ya) Gare) ~< Ye is rete: ‘s vi i) fap : aoe ma 7 iL? A PROCEEDINGS OF THE California Academy of Sciences FOURTH SERIES Vol. XXVIII SAN FRANCISCO PUBLISHED BY THE ACADEMY 1953-1956 COMMITTEE ON PUBLICATION Dr. Robert C. Miller, Chairman Dr. G. F. Papenfuss Dr. Edward lL. Kessel, Editor No. No. No. No. No. =“ 10. fi 12. CONTENTS OF VOLUME XXVIII YonGE, C. M. Observations on Hipponix antiquatus (Lin- naeus) (9 text figures). Published July 15, 1953.... Gressirt, J. LINSLEY. The California Academy-Lingnan Dawn-Redwood Expedition (135 text figures). Pub- Hislvedecilyelonne lO Do ien weewse ergs Olen s tses aa cysalfvei's oto 70 Essic, E. O. Some New and Noteworthy Aphidae from Western and Southern South America (Hemiptera- Homoptera) (56 text figures). Published July 15, TI QIBie e cttottierad ait fat iit Md a Reet ies Psat drut arena ng Gennes pe Orr, Rosert T. Natural History of the Pallid Bat, An- trozous pallidus (LeConte) (28 text figures). Pub- kishedtvanuaryats PIots tae Pas ee ease Ss are EOS Sasa, Manabu, and E. W. JAMESON, JR. The Trombi- culid Mites of Japan (45 text figures). Published BU EDENDT MIEN ea an) oy ae Meet ya rea Ng ech Sr cceve sete ve loncis) se eahielsl oye, LOUKASHKIN, ANATOLE S., and NORMAN GRANT. Further Studies of the Behavior of the Pacific Sardine (Sar- dinops caerulea) in an Electrical Field (5 text fig- Mes a ublishted! ae 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. *5. C—C, extent of ctenidium (dotted where viewed through the mantle wall); WM. mouth; MG, mid-gut; OD. opening of oviduct; OF, oesophagus; PC, pericardium; R, rectum; SS, style-sac; ST, stomach. Other let- tering as before. 6 CALIFORNIA ACADEMY OF SCIENCES [Proc. 41H Ser. MANTLE CAVITY After removal from the shell, the animal may be viewed from the dor- sal aspeet, as shown in figure 3. Owing to the forward extension of the shell muscles and the consequent constriction of the opening of the mantle cavity, the anus (A) has been displaced dorsally, opening slightly to the right of the mid-line. The anterior end of the etenidium (C) is similarly carried round dorsally, terminating just to the left of the anus. Figure 4. Hipponix antiquatus, dorsal view of organs in the mantle cavity after opening along the right side. 5. K, 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. r2—wz, cut ends of mantle edge; y—y, cut surfaces, posterior wall of mantle; ¢—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 alone 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 Hipponir. In archaeogastropod limpets, such as Acmaea or Patella (Yonge, 1947), and also in the mesogastropod Capulus, which is closely related to Hipponir, the head oceupies the center of the mantle ‘avity. This is a consequence of secondary symmetry associated with loss of coiling in the shell and visceral mass. In all of these limpets height is seldom less than breadth, it is often greater. In Hipponir, on the other hand, height is always less than half the breadth. As a result of this and Vom. XXVIII] YONGE: OBSERVATIONS ON HIPPONIX ANTIQUATUS 7 of the reduction of the pedal tissues (with which this compression 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 shallow. Hence the head, although it projeets forward out of the middle of the opening of the mantle eavity (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 head (figs. 4 and 7B: KR, DR). Apart from this flattening and extension posteriorly, the mantle cavity is that of a typical pectinibranch prosobraneh, having the same general disposition of the pallial organs as, for instanee, in Buccinwm (Yonge, 1938). The pericardium, as revealed by dissection and in see- tions, is situated far to the left at the base of the etenidium (figs. 3 and 7C: 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-line. Internally it is unusually capacious, as shown in figure 5. It communicates with the pericardium by way of a long reno-pericardial eanal (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 ovidueal (figs. 3 and 4 OD). 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 ectenidium (C) occupies the > Figure 5. Hipponix antiquatus. longitudinal section. X11. Bb, 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; SS, 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 eurrent (fig. 3, 1) extends the linear osphradium (figs. 3; 4; 8 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H Ser. and 7A, O). Between the ctenidium and the rectum, the roof of the cavity is covered by the extensive tissues of the hypobranchial gland (figs. 4; 5; and. (; 1G). CILIARY CURRENTS The lateral cilia on the ctenidial filaments create an inhalant cur- rent (1) which enters the cavity on the left side, impinging first on the osphradium. The exhalant 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 Vermetus novae-hollandiae, Capulus ungaricus and, to a striking extent, Crepidula fornicata (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 cila which are here concerned with food collection. In Hipponir there is only a moderate inhalant current, adequate for the lmited respiratory needs of the animal, while the frontal cilia retain their primitive funetion of cleansing. In ciliary feeding species, moreover, collected particles are conveyed to the mouth along the tips of the elongated filaments, and by way of special food grooves, to the mouth. This has been deseribed in various of the Vermetidae (Yonge, 1932, 1938; Morton 1951b), in Turritella com- munis (Graham, 1938), in the Struthiolaridae (Morton, 1951a)—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 the mantle cavity is essentially as in typical pectinibranchs. As de- scribed elsewhere (Yonge, 1938), there are three currents concerned with rejection of sediment, (4) those carrying heavier particles to the inhalant opening; (B) those carrying medium particles across the floor of the mantle cavity; (C) those earrying the finest particles over and between the etenidial filaments for later consolidation dorsally in the mucus from the hypobranechial 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). Vou. XXVIII] YONGE: OBSERVATIONS ON HIPPONIX ANTIQUATUS 9 In Hipponix the only modification is due to the constriction of the opening of the mantle cavity which has had the effeet of carrying the greater part of the etenidium 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 A on the floor of the inhalant aper- ture (fig. 2). The current then impinges on the osphradium (O) and passes between the filaments. Larger particles are carried to the tip of the filaments by the frontal cilia and are rejected by cilia on the surface of the head, on the sides of the tentacles and on the