroke is considered to be less 
efficient in the bat's wing than in the bird's wing. The feathers of the 
latter can be separated to minimize air resistance at such times while 
any comparable action is impossible for the wdng of a bat. Other ac- 
tions on the part of a bat in flight, however, compensate for this loss 
in efficiency. In straight flight the pallid ])at makes 10 to 11 wing 
strokes per second. 


Members of this species emerge rather late in the evening. In early 
June the illumination was less than 0.1 foot candles when the bats 
emerged. In the latter part of September they were recorded emerg- 
ing earlier in relation to sunset. 

Observations in the field as well as on captive bats kept in outdoor 
cages indicate that in central California pallid bats copulate at least 
during the months of October and November. Copulation was observed 
in captive bats maintained in indoor laboratory cages between the 
months of November and February, inclusive. Nothing is known re- 
garding the sexual behavior of these bats in the wild in winter. 

Early winter ovulation was induced in a female that was placed 
in a warm room on December 5 and kept there until she gave birth to 
a normal young 83 days later on February 26. Another female that 
was known to have copulated was isolated in an out-of-door flight cage 
on December 29. Nearly 5 months later she gave birth to one young, 
showing that in this instance later winter or early spring insemina- 
tion was not necessary for fertilization. Vaginal smears of captive fe- 
males revealed the ]n-esence of nucleated epithelial cells all year around. 
Cornified cells were most abundant from the end of January until 
early in May. Leucocytes were noted only in March and April and 
their presence was thought to be associated with ovuhition. The gesta- 
tion period is estimated to average about 9 weeks. The average num- 
ber of young for 28 females was 1.8. Young are born between early 
May and the middle of June, the time varying with year, locality, 
colony, and with individual females in a colony. The sex ratio at birth 
is nearly equal. 

Females hang u]>right at time of parturition and the young are 
born by breech presentation. It would appear that under normal cir- 
cumstances the female cuts the umbilical cord and eats the placenta. 
Young pallid bats have their eyes closed at birth and their ears are 
folded down tightly against the sides of the head. The body is essen- 
tially naked. The eyes o])en lietween the eighth and the tenth day 
of jiostnatal life, at which time the ears can be erected. Fear is ex- 
hibited by the tenth day. Fur is evident on parts of the body by the 
fourth day and has nearly grown out by the end of one month. Cap- 
tive-born young were observed flying when 7 weeks old but it is be- 
lieved that young in the wild can fly at an earlier age. 

The deciduous incisors and canines have erupted through the gums 
at time of birth and the premolars generally make their appearance 
when the young are 3 to 5 days old. The first deciduous teeth to be lost 
are the premolars and the last are the canines. Many of the perma- 
nent teeth appear before their deciduous predecessors are lost. 

Large orthopterous insects and beetles appear to be important in 


the diet of pallid bats. Some of the insects eaten are flightless species 
which must be picked up from the ground. 

Mites of the genera Sinnturnix and Steatonyssus, ticks, and wingless 
dipterans belonging to the family Nycteribiidae are common ectoparasites 
on pallid bats. Less common are fleas and bat bedbugs. These bats 
have few known enemies. If disturbed enough to fly during the day- 
time they may be attacked by small hawks. There is one record of a 
pallid bat found in the stomach of a snake, and another of remains of pallid 
bats found in owl pellets. 


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1949. Sex ratios in Indian bats. Journal of the Bomhay Natural History 
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1944. Notes on tlie winter occurrence of bats in Nevada. Journal of Mam- 
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Allen, G. M. 

1939. Bats, x + 368 pp., frontispiece + 57 text figs. Harvard University 
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Allen, J. A. 

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1935. Bird flight, xii + 294 pp., illustrated. Dodd, Mead, and Company, New 


Bailey, V. 

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24 text figs., 1 map. 
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1936. The mammals and life zones of Oregon. United States Department of 

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1936. The seasons in a tropical rain-forest (New Hebrides). — Part 4. Insec- 
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Vol. XXVIII, No. 5, pp. 247-321, Figs. 1-45 January 7, 1954 




Institute for Infectious Diseases 
University of Tokyo 



Department of Zoology, University of California, 
Davis, California 

The classical studies of tsutsugamushi disease in Japan long ago drew 
first attention to the chigger (trombiculid mite) fauna of that country, and 
for many years this disease and the associated chiggers were investigated 
and reported upon by Japanese workers. Because many of their reports 
are difficult to obtain and because of the previous lack of interest in chig- 
gers, the western world had only a vague understanding of the Japanese 
species. Our knowledge of Oriental chiggers was greatly advanced during 
World War II : many species were described and the status of many pre- 
viously named species was clarified. Following the war, there has been a 
resurgence in the taxomomic study of chiggers in Japan, and many new 
species have been described from this small country. Unfortunately, most 
of these descriptions are in Japanese and in journals of a limited distri- 
bution outside of Japan. 

Studies upon which this paper is in part based were conducted under 
contract no. DA-49-007-MD-242, between the Regents of the University 
of California and the Medical Research and Development Board, office of 
the Surgeon General, Department of the Army. The work was initiated 
while the junior author was associated with the 406th Medical General 
Laboratory and Far East Medical Research Unit in Japan during the period 
January-October, 1952. Logistical support for the preparation of the 
illustrations was supplied by the U. S. Army Hospital, 8164 Army Unit, 
APO 9, by special arrangement with the above-mentioned organizations. 

This paper presents a brief summary of the chigger fauna of Japan. 
Omissions are inevitable for new species are being described from Japan 



almost every month; and before publication of this paper our list will be 
outdated. Nevertheless, we hope that the keys, diagnoses, and data with 
these illustrations will simplify the identifications of Japanese chiggers. 

It would be premature to discuss the Japanese trombiculid fauna with 
regard to chiggers elsewhere; but from the list of chiggers already known 
from Japan, however, some comments are justified. We have listed here 6 
genera and 45 species of chiggers in the Japanese fauna. Of the 45 species 
known from Japan, 27 are endemic : in view of the widespread geographic 
distribution of species of chiggers generally, it is remarkable that more 
than half of the known species in this tiny country seem to be confined to it. 

The most important genus is Tromhicula. Worthy of comment is the 
abundance both in numbers of individuals and in species of the subgenus 
Leptot romhidium : this group contains 18 species in Japan, and 14 of these 
are apparently confined there. Although the subgenus Lepiotromlndium 
is well represented elsewhere in the Orient, there seems to be no such con- 
centration of species in such a small area. This situation presents an inter- 
esting contrast to the fauna of adjacent North America where the group is 
rather scarce. The subgenus TromhicuJmdus, containing one species in 
Japan, is represented by four other species in the tropical Orient, and thus 
seems to be an austral element. The subgenus Miyatromhicula contains a 
single species (T. (31.) kochiensis) found on the islands of Skikoku, Kyushu, 
and southern Honshu. In contrast to the other groups of Tromhicula, the 
subgenus NeotronihicuJa, with six species in Japan, contains two species 
that are also found in North America, and four species that occur on the 
adjacent Asiatic mainland. The single species of the subgenus Eutromhi- 
cula (ivichmanni) in Japan is widespread and common in the southern 
Orient; avian hosts of this species make a wide geographic distribution 
easily understandable. 

In Japan the genus Euschdngastia is scarce, and known from only four 
species; one of these (E. ikaoensis) has been found also in southern Korea. 
This state of affairs is vastly different from what obtains in North America 
where Euschdngastia is one of the dominant groups. 

The genus Neoschongastia, its members characteristically parasites of 
birds, is represented by six species in Japan; five of these occur elsewhere 
in the world, two of them having been described from North America. 
Theoretically, of course, they can be as widely distributed as their hosts; 
and because most species of Neoschongastia appear not to be confined to 
a single species of host, they may easily be more widely distributed than 
any one host. 

When the chigger faunas of the world are more completely known, we 
can better classify that of Japan. As we know it today, on the specific level 
the trombiculid fauna of Japan seems not closely allied to that of any other 



region : the large number of endemic species justifies labelling the fauna as 
typically Japanese : that is, the species seem no more closely related to 
Oriental species than they are to Xearctic species. Generically, Japanese 
chiggers seem more typically Palaearctic than Oriental in affinities; many 
genera of the tropical Orient are not known to occur in Japan. 

The illustrations in this paper were made in Kyoto, Japan, and repre- 
sent the holotype wherever such was available. The staff of artists was 
under the direct supervision of Mr. K. Yamazaki; Mr. Yamazaki was 
assisted by Mr. A. Shimazoe, Mr. K. Daishoji, Mr. S. Shibata, Mr. M. Endo, 
Miss F. Tamara, and Miss C. Tanaka. For the excellence of their pains- 
taking labors we are deeply appreciative. We wish to thank also Miss Y. 
Yoshida, technician and interpreter, who prepared the bulk of the material 
collected during the course of this study. 

Dr. Mitosi Tokuda was of great assistance in collecting many of the 
specimens used in this paper. Special thanks are due Mr. Seiichi Toshioka 

Fig. 1 Acomatacarus yosanoi 


for his invaluable service in cheeking many of the scutal measurements 
used in this paper; and for collecting a large amount of the data on host 
and geographic distribution of Japanese chiggers. 

We are greatly indebted to Dr. James M. Brennan for much council 
given during the preparation of this paper; and to Major Paul W. Oman, 
MSC, under whose administration this work was done. 

Key to Genera and Subgenera of Chiggers in Japan 

1. Coxa I with two setae genus Acomatacarus 

— Coxa I with one seta 2 

2. With no anteromedian seta genus Gahrliepia 3 

— With one anteromedian seta 4 

3. Scutum with four marginal setae subgenus Walchia 

— With more than four setae, some not marginal subgenus Gahrliepia 

4. Sensillae expanded distally 5 

— Sensillae flagelliform genus Trombicula 7 

5. Scutum partly submerged beneath the integument genus Neoschongastia 

— Scutum entirely on the surface of the integument 6 

6. Coxa II with a single seta genus Euschongastia 

— Coxa II with more than a single seta genus Doloisia 

7. Posterolateral setae greatly expanded, foliate subgenus Trombiculindus 

— Posterolateral setae slender 8 

8. Coxa III with several setae subgenus MiyatromMcula 

— Coxa III with a single seta 9 

9. With one or more mastitarsalae III 10 

— With no mastitarsalae III 12 

10. Scutum roughly pentagonal; palpal claw three-pronged. .subgenus NeotromMcula 

— Scutum rectangular; palpal claw two-pronged 11 

11. Axial prong (of palpal claw) external (or dorsal) ; with 20 or 22 dorsal 
setae subgenus Eutrombicula 

— Axial prong (of palpal claw) internal (or ventral); with 30 or more dorsal 
setae ungrouped species 

12. Palpal femoral and genual setae nude; galeal seta feathered; no mastitar- 
salae III; sternal setae 2-2; coaxal setae 1-1-1; scutum more or less 
rectangular subgenus Leptotrombidium 

— Without the above combination of characters ungrouped species 

Acomatacarus yosanoi Fukuzumi and Obata, 1953 
(Figure 1) 

Acoinatacarus yosanoi Fikuzumi and Obata, 1953, Kitasato Archives of Experi- 
mental Medicine, vol. 26. no. 2, pp. 1-22. 

Diagnosis: All palpal setae feathered; ventral tibial seta with long 
branches, other palpal setae very short, fine branches. Palpal claw four- 
pronged. Galeal seta with fine branches. Chelicera with a row of about 



seven small teeth along the dorsal edge. Scutum with general conformation 
and characteristics of the genus; posterior margin more or less three sided. 
Sensillary bases about on a line with the posterolateral setae. Bases of 
sensillae with a small number of very small barbs; sensillae nude distally. 
With 80 or more dorsal setae, more or less scattered. A single pair of sternal 


Fig. 2 GaJirliepia ogatai 



[Proc. 4th Ser. 

setae between coxae III. Coxal setae 2-1-1. Seutal measurements of holo- 
type: AW-66, PW-76.5, SB-24, ASB-27, PSB-20, SD-47.5, AP-24, 
AM^6, AL-46, PL-64, S-72. 

Fig. 3 Gahrliepia saduski 



Distribution and hosts : Known only from the type collection. 

Type data : From Rattus rattus from Miyake Island (Izu Islands, south 
of Tokyo), January 18, 1952; holotype and one paratype deposited at the 
Kitasato Institute, Tokyo. 

F'ig. 4 Doloisia okabei 


Gahrliepia (Walchia) ogatai (Sasa and Teramura, 1951) 

(Figure 2) 

Walchia ogatai Sasa and Teramura, 1951, Tokyo Iji Shinshi, vol. 68, no. 5, pp. 9-10. 

Diagnosis: Galeal seta forked (not nude as in the original description) ; 
palpal setae all forked or feathered except lateral tibial seta which is nude. 
Scutum small, bluntly or sharply pointed posteriorly. Two pairs of humeral 
setae. Coxal setae 1-1^ or 5. Sternal setae 2-2. Scutal measurements of 
holotype: AW-34, PW-57, SB-27, ASB-26, PSB-27, AP-36, AL-31, 
PL-30, S-28. 

Distribution and hosts.:. Mogera ivogura and Apodemus speciosus from 
Akita and Shizuoka prefectures. 

Type data : Holotype and four paratypes taken from Mogera wogura, 
Uchitomo, Akita Prefecture; November 3, 1950. Deposited at the Institute 
for Infectious Diseases, University of Tokyo. 

Gahrliepia (Gahrliepia) saduski Womersley, 1952 

(Figure 3) 

Gahrliepia (Gahrliepia) saduski Womersley. 1952, Records of the South Australia 
Museum, vol. 10, pp. 301-302. 

Diagnosis: Galeal seta feathered; palpal setae feathered, except dorsal 
and lateral tibial setae which are nude. Scutum unsculptured and with no 
punctae; with 2-5 setae behind PL. Coxal setae 1-1-3, 4, or 5 (some speci- 
mens have 3 setae on coxa III on one side and 5 on the opposite side). 
Scutal measurements (mean of seven specimens given by the describer) : 
AW-48.9, PW-63.5, SB-41.4, ASB-22.4, PSB-65.4, sb-87.8, AP-38.4, 
AL-36.7, PL-35.2, S-38.4. 

Distribution and hosts : Known from many localities in Honshu, Shi- 
koku, and Kyushu; from Mus. sp., Apodemus speciosus, A. geisha, Cleth- 
rionomys smithii, Microtus montehelloi, Urotrichus talpoides, and Passer 

Type data : Holotype and two paratypes from Mus. sp., Misinobe (prob- 
ably a misspelling of Mizonobe) near Yachi, Yamagata Prefecture; October 
28, 1945. Holotype at South Australia Museum, Adelaide. 

Doloisia okabei Sasa et al., 1952 

(Figure 4) 

Doloisia okabei Sasa, Hayashi, Kawashima, Mitsutomi, and Egashira, 1952, Tokyo 
Iji Shinshi, vol. 69, no. 3, p. 12. 

Diagnosis: The only species of the genus known from Japan. Coxal 
setae 2 — 4(4^5) — 8(7-9). Sternal setae 2-2. Galeal seta nude. Palpal 



femoral seta feathered; palpal genual and dorsal and lateral palpal tibial 
setae nude; ventral tibial seta feathered. Palpal claw three-pronged. Scutal 
measurements of holotype: AW-28, PW-63, SB-23, ASB-23 PSB-20 
AP-30, AM-30, AL-21, PL-40, S-35. 

Distribution and hosts: Known only from the original collection. 

Fig. 5 Euschongastia alpina 



[Proc. 4th Ser. 

Type data: Holotype and two paratypes from Urotrichus talpoides, 
taken at Toshita, Kumamoto Prefecture; October 21, 1951. Holotype and 
two paratypes deposited at the Institute for Infectious Diseases, University 
of Tokyo. 

Fig. 6 Euschdngastia ikaoensis 


Genus Euschongastia Ewing, 1938 

Only four species of Euschongastia are known from Japan and they are 
relatively uncommon. To date only one species (E. ikaoensis) is known to 

Fig. 7 Euschongastia kitajimai 


occur on the adjacent Asiatic mainland; this species has been taken in 
southern Korea. Euschongastia differs from Neoschongastia in the charac- 
ter of the scutum and in the host distribution : in Euschongastia the scutum 
lies entirely on the surface of the integument and the hosts are mammals, 
whereas in Neoschongastia at least some parts of the scutum lie beneath 
cuticular striae and the hosts are almost always birds. 

Key to Japanese Species of Euschongastia 

No mastitarsalae III 2 

One mastitarsala III E. kitajimai 

A single seta on coxa III 3 

Two setae on coxa III E. miyagawai 

Two pairs of humeral setae; lateral tibial seta of palpus feathered; galeal 
seta nude E. ikaoensis 

A single pair of humeral setae; lateral tibial seta of palpus nude; galeal 
seta branched E. alpina 

Euschongastia alpina Sasa and Jameson, new species 

(Figure 5) 

Gnathosoma: Chelicera with a subapical dorsal tooth; cheliceral base 
with no punctae. Capitular sternum with a single feathered seta and a 
few punctae. Palpal genual and femoral setae feathered; dorsal and ven- 
tral tibial setae feathered, lateral seta nude. Palpal claw five-pronged. 
(Jaloal seta branched. 

Legs: All coxae unisetose; the seta on coxa III distinctly behind the 
anterior margin of the coxa. Specialized (nude) setae: Leg I, 2 genulae, 1 
microgenuala, 2 tibialae, 1 microtibiala, 1 spur, 1 microspur, 1 subter- 
minala, 1 parasubterminala, 1 pretarsala; Leg II, 1 genuala, 2 tibialae, 1 
spur, 1 microspur, and 1 pretarsala; Leg III, 1 genuala. 

Scutum : Broadly rectangular, more than three times as wide as long, 
concave anteriorly, convex posteriorly, and with the lateral margins diverg- 
ing posteriorly. Posterior corners angular, posterolateral setae placed in 
the corners. Sensillary bases behind a line connecting posterolateral setae. 
Sensillae pyriform. Scutum with few or no punctae. No diagonal lines in 
front of sensillary bases. Scutal measurements of holotype: AW-63, 
PW-95, SB-36, ASB-26, PSB-9, AP-20, AM-35, AL-35, PL-75, S^O. 

Setae : Humeral setae similar to posterolateral setae ; other dorsal setae 
slightly shorter. Dorsal setal formula 2-10-12-8-6-2-2. Sternal setae 2-2. 
About 44 small setae ventrallv behind coxae III. 



Type: Holotype from Apodemu^i geisha, Kamikochi (1600 meters, ele- 
vation), Nagano Prefecture; July 12, 1952. One paratype from A. specio- 
sus, Mt. Hiei, Shioa Prefecture; March 27, 1952. Both specimens collected 
by Mitosi Tokuda and E. AV. Jameson, Jr. Deposited in the United 
States National ^Museum. 

Euschongastia ikaoensis (Sasa et al., 1951) 
(Figure 6) 

Neosclwngastia ikaoensis Sasa, Sawada, Kano, Hayashi, and Kumada, 1951, Tokyo 
Iji Shinshi, vol. 68, no. 4, p. 7. 

Diagnosis: All palpal setae feathered. Galea! seta nude. Palpal claw 
5-7 pronged. Sternal setae 2-2. Coxal setae 1-1-1. Bases of sensillae about 
as far from each other as from their adjacent lateral margins of the scutum; 
sensillae pyriform. Two pairs of humeral setae. Scutal measurements of 
holotype: AW-61.5, PW-67, SB-24, ASB-18, PSB-15.5, AP-21, AM-30, 
AL-30, PL-39, S-33. 

1 Mt f . 


Fig. 8 Euschongastia miyagawai 



Distribution and hosts: From Kanagawa, Gumma, Oita, Yamanashi, 
Shiziioka, Kiimamoto, and Shimane prefectures from Urotrichus talpoides, 
Apodemus speciosus, CJethrionomys sniithii and 3Ius molossinus; this chig- 
eer also occurs in southern Korea. 

Fig. 9 Neo.schongastia asakawai 



Type data: Holotype from Apodemus speciosus from Ikao, Gumma 
Prefecture; November 27, 1950. Two paratypes from the same host from 
Yamanaka, Yamanashi Prefecture; November 22, 1950. Holotype and para- 
types deposited at Institute for Infectious Diseases, University of Tokyo. 

Euschongastia kitajimai Fukuzuki and Obata, 1953 

(Figure 7) 

Euschongastia kitajimai Fukizumi and Obata, 1953, Kitasato Archives of Experi- 
mental Medicine, vol. 26, no. 1, pp. 1-22. 

Diagnosis: Palpal femoral seta feathered; palpal genual seta and pal- 
pal tibial setae feathered or branched. Galeal seta nude. Palpal claw 
three-prono-ed. Sensillary bases closer to their adjacent scutal margins than 
to each other, midway between anterolateral and posterolateral setae. Sen- 


Fig. 10 Neoschongastia posekanyi 


sillae about five times as long as wide, with several rows of long barbs. 
Sternal setae 2-2. Coxal setae 1-1-1. One mastitarsala III. Scutal meas- 
urements of holotype: AW-52, P\V-62.5, SB-24, ASB-22.5, PSB-26, 
AP-33, AM-31, AL-25, PL-36, S-37. 

Distribution and hosts: Known only from the original collection. 

Type data: From Rattus rattus, Miyake Island (Islands of Izu, south 
of Tokyo) ; January 18, 1952; deposited at Kitasato Institute, Tokyo. 

Euschongastia miyagawai Sasa et ah, 1951 

(Figure 8) 

Euschongastia miyagaivai Sasa, Kumada, and Miura, 1951, Tokyo Iji Shinshi, vol. 

68, no. 8, p. 19. 
fNeoschongastia covelli Rauforu, 1953. Parasitology, vol. 42, nos. 3 and 4, pp. 


Diagnosis : Palpal genual and femoral setae feathered ; dorsal tibial seta 
branched or nude, lateral tibial seta branched, and ventral tibial seta feath- 
ered. Galeal seta nude. Palpal claw 3-5 pronged. Sensillary bases very 
close together; sensillae globular. Sternal setae 2-2. Coxal setae 1-1-2. 
Dorsal setae in indistinct rows, between 50 and 60 setae. Scutal measure- 
ments of holotype: AW-62, PW-81, SB-8, ASB-28, PSB-11, AP-26, 
AM-40, AL-61, PL-57, S-31. 

Distribution and hosts : Known from Apodemus speciosus and Mustela 
sp. and several kinds of wild birds from Kanagawa, Yamanashi, and Kofu 
prefectures, and from Miyake Island, south of Tokyo. 


Type data: Holotype and two paratypes from Apodemus speciosus 
from Okuyugawara, Kanagawa Prefecture; March 5, 1951; deposited at the 
Institute for Infectious Diseases, University of Tokyo. 

Remarks: The original illustration is in error in showing the seta on 
the first palpal segment (palpal coxal seta) nude; this seta is feathered. 
Neoschongastia coveUi Radford, 1953 closely resembles E. mnjagawai and 
may well be a synonym. N. covelli was described from a squirrel (Cal- 
losciurus pygenjthrus) from Imphal, Manipur State, India. 

Genus Neoschongastia Ewing, 1929 

In Japan there are six species of Neoschongastia, all parasites of birds; 
undoubtedly additional species will be found in the future. The condition 
of the palpal setae, galeal seta, number of prongs on the palpal claw, pres- 
ence or absence of mastitarsalae III, number of setae on coxa III, and the 
number of sternal setae are all variable in this genus, and are specific char- 
acters. The almost unique feature of Neoschongastia is the position of the 


scutum, being partly submerged beneath the integument so that the cuti- 
cular striae can be seen above the scutum. 

Key to Japanese Species of Neoschongastia 

1. Coxa III with more than a single seta 2 

— Coxa III with a single seta 3 

2. Fifteen or more sternal setae N. carveri 

— Two pairs of sternal setae N. americana solomonis 

3. Sternal setae 2-2-2 N. paenitans 

— Sternal setae 2-2 4 

4. Sensillae spatulate, with long, slender scales N. monticola 

— Sensillae globular, with short scales 5 

5. One mastitarsala III N. posekanyi 

— Two to six mastitarsalae III N. asakawai 

Fig. 11 Neoschongastia americana 



[Proc. 4th See. 

Neoschongastia asakawai Fukuzumi and Obata, 1953 
(Figure 9) 

Neoschongastia asakawai Fukuzumi and Obata, 1953, Kitasato Archives of Experi- 
mental Medicine, vol. 26, no. 1, pp. 1-22. 

Diagnosis : This species bears some resemblance to N. posekanyi, but is 
distinguished by the greater number of mastitarsalae III and two or three 
pairs of humeral setae. In N. asakawai there are many more dorsal setae 
than in N. posekanyi. The variation in the number of mastitarsalae III 
(Fig. 9) is quite uncommon in Neoschongastia; these nude setae numbered 
from two to six. Seutal measurements of holotype: AW-50, PW-68, 
SB-41.5, ASB-22.5, PSB-30, SD-50.5, AP-40, AM-tl.5, AL-51, PL-46, 

Distribution and hosts : Collected from birds (Scolopax rusticola, 
BamhiisicoJa thoiacica, Siphia mugimaki, Pi'unella ruhida, Emberiza clioides, 

Fig. 12 Neoschongastia paenitens 



E. spodecephala, Turdus obscurus, and Passer montanus); known from 
Miyake Island (Tokyo-to) and Yamanashi Prefecture. 

Type data: Holotype from Scolopax rusticola, Miyake Island (Islands 
of Izu, south of Tokyo) ; January 20, 1952; deposited at Kitasato Institute, 

Fig. 13 Neoschongastia carveri 


Neoschongastia posekanyi Wharton and Hardcastle, 1946 

(Figure 10) 

Neoschdngastia posekanyi Wharton and Hardcastle, 1946, Journal of Parasitology, 
vol. 32, p. 302. 

Diagnosis : Head of sensillae globular, with small scales. Coxae uni- 
setose. Sternal setae 2-2. One mastitarsala III. One pair of humeral setae. 
Galeal seta nude (in original description) or 2-3 forked (in Japanese speci- 
mens). Palpal femoral and genual setae branched; dorsal tibial seta nude; 
lateral tibial seta forked; and ventral tibial seta feathered. Scutal meas- 
urements (mean of five specimens) given by the describers: "AW-75, 
PW-80, SB-47, ASB-25, PSB-34, AP-29, AM-49, AL-73, PL-52, S-30." 

Distribution and hosts: Collected from birds (Turdus celaenops, 
Monticola solitarius, Streptopelia orientaJis, and Columha janthina) from 
Hachijo and Koshima islands, south of Tokyo; Saxicola torquata and 
CMoris sinica, Schizuoka Prefecture. 

Type data : Holotype from Streptopelia orientalis, Hianza Island, Oki- 
nawa. Deposited in the United States National Museum. 

Neoschongastia americana solomonis Wharton and Hardcastle, 1946 

(Figure 11) 

Neoschongastia americana solomonis Wharton and Hardcastle, 1946, Journal of 
Parasitology, vol. 32, p. 289. 

Diagnosis : Coxal setae 1-1-3. No mastitarsalae III. Sternal setae 1-1. 
One pair of humeral setae, (^aleal seta branched, and all palpal setae 
branched or feathered. Palpal claw three-pronged. Sensillary bases far 
apart; nearer their adjacent scutal margina than to each other. Sensillae 
globular, with many fine scales. Scutal measurements (mean of five speci- 
mens from Okinawa) as given by the describers: "AW-51, PW-70, SB-37, 
ASB-25, PSB-30, AP-36, AM-44, AL-56, PL-59, S-24." Scutal measure- 
ments of a specimen from Japan: AW-50, PW-64, SB-34, ASB-34 
PSB-31, AP-36, AM-42, AL-55, PL-52, S-25. 

Distribution and hosts: From Hachijo Island from Monticola soli- 
tarius. Originally described from Bougainville, Guam, and Okinawa. Neo- 
schlmgastia a. americana is known from North America. 

Type data : Collected from Hirundo taJiitica, Cape Torokina, Bougain- 
ville; July 3, 1944. Holotype deposited in the United States National 



Eemarks : In scutal measurements the Japanese specimens resemble N. 
a. solomonis more closely than N. a. americana, but the difference (length 
of the scutum) may well prove to be clinal. 

Pig. 14 NeoscJiongastia monticola 



Fig. 15 Tronihicula kansai 



Fig. 16 Tromhicula kochiensis 


Neoschongastia paenitens Brennan, 1952 
(Figure 12) 

Neoschongastia kohlsi Brennan, 1951, .Journal of Parasitology, vol. 37, no. .579; pre- 
occupied by N. kohlsi Philip and Woodward, 1946. 

N. jyaenitens Brennan, 1952, Proceedings Entomological Society of Washington, 
vol. 54, p. 137. 

?N. okumurai FiTKizrMr and OnAXA. 1953, Kitasato Archives Experimental Medi- 
cine, vol. 26. no. 1, pp. 1-22. 

Diagnosis: Coxal seta 1-1-1. Two almost nude setae on tarsus III. 
Femoral, genual, and ventral tibial setae of palpus branched or feathered. 
Lateral and dorsal tibial setae of palpus nude. Galeal seta nude. Sensil- 
lary bases closer to each other than to their respective scutal margins; sen- 
sillae globular, with many fine scales. Scutal measurements of holotype 
(Brennan, 1951: 580) : "AW-52, PW-76, SB-25, ASB-25, PSB-20, AP-38, 
AM— to, AL-54, PL-49, S-31." Scutal measurements of a Japanese speci- 
men: AW-49, PW-71, SB-21, ASB-25, PSB-18, AP-32, AM-31, AL-50, 
PL-53, S-28. 

Distribution and hosts : Previously known only from Montana, U.S.A. 
In Japan known from Aomori Prefecture and Hachijo Island from Monti- 
cola solitarius. 

Type data : Collected from Petrochelidon pyrrhonota, Granite County, 
Montana; July 17, 1950. Holotype deposited in the United States National 

Comment: N. okumurai Fukuzumi and Obata seems closely related to 
paenitens and is probably a synonym. 

Neoschongastia carveri AVharton and Hardcastle, 1946 

(Figure 13) 

Neoschofigastia carveri Wiiakton and Hardcastle, 1946, Journal of Parasitology, 
vol. 32, p. 313. 

Diagnosis : This species is distinct in the large number of sternal setae 
(20-24), the numerous setae on coxa III (5-8), the excessively long spur 
on leg I (more than half as long as the tarsus), and the presence of at 
least two genualae on each leg. Scutal measurements (mean of five speci- 
mens) as given by the describers: "AW-80, PW-84, SB^3, ASB-30, 
PSB-37, AP-45, AM-71, AL-74, PL-141, S-40." Scutal measurements of 
a Japanese specimen: AW-83, PW-84, SB-36, ASB-31, PSB-38, AP-46, 
AM-46, AL-76, PL-120, S-42. 

Distribution and hosts: Originally described from Guam, Okinawa, 
Ulithi and Peleliu from numerous birds; in Japan from MonticoJa solitarius 
from Hachijo Island. 



Type data: Collected from Demigretta sacra, Port Ajayan, Guam; 
July 16, 1945. Deposited in the United States National Museum. 

Fig. 17 Tromhicida japonica 



Fig. 18 Trovibicula nagayoi 


Neoschongastia monticola Wharton and Hardcastle, 1946 
(Figure 14) 

Neoschongastia monticola Wharton and Hardcastle, 1946, Journal of Parasitology 
vol. 32, p. 301. 

Diagnosis : Distinctive in the narrow sensillae which are provided with 
long hairs. Coxae nnisetose. Sternal setae 2-2. One mastitarsala III. One 
pair of humeral setae. Seutal measurements (mean of five specimens) 

Fig. 19 TromMcula mitamurai 



given by the deseribers: ''AW-56, PW-78, SB-32, ASB-24, PSB-26, 
AP-27, AM-43, AL-35, PL-75, S-71." Scutal measurements of a Japanese 
specimen: AW-57, PW-83, SB-34, ASB-25, PSB-28, AP-29, AM-43, 
PL-36, PL-76, S-70. 

Distribution and hosts: Originally known only from Okinawa. As 
stated by the deseribers, this chigger seems to be confined to birds of the 
genus Moniicola. In Japan it occurs on MonticoJa solitarius on Miyake 
Island and Hachijo Island. 

Type data: From 31 onticola solitarius, Takabannase Island, Okinawa; 
June 10, 1945. Holotype in the United States National Museum. 

Genus Trombicula Berlese, 1905 

The genus Trombicula is based on adult morj^hology; the larva of the 
genotype {Trombicula minor Berlese) has not yet been associated with the 
adult, and therefore the current definition of the larval stage of Trombicula 
is tentative, a clarification pending the discovery of the larva of Trombicula 
minor. This interesting situation has been discussed at length by various 
authors. In Trombicula we include those chiggers with seven segments in 
each leg, the scutum with five setae and a pair of flagelliform sensillae, and 
in which the empodium is claw-like. There are several subgenera and num- 
erous species that do not fit into any established subgenera. 

Subgenus Trombiculindus Radford, 1948 

T rombiculindus was established as a genus for a chigger in which the 
posterolateral setae and some of the body setae are expanded and foliate. 
This is the only distinctive feature of this group, and we consider it as a 
subgenus in this paper. One species is known from Japan; the other species 
of Trombiculindus are recorded from the tropical Orient. 

Trombicula (Trombiculindus) kansai Jameson and Sasa, 1953 

(Figure 15) 

Trombicula (TromhicuUndus) kansai Jameson and Sasa, 1953, Journal of Para- 
sitology, vol. 39, pp. 247-249. 

Diagnosis: With the foliate posterolateral setae characteristic of the 
subgenus. Palpal genual, femoral, and lateral and ventral tibial setae nude; 
dorsal tibial seta feathered. Galeal seta feathered. Sensillary bases slightly 
behind posterolateral setae; sensillae nude basally, with barbs on the distal 
two-thirds. Posterolateral setae with small barbs on their surface. Sternal 
setae 2-2. Coxal setae 1-1-1. Two pairs of humeral setae. All dorsal setae 
and approximately half of the ventral setae foliate. Scutal measurements 


of holotype : AW-70, PW-82, SB-33, ASB-29, PSB-17, AP-20, AM-79, 
AL-44, PL-58, S-64. 

Fig. 20 Trombicula tamiyai 


Distribution and hosts : Most of the specimens were collected from the 
shrew-mole, Urotrichus talpoides; they usually attached at the base of the 
tail. A few specimens were found on the mice, Apodemus geisha and A. 
speciosus. Kyoto, Ilyogo, Mie, Shiga, Kumamoto, and Oita prefectures. 

Type data: Holotype from Urotrichus talpoides, Ohara Area (near 
Kyoto City), Kyoto Prefecture; March 8, 1952. Holotype and ten paratypes 
deposited in the United States National Museum. 

Subgenus Miyatrombicula Sasa, Kawashima, and Egashira, 1952. 

This monotypic subgenus is characterized by the multisetal condition 
of coxa III, the pentagonal scutum, and the presence of a short mastitarsala 
III. It appears to be allied to Tromhiciila cynos Ewing of North America, 
and a comparison of these two species should be made. 

Trombicula (Miyatrombicula) kochiensis Sasa et al., 1952 

(Figure 16) 

Trombicula (Miyatronibicula) kochiensis Sasa, Kawashima, and Eciashira, 1952, 
Tokyo Iji Shinshi, vol. 27, no. 6. p. 5. 

Diagnosis: All palpal setae feathered. Palpal claw three-pronged. 
Galeal seta feathered. Scutum pentagonal, posterior margin projected and 
rather pointed. Sensillary bases slightly behind a line connecting postero- 
hiteral setae. Bases of sensillae nude; distal half or two-thirds with short 
barbs. Scutal setae densely plumose. Dorsal setae 80 or more, in poorly 
defined rows. Sternal setae 2-2. Coxal setae 1-1-6(6-9). One very short 
mastitarsala III. Scutal measurements of holotype: AW-63.5, PW-76, 
SB-26.5, ASB-38, PSB-35, SD-73, AP-28, AM^O.5, AL-47.5, PL-58.5, 

Distribution and hosts: Kochi, Tottori, Shimane, Kumamoto, Oita 
and Hyogo prefectures from Apodemus speciosus, Clethrionomys smithii 
and Rattus norvegicus. 

Type data: From Rattus norvegicus, Kochi City Park, Kochi Prefec- 
ture; November 14, 1951; collected by F. Kawashima; deposited at the 
Institute for Infectious Diseases. 

Subgenus Neotrombicula Hirst, 1925 

This group of species is easily recognized by several morphological 
characters: scutum more or less pentagonal with the posterior margin 
broadly projected (rounded or bluntly pointed), leg III with at least one 
mastitarsala, unisetose coxae, and sternal setae 2-2; the palpal claw is 



three-pronged, and the lateral prong is usually conspicuously divergent. 
Brennan and Wharton (1950) recently discussed the North American spe- 
cies of Neotromhicula, and separated them into several groups: the "au- 
tumnalis", "hisignata", and "microti" groups of Brennan and Wharton are 

Fig. 21 TromhicuTa microti 


all represented in Japan. They are parasitic on a variety of small mammals 
and seem to have no well developed host specificities. Of the six species of 
the subgenus Neotromhicula known from Japan, five (T. (N.) tamiyai, T. 
(N.) pomeranzevi, T. (N.) microti, T. (N.) nagayoi, and T. (N.) japonica) 
also occur on the adjacent Asiatic mainland, and two (T. (N.) pomeranzevi 
and T. (N.) microti) are known to occur in North America, 

Key to Japanese Species of Neotrombicula 

1. One mastitarsala III (-autumnalis" group) 2 

— One mastitibiala III and two or three mastitarsalae III 3 

2. Sensillary bases in advance of posterolateral setae; usually one pair of 
humeral setae T. (N.) nagayoi 

— Sensillary bases on a line with posterolateral setae; usually two pairs of 
humeral setae T. (N.) japonica 

3. One mastifemorala III, one mastitibiala III, and two mastitarsalae III 
{''microti" group) 4 

No mastifemorala III, one mastitibiala III, and three mastitarsalae III 

{"Hsignata" group) 5 

4. More than 50 dorsal setae; palpal genual and femoral setae feathered 

T. (N.) pomerayizevi 

Fewer than 30 dorsal setae; palpal genual and femoral setae nude 

T. (N.) microti 

5. Sensillae branched on middle third only; coxa III oval T. (N.) mitamurai 

— Sensillae branched on distal two-thirds ; coxa III elongate T. (N.) tamiyai 

Trombicula (Neotrombicula) japonica Tanaka et al., 1930 

(Figure 17) 

Trombicula autumnalis japonica Ta.naka, Kaiwa, Teramira, and Kagaya, 1930, 
Zentralblatt Bakteriologie, Abt. 1, Band 116, p. 353. 

Diagnosis : Galeal seta usually nude, sometimes forked. Palpal femoral, 
genual, and ventral tibial setae feathered; lateral tibial seta forked or 
nude; dorsal tibial seta nude. Basal third of sensillae nude; distal two- 
thirds with moderate branches. Sensillary bases approximately on a line 
with posterolateral setae. Usually two pairs of humeral setae. Dorsal setae 
rather long, with short barbs. Scutal measurements of a topotype : AW-81.5, 
PW-100, SB-34, ASB-37.5, PSB-24, AP-31, AM-70, AP-67.5, PL-82.5, 
S-80.0 (sensillae measured on a different, but topotypical specimen). 

Distribution and hosts : Known from Akita, Niigata, Yamagata, Yama- 
iiaslii, Ishikari, Nagano, Kyoto, Kumamoto, and Oita prefectures; collected 



from Apodemus geisha A. speciosus, Clethrionomys rufocanus, and Microtus 
monteheUoi. Known also from southern Korea. 

Type data: Described from Microtus monteheUoi (?) taken at Yuzawa, 
Akita Prefecture. 



Fig. 22 TromMcula pomeranzevi 


Trombicula (Neotrombicula) nagayoi Sasa et al., 1950 

(Figure 18) 

Tromhicula nagayoi Sasa, Hayashi, Sato, Miura, and Asahina, 1950, Tokyo Iji 
Shinshi, vol. 67, no. 12. p. 14. 

Diagnosis : The gnathosomal features of T. nagayoi are similar to those 
of T. japonica, but generally there are fewer branches on the setae. Palpal 
femoral seta feathered; palpal genual seta and ventral tibial seta branched; 
dorsal and lateral tibial setae nude. Galeal seta nude. Sensillary bases 
distinctly forward of the posterolateral setae. One pair of humeral setae. 
Dorsal setae shorter than in T. japonica and with more barbs. Of the three 
scuta illustrated (fig. ]8), the uppermost is from a paratype slide; the 
lower two scuta were from a collection in which the scuta were slightly 
larger and in 12 of 24 specimens bore supernumerary scutal setae. Such 
aberrations are not infrequent and T. nagayoi is especially variable in this 
respect. Scutal measurements of holotype: AW-72, PW-89, SB-30, 
ASB-27, PSB-29, AP-31, AM^5, AL-41, PL-57, S-75. 

Distribution and hosts: Yamanashi, Iburi, Kyoto, and Eniwa pre- 
fectures from Apodcmus speciosus, Clethrionomys mfocanvs, and Microtus 
monteheUoi. Known also from southern Korea. 

Type data: From Apodemus speciosus, Yamanaka, Yamanashi Pre- 
fecture. Deposited at Institute for Infectious Diseases, University of Tokyo. 

Trombicula (Neotrombicula) mitamurai Sasa ef al., 1950 

(Figure 19) 

Trombicula viitamurai Sasa, Hayashi, Ktjmada, and Teramura, 1950, Tokyo Iji 
Shinshi, vol. 67, no. 11, p. 18. 

Diagnosis: All palpal setae branched or feathered. Galeal seta feath- 
ered. Cheliceral base and segments of palpi dorsally with small punctae. 
Sensillary bases behind posterolateral setae. Sensillae nude basally and 
distally, with 4-6 branches along middle third. With one or two pairs of 
humeral setae. Coxa II shaped as illustrated, distinctly more oval than 
that of T. tamiyai. Scutal measurements of holotype: AW-59, PW-84.5, 
SB-26, ASB-31, PSB-29, AP-26, AM-48, AL-54.5, PL-58, S-81. 

Distribution and hosts: This species is known from Kanagawa, Shi- 
zuoka, Oita, and Yamanashi prefectures from Urotrichus talpoides, 
Apodemus speciosus, and Microtus monteheUoi. 

Type data : Holotype and one paratype from Urotrichus talpoides at 
Yamanaka (slope of Mt. Fuji), Yamanashi Prefecture; deposited at the 
Institute for Infectious Diseases, University of Tokyo. 


Trombicula (Neotrombicula) tamiyai Philip and Fuller, 1950 

(Figure 20) 

Trombicula tamiyai Philip and Fcller, 1950, Parasitology, vol. 40, p. 51. 

Diagnosis : Generally quite similar to T. mitamurai Sasa et al., but with 
the distal half or two-thirds of the sensillae with eight or more branches. 

A /'^■^•^ 

Fig. 23 Trombicula iHilpaUs 


Coxa III more elongate than in T. mitamurai. Scutal measurements of type 
series as given by describers: "AW-65.5, PW-80, SB-25.6, ASB-28.8, 
PSB-30, AP-19.2, AM-48, AL-t2.5, PL-60.8, S-73.6." 

Distribution and hosts: Yamagata, Niigata, and Akita prefectures; 
from Microtus montebelloi. Known also from southern Korea, from a 
variety of small rodent hosts. 

Type data: Holotype from Microtus monteheUoi from Okiage Village, 
Yamagata Prefecture; September 17, 1920. Type in the United States 
National Museum, 

Trombicula (Neotrombicula) microti Ewing, 1928 

(Figure 21) 

Trombicula microti Ewing, 1928, Proceedings of the Entomological Society of 
Washington, vol. 30, pp. 77-80. 

Diagnosis: Similar in some respects to T. pomeranzevi Schluger, es- 
pecially in the vestiture of the legs. Palpal genual and femoral setae nude; 
dorsal seta on palpal tibia branched, lateral seta nude, and ventral seta 
branched. Galeal seta branched or feathered. Sensillary bases slightly in 
front of posterolateral setae; sensillae with scattered barbs along most of 
their lengths. Twenty-four to twenty-six dorsal setae in Japanese speci- 
mens. Scutal measurements of a specimen from Honshu : AW-47, PW-90, 
SB-28, ASB-33, PSB-28, AP-32, AM^6, AL-52, PL-56, S-84. Japanese 
specimens differ from North American specimens chiefly in the distribution 
of barbs along the sensillae. 

Distribution and hosts : In Japan this species has been collected from 
Aomori Prefecture on Honshu and Daisetsu, Hokkaido. Known from many 
kinds of small rodents and insectivores from many localities in the 
United States and Canada. Dr. Kiyoshi Asanuma has collected this 
species from eastern Manchuria. 

Type data : Type from Microtus richardsoni, Lincoln County, Wyo- 
ming, U. S. A.; August 13, 1927; deposited in the United States National 

Trombicula (Neotrombicula) pomeranzevi Schluger, 1948 

(Figure 22) 

Tromhicula pomeranzevi Sciiluoer, 1948, Entomologicheskoe Obozrenie, (Moscow), 

tome 30, p. 157. 
Tromhicula alaskensis Brennan and Whartox, 1950, American Midland Naturalist, 

vol. 45, p. 178, neic synonymy. 



Diagnosis: All palpal setae (except ventral tibial seta) feathered; 
galeal seta nude. Sensillary bases slightly behind posterolateral setae. 
Sensillae with minute barbs on the basal half. Fifty or more dorsal setae. 
There appear to be no characters by which T. alaskensis can be separated 

Fig. 24 Tronibicula yasnokai 



from T. pomeranzevi. North American specimens of T, alaskensis were 
compared with T. pomeranzevi from Hokkaido, and there are no differences 
of specific merit. Dr. Brennan examined some of the Japanese specimens 
of T. pomeranzevi and concurred with our opinion. Scutal measurements 




Fig. 25 Trombicula himizu 



of a specimen from Hokkaido: AW-85.0, PW-112.5, SB-32.5, ASB-42.5, 
PSB-32.5, AP-36, AM-62.5, AL-65, PL-75, S-112.5. 

Distribution and hosts: Known from Clethrionomys rutilis and 
Apodemus speciosus from Aomori and Ishikari prefectures. Known also 
from U.S.S.R. and North America. 

Type data: Described from Clethrionomys rufocanus and Apodemus 
speciosus from South Shore, vicinity of Barrabasha, U.S.S.R.; June 19, 
August 5, and September 24, 1940. 

Fig. 26 Trombicula miyajimai 


Subgenus Leptotrombidium Nagayo et al., 1916 

In Leptotrombidium the palpal femoral and genual setae are nude; the 
palpal claw is three-pronged (the prongs being closely appressed) ; the 
galeal seta is feathered; the scutum more or less rectangular; the sternal 
setae 2-2; the coxal setae 1-1-1; and there are no mastitarsalae III. Im- 
portant specific characters in this group are the shape of the scutum, posi- 
tion of sensillary bases, character of the sensillae and scutal setae, number 
and character of dorsal body setae, and the position of the seta on coxa III. 
There are 18 species of Leptotrombidium in Japan, a rather large con- 
centration of forms: this figure does not include several probable syno- 
nyms; and, as there are doubtless others to be discovered, our list is con- 
servative. Only four of these species {T. palpalis, T. akamushi, T. pallida, 
and T. fxLJi) are known to occur outside of Japan. The species of Lepto- 
trombidium are usually common within their geographic range: T. (L.) 
fuji, T. (L.) kitasatoi, T. (L.) palpalis, T. (L.) intermedia, and T. (L.) 
scutellaris have a rather broad geographic distribution and are commonly 
collected. Trombicula akamushi, so frequently mentioned in accounts of 
tsutsugamushi disease and the chief vector of this disease in Japan, is ac- 
tually confined to a small area in Japan and only locally common. Although 
most species seem to have no well developed host specificity, a few species 
seem closely associated wdth one kind of mammal: Trombicula himizu is 
typically a parasite of the shrew-mole (Tfrotrichus talpoides) as is T. 
miyairii and T. tenjin, and bats (Myotis macrodactylus) are the only known 
host of T. toshiokai. 

Key to Japanese Species of Leptotrombidium 

1. Ventral palpal tibial seta feathered 2 

— Ventral palpal tibial seta nude 5 

2. Lateral palpal tibial seta feathered T. (L.) miyajimai 

— Lateral palpal tibial seta nude 3 

3. Anterolateral and posterolateral setae similar T. (L.) palpalis 

— Posterolateral setae much heavier than anterolateral setae 4 

4. Posterolateral setae rasp-like, barbs short T. (L.) yasuokai 

— Posterolateral setae pectinate, barbs long T. (L.) himizu 

5. Seta on coxa III on anterior margin of coxa 6 

— Seta on coxa III markedly behind the anterior margin of coxa 7 

6. First post-humeral row of setae usually ten T. (L.) tanaka-ryoi 

— First post-humeral row of setae usually eight T. (L.) fuji 

7. Posterolateral setae in the angulate corners of the scutum 8 

— Posterolateral setae in the anterior part of the rounded posterior corners 

of the scutum 9 



8. Sensillary bases in advance of the posterolateral setae; posterior margin 

of the scutum more or less straight T. (L.) aJcamushi 

— Sensillary bases about on a line with posterolateral setae; posterior margin 
of scutum evenly rounded t. (L.) scutellaris 

Fig. 27 Trombicula akamushi 


9. Bases of sensillae nude lOi 

— Bases of sensillae barbed 15 

10. Sensillary bases about on a line with posterolateral setae 11 

— Sensillary bases clearly behind a line connecting posterolateral setae 13 

11. Ventral palpal tibial seta shorter than the palpal genual seta; first post- 
humeral row of setae usually eight T. (L.) kuroshio 

— Ventral palpal tibial seta as long or longer than the palpal genual seta; 
first post-humeral row of setae usually ten 12 

12. Scutum about twice as wide as long; dorsal setae rather long, with the 
barbs rather appressed T. (L.) intermedia 

— Scutum about two and one-half times as wide as long; dorsal setae i-ather 
short and bushy T. (L.) teramurai 

13. Scutum more than twice as wide as long; usually two pairs of humeral 
setae T. (L.) miyazakii 

— Scutum less than twice as wide as long; a single pair of humeral setae 14 

14. Dorsal setae with barbs short T. (L.) tenjin 

— Dorsal setae with barbs very long (and few in number) T. (L.) miyairii 

15. Basal barbs of sensillae divergent, conspicuous 18 

— Basal barbs of sensillae delicate and appressed 16 

16. Sensillary bases markedly behind a line connecting the posterolateral 
setae T. (L.) miyairii 

— Sensillary bases approximately on a line connecting the posterolateral 
setae 17 

17. Scutum slightly less than twice as wide as long; ventral and lateral palpal 
tibial setae subequal T. (L.) toshiokai 

— Scutum slightly more than twice as wide as long; ventral palpal tibial 
seta much longer than lateral palpal tibial seta T. (L.J intermedia 

IS. First post-humeral row of setae 12-15; dorsal setae shorter than length of 
scutum - T. (L.) iKillida 

— First post-humeral row of setae 8-10; dorsal setae longer than scutum 19 

19. Dorsal setal formula 2-10-8-8-6-4-2 T. (L.) tosa^ 

— Dorsal setal formula 2-8-6-6-2-2 T. (L.J kitasatoi 

Trombicula (Leptotrombidium) palpalis Nagayo et al., 1919 

(Figure 23) 

Tromhicula palpalis Nagayo, Mitamira, and Tamiya, 1919, Jikken Igaku Zasshi, 
vol. 3, no. 4, pp. 265-312. 

Diagnosis : The ventral palpal tibial seta is feathered. Sensillary bases 
about on a line with the posterolateral setae. Sensillae basally nude (or 

1. Species in which the barbs are very minute will key out in both halves of this couplet. 

2. Possibly a synonym of T. kitasatot. 



with a few very small barbs), with numerous barbs on the distal half. 
Scutum approximately twice as broad as long; posterior corners well 
rounded, and posterolateral setae in the anterior part of these corners, well 
in advance of the posterior margin of the scutum. Specimens from moun- 
tain areas have a tendency to have more dorsal setae than specimens from 

Fig. 28 Trombicula pallida 


valleys and low grasslands. In such slightly aberrant mountain specimens, 
the rows of setae will be crowded and irregular. 

Distribution and hosts: Saitama, Nanagawa, Gumma, Yamagata, 
Akita, Okayama, Yamanashi, and Ishikari prefectures from Microtus 
monteheUoi, Clethrionomys rufocanus, C. smithii, Apodemus speciosus, 
Rattus norvegicus, Urotrichus talpoides, and Passer montanus. This chig. 
ger has also been collected in southern Korea. 

Type data: A leetotype has been selected from specimens collected 
from Microtus monteheUoi by the original authors at Yachi, Yamagata; 
March 27, 1919; deposited at Institute for Infectious Diseases, University 
of Tokyo. 

Trombicula (Leptotrombidium) yasuokai Sasa et al., 1952 

(Figure 24) 

Trombicula yasuokai Sasa, Kawashima, and Hiromatsu, 1952, Tokyo Iji Shinshi, 
vol. 69, no. 1, p. 43. 

Diagnosis : Ventral tibial seta of palpus feathered. Cheliceral base with 
scattered punctae. Sensillary bases markedly behind posterolateral setae. 
Sensillae basally nude, with scattered barbs on distal half. Hind margin of 
scutum more or less three sided : straight immediately behind and between 
the sensillae, and directly anteriorly near the posterolateral setae. Postero- 
lateral and anteromedian setae distinctly heavier than anterolateral setae 
and with rasp-like, heavy barbs. Scutal measurements of holotype: 
AW-62.5, PW-81, SB-35, ASB-30, PSB-11.5, AP-19.5, AM-69.5, 
AL-43.5, PL-72.5, S-66.5 

Distribution and hosts : Knowli from Kochi Prefecture from Mogera 

Type data : Holotype and one paratype from Sakiyama, Muroto-maehi, 
Kochi Prefecture from Mogera ivogura; collected by Y. Hiromatsu, August 
26, 1951. Holotype and paratype deposited at Institute for Infectious 
Diseases, University of Tokyo. 

Remarks: The original illustration shows the sensillary barbs to be 
sliort and scale-like, and the scutum to be evenly rounded on the hind mar- 
gin near the posterolateral setae. The actual condition of the holotype is 
shown in the accompanying figure. 

Trombicula (Leptotrombidium) himizu Sasa et al., 1951 
(Figure 25) 
Trombicula himizu Sasa, Kumada, Hayasiii, Enomoto, Fukuzumi, and Obata, 
1951, Eisei Doobutsu, vol. 2, no. 1, pp. 1-5. 

Diagnosis: Ventral tibial seta of palpus feathered. Sensillary bases 
behind posterolateral setae. Sensillae basally nude, with scattered barbs 



distally. Posterolateral setae in advance of the rounded posterior corners 
of the scutum. Anteromedian and posterolateral setae with very heavy 
shafts and rather stout barbs. The numerous dorsal setae similar in nature 
to the anteromedian and posterolateral setae of the scutum; the small 
number and large size of the barbs on these setae serve to separate T. (L.) 
himizu from the other species of Leptotromhidium, with a feathered ventral 
tibial seta of the palpus. Scutal measurements of the holotype: AW-60.5, 
PW-72, SB-32, ASB-27.5, PSB-10.5, AP-19, AM-67, AL^4.5, PL-60.5, 

Distribution and hosts : Kanagawa, Shizuoka, and Yamanashi prefec- 
tures, from TJrotrichus talpoides, Apodemus speciosus, and Microtus monte- 

Type data : Holotype from Mt. 0.yama, Kanagawa Prefecture, from 
TJrotrichus talpoides; November 23, 1950. Three paratypes from the same 
host from Yamanaka, Yamanashi Prefecture, November 20, 1950. These 


Fig. 29 Trombicula tanaka-ryoi 


specimens are deposited at the Institute for Infectious Diseases, University 
of Tokyo; one specimen (paratype) deposited at the Kitasato Institute, 

Trombicula (Leptotrombidium) miyajimai Fukuzumi and Obata, 1951 

(Figure 26) 

Tromhicula miyajimai FrKUZUMi and Obata, 1951, Kitasato Arcliives of Experi- 
mental Medicine, vol. 23, p. 5, 

Diagnosis: This is the only species of Leptotrombidium in Japan in 
which all the palpal tibial setae are feathered. The cheliceral base and, 
dorsally, the segments of the palpus bear numerous punctae. Sensillary 
bases about on a line with posterolateral setae. Posterolateral setae set in 
the rather sliarp corners of the scutum. Hind margin of the scutum pro- 
jecting slightly in the area behind and between the sensillae. Sensillae 
basally nude and distally with 8-12 barbs. Posterolateral setae rather 
heavy and with sliort fine barbs. Dorsal setae long and similar to postero- 
lateral setae of scutum. Scutal measurements of holotype: AW-71.8, 
PW-86.8, AP-28.3, ASB-35, PSB-14, SB-31.7, AM-57.9, AL-46.1, 
PL-80.4, S-64.2. 

Distribution and hosts: Kanagawa, Kyoto, Chichibu, Okayama, 
Gumma, and Kochi prefectures from Apodemus speciosus, A. geisha, Cleth- 
rionomys smithii, and Microtus monteheUoi. 

Type data: Holotype and five paratypes from Apodemus speciosus 
from Matsuda, Kanagawa Prefecture; October 9, 1950. Holotype and para- 
types deposited at the Kitasato Institute. 

Trombicula (Leptotrombidium) akamushi (Brumpt, 1910) 

(Figure 27) 

Trombidium akamushi Brumpt. 1910, Precis de Parasitologie. 2nd ed.. p. 506. Paris. 

Diagnosis: Palpal femoral and genual setae nude; lateral and ventral 
palpal tibial setae nude; dorsal palpal tibial seta feathered. Galeal seta 
feathered. Cheliceral base and palpal femur with numerous small punctae. 
Sensillary bases well in advance of the posterolateral setae {T. (L.) aka- 
mushi is unique in this respect among Japanese species of Leptotrom- 
bidium). Sensillae nude basally, plumose distally. Posterolateral setae 
placed in the angulate corners of the scutum. Seta of coxa III markedly 
behind the anterior margin of coxa. Scutal measurements (mean of five 
specimens from Akita Prefecture, from Fukuzumi and Obata, 1951, Kita- 
sato Archives of Experimental Medicine, 23:8): AW-65.5, PW-73.3, 



AP-28.6, ASB-18.9, PSB-17.1, SB-28.9, AM-53.3, AL-42.9, PL-51.7, 

Distribution and hosts: This species in Japan seems to occur only 
along the grassy areas in the valleys of the four main rivers in northwest- 

Fig. 30 Trombicula fuji 


ern Honshu : the Shinano and Aoano rivers in Xiigata Prefecture, the 
Mogami River in Yamagata Prefecture, and the Omono River in Akita 
Prefecture. The most common host is Microtus monteheUoi, but Rattus 
norvegicus, Apodemus speciosus, and several species of wild birds are also 

Type data: From man; no type locality was designated, but Tanaka 
collected his original material from Yuzawa, Akita Prefecture 

Remarks: As Philip (1947, American Journal of Hygiene, vol. 46, p. 
60) pointed out, the original description of T. akamushi consists of a poorly 
executed drawing. It is uncertain what species is intended from the illus- 
tration, but the species illustrated here is one that workers today generally 
agree to call Tromhicula akamushi. It has long been accepted in the liter- 
ature as the vector (and more recently, the major vector) of tsutsugamushi 
disease in Japan; there is nothing to be gained by a change of names. 
Actually, the identification of the species shown by Brumpt is probably no 
more difficult tlian that of many original descriptions of that period. The 
feathered condition of the palpal femoral and genual setae, clearly shown 
in the original figure, indicate that the species shown was not a species of 
LeptotromMdium. Quite possibly the species shown is Tromhicula (Neo- 
tromhicula) joponica. Apparently early workers identified what we now 
call T. akamushi without referring to the original description, thus giving 
rise to a misidentification which workers have followed ever since. The 
present assignment of names should stand. 

Trombicula (Leptotrombidium) pallida Nagayo et al., 1919 

(Figure 28) 

Tromhicula pallida Nagayo, Mitamira, and Tamiya, 1919, Jikken Igaku Zasshi, 

vol. 3, pp. 265-312. 
TromMcula burnsi Sasa, Teramura, and Kano, 1950, Tokyo Iji Shinshi, vol. 67, 

no. 10, p. 22. Neiv synonymy. 
TromMcula murotoensis Sasa and Kawashima, 1951, Tokyo Iji Shinshi, vol. 68, 

no. 12, p. 16. Nexo synonymy. 

Diagnosis: Scutum more or less rectangular, with the posterolateral 
setae well in the anterior part of the rounded posterior corners. Sensillary 
bases slightly behind a line connecting the posterolateral setae. Sensillae 
with conspicuous, and divergent (though small) basal barbs, distal two- 
thirds with branches. The dorsal setae in T. pallida are numerous, with 12-15 
in the first post-humeral row; they are distinctive in being rather heavy 
and short with numerous barbs. Seta on coxa III placed well behind the 
anterior margin of the coxa. Tromhicula pallida is highly variable, espe- 
cially with regard to the number of dorsal setae. Scutal measurements of 



an average specimen : AW-72, PW-78, SB-36, ASB-33, PSB-15, AP-20, 
AM-54, AL-45, PL-72, S-60. 