floor of the mantle cavity to the right, that is current 6 (fig. 4). The finest particles are carried between the filaments and are then consolidated in the mucus from the hypo- branehial gland. Cilla of current C then earry 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 1s foreed out when the shell muscles contract. It was 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, 1958; Yonge and Iles, 1939; Morton, 19516) while Capulus 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 etenidia are carried in a ciliary tract to the upper surface of the propo- dium (much larger than that of Hipponir) where the proboscis collects the mucus-laden masses by means of the radula. Orton (1949) has pointed out that Capulus may also live on the shells of lamellibranchs, such as Modiolus and Monia, and probably takes some of the food of these ani- mals by inserting the proboscis into the mantle cavity. But in Hipponie there is no doubt that etenidium and ciliary rejection currents are in no way modified for feeding. The presenee 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. 10 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SrEr. ALIMENTARY CANAL The mouth consists of a vertical slit at the end of the proboscis and at the base of the two lateral flaps in which this terminates. There is a small, and certainly funetional, radula and small salivary glands. The bueeal cavity leads into what is termed by Graham (1939) the anterior oesophagus and which possesses in Hipponir 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 glandular 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. X10. D, opening of duct into digestive diverticula (ventral) ; MG, opening into mid-gut (dorsal) separated by typhlosole (TY) from style-sac (SS); OF, opening of oesophagus (ventral). gq, g, 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-eut (MG) which is separated by a conspicuous typhlosole (TY) from an associated style-sac (SS), leaves on the same side but dorsally (fig. 5). 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 gastrie 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 caecum which VoL. XXVIII] YONGE: OBSERVATIONS ON HIPPONIX ANTIQUATUS il 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 around 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 mm. these ter- minal regions of the gut pulled out to a length of some 22 mm. Foop AND FEEDING The only previous account of feeding in Hipponix appears to be that of Risbee (1935) on H. australis. This species lives on the shells of other gastropods, usually of species of Turbo, 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. Hipponix antiquatus feeds in essentially the same manner but on fragments of material—organie detritus, pieces of algae, ete——that are earried within the very limited area in front of the shell where alone the proboscis can browse. 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 earried on by a sedentary animal if food supplies are constantly being renewed by water movements. The proboscis itself is muscular and very active. The terminal lobes, with the mouth which they flank, were frequently seen to open widely and grope forward in apparent search for food. The odontophore was then seen to protrude from the mouth opening. Somewhat similar ob- servations were made on C. ungaricus after mucus-laden food masses had been carried on to the surface of the propodium (Yonge, 1938). In this species, however, the proboscis, which is grooved anteriorly, represents 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 caleareous coral- line algae up to 2 mm. long and 0.5 mm. wide. The radula must convey 12 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4tTH Ser. these fragments into the bueeal mass and anterior oesophagus the power- ful ciliation of which, possibly aided by some muscular action, serving to carry them into the stomach. Within the stomach material would appear initially to pass to the right, into the caecal extension, as indicated in figure 6. This serves as ¢ store and perhaps to some extent also as a gizzard although the triturition which must occur before the larger particles can pass into the mid-gut is probably 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 intact 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 » long and about 500 » 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 opening, normally small, is greatly distended at defecation. The exhalant current must aid in ear- rying the pellets 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) mixine of food with the style substance, (2) sorting of fine particles for passage into the digestive diverticula, (3) removal of larger particles and waste from the digestive diverticula into the mid-gut, (4) rotation and passage imto 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] YONGE: OBSERVATIONS ON HIPPONIX ANTIQUATUS 13 REPRODUCTIVE ORGANS Every specimen examined in life or disseeted was female. One small specimen of which longitudinal sections were made showed no sien 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 Hipponiz.* In C. ungaricus the reproductive sys- tem is simple, consisting, in the male phase, of a testis with associated am- pullae and a duct leading into a pouch with which a receptaculum com- munieates by a 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 tip 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 now becomes what Giese deseribes as a uterus into which the receptaculum continues to open. The sperm groove and the penis disap- pear but the female reproductive aperture, although longer, remains in the original position. Presumably the males, being vounger and probably more active, are able to move on to and copulate with the larger and probably completely immobile females. Jones (1949) deseribes the presence of small speci- mens of C. ungaricus on the shells of larger ones, and these may well have been males and females respectively. After copulation, sperms would presumably be stored in the receptaculum and later fertilize the egos when they entered the “uterus.” The latter is thick-walled and glandular and responsible for the formation of the protective capsule dis- eussed in the next section. The small specimen of Hf. antiquatus without reproductive organs was some 6 mm. in diameter and was attached to rock. If this animal was indeed immature and if all members of this species are protandric her- maphrodites then they cannot be mobile in a subsequent male phase. Cer- tainly no small mobile individuals were observed and in the very turbu- lent conditions where H. antiquatus lives, permanent attachment in early life would seem to be almost essential. This matter cannot be regarded as settled but all available evidence, admittedly very limited, indicates that the species, almost certainly a protandrie hermaphrodite, is not mo- bile in the male phase and so copulation cannot occur. Tf this be so, then fertilization may occur in one of two ways. The first possibility is that the spermatazoa are liberated freely into the sea and that fertilization occurs after these have been carried into the mantle eavity and have entered the genital aperture of a female. Such ‘‘eurrent fertilization’’ is known to oeceur in Turritella communis *The parasitic Thyca stellasteris, according to Koehler and Vaney (1912), is not hermaphrodite al- though the male is smaller than the female. 