Distribution and hosts : This is a common and widely distributed spe- 
cies in Japan, and is known to occur also in southern Korea. Yamagata, 
Kanagawa, Saitama, Akita, and Kyoto prefectures, and Oshima and Miyake 
islands from Crocidura dsi-nezumi, Apodemus speciosus, Clethrionomys 
smithii, Rottus norvegicus, Microtus montehelloi, and small birds. 

Type data : Lectotype selected from specimens collected by the original 
authors at Yachi, Yamagata Prefecture; March 27, 1919, from Microtus 
moniebeUoi; deposited at tlie Institute for Infectious Diseases, University 
of Tokyo. 

Remarks: Tromhicula hurnsi and T. murotoe7isis are regarded as syno- 
nyms of T. pallida. 

* 1/ '■ 

Fig. 31 Tromhicula kitasatoi 


Trombicula (Leptotrombidium) tanaka-ryoi Kawashima and Sasa, 1952 

(Figure 29) 

Trombicula tanaka-ryoi Kawashima and Sasa, 1952, Tokyo Iji Shinshi, vol. 69, 

no. 2, p. 21. 
Trombicula tanaka-ryoi Mitsitomi, Sasa, Hayasiii, Eto. Kobayasiii, and Kitahaha. 

1952, Tokyo Iji Shinshi, vol. 69, no. 4, p. 18 (Original illustration). 

Diagnosis: Very close to T. fuji Kuwata et al., 1950. TromhicuJn 
tanaka-ryoi differs in the shorter genual palpal seta, the larger scutum, and 
the dorsal setal formula. The scuta of the two species are remarkably 
similar, but in T. tanaka-ryoi the scutum is approximately twice as wide 
as long whereas in T. fuji it is less than twice as wide as long. The sensil- 
lary bases in both species are far behind a line connecting the posterolateral 
setae, and the sensillae possess conspicuous, divergent basal barbs in both 
species. In T. tanaka-ryoi the first post-humeral row of dorsal setae is 
usually ten and in T. fuji it is usually eight. In both species the seta on 
coxa III arises from the anterior margin of the coxa. Scutal measurements 
of holotype : AW-64, PW-64, SB-29, AP-17.5, SD-43, PSB-13, ASB-30, 
AM-44.5, AL-36, PL-62.5, S-56.5. 

Distribution and hosts: Okayama, Kochi, Kyoto, Kumamoto, Oita, 
Shiga, Ehime, Mie, Tottori, Shimane, and Nara prefectures from Crocidura 
dsi-nezumi, Apodenius speciosus, and Clethrionomys smithii. 

Type data : Holotype from Mt. Ishiguchi, Ehime Prefecture, from 
Clethrionomys smithii; collected by Eyo Tanaka; September 10, 1951, One 
paratype with the same data, and three paratypes from Ajjodemus speciosus 
with the same data. Deposited at the Institute for Infectious Diseases. 

Trombicula (Leptotrombidium) fuji Kuwata et al., 1950 
(Figure 30) 

Trombicula (Leptotrombidium) fuji Kuwata, Berge, and Philip, 1950, Journal of 
Parasitology, vol. 36, p. SO. 

Diagnosis: The very long palpal genual seta of T. fuji is distinctive. 
The scutum is small and the sensillary bases are placed far behind a line 
connecting the posterolateral setae. The first post-humeral row of setae is 
almost always of eight setae. The seta on coxa III arises from the anterior 
margin of the coxa. ( See also diagnosis for T. tanaka-ryoi. ) Scutal measure- 
ments of holotype: AW-50.5, PW-52, SB-24, AP-14, SD-30.5, PSB-11, 
PL-SB-10.5, AM-29.5, AL-28.5, PL-51, S-40.5. 

Distribution and hosts: Aomori, Akita, Yamagata, Niigata, Gumma, 
Saitama, Chiba, Tokyo, Kanagawa, Yamanashi, Nagano, Hyogo, Okayama, 
Shimane, Oita, Kumamoto, Shizuoka, Kyoto, Mie, Shiga, Fukuoka, and 
Kochi prefectures from Apodemus speciosus, A. geisha, Rattus norvegicus, 



R. rattus, Clethrionomys smithii, and Microtus montehelloi. Known also 
from the Pescadores. 

Type data : From Apodemus speciosus from east slope of Mt. Fuji, 
Fujino Susuno (near Gotemba), Shizuoka Prefecture; October 30, 1948, 
collected by Major Trygve 0. Berge; deposited at the United States Na- 
tional Museum. 

Trombicula (Leptotrombidium) kitasatoi Fukuzumi and Obata, 1950 

(Figure 31) 

Trombicula kitasatoi Fukuzumi and Obata, 1950, Kitasato Archives of Experi- 
mental Medicine, vol. 23, no. 3, p. 79. 

Diagnosis: Scutum more than twice as wide as long; posterolateral 
setae in the anterior part of tlie rounded corners of the scutum. Sensillary 

Fig. 32 Trombicula tosa 


bases about on a line with posterolateral setae; basal third of sensillae 
with conspicuous, divergent barbs. First post-humeral row of dorsal setae 
usually eight. Seta of coxa III markedly behind the anterior margin of 
the coxa. Scutal measurements of holotype: AW-70.4, PW-79.2, AP-22.0, 
SB-33.9, ASB-25, PSB-15, AM-51.7, AL-40.0, PL-73.3, S-53.2. 

Distribution and hosts: Common and widely distributed. Saitama, 
Yamagata, Fukushima, Kanagawa, Chiba, Gumma, Kyoto, Shiga, Mie, 
Okayama, Fukuoka, Kagoshima, Kumamoto, Oita, and Kochi prefectures 
from A2)odemus speciosus, A. geisha, Rattus norvegicus, R. rattus, Cleth- 
rionomys smithii, and Microtus montehelloi. 

Type data: From Ai)odcmus speciosus from Tsurumi, Kanagawa Pre- 
fecture; September 9, 1950; collected by the original describers; deposited 
at the Kitasato Institute, Tokyo. 

Trombicula (Leptotrombidium) tosa Sasa and Kawashima, 1951 

(Figure 32) 

TromMcula tosa Sasa and Kawashima, 1951, Tokyo Iji Shinshi, vol. 68, no. 10, p. 10. 

Diagnosis: Very similar to T. kitasatoi, differing only in the dorsal 
setal formula: in T. tosa it is 2-10-8-8-6-4-2 and in T. kitasatoi it is 
2-8-6-6-2-2. In the original description the enlarged drawing of the 
scutum shows the hind margin to be evenly rounded but the holotype is 
as illustrated in the accompanying figure in this paper. Scutal measure- 
ments of holotype: AW-70, PW-78, SB-32, ASB-29, PSB-13, SD-42, 
AP-22, TM-52, AL-37, PL-65, S-56. 

Distribution and hosts: From Kochi Prefecture from Rattus norve- 
gicus and R. Rattus. This species is abundant in sweet potato fields and 
residential areas in the coast-line area of western Kochi. Trombicula tosa 
occurs exclusively in the summer months; from near-by areas of scrub and 
forest, T. kitasatoi occurs from autumn to spring. 

Type data : Holotype and five paratypes from Rattus norvegicus from 
Ida, Kochi Prefecture; June 29, 1951; deposited at the Institute for Infec- 
tious Diseases. 

Remarks : When the biology and complete life histories of T. kitasatoi 
and T. tosa are better known, the latter may prove to be a biological variant 
or a synonym of T. kitasatoi. Among a series of specimens mounted on a 
single slide, five clearly belong to T. tosa and one; definitely to T. kitasatoi. 

Trombicula (Leptotrombidium) miyairii Sasa et aJ., 1952 

Figure 33) 

TromMcula miyairii Sasa, Hayashi, Kawashima, Mitsutomi, and Egashira, 
1952, Tokyo Iji Shinshi, vol. 69, no. 3, p. 11. 



Diagnosis : Gnathosoma similar to that of T. akamushi. Scutum small, 
about twice as wide as long; with posterior margin projected consid- 
erably behind the posterolateral setae. Sensillary bases considerably 
behind posterolateral setae. Bases of sensillae with minute barbs; distal 
half with two or three very long branches. Posterolateral setae heavy, 
with the shaft thick and the branches heavy and long. Dorsal bodv setae 

Fig. 33 TroniMcula m.iyairii 


similar to posterolateral setae. Seutal measurements of holotype : AW-59, 
P\V-60, SB-20, ASB-31, PSB-14, AP-15, AM^O.5, AL-35, PL-62, S-52. 

Distribution and hosts : To date all specimens have been collected from 
Urotrichus talpoides from Kumamoto, Oita, and Ehime prefectures. 

Type data: Holotype from Urotrichus talpoides from Toshita (near 
Mt. Aso), Kumomoto Prefecture; October 21, 1951; three paratypes with 
the same data; two paratypes from Mt. Ishijuchi, Ehime Prefecture; Sep- 
tember 8, 1951. Holotype and all paratypes deposited at Institute for 
Infectious Diseases. 

Trombicula (Leptotrombidium) tenjin Sasa et ah, 1951 

(Figure 34) 

Trombicula tenjin Sasa, Hayashi, Kumada, and Mu'ra, 1951, Tokyo Iji Shinshi, 
vol. 68, no. 3, p. 17. 

Diagnosis: Gnathosoma similar to that of T. akamushi; the branched 
palpal setae with fewer and longer branches. Scutum about twice as wide 
as long; with numerous scattered punctae; posterior margin projected far 
behind posterolateral setae, with a slight mesal concavity. Sensillary bases 
far behind a line connecting the posterolateral setae. Sensillae rough and 
rasp-like basally, with six to eight rather long barbs on the distal half or 
two-thirds. Barbs on seutal setae rather heavy, not especially long. Dorsal 
setae similar to seutal setae. Seutal measurements of holotype: AW-63.0, 
PW-70.0, AP-21.5, ASB-28.0, PSB-10.0, SB-30.5, AM-41.0, AL-38.0, 
PL-80.5, S-48.5. 

Distribution and hosts : Known from Kanagawa Prefecture from Uro- 
trichus talpoides. 

Type data: From Urotrichus talpoides from Okuyugawa, Kanagawa 
Prefecture; November 12, 1950; deposited at the Institute for Infectious 

Trombicula (Leptotrombidium) teramurai Sasa et al., 1951 

(Figure 35) 

Trnmhicnla teramurai Sasa, Kumada, and Teramura, 1951, Tokyo Iji Shenshi, vol. 
68, no. 9, p. 8. 

Diagnosis : Gnathosoma similar to that of T. akamushi. Scutum more 
than twice as wide as long (almost three times as wide as long); lateral 
margins markedly concave; posterior margin projected considerably behind 
posterolateral setae, but slightly concave mesally; with scattered punctae. 
Sensillary bases slightly behind a line connecting posterolateral setae. 



Bases of sensillae nude; distal two-thirds plumose. Dorsal setae rather 
short and plumose, as illustrated. Scutal measurements of holotype: 
AW-70, PW-80, SB-31, ASB-28, PSB-14, AP-17, AM-59, AL-38, 
PL-58.5, S-67. 

Fig. 34 Trombicula tenjin 


Distribution and hosts: From Niigata, Akita, and Yamagata from 
Microtus montehelloi, Apodemus speciosus, and TJrotrichus talpoides. 

Type data : Holotype and twelve paratypes from Microtus montehelloi 
from Akakura, Niigata Prefecture; April 24, 1951; deposited at Institute 
for Infectious Diseases. 

Eemarks: The original illustration of the scutum errs in showing it 
too narrow; the illustration in this paper, made from the holotype, is 

Trombicula (Leptotrombidium) kuroshio Sasa and Kawashima, 1952 

(Figure 36) 

Trombicula kuroshio Sasa and Kawashima, 1952, Tokyo Iji Shinshi, vol. 68. no. 12, 
p. 15. 

Diagnosis: Gnathosoma similar to that of T. akamushi; genual seta 
distinctly longer than ventral tibial seta. Scutum more than twice as wide 
as long; posterior corners more or less rounded. Sensillary bases distinctly 
behind a line connecting posterolateral setae. Base of sensillae nude, distal 
two-thirds plumose. The posterior margin of the scutum is usually more 
or less three-sided, sometimes irregularly rounded. Scutal measurements 
of holotype: AW-55, PW-63, SB-24, ASB-27, PSB-10, AP-16, AM-37, 
AL-32.5, PI-59, S-47. 

Distribution and hosts : From Kagoshima, Kumamoto, Oita, and Shiga 
prefectures, from Apodemus speciosus and Clethrionomys smithii. 

Type data: Holotype and seven paratypes from Apodemus speciosus 
from Maruyama, Muroto-machi, Kochi Prefecture; August 26, 1951; de- 
posited at the Institute for Infectious Diseases. 

Trombicula (Leptotrombidium) toshiokai Sasa and Jameson, new species 

(Figure 37) 

This species is one of two kinds of chiggers collected from bats in Japan; 
it is known only from this host. 

Gnathosoma: Cheliceral base with a few punctae; a few small punctae 
on dorsal part of palpal femur. Chelicera with a subapical dorsal tooth. 
Palpal setae essentially as in T. akamushi : femoral, genual, and lateral and 
ventral ti1)ial setae nude; dorsal tibial seta feathered. Galeal seta feathered. 

Legs: All coxa unisetosa: seta on coxa III distinctly behind the ante- 
rior margin of the coxa. Specialized (nude) setae : Leg I, 2 genulae, 1 micro- 
genual a, 2 tibialae, 1 mici'otibiala, 1 spur, 1 microspur, 1 subterminala. 



1 parasubterminala, 1 pretarsala; Leg II, 1 genuala, 2 tibialae, 1 spur, 1 
pretarsala; Leg III, 1 genuala, 1 tibiala. 

Scutum: About twice as wide as long (in T. intermedia and T. miya- 
zakii the scutum is at least slightly more than twice as wide as long). Sen- 

Fig. 35 Tromhicula teramurai 



[Proc. 4th See. 

sillary bases slightly behind a line connecting the posterolateral setae. 
Sensillae with minute and inconspicuous basal barbs, distal half plumose. 
Posterior margin of the scutum rounded, concave mesally. Punctae few. 
Scutal measurements of holotype: AW-91, PW-110, SB^8, ASB-42, 
PSB-18, AP-35, AM-77, AL-59, PL-84, S-70. 

Fig. 36 Trombicula kuroshio 




Setae: Dorsal setae quite long, with few barbs. Dorsal setae formula: 
2-14 (12-15) -12-10 ( 10-12 )-8-t. S.ternal setae 2-2. About 45-50 small 
setae behind sternal setae, ventrally. 

Type data: Holotype and ten paratypes from Myotis macrodactylus 
from the bat caves at Nikko, Tuchigi Prefecture; collected by Seiichi Tosh- 
ioka and William Suyemoto, September 4, 1952; two additional specimens 
from the same host from Kyoto Prefecture collected by Yukio Shogaki and 
J. McClendon. Holotype and five paratypes deposited in United States 
National ^Museum; one paratype deposited in Institute for Infectious Dis- 
eases, Universitv of Tokyo. 

i fii-'^'-y^ 

Fig. 37 Tromhicula toshiokai 



Trombicula (Leptotrombidium) miyazakii Sasa et al., 1951 

(Figure 38) 

Trombicula miyazakii Sasa, Sawada, Kano, Hayasiii, and Kumada, 1951, Tokyo 
Iji Shinshi, vol. 68, no. 4, p. 7. 


Fig. 38 Trovihicula miyazakii 



Diagnosis : Gnathosoma similar to that of T. akamushi. Scutum slightly 
more than twice as wide as long; posterolateral setae in the anterior part 
of the rounded posterior corners. Sensillary bases slightly behind a line 

Fig. 39 T7-ombicula inter^nedia 


connecting the posterolateral setae. Basal third of sensillar nude, distal 
two-thirds plumose. First two post-humeral rows of dorsal setae very close 
together, together with about twenty setae. Two pairs (usually) of humeral 
setae. Dorsal setae much more numerous than in most Japanese species of 
Leptotromhidium. Scutal measurements of holotype: AW-80, PW-91, 
AP-25, SB-40.5, ASB-25.5, PSB-17.5, AM-64, AL-46.5, PL-71.5, S-76. 

Distribution and hosts: Gumma, Kanagawa, Saitama, Gifu, Nagano, 
and Ehime prefectures from Apodemus speciosus and Clethrionomys 

Type data : A holotype and a paratype from Apodemus speciosus from 
Ikao, Gumma Prefecture, November 27, 1950; deposited at the Institute for 
Infectious Diseases, University of Tokyo. 

Trombicula (Leptotrombidium) intermedia Nagayo et al., 1920 

(Figure 39) 

Trombicula intermedia Nagayo, Mitamira, and Tamiya, 1920, Verhandelingen 
Japanische Pathologische Gesellschaft, vol. 10, p. 147. 

Diagnosis: Gnathosoma similar to that of T. akamushi, but with the 
ventral tibial seta of the palpus very long. Scutum with the rear margin 
convex with a slight mesal concavity; hind corners rounded; the outline of 
the rear margin is quite variable. Sensillary bases about on a line with the 
posterolateral setae. Basal third of sensillae with minute, inconspicuous 
barbs; distal two-thirds of sensillae plumose. Dorsal setae rather long, 
with barbs short. Scutal measurements (from Fukuzumi and Obata, 1950, 
Kitasato Archives of Experimental Medicine, 23:81) : "AW-68.6, PW-80.7, 
AP-25.3, ASB-20.7, PSB-12.8, SB-36.5, AM-50.6, AL-44.0, PL-62.5, 

Distribution and hosts: This species is very common and widely dis- 
tributed throughout the mountain areas of eastern Honshu and Kokkaido: 
Akita, Yamagata, Niigata, Aomori, Ishikari, Gumma, Yamanashi, Shizuoka, 
Nagano, Kyoto, Shiga, and Oita prefectures. It is known from Urotrichus 
talpoides, Apodemus speciosus, Clethrionomys smithii, C. rufocanus, and 
Microtus montehelloi. 

Type data : The original description is based on specimens collected at 
Arato, Yamagata Prefecture. A lectotype (remounted) selected from the 
original series, was collected from Microtus montehelloi; Arato, Yamagata 
Prefecture; October 20, 1920; deposited at the Institute for Infectious Dis- 
eases, University of Tokyo. 




Trombicula (Leptotrombidium) scutellaris Nagayo et ah, 1921 

Figure (40) 

Trombicula scutellaris Nagayo, Miyagawa, Mitamura, Tamiya, and Tenjin, 1921, 
American Journal of Hygiene, vol. 1, pp. 569-591. (In nos. 5 and 6, Septem- 
ber-November, 1921.) 

Trombicnla scutellaris Nagayo, Mitamura, Tamiya, and Tenjin, 1931, Verhande- 
lingen Japanische Pathologische Gesellschaft, vol. 11, p. 471. (December, 

Diagnosis : Gnathosoma similar to that of T. akamushi; in T. scutellaris 
the ventral tibial seta of the palpus is very long whereas this seta in aka- 
mushi is quite short. Scutum similar to that of T. akamushi in that the 
posterolateral setae are placed in the rather angulate corners of the scutum; 
the sensillary bases are, however, about on a line with the posterolateral 

Fig. 40 Tromhicula scutellaris 


setae (in T. akamushi the sensillary bases are distinctly in advance of the 
posterolateral setae). The rear margin of the scutum is convex and evenly 
rounded. The bases of the sensillae are nude, the distal half plumose. Seta 
on coxa III distinctly behind the anterior margin of the coxa. Scutal meas- 
urements of a topotype (from Kuwata et al., 1950, Journal of Parasitology, 
vol. 36, p. 82) : "AW-69.5, PW-82.5, AP-31.5, SB-31.5, PSB-19.5, AM-62, 
AL-52.5, PL-64, S-76.5." 

Distribution and hosts: Known from Yamagata, Nagano, Niigata, 
Kagoshima, and Shizuoka prefectures and from the Izu Islands (south of 
Tokyo), from Microtus montehelloi, Apodemus speciosus, Rattus norvegi- 
cus, and several species of wild birds. 

Type data: This species was first recognized as a distinct species by 
Kawamura in Niigata, but was described as new by Nagayo et al., based on 
specimens collected at Yachi, Yamagata Prefecture, from Microtus monte- 
belloi; October 24, 1920. A holotype is not preserved. A lectotype (re- 
mounted separately) is deposited at the Institute for Infectious Diseases, 
University of Tokyo. 

Remarks: In the Izu Islands T. scutellaris is the suspected vector of 
Shichito fever, a form of tsutsugamushi disease. 

Subgenus Eutrombicula Ewing, 1938 

Species of EutromhicuJa commonly infest birds and reptiles, and not 
infrequently attack man, causing an irritating dermatitis. The palpal claw 
is two pronged, the axial prong being dorsal (or external) ; there are 20 or 
22 dorsal setae; and with one or more mastitarsalae III. Only one species 
is known from Japan. 

Trombicula (Eutrombicula) wichmanni (Oudemans, 1905) 

(Figure 41) 

Thrombidium wichmanni Oudemans, 1905, Entomologische Berichten, vol. 1, no. 

22, p. 217. 
Trombicula ivichmanni Sasa and Kano, 1950, Tokyo Iji Shinshi. vol. 67, no. 4, p. 9. 

Diagnosis: Palpal femoral seta feathered; palpal genual seta nude; 
dorsal and lateral setae of palpal tibia nude, and ventral tibial seta feath- 
ered. Palpal claw two pronged, the axial prong dorsal (or external). Spur 
on palpal thumb rather short. Galeal seta nude. Sternal setae 2-2. Coxal 
setae 1-1-1. One mastitarsala III. Sensillary bases well in advance of 
posterolateral setae. Sensillae basally nude, with 5-8 barbs on distal half. 



Twenty or twenty-two dorsal setae. Seutal measurements of a specimen 
from Japan : AW-90, PW-108, SB-i5, ASB-36, PSB-40, AP-39, AM-46, 
AL-45, PL-56, S-51. 


Fig. 41 Trombicula wichmanni 


Distribution and hosts: This chigger is widely distributed in the 
warmer regions of the Orient. In Japan it occurs on Koshima and Hachijo 
islands (south of Tokyo) on Rattus norvegicus, wild birds, dogs, cats, and 

Type data : Taken originally from the crowned pigeon from New Guinea 
and from man in North Celebes. 

UngTouped Species of Trombicula 

The following species are placed together for convenience; they are not 
necessarily closely allied. It would create an erroneous impression to place 
them in a subgenus together, although some authors have so treated those 
species of Tromhicula that do not fit into any of the established subgenera 
of TromhicuJa. 

The first two species (T. anous and T. shiraii) would fall into the genus 
(or subgenus) Acariscus Ewing which we do not recognize; these two species 
seem both biologically and morphologically allied. Tromhicula koomori and 
T. hasegawai are unrelated to each other and to other chiggers in Japan. 

Key to Ungrouped Species of Trombicula in Japan 

1. Palpal claw two-pronged; 1 mastitarsala III 2 

— Palpal claw three-pronged; no mastitarsala III 3 

2. Sternal setae 2-2 T. anous 

— Sternal setae 2-4 T. shiraii 

3. Posterior margin of scutum straight ; bases of sensillae nude T. hasegawai 

— Posterior margin of scutum bluntly pointed; bases of sensillae with con- 
spicuous barbs T. koomori 

Trombicula anous (Wharton, 1945) 
(Figure 42) 

Acariscus anous Wharton, 1945, Journal of Parasitology, vol. 31, p. 403. 
Tromhicula anous, Kano and Sasa, 1952, Tokyo Iji Shinshi, vol. 69, no. 10, p. 15. 

Diagnosis: Palpal femoral and genual setae feathered; palpal tibial 
setae nude. Spur on palpal thumb very long and slender. Palpal claw two 
pronged, axial prong internal (or ventral). Galeal seta nude. Sternal 
setae 2-2. Coxal setae 1-1-1. Dorsal setae 32. Sensillae basally nude, with 
5-7 barbs on the distal third. Sensillary bases well in advance of postero- 
lateral setae. One mastitarsala III. Scutal measurements (mean of five 
specimens) as given by the describer: ''AW-93, PW-101, SB-37, ASB-32, 
PSB-19, AP-30, AM-32, AL-53, PL-75, S-73." Scutal measurements of 



a specimen from Miyake Island, south of Tokyo : AW-84, PW-89, SB-28.5, 
ASB-38, AP-32, AL-49, PL-81, S-84. 

Fig. 42 Trombicuhi anous 



[Proc. 4th Ser. 

Distribution and hosts: Originally known from Anous stolidus and 
Heteroscelus iiicatius from Guam. Two specimens from Pluvialis dominicus 
from Miyake Island, Japan. 

Fig. 43 Trombiciila shiraii 



Type data : Type host not indicated ; from Ypao Point, Guam, Mariana 
Islands. Holotype in the United States National Museum. 

Trombicula shiraii Sasa, Kano, and Ogata, 1952 
(Figure 43) 

Trombicula shiraii Sasa, Kano, and Ogata, 1952, Tokyo Iji Shinshi, vol. 69, no. 10, 
p. 14. 

Diagnosis: Femoral seta of palpus feathered; palpal genual and tibial 
setae nude. Palpal claw two pronged, axial prong internal (or ventral). 
Spur on palpal thumb long and slender. Galeal seta nude. Cheliceral base 
and palpal femur and genu with numerous punctae. Scutum more or less 
rectangular, about twice as broad as long. Sensillary bases in advance of 
posterolateral setae. Sensillae with basal half nude, with 5-7 barbs dis- 

Fig. 44 Trombicula hasegawai 


tally. Scutum with numerous punetae. Sternal setae 2^. Coxal setae 
1_1_1. One mastitarsala III. About 90 dorsal setae. Scutal measurements 
of holotype: AW-80.5, PW-89, SB-29.5, ASB-35, AP-32.5, AM-61, 
AL-49.5, PL-82, S-82.5. 

Distribution and hosts: Known only from the original collection. 

Type data: Holotype and one paratype from Charadrius dominicus 
from Tokyo; May 1, 1952. Holotype at Institute for Infectious Diseases, 
University of Tokyo. 

Trombicula hasegawai Sasa et al., 1953 

(Figure 44) 

TromMcula hasegaicai Sasa, Hayashi, and Kawashima, 1953, Tokyo Iji Shinshi, 
vol. 70, no. 4, p. 15. 

Diagnosis: Palpal femoral and genual setae feathered; palpal tibial 
setae nude. Galeal seta nude. Palpal claw three-pronged. Cheliceral base 
and palpal femur and genu with fine punetae. Scutum rectangular, with 
posterior margin straight. Posterolateral setae in the rather angular cor- 
ners of the scutum. Sensillary bases in advance of a line connecting the 
posterolateral setae. Bases of sensillae nude; distal half plumose. Two pairs 
of humeral setae. First two post-humeral rows of setae close together and 
appearing as a single irregular row of setae; fifty or more dorsal setae. 
Sternal setae 2-2. Coxal setae 1-1-1. Scutal measurements of holotype: 
AW-71.5, PW-80, SB-22, ASB-35, PSB-13.5, AP-35.5, AM-52, AL-48.5, 
PL-61, S-56. 

Distribution and hosts : Known only from the type locality from Tur- 
dus celaeonops, Passer montanus, Strejitopelia orientalis, Janthoenas jan- 
thrine, and Monticola soUtarius. 

Type data : Holotype and 10 paratypes from Turdus celaeonops, from 
Hachijo Island (south of Tokyo); September 4-10, 1952. Deposited at 
Institute for Infectious Diseases, University of Tokyo. 

Trombicula koomori Sasa and Jameson, new species 
(Figure 45) 

This species appears to have no close relatives in Japan, and is not very 
much like other known bat chiggers. 

Gnathosoma : Cheliceral base with few punetae. Palpal femoral, genual, 
and lateral and ventral tibial setae all feathered; dorsal tibial seta nude 
and very long (exceeding the tip of the palpal claw). Palpal thumb with 




four feathered setae; the spur rather short, its length about equal to the 
width of the palpal thumb. Palpal claw three-pronged. Galeal seta nude. 
Chelicera twisted apieally, with no projections laterally or subapically. 

Fig. 45 Trombicula koomori 


Scutum: Slightly wider than long; posterior margin produced and 
somewhat pointed; with conspicuous scattered punctae. One of the three 
original specimens possesses two anteromedian setae. Sensillary bases in 
advance of j^osterolateral setae. Sensillae with conspicuous and somewhat 
divergent basal barbs, and with 8-10 long branched on the distal two- 
thirds. Scutal measurements of holotype : AW-62, PW-64, SB-23, ASB-32, 
PSB-23, AM-38, AL-42, PL-60, S-54. 

Legs: Coxae with punctae; other segments with few punctae or none. 
Coxae unisetose. Nude setae on legs: Leg I, 2 genualae, 1 microgenuala, 
2 tibialae, 1 spur, 1 microspur, 1 subterminala, 1 parasubterminala, 1 pre- 
tarsala; Leg II, 1 genuala, 2 tibialae, 1 spur, 1 microspur, 1 pretarsala; 
Leg III, 2 genualae, 1 tibiala. 

Setae : Dorsal setae slender and feathered, similar to the posterolateral 
setae. Dorsal setae in seven rows; dorsal setal formula: 2-14-10-8-8-6-4. 
Sternal setae 2-2. Ventral setae behind sternal setae 50-55. 

Distribution and hosts: In Japan known only from the type collec- 
tion. Taken also in southern Korea from the type host. 

Type data : Holotype and two paratypes from Rhinolophus ferrum- 
equinum (Schreber); near Ohara, Kyoto Prefecture; August 26, 1952; 
collected by Y. Shogaki and J. McClendon. Holotype in the United States 
National Museum; paratypes in Rocky Mountain Laboratory (Hamilton, 
Montana) and Institute for Infectious Diseases, University of Tokyo 

Remarks : The specific name is the Japanese word for bat. 


Brennan, James M. 

1951. Two new species of Neoschongastia with a key to tlie species of the world 

(Acarina: Trombiculidae). Journal of Parasitology, vol. 37, no. 6, pp. 


1952. The genus Pseudoschongastia Lipovsky, 1951, with the description of two 

new species and a key to the world species, also Neoschongastia paeni- 
tens, new name for Neoschongastia kohlsi, Brennan, 1951, preoccupied 
(Acarina, Trombiculidae). Proceedings of the Entomological Society 
of Washington, vol. 54, no. 3, pp. 133-137. 

Brennan, James M. and George W. Wharton 

1950. Studies on North American chiggers. No. 3. The subgenus Neotrom- 
bicula. American Midland Naturalist, vol. 44, no. 1, pp. 153-197. 


Brumpt, E. 

1910. Precis de Parasitologie, 2nd Edition. Masson, Paris, 915 pp. 

EwiNG, H. E. 

1928. A preliminary key to the larvae of fifteen species of the mite genus 
Trombicula, with descriptions of four new species. Proceedings of the 
Entomological Society of Washington, vol. 30, no. 2, pp. 77-80. 

FiKizuMi, Sadakiciii and Yosmo Obata 

1950. A new species of trombiculid mite from the vicinity of Yokohama in 

Japan. Kitasato Archives of Experimental Medicine, vol. 23, no. 3, 
pp. 79-81. 

1951. A new species of trombiculid mite found in Kanagawa Prefecture in 

Japan. Kitasato Archives of Exjierimental Medicine, vol. 23, no. 4, 
pp. 5-9. 
1953. On five new species of trombiculid mites (Acarina) recovered in Miyake- 
jima Island. Kitasato Archives of Experimental Medicine, vol. 26, no. 
1, pp. 1-22. 

Jameson, E. W., Jr. and Manabu Sasa 

1953. Trombicula (Tt'ombiculindus) kansai, a new chigger from central Hon- 
shu, Japan. Journal of Parasitology, vol. 39, no, 3, pp, 247-249. 
Kawashima, Fumio and Manabu Sasa 

1952. Study of tsutsugamushi. No. 19. On new trombiculid mites collected in 

Shikoku, Japan. Tokyo Iji Shinshi, vol. 69, no. 2, p. 21. 

KuwATA, TsiGCo, T. O. Berge, and Cornelius B. Philip 

1950. A new species of Japanese larval mite from a new focus of tsutsugamushi 
disease in southeastern Honshu, Japan. Journal of Parasitology, vol. 
26, no. 1, pp. 80-83. 

MiTSLTTOMi, Shingo, Manabu Sasa, Shigeo Hayashi. Masayoshi Egashira, Kasuo 
KoBAYASHi and Keita Kitahara 
1952. Study of tsutsugamushi. No. 21. On chigger mites in the mountain area 
of Kyushu. Tokyo Iji Shinshi, vol. 69, no. 4, p. 18. 

Nagayo, Mataro, Tokxjsiiiro Mitaml'ra and Takeo Tamiya 

1919. Three species of the tsutsugamushi. Jikkan Igaku Zasshi, vol. 3, no. 4, 

pp. 265-313. 

1920. On the species of tsutsugamushi and their significance to tsutsugamushi 

disease. Verhandelingen Japanische Pathologische Gesellschaft, Bd. 
10, pp. 142-147. 

Nagayo, Mataro, Yoneji Miyagawa, Tokutaro Mitamura, Takeo Tamiya, and 
Satoru Tenjin 

1921. Five species of tsutsugamushi (the carrier of Japanese river fever) and 

their relation to the tsutsugamushi disease. Aitierican Journal of 
Hygiene, vol. 1, nos. 5 and 6, pp. 569-591. 

Oudemans, a. C. 

1905. Acarologische Aanteekeningen, 16. Entomologische Berichten, Bd. 1, 
pp. 216-218.^ <2i . 


Philip, Cornelus B. 

1947. Observations on tsutsugamushi disease (mite borne or scrub typhus) 
in northwestern Honshu Island, Japan, in the fall of 1945. I. Epide- 
miological and ecogical data. Ameriraii Journal of Hygiene, vol. 46, 
no. 1, pp. 45-59. 

Philip, Cornelius B. and H. S. Fi'ller 

1950. The harvest mites ('Akidani') of Japan and the Far East and their re- 

lationship to the autumnalis group of trombiculid mites. Parasitology, 
vol. 40, nos. 1 and 2, pp. 50-57. 

Sasa, Manabu, Siiigeo Hayaisiii and Fumio Kawashima 

1953. Study of tsutsugamushi. No. 32. New tsutsugamushi from birds from 
the vicinity of Hachijo. Tokyo Iji Shinshi, vol. 70, no. 4, pp. 15-17. 

Sasa, Manabu, Siiigeo Hayashi, Fumio Kawashima, Shingo Mitsitomi, and 
Masayoshi Egashira 

1952. Study of tsutsugamushi. No. 20. Two new species collected from moles 
in Kyushu and Skikoku. Tokyo Iji Shinshi, vol. 69, no. 3, pp. 11-13. 

Sasa, Manabu, Shigeo Hayashi, Nobuo Ki mada, and Akiko Miura 

1951. Study of tsutsugamushi. No. 6. On Trombicula tenjin, n. sp. in Japan. 

Tokyo Iji Shinshi, vol. 68, no. 3, pp. 17-18. 

Sasa, Manabu, Shigeo Hayashi, Koji Sato, Akiko Miura, and Siiozo Asaiiina 
1950. Study of tsutsugamushi. No. 5. Chigger mites collected in the Mt. Fuji 
Area. Tokyo Iji Shijishi, vol. 67, no. 12, p. 14. 

Sasa, Manabu, Siiigeo Hayashi, Nobuo Kumada, and Seiyu Teramura 

1950. Study of tsutsugamushi. No. 4. Chigger mites collected in the Mt. Fuji 
Area. Tokyo Iji Shinshi. vol. 67, no. 11, pp. 17-18. 

Sasa, Manabu, and Rokuro Kano 

1950. Study of tsutsugamushi. No. 1. Trombieula unchmanni discovered in 

Japan. Tokyo Iji Shiyishi. vol. 67, no. 4, pp. 9-10. 

Sasa, Manabu, Rokuro Kano, Nobuo Kumada, and Meiiciii Ueda 

1951. Study of tsutsugamushi. No. 14. The chigger mites in Hokkaido. Tokyo 

Iji Shinshi, vol. 68, no. 9, p. 9. 

Sasa, Manabu, Rokuro Kano, and Takayuki Obata 

1952. Study of tsutsugamushi. No. 28. On trombiculid mites collected from 

birds in the Tokyo Area. Tokyo Iji Shinshi, vol. 69, no. 10, pp. 13-15. 

Sasa, Manabu and Fumio Kawashima 

1951. Study of tsutsugamushi. No. 15. A new species found in Kochi Prefec- 
ture. Tokyo Iji Shinshi, vol. 68, no. 10, pp. 9-10. 

1951a. Study of tsutsugamushi. No. 17. Two new species found in Kochi Pre- 
fecture. Tokyo Iji Shinshi, vol. 68, no. 12, pp. 15-17. 


Sasa, Manabu, Fumio Kawashijia, and Masayoshi Egashira 

1952. Study of tsutsugamushi. No. 25. On Trombicula (MiyatromMcula) 
kochiensis, new subgenus, new species, from western Japan. Tokyo 
Iji Shinshi, vol. 69, no. 6, pp. 5-6. 

Sasa. Manabc, Fimio Kawashima, and Seiichiro Hikomatsu 

1952. Study of tsutsugamushi. No. 18. A new species collected from a mole in 
Kochi Prefecture. Tokyo Iji Shinshi. vol. 69, no. 1, p. 43. 

Sasa, Manabu. Nobuo Kumada, and Akiko Mivua 

1951. Study of tsutsugamushi. No. 12. On the genus Euschongastia. Tokyo 
Iji Shinshi, vol. 68, no. 8. p. 19. 

Sasa, Manabu, Nobuo Kumada, and Seiyu Teramura 

1951. Study of tsutsugamushi. No. 13. New species collected in Niigata and 
Akita Prefectures. Tokyo Iji Shinshi, vol. 68, no. 9, pp. 8-9. 

Sasa, Manabu, Nobuo Kumada, Shigeo Hayasiii, Yoshibumi Enomoto, Sadakichi 
FuKuzi Ki, and Yoshio Obata 
1951. A new larval mite, Trombicula himizu, n. sp. from Japan. Eisei Doobutsu. 
vol. 2, no. 1, pp. 1-5. 

Sasa. Manabu, Toshisada Sawada, Rokuro Kano, Shigeo Hayashi, and Nobuo 
1951. Study of tsutsugamushi. No. 7. Two new species collected in Gumma 
Prefecture. Tokyo Iji Shinshi. vol. 68, no. 4, pp. 7-8. 

Sasa. Manabu and Seiyu Teramira 

1951. Study of tsutsugamushi. No. 8. A new species of Walchia, previously 

unrecorded from Japan. Tokyo Iji Shinshi, vol. 68, no. 5, p. 10. 

Sasa. Manabu, Seiyu Teramira, and Rokiro Kano 

1950. Study of tsutsugamushi. No. 2. Chigger mites from Hachijo Island. 
Tokyo Iji Shimihi, vol. 67, no. 10, pp. 22-23. 


1948. New red mites (Acari. Trombidiidae) of the fauna of the U.S.S.R. 
Entomologicheskoe Obozrenie. i. 30, nos. 1 and 2, pp. 157-164. 

Tanaka, Keisike, J. Kaiwa, S. Teramura. and I. Kagay'a 

1930. Beitrage zur japanischen Kedani-Krankheit. Zentralblatt fiir Bakterio- 
logie, Bd. 116, pp. 353-385. 

Wharton, G. W. 

1945. Two new species of Acariscus: A. pluvius and A. anous (Acarinida: 

Trombiculidae). Journal of Parasitology, vol. 31, no. 6, pp. 401-405. 

Wharton. G. W. and A. B. Hardcastle 

1946. The genus Neoschongastia (Acarinida: Trombiculidae) in the western 

Pacific area. Journal of Parasitology, vol. 32, no. 3, pp. 286-322. 


1952. The scrub-typhus and scrub-itch mites {Trombiculidae. Acarina) of the 

Asiatic-Pacific Region. Records of the South Australia Museum, vol. 
10, pt. 1 (text), 435 pp., pt. 2 (plates), pp. 438-673. 




Fourth Series 
Vol. XXVIII, No. 6, pp. 323-337, figs. 1-5 July 9, 1954 






California Academy of Sciences 

Tlie first stage of experimental studies on galvanotropie responses 
of the Pacific sardine {Sarflinops caerulea), using pulsating direct cur- 
rent of low frequencies and a triangular wave form, was described by 
(4roody, Loukashkin, and (Irant (1952). That account was concluded 
(pages 317-322) by a study of the effects of the three variables in- 
volved, namely, maximum current density, ratio of current-on to current- 
oft' periods, and frequency of pulsation. During this experiment, 900 
tests were made. 


The experimental tank and electrical design used in the present 
study are shown in figure 1 (after Groody, et ah, 1952, p. 312). 

Tlie following definitions, adopted in the earlier report {op. cit.), 
were used. 

Perfect. AVhen all four fish responded readily to each reversal of 
the poles and displayed a directional reaction toward the positive pole. 

(iood. AVhen three of the four fish displayed the reaction given 
above or when all four fish failed to respond to one of the three re- 
versals of poles. 

Fair. AVhen two of the fish readily responded to all pole reversals, 
or when all fish responded to at least one reversal. 

[ 323] 



[Proc. 4x11 Si:r. 

Middle lOf tank 
Aerator | 

Protective plastic screen barriers 



90 V. D.C. 

controlled by electronic timer 

Figure 1. Diagram of experimental wooden tank and electrical circuit for 
triangular wave form of pulsating direct current. Reversing switch not shown 
(after Groody, Loukashkin, and Grant, 1952). 

None. When one or none of the fish displayed directional reactions. 
The first two grades were classified as satisfactory and the last two 
grades as unsatisfactory (table 1). 

In addition to the above definitions for recording behavior, records 
were made of the fishes' response to the current relative to fright reac- 
tions to stimuli and to apparent control of their own swimming move- 
ments. For these records, the following symbols were used : 

(— ). Swimming movements apparently under control of the fish, 
no directional response, and all reactions to fright stimuli retained. 

(1/^+). The avoidance reaction to the barrier was retained, but the 
fish did not respond to vibrations of the tank with fright reactions. 

( + ). Directional responses apparently controlled by electrical cur- 
rent density only. Normal fright reactions to the barrier or to other 
stimuli completely lost. 

(++). In addition to the reactions above under ( + ), the fish re- 
mained in contact Math the protective screen next the positive pole as 


s e 

g "■ 



•^ + 

o o o o o 

O O O O o 

o o o o o 

OJ l> CO (N 00 

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N •* »-l O «£> 

oi -ef th o ui 

oo CO t^ oo e<i 

LO 00 CO rH CO 

CO in ^H t- Oi 

O C: O O O 
O O O O O 
O O O O O 





















S *.i 

to to 






"5 s 

ec to 






1 b 

















S a 

o «C 




















O i-H 00 OO 00 

Tf -* eg (M tH 

in o ■*! o — I 

o in o in o 
T-i i-H eg (M eo 

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s c 

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o o o o o 
ii' d <::> (d: d 
o o o o o 

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iH CO IC CO t- 

00 rf i>- CO eg 

o o i-H o eg 
d d T-J d eg 
tH eg CO CO eg 

t- CO eg t- CO 

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o o o o o 
o <:6 d d d 
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lij iij 

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Figure 2. Triangular and square wave forms of direct current pulsating at 
low frequencies with a pulse of 30 milliamperes of maximum current density 
and an "on" to "off" ratio of 2:1. The frequency of pulsation is 5 per second. 
Timing base is 60 cycles per second. 

long as the current was on. This was, of course, the most pronounced 
reaction to the current short of stunning and death. 

( — ) and (1/^+) were classified as unsatisfactory, ( + ) and (+ +) as 
satisfactonj (tahle 2). 


The present studies centered on the stimulating effects on sardine 
hehavior of: (1) the .square wave form of direct current pulsating at 
low frequencies; (2) application of higher frequencies of pulsating direct 
current using triangular and square wave forms; (3) application of half- 
wave rectified 60-cycle alternating current; (4) application of quarter- 
wave rectified 60-cycle alternating current; and (5) application of the 
condenser discharge impulse. 

(1) Effects of the square wave form of direct current pulsating at 
low frecjuencies. Using the same facilities and equipment and following 
the same procedure and recording as described in the preliminary re- 
port cited above, a series of 900 analogous tests were made in order to 
investigate the effects of the square wave form of direct current pul- 
sating at low frequencies. 

The s(iuare wave impulse was obtained by omitting an inductance 
from the circuit diagrammed in figure 1. Contrary to the triangular 


Triangular wave form 
100 , Square wave form 

90 . 

80 - 




c 50 


30 U 







Maximum currenf densilies in milliamperes 
per square inch of cross-seciional area of wafer. 

Figure 3. "Satisfactory" directional reactions (in per cent) of the sardines 
in an electrical field produced by using triangular and square wave forms of 
direct current pulsating at low frequencies. Percentages are based on 180 ob- 
servations for each of the five maximum current densities of each wave form tested. 

wave form, the square wave impulse rises instantly to its maximum, 
maintains this density for the desired period and then, like the tri- 
angular wave, it returns abruptly to zero (figure 2). The results of 
these tests are shown in tables 1 and 2. 

The tests indicate that current density is the most critical factor. 
When the maximum current density increases from 10 to 30 milliamperes 
per square inch of cross-sectional area of water, there is a well-displayed 


trend toward increase of "satisfactory" results. As is shown in figures 
2 and 3, this is true of both wave forms but is more marked in the 
triangular wave. 

A comparative anal.vsis of the results obtained indicates that the 
triangular wave form produced comparatively better directional responses 
than did the square wave, while "satisfactory" control of fish move- 
ments was obtained more frequently with the square wave. However, 
the difference does not appear to be significant. It can be concluded 
that both wave forms of direct current pulsating at low frequencies* are 
"satisfactory" in producing a directional response and in control of the 
sardine movements. 

(2) The appUcation of higher frequencies of pulsating direct cur- 
rent using triangular and square wave forms. In order to obtain pul- 
sating direct current of the higher frequencies, a high-speed mechanical 
interrupter was introduced into the circuit diagrammed in figure 1. Using 
both types of current wave forms and applying a 1:1 ratio of current- 
on to current-off periods (figure 5: A and B), the following frequencies 
w^ere tested: 5, 20, 35, 50, 65, and 80 per second. 

The experiments disclosed that the average current density of pul- 
sating direct current is a more important factor in producing directional 
swimming and control of fish movement than the maximum or peak 
current density. Also, it was found more convenient to express current 
density as an average value since this can be read directly from the 
ammeter dial. This average value is dependent on the ratio of the "on" 
to "off" periods.** For example, using a square wave form and an 
"on" to "off" ratio of 1 :1, the average current density would be one 
half of the maximum or peak current value attained during the "on" 
period. Applying the same value of the maximum current density and 
changing the "on" to "off" ratio from 1:1 to 2:1 and 1:2, the average 
current density will be higher when a 2:1 ratio is used, lower when a 
1 :2 ratio is used. This statement differs from, but is not inconsistent 

*The table of sardine reactions to variations in current pulsation frequency is omitted because the rate of 
current pulsation (within the range of 2, 4, and 6 times per second) seemed to have no significant effect 
on directional swimming and control of fish movements. This observation is in full accord with the 
records for the triangular wave form given in the preliminary report (Groody, et al., 19S2, table 2). 

*When straight direct current (i.e., not pulsating) is used, the values of maximum and average current 
densities are equal. 

Figure 4. Numbers of classified directional reactions of the sardines per 
current density tested using triangular and square wave forms of direct current 
pulsating at low frequencies. Bach column for each maximum current density 
represents 180 observations. 


Triangular wave 












Square vjcwie. 


20 25 30 

20 25 30 

Maximum current densities in Maximum current densities in 

mi II iamperes per square inch milliamperes per square inch 


<^ Good 







with, that in our earlier paper (Groody, et al., 1952, p. 318, and table 
3). It was shown there that control of movement was inereasinjily suc- 
cessful with increasinti' ratio of current-on to current-off periods. It will 
be ol)vious that this result is satisfactorily explained as an increase 
in average current density. 

Using a triangular or any other wave form and applying the same 
values of tlie maximum or peak current density and the same ratios of 
the "on" to "off" periods, the average current density would be always 
less than that of the square wave form. 

Use of the higher frequencies in this experiment produced "perfect" 
directional swimming and full (+ +) control of fish movements at con- 
siderably reduced average current densities. The current density re- 
quired to produce highly "satisfactory" results at a frequency of 65 to 
80 pulses per second was only 50 per cent of that needed at a frequency 
of five pulses per second. 

Optimal ranges of the average current density estal)lished by the 
above experiments for the Pacific sardine 200-300 mm. in standard length 
are shown in table 3. 


The Effect of Frequency of Pulsation Upon Optimal Average Current Density 

(240 Observations) 

Values of frequencies of pulsating Values of the optimal average current 

direct current, using square or tri- densities which produce "perfect" di- 

angular wave forms and 1 : 1 ratio of lectional swimming and full (++) 

current-on to current-off periods. control of fish movements. 

5 times per second From 8 to 12 milliamperes/sq. inch 






(3) The appUcation of half-wave rectified GO-cycle alternating current. 
As frequencies of pulsating direct current as high as 65 to 80 per second 
were found to be more effective than the low frequencies, in regard to 
reducing the amount of average current density required to produce di- 
rectional swimming and control of fish movements, it was then decided 
to test the sardines in an electrical field of half-wave rectified 60-cycle 
alternating current, using a standard mercury vapor rectifier tube. This 
produces a rounded and evenly sloped current wave form with a 1:1 ra- 
tio of current-on to current-off' (figure 5C). 

•om 8 



















o _^ 

•2 -a 

> O 1^ 

^ S S-, 

o o o o o 

IP 5 ^ i^ 

i. =t-i CS O) 

0. ^ V °^ 

iS _v H <i> 

■— "O '^ o, 

? O) . 

^ S W tc 

^ ^ '^ 

.2 "^ • - ?3 

o; ^ 8 ^ 


>^ ^ r -^ o 



























































































































• rH 








^ — 

















































Using the facilities and e(iiiipnient shown in figure 1 with the ex- 
ception that the inductance and contactor were left out of the circuit, 
a series of more than 300 tests were made, using groups of four sar- 
dines 200-230 mm. in standard length, the procedure and recording 
being the same as in previous experiments. 

From these experiments it was found that the optimal average cur- 
rent density required to produce "perfect" directional swimming and 
fully controlled fish movements varied within the range of 3.3 to 5.0 
milliamperes per square inch of cross-sectional area of water. The higher 
average current densities from 6 to 9 milliamperes showed a tendency 
to stun the fish and it appears that this stunning effect is directly re- 
lated to the increase in average current density. 

Following the experiments with the sardines, topsmelt {Atherinops 
(tffinis), 100-110 mm. in standard length, were subjected in groups of 
four to the stimulation. They were found to respond in a similar 
manner to the sardines, the only difference being that the optimal aver- 
age current density requirement was found to be almost twice as high 
(table 4). This is in accord with a statement made in the earlier study 
(Oroody, et al., 1952, p. 317) concerning the api)arent inverse relationship 
between the amount of current density needed to produce directional swim- 
ming and the size of the fish. It should be pointed out, however, that 
these observations were made on different species of fish. Unfortunately, 
the species of fish used were available in only one size. Interestingly, 
McMillan (1929, \). 102), who studied the effect of voltage gradient re- 
quired to produce paralysis on rainboW' trout (Sohno irideus) and Chi- 
nook salmon {0)icorhynchus tschawytscha) that varied in length from 
1.87 to 12.50 and from 3.10 to 31.75 inches respectively, states ". . . the 
voltage gradient required to produce paralysis is inversely proportional 
to the length of the fish. In other words, the long fish require a much 
lower field strength to paralyze them than the short ones." 

Further, to check this relationship between length of fish and current 
density required, jacksmelt {Atherinopsis californiensis) 300-330 mm. 
in standard length, were similarly tested in groups of four. This fish, 
too, reacted as did the sardines but the optimal average current density 
requirement was as low as 2.5 to 3 milliamperes per square inch of 
cross-sectional area of water. Higher current density caused the fish 
to roll over and remain stunned until the current was turned off. 

In each of the three species tested with average current densities 
above the optimal range, fish have been stunned to various degrees 
but all recovered as soon as the current was turned off. When returned 
to their holding tanks, the fish rejoined schools and apparently suffered 
no ill effects from the experiments. 


In the case of all three species of fish, the half-wave rectified 60-cycle 
alternating current caused a higher speed of swimming than the tri- 
angular or the square forms. Other observations indicated that, under 
the influence of the half-wave rectified A.C., the fish reacted with con- 
siderably more ease and showed less distress during forced swinuning 
toward the positive pole. This is not easy to establish on a quantitative 
basis, but was a fact of observation that was confirmed by several 

Typical behavior of the fish in a field of half-wave rectified 60-cycle 
A.C. is summarized in table 4. 

Table 4 

Reactions and Optivtal Average Current Densities Using Half-Wave Rectified 

60-cycle Alternating Current 

Species of Fish Stimulated 

Standard Optimal Range of the Average 

Lengtli of Current Density Required to 

the Fish Produce "Satisfactory" reac- 

Tested in tions as Described in the 

Millimeters Lower Part of This Table 

Pacific Sardine 

Sardincjps caerulea 


Atherinops affinis 


A therinopsis ca lifornien s is 


3.3 to 5 milliamperes 
7.0 to 9.0 
2.5 to 3.0 

Description of fish behavior in an electrical field within the optimal range of 
the average current densities: 

Orientation instantaneous and simultaneous. Directional swimming "perfect"; 
response to reversals of polarity immediate and coordinated. Control of move- 
ment full (+-I-): fish cannot avoid barrier inserted between swimming fish 
and positive pole: all natural fright reactions to disturbing stimuli entirely lost; 
upon I'eaching protective screen (two feet in front of the positive electrode) 
fish cannot swim away, being strongly attracted toward the electrode. Swim- 
ming very rapid, sometimes performed as a rapid gliding over the water sur- 
face toward positive pole. Easy on fish. 

This half-wave rectified 60-cycle alternating current was also inter- 
rupted 30 times a second by using a thyratron tube, and the wave form so 
obtained is shown in figure 5D. 

The experiments carried out on the same three species of fish re- 
sulted in obtaining "perfect" directional swimming and fully controlled 
movement (+ +) of the fish as described above, however, the amount 
of the average current density required to produce these reactions was 


about one-half of that established I'or the pure (uninterrupted) half- 
wave rectified 60-cyele alternating current. 

(4) T'he applkation of the quarter-wave rectified 60-cycIe alternat- 
ing current. For experiments using a wave form that starts wdth the 
maxiinuiu cun-eut and slopes gradually to zero as shown in figure 5E, 
the autliors used a mercury vapor thyratron tube and phase-changing 
ecpnpment. It is actually a quarter-wave impulse and the reverse of 
the originally tested triangular wave form of pulsating direct current. 

Use of this quarter-wave form and the same optimal range of the 
average current densities and species of fish as in the experiments with 
lialf-wave rectified 60-cycle A.C. (table 4), resulted in fish reactions that 
were less satisfactoi-y. In this experiment, over 200 tests were made. 

Although the fish did orient and swim toward the positive pole, their 
direction of swinmiing being fully under electrical control, it was obvious 
that they exi)erienced certain difficulties and attempted to escape from 
the field by raising their lieads out of water and trying to jumi) over 
the sides of the tank. There were several cases of strong stunning effects, 
even when the lower values of the optimal range of the average current 
density were applied. It also required a longer period for the fish to 
relax and resume normal behavior after these tests than it did after 
any of the previous experiments. These effects were observed in the 
responses of each of the three species used. 

It is assumed that in order to obtain an oi>timal range of the aver- 
age densities established for the half-wave current, the magnitude of the 
w^ave peak must be much greater in the quarter-wave impulse than in 
the former. In all prol)al)ility, this is the cause of a strong and lasting 
stunning effect observed on the fish tested. 

(5) The application of the condenser discharge impulse. The present 
investigations of the effects of various wave forms and frequencies of 
pulsating direct current w^ere completed by experiments with condenser 
discharge jiulses having a frequency of approximately 14 per second. 
This was produced by charging a 50 MF condenser from a source of 
half-wave rectified 120-volt alternating current and discharging through 
a mercury vapor thyratron tube. The wave form thus obtained is dia- 
grammed in figure 5F. 

A series of more than 300 tests showed that this type of current im- 
pulse was also effective in producing directional swimming of the sardine. 
The optimal range of the average current densities was found to be 
as low as 0.4 to 0.8 milliampere per square inch of cross-sectional area 
of water.* It is evident from these tests that the amount of electrical 

*The negative results of the effects of the condenser discharge mentioned in the preliminary report (Groody. 
et al., 19S2) were probably due to the discharge of condenser through mechanical contactor instead of 
the thyratron tube. 


energy required to produce directional swimming of the sardine is much 
less using condenser discharge than by employing other wave forms tested. 

When optimal average current densities were used the sardines' re- 
actions sliowed that while forced directional swimming was clearly dis- 
played, the fish experienced considerable difficulty in reorienting when 
the polarity of the electrical field was reversed. They usually continued 
swimming in the original direction for a short distance. It was observed 
that while they were attempting to turn, their bodies vibrated at what 
appeared to be the pulse frequency. It also appeared that this type of 
current impulse caused the fish to become exhausted in a short time: 
they were easily picked up by hand for transferring in a bucket back 
to their holding tank. 

Similar effects were observed with jacksmelt. Their optimal range was 
found to be ])etween 0.4 to 0.6 milliampere per square inch. No topsmelt 
were available at the time of the experiments. 

Although controlled directional swimming was clearly displayed, the 
full control of fish movements (++) was not immediately obtained: 
both the sardine and jacksmelt were able to leave the positive pole and 
swim away toward the negative pole; however, after several such escap- 
ing attempts the fish finally returned to the positive pole. 

Application of the higher values of the average current density above 
optimal range in order to produce an immediate and full control of the 
fish movements resulted in a strong stunning effect on the fish as soon 
as the current was turned on. One of the sardines used in these tests 
became totally blind, and it died 12 hours later. 

The saving in current obtained by the use of condenser discharge 
is i)articularly significant in view of the fact that one of the most im- 
portant problems in the development of electrofishing is the large amount 
of power needed to establish suitable current densities in sea water. 


Observations made during the course of experimental studies of the 
behavior of the Pacific sardine carried on in the California Acad- 
emy of Sciences since 1950, have established that this species is highly 
susceptible to stimulation by electrical currents. The fish usually begin 
to feel the presence of an electrical field even when the lowest values of 
average current density are ajiplied. 

The sensitivity of the sardine to changes in average current density 
and the degree of stimulating effect that has been once established for 
each density value remain con.stant. This is true for each particular 
type of pulsating current tested. As soon as the optimal range of aver- 
age current density required to produce directional swinuuing and con- 


trol of fish movoincnt has been found, the sardine's reactions will always 
remain identieal regardless of the number of times the tests are repeated 
or the sequence of the tests. 

The observations present another fact, namely, the sardines just cap- 
tured or the ones that have been kept in captivity for an extended 
period of time display identical sensitivity as described above. No con- 
ditioninii' has ever been observed despite the fact that the experimental 
fish were confined in the Steinhart Aquarium's holding tanks for more 
than two and one-half years and were used twice a week on the average 
in electrical stimulation experiments. In contrast to the topsmelt and 
jacksmelt, their peculiar reactions to each particular value of the average 
current density applied, have been invariably the same. Topsmelt and jack- 
smelt usually became conditioned after 5-10 successive stimulations, and 
therefore in the experiments here reported these fishes have often been 
replaced by the fresh specimens not previously subjected to the electri- 
cal field. 

The results of the effects of various wave forms and currents and 
of diiTerent frequencies recorded during the course of experimental studies 
have been checked many times. These repeated tests have proved that 
the optimal ranges of average current density experimentally deter- 
mined for the three species of fish for each different type of electrical 
current, have always remained essentially the same. 

The observations also indicate that, when a proper value of average 
current density is applied and the fish movements are fully (+ +) con- 
trolled, their natural fright reactions to disturbing stimuli are entirely 
suppressed and the fish can be easily picked up by hand. This observation 
applies to all three species used in the present investigations. 


1. The five types of pulsating direct current wave forms used by 
the experimenters are effective in producing forced directional swim- 
ming of the Pacific sardine {Sardinops caerulea) and also the jacksmelt 
{Atherinopsis calif orniensis), and the topsmelt {Atherinops affinis) that 
were used as controls. 

2. All wave forms tested can produce full (+ +) control of fish 
movements and force them to the positive pole where they can be held 
until the current is turned off. 

3. The electrically stimulated reactions can be obtained and will re- 
main constant if a proper optimal range of average current density is 
determined for each particular species and size. 

4. Current density is the most critical factor in producing forced 
directional swimming and control of fish movements. 


5. Average current densities above the optimal i-ange may cause 
temporary paralysis or even death of the fish whereas those below pro- 
duce a slight directional response or none at all. 

6. The optimal average current density required to i)roduce "satis- 
factory" directional swimming and controlled (+ +) movement of fish 
appears to vary inversely with the size of the fish. 

7. Full control of fish movement and forced directional swimming 
can be obtained with any frequency of current pulsation from 2 to 80 
per second; the frequencies above 80, however, have not been tested. 

8. The use of a pulse frequency as high as 60 to 80 per second re- 
duced the optimal average current density to 50 per cent of the amount 
required at a frequency of 5 per second. 

9. The most effective and "satisfactory" results, as far as the smooth- 
ness of performance and school coordination are concerned, were re- 
corded when either continuous or interrupted half-wave rectified 60-cycle 
alternating current was used. 

10. Condenser discharge pulse produced the "satisfactory" reactions 
at very low average current densities (0.4 to 0.8 milliampere). This rep- 
resents a substantial decrease in power requirements. 


This work has been carried on with funds provided by the Marine 
Research Committee as a part of the California Cooperative Oceanic 
Fisheries Investigations. The authors express their appreciation to Dr. 
Robert C. Miller, director of the California Academy of Sciences, for 
his keen interest in the studies and for encouragement and valuable sug- 
gestions during the progress of the research; and to Dr. Earl S. Herald, 
curator of aquatic biology, California Academy of Sciences, for his will- 
ing cooperation and valuable counsel. Acknowledgment is made also to 
Drs. Roger Revelle and Carl L. Hubbs, of the Scripps Institution of 
Oceanography; Dr. Francis N. Clark, director of the California State 
Fisheries Laboratory, and Mr. John C. Marr and Dr. E. H. Ahlstrom, 
of the U. S. South Pacific Fisheries Investigations, for helpful sugges- 
tions and constructive criticism of the manuscript. 

Groody. Tom, Anatole Loukashkin, and Norman Grant 

1952. A preliminary report on the behavior of the Pacific Sardine (Sardinops 
caerulea) in an electrical field. Proceedings of the California Academy 
of Sciences, vol. XXVII, no. 8, pp. 311-323, 4 text figures. 

McMillan, F. 0. 

1929. Electric fish screen. Bulletin of the United States Bureau of Fisheries, 
vol. XLIV (1928), pp. 97-128, 21 text figures. 




Fourth Series 
Vol. XXVIII, No. 7, pp. 339-353, figs. 1-3 June 30, 1955 





California Academy of Sciences 


Tlie behavior of marine fishes under laboratory conditions, particularly 
that of members of the Clupeoidea, has been very little studied. Parr (1927) 
offers this explanation: "It is most unfortunate that the species showing 
the schooling performances most clearly, as for instance herrings, spratts, 
and mackerels, usually are of such delicate nature that it is practically im- 
possible to keep them alive for any length of time." Further reference to 
the subject is made hy Spooner (1931) who states: "Choice of schooling- 
fish suitable for observation in captivity is limited to a very few species. 
Small mackerel, herring, and spratt are excluded on account of difficulties 
involved in keeping them in confinement." 

A number of behavior studies under laboratory and field conditions 
have been made, however, on various si)ecies of herrings (Newman, 1876; 
Shelford and Power.s, 1915; Powers, 1921; Breder, 1929), and on several 
unrelated marine fishes, including the great blue shark Prionace glauca 
(Ilubbs, 1948), and the tarpon Tarpon atlanticus (Shlaifer and Breder, 
1940; Shlaifer, 1941). 

Papers on the Pacific sardine Sardinops caeruJea (Girard) have been 
confined largely to scattered field observations by Cornish (1883) and 
Allen (1920, 1930). 

"Submitted February 10, 1953. 

[ 339 ] 

340 CALlFOh'MA ACAltKMY OF SCIENCES [Piux . 4tii Sek. 

Fishes of tlic suborder (Mn])eoidea are aiiatoiuically, ])liysio]og'ically, and 
temperainenlally anionii' the most delicate of marine fishes. This is espe- 
cially true oi' the Facitic sai'dine, whose nervous temperament, so familiar 
to men en<iai>ed in the sai'dine fishery, renders it a difficult subject for 
laboratory study. The ease with whicli the sardine is injured through han- 
dling, coupled with its sensitivity to environmental changes in salinity, tem- 
perature, and oxygen may well account for the absence of laboratory studies 
on this economically important fish. 

Since these fishes rank among the most important from the standpoint 
of the worhVs economy and resources, it was deemed necessary to establish 
the conditions under which they should be transjiorted to a laboratory and 
thei-e maintained as objects for detailed ex])erimental investigations. 

Methods of Handling 


Altogether, seven shipments of live adult sardines, totalling 751 speci- 
mens, were delivered to the Steinhart Aquarium during 1949-51. The first 
three lots, purchased from bait-fishermen, came from Monterey Bay. 

The first lot of 57 sardines was delivered IMay 19, 1949. All the fish died 
off rapidly. The second lot of 37 sardines was delivered August 4, 1949. By 
the next day 27 fish (73 per cent) had died; the rest of the fish survived a 
few days. The third shipment of 24 sardines was made on December 22, 
1949. Heavy casualties M'ere incurred in the first 48 hours after delivery; 
however, five fish survived until the end of May, 1950. 

This excessive mortality among the sardines from Monterey Bay was 
considered to be due to the long travel time (about three hours) in the col- 
lecting truck; so all subsequent collections were brought to San Francisco 
from the Los Angeles area al)oard the ^I.S. YeUotcfin, research vessel of the 
California Department of P^ish and (Jame. 

The sardines from the Los Angeles area were also purchased from bait- 
fishermen and were kept in the wooden bait-tank of the YeJlowfin, which 
tank contained 1,125 gallons of circulating sea-water. The vessel's voyage 
between Los Angeles and San Francisco was usually nuide in 50-52 hours, 
and sometimes the ship exjierienced rough weather. Delivery to the Acad- 
emy's Steinliart A(iuarium from aboard ship, a distance of eight miles, was 
made in a collecting truck e(iui])])ed with 45- and 100-gallon cans (fig. 1) 
that have air compressors and inside lights. Two or three truck trips, each 
of which took from 15 to 40 miiuites, were required to unload each ship- 
ment. Dip-nets and two-gallon buckets were used for transferring the 
sardines, both at the dock and the Aquarium. 

The foui'tli shi])ment of 128 sardines, from Long Beach, was delivered 
on -lune 9, 1!)50. In f)6 hours after delivery, 85 sardines (66.5 per cent) 


Figure 1. Drawing of a 100-gallon trinispoi'ting caii. 

had died; ))y tlio end of the thii'd week only 13 fish survived. The fiffh ship- 
ment of 114 sardines, from Los Ansieles harl)or, was delivered on October 
4, 1950. In 96 lionrs of confinement in the Aqnarinm's liolding tanks, 54 
(47.4 per cent) sardines died off; l)y the end of the third week the cumula- 
tive mortality was 84.2 per cent. On tlie sixty-second day of captivity only 
16 fish survived. The sixth shii)ment of 116 sardines,' from Los Angeles 
harbor, was received on December 3, 1950. In a 96-hour period 42.2 per 
cent of the fish died off; at the end of the third week the cumulative mor- 
tality was 67.2 per cent. Toward the end of October, 1951 (13 months after 
delivery), 32 sardines were still living. The seventh shipment of 275 sar- 
dines, from Los Angeles harbor, was delivered on October 22, 1951. In a 
96-hour ])eriod only 38.9 per cent died; at the end of the third week the 
cumulative mortality was only 42.9 i^er cent. A count made on January 8, 
1953 (14 months and 17 days after delivery), revealed that 43 sardines 
successfully survived, together with 10 fishes of the sixth shipment. 

Maintenance : 

At the Steinhart Aquarium, the experimental fish were confined in 
holding tanks having a capacity of about 1,000 gallons (table I) which, as 


(mr experience indieates, were adequate for about 50 adult fish. The 
Aquarium salt water is pumped from the ocean to a storage tank and the 
rate of flow through the 1,000-gallon holding tanks is 86 gallons per hour. 
Each tank is aerated but in most cases temperature and pH are not con- 
trolled. In most tanks, the temperature range was 11.0-20. 5'^C. and that 
of the pU, 7.0-8.0. 

Degree of Survival 

In transit: 

Data on mortality during the sea voyage to San Francisco from Los 
Angeles were not recorded in the ship's log of the Yellowfin. However, 
according to John Radovich (in lit.), "sardines have been kept alive for an 
entire cruise on several occasions with a very small loss in the vessel's bait- 

From our observations it is readily apparent that sardines of the firet 
load removed from the vessel's bait-tank and transferred to holding tanks 
of the Aquarium, as a rule, incur heaviest losses during the first three-week 
period in comparison with those of the other loads of the same shipment 
that were transferred later. This is especially true of the seventh shipment 
of October 22, 1951, when 275 sardines were transported in three consecu- 
tive truck trips, and distributed in three separate holding tanks at the 
Aquarium. A very accurate record of mortality was kept, and a summary 
is presented in table II. 

Duta on Holding Tanks in Steinhart Aquarium 

Capacity 1057 gallons 

Inside dimensions 74" x 75" x 44" 

Operating water volume 1,000 gallons 

Rate of water turnover 86 gal./hr. 

Air introduced by suction through opening 

in water inlet tube variable rate 

Annual temperature range for tank with non- 
regulated water temperature 11.0°C. to 20.5°C. 

Greatest temperature fluctuation in 24 hours 1°C. 

Observation indicates that fisli of tlie first load are those whicli have 
suffered most during the voyage or from previous handling by the bait- 
fishermen. They usually swim about close to the surface in the shii)'s ])ait- 
lank and are the first to be netted out for transi)ortation, while healthy fish 
dive to the bottom of the tank and stav there until netted. 



Mortality among sardines of various loads transported in a truck from the 
Yellowfin bait-tank to the 8teinhart Aquarium holding tanks on October 2i, 19ol. 
Shipment consisted of 21.') adult sardines. 

Percentage of 

•c: e 








a S 

1^ ^ 




Total daily for 










4i >i 


2 1 136 80 6 58.8% 9.8% ^^ mortalit.v after 6th day 

until end of 3-week period. 

42 2 47 

^_^ One more fish died between 

112 28 10 25.0'^o 2.5% 11th da.v and end of 3-week 


41 3 27 5 8 18.59;^ 2.3% ^^ mortality after 8th day 

until end of 3-week period. 

Improvements in methods of handling and transporting the fish, sucli 
as less crowding and water movement in the truck's transporting cans and 
the use of artificial light, lowered the mortality significantly (table III 
and fig. 3). The last three shipments showed a drop in mortality from 
84.2 per cent to 42.9 per cent. 

In the Aquarium : 

The high mortality, as a rule, occurs during the first three weeks of 
confinement in the Aquarium. In large measure, it reflects the effect of 
transportation from aboard ship to the Aquarium. In regard to fish sur- 
vival, these three weeks are considered as a critical period. As our records 
show, at the end of the first tliree weeks mortality in the Aquarium hold- 
ing tanks due to transporting and handling was zero. 

Closely comparable results were obtained by Janssen and Alpin (1945). 
They state that mortality attributable to catching and handling among the 
sardines kept in floating boxes for tagging experiments was high for about 
the first six days of confinement (about 55 per cent for the control fish, 
and 70-75 per cent for the tagged fish). Similarly, after 20 days of con- 
finement there were few deaths. 

During the 44-month period, out of the 751 sardines delivered to the 
Aquarium, 520 fish or 69.2 per cent died during the first three weeks of 
captivity in the Aquarium. Approximately 178 fish (23.7) per cent from 
which groups of specimens were drawn for experimental purposes, died 
thereafter because of accidents, starvation, or other causes of natural mor- 

344 I'AfJFOh'MA ACADKMY OF SCIENCEf^ [Pkoc. 4tii Skr. 

tality, while 53 fish (7.1 per cciil ) wei-e still ]ivin<>- on .Jamiary 8, 1953 
(table VI). 

It is shown in labK' V lluit only al)oiit 44 i)er cent of the mortality can 
he attributed to natuial causes. Followinu' the critical period, the mortality 
rate is oxeeedin<iiy low. i^'or exami)le, of the last or seventh shipment of 
275 .sardines, 157 survived the critical three-week period after which mor- 
talities were obsci'ved on the average (n = 9) at intervals of only 57 days 
(interval range, 4-109 days). In one instance, 10 sardines survived 768 
days but a])])eared to be starved or suffering from a pathological condition 
at the end of the exi)erimeut. 





Last Three Shipvients Transported from Los Angeles to »S'on Francisco 
Aboard the "Yelloiriin" 

n', . 

fish delivered 

lU) -fl.- 

ih delivered 

275 flf 

ih del 



Oct. .',. 1950 

on D 

ec. 3. 1950 

on Oct. 22, 








































































9 th 























67. 2 




























































100.0 . 

*The first shipment from the Los Angeles area which incurretl 90 per Cent loss at the of the first 
three weeks is not included due to incomplete daily recording. 


Figure 2. Showing e.xtreme examples of external injury attributable to transporting. 



[Proc. 4tti Skk. 



Mortality due 
to transport- 

Survival of fish of: 












Date of delivery 


first 3 Weeks 








- Per 



















May 19, 1949 





August 4, 1949 




*26 of the 


December 22, 1949 






6th ship- 


June 9, 1950 







ment and 


October 4, 1950 







4 of 


December 3, 1950 







the 5th. 


October 22, 1951 














Factors Causing Mortality 

In Transit: 

Examination of the dead fish immediatelj^ following delivery has indi- 
cated that all of them had lost most of their scales, in some instances, 90 
per cent (fig-. 2). The first fish to die were those with head or abdominal 
injuries causing internal hemorrhages of the brain, heart region, or intes- 
tinal tract. Fish with badly damaged snout areas and fins survive a little 
longer, while those with pinpoint subcutaneous hemorrhages over the entire 
body may live much longer. Fish in the last category with scales intact may 
even recover and survive the three-week critical period. 

When a ship carrying sardines in her bait-tank is preparing to enter a 
harbor which is fed by fresh water, extreme care should be taken in regard 
to the time of entrance. From sad experience we know that the rate of mor- 
tality is high when the salinity is markedly reduced by the circulation of 
brackish water through the vessel's bait-tank. 

To avoid mortality due to changes in salinity or to pollution, sardine- 
carrying vessels should enter port at the beginning of flood tide and unload 
before the ebb. Also, transporting cans to carry the sardines from the 
docked vessels should contain a fresh supply of sea-water for each truck trip. 
This is necessary not only because of the salinity factor, but also because 
of the copious slime secretion and scale .sliedding, both of which interfere 
with normal respiration of the fish. 


Uninterrupted aeration and a temperature differential not exceeding 
S^C. when fish are moved from a vessel to the Aquarium must be main- 
tained to reduce mortality. Also, the use of dip-nets in handling the fisli 
was discouraged in favor of two-gallon buckets with a maximum of four 
sardines per bucket. 

Both external and internal body injuries and extensive shedding of 
scales result from excessive water motion in the truck cans during trans- 
port. When, in 1949, sardines were transported from Monterey Bay to 
San Francisco, 130 miles in 3.5 hours, it was demonstrated that a single 
100-gallon truck can is more satisfactory than four 45-gallon cans. 

Mortality in transport was further reduced by providing adequate arti- 
ficial lighting in the shipping cans so as to reduce mechanical injury of 
the fish due to contact with the sides of the cans. Overcrowding of buckets 
and shipping cans is a distinct limiting factor. This study has shown that 
not more than four adult sardines should be carried in two-gallon buckets, 
10 in 45-gallon cans, and not more than 25 in 100-gallon containers. 

In the Aquarium: 

As stated above, our experience indicates that the number of adult fish 
in the 1,000-gallon tanks of the Aquarium should not exceed 50. Further- 
more, it has been demonstrated that the ill-effects of transportation and 


Mortality Due to Causes Other Than Transporting and Handling 

Mortality Causes t-t „. * t7<- u t» /-. ^ 

" Number of Fish Per Cent 

Experimental and Accidental: 

A. Exposure to anesthetic test 5 2.8 

B. Overexposure to electrical field 25 14.1 

C. Removal tor physiological examination 

after hormone injection 19 10.7 

D. Leaping from holding tanks 12 6.7 

E. Increase of water temperature in 

summer months, 1949, 1950 39 21.9 

Total 100 56.2 


F. Starvation 22 12.3 

G. Other factors (overfeeding, pathological 

causes, etc.) 56 31.5 

Total 78 43.8 

Grand Total 178 100.0 


Sardine Longevity in Confinement 

[Pkoc. 4tii Sf.k. 


, o 






Cj '^ p* 

Recorded for 

of Con- 

"W J- 

Date of 
Sardine Delivery 

Sardine Survival 
After Critical 




3-Week Period 


•^ C 

S "S 

w s s^ 































































































"■The last survivor of the seventh shipment died on May 16, 1954, after having been kept 937 days 
in captivity. 

handling" are felt only during the first three weeks following arrival at the 
Aquarium. During tliis critical period, tlie fish either die or become ada])ted 
to the artificial environment, and the survivors may be kept for relatively 
long periods of time (table VI). 

Temperature is a critical factor in maintaining sardines under aquarium 
conditions. In the Aquarium's tanks without temperature control, water 
temperatures ranged from 11°C. in winter to 20.5°C. in summer. The 
greatest mortality, i)resumably due to temperature, occurred during June 
and July when the temperature rose to above 20°C. 

Our experiments suggest that sardines may be safely kept at aquarium 
temjieratures not exceeding 18 °C. 

These temperature findings are in accord with available information on 
the temperature conditions under wliich sardines exist in tlie sea. The range 
of water temperatures at which sardine schools were obtained on scouting 
surveys conducted by the California Department of Fish and Game was 
from 11°C. to 20.4°C. (California Cooperative Sardine Kesearch Program, 
1951). Daniel Miller* concludes on a basis of field records that "from the 
general ])ictnre it w(mld seem that sardines might prefer temperatures in 
the range of from 13°C. to 18°C. It be remembered, however, that 
these data were collected in all months of the vear and within the entire 

*Scripps Institution of Oceanography 
1952 (mimeographed). 

Rcporl on the 1952 Sardine Conference, page 5, October 30, 



r o 

A^jioijo^ P 95Diua3Jad 9Auo|nujno 

aj "cc a: 

3 o ■;:; 


area surveyed off our coast." Schmidt (1936), who studied the iwassi, a 
Japanese sardine, Sardinops sagax melanostictus, states, "Iwassi is very 
sensitive to temperature and it prefers to keep to regions where the tem- 
perature of the surface water is between 12° and 19° C. The sardines beoin 
to advance to our shores in the Peter the Great Bay when water is warmed 
to a temperature of 8°-10°C., which is usually by the end of May, or in the 
beginning of June. When water temperatures have reached 12°C., the 
catches of sardines become abundant. With the increase of water tempera- 
ture up to 18°-19°C., the sardines move northward in search of cooler 
water." Suehiro (1951) experimented on the tolerance of the Japanese 
sardine to sudden temperature changes. He writes, "when sardines which 
at the time of the experiment were living in sea water of about 22 °C. were 
transferred directly into an experimental tank having a water temperature 
of 28°C. there was only 10-15% mortality in the course of three hours." 
When fish from normal temperature water were transferred directly to 
water with 30°C., "there was within 1 hour and 30 minutes a mortality 
of 60-100 per cent, a result almost equivalent to total destruction." 

During our studies, the hydrogen ion concentration of the circulating 
water in the holding tanks varied from 7.0 to 8.0 pH values. Tests on eight 
adult sardines, 200 mm. in standard length, showed that a 2^H of 6.0 may 
be tolerated for 24 hours without visible harm. 

Artificial light in the transporting cans has been found to reduce mor- 
tality. John Radovitch (in lit.) states that "A night light is necessary to 
keep the fish milling instead of smashing into the sides of the tank. A sur- 
prisingly large number of fish may be confined in a tank if the tank is 
large enough to allow the fish to mill in a circle." In the Aquarium, it is 
not necessary to keep the tanks lighted at night after the critical three- 
week period. By then, the fish have become sufficiently adjusted to the 
space factor to avoid injuring themselves against the walls of the tank. 

Regarding the food of the sardine, Blin (1923), De Buen (1927), and 
Parr (1930; commenting on a paper by Lewis 1929), stress the impor- 
tance of copepods. Dr. Cadet Hand* found that 30.8 mg. of organic matter 
consisting of seven groups — diatoms, dinoflagellates, small copepods, large 
copepods, euphausiids, chaetognaths, and fish eggs — constitutes an average 
stomach content for the Pacific sardine. Significantly, the copepods com- 
prised 25.7 mg. or 83.7 per cent of the total weight. Copepods not being 
readily available, the brine-shrimp, Artemia salina was used for food, and 
it proved to be ideal. For a long period sardines rejected other foods used 
for different fishes of the Steinhart Aquarium, but after confinement of over 
a year they readily consumed finely chopped or ground horse heart, fish, and 
even commercially manufactured dry fish pellets containing protein. 

*Scripps Institution of Oceanography, Report on the 1952 Sardine Conference, p. 18, October 30, 
1952 (mimeographed). 


Interestingly, sardines coexist in the Aqnarinm's tanks with other kinds 
of fishes, such as grunts, Brachydeuterus axillaris; bonefish, Alhula vulpes; 
starry flounders, PUitichthys stellatus: lingcod, Opiodon elongatus; staghorn 
sculpins, Leptoiottus annatus; and buffalo seulpins, Enophrys bison. Usu- 
ally, the sardines occupy the upper, and the remaining species the lower 
water layers of the tanks. No intermingling of the two groups has been 

In conclusion, this study has shown that, despite its delicate physical 
structure and high sensitivity to handling, the Pacific sardine may be satis- 
factorily maintained under aquarium conditions, and that it may, therefore, 
be subjected to experimental investigations. 


This work has been carried on with funds provided by the Marine Re- 
search Committee as a i)art of the California Cooperative Oceanic Fisheries 
Investigations. The authors express their appreciation to Dr. Robert C. 
Miller, director of the California Academy of Sciences, for his keen interest 
in the studies and for encouragement and valuable suggestions during the 
progress of the research; and to Dr. Earl S. Herald, curator of aquatic 
biology, California Academy of Sciences, for his willing cooperation and 
valuable counsel; and Dr. Frances N. Clark, director of the California State 
Fisheries Laboratory, California State Department of Fish and Game; and 
Mv. .John Radovich of the same organization, for helpful suggestions and 
for their interest in tlie study. Acknowledgment is made also to Dr. Rolf 
Bolin. of the iroi)kiiis Marine Biological Station; and Messrs. John C. Marr 
and Ted AVidrig, and Dr. E. H. Ahlstrom, of the U.S. South Pacific Fish- 
eries Investigations, for hol]iful suggestions and constructive criticism of 
the manuscript. 


Allen, W. E. 

1920. Behavior of loon and sardines. Ecology, 1:309-310. 
1930. Elusive sardines. Catalina Islancler, 17(4) :10. 


1923. Note sur I'alimentation de la sardine: Eiiterpes et sardines, ^ociete 
Zoolocilque de France, Bulletin //8(2-3); 99-105, 4 figs. 

Bkedek, C. M. Jk. 

1929. Certain effects in the habits of schooling fishes as based on the observa- 
tions of Jenkinsia. Aviej-icctJi Museum Novitates, 382:1-5. 

BuEX, Fernando de 

1927. Notes preliminares sobre la biologia de la sardine. Inst it ut a espanol de 
Oceangrafia, Notas y resitmenes. ser. 2, no. 15, 55 pp., 7 figs. 


California C»h)ii:hativi, Sakdink Reseakoh Program 
1950. Progress Report. 1950, p. 45. 

COKNISII, Thojias 

18S3. Habits of the pilchard. Zoologist. London, ser. 3,7:505-506. 

HlHB.S. Caui, L. 

1948. "Leapfrogging" by topsnielt and shark. Copcia, 4:298. 

Ja.xssi:n. Jr.. Joii.n F., and J. Ai.frkd Alpin 

1945. The Effect of Internal Tags Upon Sardines. California Fish and Game, 
Fish Bulletin. 61:43-62. 

Lkwis, R. C. 

1929. The food habits of the California sardine in relation to seasonal distribu- 

tion and microplankton. Bulletin of So-ipps Institution of Oceanogra- 
phy, Technical Series, 2(3) : 155-180, 2 figs. 

Newman, E. 

1876. Mr. Saville Kent's lecture, at the Society of Arts, on "The Aquarium: 
construction and management." Zoologist. London, ser. II, 11:4853- 


Parr. A. E. 

1927. A contribution to the theoretical analysis of the schooling behavior of 
fishes. Occasional Papers of the Bingham Oceanographic Foundation. 
1:1-32, 3 figs. 

1930. Is the presence of phytoplankton in the stomach contents of California 

sardine caused by special pursuit or merely due to incidental inges- 
tion? Ecology. 11(2) : 465-468. 

Powers. Edwin B. 

1921. Experiments and observations on the behavior of marine fishes toward 
the hydrogen-ion concentration of the sea water in relation to their mi- 
gratory movements and habitat. Puhlications of the Puget Sound Bio- 
logical Station 3:1-22, 4 pis. 

Schmidt, P. J. 

1936. Migration of fishes. First Edition, Moscow-Leningrad, pp. 90-96 (on 
Japanese sardine), in Russian. 

Siielfori). V. E., and E. B. Powers 

1915. An experimental study of the movements of herring and other fishes. 
Biological Bulletin. 28:315-334, 1 fig., 2 charts. 


1941. Additional social and physiological aspects of respiratory behavior in 
small tarpon. Zoologica. 26:55-60. 

Shlaifek. a., and C. M. Breder, Jr. 

1940. Social and respiratory behavior of small tarpon. Zoologica. 25:493-512. 
2 pis., 1 fig. 


Spooner, G. M. 

1931. Some observations on schooling in fish. Journal of Marine Biological 
Association of the United Kingdom, new series, 17(2) : 421-448, 12 figs. 


1951. A study of the causes of death of bait fishes. Translated from the 
Japanese language by W. G. Van Campen. Washington, D.C., U.S. Fish 
and Wildlife Service. Special Scientific Report: Fisheries no. 62, 57 
pp., 1 pi., 15 figs. 




Fourth Series 
Vol. XXVIII, No. 8, pp. 355-392, figs. 1-28 June 30, 1955 




Curator, Department of Herpetology 
California Academy of Sciences 

The following paper is based upon three separate collections made dur- 
ing the years 1929-1930 (starting on Augnst 1); 1936-1937 (starting on 
October 3); and 1947-1948 (starting on Se])tenib('r 10). 

Practically the full six months allowed on a visitor's visa was spent in 
the field on each trij). This was made ])ossible through the courtesy of the 
director of the Australian Museum, Dr. Charles Anderson; Dr. A. B. 
Walkom who succeeded him; and Mr. J. R. Kinghoi'u, assistant to the 
director and curator of l)irds, reptiles, and am]ihil)iaiis, who attended to 
the various formalities liefore I arrived in Australia, thus saving much 
valuable time. 

To mention all those whose hospitality and hel]) I enjoyed at the various 
cattle and sheep stations, on which most of the collecting was done, would 
l)e next to impossible. I cannot, however, omit those who "opened the door" 
to Australia by starting me on my way: Mr. -lames Burns, of Sydney; Mr. 
William Kelly, of Brisbane; and ^Ir. Charles Craig, of Pci'th, who attended 
to all \\\y wants in AVest Au.stralia. To these gentlemen I am deei)ly in- 
del)ted for arranging my stops on the various stations whose owners or 
managers did everything ])ossible to make my efforts a success. Air. and 
Airs. Percy Allan, of the Ketro Station, and Air. and Airs. Charles Barnard, 
owners of Coomooboolaroo, cannot Ix^ left unmentioned. 

AVeather conditions were ideal on all three trijis. On the third I was 
fortunate enough to encounter a few light rains which brought out a variety 
of am]>hil)ians not met with before. AVorld AVar II, however, still had a 
lingering and disastrous effect on travel conditions and these greatly hin- 



dered field a('ti\itie.s so tiiat it was iiiii)()ssil)le to control one's movements or 
take along all of the necessary collecting equipment. For this reason much 
valuable time was lost in AVest Australia at the conclusion of the third 

During the study of the material collected it was found that there were 
three amphibians which appeared to be new. In order not to run the risk 
of adding to synonymy these were sent to Mr. Arthur Loveridge to compare 
with specimens in the almost complete collection of Australian amphibians 
in the Museum of Comparative Zoology at Harvard. jVIr. Loveridge very 
kindly set aside his own work and spent considerable time studying this 
material and agreed with me that they were new species. These were origi- 
nally described in the Proceedings of the Biological Society of Washington, 
vol. 63, pp. 131-138, December 29, 1950. 



Callandoon: September 23, 1947-October 3, 1947. A sheep station in 
the vicinity west of Goondiwindi, southeastern Queensland. This is a thinly 
wooded area and contains grasslands with scattered ring-barked gum trees 
of several species and patches of brigalow scrub. 

Clermont: August 14, 1929. A town in eastern Queensland at the ter- 
minus of the Emerald-to-Clermont railway. It is typical sheep country 
with open grasslands and scattered gum trees. 

Coomoohoolaroo: September 15-October 26, 1929, and October 10- 
November 15, 1936. A cattle station fifteen miles south of Duaringa, east 
central Queensland. This area is heavily wooded with iron bark trees and 
much ring-barked country. Several lagoons are on the station but there is 
no running water, although there are a few stream beds which may have 
.some at long intervals. Many fallen and many dead but standing trees 
are present. Tliis was one of the best collcM-ting grounds encountered. 

Du(trin<i(t: November 13, 1936. A vibage on the Rockhani])ton-to- 
Emerald railway fifty-five miles west of Rockham]iton. 

Gaeta: December 29, 1936, and .Jaiuiary 11, 1937. A cattle staticm ten 
miles west of Kolonga Creek Station. It consists of heavily wooded and a 
considerable number of rocky areas. Koh)nga Creek runs through this 


Goondiwindi: September 28, 1947. A town on the Brisbane-to-Dirran- 
bandi railway 140 miles west of Brisbane. 

Kolonga Creek: December 24-29, 1936, and January 1-13, 1937. A 
cattle station twenty-five miles north of GinGin. This area consists of open 
rolling grasslands with many large gum trees, many of them ring-barked. 
A moderate-sized and permanent stream (Kolonga Creek), with some deep 
l^ools along the banks, runs through the property. 

Meirgenta: November 29, 1936. A sheep station in the vicinity east of 
Retro and similar in character to it (see below). 

McPherson Baneje: October 3-7, 1936. A wooded mountain range (alti- 
tude 2,500 feet) with clearings on the ridges. Some lumbering has been done 
and there are roads thrcmgh the virgin forest. On the clear ridges are many 
fallen trees and burned out logs and stumps of the large gums. This range 
is located fifty miles inland on the New South Wales-Queensland border. 

Noondoo: September 10-20, 1947. A sheep station south of the Noondoo 
siding ten miles east of Dirranbandi, southeastern Queensland. There are 
scattei-ed small trees and much open country with artesian-well water. A 
peculiar formation not seen elsewliere was a number of sand bars parallel- 
ing each other across some of the open grasslands. 

Retro: August 1-September 10, 1929, and November 30-December 12, 
1936. A sheep station twenty miles west of Capella, eastern Quensland. 
Here are open grasslands and scattered gum trees with moderately heavy 
groves of the tea tree and an occasional bottle tree. Both black and red soil 
are in evidence. Though the surrounding country did not look attractive 
as a collecting ground it proved to be excellent, with a large number of 

Tcdafa: November 17-28, 1936. A sheep station twenty miles south of 
Emerald. It is partly wooded with a heavy growth of brigalow scrub but 
includes much open grassland. Bottle trees are much in evidence on this 

New South AVales 

Blue Mountains : Localities visited in this region were Cox's River, Feb- 
ruary 15, 1930; Hampton, February 14, 1930; and the Hampton-Lithgow 
area, January 19-23, 1948. 


Bundy: October 23-November 15, 1947. A shee]) station twenty miles 
southeast ot* Moree, northeastern New South Wales. This is open grassland 
with considerable brigalow scrub. A small creek, dried up in the summer 
months, runs through tlie property. A small amount of fallen timber and 
ring-barked trees characterizes this area. 

Carroll: November 2, 1947. A sheep station ten miles west of Moree. 
This consists of the same type of country as that around Bundy Station. 
It also has a creek whicli is dried up in the summer months. 

Keera: October 14-20, 1947. A cattle and sheep station fifteen miles 
southeast of Bingara, northeastern New South Wales. This is a wooded, 
hilly country with tlie Gwydir River, a fast-flowing stream, running through 
the property and giving an abundant and permanent supply of water. 
This region also includes a considerable area of open grasslands with 
scattered trees. 

Sydneij: November 7, 1929, February 9, 1930, February 19, 1948. The 
metropolis of New South Wales. 

Ulong: January 27-February 15, 1948. A lumber mill town twenty- 
eight miles inland from Coff's Harbor, northeastern New South Wales. 
This is hilly open country with scattered trees and includes several streams 
of water and an area of virgin forest in which lumbering operations are 
much in evidence. Numerous log-roads allow access to the untouched areas. 


Ned's Corner: November 20-December 6, 1929. A sheep station on the 
banks of the Murray River, thirty-seven miles west of Mildura. It is 
characterized by open grassland and red soil. There is, on the banks of 
the river, a heavy growth of large gum trees. 

West Australia 

Geraldton: January 5, 1948. A town on the coast of West Australia. 
It is situated between 28 and 29 degrees south latitude. 

Nangahrook: January 8-22, 1930. A lumber mill in the heart of the 
giant gum forests, eighteen miles east of Yarloo}) on the main southwestern 
line from Perth to Pemberton. It consists of a virgin forest of giant gum 
trees with a heavy growth of ferns and grass trees, or "black boys" as 
they are sometimes called. 


Pelsart Island 

This island, visited December 1-30, 1947, is a coral reef beloiifjiiig to 
the southern group of the Abrolhos Islands, forty miles west of Geraldton. 
Latitude 29° south, longitude 114° east. It is approximately six miles long 
and three-quarters of a mile wide at the widest part, and is composed of 
large flakes and small pieces of broken coral, covered in places with low 
scrub vegetation and a few mangroves in a lagoon in the reef. 



Mixophyes fasciolatus fasciolatus Giinther 

This name is ai)i)]ied to five specimens, 2 (C.A.S. Nos. 77824-77825) 
from Cox's River in the Blue Mountains and 3 (C.A.S. Nos. 82049-82051) 
from Ulong, New South Wales. Loveridge^ has given the subspecific name 
scheviUi to a form from northern Queensland having only two phalanges 
of the fourth toe free of web, instead of three as in typical M. f. fasciolatus. 
Two specimens from Ulong tend to show this character, but a third (juve- 
nile) has three jiha hinges of the fourth toe entirely free of web. 

In the adpressed limb the til)io-tarsal joint reaches the snout; vomerine 
teeth in two prominent straight series between the choanae; tongue nitched 
behind; a large oval tymjnnium; subarticular tubercles prominent; a large 
inner metacarpal tubercle and a ])r()minent inner metatarsal tubercle; a 
prominent crescent-sha])ed fold of skin over the lym]>anum. 

The color, in life, is grayish or brownish, with irregular patches of a 
darker shade; limbs of adults with broad crossbars, obsolete in some indi- 
viduals. The juvenile from Ulong shows distinct narrow bars of black. A 
characteristic heavy dark-brown bar between the eyes is absent in No. 
82050 from Ulong. The thighs are marbled with black and the gi'oin is 
whitish with black spots. Under surfaces are whitish. 

The largest specimen is in the Ulong series, a female (82050) 92 mm. 
in length. It was dug out of a rotten stum]) on a grassy hillside. 

Cyclorana alboguttatus (Giinther) 
(Figure 1) 

Six specimens (C.A.S. Nos. 77973-77977) from Coomooboolaroo and 1 
(C.A.S. No. 77978) from Kolonga Creek. 

1. Occasional Papers of the Boston Society of Natural History, S, p. 55, 1933. 



[Proc. 4th Skb. 

Three individuals show the vomerine teeth in two straight series be- 
tween the ehoanae. In one the series touch on the median line and one 
(No. 77977) has a very prominent single series. Tongue slightly nicked 
behind; tympanum prominent; two moderate-sized metacarpal tubercles; a 
large elongate metatarsal tubercle; subarticular tubercles prominent; toes 
about one-third webl)ed; in adpressed limb tibio-tarsal joint reaches eye, 
or slightly beyond. 

Skin smooth, with scattered flattened tubercles; a prominent crescent- 
like fold over tympanum; lateral folds prominent; belly and back of thighs 
granular; throat smooth. 

Color, in life, dark slate to ashy gray; back of thighs covered with small 
round spots of white; sides mottled with black; belly w^hitish; a more or 
less obscure dorsal stripe; throat mottled or clouded with dark gray. 

The Kolonga s])ecimen was taken in a puddle under a cattle guard, 
those from Coomooboolaroo from crevices between the rocks lining a well. 

A si^ecimen from Coomooboolaroo measures 60 mm. from snout to vent. 

Cyclorana australis (Gray) 

Nine specimens (C.A.S. Nos. 77828-77836), the largest species of the 
genus encountered, were taken at Coomooboolaroo in Octol)er and Novem- 
ber, 1936, and a tenth one from Kolonga Creek, .January 1, 1937. 

Fig. 1 (left). Cyvlorana (ilhofjutlatiis C.A.S. No. 77977. Coomooboolaroo Cattle 
Station, 15 miles south of Duariuga, Queensland. Natural size. 

Fig. 2 (right). Cytloranu hrevipes C.A.S. No. 77827, Coomooboolaroo Cattle 
Station, 15 miles south of Duaringa. Queensland. Natural size. 


Vomerine teeth in two prominent series between the choanae; tongue 
slightly nicked behind; tympanum distinct, but moderate in size; two large 
metacarpal tubercles; a ]>rominent elongate and free-edged metatarsal tu- 
bercle; subarticular tubercles prominent; in adpressed limb tibio-tarsal 
joint reaches midway between tympanum and eye or to the eye; toes 
slightly webbed. 

A very prominent dorsolateral ridge is present. A distinct fold of skin 
passes from the front of the nostril over the eye and the tympanum to the 
corner of the mouth. The skin is granular in males and smooth or very 
slightly granular in females. The toes are slightly webbed. 

In life, the dorsal surface is grayish, spotted and marbled with black. 
The undersurfaces are whitish, the throat being spotted with dark gray in 
females and uniform dark gray in males. No. 77837, a breeding male, lias 
black inner metacar])al tubercles. 

The Coomooboolaroo specimens were taken at 9 :30 at night in the high 
grass alongside a pool of water. The Kolonga specimen was dug out of the 
lawn in front of the homestead. 

This species is known to the pastoralists as the "Bully Frog." 

A specimen from Coomooboolaroo measures 87 mm. from snout to vent. 

Cyclorana brevipes (Peters) 

(Figure 2) 

Two specimens (C.A.S. Nos. 77826-77827) are from Coomooboolaroo. 
Vomerine teeth in two round clusters between choanae; tongue nicked be- 
hind; tympanum distinct; two prominent metacarpal tubercles; prominent 
metatarsal tubercle with free inner edge; subarticular tuberclas prominent; 
fingers rounded; toes slightly webbed; tibio-tarsal joint in adpressed limb 
reaches to middle of tympanum. 

Skin smooth with small, flat warts; sides and belly granular; throat 
smooth, except that of males which is finely granular; a crescent-like fold 
of skin over the tympanum. 

In life, the color is gray wdth irregular dorsal spots and blotches. A 
narrow doi*sal stripe is present in the two specimens at hand. Belly whit- 
ish; throat of female (No. 77827) clouded with light gray; throat of male 
(No. 77826) slate color with the edges somewhat darker. 

The female measures 48 mm. from snout to vent. 

Both specimens were taken on the banks of a lagoon under }>ieces of 
bark half buried in the mud. 

Cyclorana inermis (Peters) 

(Figure 3) 
This was a common species at Coomooboolaroo, where 135 specimens 



[Proc. 4th Skr. 

(C.A.S. Nos. 77838-77972) were taken. Five (C.A.S. Nos. 77915-77919) 
are from Kolonga Creek. 

A froo' of slender build with a large tympanum situated very close to 
the eye and about two-thirds as large. Vomerine teeth in two small clusters 
between fronts of choanae; tongue nicked behind; two small metatarsal 
tubercles; subarticular tubercles prominent; toes two-thirds wxbbed; in ad- 
pressed limb tibio-tarsal joint reaches snout. 

Tlie skin is rugose; belly and back of thighs covered with tubercles; 
throat smooth; a gular fold present. 

In life, the color is gray, with blackish dorsal spots; back of thighs 
mottled with black; undersurfaces clear, except the throat which is dark 
in males. The lips are spotted with white. 

The entire series were taken in the garden of the homestead, where 
they were found hidden at the base of the leaves of the pineapple plants. 
A few were found hopping about the pathways after dark. 

Thii-ty-five mm. from snout to vent is a large individual. 

Cyclorana slevini Loveridge 

(Figure 4) 

Two specimens of this frog (C.A.S. Nos. 82052-82053) taken at Noon- 
doo were described by Loveridge-. Its nearest relative apj)ears to be C. 


Fig. 3 (left). Cyclorana inermis C.A.S. No. 77920, Coomooboolaroo Cattle Sta- 
tion, 15 miles south of Duaringa, Queensland. Natural size. 

Fig. 4 (right). Cyclorana slevini C.A.S. No. 82052, Noondoo, Queensland. Na- 
tural size. 

2. Proceedings of the Biological Society of Wasliington, vol. 63. pp. 131138, Dec. 29, 1950. 


oustraJis, from which it differs in interorbital width, size of tympanum, 
and webbing of toes. 

Vomerine teeth in two prominent clusters between choanae; tongue 
rounded behind; tymi^anum kirge, about the size of orbit; an elongate inner 
metacarpal tubercle; an inner free-edged metatarsal tubercle; subarticular 
tubercles present, but not prominent; toes two-thirds webbed; in adpressed 
limb tibio-tarsal joint reaches tympanum; lateral folds absent; belly and 
anal region granular. 

In life, the color is grayish, with darker spots or reticulations. The 
undersurfaces are yellowish-white. 

Both specimens were taken half buried, in sandy soil, under old slabs 
of bark. The larger one, a female, is 48 mm. in length from snout to vent. 

Limnodynastes convexiusculus (Macleay) 

A single specimen of this species (C.A.S. No. 78606) was taken at 
Kolonga Creek. 

Vomerine teeth in a long, straight series extending well beyond the 
outer edge of choanae; tongue oval, tapering toward end and slightly nicked 
behind; tympanum very indistinct; a prominent inner and slightly smaller 
outer metacarpal tubercle with an elongate tubercle between; an elongate 
metatarsal tubercle; subarticular tubercles very prominent and conical in 
shape; fingers and toes long and rounded, without enlarged terminal disks; 
a rudimentary web between the third and fourth and the fourth and fifth 
toe; in the adpressed limb tibio-tarsal .joint reaches back of the eye. 

The skin is smooth, with the back of the thighs slightly granular. 

In life, the ground color is light reddish-brown; a wide blackish band ex- 
tending from the tip of the snout to the forearm. A whitish tapering stripe 
from the back of the eye to the forelimb, widest posteriorly; a number of 
large blackish spots on the sides; three wide blackish dorsal stripes with 
undulating edges, the center one with a very narrow longitudinal stripe of 
white; undersurfaces whitish, the throat and sides of belly clouded with 
reddish -brown. 

This, a half-grown sjiecimcn, was taken under a piece of bark on the 
banks of Kolonga Creek. 

Limnodynastes dorsalis dumerilii Peters 

(Figure 5) 

This species was not met with in numbers, only three specimens (C.A.S. 
Nos. 78035 and 82196-82197) from the Hamptoh-Lifligow area in the Blue 
Mountains being taken. All agree in liaving no doi'sal s1i-i]>e and with 
the toes devoid of fringe>>. • 


Vomerine teetli in a straijiht series, with a very slight division, and ex- 
tend to middle of the choanae; tongue rounded behind, tympanum hidden; 
metacarpal tubei-cles i)r()minent; a very large s])ade-like inner metatareal 
tubercle; no outer tubercle; subarticular tubercles prominent, those at the 
base of toes being ])articularly so; toes slightly fringed and webbed; ti})io- 
tarsal joint when carried forward reaches tympanum. 

Fig. 5. Lininodyna.ites (lorsalis dur.ierilii C.A.S. No. 78035, Hampton, New 
South Wales. Natural size. 

The skin is smooth, covered with minute tubercles and small, flat, glands; 
an elongate gland extends from the back of the eye to the shoulder; a white 
lateral fold; a prominent round whitish gland at each side of the vent; tibia 
with a large, oval-shaped gland; back of thighs covered with small tubercles. 

The color, in life, is dark brown with scattered dark markings; sides 
and belly with dark brown reticulations; throat uniformly dark brown or 
light brown with reticulations of a darker shade. 

The largest specimen of the series measures 69 mm. in length. One 
(C.A.S. No. 82197) was taken towards late afternoon in a small hollow 
in the base of a tree. 

Limnodynastes dorsalis terraereginae Fry 

(Figure 6) 
A series of eight specimens; 3 (C.A.S. Nos. 78038-78040) from Coo- 


mooboolaroo; 2 (C.A.S. Nos. 78036-78037) from Kolonga, and 3 (C.A.S. 
Nos. 82198-82200) from Keera have the second finger considerably flat- 
tened and heavily fringed. 

The vomerine teeth extend to the outer edge of the choanae. The hind 
leg is slightly shorter than in its congener L. d. dumerilii, which it re- 
sembles in other respects, except coloration. 

Fig. 6. Lini nodyxasies dofsali.s terra) eginae C.A.S. No. 78038, Coomooboolaroo 
Cattle Station, 15 miles south of Duaringa, Queensland. Natural size. 

A specimen (No. 78036) was colored in life as follows: ground color 
black; throat, sides and back of jaw yellow; lower lateral surfaces yellow, 
with black si)otting and reticulations; thighs and undersurface of hind legs 
red; reddish areas on top of forelegs; undersurfaces yellowish, with sides 
and throat mottled with gray. Two specimens from Keera, two from Coo- 
mooboolaroo, and one from Kolonga show signs of a dorsal stripe. 

A specimen from Coomooboolaroo is 64 mm. in length. The Kolonga 
specimens were taken in the water under a cattle guard and those from 
Keera in newly dug postholes which filled with rain during the night. 

Limnodynastes fletcheri Boulenger 

(Figure 7) 
Twenty-nine specimens (C.A.S. Nos. 82355-82381 and 82389-82390) 



[Proc. 4tii Skk. 

from Callandoon, 2 (C.A.S. Nos. 82382-82383) from Keera, and 5 (C.A.S. 
Nos. 82384-82388) from Biuidy. 

Vomerine teeth in two short series in back of ehoanae; tongue rounded 
behind; tyin])anuin indistinet; a large inner and small outer metacarpal 
tubercle; a small elongate inner metatarsal tubercle; subarticular tubercles 
present; first finger slightly fringed, second more heavily; toes slightly 
webbed; in adpressed limb tibio-tarsal joint reaches tympanum. 

A prominent fold of skin is j^resent from below the back of the eye 
to the forearm. The top of the hind legs and the anal region are covered 
Avith small tubercles. No. 82371, a breeding male, has a swollen thumb and 
the bone protruding as in L. pevonii. 

Color, in life, slate-gray, with irregular spots and blotches over the sides 
and dorsal surface; limbs crossbarred; a black spot under the eye; usually 
a dorsal stripe of white and a pink spot over the eye; undersurfaces whit- 
ish; throat clouded with dark gray. 

In the present series of 36 specimens 10 are without the dorsal stripe 
and 6 lack the i)ink spot over the eye. 

A large male measures 5-4 mm. from snout to vent. 

All were found under old logs or bark in moist depressions in paddocks. 

Fig. 7 (left). Limnodynastes flrtcheri C.A.S. No. 82370, Callandoon Sheep 
Station, vicinity west of Goondiwlndi, Queensland. Natural size. 

Pig. 8 (right). Limnodynastes ornatus C.A.S. No. 78050, Coomooboolaroo Cattle 
Station, 15 miles south of Duaringa, Queensland. Natural size. 


Limnodynastes ornatus (Gray) 

(Figure 8) 

Eleven specimens were taken, 10 being from Coomooboolaroo and one 
from Kolonga. 

Vomerine teeth in two straight, or very slightly curved, series extend- 
ing beyond choanae; tongue slightly nicked behind; tympanum indistinct; 
a large inner and moderate-sized metacarpal tubercle; large shovel-shaped 
inner metatarsal tubercle; no outer tubercle present; subarticular tubercles 
prominent; toes slightly webbed and heavily fringed, in the adpressed limb 
the tibio-tarsal joint reaches the tympanum. 

The skin is smooth with numerous small warts; a cluster of tul)ercles 
below the vent; undersurfaces smooth; a cluster of small tubercles at the jaw. 

In life, the ground color is purplish-gray with numerous dark spots and 
reticulations; a dark streak extending through the nostril and eye to the 
forelimb; a dark band between the eyes and a large squarish spot below 
the eye; undersurfaces whitish, with the sides of the throat grayish. 

The largest specimen measures 39 mm. in length. 

All of the Coomooboolaroo frogs were taken at the waters edge of a 
shallow, muddy lagoon. The Kolonga specimen (No. 78501) has the third 
finger of each hand greatly flattened and the bone protruding from the tip. 
Boulenger^ states in a footnote that a female had on the breast two cica- 
trices evidently caused by the thiunbs of the male. The thumbs in the 
Kolonga specimen, however, show no signs of a swelling or presence of 
protruding ])one, this, as stated above, takes ]dace in the third fiinger. 

Limnodynastes peronii peronii (Dunieiil and Bibron) 
(Figure 9) 

The series at hand comprises 32 specimens from New^ South AVales, 15 
(C.A.S. Nos. 82065-S2076 and 77986-77988) from the Plampton-Lithgow 
area, 5 (C.A.S. Nos. 77981-77985) from Cox's Eiver in the Blue ^Mountains, 
and 12 (C.A.S. Nos. 82054-8206-t and 82391) from Ulong. 

Vomerine teeth in two straight series behind and extending l)eyond 
choanae; tongue rounded in back; tympanum hidden; two well-developed 
metacarpal tubercles; a small to mediiun metatarsal tubercle; subarticular 
tubercles present, except on first finger; adult females w4th two inner 
fingers flattened and distinctly fringed; adult males with a sharply pointed 
metacarpal bone protruding from a longitudinal slit on top of the first 
finger; toes w^ith only a trace of w^eb. This is shown very distinctly in 12 

3. Catalogue of Batrachia Salientia, p. 262. 



[Proo. 4th Skb. 

males of the series. In the adpressed limb the tibio-tarsal joint reaches the 
eye, or between the eve and the nostril, but in none of the specimens does 
it reacli beyond the nostril. 

Skin smooth; a distinct fold from the back of the eye to the forelimb; 
lateral folds present; back of thighs liranular; belly and throat smooth. 

In life, the color is dark to light gray with a heavy dorsal black line, 
divided by a narrow white one, from between the eyes to the vent. Promi- 
nent lateral black lines are present. The dorsal line is occasionally broken, 
which happens in two of the Blue Mountain frogs and in one from Ulong. 
In general, the Blue Mountain frogs are a much darker shade than those 
from Ulong, which are light gray, and the white line is often obsolete. 

Undersurfaces whitish, with throat, sides of belly, and undersurface 
of hind limbs spotted with gray. 

As tliese frogs have considerable variation as to striping, and liaving 

Fig. 9. Livinodi/uaste.s perojiii peronii C.A.S. No. 77983, Cox's River, New South 
Wales. Natural size. 


prominent, obsolete, or broken stripes, and vary in shade as well, Parker^ 
may be safely followed in considering L. p. Imeatus a synonym. 

All of these frogs were taken at night while hunting with a flashlight, the 
ones from Ulong being found in a small puddle with a heavy growth of grass 
surrounding it, and those from the Blue Mountains in a hillside spring. 

Sixty mm. is the average length of an adult. 

Limnodynastes salmini Steindachner 
(Figure 10) 

This species is represented by 52 specimens, 50 (C.A.S. Nos. 82077- 
82126) from Callandoon and 2 (C.A.S. Nos. 82127-82128) from Bundy. 

Vomerine teeth in two scarcely separated and slightly curved series be- 
hind choanae; tongue rounded behind; tympanum indistinct; a large inner 
metacarpal tubercle and moderate outer one; a large elongate inner meta- 
tarsal tubercle; subarticular tubercles moderate. No. 82125, a large female, 
has a trace of fringe on the second finger; toes with or without a bare sug- 

Fig. 10. Limnoclyuastrs .salinini C.A.S. No. 82115, Callandoon Sheep Station, 
vicinity west of Goondiwindi. Queensland. Natural size. 

4. Novitates Zoologicae, vol. 42, part I, p. 57, April, 1940. 


gestion of web; in the adpressed limb the tibio-tarsal joint reaches the 

Skin smooth; a prominent fold of skin from tlie back of the eye to the 
foreleg; a lateral fold present; region of vent slightly tubercular; under- 
sur faces smooth. 

In life, the ground color is gray, with an elongate black patch back of 
the eye; a small black patch under the eye; a w^hite or pink line from the 
back of the eye to the foreleg; a broken black dorsal line, with a narrow 
one of pink down its center; two broad lateral lines of pink, bordered by 
narrow, broken lines, of black; undersurfaces whitish, sides of the throat 
clouded with gray. 

Sixty mm. from snout to vent is adult size. 

The entire series from Callandoon was taken in paddocks drying out 
from recent rains, but with a few mudholes left. The frogs were under old 
logs and pieces of bark wherever there was sufficient moisture left. The 
two from Bundy were under the bark of a fallen tree. 

Limnodynastes tasmaniensis Oiinther 

(Figure 11) 

A series of 113 specimens are from the following localities: Retro, 20 
(C.A.S. Nos. 77790-78009) ; Magenta, 2 (C.A.S. Nos. 78010-78011) ; Talafa, 
13 (C.A.S. Nos. 78023-78034 and 77987) ; Coomooboolaroo, 11 (C.A.S. Nos. 
78012-78022) ; Callandoon. 45 (C.A.S. Nos. 82148-82192) ; Bundy, 2 (C.A.S. 
Nos. 82194-82195); Keera, 1 (C.A.S. No. 82193); Blue Mountains, 19 (C. 
A.S. No. 82129-82147). 

Vomerine teeth in two straight series behind choanae and somewhat 
variable in length, extending either to inner edge of the choanae, middle, or 
in some individuals slightly beyond outer edge; tongue slightly nicked 
])ehind; tympanum indistinct; metacarpal tubercles prominent; two meta- 
tarsal tubercles; inner slightly elongate; outer very small, occasionally in- 
dicated by a white spot; subarticular tubercles ]n'esent; fingers fringed; 
toes lightly fringed, with indication of web; in adjn'essed limb tibio-tarsal 
joint reaches tympanum or to the back of eye. 

Skin smooth, with flattened warts above; a prominent fold of skin from 
the back of the eye to the forelimb; anal region and ])ack of thighs with 
prominent tubercles; a dermal flap covering the vent; undersurfaces smooth. 

The ground color, in life, is slate or gray, with irregular spots or blotches 
of black; a black l)and extends from the nostril through the eye to the 
forelimb; limbs spotted or crossbarred; occasional areas of pink on the dor- 
sal surface; a dorsal stripe of yellow, ])ink, or white. The throats of males 
are suffused with dark gray or slate. 

In the ])resent series 36 have the dorsal stri]ie ])rominent; in 49 it is 
ol)solete, and in 28 absent. 



A female from Retro measures 43 mm. in length. 

In the dry sheep country such as Retro and Talafa these frogs were 
found at night about the watering troughs and tanks or sitting along the 
pipes leading from the tank to the trough. Farther south at Callandoon 
they were in the muddy parts of paddocks, where the surface still had 
some water from recent rains. 

Parker^ is being followed in considering L. pJafjicephalus and L. oliva- 
ceus Loveridge'' as synonyms. 

Lechriodus fletcheri (Boulenger) 
(Figure 12) 

This was not an uncommon frog in the virgin rain forest in the vicinity 
north of Ulong where 31 specimens (C.A.S. Nos. 82201-82231) were taken 
at an elevation of approximately 1,800 feet. 

Vomerine teeth in a straight, or very slightly curved, series extending 

Fig. 11 (left). Limnodynastes tasvianiensis C.A.S. No. 77993, Retro Sheep Sta- 
tion, 20 miles west of Capella, Queensland. Natural size. 

Fig. 12 (right!. Lechriodus Jletrheri C.A.S. No. 82201, Ulong, New South Wales. 
Natural size. 

5. Novitates Zoologicae, vol. 42, part I, p. 52, April, 1940. 

6. Bulletin of the Museum of Comparative ZooloK.v, vol. T.XWllT, Xo. 1. p. 19, January. 1935. 



to the outer edge of choanae; tongue slightly nicked behind; tympanum 
distinct and oval, with minute black tubercles scattered over surface; a 
large inner metacarpal tubercle and a smaller outer one; a moderately oval 
metatarsal tubercle; subarticular tubercles prominent; breeding females with 
first and second toes flattened; toes slightly webbed and heavily fringed; 
toes of males bordered by minute spines; in adpressed limb tibio-tarsal 
joint reaches snout. 

The skin is smooth, but so heavily covered with minute black tul>ercles 
it has the texture of a very fine sandpaper; backs of limbs similarly cov- 
ered; a prominent fold from the back of the eye to a point midway be- 
tween the legs; undersurfaces smooth. 

In life, this is a remarkably colored frog. There are two shades for the 
ground color, a light gray or a chestnut brown. A heavy black line extends 
from the back of the eye, sharp-edged on top and undulating below, en- 
compasses the top of the tympanum and extends to the corner of the mouth. 
This is occasionally bordered above by a narrow line of pink. Tops of first 
and second fingers are sometimes black; backs of thighs and undersurfaces 
of feet are black; the limbs are crossbarred with black, the hind limbs 
occasionally tinted with pink; the lower surfaces are whitish, the lower jaw 
being edged with a very narrow black line underneath. 

An average specimen of the series at hand measures 44 mm. in length 
and the largest 50 mm. 

All of these frogs were taken in the virgin rain forest where timber 
was being cut. Ruts formed by the heavy tires of the logging trucks along 
the roads and filled with water by the night rains were their favorite 
haunts. On the collecting date, February 15, these frogs were breeding 
and the puddles contained heavy patches of gelatinous froth containing 
quantities of small, black eggs. 

Adelotus brevis (Giinther) 
(Figure 13) 

This species, one of the most strikingly colored of the Australian am- 
phibians, was taken in three localities, 1 (C.A.S. No. 78052) from the Mc- 
Pherson Range, 2 (C.A.S. Nos. 78053-78054) from Kolonga, and 123 (C. 
A.S. Nos. 82232-82354) from Ulong. 

Vomerine teeth in two oblique series behind choanae ; maxillary teeth 
prominent; tongue rounded behind; tympanum hidden; prominent inner 
metacarpal and metatarsal tubercles and less prominent outer ones; sub- 
articular tubercles prominent; females with first two fingers flattened and 
fringed, with slight indication of web; toes with slight indication of web; 
in adpressed limb tibio-tarsal joint reaches tympanum. At the tip of the 
lower jaw are two tooth-like erections of skin, very prominent in males 
and much reduced or hardlv discernible in females. 



The skin is smooth with a light scattering of small warts, whieli are 
heavier and more numerous on the back of the thighs. 

In life, the ground color is light or dark gray with irregular markings 
of black; a large black blotch on the top of the head and a curved band 
from the tip of the snout, passing through the eye to the corner of the 
mouth; limbs heavily crossbarred with black; the undersurfaces are rich 
pink, heavily mottled or reticulated with black. In alcohol this pink dis- 
appears rapidly, leaving a dull white color. Some individuals have the 
throat clouded with gray and minutely spotted with pink. 

An adult measures 37 mm. in length. 

The McPherson Range specimen was found in the imprint of a horse's 
hoof in the mud at the side of a water trough, while those from Ulong were 
found under logs, bark, boards, and various types of debris. A few were 
found in shallow rain puddles hiding at the roots of the grass and covered 
with a blanket of small bubbles. No signs of eggs could be found. 

Crinia signifera signifera (Girard) 
(Figure 14) 

This highly variable species w^as taken in three localities: 36 (C.A.S. 
Nos. 78066 78101) from Xangabrook; 6 (C.A.S. Nos. 82396-82401) from 
Callandoon; 1 (C.A.S. No. 82394) from the Blue Mountains; and 1 (C.A.S. 
No. 82395) from Ulong. 

The majority of those from Nangabrook are not mature and it is dif- 

Figure 13 (left). Adelottis breris C.A.S. No. 822.53, Ulong, New South Wales. 
Natural size. 

Fig. 14 (right). Crinia signifera signifera C.A.S. No. 82394, Blue Mountains, 
New South Wales. Natural size. 


ficult to distinguish the charaeteristies of the species. However, vomerine 
teeth lacking in more mature individuals; tympanum hidden; a large outer 
and small inner metacarpal tubercle; palmar and subarticular tubercles 
very prominent; a small inner and outer metatarsal tubercle; back of thighs 
granular; belly and sides with flattened granules; dorsal surface smooth. 
In adpressed limb tibio-tarsal joint reaches close to back of eye. 

In life, the color brown or bluish-gray, some individuals with a black 
lateral stripe from tip of .snout to groin; back of thighs occasionally pink; 
a dark area on dorsal surface back of eyes and a trace of a narrow verte- 
bral stripe; undersurface whitish, spotted, or marbled with black. 