14 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47m Serr. (Fretter, 1946) and in various members of the Vermetidae (Morton, 1951b). All of these animals are, however, ciliary feeders (Graham, 1938, Morton, 1951) and so with enlarged ctenidia which create a powerful in- halant current. In addition the pallial genital duct is widely open ventrally to permit the reception of sperm carried to it in the water. All of these animals, moreover, probably live in numbers together and in compara- tively still water, ie. under conditions where a local concentration of sperm adequate to ensure fertilization could be produced. 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 Turritella 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 animal when it passes into the female phase. This would not be unprecedented in the Mollusea, having been shown to occur in the wood-boring lamelli- branch, Vylophaga dorsalis (Purchon, 1941). This animal alternates in sex but the sperms produeed in a male phase are stored in a receptaculum for use in the subsequent female phase. Yylophaga dorsalis lives largely isolated in drift wood and this is probably the only feasible method by which fertilization ean be assured. Although the habitat is different, the problem in the ease of HW. antiquatus is not dissimilar and the solution may be the same. The female reproductive organs are very similar to those deseribed by Giese in Capulus. 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 duet (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 eges are large, containing great quantities of yolk, when they leave the ovary (fie. 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 eges will be passed into the mantle cavity. ATTACHMENT OF Hag CAPSULES Sedentary Mesogastropoda must either attach the ege capsules to the inside of the shell, as do the Vermetidae (Morton, 1951b), fasten them VoL. XXVIII] YONGE: OBSERVATIONS ON HIPPONIX ANTIQUATUS 4) =. 7 CS Sahih [oc FAO Figure 7. Hipponix antiquatus, middle (B), and posterior (C) sules; N, transverse section through the anterior (A), regions of the mantle cavity. x5. HC, egg cap- nerve collar at base of proboscis; OV, ovary containing large, yolk-filled eggs; RC, reno-pericardial canal, showing opening into pericardium (PC); RS, duct from oviduct (capsule gland) into receptaculum seminis; 7, tentacle; V, ventricle. Other lettering as before. 16 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H Serr. to the rock or pebbles on which the female rests, as in Calyptraeidae such as Crepidula fornicata (Lebour, 1937), or else retain them secured to the actual body of the female. This is the case in both Capulus and Hipponw. In C. ungaricus a single large egg capsule or cocoon is attached to the large propodium of the female. Full references to literature are given by Thorson (1946) who summarizes previous work in the statement that “‘Wach female protects a single thin-walled cocoon of the form of a sausage, and each cocoon contains several eggs, 200 » across, which hatch as veligers through a fissure on the under side of the cocoon.’’ Although Risbee (1935) does not mention males and gives a very imperfect account of the female genital system in H. australis, he gives a good description 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. antiquatus. 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 saes received the eges and also vellow nutritive material of a fatty nature. He was unable to observe the role of the foot in the formation and attachment of these capsules. Conditions are essentially similar in H. antiquatus. After the comple- tion of egg-laying the mantle cavity of the female is completely filled by from six to eight ege capsules each containing up to 50 yellowish eggs. The eggs measured about 350 » in diameter, the greatest transverse diameter ot the capsules being 840 ». Each capsule was, as in H. australis, attached by an attenuated stalk to a perforated calcareous plate which in turn was attached within a membranous area situated in the depression yven- tral of the propodium (figs. 4, 70, and 8; PL). As shown in figure 8, the capsules 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 pushed 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 and attachment of the egg capsules. Owing to the sedentary habit of this animal, full details of this process could probably only be made out by examining great numbers of animals during the period of egg-laying in hope of obtaining animals in all stages. The first ege-carrying female was obtained on April 5; during the summer all animals had egg eapsules although some of these were empty. It is, however, certain that the eges are large and yolky when they leave the ovary, where they were observed in sections (fig. 7€, OV). After Vou. XXVIII] YONGE: OBSERVATIONS ON HIPPONIX ANTIQUATUS 17 Figure 8. Hipponix antiquatus, ventral aspect with egg capsules in the mantle cavity. 