The Callandoon specimens are less tubercular and have a tendency 
towards a uniform grayish, with hind limbs crossbarred with black, while 
the one from the Blue Mountains has a broad black dorsal band. 

One of the largest individuals is 24 mm. from snout to vent. 

The entire series was taken under bark slabs or at the roots of grass 
tufts close to standing water. 

Crinia georgiana Tschudi 

Eleven specimens (C.A.S. Nos. 78055-78065) were taken at Nangabrook. 

Not one of those at liand shows any signs of vomerine teeth; tympanum 
barely visible; a large outer and small inner metacarpal tubercle; a small 
inner and outer metatarsal tubercle; palmar and subarticular tubercles 
prominent; in adpressed limb tibio-tarsal joint reaches to back of tympanum. 

The dorsal surface is smooth; belly and sides heavily tuberculated; 
throat with more flattened tubercles. 

The largest specimen in the series measures 32 nun. from snout to vent. 

In life, ground color brownish with black areas over dorsal surface; a 
black lateral strii)e from tip of snout to groin, which has a rich reddish to 
pink area; back of thighs similarly colored; undersurface whitish, sprinkled 
witli black; a dark area in the middle of throat. 

This series was taken about the roots of grass tufts close to stand- 
ing water. 

Uperolia marmorata Gray 

(Figure 15) 

Only two specimens of this species were taken, 1 (C.A.S. No. 82402) at 
Noondoo, and 1 (C.A.S. No. 82403) from the Blue Mountains. 

Vomerine teeth absent; tongue rounded behind, tympanum hidden; a 
large parotoid gland present; an inner and outer metacarpal tubercle; a 
prominent elongate metatarsal tubercle and a rounded outer one; subarti- 
cular tubercles prominent. The two specimens mentioned show no indica- 
tion of webbing but have the fingers and toes rounded. In the ;ul])ressed 
limb the tibio-tarsal joint reaches the back of the eye. 



Skin smooth with longitudinal rows of small tubercles; undersurfaee of 
thighs and the anal region covered with small tubercles. 

In life, the color is olive-brown with a white groin patch; l)ack of thighs 
whitish; undersurfaces yellowish, the throat and belly are clouded with 
gray and minutely spotted with black. 

The larger specimen is from the Blue Mountains and measures 28 mm. 
from snout to vent. 

Both were taken under stones. 

Uperolia rugosa (Anderson) 
(Figure 16) 

This was found to be an abundant species in southern Queensland and 
was also found in northern New South Wales. It was taken at the follow- 
ing localities: 3 (C.A.S. Nos. 78102-78104) Coomooboolaroo; 182 (C.A.S. 
Nos. 82404-82584 and 83233) Noondoo; 83 (82585-82667) Callandoon; 5 
(C.A.S. Nos. 82668-82672) Keera; 3 (C.A.S. Nos. 82673-82675) Bundy. 

Vomerine teeth greatly reduced and barely perceptible in some indi- 
viduals; tongue oval and free behind; tympanum hidden; an inner and 
outer metacarpal tubercle; two prominent metatarsal tubercles, the inner 
compressed and the outer rounded; subarticular tubercles prominent; toes 
with an indication of fringe; in adpressed limb tibio-tarsal joint reaches to 
l)ack of eye; a prominent parotoid gland. 

The dorsal surface and the back of the thighs are covered with minute 
tubercles. These are scattered and not in longitudinal rows as in T. mar- 
morata. The bellv is smooth and the sides of the bodv finelv granular. 

^ \ 


Fig. 15 (left). Uperolia uKirniorata C.A.S. No. 82403, Blue Mountains, New- 
South Wales. Natural size. 

Fig. 16 (right). Uperolia rucjosa C.A.S. No. 83233, Noondoo, Queensland. Natu- 
ral size. 



[Proc. 4th Skk. 

Color ill life, silvery «ray, groin and back of thigh with large, rich 
orange blotch: undersurfaces whitish, shaded with dark gray; throat very 
dark gray to blackish. Preserved specimens present a very different color 
pattern. It is not until they are in alcohol that the brown to blackish mark- 
ings covering the dorsal surface appear and the orange changes to white. 
The average adult measures 25 mm. from snout to vent. 

This species was found under any type of cover, but mostly under old 
bark, stones, or logs. At Noondoo, where it was abundant, it was not un- 
common on turning over a log to find as many as six or seven individuals 
huddled together in remarkably dry, sandy soil. 

Pseudophryne bibronii Giinther 

(Figure 17) 

This little toad was not met with in numbers, though it is said not to be 
uncommon and is widely spread through southeastern Australia. In the 
Blue Mountains 4 (C.A.S. Nos. 82676-82679) were taken in the Hampton- 
Lithgow area; 3 (C.A.S. Nos. 78112-78114) on the banks of Cox's River; 
and 1 (C.A.S. No. 83239) at Bowenfalls. Six (C.A.S. Nos. 78105-78110) 
are from Coomooboolaroo and 1 (C.A.S. No. 78111) is from Kolonga. 

Vomerine teeth and tympanum absent; tongue oval and free behind; a 
large outer and medium-sized inner metacarpal tubercle; a large inner and 
medium-sized outer metatarsal tubercle; subarticular tubercles prominent; 
second and third toes with a slight indication of fringe; tip of fourth toe 
reaches the snout. 

Dorsal surface covered with minute tubercles; back of thighs granular; 
bellv smooth. 

Fig. 17 (left). Pseudophryne bibronii C.A.S. No. 82678, Blue Mountains, New 
South Wales. Natural size. 

Fig. 18 (right). P.seuelophryne coriacea C.A.S. No. 82680, Ulong, New South 
Wales. Natural size. 


From my notebook T eopy the following- description of the specimens 
taken at Coomooboolaroo. Dorsal surface cinnamon witii })lack reticuhi- 
tions; a canary-yellow ]>atcli in back of the forearm; a transverse canary- 
yellow line above the vent; undersnrfaces marbled with l)lack and wliite. 
In preserved specimens the yellow turns to white and the cinnamon loses 
its brightness and becomes a dull ])rown. 

An average individual measures 26 mm. from snout to vent. 

Five of the specimens from Coomooboolaroo were taken under a stone 
in a drying-up creek ])ed and one under an old piece of wood alongside 
a water trough. 

Pseudophryne coriacea Keferstein 

(Figure IS) 

A single specimen of this species (C.A.S. No. 82680) was taken under 
an old log in the rain forest at l^long. 

Vomerine teeth and tympanum absent; tongue oval and free behind; a 
large outer metacar])al tubercle and a mere indication of an inner one; 
moderate inner metatarsal tubercle and a minute outer one; subarticular 
tubercles prominent. With limb ad])r('ssed, ti]) of the longest toe reaches 
beyond snout. 

The dorsal surface is covci'cd with minute tubercles and the sides and 
back of the thighs are granular. 

In life, the dorsal color of this specimen was a dark orange; sides and 
belly black; undersurface of belly and limbs with irregular wliite markings. 

It measures 25 mm. from snout to vent. 

Notaden bennetti (Uinther 

(Figure 19) 

Nine specimens of this little toad were taken as follows: 2 (C.A.S. Nos. 
82681-82682) from Xoondoo; 6 (C.A.S. Nos. 82683-82688) from Callan- 
doon, and 1 (C.A.S. No. 82689) from Bundy. 

This is an easily distinguished form, the body being short and stout, 
almost globular in sha])e. IMouth very small, legs and toes short, the latter 
very much flattened and heavily fringed; a large outer metacarpal tubercle 
and a smaller inner one; a large elongate metatarsal tubercle present; a 
tubercle at the base of each tinger; tympanum hidden. 

The skin is somewhat leathery in texture. In life, the ground color is a 
light pea-green, covered with black warts forming a cross-like pattern on 
the back, these sometimes being crowned with an orange-colored spot; a 
number of .small, white warts, on the sides and anal region; back of thighs 
covered with small tuliercles. The undersurface is whitish, Ihe thi'oat being 
covered with black spots. 



[Proc. 4tii Skr, 

A specimen from Nooiidoo (No. 8268.1) measures 40 mm. from snout 
to vent. 

These toads are reported to be sometimes seen in countless numbers 
after a rain, but despite the fact that a rain did fall at Callandoon, where 
the largest series were taken, there were no signs of them while hunting 
with a light after dark. Those secured were chopped out of a small log 
half buried in the earth. The Noondoo s])ecimeris were found under logs 
in quite sandy soil. 

On account of the cross-like pattern on the l)(i('k I have heard these 
toads referred to as the "Holy Cross Toad" and "Catholic P^rog." 

Hyla adelaidensis Gray 

(Figure 20) 

This species was met with in only one locality, Nangabrook, West Aus- 
tralia, where 61 specimens (C.A.S. Nos. 78446-78506) were taken. 

A slenderly l)uilt frog with a decidedly acuminate snout; vomerine teeth 
in two rounded clusters between choanae; tongue oval, with free edges and 
nicked behind; tympanum prominent, about two-thirds diameter of the 
eye; subarticular tubercles prominent; a sharp inner metatarsal tubercle; 
fingers lightly webbed and toes fully webbed; in adpressed limb tibio-tarsal 
joint reaches snout. 

Fig. 19 (left). Notaden hennetti C.A.S. No. 82689, Bundy Sheep Station, 20 
miles southeast of Moree, New South Wales. Natural size. 

Fig. 20 (right). Hyla adelaidensis C.A.S. No. 78454. Nangabrook, West Aus- 
tralia. Natural size. 



Skill smooth aliove, oraiuilar on belly and undersurfaee oi" thi«hs; a 
])roniinent dermal fold across eiiest. 

This is a brilliantly marked species, if not the most colorful met with. 
In life, dorsal region greenish gold; a black area from the back of the eye 
to the shoulder, bordered beh)w by a thin line of yellow, which extends 
along side to a point midway between limbs; back of thighs l)lack, heavily 
spotted with yellow; nndersurfaces yellow, throat clouded with gray. Forty 
mm. is average size for an adult. 

All of this series were taken in the still waters of a slow-moving- 
stream flowing through the lieart of the giant gum forest in the vicinity 
of Nangabrook. 

Hyla aurea (Lesson) 

(Figure 21) 

Though not an uncommon Australian frog this species was met with 
only once, at Nangabrook, West Australia, where three specimens (C.A.S. 

Nos. 78507-78509) were taken. 


Fig. 21. Hyla aurea C.A.S. No. 78508, Nangabrook. West Australia. Natural size. 


Vomerine teeth between choanae in two, slij^htly oblique, transverse 
series; tongue oval with free lateral edges and nicked behind; tympanum 
prominent; two thirds the diameter of eye; subarticular tubercles promi- 
nent on fingers, less so on toes; a moderate, wedge-shaped, metatarsal tu- 
bercle; fingers with rudimentary webs; toes fully webbed. In adpressed 
limb tibio-tarsal joint reaches to front of eye. 

The skin is smooth. A very prominent fold from the back of the eye 
over the tympanum to the base of the forearm is present. The sides, belly, 
throat, and the undersurface of the thighs are strongly tubercular in the 
more mature specimens. 

In life, the three Nangabrook specimens did not show the gaudy colors 
accredited to this species, the ground color being gray with a tinge of 
green. A broad dorsal line of light gray and two lateral lines of similar 
color are present. The area between these lines is dark gray, broken up 
into large elongate spots on the sides. 

The undersurfaces are whitish, with traces of dark gray reticulations. 
The largest specimen in the series measures 67 mm. and is probably half 

Two specimens were found in still water ponds alongside a stream and 
one was dug out of a hole in the bank of a stream. 

Hyla aurea ulong'ae Loveridge^ 

A single male (C.A.S. No. 83235, Australian Museum Register No. R^ 
13817) was taken at Ulong, New South Wales. Not conforming with the 
specimens at hand and so as not to add to synonymy it was sent to Mr. 
Arthur Loveridge, at Harvard College, Cambridge, Mass., who had worked 
over the extensive Australian collection at that institution and who deter- 
mined it to be a new subspecies, being characterized by a single series 
of vomerine teeth between the choanae. 

Hyla bicolor (Gray) 
(Figure 22) 

This little hyla, of which 110 specimens were taken, was found in only 
two localities, Kolonga Creek and Ulong. From the former locality 37 
(C.A.S. Nos. 78254-78290) were taken and 73 (C.A.S. Nos. 82690-82762) 
are from the latter. 

Vomerine teeth absent; tongue more or less oval and nicked behind; 
tympanum distinct, two-thirds diameter of eye ; subarticular tubercles mod- 

7. Loveridge, A. New frogs of the genera Cyclorana and Hyla from southeastern Australia. Pro- 
ceedings of the Biological Society of Washington, vol. 63, pp. 131-138. 1950. 



erate, finger rounded with trace of web at base; toes two-thirds webl^ed; in 
the adpressed limb tibio-tarsal joint reaches to tip of snout. 

Skin smooth, with a prominent dermal fold across chest; ])elly and 
undei-surface of the thighs strongly granular, throat slightly so to smooth. 

In life, the color is pea-green with a narrow white line extending from 
a point under the eye to the forelimb; tympanic area blackish; undersur- 
faces whitish. In two individuals in the present series the white line 
extends beyond the forelimb to a point midway between the limbs. 

An individual 25 mm. in length from Ulong is the largest of the series. 

At Ulong these little frogs w'ere taken while sitting on the leaves and 
stems of wild blackberry bushes in open meadows, and at Kolonga Creek 
among vines and potted plants about the homestead. 

Hyla caerulea (Shaw) 

This species is represented by 122 specimens from Queensland and 11 
from New South Wales as follows : from Queensland, 19 (C.A.S. Nos. 78116- 
78134) Coomooboolaroo; 25 (C.A.S. Nos. 78135-78159) Talafa; 3 (C.A.S. 
Nos. 78160-78162) Magenta; 48 (C.A.S. Nos. 78163-78210) Retro; 17 (C. 
A.S. Nos. 78211-78227) Kolonga Creek, and 10 (C.A.S. Nos. 83162-83170 
and 83226) from Noondoo. From New South Wales, 9 (C.A.S. Nos. 83173- 
83179 and 83227-83228) Bundy, and 2 (C.A.S. Nos. 83171-73172) from 

Vomerine teeth in two dome-shaped clusters between and on a level with 
l)osterior edge of choanae; tongue round, with free edges, and very slightly 
nicked behind; tympanum distinct, slightly smaller than diameter of the 

Fig. 22 (left). Hyla bicolor C.A.S. No. 82740, Ulong, New South Wales. Natu- 
ral size. 

Fig. 23 (right). Hyla ewingii alpina C.A.S. No. 83187, Blue Mountains, New 
South Wales. Natural size. 


eye; a moderate-sized outer metatarsal tubercle; subarticular tubercles 
l)romiiient; fingers and toes heavily fringed; fingers one-third webbed; toes 
two-thirds webbed; tibio-tarsal joint in adpressed limb reaches eye; disks 
on fingers and toes very large, width of largest toe disk equaling the diam- 
eter of eye, 6 mm. in an individual 83 mm. in length. 

The skin is smooth, with the top of the head covered by minute pits. 
The throat, belly, and undersurfaces of the thighs are strongly granular. 

In life, the usual color of adults is a uniform light pea-green, though 
occasionally a few minute spots of white may occur on the dorsal and 
lateral regions. The undersurfaces are whitish to light-straw color. 

Juveniles may have a few white markings. Two individuals from Bundy, 
each 36 mm. in length and one from Noondoo, 38 mm. in length, are marked 
as follows: No. 83227 (Bundy) has a few dorsal and lateral white spots; 
thighs very sparsely spotted; white patch on forearm; white line from fore- 
arm along upper jaw to a point under middle of eye; a white line along 
back of forearm. 

No. 83228 (Bundy) has a minute spot on the forearm, is sparsely spotted 
on the sides, and has a short white line along the upper jaw extending to a 
point under the middle of the tympanum. 

No. 83226 (Noondoo) has two or three lateral spots; a white line along 
the back of the forearm; a small spot on the upper forearm; a short line 
in back of the tympanum on the left side and under the tympanum on the 
right side. 

These frogs were usually found about habitations, where they hide un- 
der water tanks, watering troughs, etc., wherever there is a vestige of 
moisture. The entire series of 17 sjiecimens from Kolonga Creek were taken 
out of a fireplace in the living room of the station house. Not being in use 
during the summer the blower was put on tightly and the frogs, bunched 
up on the brick floor, had an undisturbed hideout. Another favorite spot 
was under the water tanks about the pastures. The frogs hid out in the 
daytime under the tanks and around the edges, obtaining sufificient moisture 
from overflow and leaks which kept the grass damp and green. At night 
the entire population of a tank would emerge and would sit along the water 
pipes leading to the troughs, so that just about two visits would be suffi- 
cient to capture the entire population of a tank. 

The remains of a weevil were found in one of the juvenile specimens 
from Bundy. 

Hyla ewingii alpina Fry 

(Figure 23) 

Only two specimens of this frog were taken, C.A.S. No. 83186 from 
Ulong, and C.A.S. No. 83187 from the Hampton-Lithgow area in the Blue 


Mountains. This is a larger frog than //. e. verauxii and has the dorsal 
surface covered with minute tubercles. 

The color pattern is decidedly different, being grayish with an irregular- 
edged dorsal band of brown and a large diagonal brown blotch near the 
groin. The undersurfaces are whitish with grayish throats. Both are males 
with large vocal pouches. 

Both measure 34 mm. from snout to vent. 

Hyla ewingii verauxii Dumeril 

Eeight examples of this tree frog were taken as follows: 2 (C.A.S. Nos. 
78444-78445 from the Cox River, Blue Mountains; 3 (C.A.S. Nos. 83183- 
83185) Hampton-Lithgow area, Blue Mountains; and 3 (C.A.S. Nos. 83180- 
83182) are from Ulong. 

Vomerine teeth either in two rounded clusters or two elongate series 
between ehoanae; tongue oval and nicked behind; tympanum distinct; meta- 
carpal tubercles moderately prominent; a small oval inner metatarsal tu- 
bercle; toes three-fourths webbed, except the inner which is without web; 
fingers with rudimentary webs. In the adpressed limb the tibio-tarsal joint 
reaches the eye. 

The skin is smootli on the dorsal surface and the undersurface of the 
belly. Throat and thighs are granular. 

In life, the color is grayish, with broad lateral stripes of brown. A 
broad black band extends from the tip of the snout, passes through the 
eye and over the tympanum, to the forelimb. This is sometimes bordered 
Ijelow with a narrow white line. The flanks are marked with large oval 
or squarish spots. The undersurfaces are whitish, the throat sometimes 
clouded with dark gray. 

The spotting of the flanks seems to vary somewhat, the frogs from the 
Blue Mountains being heavily marked, while those from Ulong have few 
and very small spots. 

Hyla kinghorni Loveridge'^ 

One example (C.A.S. No. 83234, Australian Museum Register No. R- 
13818) is from Ulong, New South Wales. Being in the same category as a 
preceding species (Hyla aurea ulongae) it also was sent to Mr. Arthur 
Loveridge who determined it as new, being closely related to Hyla pahnata. 
It was named after the well-known herpetologist, ^Ir. J. R. Kinghorn, of 
the Australian Museum. 

8. Loveridge, A. New frogs of the genera Cyclorana and Hyla from southeastern Austraha. Pro- 
ceedings of the Biological Society of Washington, vol. 63, pp. 131-138. 1950. 



[Proc. 4th Skr. 

Hyla latopalmata (Giinther) 

(Figure 24) 

This name is api)lied to 69 specimens as follows: 45 (C.A.S. Nos. 78539- 
78577) Kolonga Creek; 7 (C.A.S. Nos. 78532-78538) Talafa; 6 (C.A.S. Nos. 
78526-78531) Coomooboolaroo, and 11 (C.A.S. Nos. 83151-83161) are from 

These frogs are similar in size and do not appear to differ from //. 
lesueurii in any way except coloration, and this is not absolutely constant. 
The length of the hind limb varies as it docs in //. lesueurii and cannot be 
used as a character. The frogs called //. latopalmata have the doi-sal sur- 
face clouded or marbled with a darker gray than the ground color in 47 
out of 69 cases. 

Though //. latopalmaia has been recognized as distinct from H. lesueurii 
the distinguishing characters given do not seem to warrant it. The chief 
difference given seems to be that the former species is more slender in 
habit, but in comparing a series of both species that are equal in size this 
character is not substantiated. As past writers have been somewhat dubious 
as to its status it may eventually become a synonym of //. lesueurii, or at 
least a subspecies. 

Hyla lesueurii Dumeril and Bibron 
(Figure 25) 

One hundred and ten specimens of this hyla were taken; 16 (C.A.S. 

Fig. 24 (left). Hyla latopalmata C.A.S. No. 78549. Kolonga Creek Cattle Sta- 
tion, 25 miles north of GinGin, Queensland. Natural size. 

Fig. 25 (right). Hyla lesueurii C.A.S. No. 78523, Coomooboolaroo Cattle Sta- 
tion, 15 miles south of Duaringa, Queensland. Natural size. 


Nos. 78510-78525) from Coomooboolaroo, 29 (C.A.S. Nos. 83058-83080) 
from Ulong, and 65 (C.A.S. Nos. 83087-83151) from Hampton-Lithgow 

Vomerine teeth in two elongate series between the ehoanae, occasionally 
joining on median line; tongue rounded and nicked behind; tympanum dis- 
tinct; two-thirds diameter of the eye; subartieular tubercles prominent; a 
small, oval, inner metatarsal tubercle; outer metatarsal tubercle minute or 
absent. Out of 23 examples the outer metatarsal tubercle is absent in six; 
represented by a mere dot in twelve and distinct in five. Fingers without 
webs, and toes two-thirds webbed. In the adpressed limb the tibio-tarsal 
joint reaches to a point between the eye and the snout in one, to the snout 
in twelve, and beyond the snout in ten. In the vinosa variety four reach 
the snout and tw^o slightly beyond. 

A fold of skin passes over the tympanum, curving downward to the 
forearm. The skin is smooth above, the belly and undersurface of the 
thighs being granular. 

The largest specimen in the Ulong sei-ies measures 42 nun. from snout 
to vent and six specimens of the vinosa variety 66, 64, 63, 62, 61, 61, 

In life, the ground color was both grayish and buff. Shown under a 
light at night the buff color was extremely light, but in daylight changed 
to a much darker shade. A heavy black band starts at the tip of the snout, 
passes through the eye and over the tympanum to the forearm. There is 
considerable black marbling in the groin and the back of the thighs. The 
undersurfaces are whitish, with the throat sparsely clouded with gray. In 
the vinosa variety there are large, white-s]>otted, black blotches in the groin 
and the back of the thighs are black with whitish rings, usually with a 
white spot in the center. 

In the daytime these frogs were found under boards and rocks and 
at night sitting on stones in the bed of a running stream. 

Hyla nasuta (Gray) 

(Figure 26) 

Kolonga Creek was the only locality where this s])ecics was found. 
Fifty-six specimens (C.A.S. Nos. 78578-78605 and 78607-78634) were taken. 

Hyla nasuta is a slenderly biult frog with nose strongly acuminate and 
long hind legs. 

The vomerine teetli are in two rounded clusters between the ehoanae. 
In a single specimen, No. 78579, they show a tendency to be elongated and 
in an oblique position. Tongue oval, with free edges, and nicked behind; 
tympanum prominent, about equal to diameter of eye; an elongate inner 
metatai-sal tubercle: a small rounded outer metatarsal tubei'cle; finoers free 



of web; toes two-thirds webbed; in adpressed limb tibio-tarsal joint reaches 
well bej'Ond snout. 

Skin smooth, with more or less obsolete longitudinal folds; undersurfaee 
of thighs and belly lightly granular; throat smooth; a prominent fold 
across chest. 

In life, the gi-ound color is grayish or brownish with three rows of 
black, elongate, oval spots down the back; two thin light dorsal stripes 
separate the middle from the lateral spots; loreal region black; a black line, 
bordered above and below by a narrower white one on the back of the thighs; 
undersurfaces whitish. The males have the throats suffused with dark gray. 

The entire series were taken in Kolonga Creek, a wide, shallow, and slow- 
moving stream, running through the station property. The frogs were 
found floating amongst the patches of grass close to the bank. The average 
length of an adult in the present series is 46 mm. 

Hyla parvidens Peters 

This name has been applied, with some hesitancy, to two immature speci- 
mens (C.A.S. Nos. 78442-78443) taken on the banks of the Cox River, 
New South Wales. Neither one shows the adult type of coloration, but 
some of the characteristics of the species are present; the clear coloring of 
the back of the tliighs, the small tympanum, and the more or less obsolete 
vomerine teeth. In the adpressed limb the tibio-tarsal joint reaches to the 

Fig. 26. Ilyla nasuta C.A.S. No. 78604, Kolonga Creek Cattle Station, 25 miles 
north of GinGin, Queensland. Natural size. 


eye. The larger one measures 22 mm. from snout to vent and is a slight 
reddish-brown, with darker marbling or spotting on the dorsal surface. 
The undersurfaces are whitish. The belly and anal region are tubercular. 

Hyla peronii (Tsehudi) 
(Figure 27) 

Fifty-three specimens of this strikingly marked hyla were taken, 18 
(C.A.S.Nos. 78228-78245) Coomooboolaroo; 3 (C.A.S. Nos. 78246-78248) 
Talafa; 5 (C.A.S. Nos. 78249-78253) Kolonga Creek; 18 (C.A.S. Nos. 
83206-83220 and 83231-83232) Noondoo; 1 (C.A.S. No. 83221) Callandoon; 
3 (C.A.S. Nos. 83237-83238) from Ulong; and 5 (C.A.S. Nos. 83222-83225 
and 83229) from Bundy. 

Vomerine teeth in two transverse series between the choanae; tongue 
rounded, with free edges and slightly nicked behind; tympanum distinct, 
about two-thirds diameter of eye; a small outer metatarsal tubercle; sub- 
articular tubercles prominent; fingers and toes heavily fringed; fingers two- 
thirds webbed and toes fully webbed; disks on fingers and toes large, the 
largest equaling diameter of tympanum; in adpressed limb the tibio-tarsal 
joint reaches to back of eye. 

The skin is somewhat rugose. A prominent fold extends from the back 
of the eye and over the tymi)anum to the forearm and a pronounced der- 
mal fold across the chest is present. The throat, belly, and the undersurface 
of the thiglis arc stronglv tubercular. 

Fig. 27. Hyla peronii C.A.S. No. 78246, Talafa Sheep Station, 20 miles south 
of Emerald, Queensland. Natural size. 


This species is a strikingly marked frog. In life, the color is dark gray 
to slate, the dorsal surfaces being clouded or marbled with black. The area 
back of the forearm, the sides, groin, and back of the thighs are marbled 
or spotted with black and a rich canary-yellow, the latter color completely 
disappearing in alcoholic specimens. A light gray, and somewhat obsolete 
dorsal stripe may be present. The undersurfaces are whitish to straw color. 

The largest specimen taken is from Ulong and measures 65 mm. from 
snout to vent. This one, however, is apparently a large specimen, the aver- 
age length in the series being around 43 mm. 

At Coomooboolaroo these frogs were found under the bark of standing 
trees and at Ulong clinging to reeds in small ponds along the edge of a 
creek running through an open meadow. The series from the Hampton- 
Lithgow area were all found under small stones in the dry portions of 
a river bed. 

Hyla rubella Gray 

(Figure 28) 

This was by far the most abundant hyla met with, 447 being taken as 
follows: 138 ("c.A.S. Nos. 78291-78428) Coomooboolaroo; 5 (C.A.S. Nas. 
78429-78433) Talafa; 6 (C.A.S. Nos. 78434-78439) Kolonga Creek; 2 (C. 
A.S. Nos. 78440-78441) Retro; 153 (C.A.S. Nos. 83230, 82780-82916 and 
83042-83057) Noondoo; 7 (C.A.S. Nos. 82917-82923) Callandoon; 136 (C. 
A.S. Nos. 83188-83205 and 82924-83041 ) Bundy. 

Vomerine teeth in two small clusters between and behind choanae; 
tongue oval with free edges; tympanum distinct, about two-thirds diameter 
of eye; subarticular tubercles prominent; a small, elongate inner meta- 

Fig. 28. Hyla rubella C.A.S. No. 78432. Talafa Sheep Station. 2U miles south 
ot Emerald, Queensland. Natural size. 


tarsal tul)oi'('le; fiugei's free of weh and toes two-thirds webbed. Loveridge*^ 
states that speeiinens preserved in formalin show the fifth toe fully weighed 
to the base of the disk, but that this condition is not found in alcoholics. 
The present series is preserved in alcohol and this character is occasionally 
met with. The disks on the fingers and toes vary considerably in size. In 
the adpressed limb the tibio-tarsal joint reaches the back of the tympanum. 

The skin is smooth above with the sides, throat, belly, and undersurface 
of thighs, granular. A prominent dermal fold crosses the chest. 

In life, the ground color is light or dark cinnamon covered with minute 
spots of black. A prominent blackish band extends from the tiji of snout, 
through eye, to back of shoulder and occasionally to groin; undersurfaces 
whitish or yellowish; throat of males suffused with black. Adults average 
35 mm. in length. 

These hylas sought almost any type of cover and were found in water 
troughs, holes in fence i)osts. and under debris and bark of fallen or 
standing trees. As many as a dozen, or even 20 were found huddled to- 
gether under a single slab of bark. 

Under a light at night the skin is a rich reddish-]unk. 


Andersson. L. G. 

1913. Results of Dr. E. Mjoberg's Swedish Scientific Expeditions to Australia 
1910-1913. Amphibia Kungl. Svenska Vetenskapsakademiens Hand- 
lingar, Band 52. N:0 4. pp. 1-26. pi. 1. 

1916. Results of Dr. E. M.loberg's Swedish Scientific Expeditions to Australia 
1910-1913. Amphibia Kungl. Svenska Vetenskapsakademiens Hand- 
lingar. Band 52, No. 9. pp. 1-20. pi. 1. 


1882. Catalogue of the Batrachia Salientia s. Ecaudata in the Collection of the 

British Museum. London, pp. 1-502, pis. I-XXX. 
1918. On the Papuan, Melanesian, and North Australian species of the genus 

Rana. Annals and Magazine of Natural History. (9) vol. 1, pp. 236-242. 

Fletcher, J. J. 

1890-1898. Contributions to a moi'e exact knowledge of the geographical dis- 
tribution of Australian Batrachia. Proceedings of the Linnean Society 
of New South Wales, vol. 5, pp. 667-676; vol. 6, pp. 263-264; voL 7, pp. 
7-19; vol. 8, pp. 524-533; vol. 12, pp. 660-684. 

Fry, D. B. 

1912. Description of Austrochaperina, a new genus of Engystomatidae from 
North Australia. Records of the Australian Museum, vol. 9, pp. 87-106, 

9. Bulletin of the Museum of Comparative Zoology, vol. LXXVIII, No. 1, p. 42, 1935. 



Fry. D. B. (Cont.) 

1913. On a Varanus and a Frog from Burnett River, Queensland, and a revi- 
sion of the variations in Limno(Ujn<istes clorsalis Gray. Records of 
the Australian Museum. Stjdiiey, vol. 10, pp. 17-34. 

1913. A re-examination of Macleay's New Guinea and Queensland frog types. 

Memoirs of tJie Queensland Museum, vol. 2, pp. 46-50. 

1914. On a collection of Reptiles and Batracliians from Western Australia. 

Records of the Western Australian Museum, vol. 1, pp. 174-210, pis. 

1915. Herpetological notes. Proceedings of the Royal Society of Queensland, 

vol. 27, pp. 60-95, pis. 1-4. 

Garman. S. 

1901. Reptiles and Batracliians from Australia. Bulletin of the Bluseum of 
Comparative Zoology, vol. 39. pp. 1-14. 

Glauert. L. 

1929. Contributions to the Fauna of Rottnest Island. Journal of the Royal 
Society of Western Australia, vol. 15, pp. 43-45. 

Harrison, L. 

1927. Notes on some Western Australian Frogs, with descriptions of new spe- 
cies. Records of the Australian Museum, vol. 15, pp. 277-287. figs. 1-5. 

Kingiiorn, J. R. 

1924. Reptiles and Batrachians from South and South-West Australia. Records 
of the Australian Museum, vol. 14, No. 3, pp. 163-183. 

1932. Herpetological Notes, No. 4. Records of the Australian Museum, vol. 18, 

No. 7, pp. 355-363. 
1945. The Simpson Desert Expedition, 1939 Scientific Reports: No. 3, Biology- 
Reptiles and Batrachians Transactions of the Royal Society of South 
Autralia. vol. 69, part 1, pp. 3-4. 


1933. Four new Crinine Frogs from Australia. Occasional Papers of the Boston 

Society of Natural History, vol. 8, pp. 55-60. 

1933. A New Genus and three new species of Crinine Frogs from Australia. 

Occasional Papers of the Boston Society of Natural History, vol. 8, 
pp.^ 89-94. 

1934. Tasmanian Amphibia in the Museum of Comparative Zoology, Cambridge 

Massachusetts, Papers and Proceedings of the Rcryal Society of Tas- 
mania. 1933, pp. 56-64. 

1935. Australian Amphibia in the Museum of Comparative Zoology, Cambridge 

Massachusetts, Bulletin of the Museum of Comparative Zoology, vol 
78, No. 1, pp. 3-60, plate. 

1936. On some reptiles and amphibians from the central region of Australia 

Transactions of the Royal Society of Soutli Australia, vol. 62, (2), pp 
1950. New Frogs of the Genera Cycloraiia and Hyla from Southeastern Aus 
tralia. Proceedings of the Biological Society of Washington, vol. 63 
pp. 131-138. 


Lucas, A. H. S. 

1892. Notes on the Distribution of Victorian Batrachians. Proceedings of the 

Royal Society of Victoria, vol. 4, pp. 59-64. 
1901. Two new species of Frogs from Victoria. Proceedings of the Royal So- 
ciety of Victoria, (2), vol. 13, pp. 175-178. 

Lucas and Li: Soukf 

1909. The Animals of Australia. Amphibia, pp. 259-297. 

Mertexs, Robkrt 

1930. Die Amphibian und Reptilien der Inseln Bali, Lombok, Sumbawa und 

Flores. Abhandlun.gen Senckenbergischen Naturforschende Gesell- 
schaft, Frankfort-am-Main, vol. 42, pp. 117-344, pis. 1-8. 

Nip:ui:n, Fk. 

1923. Das Tierreich. Vol. 46, Amphibia Anura I, pp. 1-584, text figs. 
1926. Das Tierreich. Vol. 49, Amphil;ia Anura II, pp. 1-110, text figs. 


1931. The Biology of the Amphibia. New York, pp. 1 577. text figs. 

OiiUJsY, J. D. 

1907 (1906). A new Tree Frog from Brisbane. Proceedings of the Royal So- 
ciety of Queensland, vol. 20. pp. 31-32. 

Parker. H. W. 

1926. New Reptiles and a New Frog from Queensland. Annals and Magazine 
of Natural History, (9), vol. 17, pp. 665-670, figs. 1-3. 

1934. A Monograph of the Frogs of the family Microhylidae. British Museum, 
London, pp. 1-208, 67 text figures. 

1940. The Australian Frogs of the Family Leptodactylidae. Nointates Zoologi- 
cae, vol. 42. Part 1, pp. 1-106. pi. 1. 20 text figures. 

Peters, W. C. H. 

1871. tJber einige Arten der herpetologischen Sammlung des Berliner Zoologis- 
chen Museums. Monatsberichte Akademie Wissenshaft, Berlin, pp. 

Procter. J. B. 

1924. Unrecorded characters seen in living snakes and description of a new 

tree frog. Proceedings of the Zoological Society of London, pp. 1125- 
1129, pis. 1-3. 

Spex(er, Baldwin 

1896. Amphibia. Report on the work of the Horn Scientific Expedition to Cen- 
tral Australia, part 2. pp. 152-175, pis. 13-15. 

Stirling. E. C, and A. Zietz 

1893. On the Reptiles and Batrachians collected by the Elder Exploring Ex- 

pedition. Transactions of the Royal Society of South Australia, vol. 
16, pp. 159-176, pis. 6-7. 


Van Kampen, P. N. 

1923. The Amphibia of the Indo-Australian Archipelago. Leiden, pp. 1-304, 
29 text figures. 

Waite, E. R. 

1929. The Reptiles and Amphibians of South Australia. British Science Guild. 
Adelaide, pp. 1-270, 192 text figures. 




Fourth Series 
Vol. XXVTII, No. 9, pp. 393-404, 5 tables February 17, 1956 




Associate Professor of Agricultural Economics and Associate Economist in the 
Experiment Station and on the Giannini Foundation, 

University of California, Berkeley » 

Let this discussion commence from a perspective of the rate and magni- 
tude of western population growth in the recent past. Tables I-V sum- 
marize some of the most relevant data. These data are the basis for several 
generalizations on western population growth which are essential back- 
ground for the further considerations implied in the title of this paper. 
After stating these generalizations, the paper j^roceeds to some of the inter- 
relations between popidation growth and economic expansion in the past 
and concludes with comments on the prospects of the future. 

I. Generalizations from the Demographic Data 

1. Although the populations of the eleven western states and of the 
Mountain and Pacific regions liave grown rapidly, and at rates considerably 
in excess of those of tlie nation, the state-to-state differences within the West 
are very large, ranging from many times the national average to about 
half the national average. 

2. In the first four years of this decade, a rapid rate of western growth 
has contimied, but in relation to the national rate, it has diminished. 
Whereas from 1940-1950, the West grew 2.8 times as fast as the nation at 
large, in the past four years this ratio has dropped to 2.2. 

3. Particularly notable is the suggestion in the data that the areas of 
rapid growth within the region may have shifted in the present decade. 
Growth rates in the Pacific Kegion — in Washington and Oregon particu- 
larly — have declined sharply. Growth ratios of the ^Mountain Region have 
moved upward, though not uniformly and not sharply, except in Arizona 
and Nevada. 

1. Prepared for Symposium on Natural Resources of the West. Joint Session of A.A.A.S. Section F 
and the Western Society of Naturalists, Berkeley, Calif., December 28, 1954. 

[ 393 ] 


4. Even though westerners are not noted for infertility, it is true that 
our outstanding growth rates are attained mainly by large net human im- 
port balances. The ratios to be found in table III show the role of net 
migration. Note the changes of the last four years as compared with the 
preceding decade. Whereas Washington and Oregon gained 650,000 by net 
migration in 1940-1950, these states gained only 26,000 in the first three 
years of this decade. In contrast, the three-year gains in New Mexico and 
Arizona were not far behind the entire preceding decade, and the three- 
year gain in Nevada was already ahead of it. 

5. Regional and state population growth in the recent past and in the 
prospective future is compounded out of the forces that influence national 
fertility and those that impel internal migration. Death rates, though de- 
clining gradually, are substantially invariant from year to year and the 
role of immigration has become minimal, thus leaving fertility as the almost 
exclusive determinant of national population growth. What is noteworthy 
is that even after two important erstwhile population variants have been 
nullified, the future population of the nation still remains uncertain by rea- 
son of unstal)le fertility. And for states and regions, particularly those of 
the West, future populations are much more highly uncertain because of 
the complexity of forces that influence migration. 

6. Our present high level fertility is now being accepted as an unex- 
pected reversal of a long-time downward trend. As the data in table IV 
show, earlier marriages and the bearing of children earlier in the marriage 
are responsible for the marked upturn in total births and in the crude 
birth rate since 1940. But the evidence does not yet demonstrate an upturn 
in the size of completed families. Women 20-34 years of age in 1952 had 
borne significantly more children than had their counterpart generation of 
1940. If the fertility level of this generation is sustained through the nor- 
mal age range of child bearing, this will clearly demonstrate at least a 
temporary upward revision in the concept of desirable family size, other- 
wise only that childbearing has been shifted to an earlier age. For evidence 
that is at all satisfactory on this point, we shall have to be patient until 
the end of this decade. 

7. Within the limits of plausible assumptions as to fertility, the future 
national population can be projected, and with some degree of intellectual 
comfort to the projector. As for regional and state populations in the West, 
similar projections are ventured occasionally, but usually with obvious dis- 
comfort, for the forces and occurrences that impel western migration are 
not readily predictable. The largest movements to California, for example, 
have been associated with times of unusual prosperity and with the two 
world wars. Unless one can predict wars and prosperity, he would likely 
not do well at predicting migration to California, and quite possibly not 
even then. 










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Rates of Population Growth in Wefitern States and in Regions 

as Ratios to National Rates of Population Groxcth 

19.',0 to 19.J0 and 1950 to 195.','^ 

Area 1940 to 1950 1950 to 1954 

Washington 2^48 ^7 

Oregon 2.84 1.21 

California 3.73 2.72 






New Mexico 







North Central 


United States 1.00 1.00 

;i. Based on data in Table I. 

II. Population Growth and Economic Expansion 

For rapid population growth based mainly on migration to be sustained, 
there must be proportionate economic expansion. Otherwise, per-capita 
incomes will decline and thereby, sooner or later, bring the migration to a 
halt. If it were true that people lived directly on the yield of natural re- 
sources in the area of resettlement, then only sparsely occupied frontier 
communities could absorb additional population and still avoid tlie adversity 
of declining incomes. In so uncomplicated a world, impacts of rapid popula- 
tion growth on natural resources would at some stage become all too obvious. 
But modern economies, particularly those of the western states, are based 
upon technologies and institutions that obscure the relations between people 
and basic natural resources; they have (or at least seem to have) great flex- 
ibility and versatility. For the welfare of people, these are indeed fortunate 
attributes; for the analyst who seeks full and reliable knowledge of man in 
relation to his physical environment, these attril)utes contain the seeds of 
frustration and confusion. 

During the three decades for which statistics are available, most of the 
western states have enjoyed per-capita incomes above the national average. 































Large in-migration has had little, if any, adverse eft'eet. In California where 
the volume of in-niigration has been so extraordinary, the income differential 
has apparently narrowed somewhat but still stands at 20-25 per cent above 
the national average. For a state that was already well settled and popu- 
lated to grow by one half in a decade and still maintain most of its premium 
income position, as was true of California 1940-1950, is something of a feat. 
For the booster, it is the happy motivation for newer and more lauditory 
slogans; for the scholar, it poses the challenging obligation to find out liow 
it was accomplished. If we can know something of the economic expansion 
that is concurrent with rapid population growth, we may tlierefrom also 
derive some assistance in knowing its impact on natural resources. 

Inasmuch as different states and even areas within states respond differ- 
ently in the expansionary process, it seems unwise to undertake this inquiry 
for the western region as a single aggregate. Furthermore, neither the time 
nor the required analyses are available to permit this inquiry being carried 
into all states of the West. Accordingly, for this section of my topic, I shall 


Net Civilian Migration and Ratios of Net Migration to 
Natural Incre(tsr—!!i',(i-]<i.-,i) and 1950-1953 

Net Civilian Migration Ratios of Migration to 

(Thousands of persons) Natural Increase 

ApriM,1940to April1,1S50to April1,1940to Aprill, 1950to 

Area ApriM, 1950 July 1, 1953 . April 1, 1950 July 1,1953 

Washington 354 3 1.40 .03 

Oregon 300 23 2.05 .29 

California 2,588 977 2.51 1.85 

Pacific 3,243 1,004 2.27 1.39 

Montana — 34 — 4 — .47 — .12 

Idaho — 20 — 18 —.22 —.47 

Wyoming — 7 — 2 — .17 .11 

Colorado 39 11 .24 .14 

New Me.xico 12 8 .09 .13 

Arizona 138 122 1.20 2.07 

Utah 14 — 2 .11 —.34 

Nevada 33 36 2.06 4.50 

Mountain 174 150 .23 .42 

West 3,417 1,154 1.56 1.07 

South —2,017 — 522 — .27 — .18 

North Central 50 158 .01 .07 

Northeast 615 202 .19 .14 

United States 1,976 992 .11 .13 

Source: Basic data from U. S. Bureau of the Census, Current Population Reports, Series P-25, Nos. 72 
and 97. 



[Pkoc. 4th Ser. 

consider only California. A very recent book by Dr. Margaret S. Gordon, 
Employment Expansion and Population Growth, the California Experience 
1900-1950, serves excellently as a basic reference. Additionally, I have 
drawn upon a chapter by myself on the growth and development of Califor- 
nia agriculture which is to appear in a forthcoming book on the California 
population, sponsored by the Haynes Foundation. 

California migration and economic growth are a cobweb of cause and 
effect relations. Initial movements were provoked by pre-existing economic 
attractions. For sure, the initiating influence to the forty-niners was a pre- 
existing attraction. But before very long, people began to realize that it was 
just as profitable and far more comfortal)le to mine the miners. And so it 
has been through the decades. Epochs of migration have been compounded 


Fertility of the National Pojiulation 

A. Births and Birth Rates 

B. Marriage Rate 

Per Cent of Ferrales 

Pet. Increase 

No. of Births 


Birth Rate 

Age Group 
14-19 years 



in Rate 












20-24 years 







25-29 years 







30-34 years 







35-44 years 







45-54 years 







55-64 years 







65-74 years 







75 and over 
Total, 14 ~ 




and over 




C. Number Children Ever Born D. Number Children Ever Born 

per 1,000 Women per 1,000 Women Ever Married 

Per Cent Per Cent 

Increase Inrrease 

Age Group 1952 1940 1940-1952 1952 1940 1940-1952 

15-19 years 98 68 44.1 572 567 0.9 

20-24 years 836 513 63.0 1,187 969 22.5 

25-29 years 1,527 1,090 40.1 1,742 1,408 23.7 

30-34 years 1,943 1,613 20.5 2,130 1,888 12.8 

35-39 years 2,112 2,095 0.8 2,293 2,357 —2.7 

40-44 years 2,169 2,478 —12.5 2,346 2,740 —14.4 

45-49 years 2,172 2,735 —20.6 2,352 2,993 —21.4 

50 years and over 2,707 3,014 —10.2 2,937 3.315 —11.4 

Source : U. S. Bureau of the Census, Current Population Reports, Series P-2S, No. 78 ; Series P-20, Nos. 
46 and 50. 


out of the initiating occurrences and out of the self-generationg forces in- 
herent in an expanding population. In general, it may be said that the 
initiating developments have tended to be more intimately related to natural 
resources than have the subsequent service industries that were based on the 
needs of the population. But even this point is clearer in theory than in 
fact; if it were not ultimately so, then a migrating population could settle 
anywhere and prosper merely by rendering services unto itself. Yet the 
resources serving to initiate California's migrations extend considerably 
outside the usual inventory. Shipbuilding and aircraft manufacture are 
two of the largest increments of economic activity the state has ever experi- 
enced. But the resource base of these industries is not readily identified. 
For ships, ports on the Pacific were undeniably useful but the potency of the 
state's politicians was perhaps just as fruitful a resource. For aircraft manu- 
facture, temperature, humidity, the spacing of rains, and the once-blue skies 
of the California southland were probably the outstanding attributes. Thus, 
many characteristics of climate and topography, together with whatever 
spell of magic it was that created Hollywood, must be listed along with gold, 
oil, forests, and fertile land in California's inventory of resources. 

Only in the very first years of the American epoch did agriculture and 
the extractive industries occuj^y the major portion of the population. 
Through the years, employment in these industries has not kept pace with 
population growth. In consequence, their relative position has consistently 
receded until they now have less than one tenth of all gainfully employed 
workers. Meanwhile, such industries as construction, manufacturing, trade, 
and public administration have expanded at least in proportion with popu- 
lation growth. By 1950, construction occupied about as many workers as 
did agriculture; distribution and service in total were approximately nine 
times the size of agriculture. 


The National Population to 1915 — Possible Maximum and Minimum Projections 

A Series* D Seriesf 

Percentage Percentage 

Number Increase Number Increase 

Year (Millions) Over 1950 (Millions) Over 1950 

1955 164.8 8.6 164.4 8.4 

1960 177.4 17.0 173.8 14.6 

1965 189.9 25.2 180.9 19.3 

1970 204.2 34.6 189.1 24.7 

1975 221.0 45.7 198.6 31.0 

* Assumes age-specific fertility rates of 1950-1953 will continue through to 1975. 

t Assumes age-specific fertility rates will decline linearly from 1950-1953 levels to 1940 levels by 1960 

and thereafter remain at approximately the 1940 level until 1975. 
Source: U. S. Bureau of the Census, Current Population Reports, Series P-25, No. 78. 


Those whose thinking runs in fundamentalist terms tend to assume tliat 
rapid growtli to a 12i/4-million population must have brought severe impacts 
upon California's agriculture and its farming resources. However, my inter- 
pretation of the evidence leads to a different conclusion. This conclusion 
derives basically from the fact that in volume and composition of output, 
California's agriculture has always been primarily responsive to national 
and international markets rather than to state or western markets. The 
majority of the state's farming resources have always been used in produc- 
ing export specialties. With respect to the use of these resources, growth 
in the size and buying power of the national population has been of major 
importance, but the proportion of the national population living in Cali- 
fornia or in the West has not been of much immediate consequence. With 
the growing of the state population, some of the commodities have shifted 
from the surplus to the deficit category; this is particularly true of cereals 
and livestock products. Although profitability in these commodities was en- 
hanced by larger local markets, it still was not sufficient, generally speaking, 
to induce resources away from the export specialties. Consequently, the state 
has been and remains one of agricultural surpluses and deficits, with no 
apparent strong influences toward self-sufficiency. 

It is therefore doubtful that the agriculture of California would have 
been much different from what it is had the state population remained at a 
fraction of its present magnitude, providing that national and international 
markets had been as they have. The relevant point of this speculation is 
that it indicates that the impact of population on agricultural resources, in 
an economy of well-developed exchange and transport facilities, is more a 
matter of the national than of state or regional populations. 

It is perhaps worth noting the extent to which o]^portunities in agricul- 
ture have been an initiating influence in attracting migrants to California. 
The historical record of the first two decades of the American regime, 1850- 
1870, is well marked with expression of lament that the large land holdings 
were not being divided ra])idly into family farms. Immediately following 
the completion of the transcontinental railroad in 1869, several immigration 
societies were organized for the principal purpose of stimulating the move- 
ment of small farm o]^erators to California. The campaigns that were 
launched in 1870s and 1880s lacked not in color or versatility of approach. 
Though they may have had a hand in stimulating general migration, they 
produced little if any acceleration in the growth of family farming. By 1900, 
the theme of opportunities for family farmers was largely abandoned and 
replaced by equally unsuccessful appeals to hired farm hands. The combina- 
tion of these two failures left a structure of agriculture that was heavily 
dependent on recurrent infusions of labor from foreign or external sources. 
Chinese, Japanese, Filipinos, East Indians, Mexicans, and the eiusodic Okies 
and Arkies have all had tlieir roles upon tliis stage. As each infusion lost 


itself into tlie total mix of the state's economy, a new one has been sought. 
In these indirect and somewhat obscured ways, the natural resources of 
agriculture have perhaps had a greater impact on California's population 
than has its population upon agricultural resources. 

Viewing the state in terms of its aggregate economic growth, or in terms 
of the development of its principal industries, it is difficult to identify 
intimate and unobscured relationships to natural resources. Admittedly, 
within a narrower focus, one can find ghost towns in the mining areas, oil 
wells that have ceased to flow, forests that no longer hold saw timber, and 
farm lands for which there is not sufficient irrigation water. Certainly 
these are impacts — and ostensibly devastating impacts — of population upon 
natural resources. But whether they are impacts inherently associated with 
population growth, either state, regional, or national, is another matter. 
Generally speaking, it seems to me to be more a matter of policy and atti- 
tude than of people ^^er se. Certainly, the huge mounds of gravel and 
boulders left in the wake of gold dredgers were not induced by the needs 
of a growing population but by a most dubious national gold-price policy. 
This, I admit, is not a particularly instructive example, for gold has very 
little usefulness, whether diffused in the earth's crust or peacefully resting 
in the air-conditioned and well-guarded vaults of Fort Knox. But on the 
other hand, the burden to taxpayers of acquiring and guarding the gold, 
and the havoc wrought upon streams and meadows in digging it out, are 
not only clear net losses to society, but also, unfortunately, crushing testi- 
mony of man's propensity to concede his resources to political chicanery 
just as readily as to fundamental needs. Looking at more useful resources, 
such as timber stands and oil deposits, where the issues are vastly more 
complex by reason of the conflict between present and future use, one still 
cannot avoid the conclusion that the manner of use derives less from in- 
herent needs of a growing population than of the policies and attitudes 
with which we have approached the resource. Certainly, this is true for the 
renewable resources. As for the nonrenewable deposits of coal, oil, and 
minerals, there may be nothing morally wrong in removing in a few short 
years the accumulations of the geologic ages, provided that the contempo- 
rary society is fully conscious of the risk taken that, as depletion occurs, 
fully satisfactory substitutes may not be forthcoming. In any event, there 
is a considerable range of alternatives which govern the rate of use, quite 
apart from the size of the population. 

III. Appraisal and Prospect 

What can be said in appraisal of economic expansion in the "West and 
its relation to natural resources? Truthfully, this is a point that I find 
highly confounding. As an economic accomplishment, it seems to demon- 
strate (or to redemonstrate) that the greatest natural resource known to 


man is his own resourcefulness. Evidence of this is found in two directions : 
first, in the imaginative ability to perceive resource values in obscure forms 
and combinations; second, in developing so geographically remote an area 
in close functional relationship with the national economy. 

Yet in terms of other values, this expansion may not be seen as a thing 
of beauty or of high ethical order. The western Indian fared no better 
than did his counterpart on the Atlantic 150 years before. Native plants 
and animals have fallen before the rapacious hand of the occupier. Some 
people believe with Thoreau that "This curious world which we inliabit is 
more wonderful than it is convenient ; more beautiful than it is useful ; it is 
more to be admired than it is to be used." For them the expanding economy 
of the West must be diabolical indeed. Dry wells, polluted streams, eroded 
hillsides, and smog may be geologically superficial but nonetheless persua- 
sive evidence that man's engineering abilities were given to him more 
abundantly than the wisdom to foresee their consequences, or the compas- 
sion willingly to abate their nuisances. 

Under our system of living, people and their commodities and services 
are highly mobile. Consequently, in any general consideration of people 
and resources, no state or region can be a self-contained entity; rather, it is 
better to think at least in national terms, and even more ideally, in inter- 
national terms. 

Nationally, these are days of growing conscience and concern about re- 
sources. The constituting of the President's Materials Policy Commission in 
1951 and its comprehensive reports were a major stride in giving effect to 
this concern. A second major stride was the organizing of the Ford Founda- 
tion's Resources for the Future, Inc., which sponsored the Mid-Century 
Conference on Resources for the Future last December (1953). To William 
Vogt and Fairfield Osborn these manifestations of concern must have been 

The report of the President's Commission and of the Mid-Century Con- 
ference, together with other writings on contemporary problems of popula- 
tion and resources, strongly suggest that a stage has been reached at which 
broadscale soul-searching is called for. Perhaps this conclusion is no more 
than the myopic egocentricity that has led individuals of preceding genera- 
tions to the conviction that their stage of development was somehow pecul- 
iarly epochal. Even after acknowledging and discounting this possibility, 
however, I am still convinced that, to rely on a badly eroded phrase, " a new 
turn of events is at hand." And it is not impossible, perhaps, that the years 
immediately ahead will have significance at least equal to the counterpart 
years of the Industrial Revolution. 

Basically, the problem at hand is our unbalanced natural resource 
budget. Our standard of high living requires prodigious quantities of 
materials and energy. Furthermore, sustaining the ever-improving mate- 


rial progress to which we are so deeply committed and this, for an ex- 
panding population, means rapidly accelerating demands for energy and 
materials. Except as the finiteness of the resource base is lifted or postponed 
by research or technology, the prospective relationship between resource 
demand and supply is more that of collision than of equilibrium. 

These recent reports impress one of the exciting possibilities and poten- 
tialities for research and technology. It is not at all difficult to be persuaded 
that there will be substitutes for resources as we now know and use them, 
and also substitutes for the substitutes. But this has to be largely a matter 
of faith. And even those of great faith realize we are at the point where, 
more than ever before, human abilities for research and technological de- 
velopment have to be treated as a scarce resource to be developed and 
conserved accordingly. 

Significantly, both the President's Commission and the Mid-Century 
Conference identified research and technology as an endogenous force cap- 
able of being shaped and manipulated by deliberate policies. The partici- 
pants in Section VII of the Mid-Century Conference, in discussing resources 
research, dealt with two mattei's that are likely to become increasingly 
familiar to scientists in all fields — namely, organization and communication. 
These are problems that are exceedingly intricate and delicate. I mention 
them now, not in the hope of being informative with respect to their solu- 
tions, but only to say that, notwithstanding their acknowledged existence, 
I doubt that either those having an abiding faith in science and technology, 
or scientists and technologists themselves, fully appreciate the compelling 
importance of these problems or the imagination and forbearance it will 
take to solve them. 

In closing, let me add another and important dimension to this last 
point. This is most readily done by quoting a paragraph of wisdom from 
one of the participants of the Mid-Century Conference — Stanley A. Cain 
of the University of Michigan : 

Because man is the creator of culture and is at the same time its creature, the 
problems of resource use are only partially scientific and technological. They are 
also — perhaps largely — social, economic, and political problems. We cannot definitely 
realize particular research problems in the absence of the situation in which these 
problems are created and the answers that are obtained will be applied. Only a 
deeper and wider understanding of the nature of human ecological problems can 
bring about the necessary social and financial support for their prosecution and the 
needed wedding of physical and social fields of research. Here I find the greatest 
underdeveloped area in the national research program. 

Without concurrent and integrated progress in this area of greatest un- 
derdevelopment, out of which man may gain a better understanding of 
himself in terms of his universe, it seems neither irresponsible nor sophistic 
to "view with alarm." 


We presently face two potential juggernauts — population and tech- 
nology. Conceivably, our technological future could go either to the extreme 
of being too successful or to that of niggardliness. At the first extreme, 
there is the danger that man's values may derive too largely from the be- 
guilements of the advertiser; at the second extreme, they may be instilled 
by the despot thriving upon frustration and despair. At either extreme, 
or even at an intermediate course of development; confident living will re- 
quire the deep-seated conservatism and humility with respect to himself that 
man will have only if he realizes and understands he is the creature as 
well as the creator of his culture. The conservatism that derives from this 
knowledge and understanding should be the way to avoid the pitfalls of 
endeavoring a too-thorough mastery of the universe. There is no compulsion 
for man, either by reason of his numbers or by reason of grasping for 
material comforts, to undertake to consume everything the planet has to 
offer and, in the meanwhile devote his outstanding capabilities to doing 
it faster and more thoroughly. 




Fourth Series 
Vol. XXVIII, No. 10, pp. 405-414 February 17, 1956 



Chairman and Professor, Department of Irrigation and Soils, 
University of California, Los Angeles 

The West, as herein defined, is represented by the 17 western states and 
includes all or parts of the following major drainage basins : North Pacific 
Coast (I), South Pacific Coast (II), Great Basin (III), Gulf of California 
(IV), Gulf of Mexico, exclusive of Mississippi River (V), and parts of the 
Western Mississippi River (VI), and the Hudson Bay drainage basins 

The area is large (some 1,817,000 square miles) and possesses many 
sharp contrasts : contrasts in the amount, distribution, and character of pre- 
cipitation; in the nature of its streams, from large rivers of sustained flows 
to arroyos that are only occasionally wet by flash flows; in elevation from 
peaks in excess of 14,000 feet to depressions below sea level; and in popula- 
tion, from densely populated industrial cities of the Pacific and Gulf coasts, 
to unpopulated mountainous regions. Only about 2 per cent of the area 
is irrigated. 

Increase in Water Demand 

Phenomenal increases in population and in industrialization have placed 
a heavy and continuing demand upon the water resources. These uses in- 
clude water for irrigation, industry, home, navigation, and recreation. Cer- 
tain uses do not deplete the water supply or affect its quality, such as the 
generation of electricity, but water used for irrigation is largely lost to the 
atmosphere by evaporation; while industrial and domestic uses often tend 
to degrade water quality without materially reducing the supply. 

North Pacific Coast Basin (I). This includes the drainage basin of 
the Columbia River plus a few small coastal streams, mainly in southern 
Oregon. A description of the Columbia River will characterize the water 

1 . Prepared for Symposium on Natural Resources of the West. Joint Session of A.A.A.S. Section F 
and the Western Society of Naturalists, Berkeley, Calif., December 28, 1954. 

[ 405 ] 


situation of Basin I. Tliis river, the second largest in the United States, 
has a mean annual flow of about 180,000,000 acre feet. Approximately 40 
per cent of the flow originates outside of the United States in Canada. Of 
the remaining 60 per cent, 25 per cent originates in the relatively small area 
of the Cascades, 25 per cent in the plateau area, and 50 per cent in the 
Rockies. The Cascade section has heavy winter precipitation and runoff, 
while the snow pack of the Rockies provides the principal natural flow 
of late summer. 

The Columbia River is a relatively low gradient stream, yet because of 
its large flow, has tremendous power potentialities : one-sixth of the nation's 
hydroelectric power is generated by plants on this stream; yet to date only 
about 10 per cent of the total 30 million kilowatts of potential power 
is developed. 

Presently irrigated lands of the basin do not place a heavy demand on 
the water supply, only about 5 per cent of the total water supply being 
used for irrigation. It is estimated that approximately 4 million acres more 
can be brought under irrigation. 

Some of the industries of the area require large volumes of water for 
their operation. The large water supply available at relatively low cost 
has not required the employment of water-conserving practices. 

Since the Columbia is a very important fish stream, provisions for the 
movement of migrator}- fish have to be a part of any water-development 
program of the area. 

South Pacific Coast Basin (II). With the exception of a small part 
of the upper portion of the Klamath River watershed, this is located within 
the state of California. Seven hydrographic subbasins within the state are 
recognized; all but No. 6 and part of No. 7 being within the large South 
Pacific Basin. 

The first of these areas or subbasins is a section of high winter precipi- 
tation and runoff. Forty per cent of the state's estimated annual water 
supply of about 70,000,000 acre feet is to be found in this area, which repre- 
sents about one eighth of the state's area. Because of the rugged topography 
and sparse population, there is little demand for the water locally. Plans 
for the transporation of some of the water to areas of deficient supply are 
being formulated. 

Subbasin 2, San Francisco Bay area, some 4,400 square miles in extent, 
is densely populated and highly industrialized. Average annual precipita- 
tion is slightly less than 25 inches, mainly in the form of winter rain. Since 
the average runoff of about I14 million acre feet does not equal demand, 
the two large metropolitan areas, San Francisco and the East Bay, have 
each gone to the Sierra Nevada for additional water. 

Subbasin 3 is slightly in excess of 11,000 square miles, with an annual 
runoff of about two and a half million acre feet. This is another area of 


over-all water deficiency. Except for the northern part, it would be very 
expensive to provide a supplemental water supply. 

Subbasin 4 is a densely populated, highly industrialized area where over 
half of California's twelve million people reside. From this 11,000 square 
miles, about 6 per cent of the state s area, the average annual runoff is 
about li/i milion acre feet, or 1.8 per cent of the state's total. 

The first major importation of water to southern California was made in 
1913, when water was transported from Owens Valley, on the eastern side 
of the Sierra Nevada. Recently the project has been extended northward 
to include the Mono Basin. 

During the '20s thirteen southern California cities joined to obtain Colo- 
rado River water. The main acpieduct, with a capacity of 1600 cfs, was 
completed during the '30s and n.ow most of the important cities in southern 
California, including San Diego, are members of the water district. 

Ground waters are utilized to a high degree. In fact, in certain areas 
adjacent to the coast the ground water levels have been lowered to such 
an extent that salt water intrusion is occurring. To coml)at this menace 
tests have been made on the effectiveness of an interposed fresh water dike 
between the ocean and the ground water basin to exclude the sea water. 
Water has also been obtained from the aqueduct of the Metropolitan Water 
District of Southern California to recharge the underground aquifers. 

Subbasin 5 is the site of the well-known Central Valley Project. Actu- 
ally, the valley is divided into a number of subsections, the main ones being 
the Sacramento and the San Joaquin valleys. Three eighths of the state's 
area, responsible for about a third of the total runoff, is included in this 
division. The northern part, the Sacramento Valley, appears to have excess 
water, while in the south the reverse is true. In the main, then, the problem 
is one of moving the water southward. 

The key unit in the project is Shasta Dam. It serves irrigation, power 
generation, flood control, navigation, salt water repulsion in the lower 
basins, and recreation. As water for irrigation represents about 90 per cent 
of California's water use, most emphasis has been placed on the irrigation 
aspects of the project. 

The plans provide for the use of the extensive underground basins as 
regulating reservoirs. The estimated ground water capacity of the Sacra- 
mento-San Joaquin valley, between limits of 20 to 200 feet, is 130 million 
acre feet, or more than four times the efl'ective storage capacity of Lake 
Mead, back of Hoover Dam. Joint use of surface and ground water storage 
provides a very flexible system to compensate for the irregularities in yearly 
and seasonal runoff. 

Completion of the Friant-Kern Canal came none too soon, as some of 
the ground water basins were being depleted. 

Subbasin 6 lies to the east of the Sierra Nevada, and although much of 


it is relatively high, the average rainfall is comparatively low. Runoff aver- 
ages over 3 million acre feet yearly, of which much flows into Nevada 
through the Truekee, AValker, and Carson rivers. The City of Los Angeles 
diverts water from part of this region. The hydroelectric power poten- 
tialities of the area are fairly well developed by the city and public utility 
companies. This subbasin is a part of the Great Basin and will be men- 
tioned again. 

Subbasin 7, in the southeastern part of the state, is a region of extreme 
aridity, averaging about four inches of rain per annum, and with an aver- 
age runoff of less than 200,000 acre feet, exclusive of the contributions of 
the Colorado River. Palo Verde, Imperial, and Coachella valleys receive 
water from the Colorado River, the latter two from the All- American Canal. 
The water problems of the area are mainly those of deficiency, except in 
those sections which receive water from the Colorado. 

Great Basin (III). — This is a sparsely settled region of high basin 
plains and mountains, with no drainage to the ocean. The climate is arid 
to semiarid except at the higher elevations. The major use for water in 
the area is irrigation. 

No major streams are to be found in this area. Flows from the eastern 
slopes of the Sierra Nevada in California contribute about 3 million acre 
feet per annum, while tlie Humboldt River is the principal stream in the 
state of Nevada. 

Within the Great Salt Lake subbasin are the Bear River and the Weber 
River systems. Both streams are fairly well regulated by storage reservoirs. 
Despite this fact, some one million acre feet of water enter Great Salt Lake 
yearly. While some of this can be conserved, it should be pointed out that 
in irrigation it is necessary to waste a certain percentage of water applied 
so that soluble salts will be removed from the soil. The Weber basin receives 
some water through a diversion from the Colorado Basin, as does the Sevier 
subbasin in the southwestern part of the state. 

The Bureau of Reclamation's plan for the upper Colorado provides for 
importation of water from the Colorado River Basin into Utah. 

Gulf of California (IV). This includes the Colorado River water- 
shed and a few minor streams. Except for the northern portions of the 
basin, the area is characterized as arid. Although the Colorado River water- 
shed is nearly as large as that of the Columbia River, its flow is only about 
one tenth of that of the latter. Within this basin, representing about one 
twelfth of the area of the United States, less than one per cent of the na- 
tion's population resides. 

The principal tributary of the Colorado is the Green River. The Colo- 
rado and the Green are fed mainly from snow, so have good summer flow. 
It is estimated that three fourths of the river's flow originates in the basin 
above the mouth of the Green River in southern Utah. While estimates vary 


as to the average annual flow of the river at the Mexican border, it would 
be about 17 million acre feet under natural conditions. Typical of stream 
flow in arid regions, fluctuations are wide, ranging from 5 million to 25 
million acre feet, and the flow above Lake Mead from 700 to 300,000 cfs. 
Since Hoover Dam lias Ijeen constructed, the downstream flow has varied 
from slightly under 2,000 to about 26,000 cfs. 

The great overriding problem of the Colorado Basin is the attempt to 
meet the many demands for water. These demands greatly exceed the 
supply. This has caused many controversies. In 1922 the Colorado River 
Compact was signed by the representatives of six of the interested states. 
Under this compact 7,500,000 acre feet per annum was apportioned in per- 
petuity to both the upper and lower basins, Lee's Ferry marking the bound- 
ary between them. In addition, the lower basin was appropriated a million 
acre feet. Water unappropriated after October 1, 1963, might be divided 
among the states. 

Under terms of an international water treaty witli Mexico 1,500,000 acre 
feet annually are assigned to that country. This is to come from the amount 
allotted to the lower-basin states. 

Because of aridity, farming in the basin is not practical without irri- 
gation. Some two and three-quarter million acres of land are now irrigated 
within the basin proper, plus slightly less than half a million acres in the 
Imperial Valley. Exclusive of the Imperial Valley, the area irrigated is 
about equally divided between the upper and the lower basins. However, 
because of aridity, water requirements per unit area are greater in the 
lower basin. 

Within the past score of years a series of dams have been built, includ- 
ing Hoover, Davis, Parker, Imperial, and most recently ]\Iorelos Dam, a 
structure built by Mexico just south of the border. In addition to the dams 
are the great irrigation systems, including the All-American Canal serving 
the Imperial and Coachella valleys, and the extensive system now being 
built into the Gila Valley. 

Plans by the Bureau of Reclamation for the river's development provide 
for ten large dams in the upper basin capable of storing some 48.5 million 
acre feet of water for multipurpose use. Besides providing for irrigation 
of new land in the upper basin, estimated at 380,000 acres, the development 
will provide a supplemental supply to nearly a half-million acres now 
developed, much of which lies outside the drainage basin of the Colorado 
in Utah and Colorado. Transmountain diversions of Colorado River water 
to areas east of the Rockies amount to about 200,000 acre feet annually, and 
another diversion of 300,000 acre feet is contemplated. Diversion of low-salt 
content water from the upper basin will have a deleterious effect on the 
water quality of the lower basin, the quality of which will continue to de- 
teriorate as the upper basin states increase their water use. 


The heavy sediment load of the Colorado poses the very serious problem 
of reservoir silting and stream channel clogging. High river gradient and 
poor vegetative cover over much of the catchment are important contribut- 
ing factors. Five million acre feet is provided in Lake Mead for debris 
storage. Additional control works are required upstream so that excessive 
flows, the main contributor to debris movement, will be reduced. Not all 
silt problems are confined to the stream above Hoover Dam. The desilted 
water from each of the storage reservoirs has caused a degrading of the 
stream channel immediately below the structure and an upgrading imme- 
diately above the next downstream structure. 

Present hydroelectric plants on the Colorado Kiver have an installed 
capacity of over li/o million kilowatts. The Hoover Dam plant has the 
largest output, while power from Parker will be used mainly to deliver 
water to the system of the Metropolitan Water District of Southern Cali- 
fornia against a total head of over 1,600 feet. Estimated total potential 
hydropower for the basin is about 6,500,000 kilowatts. 

Arizona is the only state in this basin that makes extensive use of 
ground water. During the period 1948-1953 some 400,000 acres were 
brought under irrigation, mainly through the use of ground water. 

Gulf of Mexico Basin (V). This is an area where the demand for 
water for irrigation and industry is expanding very rapidly. Ground water 
supplies have been developed to a high degree, in some instances overde- 
veloped. Additional supplies can be provided by storage of the flood waters 
on those streams draining the central part of Texas. In fact, some of the 
streams in the Mississippi Basin are jiotential sources of additional water 
for this basin. 

The Rio Grande River, an international stream, is the principal river 
of the basin. Rising in Colorado, it flows some 1800 miles to the Gulf of 
Mexico. It has a watershed of 185,000 square miles, of which 57 per cent 
is in the United States. Practically the entire flow of the upper portion 
of the stream, some 3 million acre feet, is used locally, mainly for irrigation. 
The downstream flow's annual average, about 5 million acre feet, is provided 
by tributaries arising in Mexico and from the Pecos River. Elephant Butte 
and Falcon are important water-impounding structures which help control 
stream flow. An important feature of stream regulation is the maintenance 
of a desirable salt balance. In order not to build u]i an undesirable amount 
of salt in the irrigated lands, a certain amount of water must be released 
so that excess salt will be carried out of the basin to the Gulf of Mexico. 

Among the streams to the east of the Rio Grande are the Nueces, San 
Antonio, Guadalupe, Colorado, Brazos, Sabine, and Trinity. Most are sub- 
ject to wide seasonal and annual variations in flow, conditions which gener- 
ally make storage expensive. The hydroelectric output of the area is not 


large, but the basin generally does not lack for power because of the great 
oil and gas resources. 

MissLssn^pi KivER (VI). This area drains all of five states and parts of 
six others, more than a half -million square miles, and extends from the 
Canadian border to the Gulf of Mexico. It includes the Missouri River 
basin plus such other important streams as the Red, White, Black, and 
Arkansas rivers. While most of the important tributaries have their sources 
in the Rockies, the major portion of the basin occupies plain lands to the east. 
For a long time the chief attention given to these streams concerned flood 
control. Now a huge multii)urpose project is underway to develop the 
water resources of the area for a wide range of uses. Within the past decade 
there has been a marked increase in irrigation. Grasslands are giving way 
to irrigated farms. 

Important tributaries of the Missouri River from the west include Milk, 
Yellowstone, Cheyenne, Platte, and Republican rivers. The Yellowstone 
is an important tributary with an average annual runoif of nearly 7 million 
acre feet. Its flow characteristics are similar to the other tributaries that 
drain the Rockies. The major portion of the runoff occurs during April, 
May, and June. 

A comprehensive federal program for the Missouri Valley is now under 
construction. It contemplates the ultimate construction of some 90 reser- 
voirs, which will have a capacity of about 46 million acre feet. Reservoirs 
on the tributaries will be used for irrigation, flood control, and power de- 
velopment. Main river storage will be chiefly for navigation improvement, 
power, and irrigation. It is estimated that the ultimate development will 
bring under irrigation about 5 million acres, and produce nearly four bil- 
lion kilowatt hours of electricity annually. 

Surely this area is undergoing a transition period. 

Increasing the Effectiveness of Our Water Resources 

Cloud seeding and sea water reclamation are two proposals that often 
appear in the news. To what extent precipitation can be increased by cloud 
seeding is still a question upon which there are many opinions. Extensive 
experiments on demineralization of sea and less saline waters are now being 
conducted. Results to date show promise for the reclamation of waters 
which are slightly too saline for irrigation use, but the problem of reclaim- 
ing sea water at a cost which is currently considered reasonable is yet to 
be accomplished. 

The avenue that offers the greatest opportunity to increase the effective- 
ness of our water resources is to increase the use-efficiency. Farmers can 
adopt better means of water application, industry can, through re-use, mate- 
rially reduce the requirements for large flows; and through the elimination 
of large areas of noneconomic, high-water-consumption plants, water can be 


conserved. A conscious effort should be made not to pollute the water sup- 
ply unnecessarily, so that it will be available for re-use. Surface and ground 
water storage of flood and other surplus water, although often expensive, 
has to be resorted to in the semiarid and arid sections. 

Water Resources of the Future 

I have no crystal ball, nor am I the seventh son of a seventh son, but cer- 
tain trends in the development and use of our water resources do seem fairly 
clear. For the Northwest Basin, the development of the tremendous power 
potential of the Columbia River will continue, giving rise to a great indus- 
trial development. Irrigated acreage will increase, but despite all local de- 
mands, some 150,000,000 acre feet of fresh water will pass- annually into 
the sea. 

In the South Pacific Basin great efforts will be made to equalize the geo- 
graphic disparity between water resources and water demand. Maintenance 
of acceptable water quality, especially in the ground water basins, will be 
a problem. Careful use of water by farmers and industry will increase 
water use efficiency, including re-use of some of the water now wasted to 
the sea by coastal cities. Some of the isolated ground water basins will be 
pumped to the economic limit and then abandoned so far as irrigation 
is concerned. 

As for the Great Basin, it is difficult to visualize great water resource 
potentialities despite the proposed diversions from the Colorado River. The 
basin is rich in mineral wealth but not in water. 

The Gulf of California Basin will always be an area of low water supply. 
Upper Colorado River development will harness the water for multipurpose 
projects. This will result in a general degrading in water quality down- 

In the Gulf of Mexico Basin west of the Mississippi, especially in the 
eastern and south central portions, the indications are that there will be a 
strong demand for the harnessing of local water supi^lies, together with a 
move to bring water from the northeast. The great industrial potential of 
the coastal section is in no small measure associated with good water 

The western Mississippi River basin will undoubtedly undergo marked 
changes during the next few decades. Available water during late summer 
and low-cost power will favor expansion of irrigation and encourage indus- 
trial development. Harnessed water resources will pei'mit higher use of 
much of what the American people have come to know as the grasslands. 



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United States Congress 

1952. II. The Physical Basis of Water Supply and Its Principal Uses. Interior 

Insular Affairs Committee. Pp. 1-93. 

1953. III. Ground Water Regions of the United States — Their Storage Facili- 

ties. Interior Insular Affairs Committee. Pp. 1-78. 

United States Department of Agriculture 

1948. Irrigation Agriculture in the West. Miscellaneous Publication 670, pp. 

United States Department of Commerce 

1950. Irrigation of Agricultural Lands. United States Census of Agriculture. 
United States Government Printing Office. Washington, D. C. 


United States Department of Interior 

1946. The Colorado River, a Natural Menace Becomes a National Resource. 

Bureau of Reclamation. Pp. 1-295. 

1947. The Columbia River. Bureau of Reclamation. Volume 1, pp. 1-399. 

1948. Report on Central Arizona Project. Bureau of Reclamation. Mimeo- 

1950. The Columbia River. Bureau of Reclamation. Pp. 1-690. 

United States Geological Survey 

Water-Supply Papers, Surface and Ground Water. Numerous — United 
States Printing Office. Washington, D. C. 

United States Senate 

1944. Missouri River Basin, Conservation, Control and Use of Water Resources. 
Senate Document 191. 78th Congress, 2nd Session. Pp. 1-211. 

University of Oregon 

1953. Research in the Policy and Administration of Natural Resources. De- 
partment of Political Science. Eugene, Oregon. Pp. 1-68. 




Fourth Series 
Vol. XXVIII, No. 11, pp. 415-423 February 17, 1956 



Executive Vice President, Forest Genetics Research Foundation 

This discussion pertains to the future productivity and management of 
the commercial forest lands of the ten western states, the area extending 
from the eastern borders of Montana, Wyoming, Colorado, and New Mexico 
to the Pacific Ocean. Commercial forest land is defined as "forest land bear- 
ing or capable of bearing timber of commercial character and economically 
available now or prospectively for commercial use and not otherwise with- 
drawn from such use" (1). Noncommercial forest land is chiefly valuable 
for purposes other than timber production. Since its management differs 
markedly from that of commercial forest land, it lies outside the scope of 
tliis paper. 

The western states contain some 108 million acres of commercial forest 
land, mostly in Montana, Idaho, Washington, Oregon, and California (2). 
These lands bear 98 per cent of the nation's virgin timber. The western 
forests are notable for the presence of such commercially valuable species 
as Douglas fir {Pseudotsuga menziesii (Mirb.) Franco), ponderosa pine 
{Pinus ponderosa Laws), sugar pine {Pinus lumhertiana Dougl.), western 
white pine {Piiius monticola Dougl.), and redwood (Sequoia sempervirens 
(D. Don) Endl.), not one of which occurs elsewhere in the United States. 
Some of these forests, notably the Douglas fir, spruce-hemlock, and redwood 
types of western Washington, Oregon, and northwestern California, have 
growth rates probably equal to any in the northern hemisphere. In contrast, 
some forests of the ponderosa pine type that occur in localities having low 
precipitation grow very slowly, but if allowed to mature for several cen- 
turies produce timber of high value. About two thirds of the western com- 
mercial forest area is in public ownership, mostly federal. The privately 
owned lands, constituting those selected out of the public domain prior to 
the establishment of the national forests, are generally more accessible and 

1. Prepared for Symposium on Natural Resources of the West. Joint Session of A.A.A.S. Section F 
and the Western Society of Naturalists, Berkeley, Calif., December 28, 1954. 



Tlie commercial forests of the West are currently producing nearly 40 
per cent of the nation's sawtimber, and timber production is, of course, one 
of their principal uses. The other values inherent in these forests, however, 
are of critical importance to the West, and must be given due weight in 
management practices. I refer primarily to water yield and the production 
of forage that provides grazing, mostly during the summer season, to sub- 
stantial numbers of domestic livestock. The use of commercial forests for 
recreation, including hunting and fishing, does not basically affect manage- 
ment practices. 

The principal tasks that lie ahead in making the western commercial 
forests fully productive of timber can be grouped under four general ob- 
jectives: protection; conversion of virgin forests into well-stocked growing 
stands; prompt establishment of adequate regeneration; and improvement 
in quality and yield of wood. 

The first objective, protection, is designed to decrease losses from fire, 
insects, and disease. The western forests suffer an annual loss of more than 
one billion board feet of timber from these causes. About one fourth is due 
to fire, and three fourths to insects and disease. This loss can never be en- 
tirely prevented, but it can be materially reduced. 

Eeduction of fire losses will be achieved by fire prevention, largely 
through public education; increase in specialized, highly skilled personnel; 
development of improved equipment; and aggressive research upon the na- 
ture of and relationships between the ground and atmospheric conditions 
affecting forest fire. 

Progress is being made in reduction of losses from parasitic insects. 
New, powerful insecticides, applied to large forest areas by airplane, are 
proving increasingly efficaceous in controlling certain types of insects. 
Other types are being attacked by silvicultural methods providing environ- 
ments favorable to tree vigor and unfavorable to the insects. 

The struggle to decrease losses from the many fungus diseases attacking 
the forest is seemingly never ending. Losses caused by wood-rotting fungi 
are often heavy in virgin forests containing a large proportion of old trees. 
As these virgin forests give way to young, vigorous stands of second-growth 
timber this ty])e of damage will decrease. Silvicultural methods seem prom- 
ising for the control of some diseases attacking juvenile trees. 

A new approach, the genetic development of tree forms resistant to in- 
sects and fungi, is now assuming importance. Current research is giving 
encouraging results. 

Attainment of the second objective, conversion of virgin forests into 
well-stocked, growing stands, is necessary to assure permanent production 
of wood products. Only a system of forestry which provides for the growing 
and harvesting of successive timber crops can guarantee a continuous tim- 
ber supply. The virgin forest is a natural heritage of great value. To the 


commercial forester, however, it can only be considered as the current crop 
of timber, to be harvested and replaced by the man-cultured young forest 
that will, when mature, constitute the next timber crop. In commercial 
forestry the tree farm will supplant the forest primeval. It is recognized, 
of course, that areas of virgin forest required for scientific study and for 
recreation will be reserved and are then, by definition, no longer consid- 
ered as commercial forest. 

Effective conversion of a virgin forest into a vigorous, well-stocked 
second-growth stand is a vexatious and difficult matter. Methods of harvest- 
ing the virgin timber have markedly different effects upon the amount of 
regeneration and the promptness of its appearance. The relative merits 
of clear cutting and selective cutting of virgin stands are frequently debated 
by foresters. Both doubtless have their place in the western forests. In the 
dense stands of the Douglas fir and spruce-hemlock types in western Oregon 
and Washington, clear cutting in small enough blocks to assure adequate 
seed dispersal from surrounding trees is proving effective. In many of the 
thinner ponderosa pine forests, where every acre may bear trees from seed- 
ling size to merchantable veterans, selective cutting often proves more 

To secure the third objective, prompt establishment of adequate regen- 
eration on cut-over areas and in poorly stocked juvenile stands, is a difficult 
task at best in many parts of the West. A substantial portion of the west- 
ern commercial forest area is characterized by high temperatures and scanty 
precipitation during much of the growing season. These conditions militate 
against the establishment of sufficient numbers of coniferous seedlings to 
form an adequate stand. The density of many virgin forests indicates that 
in the past nature has coped with the problem successfully. But nature is 
content to spend many decades in developing a dense forest; man is not. 
Many of our best virgin forests may well have resulted from a fortuitous 
sequence of such favorable circumstances as the sudden destruction of an 
old forest by fire during the late summer of a good seed year, followed by a 
year of temperatures and summer precipitation particularly favorable to 
the establishment and survival of forest tree seedlings. But commercial 
forestry cannot await the operation of so uncertain a process, so other ways 
must be found to secure prompt regeneration. 

Planting, as a solution to the problem of uncertain natural regeneration, 
has been only partially successful in the West, because adverse field condi- 
tions affect planted tree stock and natural seedlings alike. Also, it is a costly 
process. Nevertheless, it has certain advantages over natural regeneration. 
Where successful, it assures a stand of the desired species, with the trees 
properly spaced to produce high yield at maturity. And it is only by plant- 
ing that trees genetically improved by breeding can come into use. 

It is probable that forest planting in the West could be much more 


successful, despite adverse field conditions. Study of the methods commonly 
used in seed collection, nursery growing of the planting stock, and its out- 
planting in the field leads me to the conclusion that practices consistent 
with the principles of the plant sciences are frequently sacrificed in order to 
keep costs at a minimum. The cost accountant seems able to outvote the 
plant scientist in determining procedures. Analyses of current planting 
methods by plant scientists and more research on problems not yet solved 
would, I am sure, materially increase the degree of success of planting. 

Direct seeding, in contrast to the planting of nursery grown stock, offers 
some promise for artificial regeneration. Two methods are being tested: 
broadcasting of seed from airplanes and planting of the seed at regularly 
spaced intervals. The greatest deterrent to either method is the presence 
of rodents that eat the seeds or the seedlings. Temporary heavy reduction 
of the rodent population on the seeded area by the broadcasting of poison 
bait may be the answer to this problem. 

The fourth objective, improvement in quality and yield of wood, opens up 
new and interesting possibilities in forestry. It holds the promise of increas- 
ing timber production by application of the same scientific knowledge that 
has increased farm crop production so markedly. There are two possibili- 
ties: tree improvement for greater yield and better quality, and soil im- 
provement for greater yield. 

In the United States, forest tree improvement is a new and rapidly 
developing field of research from which material benefits can be expected. 
Its object is the development of tree forms specially suited to local condi- 
tions, including resistance to the insect and fungus pests of the particular 
region; good form and growth rate; and special wood qualities designed to 
meet the requirements of special uses. Two general methods of forest tree 
improvement are being used : selection of superior races and phenotj^^es, and 
development of new forms by breeding. 

Provenance studies, by which races of widely distributed species are 
tested under a variety of environmental conditions, are useful tools in selec- 
tion of races and are being widely used. In most cases a race is better suited 
to its own environment than to any other, but tests have demonstrated 
valuable exceptions. 

A practical method for selection of superior phenotypes is by securing 
seed from groups of superior trees, carefully selected for desired character- 
istics and sufficiently isolated to prevent cross-pollination from other, poorer 
forms. This can be accomplished by harvesting or roguing out the inferior 
trees on an area containing a goodly number of superior individuals and by 
removal of all trees of the same species over a surrounding area of sufficient 
size to effect the necessary isolation. Cross-pollination thus occurs only 
between superior trees. 

Increasing evidence indicates that the continued, persistent selection of 


superior races and phenotypes will bring a degree of improvement of forest 
growing stock that justifies its application to forestry. While admittedly 
slow, improvement is cumulative in effect. Selection has the advantage of 
simplicity in comparison with tree breeding and can be applied by foresters 
who do not have specialized knowledge of plant genetics. 

Forest tree breeding, however, holds the greater potentiality for forest 
tree improvement. Forest trees are one of the few widely used kinds of 
plants which have not been materially improved by genetical means, and 
the potentialities for their improvement are as great as those for any of our 
agricultural crops. Over the past century a number of natural forest tree 
hybrids were described and a few artificial hybrids were produced, but no 
attempt was made to apply the information to practical forestry. It is only 
in the last two decades that forest genetics has been recognized as a distinct 
branch of plant genetics, with its own objectives, methods, and increasing 
body of literature. 

In the United States, breeding work on many forest tree species is now 
being conducted by numerous universities and by federal, state, and private 
research institutions. The work is largely exploratory in nature, in that 
very few artificially bred tree forms are old enough to permit evaluation at 
maturity. A large number of hybrids are being produced, mostly interspe- 
cific, but with an increasing number of intraspecific crosses between well- 
defined races of several species. Some of these new forms indicate in the 
juvenile stage that desirable traits of the parents can be brought together in 
the Fi generation. A few hybrids indicate true heterosis with attending ad- 
vantages of rapid grow^th. A number of the more promising hybrids are now 
undergoing large-scale field tests, generally in the form of comparison with 
the parent species. The breeding work is developing better knowledge of 
species relationships within several important genera, which will facilitate 
future development of new forms. 

One of the greatest needs is for more research of a fundamental nature to 
provide tree breeders with information which will make their work more 
effective. Physiological and genetical studies of vegetative reproduction, 
flowering, growth, and wood structure of forest trees, and evaluation and 
measurement of the range of genetic variability of important timber species 
and their races are examples. Kecent and valuable work by Pauley and Perry 
on ecotypic variation (3) and Mirov on the gum turpentines of pines, dis- 
cussed in more detail later, are examples of the kind of research needed. 

In the western states most of the work in forest tree breeding has been 
conducted by a single organization, the Institute of Forest Genetics at Placer- 
ville, California, a branch of the California Forest and Range Experiment 
Station, U.S. Forest Service. The Institute has made no attempt to cover 
the entire field of forest tree improvement, but has limited its work to breed- 
ing and related studies of species within the genus Pinus, one of the most 


important and valuable genera in the West. This specialization has brought 
notable progress within its narrow field. There remains, however, the need 
for work on other genera. 

The Institute s principal work has been the production of pine hybrids. 
Since the inception of its breeding program in 1927, some 67 pine hybrids 
have been developed. Several exhibit marked heterosis, a phenomenon seem- 
ingly most apt to appear in the product obtained by crossing two closely 
related species of widely separated occurrence. An example is the hybrid 
P. monticoJa Dougl. X P- strohus L., which greatly exceeds either parent in 
growth. Another hybrid, thus far exhibiting satisfactory form and growth 
rate, seems to possess the very high resistance of one parent to a specific 
insect parasite. This hybrid, P. jeffryi Grev. & Balf . X P- Coulteri D. Don, 
has the commercially desirable form of P. jeffryi, is superior in growth to that 
parent, and evidently possesses the extreme resistance of P. coulteri to the 
reproduction weevil, Cylindrocopturus eatoni Buch., a destructive parasite 
occurring on young trees of several pine species, including P. jeffryi. The 
hybrid of P. strohus, susceptible to white pine blister rust, Cronartium rihi- 
cola Fischer, and of P. griffithii McClelland, highly resistant to the rust, has 
undergone seven years of exposure to the disease with no evidence of infec- 
tion. These examples are indicative of the progress possible through speciali- 
zation of forest tree breeding to a single genus. 

The Institute's investigation of the chemistry of gum turpentines of pines, 
conducted by Dr. N. T. Mirov, has proved to be of special value. The study 
has included the turpentines of most of the pine species of the world for 
which previous analyses were not available, and analyses for the species of 
the United States and Canada are now practically complete (4) . It has been 
found that the turpentine content of any species, or in some cases of a well- 
defined race, is uniform, irrespective of environmental conditions. Hybrid 
species contain all of the turpentines of both parents, but in other propor- 
tions. The resistance of the hylirid P. jeffryi X P- Coulteri to the reproduc- 
tion weevil is seemingly associated with the presence of an oleoresin origi- 
nally occurring in the resistant P. Coulteri and absent in the susceptible 
P. jeffryi. Further, the degree of resistance appears to be related to the 
amount of the oleoresin in the hybrid form, as indicated by the fact that 
the Fi hybrid, containing more of the oleoresin, is more resistant, and the 
back cross onto P. jeffryi, containing less of oleoresin, is less resistant. These 
facts justify the hypothesis that the oleoresin is the cause of the resistance. 
Classification of pine species based upon their turpentines provides a better 
basis for determining species crossability than taxonomic systems based 
upon macroscopic plant characters (5). These results from the study of 
gum turpentines illustrate the value of fundamental research to provide 
the tree breeder with information basic to his work. 

The yield of timber from a forest crop is directly related to the fertility 


of the soil upon which it grows. Tlie possibilities of increasing the yield on 
poor soils are limited. The need for soil protection and soil improvement 
will become more evident as forest geneticists develop faster growing tree 
forms that wall make severe demands upon the supply of plant nutrients 
available in the soil. 

Forest soils can become impoverished and, in fact, can be made unsuit- 
able for forest tree growth in several ways. Repeated, severe forest fires 
will adversely affect the structure, physical condition, and chemical and 
biological nature of forest soils. They will cease to possess the combination 
of properties by which we characterize forest soils. Natural processes, in- 
volving the recapture of the site by arborescent vegetation, will finally re- 
establish the forest, but a long period of time is often necessary. 

Comparable damage to forest soil can result from practices causing top- 
soil losses by erosion. Logging and road construction on steep slopes, par- 
ticularly when accompanied by almost complete destruction of the vegeta- 
tive cover, often cause heavy erosion. A degree of soil disturbance that 
merely loosens and cultivates the topsoil without heavy erosion may favor 
forest regeneration. But disturbance that results in exposure of heavy, com- 
pacted subsoil makes regeneration difficult. 

Forest soil improvement has hardly progressed beyond the stage of 
theory in the United States. We have not as yet accepted the idea that soil 
fertilization is physically or economically practicable, except in cropland 
soils. Progress in forest soil fertilization in other countries wall, I hope, 
stimulate research here. In Australia apx^lication of phosphates to forest 
soils has resulted in an increase of tree growth that may make the practice 
economically desirable, and application of small amounts of zinc to IMonte- 
rey pine seedlings has brought astonishing results (6). Research on fer- 
tilization of western forest soils, including determination of its economic 
feasibility, is a new and open field of study. Many forest soils in the "West 
are deficient in nitrogen and phosphorus, and little is knowni of the effect 
of adding these elements. Also, a study of micro-element deficiency might 
bring results of practical value. 

The water demands of the West, to meet the increasing domestic, indus- 
trial, and agricultural needs of the growing population, are constantly ex- 
panding. The commercial forest areas, together with the high mountains, 
are the source of this water. No matter how far it must be transported to 
meet the needs of population centers, its invariable sources are the moun- 
tainous areas of high precipitation. Forest management, therefore, must 
do its share to maintain and, if possible, augment water yield. 

In a thoughtful and informative paper. Dr. E. A. Colman (7) recog- 
nizes three w^ays by which forest management can contribute to water yield : 
protection, repair, and improvement of forested watersheds. Protective 
measures are designed to maintain a favorable water flow and low erosion 


rate equal to those of the virgin forest. They include fire protection; careful 
location, construction, and maintenance of roads; and the use of logging 
methods least damaging to the structure of the forest floor, the surface 
accumulation of organic matter above the mineral soil. 

Repair of damaged watersheds consists of such measures as immediate 
revegetation, preferably with valuable forest species, of forest lands de- 
nuded by fire; prevention of erosion on denuded lands by construction of 
small dams; soil stabilization on areas disturbed by logging; and clearing 
of stream channels choked with logging debris. 

To quote Dr. Colman, "The objectives of water yield improvement are 
to reduce streamflow in winter and early spring when supplies are greater 
than needs, increase the low flows of the dry season when water is in short 
supply, and decrease evaporative water losses in ways that will increase 
the quantity of usable water available downstream." Past experiments indi- 
cate that the desired regulation of streamflow may be accomplished by tim- 
ber harvesting methods favorably affecting snowpack, consequent rate of 
snowmelt, and direct evaporation of water from snow. A recent experi- 
ment in Colorado indicates the value of timber cutting in some pattern of 
strips or blocks that lessens the tree crown surface of the forest, thereby 
permitting more snow to reach the ground, yet providing the most possible 
shade. The effect is to decrease snow evaporation in the tree crowns and 
to retard, by shading, the rate of snowmelt on the ground. Similar experi- 
ments are needed in other parts of the West where other conditions exist. 

Of the 17^2 million acres of commercial forest land in California, some 
10 million acres, or 57 per cent, produce forage utilized by domestic live- 
stock, game animals, or both, and some such proportion probably holds for 
all of the western states. Some of this area has permanent grazing value 
but most of it can be grazed only during limited periods in each forest rota- 
tion. A substantial part of the forage occurs in the open meadows fre- 
quently found in the forest. Wherever occurring, forest forage must be 
protected from over-grazing, and depleted ranges must be repaired, in order 
that soil losses, destructive to timber, water yield and forage alike, may 
be prevented. 

There is at present widespread feeling that western forest management 
practices for timber, water, and forage production differ so greatly that 
they are in serious conflict. If, however, their mutual interest in common 
objectives is weighed against their differences, I see no basis for real con- 
flict. All three call for protection from fire, except where used under 
control as a tool of resource management, and from other destructive forces. 
All recognize the need for prevention of accelerated erosion and loss of top- 
soil, and for prompt revegetation of denuded areas. In all cases, sound 
management practices to attain these ends will, perforce, be based upon the 
economically feasible application of the plant and soil sciences and other 


disciplines such as hydrology and climatology. Recognition of this broad 
area of common interest will relegate conflict to the level of differences 
which can be settled by adjustment. 

Predicted increases in the population of the western states will make 
demands upon the commercial forests of the region which will justify a 
more intensive degree of management than is now being practiced. Much 
more knowledge than is at present available will be needed, and many diffi- 
cult problems will have to be solved. All that can now be said is that no 
end to the development of forest management practices can be foreseen. 
As new technologies develop new products, the kinds of raw materials 
needed from the forest may change, with consequent changes in manage- 
ment objectives. The quest for knowledge to meet increasing demands for 
forest products, old or new, will require an expanding and continuing re- 
search effort in which western universities and other research institutions 
will have an important part. The combination of continued research and 
conservative management will maintain throughout the future the produc- 
tivity of the western forests. 


1. Society of Americax Forksters 

1950. Forestry Terminology. 

2. United States Department of Agricxtlture — Forest Service 

1946. Gaging the Timber Resource of the United States. 

3. Pauley, S. S., and T. O. Perry 

1954. Ecotypic Variation of the Photoperiodic Response in Populus. Journal 
of Arnold Arboretum. XXXV: 167-188. 

4. MiRov, N. T. 

1954. Chemical Composition of Gum Turpentines of Pines of the United States 
and Canada. Journal of the Forest Products Research Society, IV: 1-7. 

5. MiRov, N. T. 

1953. Chemical Aspects of Diploxylon Pines. Zeitschrift fur Forstgenetik und 

Forstpflanzenzuchtung. 2. Band. Heft 5. 

6. Stoate, T. N. 

1950. Nutrition of the Pine. Bulletin No. 30, Forestry and Timber Bureau, 

7. COLMAN, E. A. 

1954. Water Control and Timber Management. Proceedings of Annual Meeting 

Western Forestry and Conservation Association. 




Fourth Series 
Vol. XXVIII, No. 12, pp. 425--439 February 17, 1956 



Director and Professor 
School of Fisheries, University of Washington 

As a result of the last war we have had closer contact with backward 
nations where deficient diet is the rule. This contact has been maintained 
through the work of the United Nations assistance groups with many na- 
tions in the less-developed areas of the world. All these have served to 
emphasize the growing disparity between rate of population increase and 
the present apparently limited capacity of our land masses to increase their 
productive capacity indefinitely. The gloomy predictions of Malthus appear 
to be brought nearer to realization by the success of medical science in re- 
ducing the effects of disease and raising infant as well as adult survival 
rates. Anticipating the difficulty of feeding the future world's population 
the search for new sources of food has been intensified. 

In surveying potential sources of large quantities of food one is always 
struck with the fact that whereas the oceans occupy almost three quarters 
of the earth's area, they produce at present only ten per cent of the world's 
total protein. With some areas obviously underfished and others apparently 
capable of a much liigher production, speculation has not only embraced a 
possible increase in the production of fish but has included suggestions of 
the direct use of plankton which eventually is converted into fish with a 
considerable loss of efficiency (Reay, 1954). Others have suggested that 
work should begin on the development of methods of actually farming the 
sea (Daniels and Minot, 1954). The latter would be based upon the known 
large quantities of nutrient materials in the sea that are made available 
for the use of growing plants and animals only where the ocean current 
systems bring them into the euphotic zone. Experiments on the fertilization 
of enclosed arms of the sea (Eaymont. 1947) and with artificial propagation 

1. Prepared for Symposium on Natural Resources of the V/est. Joint Session of A.A.A.S. Section F 
and the Western Society of Naturalists, Berkeley, Calif., December 28, 1954. 
Contribution Number 12, University of Washington, School of Fisheries. 



of marine fishes liave been tried with some success in Europe (Dannevig, 
1951) and salt marshes have been used for some time in Asia for the culture 
of CJmnos clianos or milk fish; but the actual use of the rich mineral deposits 
held in the ocean depths seems to be a remote possibility at the moment. 

To one who has had experience with the collection of plankton on the 
high seas, the harvest of amounts that would contribute significantly to 
our food supply seems impractical even if ways could be found to utilize 
it directly. For the present at least, it appears to be much more economical 
to permit the fish, which are especially fitted for gathering and converting 
the plankton into palatable proteins to make this harvest for us. The most 
promising line of attack in this direction appears to be the rumored search 
for methods of cultivating phytoplankton. Such a process might produce 
large quantities of desirable forms eliminating the difficulties of sorting as 
well as capture. 

Taking a more practical view, the United Nations conference on natural 
resources held at Lake Success, N. Y., in 1949, considered methods of in- 
creasing the production of fish from the sea immediately, to fill the current 
needs of food-deficient areas. It has been estimated that the total world 
production of fish in 1950 was 25,000,000 metric tons. Of this just six na- 
tions in the northern hemisphere produced 61 per cent. Japan alone before 
the last war produced 38 per cent of the world's total (Harold Thompson, 
1951). A greater disparity is shown by the 98 per cent produced in the 
northern hemisphere as compared with 2 per cent in the southern. That is, 
the more highly developed fisheries and consequently the greatest concen- 
tration of fishing effort are at present associated with the most highly indus- 
trialized countries in the northern hemisphere. In this region fish are 
produced by several countries far in excess of their own needs and have 
become an important commodity in their foreign trade. Consideration of 
all the possible sources of fish known to the participants led the United 
Nations conference to anticipate increases of from 22 per cent (II. Thomp- 
son, 1951) to 36 per cent (F.A.O. United Nations, 1953) by 1960 over the 
1950 level. LeGall (1951) stated that present needs are double the amount 
of fish now produced and that by 1960 they will be three times this figure or 
about 75,000,000 metric tons. The discrepancy between the growing need 
and predictions of what might be taken are striking and the greatest need 
for this increase is in tlie most densely populated areas where living stand- 
ards are low and diets are especially deficient in protein. 

At the present time the fishing industry does not hold such an important 
position politically or economically in the United States as it does in other 
countries with comparable levels of fish production. For example, total pro- 
duction of all meats in this country amounted to about 23 billion pounds 
in 1952 whereas fish production totaled about 4 billion pounds in the same 
year, of which approximately 25 per cent was used for food. Per capita con- 


sumption ol" fish in the United States is only about 11.5 pounds (1948) per 
year as compared witli three times that amount, 33.3 pounds in the United 
Kingdom, 83.3 in Japan, and 59.3 in Norway (Tressler and Lemon, 1951). 
Another situation that affects our fisheries adversely is that several of our 
allies are deficient in other natural resources and now use fish as one of their 
export products which help to maintain their balance of trade with the 
United States. The present need to help our allies may require relaxing 
tariff restrictions on such products, which would aggravate the competition 
our fisheries must face from countries in which production costs are low 
compared to ours. 

However, the fisheries of the United States must increase their impor- 
tance in the future, Ijotli as a source of living to those associated with them 
and also as a source of protein food. Our population growth is following 
the same course as that of other countries, and is continually increasing 
the ratio of population to acres of food-producing land. Like other coun- 
tries, we will eventually have to look to the sea for more of our food. Even 
now the United States is one of the best markets for higher-priced fish 
products and the sale of other fish could be greatly expanded if increased 
production efficiency widened the price differences between fish and compet- 
ing products such as meat. Additional sales would undoubtedly develop if 
our fisheries industry could learn to market top-quality material in as pleas- 
ing form and good condition as prevails for most meats. AVith a growing 
interest in technology and marketing, these developments can be expected. 
Moreover, if the potential demand of large populations, now prevented l)y 
poverty from buying food, can be released, the full productive capacity of 
the fishing fleets of the entire world will be required. 

In spite of the relatively low position of fisheries in the United States, 
we are now the second greatest fishing nation in the world in total produc- 
tion. Of the total fish produced in this country an average of 43.4 per 
cent came from the West Coast and Alaskan areas over the period of 1940 
to 1951. For this, as well as for other reasons that will be made apparent 
below, our Pacific Coast fisheries can be expected to play an important role 
in the future. We should therefore examine their present condition and 
attempt to foresee what ]iossibilities they offer for future production. 

Present Condition of West Coast Fisheries 

The existing fisheries of the West Coast of the United States and Alaska 
do not on the whole present an optimistic picture. In Alaska, the salmon 
fisheries have completed a second poor season with a pack of slightly over 
3,000,000 48-pound cases of canned salmon as compared with 2,900,000 
cases last year which in turn may be compared with an average of almost 
4,000,000 cases for the last 10 years. The trend has been downward since 
1934. Off the coast of California the sardine fishery has been practically 


nonexistent for two years and the Pacific mackerel seems to be following 
the same path. The salmon fisheries of the Columbia River in 1952 and 
1953 showed signs of furtlier decline beyond the drop that occurred in 1943. 

In contrast, however, the halibut fislieries under the International Pa- 
cific Halibut Commission and the sockeye of the Fraser under the control of 
the International Pacific Salmon Fisheries Commission are increasing. The 
other major West Coast fishery, the California tuna industry, is suffering 
primarily from economic competition with other tuna-producing countries. 

Our West Coast fisheries therefore present a variety of problems the 
solution of whicli will determine the extent to which we will realize their 
full potentialities. They range from the need for expansion and stabiliza- 
tion of some fisheries through the combined efforts of fisheries biology, 
hydrography, and exploratory fishing, to the obvious need for fisheries 
management of some sort in several declining fisheries. They include the 
problems of adjusting the uses of water for power and irrigation to the 
needs of the anadromous fishes, as well as the need for development of a 
basis for handling oceanic fisheries problems that arise from international 
competition for the fisheries of the high seas. They also involve the develop- 
ment of more efficient methods of fishing and of handling, processing, and 
marketing, as well as conserving and possibly culturing our fishes. 

It is not surprising that the general picture of the present condition of 
our fisheries has become confused by a combination of economic and bio- 
logical factors that have not always been well identified. This confusion is 
understandable in view of the difficulties of studying marine fish popula- 
tions which are not subject to direct observation. Problems associated with 
the measurement of productivity of certain areas or fish stocks so far have 
not been susceptible to controlled experimentation and their study has neces- 
sarily been confined to field observations which, even though they may be 
well designed, still leave many essential varial)les unmeasured. 

The burden of solving the problems of conserving, developing, and en- 
suring the highest perpetual yield of our fisheries falls on the shoulders of 
our fisheries biologists. The purpose of fisheries biologists should be to 
determine the relationship of fish populations to tlie fisheries that utilize 
them and if necessary, either to discover metliods of adjusting the fishery 
to the productive capacity of the fish population or to develop techniques 
of increasing productivity to satisfy the needs of tlie fishery. Unfortunately, 
adjustments have sometimes l^een attempted without knowing tlie i-elation 
of the fishery to the fish. In other cases it lias been maintained, again with- 
out convincing evidence, that natural causes control the size of the fish 
populations and that adjustment of the fishery w^ould serve no purpose. 
Complete and careful examination of all published data on these problems 
is not possible here, but the resolution of opposing views on this point wall 
probably control the course of much fisheries work in the near future. 


Our principal West Coast fisheries may be divided roughly into three 
types according to the nature of the species involved and of the problems 
associated with their productivity. These are first the oceanic fisheries, 
primarily for the various species of tunas. In a second group are the coastal 
fisheries which include such species as the sardines, mackerel, anchovy, rock- 
fish, halibut, and other bottom fishes. The third group includes the ana- 
dromous fishes which in commercial value and volume are dominated by 
the salmon. The most valuable fisheries are the tuna and salmon although 
the largest in volume was that for the sardine. 

The condition of each one of these major fisheries is different, as are 
the problems and possibilities for their future development and use, al- 
though some nonbiological problems affect them all. Most noteworthy of 
these is that of territorial waters and the problem of ownership of marine 
fisheries. Although this may be considered primarily a legal and political 
problem, most political boundaries do not agree with biological boundaries 
and the eventual solution of even these problems will have to be harmonized 
with the basic biology of our fisheries. For this reason international treaties 
have been created to solve some of the more immediate international prob- 
lems of conservation and use of the lialibut, salmon, tuna, and other fishes. 
These problems are made more difficult by the fact that marine fishes are 
taken in offshore waters which traditionally have been considered open to 
men of all nations. IVIajor adjustments in some recent concepts of terri- 
toriality and national rights will be necessary for satisfactory solution of 
either the biological or political aspects of our marine fisheries problems. 

Oceanic Fisheries 

Those problems which are important in oceanic fisheries are the defini- 
tion of the biological limits of stocks, some of which appear to range the 
whole Pacific, the determination of potential yield of these wide-ranging 
stocks, and adjustment of the fishery to it. If our fishermen are to share 
in this resource we will have to determine where these oceanic fishes can 
be taken in commercial quantities and will have to develop fishing tech- 
niques that will enable our men to compete with the government-supported 
fishermen of other countries. In this field for the first time in history we 
find fishermen, fisheries biologists, and physical and biological oeeanogra- 
phers working together in the well co-ordinated program of the Pacific 
Oceanic Fishery Investigations which operates out of Hawaii. 

The results of this program are striking. The existence of large stocks 
of tuna in the north equatorial Pacific Ocean has been established, and the 
proven ecological relationships between tuna and the ocean currents indi- 
cate the possibility that other stocks of tuna exist in the South Pacific, 
Atlantic, and Indian oceans. The oft-stated claim of biologists that tropi- 
cal ocean waters are generally unproductive seems to have been pretty w^ell 


exploded, though apparently it has not yet been abandoned. The possibili- 
ties of developing large tuna fisheries appear to be certain and their realiza- 
tion will depend primarily on developing more economically feasible 
methods of capturing and preserving this most perishable food far from 
home ports. 

The work of P. O.F.I, is now being extended north of the Hawaiian 
Islands and will probably be co-ordinated with other programs, including 
that of the North Pacific Fisheries Treaty between the United States, Japan, 
and Canada. One result should be the expansion and stabilization of our 
West Coast albacore fishery, w^liich has so far been confined merely to the 
eastern fringe of the range of that species. 

The need for conservation of our oceanic species has already been indi- 
cated by the Inter- American Tropical Tuna Commission. It has found that 
probably the relatively local stock of Yellowfin tuna fished by the California 
fleet off the Pacific Coast of Central America and northern South America 
has reached its maximum level of yield and will not stand a greater take. 
This does not seem to hold for the Skipjack tuna fished in the same areas 
which appears to be capable of an even larger production than its present 
80 to 125 million pounds per year (Pacific Fisherman, November 1954, page 
53). These conclusions must be accepted with reservations until the extent 
and magnitude of the tuna stocks involved are evaluated. Such a survey 
will present great problems and will require a much greater expenditure 
of funds and energy than has .so far been available. 

The most sensible approach to conserving these oceanic fishes appears to 
be through international treaties, several of which have already demon- 
strated their soundness and efficiency. For some years the United States 
tuna fisheries have faced problems arising from differing interpretations 
of the extent of territorial waters, and from the controls certain countries 
have imposed on fisheries where both the bait and tuna fish are caught. 
In 1945 President Truman proclaimed a new policy for conserving oceanic 
fisheries (Proclamation 2668, 1945). Subsequently Peru, Chile, and Ecua- 
dor have given force to their own interpretation of this act by ])roclaiming 
ownership of the sea to a distance of 200 miles from their coasts. While 
the United States, as well as other countries, has not recognized these 
claims, there seems to be a move toward extending territorial waters, espe- 
cially by smaller nations off whose shores the ships of other countries fish. 
Some answer to these claims of extended territorial controls must be found 
which will protect both the rights of the adjacent countries, and of the fish- 
ermen who have developed the fisheries, and at the same time will protect 
the fish. 

Coastal Fisheries 
Our coastal fisheries are concerned with a wide variety of species and 


fishing techniques as well as of problems. In value and volume the sardine 
{Sardino2)s caerulea) fishery has dominated this field, which includes the 
now-famous halibut fishery of the North Pacific. The condition of these 
fisheries cannot be examined in this brief survey but may be indicated by 
an outline of three typical situations. 

Sardine Fishery 

The sardine fishery of the Pacific Coast did not begin to develop until 
after World War I and only grew to its full volume after 1930 with the 
development of the fish meal and oil markets. Heaviest landings occurred 
in the season of 1936-1937 when offshore floating reduction plants had added 
their processing capacity to that of the shore plants. After the elimination 
of the floating plants during the next few years, the capacity of both the 
shore stations and of the fishing fleet increased; but even though the fleet 
increased steadily in efficiency during these years, the landing record of 
1936-1937 was never again equaled. A decline in average catch per boat 
began in the 1935-1936 season but it seems to have been relatively stable 
from 1937 to 1945, after which it again dropped sharply (Clark and Daugh- 
orty, 1950). This irregular decline in catch per boat was accompanied by 
the progressive disappearance of the older age groups and thus the fishery, 
first from the coast of British Columbia in 1946, then from Washington 
waters in 1948, and from Oregon in 1949. This was finally followed by the 
disastrous years of 1951 and later off^ San Francisco and Monterey. Al- 
though production continued in 1951 and 1952 off San Pedro with a greatly 
increased fleet, which was augmented by boats which had followed the fish- 
ery southward, even this last stronghold of the sardine gave out in 1952 
and 1953. By this time even the most optimistic should have begun to wonder 
if the sardine schools had become scarce and to doubt that they had only 
moved farther offshore or into deeper water where boats and nets could 
not find them. 

The decline in the fishery was associated with a deficiency in the num- 
bers of young fish. The last very-large-year class was produced in 1939 
(Felin and Phillips, 1948) and the production of young since then has not 
been sufficient to maintain the fishery. Because of the lack of large year- 
classes, and also because of correlations that have been found between ocean 
conditions and size of sardine year-classes, some have been led to deprecate 
the role played by the commercial catch in producing the present situation. 
It is interesting that in a brief discussion of this problem Schaefer (1954) 
was able to show, on the basis of a theoretical relationship between fish 
abundance and fishing yield, based on Verhulst's equation of growth, that 
the fishery had been a primary factor in the decline of sardine abundance. 
Unfortunately, although much money and effort have been expended inves- 
tigating the Pacific sardine, no one agency has had sufficient funds or au- 


thority to develop a broad enough program to cover the entire range of 
the species, which extends from southeastern Alaska to Mexico. As a result, 
many obscure points still remain in the sardine picture which even the 
present augmented efforts are unlikely to clarify. 

There is no doubt that major fluctuations in abundance of sardine have 
occurred in the past as a result of the varying survival of individual year- 
classes. However, it seems presumptuous to ignore the effect that the great 
reduction in abundance of mature fish, which formerly supported the fish- 
ery north of California, must have had on the production of young. 

Even though variations in abundance may occur naturally, it is prob- 
able that the level around which the sardine population varies has been 
lowered by the fishery until in the last few years the lower points in the 
cycles of abundance have become disastrous for the industry. It is also 
probable that with flexible control of the fishery these lower levels of abund- 
ance might be raised by the maintenance of a larger residual population. A 
larger population would also increase the probability that abundant year- 
classes would be produced. 

Bottom Fisheries 

A different type of problem is encountered in the coastal fisheries for 
bottom-dwelling species. While some species such as the halibut are now 
fished at about the limit of their productive capacity, there are other species 
which could produce much larger quantities of fish, especially along the 
coast of Alaska. An example is the true cod, Gadus macrocephalus. This 
species is taken along the entire coast from Oregon to the Bering Sea 
where in years past a considerable fishery was developed for salt cod. As 
the market for this product disappeared, the Bering Sea cod fishery had 
ceased to exist. In the last two years, however, the industry has introduced 
fish into the market in a new form known as "fish sticks." This new product 
has increased rapidly in popularity and has created a new market for fish 
which are suitable for conversion into this form. Cod seems to be ideal for 
this purpose and as a result, landings have increased with the demand until 
this year an estimated 12,000,000 pounds have been landed in Washington 
alone as compared with 7,000,000 pounds in 1950. Obviously the fish are 
available to the fishermen and all that is required is a market that will 
purchase their catch for enough to make operation worthwhile. A similar 
increase in rockfish landings resulted from the increased popularity of 
"Ocean Perch" filets in the last few years. 

The opposite picture is seen in the rapid rise and fall of the shark fishery 
off this coast. Beginning about 1940 the demand for vitamin A rose rapidly. 
Dogfish shark or grayfish were found to have liver oil several times richer 
in this vitamin than codfish-liver oil. The demand was increased by the 
wartime closure of Norway as a source of the latter. Another species of 


shark — the "soup fin," Galeorhinus zyopterus, was found to yield even 
greater quantities of vitamin A than the dogfish. Landings of these sharl^s 
rose so rapidly with the price of the oil that Kipley (1946) found evidence 
of a sharp decline in soupfin abundance by 1943. With the development 
of synthetic vitamin A after the war and cheaper foreign sources of fish 
oil the American landings declined and the fishery has now been practically 
abandoned. Information obtained from fishermen shows that both soupfin 
and dogfish sharks are now abundant off our shores but with no market for 
cither the oil or carcasses, except for reduction, these fish are generally 
avoided if possible or are discarded at sea. Landings fell to 1,287,000 
pounds in 1953 as compared with a high of almost 53 million pounds in 1944. 
There is a large potential sui>ply of these as well as several other species, 
l)ut some method must be found of using them that will permit an adequate 
return to the fishermen. If such a market could be developed and a good price 
offered it has been estimated that a stable production of between one and 
two billion pounds could be produced each year from all of tlie l)()ttom 
species (Chapman, 1943). 

With limited markets and an abundant supply of raw produce it is not 
surprising to find wasteful practices where the fishermen are sent out with 
orders to bring in limited amounts of only those species the processors can 
profitably market at that time. Catch limits are normal in our trawl fish- 
eries and combined with regulations which forbid the use of food fish for 
reduction purposes result in the discard of as much as 70 per cent or more 
of the total catch taken in trawls (Pruter and others, 1953). While some of 
the discarded fish are below minimiun size limits set by state law in the 
state of Washington, many are of species that cannot be marketed at pres- 
ent. Such species as the raffish, Hydrolagus coJJiei, and hake, Merluccius 
productus, though available in large quantities, have never been marketed 
on the West Coast except to reduction plants. Develo])ment of uses and 
consequently markets for fish that are now discarded would not only elimi- 
nate substantial waste; it would also avoid a possible disturbance of the 
interspecies balance, wliich might add to the strain already imposed on more 
valuable species by selective fislieries. The wasteful practices engendered by 
the lack of markets is evidence of the relatively undeveloped state of some 
of our West Coast fisheries, biologically, technologically, and economically. 
As a result of an abundance of ]n-otein and with a predominant indifference 
to fish as an article of diet, the productive capacity of our fishing fleet is not 
being utilized, and little incentive has been offered to our technologists as 
yet to develop uses for "nuisance" species. If survival of our fishing indus- 
try can be considered a worthwhile objective (and I believe it is one) condi- 
tions call for an intensified interest in technology to increase further the 
efficiency of our fishiuQ-, processing, and marketing. 

Full utilization of the fishing fleets now in existence in the U.S. inevit- 


ably would raise the problem of management. The only two examples of 
successfully managed stocks of marine and anadromous fish are to be found 
on the North Pacific Coast of America. Both of these fisheries, the halibut 
and sockeye salmon of the Fraser River, were placed in the hands of interna- 
tional commissions and both operate under treaties between the United 
States and Canada. So much has been written concerning the outstanding 
work of the International Pacific Halibut Commission and the International 
Pacific Salmon Fisheries Commission that it will not be reviewed. In spite 
of the marked success of their management there still remain a few biologists 
who either maintain that these two outstanding examples are the result of a 
chance agreement of management practices with a favorable natural in- 
crease in stocks, or that these two examples are quite different from any 
other fisheries problem. These two successful cases of fisheries management 
do, as a matter of fact, differ markedly from most other attempts, in that a 
sound foundation of biological knowledge of the species involved was in each 
case accumulated before management was undertaken, and in the Fraser 
River before work was begun on the rectification of unfavorable environ- 
mental factors. In both fisheries, too, this sound background has been used 
as a basis for regulation and, so far as possible, a system of continuous 
checks on the biological results of these regulations is maintained. It may be 
predicted with confidence that both of these fisheries will continue to main- 
tain a high level of production as long as regulations are soundly based and 
continuously readjusted according to constant observation of reaction of the 
stocks. It may be predicted just as confidently that neglect of that biological 
background will lead to the eventual failure to maintain the fisheries. 

The success of these two ''experiments" in fisheries regulation indicates 
that, with sufficient knowledge of our fish populations, a type of farming 
can be imposed upon our fisheries, in some ways comparable with present 
management of forests for continiious yield. Decisions as to kind or extent 
of knowledge that will furnish a sufficient basis for successful farming of 
this nature must not be warped by blind adherence to previously conceived 
hypotheses, either as to the need for management or the techniques to be 
used in determining this need. 

Anadromous Fishes 

The International Pacific Salmon Fisheries Commission has demon- 
strated that if the salmon are given free access to their spawning grounds, 
and if the fishery is regulated to permit adequate escapement of all parts 
and especially the peak section of each race, the runs can be rebuilt and 
maintained. No dams block access to the spawning grounds of the Fraser 
River Sockeye and the natural and artificial blocks to their movements 
have been removed. This condition is quite different from that found on 
most salmon rivers in the western United States, a typical example of which 


is the Columbia liiver. Kecently publicized plans for dams on the Copper 
and Yukon rivers in Alaska portend that conditions like those on the Colum- 
bia may develop on all salmon streams. 