5. Lettering as before. fertilization they presumably each receive a protective covering while in the capsule 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 fornicata, a stalked capsule is secreted around the eges by such a eland in the propodium. The calcareous plate and its investing membrane may be formed within the pocket ventral to the propodium ; 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 Risbee. The embryos develop into fully shelled larvae but whether these actually crawl away after emergence from the capsules, as described by Risbee for HH. australis, was not seen. In view of the very specialized habitat, it is more than probable that, in H. antiquatus also, the young crawl away from the parent to settle in due course permanently on the adjacent rock surface. DISCUSSION 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 18 CALIFORNIA ACADEMY OF SCIENCES [Proc. 41m Serr. normally oecur. It is therefore not surprising that the limpet form, with its secondary symmetry (see Yonge, 1947, p. 490, fig. 31) has independ- ently been evolved on a number of occasions. The particular conditions that exist in Hipponix are most suitably discussed by comparing them with those found in examples of the other chief types of marine limpets. In figure 9 comparison is made between four types of limpets, (A) Diodora (Fissurella) and (B) Acmaea are both archaeogastropod limpets the former having paired, i.e. zygobranchous, aspidobranchiate ctenidia, the latter having a single aspidobranch ectenidium. (C) Hipponir is a mesogastropod limpet with a pectinibranch etenidium and (D) Siphonaria is a pulmonate with a secondary gill in what has been a pulmonary mantle eavity. A eap-like shell and a horseshoe-shaped shell muscle (with the Cc D Figure 9. Diagrammatic comparison, from dorsal aspect, of four types of limpets, showing mantle cavity (WC) with inhalant (7), exhalant (#) 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, Hipponix (Mesogastropoda) with pectinibranch ctenidium, exhalant current an- terior; D, Siphonaria (Pulmonata) secondary gill, exhalant current on right. VoL. XXVIII] YONGE: OBSERVATIONS ON HIPPONIX ANTIQUATUS 19 opening on the right in Siphonaria) are common to all. The course of the respiratory and cleansing currents may be briefly summarized. The anterior mantle cavity of the prosobranehs (A-—C) has a generalized an- terior inhalant current (I) in A but, with the loss of the right etenidium, this is confined to the left in B and C. In A the exhalant current (I) 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 colleeted in the mantle (nuchal) cavity and in the pallial grooves. From such a condition have been derived those in the more specialized Lottia, 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 typical pectinibranch. 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 posterior to the inhalant opening. It is in the matter of feeding that Hipponir, and the mesogastropod limpets in general, differ from these archaeogastropod and pulmonate lim- pets. The latter all browse on encrusting vegetation, moving very slowly over the rock and seraping this with the broad radula. The mesogastropod limpets may be divided into (1) those in which the etenidial filaments have been greatly elongated and which feed exclusively by cillary cur- rents, Le. Crepidula and Calyptraea (Calyptraeidae) and (2) those in which there is a pronounced proboscis. 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 Capulus, which is the least specialized, the grooved proboscis, formed by prolongation of the terminal lobes in Hipponir, may be used to take in food collected by the enlarged etenidia (Yonge, 1938) but it may probably be also used to take in material similarly collected by a lamellibranch (Orton, 1949). In both Hipponir australis and H. anti- quatus the etenidium 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. ungaricus, but H. antiquatus is not dependent on any other animal. Although cemented to the substratum, yet it uses the proboscis to collect 20 CALIFORNIA ACADEMY OF SCIENCES [| Proc. 4TH SER. food in front of the shell. Species of a third genus, Thyca, have also lost the power of movement but in association with a completely parasitic life. They are ectoparasitie on echinoderms, the foot being reduced but the long proboscis penetrating deep into the tissues of the host (Sehepman and Nierstrasz, 1909; Koehler and Vaney, 1912). The dise of attachment is formed by the proboscis, the columellar 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 flat- 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 seeretion 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 H. anti- quatus lives, namely within crevices among rocks exposed to the full and almost unvarying foree of the Pacifie surf. Under such conditions food will constantly be renewed so that exclusive dependence on the probescis is possible. This is not the case in C. ungaricus which lives, sublittorally, in very much quieter water. Again, members of both of these groups of limpets are protandrous 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 Hipponia 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 precise manner in which the egg capsules are attached to the underside of the propodium. There is finally the question of classification. Thiele (1931) places Amalthea (—Hippomx) in the Amaltheacea but Capulus and Thyca 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 Capulus 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 serutiny. VoL. XXVIII] YONGE: OBSERVATIONS ON HIPPONIX ANTIQUATUS 21 SUMMARY Hipponix antiquatus is a mesogastropod limpet of particular interest owing to its sedentary habits. It lives cemented to the substratum in crevices among rocks often fully exposed to the Paecifie surt. 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, seereted by the mantle. The margins of the two make perfect contact when the horseshoe-shaped shell muscle contracts. The opening of the mantle cavity is constricted causing displacement dorsally of anus and etenidium; the shell is also much flattened with consequent 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 with respiration. Feeding is by means of a muscular proboscis which is extruded from the mantle cavity and swallows relatively very large masses of organie 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 probable that spermatozoa produced during the male phase are stored in the receptaculum for fertilization of eges produced in the subsequent female phase. Large ege capsules, each containing a number of large, yolky eggs, are attached to a caleareous plate embedded ventral to the propodium. The mantle cavity is largely filled by these capsules during spring and early summer. In comparison with archaeogastropod 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). With the exception of Thyca, all of these limpets are also pro- tandrie hermaphrodites. 22 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4rm Ser. “= REFERENCES Cook, P. M. 1949. A ciliary feeding mechanism in Viviparus viviparus (L). Proceedings of the Malacological Society of London, vol. 27, pp. 265-271. FRETTER, V. 1946. The genital ducts of Theodorus, Lamellaria and Trivia, and a discussion on their evolution in the prosobranchs. Journal of the Marine Bio- logical Association of the United Kingdom, vol. 26, pp. 312-351. GIESE, M. 1915. Der Genitalapparat von Calyptraea sinensis Lin., Crepidula unguiformis Lam. und Capulus ungaricus Lam. Zeitschrift fiir wissenschaftliche Zoologie, Bd. 114, pp. 169-231. GRAIIAM, A. 