The use of water for irrigation and power developed gradually on the 
Columbia until the growing industrialization of the Northwest and the in- 
creased need for power resulted in a great surge of dam building during 
the last twenty years. The gradual development had not been met with 
sufficient funds and staff to enable fisheries biologists to solve even the cur- 
rent problems, and people working in that field during the late twenties 
and thirties were fortunate to prevent the complete loss of most of our major 
West Coast runs. With the help of federal aid funds during this period, 
definite progress was made in screening irrigation ditches to prevent the 
loss of young salmon on their way to sea. Some progress was also made in 
building fish ladders to enable salmon to surmount dams but study of the 
basic biology and reaction patterns of both young and adults as well as of 
the details of fresh-water life history lagged behind, largely because of an en- 
forced economy in appropriations for fisheries work acceded to by an unin- 
formed public. The sudden increase in rate of dam construction as well as 
in the magnitude of projects sponsored by the federal government caught 
the fisheries biologists with neither sufficient facts to solve the problems 
which suddenly confronted them nor with sufficient funds to gather those 

One result of the struggle between power and irrigation interests on the 
one hand and fisheries interests on the other has been a gradual though very 
grudging acceptance of the fact that the fish do have a place in our rivers 
and that something should be done to save this major source of food. The 
principal problems of handling both upstream and downstream migrating 
salmon at dams were out of reach of the relatively small appropriations of 
the fisheries agencies until in 1951 when the United States Army Engineers 
agreed to support a program devised by salmon biologists of the Northwest 
in an attemjit to find out how to adjust conditions at dams to the needs of 
the salmon. Participating in this program are seven different fisheries agen- 
cies in the Northwest. The program now includes the study of practically 
every phase of the design of the fish ways, the relation of height of dam 
to the probable success of salmon surmounting it, and the effect of flooding 
spawning beds, including a more thorough survey of the present location of 
spawning and of areas that will be flooded by proposed dams. The down- 
stream migration period of the young of different species and their distri- 
"bution in the streams and reservoirs are also being examined in different 
areas, so that it may be possible to apply new techniques being developed 
for guiding the young around dams or through safe passages and thus re- 
duce losses that now occur in some turbines and at some spillways. 

The success or failure of this program may well determine whether or 


not we will continue to have our major runs of salmon. Its imi)ortance may 
be appreciated if we consider that the Columbia Kiver in 1883 produced 
629,400 cases of chinook salmon which at present market prices would be 
worth over $23,000,000 (Pacific Fisherman Yearbook, 1954). During 1953 a 
total of only 151,000 cases of all species of salmon were packed in this area. 

The need for this w^ork becomes clear when the life history of the salmon 
is examined. Omitting details and differences between species, all recpiire 
unpolluted water of moderate temperatures (45° to 55° F.) in which to 
spawn, in which the eggs can develop, and the young can live and grow to 
migrating size. When this time is reached, the young must have free access 
to the sea where they put on the major portion of their growth. On reach- 
ing maturity the adults return to the same stream in w^hich they were 
deposited as eggs and unless they have free access to that stream, or unless 
conditions very similar to their native stream are provided, that population 
of salmon will be eliminated. Success of the runs recpiires a sufficient area 
of suitable spawning gravel covered by clean cool running water, a suffi- 
cient area for rearing the young (in those species that spend a year or more 
in fresh water), and free access for both upstream and downstream mi- 
grants. To provide these conditions it will be necessary in some cases to 
modify or compromise some of the plans now outlined for the use of the 
streams for power and irrigation. So far most of the modifications and 
compromises have been made at the expense of the salmon. 

Great cumulative losses in spawning and rearing areas for salmon have 
been suffered in streams of the West Coast of the United States. It has been 
estimated that between 50 and 75 per cent of the original salmon spawning- 
grounds in the Sacramento-San Joaquin river systems have been lost to these 
species by impassable dams (Van Cleve, 1954). Some 70 per cent of the 
Columbia River watershed has also been lost (Washington State Department 
of Fisheries, 1947) and unknown but additional losses will occur as more 
sections of the river are flooded by dams now being built or planned for that 
stream. Every effort is being made to alleviate these losses, not only by 
attempting to eliminate or at least reduce the losses of adults and young at 
dams, but also by opening up productive stream areas previously unavail- 
able to salmon because of natural blockades (Washington State Department 
of Fisheries, 1954) and further, by attem]itin'': to develop efficient methods 
of artificial propagation to counteract the continued loss of spawning areas. 

The salmon versus dams problem is still far from solution but at least 
it is being attacked vigorously and on a sufficiently broad front that there 
is now a very good chance that at least a sustained though limited produc- 
tion of salmon can be anticipated in streams that are developed for other 

T^rospects are bright for obtaining a considei-able increase in production 


of fish from the sea but the need for preserving the productive capacity of 
our currently heavily exploited stocks must not be forgotten. The combined 
efforts of fisheries biologists, hydrographers, and marine biologists should 
result in the development of potentially large oceanic fisheries but consider- 
able technical improvements will be required to enable our fishermen to 
operate economically on these widely scattered stocks. 

On the other hand, the development of adequate markets and more ad- 
vanced and efficient marketing methods should result in the development 
of a large fishery along the west coast of North America on stocks which 
are essentially not now utilized. The best use of this fishing region requires 
some new means of using many species that cannot now be marketed. 

With the full development of our West Coast fisheries it will be neces- 
sary to anticipate the need to develop conservation measures along such 
lines as have been proven so effective by the International Pacific Halibut 
Commission in their rehabilitation of the north Pacific halibut stocks. 

The maintenance of our anadromous fish runs will involve the solution of 
the many problems encountered in handling upstream as well as down- 
stream migrants at dams. Some means will also have to be found to com- 
pensate for spawning and rearing areas that will be, or have already been 
lost by flooding or by closing them to salmon by obstructive dams. Opening 
new streams by laddering impassable natural barriers and developing more 
efficient techniques of artificial propagation are both supplementing efforts 
to preserve the original salmon spawning grounds. Conservation measures 
similar to those used so effectively by the International Pacific Salmon 
Fisheries Commission can insure continued productivity of our salmon, pro- 
vided the fresh water environment can be preserved. 

Finally, settlement of the widely divergent views on ownership of deep 
sea fisheries must be reached if their development and conservation are to 
be successfully accomplished. Treaties appear to be the best means devised 
so far to protect and develop our fisheries through the co-operative effort 
of interested nations. 



Chapman, W. M. 

1942. The latest fisheries of Washington and Alaska. California Fish and Game, 
vol. 28, no. 4, October 1942. 

Clark, F. N., and A. E. Daugherty 

1950. Average lunar month catch by California sardine fishermen, 1932-33 

tlirough 1948-49. California Division Fish and Game, Fish Bulletin 
no. 76, 1950. 

Daniel, Hawthorne, and F. Minot 

1954. The Inexhaustible Sea. Dodd, Mead and Co., N. Y. 1954. 

Dannevig, Alf. (United Nations Report) 

1951. Propagation and transplantation of marine fish in Europe. UN8CCUR 

Proceedings, vol. 7, Wildlife and Fish Resources, 1951. 

Felin, F. E., and J. B. Phillips 

1948. Age and length computations, etc. California Fish and Game, Fish Bulle- 
tin, no. 69, 1948. 

Food and Agriculture Organization of the United Nations 

1953. Improving the fisheries contribution to world food supplies. F.A.O. 
Fisheries Bulletin, vol. 6, no. 5, 1953. 

LeGall, Jean (United Nations Report) 

1951. The present world problem of sea fisheries. UNSCCUR Proceedings, 
vol. 7, Wildlife and Fish Resources. 

Pacific Fisherman Year Book, 1954, page 117. 

Pacific Fisherman — November, 1954, page 53. 

Proclamation 2668 

1945. Policy of United States with respect to coastal fisheries in certain areas 

of high seas, proclamation. Sept. 28, 1945. (Proclamation 2668) 
Federal Register, vol. 10, no. 193, October 2, 1945, p. 12304. 

Pruter, a. T., D. L. Alverson, J. Jurkovich, and A. T. Palmen 

1953. Progress Report on Otter-Trawl Studies, April through June, 1953. 

Marine Fisheries Investigations, Progress Report, Washington State 
Dej)artment of Fisheries, April-June, 1953. 

Raymont, J. E. G. 

1947. A fish farming experiment in Scottish sea locks. Journal Marine Re- 
sources, vol. 6, no. 3, pp. 219-227. 

Reay, G. a. 

1954. The ocean as a potential source of world food supply. Food Technology, 

vol. 8, no. 2, Feb. 1954. 

Ripley, W. E. 

1946. The soupfin shark and the fishery. California Division of Fish and Game, 

Fish. Btilletin no. 64, 1946. 



1954. Some aspects of the dynamics of populations important to the manage- 
ment of commercial marine fisheries. Inter-American Tropical Tuna 
Commissi07i Bulletin, vol. 1, no. 2, 1954. 

Thompson, Harold (United Nations Report) 

1951. Latest fishery resources and means for their development. UNSCCUR 
Proceedings, vol. 7, Wildlife and Fish Resources. 

TitESSLER, D. K., and J. M. Lemon 

1D51. Marine Products of Commerce. 2nd Ed.— 1951, Reinhold Publishing 

Van Cleve, R. 

1945. Program of the Bureau of Marine Fisheries. California Fish and Game, 
vol. 31, no. 3, July, 1945. 

Washington Department of Fisheries and Oregon Fish Commission 

A program of rehabilitation of the Columbia River Fisheries, prepared 
by the Washington Department of Fisheries and Oregon Fish Com- 
mission. 1947. 




Fourth Series 
Vol. XXVIII, No. 13, 441-527, figs. 1-26 February 21, 1956 



Imperial College of Tropical Agriculture 
Trinidad, British West Indies 

This report deals witli three collections of insects from China. The largest 
of these, comprising 241 s])ecimens, was made by Dr. J. L. Gressitt in Sze- 
chiian and western Hupeh; tlie second (201 specimens) was made by Mrs. 

D. E. Wright in Chekiang, mainly in the neighborhood of IMokansan; the 
third (156 specimens) of material from other areas, chiefly in Kwangtnng 
and around Hong-Kong, was made by various staff members of Lingnan 
University, including, in addition to the foregoing, W. E. Hoffman and 

E. Ii. Tinkham. The whole of the material, including all type specimens, has 
been returned to the California Academy of Sciences. 

The writer tenders his warmest thanks to Dr. Edward S. Ross, curator 
of the insect collection of the Academy of Sciences, and to Dr. .J. L. Gressitt 
for the privilege of studying this material. He also pays a well-merited 
tribute to the collectors whose personal efforts have made such a useful 
sample of this little-known fauna available for taxonomic study. 

Collections of Chinese fulgoroid homoptera are too small and too scat- 
tered, and reports on them too insufficiently detailed, to afford adequate 
opportunity for any careful study of speciation in this important zoogeo- 
graphical area. The object of the present report, therefore, has been limited 
to presenting the data in a usable manner; little attempt has been made 
to collate existing records, or to review and analyze the fauna. 

Many facts of interest have emerged from a study of the present collec- 
tions: the first record of Ptoleria in Asia north of the Himalayas; the 
substantial extension of the known distribution of Nesopompe tsoui Muir 
westward to Hupeh and northeastward to Japan; the establishment of 
new western limits for the distribution of Kinnara fumata (Mel.), Los- 




[Peoc. 4x11 Ser. 

baTiosia hakeri Muir, Diostvomhus politus Ulil., Zoraida kirkoldyi Muir, 
Pamendanga sauteri Muir, and Blegatropis formosana (Mats.); of a new 
eastern limit for Knmendaka nif/romacuhita (Dist.), and a new northern 
limit for VivaJia facialis (Dist.). Two informative captures were those of 
Caristianus ulysses Fenn. and Ommatissus lofouensis Muir. The record of 
the former, described from Bornean material, in Yunnan, has not only 
extended the known range of this species, but has strengthened the case 
for assigning a new species of achilid, taken in the same general area, to Za- 
tJiauma, a hitherto monobasic genus known only from Borneo. The identifica- 
tion of Ommatissus lofouensis has served to corroborate the accuracy of Muir's 
generic assignment, which was doubted by Melichar (1914:214). 

In general the fulgoroid fauna of south and southwestern China appears 
to be made up of generic elements of the palaearctic fauna mixed with 
others from the Burman, Indochinese, and Indonesian faunas. This was 
noted by Noualhier (1896: 251), and there is little more to be said on this 
point today. An interesting consideration has been brought to the front 
by the known great antiquity of the conifer Metasequoia glyptostrohoides. 
A new achilid species, assigned below to Magadha Distant, is associated, 
probably rather loosely, with this tree, and, as is general in the family, 




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undoubtedly spends its nynipluil life below bark and inside crevices of 
rotting branches and tree trunks. The genus Magadha was described from 
northern India and is represented in Ceylon, Assam, and Formosa. It 
is clearly headquartered in southeastern Asia, and there appears to be no 
o])vious reason why species of this genus, both in the nymphal and adult 
stages, should not have been associated with Metasequoia through an appre- 
ciable length of geological time. 

Family CIXIIDAE Spinola 


(Adapted from Muiv) 

(1) (2) A subantennal process present on genae Borysthenes Stal 

(2) (1) No subantennal process (3) 

(3) (4) Sc, R, and M in tegmina arising separately from base, not forming a com- 

mon stalk Andes Stal 

(4) (3) Sc, R, and M not arising separately from basal cell, two or more united in 

a common stalk (5) 

(5) (6) Tegmina in repose steeply tectiform, with apical margins meeting or 

nearly so, body laterally compressed. Ovipositor with valvulae promi- 
nent, more or less ensiform, curved, often accommodated in a longi- 
tudinal sulcus (") 

(6) (5) Tegmina in repose shallowly tectiform and with apical margins not 

apposed; abdomen not laterally compressed, sometimes moderately 
dorsoventrally depressed (9) 

(7) (8) Vertex distinctly angularly emarginate at apex Kirbyana Dist. 

(8) (7) Vertex truncate at apex, distinctly broader than long Ptoleria Stal 

(9) (10) Tegmina with the first (basal) fork of M closer to the fork of M3 and M4 

than to the fork of Mj and Mo Mnemosyne Stal 

(10) (9) First fork of M in tegmina closer to forking of Mj.o than to forking of 

Mo,^ (11) 

(11) (12) Mesonotum with live carinae Oliarus Stal 

(12) (11) Mesonotum with three carinae (13) 

(13) (14) Carina between vertex and frons obsolete, median frontal carina absent 

or only present on apical portion (15) 

(14) (13) Carina between vertex and frons and median frontal carina distinct.... (17) 

(15) (16) Tegmina with M 5-branched at apex, Cuj 3-branched; wings with CU| 

3-branched; frons as long as discal portion of clypeus or longer 

Kuvera Dist. 

(16) (15) Tegmina with M 4-branched at apex, Cui 2-branched; wings with Cu^ 

2-branched; frons distinctly shorter than discal portion of clypeus 

Betacixius Mats. 

(17) (18) Pronotum laterally carinate between eyes and tegulae....Jfocrocixij/s Mats. 

(18) (19) Pronotum not laterally carinate, lateral discal carinae following hind 

margin of eyes Cixius Latr. 



Genus Andes Stal 
Stal, 1866:166. Logotype, Andes unchilatus Stal, 1870:747. 

Andes uncinatus Fennah, new species. 
(Figure 1, A-C.) 

Male: length, 3.5 mm.; tegmen, 6.2 mm. 

Oehraceous; head, except lateral frontal margins, pronotum inmiediately 

Fig 1. Ancles unci7iatus, new species: A, tegmen; B, aedeagus, left side; C, anal 
segment, hind margin of pygofer, and right genital style, side view. Abides otlirepte, 
new species; D, tegmen; E, aedeagus, right side; F, anal segment, hind margin of 
pygofer, and right genital style, side view. Andes noctua, new species: G, tegmen; 
H, aedeagus, right side; I, anal segment, hind margin of pygofer, and right genital 
style, side view. 


behind eyes, disc and disc of mesonotum, and abdominal tergites, fuscous; 
lateral fields of mesonotum brown. Tegmina translucent, basal area to level 
of union of claval veins, a narrowing curved fascia from middle of costal 
cell to near apex of clavus, three short stripes more or less in line from 
distal stigma to apex of clavus, two wedges near apical angle, and a suffu- 
sion across apical cells fuscous, a faint band from node to apex of clavus, 
and a suffusion across membrane yellow. 

Anal segment moderately long, bilaterally symmetrical, anal foramen at 
distal third, apical margin rounded-truncate, latero-apical angles rounded, 
not produced. Pygofer with lateral margins strongly convex, medioventral 
process triangular, acute at apex, with a short spine on each side at base. 
Aedeagus tubular, an irregular denticulate ventral keel, swollen on left 
side near base and produced in short curved tooth at base, directed basad, 
distally a foliate lobe on left enwrapping distal left side and apex, flagellum 
comprising a long slender ])lade-like spine, slightly denticulate, rising at 
apex, curved to right then cephalad, completely looped distally, a broad 
and short lobe on left with a strongly sinuate upper margin, a shortly 
tapering membranous lobe dorsally. Genital styles of subequal width 
throughout, rectangulately bent at distal third, obliquely truncate at apex. 

One male (the type), 700-900 m. Kan-Lin-San, Lien-p'ing District, 
Kwangtung, S. China, J. L. Gressitt, April 23, 1940. This si)ecies dift'ers in 
genital structure from any of those reviewed by Muir (1925), and in teg- 
minal markings from any known to the writer. In the latter character it 
perhaps most closely approaches Andes migrntorius (Dist.). The spines 
laterad of the medioventral process of the pygofer are exceptional. 

Andes othrepte Fennah, new species. 
(Figure 1, D-F.) 

Male: length, 3.6 mm.; tegmen, 5.0 mm. 

Castaneous-brown, lateral carinae of frons and clypeus, an oblique stripe 
between eye and margin, antennae, pronotum, tegulae, rostrum, and legs 
testaceous to stramineous. Tegmina subhyaline with greyish powdering; 
a broad even band from basal cell to middle of clavus and a broad oblique 
band from costal cell basad of stigma to apex of clavus, dull yellow; a 
short stripe bounding latter band on its basal margin, and a more or less 
complete stripe bounding its distal margin, an obliquely L-shaped mark 
across middle of clavus, a spot just basad of stigma and a slightly larger 
s])ot across all Cuib and apex of clavus fuscous; stigma and membrane pale 
brown, veins concolorous, minutely granulate with fuscous on corium. 
Wings translucent, powdered sordid white, with fuscous veins. 

Anal segment of male moderately long and rather broad, lateroapical 
angles acutely produced ventrad, anal foramen slightly distad of middle. 
Pygofer laterally strongly convex on hind margin, medioventral process 


siibequilaterally triangular. Genital styles elongate, in profile angulately 
bent at basal third, in posterior view distal portion wider than basal, of 
subequal width throughout, shallowly rounded at apex. Aedeagus tubular, 
a triangular flange on ventral surface near base, directed to left; on right 
ventrally, two-thirds from base, a stout spine directed to left below aedeagus 
then dorsad; slightly distad of this, ventrally on right, a short stout spine 
directed to right, then dorsad; at apex a long, rather slender, sinuate 
process directed cephalad above aedeagus then curved to left; flagellum 
with two slender unequal spines arising near base, the dorsal slightly 
curved, the ventral in form of a shallow spiral. 

One male (the type), Hong Kong Island, Aug., 1933, W. E. Hoffman. 
This species recalls A. unduJatus Stal, but differs from the female holotype 
in tegminal markings, and from the male, as identified by Muir, in the 
shape of the genitalia. 

Andes noctua Fennah, new species. 
(Figure 1,G-I.) 

Male: length, 3.2 mm.; tegmen, 4.5 mm. Female: length, 5.1 mm.; teg- 
men, 6.0 mm. 

Dark fuscous: a spot on sides of head above eyes piceous; a series of 
frequent and fairly even sublinear interruptions on lateral margins of 
vertex and frons, and a ring round base of antennae pallid stramineous, 
antennae, lateral fields of pronotum, sometimes interruptedly, rostrum, 
lower side of thorax and legs stramineous, more or less suffused fuscous. 

Tegmina translucent, powdered ash-grey : a narrow irregular stripe from 
basal third of costal cell to union of claval veins, a zig-zag narrow stripe 
subparallel to preceding from middle of costal cell to claval suture at apical 
fifth, a broader oblique band from apex of costal cell to subapical trans- 
verse line at M, a band overlying transverse veinlets from M to apex of 
clavus, apical cells fuscous; a spot at distal edge of stigma and on marginal 
dilation at apex of clavus fuscous-piceous. Wings translucent, distally suf- 
fused fuscous, powdered grey, veins fuscous. 

Anal segment elongate, distally shallowly deflexed, apical margin very 
short, slightly excavate, lateroapical angles scarcely produced, anal fora- 
men in distal quarter. Pygofer with laterodorsal angles obtuse but well- 
defined, lateral margins convex, medioventral process triangular, flanked 
on each side at base by a distinct spine-like process. Genital styles short, in 
profile subrectangulately bent at middle, slightly dilated and rounded- 
truncate at apex. Aedeagus tubular, a small flange on right near base, 
directed laterad and minutely denticulate on margin; mesad of this a 
triangular vertical flange directed ventrad; a broad submembranous 
trough-like lobe, attached at apex, free at cephalad end, loosely enwrap- 
ping aedeagus for most of its length, this lobe truncate at anterior margin 


on left, on right (ventrolaterally) produced in three short broad lobes, 
the middle lobe denticulate on margin. A long strongly sinuate spine 
arising at apex of aedeagus directed cephalad above aedeagus; flagellum 
short, unornamented. 

Two males (one the type) and 8 females, 1,000 m. Suisapa, Lichuan 
District, W. Hupeh, China, July 23, 1948, Gressitt. 

In gross structure this species appears to be allied to A. marmoratus 
(Uhl.). The general conformation of the aedeagus is as found in A. mar- 
moratus, but the details of its outline are different, while the anal segment 
is relatively longer basad of the anal foramen and the genital styles differ- 
ently shaped at the apex. The tegminal markings are less clearly defined 
than in A. marmoratus and in the region of the claval suture distinctly 
different. The apical venation of R and M described by Muir is not found in 
the present species. 

Andes lachesis Fennah, new species. » 

(Figure 2, M, N.) 

Male: length, 4.0 mm.; tegmen, 5.2 mm. Female: length 4.1 mm.; tegmen, 
6.2 mm. 

Stramineous to testaceous, disc of frons and clypeus, genae and sides 
of head above eyes, except for an oblique pallid stripe, vertex, median por- 
tion of pronotum and mesonotum, castaneous-fuscous. Tegmina translucent 
pallid; inner angle of clavus, a broad band from basal cell between Cui and 
first claval vein to near apex, and two suffused areas on M and Cui fork 
respectively, yellow; an L-shaped mark near union of claval veins, an 
arcuate broad fascia from basal third of costa, where it is once interrupted, 
to clavus near apex, a spot at apex of costal cell and another just distad 
of stigma, a short stripe across subapical cell Mi-o and a spot on margin 
distad of claval apex, dark fuscous; membrane pale fuscous with two 
elongate-ovate pallid areas in apical cells. Veins concolorous with fine 
fuscous granulation. Tegmina sometimes suffused with pale fuscous in all 
lighter areas. "Wings sordid white, veins pale fuscous. 

Anal segment of male moderately long, relatively broad, with subparallel 
sides, apical margin excavate, lateroapical angles asymmetrically produced, 
anal foramen in distal half. Pygofer with posterior lateral margins broadly 
convex, medioventral process triangular. Aedeagus with a large vertical 
triangular keel below, which is continuous above with a loose wide shagreen 
sleeve which surrounds the aedeagal duct. This loose sleeve bears dorsally 
a short slender spine directed to right, and below this, a little more cephalad, 
a small elongate-triangular shagreen lobe. At apex of aedeagus a long stout 
sinuate spinose process directed cephalad above aedeagus, distally curving 
abruptly to right and ventrad. Flagellum coarsely shagreen or sub-fimbriate. 

Five males (one the type) and two females, Mokansan, Che-Kiang 



[Proc. 4tii Skr. 

Province, Se]it. 19-28, 1927, Mrs. D. E. Wright. A female taken at 1,000 m., 
Suisapa, Liehiian District, W. Ilupeh, China, Aug. 21, 1948, Gressitt, is 
placed here. 

In the denticulate process of the aedeagus this species can be compared 
with A. pseudohrunneus Muir and A. hrunniceps Muir; it differs from 
both in the deep ventral carina below the aedeagus, and in the position and 
shape of the spinose processes, in the shape of the anal segment and genital 
styles, and in the color pattern of the tegmina. 

Genus Ptoleria Stal 
Stal, 1859a: 321. Haplotype, Ptoleria arcuigera Stal, 1859a: 321. 

Ptoleria indica (Distant). 
Caneirona indica Distant, 1916:39. 

One female, 1,000 m. Suisapa, Lichuan District, W. Hupeh, Aug. 23, 
1948, Gressitt, is generally similar to Distant's type but with greater fus- 
cous suffusion over the hind portion of the membrane. 

Fig. 2. Cixius scrupeus, new species: A, aedeagus, left side; B, aedeagus, right 
side; C, left genital style; D, head and thorax; E, frons and clypens. Cixius 
phonascus, new species; P, aedeagus, left side; G, anal segment of male, right side; 
H, right genital style, inner aspect; I, left genital style, outer aspect. Cixius galcoJa. 
new species: J, aedeagus, left side; K, anal segment of male, left side; L, left genital 
style. Andes lachesis. new species: M, aedeagus, left side (process of left side also 
shown detached); N, anal segment, pygofer and left genital style. 


Genus Cixius Latreille 

Latreille 1804:310. Logotype, Cicada nervosa L. 1758 Syst. Nat. 10:437. 

Cixius phonascus Fennnh, new species. 
(Figure 2, F-I.) 

Male: Length, 5.0 mm.; tegmen, 6.1 mm. 

Vertex broader across base than long in middle, width of apex more 
than half of base, both transverse carinae convexly angulate. Dark reddish 
brown; carinae of pronotum and vertex and legs testaceous. Tegmina trans- 
lucent, more or less brown with pale areas; basal area usually suffused 
Ijrown, a diffuse fascia from union of claval veins transversely across to 
costa, a suffusion in membrane yellowish-brown, apical cells at base and 
apex fuscous, veins yellow. AVings infuscate distad of transverse veins, 
veins fuscous. 

Tegmina with 1 or 2 Sc veins at apex distad of stigma, 3 R's and 5 M's. 

Anal segment long, tubular, deflexed at anal foramen in apical quarter, 
apical margin deeply rounded. Pygofer short dorsally, long ventrally, with 
lateral margins obtusely angulately produced, medioventral process short, 
triangular. Genital styles short, expanding distally, curved through about 
110° at middle, angle at apex of dorsal margin slightly produced, acute. 
Aedeagus tubular, more or less straight, a moderately long spine arising 
ventrally at apex, shallowly sinuate, directed cephalad below aedeagus, on 
left near apex a pair of short curved spines, a short upcurved spine on 
right distally; flagellum more or less tubular, sinuate, directed cephalad. 

One male (the type), Loh Fau Shan, Poh-lo District, Kwangtung', S. 
China, April 6-8, 1934; one male and one female, Yaoshan, Lin-hsien, 
Kwangtung, S. China, May 6-10, 1924. 

This species differs from C. gravelyi Muir in its larger size, paler body- 
color, infuscate distal area of the wings, and in the shape of the genital 
styles; from C. laticeps Mete, in almost every detail of coloration. In teg- 
minal markings it is perhaps nearest to C. velox Mats., from which it is 
separated by the proportions of the vertex: from C. kuyanianus Mats, it 
dift'ers in tegminal markings and darker body -col or, and, apparently, in the 
shape of the apical portion of the genital styles. 

Cixius galeola Fennah, new species. 
(Figure 2, J-L.) 

Male: length, 4.9 mm.; tegmen, 6.0 mm. 

Vertex only slightly broader across base than long in middle line, width 
of apex equal to half width between basal angles. 

Testaceous; mesonotum castaneous. Tegmina yellowish hyaline; a faint 
suffusion along costal cell, brown; a bold broad spotted fascia obliquely 
across membrane from apical cell Cui to apex of Sc; veins yellow. Wings 


hyaline, a broad suffusion along margin extending inward to cover apical 
cells, fuscous, veins concolorous. 

Tegmina with 2 Sc veins at apex distad of stigma, 3 R's, 5 M's, Sc + R 
fork basad of Cui fork, which is basad of union of claval veins. 

Genitalia similar to preceding, distance between anal foramen and lateral 
margin shorter than in preceding. Aedeagus tubular, straight, a very short 
stout curved spine arising ventrally at apex, curved to right, on left side 
near apex two spines, the ventral spine about half as long as the dorsal, 
strongly curved anteriorly and cephalad, dorsal spine sinuate, directed 
anteriorly, on right side near apex a moderately long spine directed ceph- 
alad, curved dorsad at its apical third, flagellum as in preceding species. 
Genital styles as figured. 

One male (the type), Keung-Tin-Heung-, Lin-Hsien District, Kwang- 
tung, S. China, July 13-14, 1934. 

Cixius galeola differs from C. piloseUns Mats, in that the anterior trans- 
verse carinae of the vertex are not acutely angulate, and the clavus is devoid 
of fuscous markings: from C. nawae Mats., apart from differences in teg- 
minal marking, it is separated by the shape of the medio-ventral process 
of the pygofer, which in C. nawae is narrow and oblong: from C. nitohei 
Mats., to which it is perhaps nearest, it differs in the tegminal markings. 

Cixius scrupeus Fennah, new species. 
(Figure 2, A-E.) 

Male: length, 4.9 mm.; tegmen, 5.6 mm. 

Piceous; carinae and margins of head, pronotum and tegulae, femora at 
apex, yellow or testaceous; post-tarsi and longitudinal stripes on post-tibiae 
pale fuscous. Tegmina greyish-hyaline, a suffusion at base, one or two in 
middle of clavus and a spot near its apex brownish fuscous; stigma, except 
at base, and veins of membrane, fuseous-piceous, veins otherwise stramine- 
ous with granules darker. Wings greyish hyaline, veins yellowish, distally 

Anal segment elongate-ovate, distal margin shallowly convex, latero- 
apical angles not produced. Pygofer short dorsally and ventrally, lateral 
margins broadly produced, medioventral process broadly triangular. Genital 
styles narrow at base, expanded and curved dorsad distally. Aedeagus tubu- 
lar, slightly expanding distally, a long stout spine on left side one-third 
from apex, directed dorsad and curved cephalad at tip, a short spine near 
apex on right, slightly curved, directed laterad then obliquely dorso-ceph- 
alad; flagellum sinuate, distally with a stout spine directed cephalad, apical 
portion of flagellum in form of an even curved tube. 

One male (the type), Arisan to Hoshe, Tainan-Taichu District, Formosa 
July 14, 1948, Gressitt. 

In general coloration this species can only be compared with C. hakonensis 


IMats. It differs in that the frons and vertex are yellowish only at the mar- 
gins, not entirely, while the legs in C. hakonensis are yellow, not mostly 
pieeous as above. In C. hakonensis three piceous spots lie close to the 
costal margin; the medioventral process of the pygofer is narrow and peg- 
like, while the distal margin of the male anal segment is shallowly excavate. 

Cixius spp. 

One mutilated specimen, 2,300 m. Ali Shan (Arisan), Tainan District, 
Mar. 10, 1948, L. Gressitt. This species is near C. kuyanianus Mats. One 
female, with a relatively elongate and porrect ovipositor, taken by the above 
collectors in the same locality on August 20, 1947, is close to C. arisanus 

One mutilated specimen and one female, Yaoshan, Lin-hsien District, 
Kwangtung, S. China, May 6-10, 1934. In this pair the vertex is broader 
than long and almost as wide at apex as at base. 

Genus Oliarus Stal 
Stal 1862:306. Logotype, Oliarus ivalkeri Stal 1859b: 272 

Oliarus kurseongensis Dist. 

(Figure 4, G-H.) 

Distant, 1911, Ann. Mag. Nat. Hist. (8)8:737. 

One male, 1,000 m. Suisapa, Lichuan District, W. Hupeh, July 27, 1948, 

Oliarus nigronervatus Fennah, new species. 

(Figure 3, A-F.) 

Male: length, 5.5 mm.; tegmen, 8.0 mm. 

Vertex longer than broad (1.26:1), lateroapical aerolets separated by 
a rectangular median fossette, frons with disc markedly hollowed on each 
side of median carina, lateral margins obliquely subfoliate, median ocellus 
visible, though obsolescent. Rostrum slender, apical segment attaining post- 
coxae, slightly shorter than subapical. Post-tibiae feebly trispinose, apically 
with five short spines and one long spine, basal metatarsal segment with 7 
teeth, second metatarsal with six. Tegmina with Sc + R fork level with 
Cui fork. 

Castaneous-piceous; carinae of head and pronotum, subapical segment 
of rostrum, femora, and tibiae at apex testaceous; a short bar laterally on 
frons at apex, extending across genae to antennae, stramineous. Tegmina 
milky-hyaline, suffused fuscous near apex; veins and margins, except at 
stigma and apex of clavus, piceous. Wings milky-hyaline, suffused fuscous 
near distal margin, veins piceous, margins dark fuscous but pale at node. 

Anal segment elongate, asymmetrically ovate. Pygofer with laterodorsal 
angles asymmetrically produced, medioventral process basally parallel- 
sided, distally acute. Aedeagus narrowly ring-like at base, with three narrow 



[Proc. 4tii Ser. 

processes, one of them ai)proximately T-shaped, directed caudad as figured. 
Genital styles long and narrow, more or less straight, slightly swollen 
before apex. 

One male (the type), 1,000 m. Suisapa, Lichuan District, W. Hupeh, 
China, Aug. 19, 1948, Gressitt. This species is distinguished by the shape 
of the male genitalia. It differs from 0. ivaJkeri Stal in size and in the 
presence of a median fossette at the apex of the vertex; in the shape of the 
vertex from 0. caudatus (Wlk.), 0. hodgarti Dist., 0. siniJae Dist., 0. singu- 
laris Muir, 0. geniculatus Stal, 0. angusticeps Ilorv., 0. trifasciatus Mats. 
and ^latsumura's species 0. horishaniis, 0. iguchii, 0. qnadricinctus, 0. 
pachijceps, (). honinensis, 0. tappanus, 0. speciosus, 0. mori, 0. Jiopponis, 
0. artemisiae, and 0. kagoshimcnsis; in the shape of the male genitalia from 
all Oriental and Pacific species known to the writer or figured by Muir, and 
in coloration from 0. stigma Motsch., 0. tahrohanensis Mel., 0. fusconehido- 
sus Dist., 0. hinghami Dist., 0. indicus Dist., 0. greeni Dist., 0. annandalei 
Dist., 0. ijrolongulus Muir, 0. harwiaensis Mats., 0. hachijonis Mats., 0. 

Fig. 3. Oliarus nigronervatus, new species: A, vertex; B, aedeagus, right side; 
C, aedeagus, ventral view; D, medioventral process of pygofer, and genital styles, 
ventral view; E, male genitalia, left side; F, anal segment and pygofer, dorsal 
view. Oliarus cucullafun Noualh.: G, vertex; H, aedeagus, left side, small ventral 
spine viewed by transparency. Oliarus insetosus Jac. : I, vertex; J, aedeagus, dorsal 
view; K, anal segment of male; L, sketch of margin of pygofer, right side; M, sketch 
of margin of pygofer, left side; N, genital styles and medioventral process of 


ogasawareyisis Mats., and 0. apicalis Uhl. In the shape of the male genital 
styles it differs from 0. formosanus Mats., 0. velox Mats., and 0. oryzae 

Oliarus cucullatus Nonalh. 
(Figure 3, G, H.) 
Noualhier, 1896:255. 

Male: length, 4.0 mm.; tegmen, 4.8 mm. Female: length 4.5 mm.; teg- 
men, 5.5 mm. 

Vertex narrow, 1.6 times as long in middle line as broad across base, 
lateroapical areolets long, narrow, extending eaudad to one-fifth from base, 
lateral margins foliate, disc deeply hollowed out, frons and clypeus nar- 
rowly lozenge-shaped, disc of frons only shallowly hollowed out on each 
side of middle, median ocellus not visible, rostrum attaining post-trochan- 
ters, with apical segment as long as subapical. Post -tibiae laterally bispinose, 
apically six-spined. Basal metatarsal segment seven-toothed, second segment 
five-toothed. Tegmina with Sc + R fork level with Cui fork. 

Castaneous-fuscous; carinae broadly, tegulae, femora and tibiae at apex, 
post-tarsi, and hind margins of abdominal ventrites stramineous; two spots 
on side of head above eyes greyish-white. Tegmina of male translucent, sub- 
opaque, powdered sordid white, veins yellow except at apex, obscurely 
granulate, cross- veins and apical veins fuscous; tegmina of female either 
as in male or yellowish-hyaline at base of clavus, otherwise f uscous-castaneous, 
except for yellow proximal and ventral margins of stigma, veins concolorous 
or fuscous. Wings wholly greyish-white with brown veins (male) or distally 
infuscate with fuscous veins (female). 

Anal segment of male asymmetrical, right margin excavate. Pygofer with 
lateral margins asymmetrically produced into a Ijroadly rounded lobe on 
left side, and a turbinate subacute lol)e on right, medioventral process short, 
acute. Aedeagus tubular, a broad triangular lobe, terminating in a spine 
on right near apex directed ventrad, flcigellum terminating in a pair of 
short spines directed slightly to right, on left two spines. Genital styles in 
ventral view parallel-sided, at apex narrowly and acutely produced laterad, 
and feebly roundly produced mesad. 

Anal segment of female moderately short, lateral margins shallowly 
rounded, apical margin truncate. Posterior margin of seventh sternite very 
shallowly excavate in middle. Ovipositor with first valvulae much reduced, 
shortly triangular, third valvulae about as long as anal segment, elongate- 
triangular, flattened and blade-like. 

One male, Hangchow, China (May 22, 1923) E. C. Van Dyke, 1 female, 
Sui-Kwan San, Tin-tong, Loh-chan District, Kwangtung, S. China, Aug. 
1947; 13 males, 14 "pale" females and 15 "dark" females, 1,000 m. Suisapa, 
Lichuan District, W. Hupeh, China, July 19-25, 1948. One mutilated male. 


Pan-yii District, Ilonam Island, Canton, S. China, May 1-15, 1934, W. E. 
Hoffman, and a mutilated specimen from Ilonam Island, May 16, 1933, 
W. E. Hoffman, are assigned to this species. This species recalls 0. simlae 
Dist. but differs from the type in the shape of the vertex : it is highly prob- 
able that Mnemosyne ( ?) sinica Jac. is conspecific, but the evidence is too 
weak to justify the suppression of Jacobi's trivial name. 

Oliarus insetosus Jac. 

(Figure 3, I-N.) 
Jacobi, 1944:13. 

Male: length, 3.6 mm.; tegmen, 4.0 mm. Female: length, 3.2 mm.; teg- 
men, 4.3 mm. 

Vertex longer in middle line than broad (1.5: 1), lateroapical areolets 
extending backward to one-third from base, contiguous at apex. Frons 
with disc very shallowly tectiform, scarcely concave between carinae; 
median carina forked at base, rostrum with apical segment slightly shorter 
than subapical, attaining postcoxae. Tegmina with Sc + R fork much 
distad of Cui fork. Post-tibiae feebly trispinose laterally, with six spines 
at apex, basal metatarsal segment with 7 teeth at apex, second metatarsal 
segment with five teeth. 

Piceous, carinae and margins of head and pronotum, femora at apex, 
post-tibiae and all tarsi, margins of abdominal ventrites testaceous or ochra- 
ceous. Tegmina subhyaline, powdered greyish-white, stigma, distal trans- 
verse veins, and apical veinlets piceous, veins otherwise stramineous. Wings 
greyish white, distal margin and apical veins narrowly fuscous, veins other- 
wise pale yellowish-brown. 

Anal segment of male obcordate-ovate, apical margin deflexed and shal- 
lowly excavate. Pygofer with lateral margins slightly asymmetrical, that 
on left in profile with laterodorsal angle roundly produced, margin sub- 
rectangulate below this: margin of right side very slightly notched near 
lateroapical angle, apical margin oblique, medioventral process moderately 
long, narrow, acute, in side view shortly bladelike, striate. Aedeagus dorso- 
ventrally compressed, a short spine below, near middle directed to right, 
two longer curved spines on left near apex directed to left, at apex a 
slightly shorter curved spine directed to left, subparallel to preceding, a 
broad falcate lobe arising on left at apex, directed dorsad and cephalad, 
opposite this on right a membranous flagellum, slightly shorter. Genital 
styles as figured. 

Pregenital sternite of female with hind margin only very slightly pro- 
duced, and very shallowly excavate medially. 

One male, P'an Yu District, Honam Island, Canton, S. China, April 20, 
1934, W. E. Hoffman; one female, Un-long, New Territories, Hong-Kong, 
Sept. 19, 1940, J. L. Gressitt; four males and one female, 800-1,000 ft. trail 


between Lau-Tau-Di and Chang-Tau-Ching, Szechwan, China; one male, 
Sang-Hou-Ken, Hnpeh-Sze Border, China, July 19, 1948, Gressitt. This 
species differs from 0. walkeri Stal, which it closely resembles, in size, in 
the distinctly longer lateroapical facets of the vertex, and in the shape of 
the male genitalia. 

Oliarus petasatus Noualh. 

Noualhier, 1896:255. 

Length about 4.5 mm.; tegmen, 6.0 mm. 

Median ocellus present. Kostrum with apical segment sliglitly exceeding 
sub-apical, attaining post-coxae. Post-tibiae 3-5 spined (base and supernu- 
merary feeble), apically 6-toothed, basal metatarsal segment 7-toothed, 
second metatarsal 5-toothed. 

Venation as in 0. cucuUatus; Sc + R forked slightly distad of Cui, one 
apical Sc vein beyond stigma, 3 Es, 5 Ms. Castaneous-piceous; carinae and 
margins of head, and tegulae, brownish-yellow, tibiae and tarsi paler. Teg- 
mina hyaline, stigma yellowish-brown, veins stramineous; cross-veins and 
forks of clavus, Cui, and a minute spot in middle of vein M on corium 

One mutilated specimen, 1,900 m. Kunming, Yunnan-fu, Yunnan, S. W. 
China (July 4, 1940), Gressitt. 

This species superficially appears to be closely related to 0. cucuUutus, 
but it stands apart in the shape of the vertex and the presence of a func- 
tional median frontal ocellus. 

Genus Nesopompe Kirkaldy 
Kirkaldy, 1907:107. Oithotype, Oliarus felis Kirkaldy, 1906:399 

Nesopompe tsoui Muir. 

Figure 4, A-F.) 

Oliarus tsoui Muir, 1925:365. 

Two males, 1,000 m. Suisapa, Lichuan District, W. Ilupeh, China, June 
23, 1948, (Gressitt); 1 male, Mimasaka, Japan, July, 1912, (J. C. Thomp- 

Genus Betacixius Matsumura 

Matsumura, 1914:412. Orthotype, Betacixius ocellatus Mats. 1914:412 


(1) (2) Tegmina with a large, ocellate black spot apically (3) 

(2) (1) Tegmina without such a spot distally (5) 

(3) (4) An oblique brown band extending from clavus part way across middle 

of corium B. tonkinensis Mats. 

(4) (3) Corium of tegmina without such a band B. ocellatus Mats. 

(5) (6) Tegmina with a tapering oblique dark band extending from stigma along 

nodal line of cross veins to Cu (7) 



[Pkoc. 4tii Ser. 

(6) (5) Tegmina without such a band (15) 

(7) (8) Tegmina with apical cells of M and Cu strongly infuscate 

B. trmisversus Jac. 

(8) (7) Tegmina with apical cells not infuscate (9) 

(9) (10) Tegmina with apical margin black or obviously very dark — (11) 

(10) (9) Tegmina with apical margin fuscous or not especially darkened (13)- 

(11) (12) Frons with a pallid spot at middle of lateral margins, clypeus dark, meso- 

notum testaceous B. kumejimae Mats. 

(12) (11) Frons without such spots; mesonotum, except scutellum, castaneous- 

piceous B. euterpe, new species 

(13) (14) Tegmina with an oblique dark band extending from clavus into middle 

of corium, little distad of level of union of claval veins. ...B. imllidior Jac. 

(14) (13) Tegmina with a spot near sutural margin of clavus near union of claval 

veins, no oblique dark band at this level extending into corium 

B. obliquus Mats. 

(15) (16) Tegmina greyish hyaline, only infuscate at stigma 

male B. nigromarginalis, new species 

(16) (15) Tegmina with a dark spot or line in clavus (17) 

(17) (18) Tegmina with apical margin black or very dark (19) 

(18) (17) Tegmina with apical margin not especially dark (25) 

(19) (20) A V-shaped dark mark in stigma, an oblique band from middle of clavus 

extending into corium B. nigromarginalis, new species 

(20) (19) No such oblique fascia from clavus into disc of corium (21) 

Fig. 4. Islesoijonipe tsoui Muir: A, vertex; B, fossette of vertex; C, anal seg- 
ment, hand margin of pygofer, and genital style, side view; D, anal segment of male, 
dorsal view; E, aedeagus; F, medioventral process of pygofer. Oliariis kurseon- 
genesis Dist. : G, aedeagus, left side; H, aedeagus, right side. 


(21) (22) Costa black, a curved black line in stigma, a spot in clavus 

- B. rinkihoni Mats. 

(22) (21) Costa not especially dark, posterior margin of clavus dark (23) 

(23) (24) Clypeus distally black, pronotum laterally pallid, tegulae sordid yellow, 

their posterior edge dark B. clypealis Mats. 

(24) (23) Clypeus distally brown, pronotum laterally brown, pallid only at margin, 

tegulae light brown B. hrunneus Mats. 

(25) (26) Stigma and first apical cell of tegmen piceous B. rohustus Jac. 

(26) (25) Stigma dark, a dark suffusion over all apical cells and across base of 

tegmina B. nelicles, new species 

Betacixius nigromarginalis Feniiah, new species. 

Male: length, 3.2 mm.; tegmen, 4.2 mm. Female: length, 3.4 mm.; teg- 
men, 5.0 mm. 

Reddish-brown; frons lateroapieally, lateral margins of pronotnm (occa- 
sionally whole pronotum) and trochanters, stramineous-yellow; pronotal 
disc, mesonotum and abdominal tergites castaneous to piceous. Tegmina 
of female hyaline with greyish bloom, stigma with narrowly V-shaped mark 
fuliginous, Sc in stigma and vein round apical margin black; a narrow 
transverse fascia across clavus at apical third, extending into corium to 
cell M, a tinge at base of clavus reddish-brown; basal margin of clavus 
yellowish. Tegmina of male usually dark only at stigma, otherwise hyaline 
with stramineous veins. Wings hyaline with stramineous veins. 

Anal segment of male moderately long, telson slightly basad of middle, 
apical margin transverse-concave, apical angles acuminately produced. 
Pygofer with lateral margins strongly convex, medioventral process sub- 
triangular, rounded at apex. Genital styles S-shaped, apposed ventral mar- 
gins enclosing a broadly ovate space, distal portion of each style falcate. 
Aedeagus with a circularly curved spine on left near apex and a short ledge 
in a similar position on right; flagellum lying above left margin, sides par- 
allel for most of length, distally a short curved spine directed cephalad, and 
below this a subquadrate plate with a stout spine directed ventrad. 

Described from 8 males (one the tjq^te) and 20 females, Suisapa, Li- 
chuan District, W. Hupeh, China, on ridge about 1,200 to 1,500 m. (Gres- 
sitt, July 19-25, 1948). This species is distinguished by the shape of the 
male genitalia and by coloration. 

Betacixius nelides nelides Fennah, new species and new subspecies. 

Male: length, 3.5 mm.; tegmen, 4.2 mm. Female: length, 3.5 mm.; teg- 
men, 4.8 mm. 

Fuscous; carinae and median area of frontal disc reddish-brown, hind 
margin of ])ronotum, hind tibiae, and tarsi stramineous; lateral fields of 
pronotum, fore and middle legs, and post-femora yellow with pale fuscous 
suffusion; clypeus, mesonotum, pro- and mesocoxae and mesopleura, and 


abdomen castaneous-piceous. Tefjmina hyaline with greyish bloom, a faint 
suffusion at base, stigma, a small spot ovei-lying union of elaval veins, and 
sometimes a faint suffusion over membrane, fuscous, veins testaceous. Wings 
hyaline, with greyish bloom, veins fuscous. 

Anal segment moderately short, asynunotrical, subovate, acutely rounded 
at apex, right margin produced and lateroapically deflexed in profile, anal 
style situated at middle; ])ygofer with lateral margins symmetrically con- 
vex, medioventral jn-ocess lu'oader across base than long, distally rounded. 
Genital styles narrow and cui'\"ed basally, apically subreniform in outline, 
axis of apical area at right angle to basal stalk. Aedeagus with a large 
curved spine ventrally near apex, arising on left, curved below aedeagus 
and directed to right, a stout spine near apex, closely adpressed to left side 
and directed cephalad, flagellum with a long curved spine at base directed 
cephalad but decurved at abruptly-tapering apex, and a more slender 
curved spine directed cephalad. 

Three nuiles (one the type) and 4 females, Tung-lu, Che-Kiang Prov- 
ince, China, (Mrs. D. E. Wright, May 22, 1926). Two females from Mo- 
kansan in the same province (Mrs. D. E. Wright, Aug. 24, 1927) are doubt- 
fully placed here. 

This species is distinguished by the combination of characters given in 
the synopsis and by the form of the male genitalia. 

Betacixius nelides atrior Fennah, new subspecies. 

Male: length, 4.2 mm.; tegmen, 5.0 mm. Female: length, 3.7 mm.; teg- 
men, 5.0 mm. 

Fuscous-piceous ; basal half of frons reddish brown; rostrum, except at 
apex, pro- and mesochanters, post-tibiae, and tarsi stramineous. 

One male and 1 female, Hangchow, China, (E. C. Van Dyke, May 19, 
1923). This geographical subspecies is distinguished by its darker col- 

Betacixius euterpe Fennah, new species. 

Male: length, 3.6 mm.; tegmen, 4.8 mm. 

Testaceous; pronotal disc pale yellow; mesonotum, except scutellum and 
middle portion of abdominal tergites, castaneous-piceous. Tegmina hyaline 
with greyish bloom, a faint suffusion at base, a fascia from sutural margin 
at middle of clavus across Sc + R, another from stigma along nodal line 
to Cu, a suffusion just distad of apex of clavus, and apical margin, fuscous- 

Anal segment of male bilaterally symmetrical, apical margin transverse, 
lateroapical angles acuminate. Pygofer with lateral margins convex, medio- 
ventral process triangular, acute at apex. Genital styles S-shaped, width 
of distal portion of stalk equal to that of vertical apical portion. Aedeagus 
with a stout spine arising laterally on left near apex, directed cephalad, 


slightly deeurved at tip, a curved spine directed to right lying below fiagel- 
lum in its basal half, a more slender curved spine, directed ventrally, 
arising on right side of flagelluni near apex. 

One male (the type), Hau-leng, Tin-tong", Loh-chan District, Kwang- 
tung- Province, S. China, (Aug. 1, 1947). 

This species is distinguished by the combination of characters given in 
the synopsis and by the form of the male genitalia. 

A female from Yaoshan, Liu-hsien District, Kwangtung (May 10, 1934), 
(length 3.1 mm.; tegmen 4.3 mm.), provisionally placed here, but probably 
representing another species, differs from the male of B. euterpe in having 
the lower part of the genae and lateroapical areas of the frons pale yellow, 
apical part of clypeus, procoxae, and mesopleura and basal part of meso- 
coxae castaneous-piceous, and legs fuscous; tegmina with a spot in clavus 
slightly distad of union of veins, a fascia from stigma along nodal line to 
Cu, all apical cells in M and Cu fuscous, margin between stigma and suf- 
fused apical area, testaceous-brown. 

Genus Macrocixius Matsumura 
Matsumura, 1914:393. Ortliotype, Macrocixius giganteus Mats. 

Macrocixius giganteus Mats. 
Matsumura, 1914:394. 

One female, 1,000 m., Musha (AVuse) to Bandai, Taichung District, For- 
mosa, Aug. 24, 1947, Gressitt. 

Genus Borysthenes Stal 
Stal, 1866:165. Logotype, Cixim finitus Stal, 1866:392. 

Borysthenes maculatus (Mats.). 
Barma maculata Matsumura 1914:430. 

One male, Chizuka, Okinawa, July-Sept., 1945, Bohart and Harnage. 
Borysthenes acuminatus Fennah, new species. 
(Figure 5, A-E.) 

Male: length, 4.1 mm.; tegmen, 6.0 mm. 

Testaceous; middle portion of frons and clypeus except at apex, all coxae 
and hind femora and tibiae faintly suffused fuscous; mesonotum and post- 
tarsi castaneous; clypeus at apex, apical segment of rostrum, fore and mid- 
dle legs distad of trochanters and abdominal ventrites dark fuscous to fus- 

Tegmina subtranslucent, ivory white on most of corium, greyish white 
in costal cell and membrane, three broad fasciae, one at base, another areu- 
ately across middle, and the third over most of membrane as figured, choco- 
late brown. Wings pallid with cross veins and a broad submarginal fascia, 
sepia brown. 



[Pkoc. 4tii Ser. 

Anal segment of male moderately short and broad, lateroapical angles 
not symmetrical. Pygofer with lateral margins slightly asymmetrical, medio- 
ventral process distally scmicircularly rounded. Genital styles with inner 
and outer margins (in posterior view) subparallel, apical margin straight 
and strongly oblique, occasionally slightly reflected. 

Described from 4 males (one the type), 1,000 m. Lichuan District, W. 
Hupeh, China, July 23-25, 1948, Gressitt. 

This species is distinguished by tegminal pattern and by the shape of 
the male genitalia. 

Borysthenes deflexus Fennah, new species. 
(Figure 5, I, J.) 

Male: length, 4.1 mm.; tegmen, 6.0 mm. Female: length, 5.0 mm.; teg- 
men, 7.0 mm. 

Ochraceous-testaceous; mesonotal disc and abdomen slightly infuscate. 
Tegmina translucent, powdered greyish, an arcuate irregular band from 
middle of clavus to basal third of costal cell, a more deeply convex band 

Fig. 5. Borysthenes acuminatus. new species: A, anal segment of male, right 
side; B, pygofer, right side (basal margin uppermost) ; C, aedeagus, left side; 
D, genital style, left side; E, ditto, ventral view. Borysthenes emarginatus, new 
species: F, tegmen; G, anal segment, aedeagus, and left genital style; H, pygofer, 
left side. Borysthenes deflexus, new species: I, anal segment, hind margin of 
pygofer, aedeagus, and right genital style, side view ; J, distal outer margin of 
genital style, posterior view. 


from middle of costal cell to distad of apex of clavus, an interrupted band 
from node to sutural angle, an ovate spot in apical cells near apical angle, 
fuscous. Wings translucent with a suffusion on costa near base and two 
transverse bands distally. 

Anal segment of male symmetrical, acuminate at apex, in profile deflexed 
through 90° distad of anal foramen. Pygofer with lateral margins subree- 
tangulately convex, medioventral process elongate-triangular. Aedeagus 
tubular, a short straight spine on right near apex directed ventro-cephalad, on 
left side near apex a short spine curved ventrad and a long porrect spine 
directed ventro-cephalad, and decurved at tip, a long slender spine arising 
dorsally at apex directed cephalad, flagollum membranous, apex subspinose 
and curved dorsad. 

Genital styles in profile curved dorsad distally, upper and lower margins 
subparallel, a rounded ledge on outer face one-fifth from apex, apex acutely 
rounded, upper distal margin truncate. 

One male (the type), Tso-kok-wan, Lungtau Shan, altitude 250-350 m.. 
Kwang'tung', June 5, 1947, C4ressitt; 1 female, Keung Tin Heung, Lin- 
hsien, Kwangtung, S. China, July 16-17, 1934. 

This species is distinguished by the tegminal pattern and by the shape of 
the male genitalia. 

Borysthenes emarginatus Fennah, new species. 

(Figure 5, F-H.) 

Male: length, 4.1 mm.; tegmen, 5.6 mm. 

Testaceous; middle portion of frons, clypeus, all coxae, postfemora, and 
tibiae very slightly darker; pronotum around disc, and mesonotum, infus- 
cate, rostrum at tip piceous. 

Tegmina subtranslucent, sordid pallid yellow, a > -shaped cloud near 
base reddish-brown; a broad fascia from costal cell near apex to sutural 
margin distad of apex of clavus, a broader transverse band across middle 
of membrane, apical cells of R and Mi fuscous, veins concolorous. Wings 
translucent powdered sordid white, veins fuscous. 

Anal segment of male in profile slightly deflexed distad of middle, apical 
margin deeply excavate to level of anal foramen. Pygofer with lateral mar- 
gins subangulately convex. Aedeagus tubular, a long slightly curved spine 
on right at apex, directed cephalad, flagellum twisted through 360°, mi- 
nutely denticulate on margin distally. Genital styles in profile strongly 
angulately bent just distad of middle, an eminence distally on inner margin. 

One male, White Cloud Mountain, P'an Yu District, Canton, S. China, 
May 5, 1934. This species is distinguished by coloration and by the shape 
of the male genitalia. 



[Proc. 4th Ser. 


(1) (2) Post-tibial spur awl-shaped, circular in cross section; mesonotum with 
five carinae Ugyops Guer. 

Spur not as above; mesonotum tricarinate (3) 

Spur thick, flattened or concave on inner face, margin witliout teeth.... (5) 

Spur thin, usually deeply concave on inner face, margin with or without 
teeth (15) 

Lateral carinae of frons and vertex only moderately developed (7) 

Lateral carinae of frons and vertex deeply foliate Purohita Dist. 

Vertex subtriangular with sides slightly convex, sometimes elongate 
Tropidocephala St§,l 

Vertex quadrate (9) 

First segment of antennae not more than half as long as second (11) 

First segment of antennae at least two-thirds length of second (13) 

Mesonotum shorter than head (in dorsal view) and pronotum together 

Eurysa Fieb. 

Mesonotum longer than vertex and pronotum together. ...Pundaluoya Kirk. 

Frons at least twice as long as broad, clypeus medially carinate 

Arcofacies Muir 

Frons not nearly twice as long as broad, clypeus devoid of median carina 

A7'cofaciella. new genus 

Basal segment of antennae subtriangular or sagittate, widening distad 

- (17) 

Antennae with basal segment not as above, cylindrical or slightly com- 
pressed (21) 

Antennae with basal segment sagittate; clypeus in profile feubrectangulate 

at middle; median carina of frons forked at extreme base 

Belocera Muir 

Antennae with basal segment triangular but not sagittate; clypeus in 

profile not angulate at middle (19) 

Frons with paired submedian carinae Pseudaraeovius Kirk. 

Median carina of frons forked near level of lower margin of eyes 

Perkinsiella Kirk. 

Basal segment of post-tarsus with one or more spines on side 

Nilaparvata Dist. 

Basal segment of post-tarsus devoid of spines on side (23) 

Basal segment of antennae short, as long as broad or a little longer... (25) 

Basal segment of antennae not very short, longer than broad (35) 

Vertex distinctly longer than broad (27) 

Vertex at most only slightly longer than broad, often equal to width or 

even shorter (31) 

(27) (28) Oblique carinae of vertex meeting at apex, or only slightly before it.... (29) 


(28) (27) Oblique carinae of vertex meeting much before apex and continued distad 

as a single median carina Saccharosydne Kirk. 

(29) (30) Second antenna! segment not more than twice length of basal segment, 

median fossette on vertex subtriangulately elongate Sardia Mel. 

(30) (29) Second antennal segment three times length of basal segment, medio- 

apical fossette about as long as broad, polygonal-rounded 

Stenocranus Fieb. 

(31) (32) Median carina of frons forked at about one-third from base, or distad 

DicranoU'opis Fieb. 

(32) (31) Median carina of frons simple or forked only at extreme base (33) 

(33) (34) Vertex at base more than twice as broad as an eye in the same line 

Eoeurysa Muir 

(34) (35) Vertex not twice as broad as an eye Delphacodes Fieb. 

(35) (36) Clypeus fully three-quarters as long as frons; frons three times as long 

as broad Sogata Dist. 

(36) (35) Clypeus scarcely two-thirds as long as frons; frons two and a half times as 

long as broad (37) 

(37) (38) Median carina of frons forked at extreme base of frons; basal segment 

of antennae more than twice as long as broad at apex 

- Ukanodes, new genus 

(38) (37) Median carina of frons forked near level of middle of eyes; basal segment 

of antennae relatively shorter 

Chloriona Fieb. subg. Sogatella, new subgenus 

Subfamily ASIRACINAE Fieber 

Genus Ugyops Guerin-Meneville 

Guerin-Meneville, 1834:477. 

Haplotype, U(jyo2)s percheronii Guerin-Meneville, 1834:477. 

Ugyops vittatus ( Mats. ) . 

(Figure 6, A, C, F, G, J.) 

Bidis vittata Matsumura, 1906:31, pi. 1, fig. 5. 

Frons longer than broad (3:1) ; basal segment of antennae slightly less 
than two-thirds of length of second; genae not inflated below level of an- 
tennae. Aedeagns, viewed from left, with basal third of flagellum sinuate, 
the upper and lower margins subparallel. 

One male, Chizuka, (Jkinawa, July-Sept., 1945, Bohart and Harnage. 
This species differs from U. kinhergi Stal from Ponape in the relatively 
shorter frons and smaller size. 

Ugyops zoe l^Vnnah, new species. 
(Figure 6, B, D, E, H, I.) 

Male: length, 6.3 mm.; tegmen, 8.5 mm. 

Frons longer than broad (3.3: 1), submedian carinae uniting one-third 
from apex, genae distinctly inflated below level of antennae; basal segment 



of antennae two-thirds leng-th of apical segment; vertex with posterior 
margin almost level witli middle of eyes. Pronotal disc relatively broad, 
its lateral carinae distinctly convex. 

Testaceous; common carinal eminence at base of frons and apex of ver- 
tex, intercarinal areas of anterior half of vertex, a weak intercarinal stripe 
on each side of frons at base, a small suffusion on sides of head above eye, 
carinae of vertex finely, median carina of pronotum, posterior margin and 
a faint cloud behind eyes, and mesonotal carinae, castaneous; second seg- 
ment of antennae, protibiae, and tarsi fuscous. Tegmina sordid hyaline, 
lightly waxed grey, veins castaneous interrupted testaceous, apical portion 
of cells at distal margin infuscate; a broad infuscate Y-shaped band extend- 
ing from apical margin between R and M2 to cell M3, where one limb 
passes to the margin at Cuib and the other to the nodal line between Cu 
and M. 

Anal segment of male moderately short, bilaterally symmetrical. Pygofer 
with lateral margins convex caudad, rather angulately excavate below. 
Aedeagus, viewed from left, with basal third of flagellum markedly dilated, 
with ventral margin more convex than dorsal. 

Holotype male, Tai-pin-t'suen, Lam-ka-heung, Lai-mo-ling-, Kiung Shan 
District, Hainan Island, July 20-21, 1935; one male, Tsai-Chau (Tinhosa) 
Island, June 2, 1932, Hoffman; and one mutilated specimen, Tung-Chung, 
Lan-Tau Island (near Hong-Kong), Aug. 16-19, 1934. 

JJgyops zoe broadly resembles U. vittatus (Mats.). The genae are inflated 
below the eyes (not in U. vittatus), the lateral carinae of the pronotal disc 

Fig. 6. Ugyops vittatus Mats.: A, head and thorax, dorsal view; C, head in 
profile; F, frons and clypeus; G, pygofer and right genital style, side view; J, 
aedeagus, left side. Ugyops zoc, new species: B, head and thorax; D, head in profile; 
E, frons and clypeus; H, pygofer and right genital style; I, aedeagus, left side. 


are convex (concave in U. vittntus), and the tegmina are longer and dif- 
ferently marked. This species has a ninch shorter f rons than TJ. kinhergi Stal, 
from which, as from U. vittatus, it differs in the shape of the male genitalia 
and in size. It differs from U. pictifrons Stal, TJ. impictus Stal, and TJ. per- 
cheronii Guer. in coloration and marking (the genitalia of the last three 
were not available for comparison ) . 

Subfamily DELPHACINAE Jensen-Haarup 

Tribe Tropidocephalini Muir 

Genus Arcofacies Muir 

Muir, 1915:319. Orthotype, Arcofacies fullawayi Muir 

Arcofacies fullawayi Muir. 
Muir, 1915:320. 

One female, 300 m. Pe-poi, N. of Chung-King, Sze-chuan, W. China, July 
27, 1940, J. L. Gressitt; two females, Hong-Kong, China, Oct., 1895, Koebele. 

Genus Eoeurysa Muir 
Muir, 1913:249. Orthotype, Eoeurysa flavocapitata Muir 

Eoeurysa flavocapitata Muir. 

Muir, 1913:249. 

Four males and two females on sugar cane, Pan-yu District, Honam 
Island, Canton, S. China, May 23, July 26-31, 1935; Jan. 10, 1937, W. E. 

Genus Tropidocephala Stal 
Stal, 1853:266. Haplotype, Troijidocephala flaviceijs Stal, 1855:93 

Tropidocephala brunnipennis Sign. 
Signoret, 1860:185. 

One male, Sam-ah-Kong, Yai-hsien District, Hainan Island, S. China, 
Feb. 1, 1935; one female, Honam Island, Canton, China, April 7, 1933; one 
female, Loh Fau Shan, Poh Lo District, Kwangtung, S. China, April 
6-8, 1934. 

Tropidocephala f estiva (Distant). 
Smara f estiva Distant, 1906:478. 

One female, Cheung-nga-San, Tin-tong, Loh-Chang District, Kwangtung, 
Aug. 16, 1947, Tsang, one female. Yen-ping, Nan-ping District, Fukien, 
June and July, 1933; one female, Yaoshan, Lin-hsien District, Kwangtung, 
April 27-28, 1934; one female, Hau-leng, Tin-tong, Loh-chang District, 
Aug. 1, 1947. 


Tropidocephala speciosa Bierm. 
Orchesma speciosa Bierman, 1908:29. 

One female, Yaoshan, Lin Hsien District, Kwangtung, S. China, April 
24-26, 1934. 

Tropidocephala breviceps Mats. 

Matsumura, 1907:58. 

One female, Hoi-IIow, Kiimg-Shan District, Hainan, S. China, 1932, 
W. E. Hoffman. 

Tropidocephala signata Dist. 

Distant, 1912:192. 

One female, Riviere de Hue, Anam, March 16, 1927, Mrs. D. E. Wright. 

Genus Pundaluoya Kirkaldy 
Kirkaldy, 1902:52. Orthotype, Delphax ertiesti Kirby, 1891:140 

Pundaluoya sp. 

One female, testaceous, with yellowish-hyaline tegmina, slightly infumed 
on membrane, 1,000 m. Suisapa, Lichuan District, AV. Hupeh, China, July 
24, 1948, Gressitt. 

Arcofaciella Fennah, new genus 

Head with eyes as wide as pronotum. 

Vertex more than three times as broad as long in middle, anterior and 
posterior margins transverse, lateral margins slightly convex, strongly 
converging anteriorly, median carina simple, straight, frons inclined an- 
teriorly in profile, slightly longer than broad, broadest at level of lower 
margin of eyes, lateral margins convex, median carina simple, shortly 
forked at base, area within fork apparently depressed; clypeus in profile 
more or less at right angle to base of frons, lateral carinae straight, median 
carina obsolete. Rostrum with subapical segment exceeding apical, apex 
reaching mesotrochanters. Antennae short, stout, second segment markedly 
longer than first, but both together not exceeding length of eye. Pronotum 
more than twice as long as vertex, anteriorly shallowly convex, posteriorly 
shallowly excavate, median carina distinct, lateral carinae developed only 
in anterior portion in line with lateral margins of vertex; mesonotum 
strongly convex, almost gibbous, tricarinate, mesoscutellum horizontal. Legs 
relatively short and stout, profemora not longer than procoxae, post-tibiae 
with a small spine laterally at base, another about a third from apex, five 
teeth at apex, spur short, convex, devoid of teeth except for a minute tooth 
at apex, basal metatarsal segment with about 8 small even teeth. Tegmina 
long, corium enfolding abdomen at region of node, costal margin sinuate, 
concave distad of node, apical margin shallowly undulate, forks of Sc-R and 
Cui distad of union of claval veins, eight cells at apex, excluding stigmal cell. 



Arcofaciella verrucosa Fennah, new species. 

(Figure 7, A-F.) 

IIoLOTYPE, Female : length, 2.5 mm. ; tegmen, 4.3 mm. 

Greenish-stramineous; two spots on vertex and impression in fork of 
median carina of frons orange. Tegmina translueent-ochraceous, a small 
round callus near Sc-R fork and another near Cui fork piceous, a faint 
transverse stripe across middle of clavus, and apical veins at margin, brown. 
Wings yellowish hyaline, veins coneolorous. 

Third abdominal segment of male produced laterally in a narrowly 
conical process which is bent caudad. Anal segment very short, ring-like. 
Pygofer moderately long, longest at middle, laterodorsal angles not pro- 
duced, posterior lateral margins broadly sinuate, ventral margin convex 
and oblique in profile, medioventral process absent; diaphragm sclerotised 
only laterally, devoid of armature in middle and not developed as a sclero- 
tised bridge above base of genital styles. Aedeagus comprising a laterally- 
compressed plate, narrowing distad and strongly defiexed, recurved caudad 
at blunt apex; on right side of this a long slender spinose process, evenly 
curved ventrad and mesad distally, crossing the former process at apex. 
Genital styles rather long, vertical, very slightly curved, of subequal 
width from base to near apex, then abruptly bent caudomesad and slightly 
narrowed; apical margin shallowly concave, distal angles prominent, the 
lower especially so. 

Holotype female, Hong Kong, China, Koebele, October, 1895, deposited 
in the California Academy of Sciences. Allotype male, and paratype fe- 
male. Hong Kong, China, Koebele, deposited in the U. S. National Museum. 
This genus recalls Arcofacics but differs in the shape of the frons and in 
the relative size of the antennae, in the gibbous mesonotum, in the rela- 

Fig. 7. Arcofaciella vei-riicosa, new genus and species: A, frons and clypeus; 
B, head and thorax, side view of dorsal half; C, vertex and pronotum; D, striated 
lobe anterodorsolaterally on third (apparently first) abdominal segment; E, teg- 
men; F, post-tibial spur. 


tively lartier ])ronotiiiii, in the shai)e of the tegmina and in rostral propor- 
tions, and in the absence of a median carina on the clypeus. 

The holotype and allotype of Arcofacies peyiangensis Miiir are congeneric 
with A. verrucosa and this species must be termed ArcofacieUa penangensis 
(Muir) comb. nov. It differs from A. verrucosa in its much larger size and 
in the shape of the frons, which is 1.6 times as long as broad, as contrasted 
with 1.4; in the less acute apex of the tegmina, and in the following details 
of the male genitalia : Anal segment with posterior lip deep, with latero- 
apical angles acute, as contrasted with lip shallow and angles obtuse and 
less produced; aedeagus with a deep transverse lobe dorsally at base (the 
corresponding lobe in A. verrucosa is much smaller) ; genital styles in pro- 
file strongly constricted in distal third before incurved apex (whereas in 
A. verrucosa from the same viewpoint they appear parallel-sided). The 
two species differ also in color, A. penangensis being much darker. 

Tribe Delphacini Lambertie 

Genus Sardia Melichar 

Melichar, 1903:96. Haplotype, Sardia rostrata Mel. 

Sardia rostrata Melichar. 
Melichar, 1903:96. 

One female, Vinglon District, Cochin-China, French Indo-China, Aug. 
6-10, 1934; one female, Chizuka, Okinawa, July-Sept., 1945, G. E. Bohart 
and C. L. Harnage. 

Genus Nilaparvata Distant 

Distant, 1906:473. Orthotype Delphax Jugens Stal {=^Nilaparimta greeni Dist.) 

Nilaparvata lugens (Stal), 

Delphax lugens Stal, 1854:246. 

Two males, Nam-ting-tseun (10 m. N.E. of Sam-ah-Kong), Yai-hsien 
District, Hainan Island, S. China, Feb. 10-11, 1935. A female from Mai- 
chan (1 hour by bus more or less west of Ch'ui-man), Suwen District, July 
26, 1932, W. E. Hoffman, is referred to this species; one macropterous 
male, 1,000 m., Suisapa, Lichuan District, W. Hupeh, China, Aug. 20, 1948, 
Gressitt; eight macropterous males and eighteen macropterous females, 
Mokansan, Che-Kiang Province, Sept. 6, 1927, Mrs. D. E. Wright, are 
placed here; the genitalia also agree with figures of N. oryzae Mats. Two 
females, Riviere de Quangtri, Anam, April 30, 1927, Mrs. D. E. Wright, 
have the median carina of the frons interrupted in the middle of the disc 
by a transverse sulcus, and may possibly belong to another species. 


Nilaparvata muiri China. 

(Figure 8, S.) 

Nilaparvata (?) muiri China, 1925:480. 

One male, Chu-Chon Fn, Che-Kiang, China, Sept. 6, 1926, Mrs. D. E. 
Wright. A female, taken at 1,000 m., Suisapa, Lichnan District, W. Hupeh, 
China, Ang. 20, 1948, by Gressitt is doubtfully assigned to this species. It 
differs from N. muiri China in tlie number of lateral spines on the basal 
segment of the post-tarsus (3) and in the venation of Cu in the tegmen, 
which is normal. The mesonotum just outside the base of each lateral 
carina is tumid. 

Genus Phyllodinus Van Duzee 
Van Duzee, 1897:240. Haplotype, Eurysa nervata Van Duzee, 1894:191 

Phyllodinus luzonensis Muir. 

(Figure 8, N, 0.) 
Muir, 1916:383. 

One male and two females, all brachypterous, Riviere de Hue, Anam, 
Mrs. D. E. Wright, Mar. 16, 1927. 

Phyllodinus macaoensis Muir. 

(Figure 8, H, I.) 
Muir, 1913:246. 

One macropterous male and three macropterous females. Riviere de 
Quangtri, Anam (April 30, May 5, 1927) : one brachypterous female, 
Riviere de Hue, Anam (Mar. 16, 1927), Mrs. D. E. Wright. This series 
agrees in all details with Muir's description of P. macaoensis : the genitalia 
differ from those of P. nigromaciilosus Muir in the more tumid lateroipical 
areas of the anal segment, and in the shorter and stouter and markedly 
S-shaped genital styles, and in the coloration of the distal portion of the 
tegmina. The venation of the macropterous tegmina agrees with that de- 
scribed by Muir, though in one specimen the number of branches of Cu is 
two, not three. The position of the forking of the veins on the corium is 
variable, and anastomosis in various degrees may occur between Sc and R. 

Genus Dicranotropis Fieber 

Fieber, 1866:530. Logotype, Delphax hamata Eoheman, 1847:45 

Dicranotropis huensis. 

(Figure 8, E-G.) 

Male : length, 2.8 mm. Female : length, 2.8 mm. Bi-achypterous form : 
Stramineous; disc of frons and intercarinal areas of pronotal disc pale 
brown; genae below eyes, clypeus, lateral fields of mesonotum, pro- and 
mesocoxae except at base, a spot on metapleura, abdomen, except laterally, 
on eighth ventrite, distal margin of pygofer and tenth segment castaneous- 



[Proo. 4th Ser. 

pieeous. Tegmina dark eastaneous-ti-aiisliicent, a spot at middle of claval 
margin, another at apex of clavus, and a narrow area overlying cross-veins 
between node and M hyaline or pallid. 

Anal segment large, broad, ventral margin transverse, lateral angles 
produced ventrad in a spine. Pygofer with hind lateral margin in profile 
sinuate, more or less vertical, medioventrally a small tongue-like process 
from posterior margin on each side of middle line. Genital styles strongly 
S-shaped, twisted, broadest a third from apex. Aedeagus strongly laterally 
compressed, distal portion reflected anteriorly. Diaphragm devoid of orna- 
mentation, dorsal margin shallow! y concave. 

One male and two females. Riviere de Hue, Anam, Mar. 16, 1927, Mrs. 
D. E. Wright. This species is distinguished by the shape of the genitalia. 

Fig. 8. Unkanodes sapporoiia Mats.: A, male genitalia; P, anal segment of male 
in profile; Q, aedeagus; R, genital style, lateral view. Delphacodes inachtis, new 
species; B, male genitalia; C, aedeagus, lateral view. Delphacodes shirozui Ish.: 
D, male genitalia. Dicranotropis huensis, new species: E, male genitalia; F, apical 
portion of aedeagus; G, apical portion of genital style, lateral view. Phyllodhius 
macaotnsis Muir: H, male genitalia, posterolateral view; I, apical portion of geni- 
tral style. Chloriona (s.) sirokata M. & I.: J, male genitalia; K, anal segment of 
male, posterior view; L, M, aedeagus, left and right sides. Phyllodinus luzonensis 
Muir: N, O, male genitalia , posterior and posterolateral views. Unkanodes sap- 
porona (Mats.): P, anal segment of male in profile; Q, aedeagus, left side; R, left 
genital style. Nilaparvata muiri China: S, male genitalia. 


Geims Chloriona Fieber 
Fieber, 1866:519. Haplotype, Delphax iinicolor H.-S., 1835:66 

The writer recognizes that a wholly satisfactory subdivision of the con- 
cept Chloriona Muir, which embraces such different species as Delphax 
unicolor H.-S., Delphax vitticollis Stal {= Chloriona turneri Muir), and 
Lihurnia slossoni Ball is likely to prove very difficult to achieve: to judge 
by material so far examined, it seems likely that one or two segregates can 
be made, but that these may show the same degrees of variation as found 
in TJgyops. In view of the frequency with which Delphax furcifera Ilorv. 
appears in literature it seems desirable to anticipate adequate revisionary 
treatment by segregating this species from the narrow concept of Fieber 
based on Delphax unicolor. It is likely to prove, on critical study, that 
the gap separating the two concepts is wide, and of generic significance: 
indeed, if it were merely a case of comparing the respective type species, 
this could be shown forthwith. As, however, it is not at present possible 
to bring forward a considered statement of the position of all the species 
involved, the writer here proposes to go no farther than to establish a new 
subgenus. For the convenience of students this subgenus is compared with 
certain existing genera some of which are not represented in the Chinese 

Sogatella Fennah, new subgenus 

Head little narrower than pronotum. Vertex slightly longer than broad, 
its width at base subequal to width of eye in same line, and exceeding two- 
thirds of its length, apical margin transverse interrupted by projecting 
submedian carinae of frons; carinae of vertex and frons slender and dis- 
tinct. Frons longer than broad with median carina forked approximately 
at level of middle of eyes, lateral margins straight, subparallel. Antennae 
cylindrical, moderately short, basal segment distinctly longer than broad, sec- 
ond segment longer than first. Rostrum not attaining post-trochanters. Length 
of pronotum and mesonotum combined scarcely as long as maximum width of 
latter. Pronotum tricarinate, lateral discal carinae almost straight, strongly 
diverging basad, not reaching hind margin; not parallel with mesonotal 
carinae. Mesonotum tricarinate, longer than vertex and pronotum together. 
Legs terete, not at all compressed, rather slender, post-tibial calcar with 
about twenty small teeth, basal segment of post-tarsus devoid of spines 
along side. 

Sogatella differs from tiie typical subgenus in the relatively narrower 
vertex, which in Chloriona unicolor H.-S. considerably exceeds the width 
of an eye, in the parallel lateral margins of the frons, in the proportions of 
the frons, in the slightly shorter combined length of the pronotum and meso- 
notum, and in the fewer teeth on the post-tibial spur, of which there are thirty 


in the typical subtienotyi')e. Leptodelphax Haiipt has a relatively longer com- 
bined pronotum and mesonotum, while the medial carina of the frons is 
broadened basally, not forked. CaUigypona has a relatively longer rostrum. 
KeHsia lias a relatively shorter first antennal segment and slightly curved 
lateral frontal margins, while Prokelisia differs entirely in the shape of the 
head and pronotum. 

Type of subgenus, DeJphax furcifera Horv. 

Chloriona (Sogatella) furcifera (Horvath). 
Delphax furcifera Horvath, 1899:372. 

Five males, nine females, and one nuitilated specimen, 1,000 m. Suisapa, 
Lichuan District, W. Hupeh, China (Aug. 19-24, 1948; 1 male, Riviere de 
Hue, Anam, Mar. 16, 1927, Mrs. D. E. Wright; 1 male, Tunglu, Sept. 8, 
1926, 7 females Mokansan, Che Kiang Province, China, Sept. 6-8, 1927, 
Mrs. D. E. Wright. 

Chloriona (Sog-atella) sirokata (M. & I.) 

(Figure 8, J-M.) 

Sogata sirokata Matsumura & Ishihara, 1945:64. 

Male: length, 2.3 mm.; tegmen, 2.8 mm. 

Fuscous; carinae of frons and clypeus, antennae, sides of clypeus, meso- 
scutellum, legs, and abdomen laterally ochraceous; posterior and ventral 
margin of pronotum ivory white. Tegmina hyaline, faintly infumed, veins 

Median carina of frons forked at extreme base on dorsal surface of head. 
Post-tibial spur with twenty minute teeth. 

Anal segment of male with a pair of diverging spines arising at middle 
of distal margin. Pygofer dorsolaterally much longer than ventrally, pro- 
duced at dorsolateral angles and incurved, the distal part expanded and 
truncate parallel with truncate edge of opposite member, medioventral 
process absent, but a pair of bluntly tooth-like eminences on margin laterad 
of base of genital styles. Diaphragm thickened and umbonate medially, 
somewhat roughened. Genital styles moderately broad and flattened, shal- 
lowly curved, outer margin sinuate, inner margin concave, outer distal 
angle broadly and roundly lobate, inner distal angle acuminate. Aedeagus 
tubular, unornamented, of equal width throughout, and with orifice apical, 

Two males. Riviere de Hue, Anam, Mar. 16, 1927 (Mrs. D. E. Wright). 

Genus Delphacodes Fieber 
Fieber, 1866:524. Logotype Delphax vrulsanti Fieber, 1866:526 

Delphacodes terryi Muir. 

Muir, 1917:334. 

Two macropterous males, Honam Island, Canton, China, May 4, 1932; 
White Cloud Mountain, Canton, China, Mar. 19, 1933. 


Delphacodes shirozui Ishihara. 
(Figure 8, D.) 
Ishihara, 1949:53. 

Median carina of frons forked at level of incipient curve into vertex. 
Post-tibial spur with sixteen teeth. 

Anal segment small a pair of long slender spines arising on each side of 
middle line on hind margin. Pygofer dorsally deeply emarginate, dorso- 
lateral angles produced caudad, distally incurved, medioventral process 
absent, diaphragm armed medially with a bispinose sclerite. Genital styles 
sinuate on outer nuirgin, concave on inner, apical angles bluntly pointed, 
distal margin shallowly concave. Aedeagus basally compressed, distally 

Two males, Lau-Chi, Che-Kiang, China, June 20, 1926, Mrs. D. E. Wright. 

Delphacodes inachus Fennah, new species. 

(Figure 8, B, C.) 

Male: length, 1.8 mm.; tegmen (brachypterous), 1.1 mm. 

Vertex with median carina forked at base, on horizontal dorsal area. 
Antennae attaining level of f rontoclypeal suture. Post-tibial spur with about 
eighteen teeth, basitarsus wdth an oblique row of five teeth and two separate 
teeth at proximal end of oblique row. Rostrum slightly surpassing meso- 

Fuscous-piceous; a few spots on apex of frons and anterior half of genae 
fuscous; carinae of vertex, frons, and clypeus, lateral fields of pronotum, 
and a broad line overlying median carina of pronotum and mesonotum 
pallid to white; disc of vertex, rntennae, posterior area of genae and sides 
of clyjieus, legs, except post-tibiae at base, abdomen at sides, and a ring 
around anal emargination of pygofer, testaceous. 

Anal segment of male small, deeply sunk in emargination, wnth a pair 
of slender spines arising near middle line, directed ventrad. Pygofer nar- 
rowly but deeply emarginate above, laterodorsal angles broadly produced 
and inflected mesad, bluntly rounded, so that the cavity of the pygofer in 
posterior view is heart-shaped, a slight lenticular swelling in middle of 
lateral margin; medioventral process absent; diaphragm with armature 
carried near ventral margin of foramen, in form of a short lobe projecting 
caudad. Styles long, vertical, slightly swollen at base, slightly curved, pro- 
duced in a spine at apex. Aedeagus V-shaped, much laterally compressed, 
widened near middle, acuminate distally. 

One brachypterous male, Suisapa, Lichuan District, W. Hupeh, China, 
July 23, 1948, Gressitt. This species is distinguished by the shape of the 


Unkanodes Fennah, new genus 

Rather slender. Head little narrower than pronotuni. Vertex longer 
than broad, its width at base not exceeding width of an eye, shallowly 
rounded at apical margin; carinae of vertex and frons distinct. Frons longer 
than broad, with median carina forked only at extreme base. Antennae 
cylindrical, basal segment two and a half times as long as broad, at least 
half as long as second. Length of pronotum and mesonotum combined 
equal to maximum width of latter. Pronotum tricarinate, lateral discal 
carinae almost straight; very weakly curved laterad, not reaching hind 
margin and not in line with mesonotal carinae. Mesonotum longer than head 
and pronotum together, tricarinate. Legs terete, not at all compressed, post- 
tibial calcar with about twenty-two teeth, Imsal segment of post-tarsus devoid 
of spines. 

Type species, Vnkana sappovoyia Mats. 

Unkanodes sapporona (Mats.). 

(Figure 8, P-R.) 

Unkana sapporona Matsumura, 1935:74. 

One male, Mokansan, Che-Kiang Province, China, probably collected 
by Mrs. D. E. Wright. The genitalia are figured. 

This species, which has recently been transferred to DeJphacodes by 
Ishihara (using Delphacodes striateUa Fall, as the basis of reference for 
generic characters), is separated from DeJphacodes by the characters given 
in the key above and from Sogata (interpreted strictly from the holotype of 
S. dohertyi Dist.) in the shorter vertex and frons and relatively very much 
shorter clypeus. 



(Adapted from Muir) 

(1) (2) Claval veins uniting near apex of clavus; first claval vein strongly gran- 

ulate; second not or weakly so, subparallel to commissural margin.. (3) 

(2) (1) Claval veins uniting near middle of clavus; first claval vein not granulate, 

second strongly so, curved Anigrus Stal 

(3) (4) Clypeus devoid of lateral carinae Nisia Mel. 

(4) (3) Clypeus laterally carinate Eponisia Mats. 

Genus Nisia Melichar 
Melichar, 1903:53. Haplotype, Meenoplus atrovenosus Leth. 

Nisia atrovenosa (Leth.). 

(Figure 9, A-C.) 

Meenoplus atrovenosus Lethierry 1888:466. 

One female, Cheung-Mu-Tsang, 50 km. northwest of Chungking, China, 



on citrus, Gressitt, July 8, 1948; 1 mutilated specimen, Riviere de Quangtri, 
Anam, Mrs. D. E. Wright, April 30, 1927; 1 male, 1 female. Riviere de Hue, 
Anam, Mrs. D. E. Wright, March 16, 1927; 7 females and 7 mutilated speci- 
mens, Tunglu, Che-Kiang Province, Mrs. D. E. Wright, Sept. 8, 1926; one 
female, same locality, Mrs. D. E. Wright, Sept. 10, 1926; 1 mutilated speci- 
men, Mokansan, Che-Kiang Province, Mrs. D. E. Wright, Sept. 10, 1927. 

Nisia suisapana Fennah, new species. 
(Figure 9, D-F.) 

Female: length, 2.2 mm.; tegmen, 3.0 mm. 

Tegmina 2.1 times as long as broad, broadest at level of stigma, anterior 
margin convex, not indented at node, anterior branch of M simple at apex; 
post-tibiae 8-spined at apex, basal metatarsal segment 7-spined, second 
metatarsal segment with 5 spines. 

Stramineous, probably powdered white in life, abdomen pale fuscous, 
eyes and spines on legs black. Tegmina sordid white marked with pale 
fuscous as figured. Wings white, veins pallid. 

Third valvulae of ovipositor in profile with dorsoapical lobe well devel- 
oped, about as broad as long. 

One female, 1,000 m. Suisapa, Lichuan District, W. Hupeh, China, 
Gressitt, Aug. 20, 1948. This species differs from all others in the tegminal 
markings: it is superficially nearest to N. alhovenosa Dist. 

Genus Eponisia Matsumura 
Matsumura, 1914:285. Orthotype, Eponisia guttida Mats. 

Eponisia guttula Mats. 

Matsumura, 1914:286. 

Post-tibiae 8-spined at apex, basal metatarsal segment 7-spined, second 
metatarsal segment 6-spined. 

One female, Mokansan, Che-Kiang Province, Mrs. D. E. Wright, Sept. 6, 

Fig. 9. Nisia atrovenosa Leth.: A, anal segment and pygofer, left side; B, ditto, 
posterior view; C, aedeagus, left side. Nisia suisapana. new species: D, anal seg- 
ment and external genitalia of female, right side; E, tegmen; F, anterior portion of 


1927, is considered to be a geograpliieal representative of this species. The 
infuscate markings on the tegmina are very pale. 

Genus Anigrus Stjil 
Stal, 1866:172. Logotype, Anigrus sordidus Stal, 1866:173 

Anigrus nigricans (Mats.). 
Paranisia nigricans Matsumura 1914:285. 

Post-tibiae 8-spined, basal metatarsal segment 6-spined, second meta- 
tarsal 5-spined. Tegmina with anterior margin yellowish-brown, apical and 
commissural margins fuscous. 

One female, Mizuho, Formosa, Gressitt, April 22, 1932. 


Genus Kinnara Distant 

Kinnara, Distant, 1906:39. Orthotype, Pleroma ceylonica Melichar, 1903:42 

Pleroma Melichar, 1903:41 

Kinnara fumata Mel, 

Melichar, 1903:42. 

One male, 1,000 m., Suisapa, Liehuan District, W. Hupeh, July 25, 1948. 

Family DERBIDAE Spinola 

(1) (2) Tegmina with clavus closed, second cubital vein reaching hind margin 

directly, frons narrow, not strongly laterally compressed (3) 

(2) (1) Tegmina with clavus open; second cubital vein curving into subapical 

transverse line of cross veins: frons strongly laterally compressed.... (5) 

(3) (4) Pronotum with a distinct median disc bounded laterally by carinae; frons 

little longer than broad, margins convex, antennae and apical repcment 
of rostrum long Vinata Dist. 

(4) (3) Pronotum without a distinct median disc, frons much longer than broad, 

margins concave, antennae and apical segment of rostrum short 

Vekunta Dist. 

(5) (6) Wings not more than half as long as tegmina, usually narrow or reduced 

with stridulary organ on reduced anal lobe (7) 

(6) (5) Wings more than half as long as tegmina, ampliate (15) 

(7) (8) Tegmina with all median sectors single, usually 6; antennae usually 

shorter than frons (9) 

(8) (7) Tegmina with five median sectors, one furcate (11) 

(9) (10) Basal median cell narrow, wings about half as long as tegmina, apex 

rounded; antennae much shorter than frons Proutista Kirk. 


( 10 ) ( 9 ) Basal median cell wide and short, not more than 3.5 times as long as broad, 

wings much less than half as long as tegmina, acute at apex 

Diostrombus Uhl. 

(11) (12) Second or third median sector forked, base of clypeus in profile straight 

Pamendanga Dist. 

(12) (11) First median sector with two to four branches (13) 

(13) (14) Posterior margin of tegmina undulate Losbafiosia Muir 

(14) (13) Posterior margin of tegmina not undulate.. Zoraida Kirkaldy 

(15) (16) Tegmina less than 2.5 times as long as broad, clavus closed or nearly so 

Rhotana Wlk. 

(16) (15) Tegmina more than 2.5 times as long as broad, clavus open (17) 

(17) (18) Media arising from radius distad of Sc + R + M fork; lateral carinae of 

vertex and frons very large Megatropis Muir 

(18) (17) Media arising from R or basad of Sc + R fork (19) 

(19) (20) Sc + R fork at or basad of middle of tegmen, subcostal cell elongate in 

profile, vertex and frons meeting in a distinct angle. ...Kamendaka Dist. 

(20) (19) Sc + R fork distad of middle, subcostal cell short, in profile head broadly 

ovate, no evident point of union of vertex and frons Vivaha Dist. 

Genus Losbafiosia Muir 
Muir, 1917:85. Haplotype, Losbafiosia bakeri Muir 

Losbafiosia bakeri Muir. 
Muir, 1917:86. 

One female, Lung-Tau Shan, N. Kwangtung, China, Gressitt, June 11, 

Genus Diostrombus Uhler 
Uhler, 1896:283. Haplotype, Diostrombus politns Uhl. 

Diostrombus politus Uhl. 

Uhler, 1896:284. 

Post-tibiae 4-spined at apex, basal metatarsal segment 7-spined, second 
metatarsal 9-spined. 

Twenty-five males and 23 females, Cheung-Mu-Tsang, 50 km. northwest 
of Chungking, China, July 8, 1948, Grressitt. It is evident from the collection 
data that adults were "swarming" at the time of collection. 

Genus Zoraida Kirkaldy 
Kirkaldy, 1900:242. Orthotype, Derbe sinuosa Boheman, 1838:225 

Zoraida kirkaldyi Muir. 
(Figure 11, A-E.) 
Muir, 1918:205. 

Lateral submargins of frons each longitudinally shallowly sulcate so that 


frons between eyes is more tlian two-thirds width of an eye in frontal view. 
Post-tibiae with 1 spine laterally, 5 apically, basal metatarsal segment 6- 
spined at apex, second metatarsal 6-spined. 

Two males and 1 mutilated specimen, 1,000 m., Suisapa, Lichuan Dis- 
trict, AV. Ilupeh, China, August 20, 1948, Gressitt. 

Genus Pamendanga Distant 

Distant, 1906b: 298. Orthotype, Pamcnilanga ruUlinea Dist, 1906:299 

Pamendang"a sauterii Muir. 

Paraproutista sauterii Muir, 1915:131. 

Aedeagus broad, shallowly scoop-like; ventrally on right a moderately 
short spine curved laterad and caudad, a large thin elongate triangular 
lobe, acuminate at tip, directed caudad at apex of aedeagus, a spine dorso- 
laterally on left a little before apex, directed anteriorly, ventrally, in 
posterior view, a pair of short triangular eminences slightly to left of 
middle line. 

Post-tibiae unarmed at sides, with 5 spines at apex, basal metatarsal seg- 
ment 4-spined, second metatarsal segment 3-spined. 

One male (mutilated) and 2 females (one mutilated), 1,000 m., Suisapa, 
Lichuan District, W. Ilupeh, China; male, July 23; females, July 23, 25, 
1948, Gressitt. 

Genus Rhotana Walker 
Walker, 1857:160. Haplotype, Rhotana latipennis Walker, 1857:160 

Rhotana maculata Mats. 

(Figure 10, D.) 
Matsumura, 1914:295. 

Post-tibiae 4-spined at apex, basal metatarsal segment 5-spined, second 
metatarsal 4-spined. 

One male, Mokansan, Che-Kiang Province, Mrs. D. E. Wright, Sept. 16, 
1927. This identification requires confirmation. The specific characters of 
the tegminal venation are figured. 

Rhotana satsumana Mats. 

(Figure 10, A, B, B.) 
Matsumura, 1914:294. 

Two mutilated specimens, Mokansan, Che-Kiang Province, Mrs. D. E. 
Wright, Sept. 6, 10, 1927. 

In the material before the writer the lateral carinae of the frons are 



strongly divergent and actually separated from the level of the lower mar- 
gin of the eye. 

Rhotana satsumana contracta Fennah, new subspecies. 

(Figure 10, C.) 

Female: length, 3.3 mm.; tegmen, 6.0 mm. 

Lateral margins of frons contiguous to well below level of eyes, not 
strongly divergent until below level of antennae. Tegmina with basal vena- 
tion of M and Cu as figured. Posterior margin of pregenital sternite tri- 
angularly iDroduced medially, subrectangulate at apex. 

Stramineous, powdered pallid. Tegmina translucent, powdered white, 
basal third except in middle of intervenal areas, an arcuate fascia from 
middle of costal margin to apex of clavus subparallel to apical margin of 
tegmen, a narrow band on each side of, though separated from, apical line 
of transverse veins passing posteriorly into a large suffusion, pale fuscous; 
veins yellow, red where transversing infuscate or pigmented areas. A yellow 
suffusion bordering subapical line of transverse veins in anterior four cells. 
Wings infumed pale fuscous, white along apical margin, a short arcuate 
dark spot interrupted l)y union of M-Cu cross vein and Cuia. 

Described from one female, 1,000 m., Suisapa, Lichuan District, W. 
Hupeh, China, July 24, 1948, Gressitt. This subspecies is distinguished by 
the sliape of the lateral carinae of the frons and by the tegminal venation. 
A single female from Hainan Island (en route Cheung-kon-ts'uen to Tai 
pin-ts'uen, Kiung-Shan District, July 19, 1935) is doubtfully ascribed to 
this species; its points of difference (in tegminal venation and the shape of 
the black spot on the wings) are figured to facilitate recognition. 

Fig. 10. Rhotana satsumana Mats.: A, tegminal venation at base of M (Hainan 
Island specimen) ; B, spot on wing (Hainan Island specimen) ; E, tegminal venation 
in M (Mokansan specimen) ; C, contracta, new subspecies, tegminal venation in M. 
Rhotana maculata Mats.: D, tegminal venation at base of M. 



[Proc. 4tii Seb. 

Genus Vivaha Distant 
Distant, 1906b: 307. Orthotype, Vivaha facialis Dist. 

Vivaha facialis Dist. 

(Figure 13, A-C.) 
Distant, 1906b: 308. 

Post-tibiae with 6 spines at apex, basal metatarsal segment 4-spined, sec- 
ond metatarsal segment bispinose. 

One male and 1 female, 1,000 m., Suisapa, Lichuan District, W. Hupeh, 
China, Gressitt, Aug. 19, 1948. 

Genus Megatropis Muir 

Muir, 1913:57. Orthotype, Megatropis coccineolinea Muir, 1913:57 

Megatropis formosana (Mats.). 

Mesotiocerus formosanus Matsumura, 1914:301. 

Post-tibiae 6-spined at apex, basal metatarsal segment 4-spined, second 
metatarsal 2-spined. 

One mutilated male and 7 females, 1,000 m., Suisapa, Lichuan District, 
W. Hupeh, China, July 23, 24; Aug. 23, 1948, Gressitt. The series agrees 
so closely with Matsumura 's description and figures that no subspecific 
differentiation is apparent. 

Genus Kamendaka Distant 
Distant, 1906b: 310. Orthotype, Kamendaka spectra Dist. 

Kamendaka spectra Dist. 

(Figure 11, F, G.) 
Distant, 1906:311. 

Post-tibiae laterally unarmed, apically with 5 spines; basal metatarsus 
6-spined, second metatarsus 5-spined. 

One male taken on ridge, 1,200-1,500 m., Suisapa, Lichuan District, W. 
Ilupeh, China, July 25, 1948, Gressitt. 

Fig. 11. Zoraida kirkladyl Muir: A, anal segment of male, side view; B, medio- 
ventral process of pygofer; C, genital style; D, aedeagus, left side; E, apex of 
aedeagus, right side. Kamendaka spectra Dist.: F, apical portion of aedeagus, left 
side; G, anal segment, pygofer, and left genital style. 



Kamendaka (Eosaccharissa) nigromaculata Dist. 

Chaprina nigromaculata Distant, 1911:645. 

Post-tibiae laterally unarmed, apically with 11 spines, basal metatarsal 
segment 9-spined, second metatarsal segment 8-spined. 

One male and two females, Riviere de Hue, Anam, Mar. 16, 1937, Mrs. 
D. E. Wright. 




















(11) (12 

(12) (11 

(13) (14 

(14) (13 


(16) (15 

(17) (18 

(18) (17 
(19) (20 

(20) (19 
(21) (22 

Genus Vekunta Distant 
Distant, 1906:8. Orthotype, Vekunta tenella Melichar, 1903:41 

Tegmina unicolorous, subopaque, pallid yellowish or creamy white.... (3) 
Tegmina infuscate or with dark suffusion, or with fuscous markings.... (7) 

Wings white, or subhyaline powdered white (5) 

Wings infuscate with darker veins. Philippine Islands 

V. palawanensis Muir 

A fuscous-piceous spot on propleura; wings white with white veins. 

Formosa? V. albipennis Mats. 

No dark spot on propleura; wings hyaline-white with veins distally grey- 
ish brown. China V. nivea, new species 

Tegmina subhyaline, sordid white, veins pallid, most cells suffusedly bor- 
dered with pale sepia-brown. Formosa V. lyricen, new species 

Tegmina not as above (9) 

Tegmina pale, conspicuously bordered fuscous or fuscous piceous along 
costal margin and sutural margin, or at least along sutural margin of 

clavus, often with apical margin also dark in part (11) 

Tegmina without distinct marginal infuscation as above, usually with a 

general rather dark ground color, occasionally of light hue (17) 

Tegmina with a piceous mark over subcostal cell and apex of costal cell. 
Formosa V. nigrolineata Muir 

Tegmina not marked as above (13) 

Tegmina with a narrow fuscous band from radial cross vein to apex, apical 

margin fuscous (15) 

Tegmina without such band in R (43) 

Tegmina with costa dark fuscous. Java Y. hyalina Muir 

Tegmina with corium and veins anterior to R milky-white 

F. nigrinervis Schmidt 

Tegmina subhyaline, tinged yellow, with yellowish veins, stigma hyaline, 

a small dark spot near apex of costa, apical veins infumed at tip, apical 

margin infuscate in female. Formosa V. maculata Mats. 

Tegmina more or less generally suffused fuscous, or very dark (19) 

Tegmina piceous, costa yellowish, a small yellow spot at stigma. Assam. 

V. flavipes Muir 

Tegmina not piceous, and not so marked (21) 

Tegmina with costal cell, at least anteriorly, pallid for most of its length; 

remainder of corium infuscate (23) 



[Pkoc. 4th See. 

(22) (21 
(23) (24 
(24) (23 
(25) (26 

(26) (25 

(27) (28 
(28) (27 

(29) (30 

(30) (29 

(31) (32 
(32) (31 

(33) (34 

(34) (33 

(35) (36 
(36) (35 
(37) (38 

(38) (37 
(39) (40 

(40) (39 

(41) (42 

(42) (41 

(43) (44 

(44) (43 

Costal cell wholly infuscate, or infuscate to near apex (27) 

A black spot on propleura laterally. Ceylon V. jninctula Mel. 

No black spot on propleura (25) 

General body color yellow ; tegmina brownish yellow, disc of vertex not 

hollowed or markedly depressed. Ceylon V. tenella Mel. 

General body color castaneous; tegmina dark brown with a pallid oblong 

fleck at node; disc of vertex rectangulately hollowed out. Formosa 

V. stigmata Mats. 

Tegmina with a dark spot ad joining a pallid spot at costal margin (29) 

Tegmina unicolorous, or if with a pallid spot, with no dark spot adjacent 

to it (31) 

A hyaline spot on costal margin at node, a piceous spot just distad of it, 

and an oblique pallid stripe adjoining. Sumatra Y. nitida Bierm. 

A yellowish patch at end of costal cell, reaching from costa to media, with 

a dark spot in middle of it V. badia Muir 

Tegmina unicolorous, translucent brown (33) 

Tegmina paler at, or near, stigma, or if not then with veins very dark, 

but paler distally (35) 

Mesopleura fuscous; transverse carinae of vertex testaceous; legs yellow 

with tibiae apically and tarsi medially and apically black. Formosa 

V. botelensis Mats. 

Pleura with a round black spot; carinae of vertex fuscous, legs yellowish. 

Formosa V. makii Muir 

Mesonotum light brown on disc, darker laterally........ (37) 

Mesonotum fuscous-piceous or black :.:.'.... (41) 

Vertex 1.5 times as long as broad; tegminal veins sordid yellow. Formosa 

V. shirakii Mats. 

Vertex as long as broad, or if not, then tegminal veins dark fuscous.... (39) 
Tegminal veins sordid yellow; anal segment of male terminating in a 

small point. Japan and China V. malloti Mats. 

Tegminal veins dark fuscous; anal segment of male slightly emarginate 

at apex. Formosa V. umbripennis Muir 

Tegmina with a small pale translucent spot at apex of costal cell; veins 

unicolorous throughout. Java V. pseudobadia Muir 

Tegmina fuscous; veins very dark, paler toward apex. Formosa 

Y. atripennis Mats. 

Clypeus with a black spot in middle; a black spot on propleura; margins 

of tegulae dark. Philippine Islands Y. lineata Mel. 

Clypeus without a black spot in middle; an oval black spot on both pro- 
pleura and metapleura. Formosa Y. kotosJwnis Mats. 

Vekunta nivea Fennah, new species. 

(Figure 12, A-E.) 

Male: length, 3.5 mm.; tegmen, 4.9 mm. Feaiale: length, 3.0 mm.; teg- 
men, 4.5 mm. 

Vertex across base 1.33 times length in middle line. Post-tibiae unarmed 



laterally, 7-spined at apex, basal metatarsal segment 6-spined, second meta- 
tarsal segment 6-spined. 

Stramineous; mesonotum and pregenital sternite tinged yellowish brown, 
a fuscous-pieeous spot on mesothoracic pleurites, a paler fuscous spot later- 
ally on metathoracic pleurites, apical segment of rostrum pale, its apical 
disc black. Tegmina hyaline, entire membrane distad of stigma faintly suf- 
fused with yellow, veins pallid, faintly yellow in membrane, anterior part 
of apical margin faintly tinged orange. Wings hyaline, veins greyish-brown 
distally. Insect in life powdered wdiite. 

Pregenital sternite of female shallow, transversely sulcate, distally pro- 
duced and directed upward in a broad subtriangular process of the shape 

Anal segment of male long, narrow, broadly longitudinally sulcate, 
apically deflexed and recurved below to point cephalad. Aedeagus dorsally 
with a short medial spine at base directed upward and to left, a long medial 
spine at apex directed cephalad with a minute tooth on left at its base; on 
right side a long sinuate spine directed caudad, at apex of flagellum a 
short tooth-like plate directed upward and cephalad. Genital styles long, 
curved dorsad and mesad distally, of subequal width throughout, a pyram- 
idal eminence applied to inner face at middle, this eminence terminating 
dorsally in a single blunt tooth, ventral angles setose-toothed. 

Described from 1 male and 4 females, Mokansan, Che-Kiang Province, 
China, Mrs. D. E. Wright, Aug. 29, Sept. 16, Sept. 22, 1927. 

This species is distinguished by the combined characters of a pitchy 
mesothoracic spot, and generally jiallid color. 

Fig. 12. Tekiinta nivea. new species: A, vertex; B, pregenital sternite of female; 
C, apex of anal segment of male, posterior view; E, ditto, side view; D, aedeagus, 
side view. Vekunta lyricen, new species: F, vertex; G, tegmen; H, pregenital 
sternite of female. 


Vekunta lyricen Feiiiiah, new species. 
(Figure 12, F-H.) 

Female: length, 3.5 mm.; tegmen, 5.1 mm. 

Reddish-brown; lateral margins of frons, apex of rostrum, a band adjoin- 
ing median carina of mesonotum on each side, abdomen and genitalia, fus- 
cous; pronotal disc, a broadening band from eye to and across tegiila, a 
longitudinal submedian band on mesonotum, pallid yellow, mesoseutellum 
salmon-pink, tibiae and tarsi testaceous-fuscous. 

Tegmina greyish-hyaline, veins pallid, most cells incompletely but clearly 
bordered sepia. AVings rather sordid white, veins concolorous basally, pale 
sepia distally. 

Pre genital sternite produced in a narrowly triangular process, almost 
semicircularly rounded at apex, directed caudo-dorsad. 

One female, 1,000 m., Musha (Wuse) to Bandai, Taichung" District, 
Formosa, Gressitt, August 24, 1947. This species does not possess any 
obvious spot on the mesothoracic pleurites, and the tegminal markings do 
not agree with those of any species so far described in the genus. 

Genus Vinata Distant 
Distant, 1906a: 8. Orthotype, Erana operosa Walker, 1857:151 

Vinata sp. nr. nigricornis Stal. 

One female, Lu-ling-paai, Yaoshan, Lin-Hsien District, Kwangtung, 
Oct. 1-2, 1924. 

Post-tibiae 7-spined at apex, basal metatarsal segment 6-spined, second 
metatarsal 6-spined. 

Family ACHILIDAE Stal 


(1) (2) Pronotal disc elongate, three-quarters length of mesonotum, no median 

carina on mesonotum ; hind wing markedly notched at Cu2 

Rhotala Walk. 

Pronotal disc relatively shorter, mesonotum medially carinate, apical 

margin of hind wing entire (3) 

Vertex less than two-thirds as wide as pronotum; tegmina with numerous 

apical longitudinal veins Fat^eyitilla Mete. 

Vertex at least two-thirds as wide as pronotum; tegmina with about seven 

apical veins at margin distad of Sc (5) 

Width of vertex measured at base of middle line at least twice length 

along middle; M^.o forking at apical transverse line 

Plectoderoides Mats. 

Width of vertex not twice length along middle (7) 

Vertex devoid of a carina across apex, or with median carina prominent 

and apical transverse carina obsolete (9) 















(8) (7) Vertex with one or more distinct carinae at apex (11) 

(9) (10) Vertex not distinctly produced before eyes, about as wide at apex as at 

base, lateral discal carinae of pronotum concave, curved laterad, not 
reaching hind margin Tangina Mel. 

(10) (9) Vertex not as above, lateral discal carinae of pronotum straight, reaching 

basal margin Akotropis Mats. 

(11) (12) Vertex with a single distinct carina across apex (13) 

(12) (11) Vertex with two transverse carinae between frons and vertex, confluent 

in middle but separating sublaterally to enclose a more or less distinct 
triangular facet (17) 

(13) (14) Vertex 3 times as wide across base as long in middle line.. Zathauma Fenn. 

(14) (13) Vertex relatively narrower (15) 

(15) (16) Vertex produced before eyes for one- to two-thirds length of eye, in profile 

meeting frons acutely, lateral carinae of frons not as eminent as median 
carina, almost meeting acutely at base Betatropis Mats. 

(16) (15) Vertex produced before eyes for scarcely half their length, in profile 

meeting frons subrectangulately, lateral carinae of frons more promi- 
nent than median carina Caristianus Dist. 

(17) (18) Vertex medially carinate throughout, disc little depressed, if at all; teg- 

mina with Sc and R together with six or seven veinlets at margin 
near stigma; no transverse callus on mesonotal disc Usana Dist. 

(18) (17) Vertex medially carinate only in basal two-thirds, or less, lateral margins 

strongly raised; mesonotum with a transverse callus on anterior third 
of disc, tegminal venation not as above Magadha Dist. 

Genus Faventilla Metcalf 
Metcalf, 1948:60. Orthotype, Cixius pustulatus Walker, 1857:146 

Faventilla spp. 

One mutilated specimen, 250-300 m., Tao-kok-wan, Lung tau Shan, 
Kwangtung Province, S. China, Sept. 6, 1947, Gressitt. Dr. China, who 
kindly compared this and the following with Walker's types in the British 
Museum, points out that this "comes closest to F. pustulata Walker but 

B m, _^^>^>n]*^''J' c 

Fig. 13. Vivaha facialis Dist.: A, tegmen; B, head in profile; C, antenna. 


differs in the venation (Sc + R stalk being mneh shorter), in the absence 
of the three black spots on the tegmen and of the large black spot on the 
side of the propleuron." 

One female, 2,800 ft.. Big Pool, Loh Fan Shan, Kwangtung, Oct. 13, 1935, 
E. R. Tinkham, "resembles F. guttifer Walker in size and color, particu- 
larly the pale greenish-yellow color. It differs, however, in the much shorter 
anteriorly-rounded vertex. In F. guttifer Walker the vertex is nearly as 
long as wide at base with the anterior margin angulate where the mid- 
facial carina meets it." 

Genus Usana Distant 
Distant, 1906:293. Orthotype, Usana lineolalis Dist. 

Usana lineolalis Dist. 

Distant, 1906:294. 

Male: length, 3.0 mm.; tegmen, 4.5 mm. Female: length, 3.5 mm.; teg- 
men, 5.0 mm. 

Stramineous to testaceous, a linear mark on each side of middle line of 
vertex, a round spot on lateral lobes of pronotum, a spot on tegulae and on 
mesopleura, a faint spot on mesonotal disc near base of lateral carinae fus- 
cous. Tegmina yellowish-translucent, powdered grey, stigmal cells and all 
apical cells at margin fuscous, veins concolorous, stigmal veins and apical 
veins near margin ivory-white. Wings slightly infuscate, veins fuscous. 

Post-tibiae 7-spined at apex, basitarsus 7-spined distally, second post- 
tarsal segment 6-spined. 

Anal segment short, broad, lateral margins convex, apical margin exca- 
vate, lateroapical angles produced and deflexed. Pygofer with a short 
finger-like process laterally on hind margin near anal segment. Genital 
styles broad bearing at middle of dorsal margin a trispinose lobe. Aedeagus 
subtubular, flattened on dorsal surface, keel-like below, a spine arising at 
apex in middle line directed ventrocephalad below aedeagus; laterally at 
apex a flange-like lobe projecting laterad; on right a pair of vertical lobes, 
one overlapping the other, both concave on anterior margin. 

Twelve males, 13 females and 3 mutilated specimens, Mokansan, Che 
Kiang Province, China, Aug. 24-Sept. 19, 1927, Mrs. D. E. Wright. 

One female, Tai-ka, Tin-tong, Loh-chang District, Kwangtung, S. China, 
Aug. 18, 1947. The series differs from the type only in tlie less definite mot- 
tling of the corium along the veins. 

Genus Magadha Distant 
Distant, 1906:290. Orthotype, Cixius flavisigna Walker, 1851:348 

Magadha metasequoiae Fennah, new species. 
Female: length, 4.8 mm.; tegmen, 6.8 mm. 
Tegmina with costal vein close to margin throughout. Fuscous; vertex, 


except for a spot in each half of disc, about seven spots laterally on frons 
and a diffuse transverse bar across middle, clypeus broadly at base, narrowly 
at apex, disc and carinae of pronotum, dorsal half of tegulae and lateral angles 
of mesonotum, femora at base and apex, tibiae at base, middle and apex, 
and posterolateral margins of abdominal ventrites, ivory -yellow to stramine- 
ous; a sprinkling of small round spots on disc of frons testaceous. Tegmina 
ivory-hyaline, about eight marginal spots in costal cell, two of which are 
included in a broad fascia extending across to second claval vein, a 
broken diffuse fascia from fork of Cui to stigma, apical cells of Sc, R and 
M at least at margin and a narrow interrupted fascia following distal cross- 
veins, fuscous or fuscous-piceous. Wing infuscate, veins fuscous. 

Seventh sternite transverse posteriorly, tumid sublaterally, and with an 
eminence medially, hollowed out on its posterior face. 

Eighth segment with ventro-lateral pieces directed medially, tapering 
to bluntly-rounded apex. 

Post-tibiae with a single spine near base, six short teeth and one long 
at apex, basal metatarsus with six apical teeth, second metatarsal also 
with six. 

One female collected from Mctasequoia gJyptostrohoides, 1,000 m. Sui- 
sapa, Lichuan District, W. Hupeh, China, Aug. 21, 1948, Gressitt. This 
species is distinguished from all described species of Magadha by colora- 
tion and from Kempiana maculata Muir by the absence of a pre-costal 
area in the tegmen, by coloration of the tegmina, and by size. 

Magadha denticulata Fennah, new species. 
(Figure 14, A-D.) 

Male: length, 3.6 mm.; tegmen, 5.0 mm. Female: length, 3.2 mm.; teg- 
men, 5.0 mm. 

Vertex about 1.5 times as broad across base as long in middle line. Teg- 
mina with Sc + R folk slightly distad of Cui fork. Mi + 2 not forked before 
distal transverse line of cross veins. Post-tibiae with a tooth laterally in 
basal third, seven-toothed at apex, basal metatarsal segment 7-toothed, 
second metatarsal 6-toothed. 

Fuscous; vertex at base and apex, six spots on each lateral margin of 
frons and a short transverse bar one-fifth from apex, clypeus at base and 
apex, carinae of pronotum, lateral angles of mesonotum and apex of scu- 
tellum, femora at apex, and tibiae at middle and apex stramineous or pallid 
ochraceous, mesonotal disc with a pair of more or less distinct ocellate 
spots at basal third. Tegmina sordid ochraceous, veins including costa, pale 
regularly spotted with fuscous, two spots in costal cell, stigma, apex of 
clavus, and apical cells at base and apex, fuscous. Wings infuscate, veins 

Anal segment of male triangular, slightly broader than long, Pygofer 


with lateral martiin with a semilunate lobe, inedioventral process bifid, each 
lobe aeiiininate. Aedeagus produced laterally on left side at base in a short 
plate, toothed on margin, medially and dorsally at base a vertical plate, 
slightly inclined to right, phallobase distally broad, produced laterally 
on right in a subtriangular process, toothed at its apex. 

Posterior margin of seventh sternite transverse. Ventrolateral angle of 
posterior margin of eighth segment in ventral view only slightly acute. 

Two males (one the type) and six females, 1,000 m. Suisapa, Lichuan 
District, W. Hupeh, Aug. 20-24, 1948, Gressitt. In this species the latero- 
basal facets of the f rons are very feebly demarcated on their ventral mar- 
gin. In the males there is a feeble round impression; the texture of the 
anterior third of the mesonotum differs from that of the posterior two- 
thirds. The species differs from typical Magadha and also from Francesca 
in venation, as Mi + 2 does not fork basad of the distal transverse line. 
From Francesca it also differs in the broader vertex. It is close to 31. for- 
mosana Mats. 

One male, 250-350 m., Tso-kok-wan, Lungtau Shan, Kwangtung Prov- 
ince, June 9, 1947, Gressitt is also placed here. 

Magadha cervina Fennah, new species. 
(Figure 15, A-E.) 

Male : length, 3.0 mm. ; tegmen, 4.0 mm. 

Laterobasal triangular facets of frons moderately distinct. Post-tibiae 
laterally unispinose, apically 8-spined, basal metatarsal segment 7-spined, 
second metatarsal 6-spined. 

Reddish-brown; six spots along lateral margins of frons, apex of clypeus, 
a spot on sides of clypeus, pronotal areolets, a stripe on post-tibiae near 
base, fuscous; hind legs otherwise stramineous. Tegmina translucent, pow- 
dered sordid greyish-yellow, veins concolorous, regularly spotted fuscous, 
a few small round spots in membrane, apical cells at base and apex fuscous. 
Wings slightly infuscate, veins darker. 

Anal segment of male much broader than long, lateroapical angles de- 
curved and produced in a stout spine. Pygofer with medioventral process 
broad, its lateroapical angles produced into short lobes, distally incurved, 
apical margin excavate with two minute eminences near middle. Aedeagus 
with phallobase four-lobed, as figured, two lobes simple, spinose, one with 
two spines and one with three and a blunt eminence; phallic appendages 
minutely denticulate over whole of distal surface. Genital styles bluntlj^ 
ovate, a large three-toothed lobe dorsally near base. 

One male 1,500 m., Shin-kai-sze, Omai Shan ; Szechuan, W. China, Aug. 
16, 1940, Gressitt. This species is distinguished by coloration and by the 
shape of the male genitalia. 



Genus Plectoderoides Matsumura 
Matsumura, 1914:281. Ortliotype, Plectoderoides maculatus Mats., 1914:282 

Plectoderoides flavovittatus Feiniah, new species. 

(Figure 14, E-I.) 

Male: length, 2.8 mm.; tegmen, 3.5 mm. Female: length, 3.0 mm.; teg- 
men, 4.8 mm. 

Post-tibiae with a spine laterally near base, seven spines at apex. Basal 
metatarsal segment with seven spines at apex, second segment with six. 

Yellow stramineous; a transverse bar on frons at base and at apex, a 
similar bar across clypeus, two bars on genae, disc of vertex on each side 
of median carina, pronotum with a stripe on each side of median carina, 
a stripe behind eyes, and another across lateral lobes, mesonotum, except 
laterally, a spot medioposteriorly on tegulae, bases of pro- and mesocoxae, 
two bands on each mesopleuron, dark reddish-brown. Tegmina translucent- 
fuscous, costal and subcostal cells to stigma, apical veins, cross veins and 
apical margin, inner half of clavus along the whole of its length, except 
for three or four spots on anterior claval vein, pale yellow, veins, apart 
from preceding, concolorous. Wings infuscate, veins darker. 

Anal segment of male moderately short, apical margin excavate, latero- 
apieal angles produced, bluntly rounded. Pygofer with lateral processes 
of each side broadly subtriangular, medioventral process long, distally lat- 
erally compressed. Phallobase with a pair of sinuate lobes dorsolaterally, 
that on left broader distally, unornamented, that on right narrower and 
more heavily sclerotised; a pair of lobes ventrolaterally, that on left sub- 
triangular distally with ventral angle directed cephalad and with small 
spines at apex, that on right elongate, narrow, minutely denticulate on 

Fig. 14. Magadha denticulata. new species: A, medioventral process of pygofer; 
B, anal segment of male, dorsal view; C, apical portion of aedeagus, ventral view; 
D, denticulate process of left side of aedeagus near base. Plectoderoides flavovit- 
tatus. new species: E, anal segment of male, dorsal view; F, lateral process of 
pygofer; G, medioventral process of pygofer; H, aedeagus (ventral side upper- 
most) ; I, left genital style, inner aspect. 



[Proc. 4th Ser. 

upper margin. Aedeageal processes long, ribbon-like, obliquely truncate 
and minutely serrate at apex. 

Genital styles elongate-triangular, a long curved knobbed process on 
inner face near base, a broad curved three-cusped lobe at middle of dorsal 

Anal segment of female short. Pregenital sternite posteriorly transverse. 
Ovipositor with first valvulae four-spined, ventral lobe triangular, acute 
at apex; third valvulae broadly ovate, incurved distally. Bursa copulatrix 
armed at entrance with a triangular closely-grooved sclerite, supported on 
a slender wide crescentic base. 