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 prosobranchs. Proceedings of the Zoological Society of London, ser. B, vol. 109, pp. 75-112. JONES, N. S. 1949. Biological note on Capulus ungaricus. Marine Biological Station at Port Erin, Isle of Man. Annual Report for 1948 (no. 61), p. 29. Keren, A. M. 1937. An abridged check list and bibliography of west North American marine Mollusea. Stanford University Press, California. DIES da, Gloel Af, 16), BAG, din. 1935. West coast shells. Stanford University Press, California. KOEHLER, R., and C. VANEY 1912. Nouvelles formes de Gastéropodes ectoparasites. Bulletin Scientifique de la France et de la Belgique, T. 46, pp. 191-217. LEcDouR, 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 Biological Association of the United Kingdom, vol. 22, pp. 105-166. Moron, J. E. 195la. The ecology and digestive system of the Struthiolariidae (Gastropoda). Quarterly Journal of Microscopic Science, vol. 92, pp. 1—25. 151b. The structure and adaptations of the New Zealand Vermetidae. Part 1: The genus Serpulorbis. Part 2: The genera Stephopoma and Pywi- poma, Part 3: Novastoa lamellosa and its affinities. Transactions of the Royal Society of New Zealand. vol. 79, pp. 1-51. VoL. XXVIII] YONGE: OBSERVATIONS ON HIPPONIX ANTIQUATUS 23 Orton, J. H. 1912. The mode of feeding of Crepidula, with an account of the current- producing mechanism in the mantle cavity, and some remarks on the mode of feeding in gastropods and lamellibranchs. Journal of the Marine Biological Association of the United Kingdom, vol. 9, pp. 444— 478. 1949. Note on the feeding habit of Capulus ungaricus. Marine Biological Sta- tion at Port Erin. Isle of Man. Annual Report for 1948 (no. 61), pp. 29-30. Purcnon, R. D. 1941. On the biology and relationships of the lamellibranch XYylophaga dorsalis (Turton). Journal of the Marine Biological Association of the United Kingdom, vol. 25, pp. 1-39. RIsBec, J. 1935. Biologie et ponte de mollusques gastéropodes Néo-Caledoniens. Bulletin de la Societe Zoologique de France, T. 60, pp. 387-417. ScHEPMAN, M. M., and H. F. NIERSTRASZ. 1909. Parasitische Prosobranchier der Siboga-Expedition. Siboga-Expeditie, no. XLIX.2 Smirn, A. G., and M. GorDON 1948. The marine mollusks and brachiopods of Monterey Bay, California, and vicinity. Proceedings of the California Academy of Sciences, vol. 26, pp. 147-245. THIELE, J. 1931. Handbuch der systematischen Weichtierkunde, Bd. 1, pp. 1-376. THORSON, G. 1946. Reproduction and larval development of Danish marine bottom inver- tebrates. Meddelelser fra Kommissionen for Danmarks Fiskeri—Og Havundersogelser. Serie: Plankton, Bind IV, pp. 1-523. WERNER, B. 1948. Uther den Laichvorgang der amerikanischen Pantoffelschnecke Crepidula fornicata L. Verhandlungen Deutsche Zoologische Kiel, 1948, pp. 262-270. YONGE, C. M. 1932. Notes on feeding and digestion in Pterocera and Vermetus, with a dis- cussion on the occurrence of the crystalline style in the Gastropoda. Scientific Report Great Barrier Reef Expedition (1928-29), British Museum (Natural History), vol. 1, pp. 259-281. 1937. The biology of Aporrhais pes-pelecani (L.) and A. serresiana (Mich.). Journal of the Marine Biological Association of the United Kingdom, vol. 21, pp. 687-704. 1938. Evolution of ciliary feeding in the Prosobranchia, with an account of feeding in Capulus ungaricus. Journal of the Marine Biological Asso- ciation of the United Kingdom, vol. 22, pp. 453-468. 24 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH Ser. 1947. The pallial organs in the aspidobranch Gastropoda and their evolution throughout the Mollusca. Philosophical Transactions of the Royal Society of London, ser. B., vol. 232, pp. 443-518. 1952. The mantle cavity in Siphonaria alternata Say. Proceedings of the Ma- lacological Society of London, voi. 29, pp. 190-199. YonceE, C. M., and KH. J. ILEs 1939. On the mantle cavity, pedal gland, and evolution of mucous feeding in the Vermetidae. Annals and Magazine of Natural History (ser. 11), vol. 3, pp. 536-556. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series Vol. XXVIII, No. 2 July 15, 1953 THE CALIFORNIA ACADEMY—LINGNAN DAWN- REDWOOD EXPEDITION BY J. LINSLEY GRESSITT Research Associate California Academy of Sciences The discovery in West Central China in 1945-46 (see Chaney, 1948) ot a living species of the ancient, fossil, coniferous genus Metasequoia, 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 Seiences and personnel and equipment by Lingnan University. The trees were known from a few loealities along the border of Sze- ehuan and Hupeh provinees, 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. We 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 glyptostroboides Tu 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 | 26 CALIFORNIA ACADEMY OF SCIENCES [| Proc. 47H Ser. Figure 1. One of the largest Metasequoia trees, located in lower Suisapa Valley near Hsiao-ho. Vou. XXVIII] GRESSITT: DAWN-REDWOOD EXPEDITION ) 4 ground). The trees are straight, and not irregular like the relative, Glypto- strobus pensilis, the ‘““water-pine’’ of the Canton delta, for which the species was named, The trunk is somewhat enlarged at the base and the bark is reddish but thin. On the smaller branehes the bark is somewhat folded or curled. The lower branches slant slightly downward, and the upper branches point distinctly upward. The wood is relatively light and soft. The folage is a conspicuous pale green. The needles are flat (fig. 2), on opposite sides of the stem, and closely resemble those of the ‘ Figure 2. Foliage of Metasequoia glyptostroboides. California coast redwood, Sequoia sempervirens. Metasequova differs 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 shwi-hsa in Chinese, whieh means ‘‘ water- fir’? or ‘‘water-Cunninghamia.’’ The tree was apparently unknown out- side its range before its recent scientific discovery by Chinese plant. col- lectors. Probably its range in recent historic times has not been mueh 28 CALIFORNIA ACADEMY OF SCIENCES | Proc. 47H SER. Figure 3. Cones of Metasequoia glyptostroboides (outer two) and Glyptostrobus pensilis (inner two). more extensive than at the present time. This is suggested by the faet that the wood is not considered valuable, and is not earried 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 popula- 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 Szechuan by the Imperial forces for failure to pay taxes may have been an important 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 existine trees have been transplanted to their present situations. For example, many ot the large trees are in straight rows up ravines, paralleline the small streams. Others are around the farmhouses. Many young and medium- sized trees are in straight rows alone the edees of rice fields bordering streams. The reason for the transplanting of the volunteer seedlings from the shaded ravines to particular situations, often in rows, is apparently based on local superstition. The mountain people have the habit of pre- dicting their crops on the basis of cone development on the trees. Thus a heavy crop of cones on the upper portions of the trees is said to 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, laequer, ete.). Thus it may actually be that the dawn redwood has been preserved from final extinction more or less by chanee. It is interesting to note that the water pine, Glyptostrobus pensilis, of southernmost China, is also VoL. XXVIII] GRESSITT: DAWN-REDWOOD EXPEDITION 29 planted for geomantie purposes, generally along old paths in the delta country. After observing the soft and flexible nature of the foliage and branches, it is easy 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 appearance, 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 elements of possible North American affinity which might have survived with Metasequoia and the other ancient trees associated with it. The latest known fossil deposits otf 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 liy- ing dawn redwoods. What is still more striking is that the present 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 contemporaneous 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 wild animals having survived very sheht, hope and emphasis 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 Kwanetung, southern Hunan, and Kwangsi provinees, 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 trip was to Chiang-chin, a few hours up the Yangtze by steamboat, the other to Pe-pei, a major war-time educational and researelt center, up the Chialing River. Pe-pei is a progressive small city with public 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 we crossed the Yangtze and then the Chialing River to take a smaller steamer, as the first ship had been taken over by the army. Leavine early the next morning, we reached Wan Hsien at dusk. The 30 CALIFORNIA ACADEMY OF SCIENCES [ Proc, 471 SER. wey , RS ~ 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 butte-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 were generally planted with corn, sometimes up to the high- est level, there often being at least three such slopes between the alter- nating cliffs. Generally there were young pine trees (Pinus massoniana) Vou. XXVIII] GRESSITT: DAWN-REDWOOD EXPEDITION 31 erowing on the tops of the mountains. In other areas more or less natural vegetation grows on the steeper, irregular slopes, though corn is often planted on slopes with as much as 65-deeree gradient. The more rolling mountains are often clothed only with 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. Melons 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. Seattered groups of citrus trees, mostly oranges, were found in the low hills and alone the Mo-tau-chi River, mostly near farmhouses. Some pomelo and tangerine yrass, or with seattered pines or c co 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-tane-chin, where we met the Mo-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 alone 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 brought from the southwest. The third afternoon we reached Mo-tau-chi after climbing 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 two 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. My 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, Glyptostrobus, of Canton, which is a relative of the bald eypress, 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 Peiping 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 32 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 41H SrEr. Figure 5. Type tree of Metasequoia glyptostroboides, located at Mo-tau-chi, Szechuan. VoL. XAVIIT] GRESSITT: DAWN-REDWOOD EXPEDITION oo) ot) small ravine guarding the border of Szechuan Province, we entered Hu- 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, flat valley to Wang-chia-ying. Not far from this market town is the largest known dawn redwood, 8% feet in diameter and 115 feet tall, but we put off seeing it until our return because of our bageage 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 eall it. lbs Figure 6. View of upper part of Suisapa Valley. The valley was our principal objective and occupied most of our time for the rest of the summer. When the last of our party left, it had become quite cool. The name Suisapa (Shui-sha-pa) refers to the local, and only Chinese, name, sui-sa, of the dawn redwood. Suisapa specifically is only a single smal] 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 Hsiaoho, the only market town in the 25-mile lone valley. It is this vallev which contains the unusual assemblage of plants of 34 CALIFORNIA ACADEMY OF SCIENCES | Proc. 47a SER. 7 Figure 7. View of the middle portion of Suisapa Valley. ancient northern affinity, many of them being among the 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 the 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 about, with dangerous rapids and narrow gorges up which boats may not be towed, before meeting the Yanetze River between Chungking and Wan Hsien. Foresters who 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 co on Vor. XXVIII] GRESSITT: DAWN-REDWOOD EXPEDITION purposes, particularly for houses, fuel, and coffins. Coffins are made from trunks of Cunninghamia and earried 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 fairly 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. The west 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- siderable drop into a large, deep valley which slopes much more steeply to the south than does Suisapa Valley. On the west side of the west ridge, however, the 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. Chaney’s visit four months earlier, the mayor’s wife and one child had died and the mayor was now sick. We therefore could not lve in his house, and had to be somewhat careful until we had made friends with the people, as they tended to aseribe this bad luck as caused by the foreigners coming and cutting down a dawn redwood for specimens. How- ever, the local people themselves eut the trees not infrequently. There is a prevalent local custom of cutting the branches off the Metasequova and Cunninghamia 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 laree 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 36 CALIFORNIA ACADEMY OF SCIENCES | Proc. 47m SEr. Figure 8. Two pruned Metasequoia trees. Vo... XXVIII| GRESSITT: DAWN-REDWOOD EXPEDITION 37 Figure 9. Metasequoia trees on floor of Suisapa Valley. 