One male (the type) and one female, Mokansan, Che Kiang Province, 
China, Mrs. D. E. Wright, Sept. 1947. This species is distinguished by the 
shape of the genitalia and by coloration. 

Plectoderoides uniformis Fennah, new species. 
(Figure 15, F, G.) 

Male: length, 2.9 mm.; tegmen, 4.2 mm. Female: length, 3.3 mm.; teg- 
men, 5.1 mm. 

Post-tibiae laterally unispinose, seven-spined at apex. 

Stramineous, head and mesonotum tinged with reddish-brown, abdomen 
dorsally fuscous. Tegmina sordid yellow, translucent, subapical and apical 





Fig. 15. Magadha cervina, new species: A, anal segment of male (lateral lobes 
spread out); B, medioventral portion of hind margin of pygofer; C, left genital 
style; D, aedeagus (phallobase) ; E, aedeagal (phallic) appendage. Plectoderoides 
uniformis, new species: F, lateral process of hind margin of pygofer; G, aedeagus, 
left side. Caristianus ulysses Fenn.: H, right genital style; I, medioventral process 
of pygofer; J, anal segment of male, dorsal view; K, aedeagus, dorsal view. Ako- 
trojiis fuviata impersonata. new subspecies: L, apical portion of anal segment, 
dorsal view; M, medioventral process of pygofer; N, dorsolateral lobe of hind mar- 
gin of pygofer ; 0, apical portion of aedeagal processes, right side. 


cells more or less completely infuscate, veins concolorous except marginal 
veinlets near stigma and transverse veins which are pallid. Wings lightly 
infuscate with darker veins. 

Pygofer with laterodorsal angles narrow and finger-like. Phallobase 
with a pair of elongate lobes dorsolaterally, each serrate on upper and 
lower margins; mesad of, and slightly below these a pair of elongate unor- 
namented lobes rounded-truncate at apex; a pair of broad smooth ventral 
lobes with ventral margins apposed, united distally and reflected ventro- 
cephalad apically in a median spine; aedeageal lobes elongate, s;vTiimetri- 
cal, acuminate, apparently slightly more compressed laterally before apex. 
Genital styles of similar profile to those of Caristianus, a small blunt emi- 
nence on inner face near base. 

One male, Mokansan, Che Kiang Province, China, Sept. 28, 1947, Mrs. 
D. E. Wright. One female, 1,000 m., Suisapa, Lichuan District, W. Hupeh, 
China, July 25, 1948, Gressitt is referred to this species. This species is 
distinguished by coloration. 

Genus Caristianus Distant 
Distant, 1916:63. Orthotype, Caristianus indicus Distant, 1916:63 

Caristianus ulysses Fenn. 
(Figure 15, H-K.) 
Fennah, 1949:600. 

Male: length, 3.0 mm.; tegmen, 3.4 mm. 

Post-tibiae laterally unispinose, eight-spined at apex, basal metatarsal 
segment seven-spined, second metatarsus with six spines. 

Dark reddish-brown, carinae of vertex, except for two stripes laterally, 
lateral margins of frons, except for four stripes, a transverse band across 
apex of frons and base of clypeus, clypeus at apex, rostrum, discal carinae 
and ventral margin of lateral lobes, median carina of mesonotum, and lat- 
eral carinae suffusedly, legs and metapleurites stramineous. Tegmina dark 
reddish brown, an irregular spot in basal half of costal cell, distal half of 
costal cell, two areas in anterior half of subcostal cell, stigma, veinlets 
of R at margin, a spot at union of claval veins, and six other small spots 
on claval margin, cross-veins in Cu faintly, pallid ivory. 

Anal segment of male about as long as broad, distal margin convex, 
notched at middle. Pygofer with each lateral margin produced near mid- 
dle in a slender curved digitate process, medioventral process deeply bifid, 
each limb twisted and compressed near apex. Aedeagus narrowly tubular, 
shortly cleft medially at apex with a short horizontal lobe dorsally at apex 
on each side, forming a slight hood. 

Genital styles subquadrate, sinuate on ventral and dorsal margins, with a 
simple triangular eminence near middle of dorsal margin. 

One male, 6,500-7,500 ft., West Hills, Yunnan fu, Yunnan, S. China, 


Aug. 21-22, 1934, Ernest 11. Tinkham. This is apparently the geographical 
representative of the Bornean species, known only from a single female: 
it differs from typical C. ulysses in the mesonotal disc being wholly dark, 
the pronotal disc narrowly infuscate between the carinae, and the lateral 
carinae of the frons transversely striped. 

Genus Akotropis Matsumura 
Matsumura, 1914:270. Logotype, Akotroins fumata Matsumura, 1914:270 

Akotropis fumata impersonata Fennah, new subspecies. 
(Figure 15, L-0.) 

Male: length, 2.9 mm.; tegmen, 3.3 mm. Female: length, 2.8 mm.; teg- 
men, 3.5 mm. 

Coloration as in typical subspecies, except for absence of any dark mark- 
ing between eyes and lateral carinae of frons. Genitalia as figured. Post- 
tibiae laterally unispinose, distally 8-spined. BavSal metatarsal segment 
seven-spined, second metatarsal six-spined. 

One male (the subspecific type) and 5 females, Mokansan, Che Kiang 
Province, China, Aug. 4-Sept. 28, 1927, Mrs. D. E. Wright; one male and 
two females, 1,000 m. Suisapa, Lichuan District, W. Hupeh, China, Aug. 
19, 1948, Gressitt. In the specimens from Suisapa the ground color of the 
tegmina is distinctly darker than in the Mokansan series. 

Akotropis flaveola Mats. 

Matsumura, 1914:271. 

One female. Riviere de Quangtri, Anam, April 30, 1927, Mrs. D. E. 
Wright, is tentatively referred to this species. There is a slight curved 
fuscous stripe in the region of the ocelli. 

Genus Zathauma Fennah 
Fennah, 1949:605. Orthotype, Zathauma cristatum Fennah, loc. cit. 

Zathauma metasequoiae Fennah, new species. 

Female: length, 5.8 mm.; tegmen, 8.0 mm. 

Vertex broader across base than long in middle line (3:1), postero- 
medial portion of disc declivous, remainder hollowed out; frons in middle 
line longer than greatest width (1.8: 1), and longer than clypeus in mid- 
dle line (1.3: 1), greatest width of frons 1.7 times width at base; rostrum 
with apical segment 1.6 times as long as subapical, attaining post-trochan- 
ters. Pronotum with disc twice as broad across base as long in middle line, 
longer in middle line than vertex in same line (1.7: 1) depressions laterad 
of disc very feeble; two carinae on each side between eye and tegula; 
mesonotum tricarinate, carinae prominent except on scutellar area, which 
is medially ecarinate, lateral carinae diverging from apex to base, anterior 


portion of mesonotiim of different textnre from posterior; tegulae large, 
not carinate; protibiae slightly exceeding profemora, post-tibiae with a 
spine at basal third, six small spines and one large spine at apex, basal 
metatarsal segment with 6 teeth, the outermost largest, second metatarsal 
segment with two large outer teeth and a row of four sliort teeth between 
them; post-coxae produced lateroposteriorly in a short spine. 

Basal half of clypeus, sides of head below antennae, all femora at apex, 
all tibiae at base and apex creamy yellow, frons except for a few pale 
fuscous spots at margins distally, vertex, pronotum, except lateral lobes, 
anterior part of mesonotal disc, mesonotum laterally, and tegulae, testa- 
ceous-stramineous; distal part of clypeus, rostrum, lower side of thorax, 
posterior part of mesonotal disc, legs, and abdomen dark fuscous. Tegmina 
creamy-white at extreme base, elsewhere fuscous-piceous sparingly mar- 
bled with grey, veins concolorous with small pallid spots, and pallid apex 
at margin, transverse veins mostly pallid, pallor most pronounced in first 
branch of Sc at node; wings smoky. 

Pregenital sternite large, fully as long as fifth and sixth ventrites com- 
bined, hind margin shallowly convex. Subvaginal plate about as long as 
broad, its sides concave, mesal margins of ventrolateral parts of eighth 
segment with a submarginal channel basally; posterior angle subrectan- 

One female, from Metasequoia fjlyptosfrohokhs, 1,000 m., Suisapa, Li- 
chuan District, W. Hupeh, Aug. 21, 1948, Gressitt. This species is larger 
than the type, has a dift'erently-shaped vertex and lacks the foliately raised 
mesonotal carina. The cephalic differences, however, are no greater than 
those in Faventilla, and in other characters, including tegminal venation 
and coloring, it generally resembles Z. cristatum. 

Genus Betatropis Matsumura 

Matsumura, 1914:274. Orthotype, Betatropis formosana Matsumura 

Betatropis formosana Matsumura. 
Matsumura, 1914:274. 

Eight males and 10 females, Mokansan, Che-Kiang Province, China, 
Aug. 26-Sept. 24, 1927, Mrs. I). E. Wright. 

Genus Tangina Melichar 

Melichar, 1903:223. Haplotype, Tangina hiimnctata Melichar, 1903:44 


(1) (2) Vertex with two black or intuscate spots at apex (3) 

(2) (1) Vertex without such spots (5) 

(3) (4) Tegmina translucent, pallid yellow. Luzon T. quadripunctata Mel.f 


(4) (3) Tegmina pale yellow with two oblique dark stripes at apex of costal cell, 

and a black spot in first apical cell of Sc, which is bounded by black 
veins. Ceylon T. hipunctata Mel. 

(5) (6) Tegmina pale, a longitudinal stripe on Sc + R and another along sutural 

margin black, membrane infumed ; a black spot on propleura. Luzon 

T. quadrilineata Mel. 

(6) (5) Tegmina tinged fuscous, a broad band along anterior margin white, a 

dark spot in first and second infumed apical cells; a black spot on 
mesopleura. China T. sinensis, new species 

t T. modesta Haupt is probably not a member of this genus. It is distinguished from T. quadripunctata 
by the absence of two piceous spots on the pronotum and its generally darker hue. 

Tangina sinensis Fennah, new species. 

Male: length, 2.5 mm.; tegmen, 3.0 mm. Female: length, 2.8 mm.; teg- 
men, 3.4 mm. 

Post-tibiae 8-toothed at apex ; basal metatarsal segment 6-toothed distally, 
second metatarsal segment 5-toothed. 

Creamy-white; a large round spot on mesoplenra piceons; mesonotum 
tinged yellow, distal half of abdomen dorsally and ventrall}^ orange-yellow. 
Tegmina translucent, pale fuscous, a band of even width along costal mar- 
gin to beyond stigma, extending inward to middle of cell Sc + R, white; 
a minute spot in first infuscate apical areole beyond stigma and a larger 
round spot in adjoining apical areole, piceous. Wings hyaline, powdered 

Pygofer with medioventral process deeply cleft medially. Genital styles 
triangular in profile, a triangular spinose process at middle of dorsal mar- 
gin, directed laterad, and a similar process at apex directed dorsocephalad. 
Phallobase relatively long, with a pair of narrow tapering lobes dorsally, 
decurved at apex, minutely denticulate on ventrolateral margin, medio- 
ventrally a long straight spine directed cephalad. 

Pregenital sternite of female posteriorly shallowly convex. 

One male and 1 female, Mokansan, Che-Kiang Province, China, Sept. 
2, 19, 1927, Mrs. D. E. Wright. This species differs from T. hijmnctata in 
the absence of piceous marks on the head, of fuscous lateral mesonotal 
fields, and in the general infuscation of the tegmina coupled with the ab- 
sence of oblique dark lines near the stigma. 



(1) (2) Femora and tibiae, or merely lower angle of femora more or less widened 

into a flange (3) 

(2) (1) Femora and tibiae not at all widened (7) 

(3) (4) Vertex fully as long as pronotum and mesonotum combined; profemora 

with a small tooth subapically ; post-tibiae with 5 spines (5) 


Vertex not nearly as long as pronotum and mesonotum combined. Post- 
femora unarmed; post-tibiae 6-7 spined Ortliopagus Uhl. 

Cephalic process with two slight constructions, its apex bombinate 

Piela Lall. 

Cephalic process not constricted nor apically bulbous Saigona Mats. 

Vertex with cephalic process more than three times as long as broad... (9) 

Vertex not more than three times as long as broad (13) 

Head much narrower than pronotum; cephalic process slender, scarcely 

widened at apex, porrect. Post-tibiae with four spines 

Thanatodictya Kirk. 

Head not greatly narrower than pronotum, cephalic process relatively 
stout (11) 

Cephalic process with lateral margins feebly defined, transversely and 
irregularly rugose on upper surface. Post-tibiae l-SY>ined.. ..Leprota Mel. 

Cephalic process with lateral margins strongly carinate, median carina 
present only near base, upper surface slightly concave longitudinally. 
Post-tibiae 4-5 spined Chanithus Kol. 

Vertex narrow throughout, 2.5-3 times as long as broad, its lateral mar- 
gins sinuate (15) 

Vertex not narrow throughout, relatively broad between eyes, not more 
than twice as long as broad (17) 

Lateral carinae of frons widest apart between eyes Togaphora Mats. 

Lateral carinae of frons widest apart near fronto-clypeal suture 

Avephora Bierm. 

(17) (18) Carinae of pronotal disc strongly developed Tropidophara Bierm. 

(18) (17) Carinae of pronotal disc almost obsolete Sinodictya Mats. 

No Chinese species are known to the w^riter which fall into the restricted 
concept of Didyophara Germar (Fennah, 194-4: 81, 82, 90). 

Genus Orthopagus Uhler 
Uhler, 1896; 278. Haplotype, Orthopagus lunulifer Uhler 

Orthopagus helios Melichar. 
Melichar, 1912:60. 

Two males, Mokansan, Che Kiang- Province, China, (Mrs. D. E. Wright, 
Sept. 8, 1927) agree with the description of 0. Jielios Mel.; 1 female, Tai- 
hanroku, agrees exactly with the description of 0. elegans Mel.; 1 male and 
1 female, Taihanrokn, Japan (H. Salter, July 22, 1908), agree exactly 
with the description of 0. helios var. diffusus Mel., which was described 
from Taihanroku material. The proportions of the vertex ( length : breadth ) 
in the above material is 1.7 : 1. Even with the present short series there 
seems little room for doubt that all the above material is conspecific. 

One male, Nai-suen, 21 m. S. E. of Naam-fung, Lin Kao District, Hainan 
Island, S. China (Aug. 31, 1932) differs from the preceding in having the 














(11) (12) 

(12) (11) 

(13) (14) 

(14) (13) 

(15) (16) 

(16) (15) 


vertex not more than 1.6 times as long as broad and the lateral margins of 
the frons relatively straight. This may prove to be specifically distinct 
from the preceding, but the constriction at the middle of the vertex and 
the proportion of the produced relative to the basal portion debar it from 
being placed in the splendens-fleicheri section of Melichar's key. 

Genus Chanithus Kolenati 
Kolenati, 1857:427. Haplotype, Flata pannonica Gennar, 1830:47 

Chanithus gramineus (F.). 

Fulgora graviinea Fabricius, 1803:4. 
Dictyophor-a sinica Walker, 1851:321. 

Eight males, 7 females, Mokansan, Che-Kiang Province, 24, Sept. 6, 1927, 
Mrs. D. E. Wright; 3 males, 2 females, Lau-Chi, Che Kiang Province, July 
9, 12, Mrs. D. E. Wright; 2 males, 1 female, Tungiu, Che Kiang Province, 
Sept. 8, 10, 1926, Mrs. D. E. Wright; 2 males and 2 females, 800-1,000 ft., 
Chang Tau Ching, Szechwan, July 18, 1948, Gressitt; 3 females, 1,000 m., 
Suisapa, Lichuan District, W. Hupeh, Aug. 19, 20, 25, 1948, Gressitt. 

The intensity of pigmentation is slightly variable, but no geographical 
subspecies can be recognized in the above series. 

Genus Tropidophara Bierman 
Bierman, 1910:15. Haplotype, Tropidophara dubiata Bierman, 1910:16 

Tropidophara javana (Lethierry), 
Dictyophara javana Lethierry, 1888:467. 

One male, between Limcliow and Kung-kon, Hoh-p'u District, Kwang- 
tung, S. China, Aug. 5, 1932, W. E. Hoffman. 

Genus Thanatodictya Kirkaldy 
Klrkaldy, 1906:392. Haplotype, Dictyophara jjraef errata Distant, 1892:279 

Thanatodictya lineata (Donovan). 

Fulgora lineata Donovan, 1800:1, pi. 8, fig. 1. 

One female, Tin T au Village, Lam Lo District, Hunan Province, S. 
China, July 29, 1934. 

Genus Avephora Piierman 
Bierman, 1910:12. Haplotype, AiTephora pasteuriana Bierm., 1910:12 

Avephora eugeniae (Stal). 

Pseudophana eugeniae Stal, 1859:271. 
Avephora pasteuriana Bierman, 1910:12. 

One male, Tai Kwong village. Lam Ho District, Hunan Province, S. 
China, July 26-28, 1934; 1 male, Hokcow (near Leokay, Tonkin), Yunnan, 


S. China, Aug. 16, 1934, Chauneey Brownall; 1 female, White Cloud Moun- 
tain, Canton, China, July 6, 1932. 

Family FULGORIDAE Latreille 

(1) (2) Cephalic process porrect, stout and distally rounded, much longer than 

pronotum and mesonotum together Pyrops Spin. 

(2) (1) Cephalic process much shorter than pronotum, or with a slender append- 

age (3) 

(3) (4) Cephalic process, at least at base, strongly recurved dorsad and overlying 

apical margin of vertex (5) 

(4) (3) Cephalic process very short, directed dorsad, with a small shallow depres- 

sion at apex to which a slender rod-like appendage is weakly attached; 
anterior femora ampliate near apex Kalidasa Kirk. 

(5) (6) Carinae of frontal disc weak, near base separated almost by twice width 

of eye, converging strongly distad Penthicodes Blanch. 

(6) (5) Carinae of frontal disc subparallel, obsolete in distal third. ...Z/T/corma Stal 

Genus Lycorma Stal 
stal, 1863:232. Logotype, Aijhana imiierialis White, 1846:330 

Lycorma delicatula (AVhite). 

Aphaena delicatula White, 1845:37. 

Post-tibiae laterally 5-spine(l, apically 7-spined. 

Three males and 2 females, 1,000 m., Suisapa, Liehuan District, W. 
Hupeh, China, Aug. 21, 1948, Gressitt. 


(1) (2) Tegmina leathery, brown, with numerous and irregular cross veins; if 

subhyaline distally, distal area not demarcated basally by a distinct 
line of transverse veinlets (3) 

(2) (1) Tegmina hyaline; if of a denser consistency then cross-veins relatively 

few and regular, or membrane abruptly and very closely reticulate.... (5) 

(3) (4) Tegmina with Cu^ forked just distad of union of claval veins; wings 

reduced Padanda Dist. 

(4) (3) Tegmina with Cui not forked as described. Wings normal. ...Olontheus Jac. 

(5) (6) Tegmina with not more than nine cells at apical margin (7) 

(6) (5) Tegmina with more than nine apical cells (15) 

(7) (8) Frons setose. Longest apical cell in tegmina longer than clavus 

Trichoduchus Bierm. 

(8) (7) Frons not setose. Longest apical cell relatively shorter (9) 

(9) (10) Frons unicarinate. Tegmina with a single row of transverse veinlets 

Ommatissus Fieb. 



[Proc. 4tii Ser. 

(10) (9) Frons tricarinate. Tegmina with two rows of transverse veinlets, or trans- 

verse veinlets few (11) 

(11) (12) Tegmina with nodal line straight, distinct, with one row of transverse 

veins distad of it Zema, new genus 

(12) (11) Tegmina with nodal line not demarcated, transverse veins rather irregu- 

lar, straight or oblique (13) 

(13) (14) So + R and Cu^ simple in basal half of tegmina Cixiojisis Mats. 

(14) (13) So + R and Cuj forked in basal half of tegmina Duriopsis Mel. 

(15) (16) Tegmina with pre-costal area traversed by distinct veinlets (17) 

(16) (15) Tegmina with costal vein at margin, or if submarginal, then without 

distinct transverse veinlets (23) 

(17) (18) Tegmina with venation of distal third irregular, densely reticulate 

Mesepora Mats. 

(18) (17) Tegmina with venation of membrane not as above (19) 

(19) (20) Tegmina with Cuj forked basad of level of union of claval veins; nodal 

line situated two-thirds of length of tegmen from base (21) 

(20) (19) Tegmina with Cuj forked distad of union of claval veins; nodal line only 

slightly distad of middle of tegmen Eodryas Kirk. 

(21) (22) Second antennal segment cylindrical, more than twice as long as first 

Catullia Stal 

(22) (21) Second antennal segment short, not as above Epora Wlk. 

(23) (24) Tegmina basad of nodal line of cross-veins distinctly thicker than in 

apical portion (25) 

(24) (23) Tegmina of uniform consistency throughout (29) 

(25) (26) Vertex longer than pronotum and mesonotum together, sides of head dis- 

tinctly constricted just before eyes Ossoides Bierm. 

(26) (25) Vertex not so produced (27) 

(27) (28) Frons weakly ampliate distally, lateral margins and median carina nor- 

mal ; lateral carinae of pronotal disc almost parallel with sides of vertex 
Tambinia Stal 

(28) (27) Frons strongly ampliate to below level of antennae, lateral margins and 

median carina at base thickened, lateral carinae of pronotal disc curv- 
ing outward toward tegulae Kallitaxila Kirk. 

(29) (30) Vertex approximately twice as long as broad in middle (31) 

(30) (29) Vertex relatively shorter, not or scarcely longer than broad (33) 

(31) (32) Tegmina with Sc -f R forked once shortly before nodal transverse line, 

M and Cu^ simple Tauropola Jac. 

(32) (31) Tegmina with Sc + R three-branched at nodal line, Cu^ forked near level 

of union of claval veins Sioezeyaria Mete. 

(33) (34) Vertex anteriorly transverse; pronotal disc large; tegmina long with 

nodal line slightly basad of middle; Cu^ forked before nodal line 

Sogana Mats. 

(34) (33) Vertex anteriorly convex or acute; pronotal disc small; tegmina with 

nodal line not basad of middle; Cu^ simple basad of nodal line 

Neommatissus Muir 


In the above synopsis, Chinese species which have been referred to Kalli- 
famhinia ^Vliiir will run to Tmnhinia Stal, where they may well be placed 
until the limits of the former genus have been more precisely defined: 
Muir's concept, as interi)reted from the type species, is restricted to forms 
witii trispinose post-til)iac and claval veins which unite distad of the middle 
of the clavus. In Tamhinia the post-tibiae are normally bispinose and the 
claval veins unite basad of the middle of the clavus. 

Nacmusius Jac. (1944: 19) runs to Padanda Dist., and the writer can 
trace no generic character in which they differ. (It may be noted paren- 
thetically that the holotype of the African Padanda denti Muir is an issid.) 
Notwithstanding its hiraciine appearance, the writer suspects that Padanda 
is related to a group of genera near Ommatissus Fieb. and including Cixi- 
opsis Mats., Duriopsis Mel. (assigned by its author to Issidae), and a new 
genus described below. Perhaps OJontheus is likewise related to this group. 

Parahiracia Ouchi (1940: 299), which was described as a hiraciine tropi- 
duchid, is here considered to belong to the Issidae. 

Genus Catullia Stal 
Stai, 1870:748. Haplotype, Catullia suMestacea Stal. 

Catullia subtestacea Stal. 

stal, 1870:749. 

One male, Cheung-nga San, Tin-tong, Loh-chang District, Kwangtung, 
Sept. 9, 1947; 1 female, Tai-ka, Tin-tong, Loh-Chang District, Aug. 20, 
1947; 1 female, Naam-kong-paai, Yao shan, Yang-shan District, Kwang- 
tung, Oct. 29-30, 1934; 1 male, Tai Kwong village. Lam Mo District, 
Hunan Province, S. China, July 26-28, 1934; 1 male, Kwei-Hsien, Kwei- 
Hsien District, Kwangsi, S. China, July 28-29, 1934, E. R. Tinkham; 2 
males and 2 females, Mokansan, Che-Kiang Province, Sept. 2, 6, 22, 1927, 
Mrs. D. E. AVright. 

Genus Kallitaxila Kirkaldy 
Kirkaldy, 1901:6. Orthotype, Kallitaxila granulata Stal 

Kallitaxila granulata Stal. 

ITaxila] granulata Stal, 1870:750. 

One female, Honam Island, P'an-yu District, Canton, June, 1935, W. E. 

Genus Sogana Matsumura 
Matsumura, 1914:268. Haplotype, Sogayia liopponis Mats. 

Sog-ana hopponis Matsumura 
Matsumura, 1914:268. 

One female, 800 m., Hori (Pull. Polisia), Taichung District, Formosa, 
Aug. 23, 1947, L. Gressitt. 


Genus Ossoides Biermaii 
Bierman, 1910:26. Haplotype, Ossoides linratus Bierman 

Ossoides lineatus Biennan. 
Bierman, 1910:27. 

One female, White Cloud Mountain, P'an-yu District, Canton, Dec. 16, 

Genus Neommatissus Muir 
Muir, 1913:267. Orthotype, Neommatissus spurcus Muir 

Neommatissus congruus (Walker) 

Brixia congrua Walker, 1870:110. 
Neommatissus spurcus Muir, 1913:268. 

One female, Sao-tio, Tin-tong, Loh-chang District, Kwangtung, Aug. 23, 

Genus Eodryas Kirkakly 

Kirkaldy, 1907:93. Haplotype, Epora subtilis Mel. not Walk. 1903: 

=^ E. melichari (Dist. ) 

Eodryas melichari Distant. 
Gen. ? melichari Distant, 1906:285. 

One female, Honam Island, Canton, July 12-14, 1932; 1 female, Tai- 
pingfu, Sung-shen District, Kwangsi, Aug. 5-6, 1934, E. R. Tinkham. 

Genus Ommatissus Fieber 
Fieber, 1875:353. Haplotype, Ommatissus Mnotatus Fieb., 1876:174. 

Ommatissus lofouensis Muir. 
Muir, 1913:267. 

Post-tibiae laterally bispinose, apically 8-toothed. Basal metatarsal seg- 
ment 7-tootlied. 

Aedeagus longer than post-femora, comprising a pair of long, slender 
ribbon-like i)rocesses directed caudad, acuminate at apex, and an even 
longer median cylindrical process gradually decurved distad but curved 
upward at distal fifth to point dorsad, apex not acuminate. Genital styles 
elongate-triangular, bluntly rounded distally, a small spine, curved laterad 
dorsally near base. 

One male, Mokansan, Che Kiang Province, Aug., 1927, Mrs. D. E. Wright. 
There is no doubt that this species is congeneric with 0. Mnotatus Fieb. 

Zema Fennah, new genus 

Vertex about twice as broad as long, anterior margin obtusely angulately 
convex, posterior margin correspondingly rounded-concave, lateral mar- 


gins slightly converging distad, median carina present only in basal two- 
thirds, an impression at each side on disc; frons in middle line longer than 
greatest width (1.3:1), basal marg^in transverse, lateral margins diverg- 
ing to below level of antennae, thence shallowly incurved; a transverse 
callus across frons at base, with median and two lateral discal carinae 
arising from it, the latter enclosing an oval which is widest about level of 
lower margin of eyes; elypeus about two-thirds as long as frons, not dis- 
tinctly carinate but with median area raised. Pronotum in middle line 
rather longer than vertex in same line, anterior margin rounded convex, 
posterior margin subangulately concave with a distinct notch at middle; 
disc tricarinate, an impression on each side of middle line, lateral carinae 
diverging caudad, anteriorly evenly curving mesad into anterior margin, 
a single carina on each side between eye and tegula; mesonotum broader 
than long, median carina reaching to seutellum, which is not divided from 
disc by a groove, lateral carinae evenly curved mesad anteriorly. Post- 
tibiae with four spines laterally, and eight small spines at apex; basal 
metatarsal segment with about nine teeth at apex, second metatarsal seg- 
ment short, with a spine at each apical angle and a convex pad between them. 

Tegmina about three times as long as broad, widest at level of nodal 
line, costal margin and sutural margin subparallel, the former only weakly 
incurved at base. Sc + R, ]\I, and (.'u united to level of nodal line, distad 
of nodal line a single distinct row of transverse veins; claval suture present, 
flexible, claval veins united at middle of clavus. Wings ample, Sc + R, M, 
and Cui each forked once. 

Ovipositor with first valvulae denticulate on ventral margin with two 
blunt teeth at apex, a single relatively large tooth at apex, and two blunt 
teeth on dorsal margin; third valvulae with two or three teeth on dorsal 
margin and about five teeth on apical margin. 

Ty]5e species, Zema gressitti, new species. 

The present concept is separated from Cixiopsis by the shape of the frons 
and the tegminal venation : in Cixiopsis the frons is narrowed in the mid- 
dle, and the lateral discal carinae unite with the median carina in a point; 
the lateral carinae are foliate, as is visible from above, and the elypeus is 
medially carinate. In the tegmina the claval veins unite two thirds from the 
base of the clavus, and there is a network of veinlets on the distal third. 
Prom Ommatissus it is separated, in addition to other characters, by the 
carination of the frons and by the shape and venation of the tegmina : in 
Omynatissus the disc of the frons is unicarinate, while the tegmina are more 
acutely curved at the apical margin; moreover Cui forks on the corium 
near the union of the claval veins. It differs from Padanda in the shape of 
the head and pronotum and in tegminal venation : in Padanda the frons 
is less narrowed basally and the lateral discal carinae begin their mesad 



[Proc. 4th Ser. 

curvature i-ather nearer the base : in the tegmina of Distant's holotype 
Sc + R and M are simple on the corium, but Cui is forked slightly distad 
of the union of the elaval veins; moreover numerous irregular transverse 
veinlets are present and form a lax reticulum. The type of P. atkinsoni is 
relatively short-winged : the elaval suture is evident, but is not a functional 
line of flexure: the wings are extremely reduced. 

Zema gressitti Fennah, new species. 
(Figure 16, A-F.) 

Female: length, 5.0 mm.; tegmen, 4.9 mm. 

Vertex anteriorly broadly rounded. Tegmina with elaval suture present 
and functional; venation regular; wings fully developed. 

Testaceous to dark tawny : intercarinal area of vertex, a spot overlying 
each impression on pronotal disc, dorsal portion of lateral lobes of pro- 
notum, mesonotum except laterally, middle of clypeus, tip of rostrum, 
thoracic pleurites and abdominal sclerites f uscous-piceous ; a broad band 
overlying fronto-clypeal suture, extending across sides of head, stramineous, 
ventral portion of lateral lobes of pronotum, ivory-white; most of frontal 
disc, stripes along all femora and tibiae, pro- and mesotarsi fuscous. Third 
valvulae of ovipositor inceous dorsally, translucent testaceous ventrally. 

One female, Lung-chi-pa, Szechuan-Hupeh border, July 19, 1948, 

Family ISSIDAE Spinola 

(1) (2) Tegmina usually brachypterous. Basal metatarsal segment with two 

spines (3) 

(2) (1) Tegmina macropterous, covering abdomen completely. Basal metatarsal 

segment with more than two spines (7) 

(3) (4) Profemora and protibiae foliate in male, compressed and weakly foliate 

in female Caliscelis Lap. 

Fig. 16. Zema gressitti. new genus and species: A, head and thorax; B, frons 
and clypeus; C, head and thorax, side view; D, tegmen; E, wing; F, pregenital 
sternite of female. 



(4) (3 

(5) (6 

(6) (5 

(7) (8 

(8) (7 
(9) (10 

(10) (9 
(11) (12 
(12) (11 
(13) (14 

(14) (13 

(15) (16 
(16) (15 
(17) (18 
(18) (17 

(19) (20 
(20) (19 
(21) (22 

(22) (21 

(23) (24 

(24) (23 
(25) (26 

(26) (25 
(27) (28 

Legs normal, or only protibiae widened distally at external angles (5) 

Second segment of antennae transverse at apex with arista inserted 

apically and projecting in same line as axis of second segment 

Ovimatidiotus Spin. 

Second segment of antennae lateroapically produced in a distinct hump; 

arista projecting almost at right angle to axis of second segment 

Conocaliscelis Mats. 

Body-form hemispherical. Tegmina strongly curved, claval suture absent 

Not as above (15) 

Wings well developed (11) 

Wings rudimentary Hemisphaerins Schaum 

Frons narrow, median carina present Darumara Merc. 

Frons broad, median carina absent (13) 

Anterior margin of vertex not carinate; lateral margins of frons straight, 
divergent distally as far as subangulate inflexure below level of an- 
tennae Gergithus St§,l 

Anterior margin of vertex carinate; lateral margins of frons roundly 
diverging to below level of antennae thence smoothly incurved, outline 
convex, not subangulate MongoUana Dist. 

Frons in profile strongly produced, anterior margin of vertex basad of 
level of anterior margin of eyes (17) 

Frons in profile straight or shallowly convex; anterior margin of vertex 
distad of anterior margin of eyes (21) 

Basal area of frons tricarinate. Protibiae and mesotibiae not foliately 
expanded Fortunia Dist. 

Basal area of frons with two carinae which unite basally. At least pro- 
tibiae subfoliately expanded (19) 

Mesotibiae foliately expanded Parahiracia Ouchi 

Mesotibiae not foliately expanded Clipeopsilus Jac. 

Vertex twice as broad as long in middle, or nearly so, frons at least as 
broad as long. Wings trilobed (23) 

Vertex not twice as broad as long, or if so then frons longer than broad. 
Wings with margin indented only once (31) 

Vertex more than three times as broad as long in middle; frons with 

lateral carinae parallel to below level of antennae. Wings absent 

Hysteropteruvi A. & S. 

Vertex not three times as broad as long (25) 

Frons with a carina completely across base parallel to anterior margin of 
vertex; median carina distinct as far as fronto-clypeal suture, sub- 
lateral carinae developed only at laterobasal angles Gelastyra Kirk. 

Frons without a complete transverse carina across base in addition to 
carina between frons and vertex (27) 

Tegmina broadest at level of claval apex. Wings with margin only shal- 
lowly indented, both postcubital and anal lobes narrow and small in 
relation to anterior lobe Kodaianella, new genus 


(28) (27) Tegmina broadest in basal half; wings with postcubital and anal lobes 

ample (29) 

(29) (30) Wings with posterior vein of anterior lobe and anterior vein of post- 

cubital lobe fused into a single thick stem lying in groove near apical 
margin Sarima Mel. 

(30) (29) Wings with these veins separate, linked only at apical margin by a very 

short oblique sclerotized strut Tetrica Stal 

(31) (32) Frons tricarinate ; vertex with anterior margin strongly convex. Teg- 

mina devoid of claval suture Neodurium, new genus 

(32) (31) Frons not carinate or only weakly medially so at base (33) 

(33) (34) Lateral margins of vertex and frons laminate. Tegmina with Sc + R 

united in a common stalk on basal quarter; wings with Sc + R forked 
at middle Tetricodes, new genus 

(34) (33) Lateral margins of vertex and frons acute but not foliate. Tegmina with 

Sc and R separate from base; wings with Sc + R not forked 

Duriopsilla, new genus 

The above synopsis has been compiled in part from literature, and obvious 
differences in published characters have been used for the separation of 
genera. 1'he result does not represent a full evaluation of the genera here 


listed. Parahiracia Ouchi and CUpeopsilus Jac. may well be congeneric and 
Ijoth may ultimately be suppressed under Fortunia Dist. The whole Hemi- 
sphaeriinae require critical study. The rather large number of monotypic 
genera of Issidae described in the Fauna of British India (Distant, 1906, 
1916), which is increased in the present report, probably reflects a paucity 
of specimens rather than incorrect delimitation of generic concepts, but 
a study of further material from eastern Asia is much to be desired. 

Genus Mongoliana Distant 
Distant, 1909:87. Haplotype, Ileinisphaerius chilocJwrides Wlk. 

Mongoliana chilochorides (Wlk.). 

Hemusphaerius chilochorides Walker, 1851:379. 

Eight males and 9 females, Okinawa, April, 1912, J. C. Thompson; 
1 female, Chizuka, Okinawa, July-Sept., 1945, Bohart and Harnage, 1945; 
1 male and 2 females, Mimasaka, Japan, July, 1912, J. C. Thompson. 

Mongoliana recurrens (Butl.). 

(Figure 17, G, H; Figure 18, A-C; Figure 19, B.) 
Heniisphaerius recurrens Butler, 1875:98, pi. 4, fig. 20. 

Frons longer than broad (1.1: 1), medially ecarinate. Tegmina almost 
smooth, with dull polish, wings distinctly shorter than tegmina. Post-tibiae 
2-spined at sides, 7-spined at apex, basal metatarsal segment with two stout 
spines and seven small intervening spines. 

Sepia brown; a transverse bar across clypeus at base, a similar bar 



across frons at apex, round mottling on frons and sides of head, eleven 
round spots along a shallow sulcus near each lateral margin of frons, an- 
terior and lateral carinae of vertex (in part), a series of small spots along 
anterior margin of pronotum and a small round spot on disc on each side of 
middle, lateral carinae of mesonotal disc and scutellum, ivory yellow. 
Clypeus, distal portions of lateral lobes of pronotum, pro- and mesocoxae, 
pro- and mesofemora, pro- and mesotibiae in basal two-thirds piceous; pro- 
and mesotrochanters, rostrum, hind legs and lower surface of abdomen 
testaceous. Tegmina uniformly very dark castaneous, a short transverse 
linear spot inward from sutural margin at level of apex of clavus. 

Anal segment short, broad, ex])anding distally, widest at truncate apical 
margin, anal foramen in distal half, lateroapical angles rounded. Pygofer 
with lateral margins convex. Aedeagus shallowly U-shaped, phallobase 
terminated dorsally in a pair of short tapering bluntly-pointed lobes, 
ventrally in a pair of short, broad, rounded lobes, laterally deeply incised, 
more so on left than right. Aedeagus with a pair of long curved blade-like 
spines arising ventrolaterally distad of middle, directed cephalad and 
slightly dorsad near apex, apex of aedeagus broadly ])ilobate, each lobe 
obliquely truncate, (lenital styles moderately narrow at base, expanding 
distally, apical margin more or less semicircularly curved, dorsal margin in 
middle produced dorsad in an oblique, transversely-compressed biscuspi- 
date process. 

Posterior margin of pregenital sternite broadly and shallowly subtriangu- 
larly produced at middle. 

One female, Lung-Tau Shan, N. Kwangtung, June 11, 1947, Gressitt; 
one male and 1 female, 1,000 m., Suisapa, Lichuan District, W. Hupeh, 
June 23, 1948, Gressitt. 

Fig. 17. Hemisphaerius ruf ovaritis Wlk.: A, head and thorax; B, frons and 
clypeus. Hemispfiaerius imitatus Mel.: C, head and thorax; D, frons and clypeus. 
Gergithus iguchii Mats.: E, head and thorax; P, frons and clypeus. MongoUana 
recurrens Butl.: G, head and thorax; H, frons and clypeus. 



[Proc. 4th See. 

Genus Gergithus Stal 

Stal, 1870:756. Type, Heniisphan-ius schaumi Stal, 1855:191 

Gergfithus rugulosus Melichar. 

Melichar, 1906:64. 

One male, Hong Kong, China, Koebele, Oct. 1895. 

Gergithus iguchii Mats. 

(Figure 17, E, F; Figure 19, A.) 
Gergithus iguchii Matsumura, 1916:98. 

Female: length, 3.5 mm.; tegmen, 4.8 mm. 

Frons slightly convex, smooth, ecarinate, polished. Post-tibiae 2-spined 
laterally, 6-spined at apex, basal hind tarsns with a spine at each angle, and 
7 spines between. Tegmina with veins prominent, sub-parallel, with a few 
cross-veins or anastomoses. 

Castaneons-fuscous; a band across base of clypeus, extending on genae 
np to antennae, and pro- and mesocoxae at base, pallid yellow; clypeus, 
femora at base and tibiae laterally (except on hind legs), piceous; meso- 
notum golden yellow. Tegmina golden yellow, costal margin broadly at 
base, narrowly distally, apical margin broadly, three large contiguous spots 
lying across basal third, two larger contiguous spots lying across apical 
third eastaneous-fuscous. Wings pallid grey with darker veins. 

Twelve females, Mokansan, Che Kiang Province, China. It is possible 

Fig. 18. Mongoliana rectirrens Butl.: A, anal segment of male; B, aedeagus, 
left side; C, pygofer, left side. Hemis])haerius signifer Wlk.: D, anal segment of 
male; E, aedeagus, right side; F, left genital style. 



that the Che Kiang population is subspecifically distinct, as the darker 
markings are rehitively hirger, but the point cannot be settled with material 
of one sex only. The anterior carina of the vertex is obsolete, and this 
species could perhaps be referred to H emisphaeroides Mel., but the latter 
genus requires re-defining before further species can be assigned to it with 

Genus Hemisphaerius Schaum 
Schaum, 1850:71. Haplotype, Issus coccinelloides Burm., 1833:305 
Hemisphaerius rufovarius Wlk. 

(Figure 17, A, B.) 
Walker, 1858:95. 

The present material differs from Walker's type only in the presence of 
two obscure small pieeous spots in the apical third of the tegmina. 

One male, Dwa Bi, Hainan Island, July 25, 1935, Cxressitt, and 1 muti- 
lated specimen, Ta Han, Hainan Island, June 24, 1935, Gressitt. 

Hemisphaerius imitatus Melichar. 

(Figure 17, C, D). 
Melichar, 1906:88. 

Female : length, 4.4 mm. ; tegmen, 4.2 mm. 

Greenish translucent. Clypeus, except for a yellow transverse band at 
base, a spot on each lateral lobe of pronotum at ventral margin, pro- and 
mesocoxae, two transverse bands on pro- and mesotibiae, post-femora except 
at apex, and both surfaces of abdomen pieeous, with hind margin of each 
segment bordered yellow. 

One female, 2,000 ft., Kepakiang, Sumatra (H. C. Kellers), presented by 
W. M. Giffard. 

Hemisphaerius signifer AValker. 

(Figure 20, A-C; Figure 18, D-F.) 
Walker, 1851:380. 

One male, 1,000 m., Suisapa, Lichuan District, W. Ilupeh, China, Aug. 
19, 1948, Gressitt. 

Fig. 19. Gergithus iguchii Mats.: A, tegmen. MongoJianu reLurrens (Butl.) 
B, tegmen. 


Kodaianella, Feniiah, new genus 

Frons broader than long in middle line (1.6: 1), lateral margins diverg- 
ing straight to below level of antennae, thence abruptly incurved through 
80° to suture, basal margin slightly angulately concave, disc slightly con- 
vex, almost flat, median carina distinct, vertex twice as broad as long in 
middle line, anterior margin obtusely angulate, posterior margin sub- 
parallel to anterior, lateral margins slightly diverging distally, disc slightly 
concave, median carina fee])le. Eostrum scarcely surpassing mesotrochan- 
ters, apical segment scarcely shorter than subapical, apex very obliquely 
truncate. Pronotum in middle line about 1.2 times length of vertex in same 
line, a small impression on each side of middle, mesonotum in middle line 
slightly longer than pronotum, disc slightly tumid, anteriorly bounded by a 
fine transverse ridge parallel with hind margin of pronotum. Post-tibiae 
laterally 2-spined. Tegmina broadening distally, costal margin convex, 
apical margin convex-truncate, Sc + R forked near base, M forked slightly 
basad and Cu forked markedly distad of union of claval veins. Wings almost 
as long as tegmina, anal lobe reduced, apical margin slightly indented 
twice in Cu. 

Anal segment of female relatively narrow, third valvulae broadly tri- 
angular, apical margin membranous, oblique. 

Type species, Kodaianella hicinctifrons, new species. 

Kodaianella bicinctifrons Fennah, new species. 
(Figure 21, A-D; Figure 22, A-C.) 

Male: length, 4.0 mm.; tegmen, 4.0 mm. Female: length, 3.9 mm.; teg- 
men, 4.2 mm. 

Frons distinctly medially carinate, submarginal carinae feeble. Post- 
tibiae 11-spined at apex, basitarsus 12-spined distally. 

Testaceous; basal half of frons and a more or less distinct band across 
middle of distal half, vertex, disc of pronotum, a band across lateral lobes 
of pronotum, and intercarinal areas of mesonotum, reddish-brown; sides of 
head below eyes and ventral portion of lateral lobes of pronotum, pallid 
yellow, hind legs and anteromedial portion of abdominal ventrites infus- 
cate. Tegmina translucent testaceous lightly sprinkled reddish-brown as 

Fig. 20. Heynisphaerius siynifer Wlk. : A, frons and clypeus; B, head and 
thorax; C, tegmen. 



figured, with a narrow irregular fascia traversing middle. Wings fuscous, 
veins darker. 

Anal segment long, relatively narrow, slightly expanding distally, apical 
margin semicirculai'ly excavate, lateroapical angles more or less acutely 
produced, anal foramen in basal half. Pygofer with laterodorsal angles 
slightly prominent, rounded, lateral margin shallowly sinuate. Genital 
styles moderately short, about as wide near apex as at base, apical margin 
more or less abruptly transversely truncate, dorsal margin at middle 
produced dorsad in a large tapering lobe with anterior margin excavate 
in its dorsal third, apex acuminate, a short stout peg-like process laterally 
below apex. Aedeagus shallowly curved dorsad. Phallobase terminating 
dorsally in a median shagreen finger-like lobe directed cephalad with a tri- 
angular sclerotised spine at each angle; and ventrally in a pair of acutely 
rounded lobes. Phallus with a pair of long stout spines arising in distal 
half, directed ventrocephalad below aedeagus. 

Posterior margin of pregenital sternite of female broadly produced 
caudad, margin of medial lobe truncate. 

One male (the type), 800-1,000 ft., Chang'-Tau-Ching-, Szechwan, July 
18, 1948, and 1 female, Sang-Hou-Ken, Ilupeh-Szechwan Border, China, 
July 19, 1948, both taken by Gressitt. 

This genus runs to Samantiga in Distant 's key (1906: 351), but differs 
in the shape of the head, tegmina, and venation; it is superficially similar 
to Kodaiana Dist., but differs in the number of post-tibial spines and in the 
shape of the ovipositor; from Sarima it differs in the shape of the vertex, 
frons, and tegmina and in tegminal venation; and from Narayana in the 
structure of the head. 

Duriopsilla Fennah, new genus 

Frons longer in middle line than broad (1.2: 1), lateral margins sinu- 
ately diverging to below level of antennae, thence moderately incurved to 

Fig. 21. Kodaianella bicinctifrons, new genus and species: A, head and thorax; 
B, frons and clypeus; C, tegmen; D, wing. 


suture, disc markedly convex, carinae and lateral pustules feeble, merely 
indicated; vertex shallowly impressed, lateral and apical margins forming 
a semi-circle, posterior margin sliallowly emarginate, slightly notched at 
middle, median carina distinct. Pronotum in middle line slightly longer 
than vertex, median carina feeble, an impression on each side of middle; 
mesonotum not quite as long as pronotum and vertex together. Rostrum 
reaching postcoxae, apical segment shorter than subapical, about three 
times as long as wide, apex transversely rounded-truncate. Pronotum with 
lateral lobes smooth; mesonotum twice as broad as long. Legs slender, 
femora not compressed; post-tibiae laterally 3-spined. Tegmina relatively 
narrow, costal and sutural margins straight, more or less parallel to level 
of node, symmetrically narrowing to acutely-rounded apex; Sc and E 
separate from base, M simple, Cui forked slightly basad of middle of claval 
suture, claval suture distinct. Wings deeply incised in Cu. 

Anal segment of female relatively short and broad, tapering distad. 
Third valvulae subequilaterally triangular, apical margin submembranous, 
tumid. Posterior margin of pregenital sternite transverse. 

Type species, Duriopsilla retarius, new species. 

Duriopsilla retarius Fennah, new species. 
(Figure 25, A-D.) 

Vertex broader than long (1.2: 1). Post-tibiae 8-spined at apex; basal 
metatarsal segment 9-spined at apex. 

Ochraceous; frons yellow; clypeus yellowish-brown, rostrum, hind mar- 
gin of pronotal disc, lateral edge of lateral pronotal lobes, mesonotal disc 
medially, legs except at joints, hind basitarsi, sometimes median area of 
abdominal ventrites, fuscous; pleurites immediately below base of costal 
margin of tegmina fuscous-piceous. 

Tegmina fuscous, heavily and uniformly covered with minute pallid 
reticulum of veinlets; veins dull greenish yellow. 

Fig. 22. Kodaianella bicinctifrons, new genus and species: A, anal segment of 
male, left side; B, aedeagus, left side; C, left genital style. 


Anal segment of male triangiilar, })roader than long, lateral angles de- 

Base of first valvulae of ovipositor as fignred. 

One male (the type) and 1 female, 1,000 m., Suisapa, Lichuan District, 
W. Hupeli, China, July 25, 1948, Gressitt. This genus is similar to Duri- 
opsis, but differs in the absence of a median carina on the clypeus, and of 
lateral mesonotal carinae; in tlie 3-spined post-tibiae, and the complete 
separation of Sc and R in the tegmina. 

Genus Gelastyra Kirkaldy 
Kirkaldy, 1904:280. Haplotype, Issiis testudinarius Stal, 1854:246 

Gelastyra biplaga (Wlk.), new combination. 

(Figure 23, E, H.) 

Issus hiplaga Walker, 1851:367. 

Female: length, 5.4 mm.; tegmen, 5.0 mm. 

Vertex broader than long (1.8:1); post-tibiae 7-spined at apex, basal 
metatarsal segment 11-spined at apex. 

Testaceous, tinged green; frons except for a round spot slightly basad of 
middle minutely and heavily speckled fuscous; anterior half and middle of 
pronotum, mesonotal disc laterally, a series of six oblique stripes on each 
side of middle line of clypeus, brown; transverse sulcus at base of frons 
piceous. Tegmina pallid, sub-translucent, basal quarter, except veins, and 
a broad band from middle of costa to sutural margin distad of apex of 
clavus, except veiiLs, chocolate brown; veins green, pallid portion of tegmina 
tinged with green except in a broadly ovate area one-third from base. 

Pregenital sternite produced posteriorly in a stout subspatulate process 
slightly broader than long. Third valvulae stout, broadly triangular, apical 
margin membranous. Anal segment narrowly ovate and laterally decurved 
distad of anal foramen. 

One female, Hong Kong, Oct., 1895, Koebele. This species, which is not 
included in Melichar's monograph, runs to the sjJectans Wlk.-latifrons 
Mel. section of Melichar's key (1906:263) but differs from both in the 
shape of the vertex and of the carinae in the basal part of the frons. 

Neodurium Fennah, new genus 

Frons in middle line longer than broad (about 1.2: 1), lateral margins 
almost straight, diverging to below level of antennae thence incurved to 
suture; disc shallowly convex, strongly depressed in middle near fronto- 
clypeal suture, leaving lateroapical areas prominent; median carina strongly 
developed on basal three-quarters, more feeble in apical quarter; a pair 
of weak but distinct sublateral carinae enclosing an elongate-oval area of 
disc, strongly incurved basally to meet transversely at middle line, vertex 



[Phoc. 4tii Ser. 

siibtiu'binate, apical margin siiljrectangiilately convex, posterior margin 
obtusely angulately excavate, lateral margins straight, slightly converging 
apically; disc hollowed ont, finely carinate throughout in middle line; sides 
of head shallowly grooved between lower margin of eye and frontoclypeal 
suture, clypeus ecarinate, convex, latero-basally slightly overhung by latero- 
apical area of frons; rostrum slightly surpassing mesotrochanters, apical 
segment shorter than subapical, very obliquely truncate at tip. Pronotum 
in middle line about as long as vertex, medially carinate with a small depres- 
sion on each side of mid-line, lateral lobes with four short parallel ridges 
near posterior margin; at least twice as broad as long, mesonotum shorter 
than combined lengths of pronotum and vertex, finely medially carinate, 
even where medially depressed, lateral carinae short, strongly anteriorly 
convergent, separated from median carina posteriorly by a ridge or con- 
vexity of disc. 

Pro- and mesofemora compressed, post -tibiae laterally with a spine near 
base and two large spines distally, apically with 8 spines, basal metatarsal 

Fig. 23. Tetricodes polyphemus, new species: A, tegmen; B, wing; C, head and 
thorax, dorsal view; D, frons and clypeus. Gelastyra hiplaga (Wlk.): E, wing; 
F, tegmen; G, head and thorax, dorsal view; H, frons and clypeus. 


segment witli about 13 spines. Tegmina with costal and sutural margins 
parallel to level of node, apical margin deeply and asymmetrically rounded, 
Sc and R arising separately from base, M simple, Cu forked basad of union 
of claval veins, claval suture absent, united el aval vein continued to sutural 
angle then curving into feeble submarginal vein. Wings large, deeply in- 
cised on apical margin into two lobes, anal lobe not present, veins simple, 
distal venation coarsely reticulate. 

Pregenital sternite of female with posterior margin shallowly excavate 
medially. Ovipositor with third valvulae stout, strongly convex, their 
apical margins tumid, polished, and bounding a lenticular cavity when 

Type species, Neodurium postfasciatnm, new species. 

Neodurium postfasciatum Fennah, new species 

(Figure 24, E-I.) 

Female: length, 5.0 mm.; tegmen, 5.1 mm. 

Vertex broader than long in middle line (1.8: 1). Profemora foliately 
expanded in apical half, mesofemora compressed, protibiae with subfoliate 
margins, basal metatarsal segment with 13 spines. 

Testaceous-brown, finely sprinkled fuscous; a pair of ocellate spots at 
base of vertex, clypeus, and a broad band near apex of profemora, fuscous 
piceous. Tegmina tawny-gold, with darker suffusion and piceous spots and 
marbling on all areas except a broad band from innnediately distad of 
node to estimated position of apex of clavus, and between hind margin 
and united claval veins. Wings translucent-fuscous with darker veins. 
Abdominal ventrites suffused fuscous; third valvulae of ovipositor ochra- 
ceous dorsally, fuscous ventrally, with a broad piceous band adjoining 
polished dull yellow tumid margin. 

Anal segment of female short, lateral margins strongly convex, apex 
subacutely rounded. Third valvulae of ovipositor, when apposed, in ventral 
view about twice as broad as long. Base of first valvulae and posterior mar- 
gin of seventh sternite as figured. 

Three females (one the type), 1,000 m., Suisapa, Lichuan District, W. 
Hupeh, China, Aug. 19, 20, 1948, Gressitt. The genus Neodurium differs 
from Duriopsis Mel., which it resembles, in shape of frons, proportions 
of mesonotum, and tegminal venation, and from Flavina Stal in proportions 
of frons, shape of legs, and absence of a claval suture in the tegmina; it 
also differs from both in the number of post-tibial spines. 

Tetricodes Fennah, new genus 

Vertex broader than long (2:1), anterior margin obtusely angulately 
produced, lateral margins slightly convergent anteriorly, posterior margin 
subangulately emarginate, disc sloping down to middle line, median carina 


absent; frons longer than broad (1.3: 1), basal margin shallowly excavate, 
lateral margins shallowly convex, disc with median carina present only 
basally, otherwise tumid in basal half, position of sublateral carinae marked 
by shallow groove, frontoclypeal suture slightly impressed, rostrum at- 
taining post-trochanters. Pronotum short, extremely narrow behind eyes, 
in middle line slightly longer than vertex; mesonotum slightly longer than 
pronotum. Post-tibiae armed with two stout spines in distal half, a minute 
tooth sometimes at extreme base, eight teeth at apex, basal metatarsal joint 
as long as other two combined. Tegmina with costal and sutural margins 
parallel, the former gradually rounding into oblique apical margin which 
is acutely bent at M4, Sc + R fork one-quarter from base, each limb simple 
to apex, M three-branched, Cui simple, union of claval veins level with 
fork of M. Wings larger than tegmina, with broadly reticulate venation, 
margin deeply cleft in Cu, anal lobe absent. 

Anal segment of female moderately elongate, parallel sided and distally 
rounded. Ovipositor with third valvulae stout, triangular, with thick, tumid, 
pellucid apical margin. 

Type species, Tetricodes polyphemus, new species. 

Tetricodes polyphemus Fennah, new species. 

(Figure 23, A-D.) 

Female: length, 5.2 mm.; tegmen, 5.5 mm. 

Tegmina with apex of R curved toward M, M3 distilly uniting with M1+2. 

Ochraceous to pallid with scattered greenish suffusion; discs of vertex 
and pronotum, except in middle line, and disc of mesonotum orange-brown 
and sepia; a polished tumescence on frons and a suffusion over distal quar- 
ter, a narrow triangle before eyes, and a spot on mesopleura, piceous; most 
of elypeus, lateral lobes of pronotum near margin, except for a few greenish 
pustules, a pair of transverse bands on femora, a suffusion on pro- and meso- 
tibiae distally, abdomen dorsally and ventrites 4 to 7, fuscous. Tegmina 
dark sepia marbled with transverse veins and parts of longitudinal veins 
emerald and pallid green. Wings fuscous. 

One female, 1,000 m., Suisapa, Lichuan District, W. Hupeh, China, Aug. 
21, 1948, Gressitt. 

In Melichar's key to Thioniinae (1903:254) this species runs to "7," 
but it differs from both alternatives; from Flavina Stal in the form of the 
head and in the number of post-tibial spines, and from Cameruniellu Hagl. 
in tegminal and wing venation. If Flavina ^ striata Dist. belongs in this 
genus, it is separated from T. polyphemus by coloration, especially of the 

Genus Tetrica Stal 
Stal, 1866:208. Logotype, Tetrica fusca Stal, 1870:757 

It appears likely that this genus, as currently recognized, is composite 



or contains groups of species which might he recognized as distinct snh- 
genera: it is also possil)le that species have been assigned to Sarima Mel. 
by their authors because they did not agree with their interpretation of 
Tetrica. Melichar's concept of Sarima is restricted to species in which the 
two veins which adjoin the first fold of the wings are fused together dis- 
tally to form a single stout rod. On account of this restriction the following 
two species fall into the looser concept of Tetrica Stal. 

Tetrica zephyrus Feiniah, new species. 
(Figure 24, A-D.) 

Female: length, 5.5 mm.; tegmen, 6.0 mm. 

Lateral margins of frons evenly incurved distally through less than 90°. 
Rostrum with apical segment markedly expanding distally in anterior 
view, at apex twice as wide as at base of subapical segment. Post-tibiae 
laterally bispinose, apically 7-spined: basal metatarsal segment with 9 
spines. Tegmina with Sc reaching to middle of costal margin, of subequal 
prominence throughout. 

Testaceous; a suffusion anteriorly on vertex, over frons except in basal 
fifth and clypeus except at sides, genae before eyes and abdominal ventrites 
fuscous, a narrow band across base of frons, excluding transverse carinae, 

Fig. 24. Teti'ica zephyrus, new species: A, tegmen; B, wing ; C, head and thorax; 
D, frons and clypeus. Neodurium postfasciatum, new species: E, tegmen; F, wing; 
G, head and thorax ; H, frons and clypeus ; I, apex of vertex, anterodorsal view. 


a spot antei'olaterally on ])i'onotiiiii underlying ])asal surface of head, two 
bands near each side of mesonotum, and tibiae at apex, fuscous-piceous. 

Tegmina greyish-translucent, a diffuse fascia from base of clavus across 
humeral eminence to costal margin, an irregular and interrupted V-shaped 
suffusion from apex of clavus to fork of Cu, thence obliquely to costal mar- 
gin near node, brown; intervenal submarginal ai'eas of costa and apex dark 
fuscous. Wings fuscous. 

Deflexed part of anal segment of female about 5 times as long as broad. 
Ovipositor with third valvulae triangular, castancous, polished, narrowly 
membranous at tip, which is subacute. 

Two females (one the type), Mokansan, Che-Kiang- Province, China, 
Aug. 24, 1927, Mrs. D. E. Wright. This species differs from T. aequa Jac. 
in coloration and from S. himaculata Mel. and S. clathrata Mel. in the po- 
sition of the union of the frontal carinae, and in the color and pattern of 
the tegminal markings, from S. mnagisana IVIel. in coloring of frontal 
carinae and tegmina, and from S. sinensis (Wlk.) in the much less angulate 
anterior margin of the vertex. 

It is just possible that it may prove to be a geographical subspecies of 
Sarima nigvifacies Jac, though it differs substantially from the described 
coloration, wliile it cannot be assumed that the species are congeneric. The 
decision to erect a new species, based only on female material, was taken 
by the writer after he had satisfied himself that the nuances of shape of 
the head and its carinae, and of the tegmina and their venation furnish ade- 
quate means of s]:)ecific recognition. 

Genus Caliscelis Laporte 
De Laporte, 1833:251. Haplotype, Fulgora honelli Latr., 1807:166 

Caliscelis chinensis Mel. 

Melichar, 1906:16. 

Post-tibiae laterally with 1 spine; apically wuth 7 short stout spines. 
Basal metatarsal segment 2-spined. 

Ovipositor with third valvulae thickened and slightly tumid on hind mar- 
gin near base. 

One female, Tunglu, Che Kiang Province, China, Sept. 10, 1926, Mrs. 
D. E. Wright. 

Fig. 25. Durioiis