38 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 41H SER. “oy . : “2 Figure 10. Metasequoia trees, south-central part of Suisapa Valley. VoL. XXVIIT} GRESSITT: DAWN-REDWOOD EXPEDITION 39 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 vew 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. Many young ones are also planted along the main stream, bordering rice paddies. In the upper portion of the valley they are scarcer. Their altitudinal range is approximately from 2,000 to 4,000 feet. There can hardly be said to be any Metasequoia forests or even moderate natural stands. Most 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 Metasequoia trees 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 warnings 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 Waneg- chia-ying that a group of bandits were trailing us, but they turned out 40) CALIFORNIA ACADEMY OF SCIENCES [ Proc. 471 SEr. to be plain-clothes guards tracking bandits. The wife of a bandit who had robbed one of our guards on a previous trip had been apprehended in a village along our route and was interrogated by the officer during our evening there. In spite of a total of six one-way trips we made )be- tween Wan 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 enerusted with cinders. Rice was cooked in a large iron pot and other things in a large econeave iron pan lke a slice from the edge of a sphere. The local food consists mainly of rice, corn, white potatoes, beans, squash, peppers, 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 were 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 buyine 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 eges, a bouillon cube, and a little pork fat, with sometimes a green vegetable. Lunch con- sisted of one or two eges and two or three small potatoes boiled and earried alone with us. Though the food of these people seemed poor, they had quite enough of it for themselves, and in some respects they were rich. Though they had next to nothing of a ‘‘eivilized’’ nature except their cooking utensils and carpentry tools, they could raise all the food they needed and sell VoL. XXVIII] GRESSITT: DAWN-REDWOOD EXPEDITION 41 some, plus plant drugs, mushrooms, and the like on the market or to the outside world. They build their own houses and coffins and also sell some of the latter. Here their coffins 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 up, but every 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 floors 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 people was rather different from proper Man- darin, or 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 lone gowns of fairly thick, blue, home-made eleth reaching almost to the ground. This was true for men, women and children, and beneath this they usually wore jackets and lone 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 appeared too warm. In the fields, cutting wood, or collecting for us, they would often tie up the lone gown around the waist. These people did not seem to be given to musi¢ as are people of many parts of China. However, one use for it was quite impressive. A type of native communism existed here, whereby quite a bit of the agricul- tural operations 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 evmbal and a drum. The leader sane out short chants, and the whole group joined in the chorus, to the accompaniment of the leader’s evmbal and the drum of his assistant. The tempo of the work was not fast, and there were frequent pauses when all rested. Though I often heard these musical work-ganes, | only onee eame within close range of one, on a steep hillside cornfield. I was prevented from eetting a pieture by curiosity toward me breaking up 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 bit 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 absorbed as the Chinese pressed into the mountains. The 42 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 47H SER. imprint of the former remain in rounder faces, thicker lips, 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 undershorts while washing clothes in the river. We also frequently found hair, flies or other foreign objects in the food, tea leaves in the rice, ete., 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 be- cause these people seemed to have less use for money. This was also re- flected in the fact that we could noi 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 eustom. 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, wildeats, 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 lone- tailed, slender, black and white animal whieh may have been a kind of VoL. XXVIII] GRESSITT: DAWN-REDWOOD EXPEDITION 43 civet. Once one of our boys saw a large dead cat at market, but could not buy it. The only specimens of mammals we got were moles, mice, and bats. 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, erow tits, scimitar babblers, quaker thrushes, other laughing thrushes, Chinese bulbul, yellow-vented bulbul, Chinese fineh-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 (Hnhydris), green grass snakes (Hurypholus), 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 Valley. isapa u SS) of mel Yk E N¢ = u OF SCIE the eastern portion _ ~ SS 7 VI i ACADE with ing LIFORNIA connect Side valley e 12. Figur Vor. XXVIII| GRESSITT: DAWN-REDWOOD EXPEDITION Figure 13. Margin of the middle portion fields and forests above. 46 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 47H SER. 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 without 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 apt to be a large Cupressus (no. 2505), ginkgo, or rarely pine (Pinus massoniana or Pinus mo: 2036). 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 massoniana) 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, | began to wonder strongly if this trip was really bringing us to the high plateaus we had observed. We had been surprised when told that the mountains about which we had inquired were 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 deseribe 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, but the others 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 Vou. XXVIIT] GRESSITT: DAWN-REDWOOD EXPEDITION 47 conspicuously and there were peaks in front of them. Returning 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 clinbing 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 by 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 ot 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 Hsiaoho 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 eame 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 erossed 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 threatened, so we decided to stop for the 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 48 CALIFORNIA ACADEMY OF SCIENCES | Proc. 47it SER. of this range. Though I sealed one, 3500 feet high, passing through tall, wet grass, cornfields, and pine groves with brambles, the view was ham- pered and it appeared that we had gone too far to the east and in too roundabout a way to reach our objective. Along the trail, between these conical peaks, were some of the serried, sharp limestone projections repre- senting peculiarly eroded, vertically tilted limestone strata. These had also been conspicuous 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 might neither of us could sleep at all, the bed- bues, 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 climbed the ridge and followed it to the end. It was amazing to see the extent of rice terracing and the number ot farm- houses and people on the steep slopes of this high ridge. Some ot 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, I had more opportunity to associate with the people. The students of the school (all boys) were interested in our insects. They all smoked, and many of the younger ones had their small bamboo pipes, of various types, 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 Lune- shen-tal 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 Metasequoia. 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 Hsiaoho temple, Just outside the south end of the valley and a mile south of Hsiaoho 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. We 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, papered, outside window. Below our room, side-by-side, were the pigpen and toilet, and below them the common receptacle for human and porcine waste products. In a dirt-floored room nearby we built two stone fire- VoL. XXVIII] GRESSITT: DAWN-REDWOOD EXPEDITION 49 places and above them, suspended from the rafters, our ropes and baskets for drying plants and insects. That afternoon we hired two new collectors and in the 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 peaks 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 community 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 pine. (Pinus no. 2536) with cones shaped something like those of knob-cone pine, though without the knobs. On my return to Wan Hsien I noticed quite a few young trees of this species just south of Mo-tau-chi, the northernmost Metasequova locality. On our way back to Hsiaoho 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 had 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 vallevs as well as on the hillsides, as the drainage was apparently not suitable for rice cultivation. After erossing a ridge, we headed south over somewhat level country into Szechuan Provinee. At Wen-tu, a market town near the border, we rested in a teahouse during the middle of the busy market day. From here 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- spicuous erect remains of others rising from lower surrounding country. We descended our ridge obliquely from 3,400 feet, coming down through 50 CALIFORNIA ACADEMY OF SCIENCES [ Proc, 4vH SER. 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-lo, 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 Ssampan terry 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 eir- 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 landslides. 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 VoL. XXVIII] GRESSITT: DAWN-REDWOOD EXPEDITION 51 night there as it was too late to reach the next market over a high 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 Cunninghamia 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 again. Descending further to rice fields we reached the hamlet of Suen-wu, where there were a few relatively wealthy-looking homes with 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.) We 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 eurved 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. We stumbled down the path for nearly two miles after dark until we reached the 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, 52 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4tH SER. oak, and other native vegetation. We breaktasted at a large farmhouse by the trail under some large cupressine (no. 2505) trees. From the pass, at 2,400 feet altitude again, we had a good view to the south. The valleys were deeper, and in part wider, but still the mountains were of the re- peated thick limestone layers, and with about the same average maxi- mum height. One prominent vertical peak some distance to the south was probably 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 question that we had been passing through our “‘high plateaus,’’ and that my dreams were unfounded. These canyons were apparently quite unsuited for the sup- port of dawn redwoods, as their bottoms were too low, their sides mostly too precipitous, and their tops too dry or too exposed. Descending again through forest, then tung groves and corn, we reached the Shanghai-Chuneking highway in a relatively wide valley bottom with a small river at the market ot Liang-wei-tan, at 1,100 feet above sea level. After a large lunch of noodles and our first eges of the trip, we ended our southward mareh and headed east alone the high- way towards Chien-chiang, to return north by another route. The highway wound alone the sides of canyons, with some switeh- backs and tunnels. Our route was partly by short-cuts, including a climb over a pass. Beyond this pass, the valley leading to Chien-chiane became fairly broad, with some large, nearly flat areas of rice fields. The popu- lation became quite dense, and there was considerable coolie transport alone the highway, though we saw no vehicles pass alone the road. Ilav- ine 20 miles to cover after our near midafternoon start, we did not reaeh the city till well after dark. We were too late to get a regular supper, so again had noodles and eggs. We then found what was apparently the best inn in the city, at the bus stop. While bathing with a basin of water in our room, and tending to some developing sores on my feet, we had several visits from the authorities who were somewhat put out be- cause | had left my passport in Wan Hsien and neither the boy nor I had brought along from IIsiaoho our official letters from the university. The only identification we had was my name ecard. That night we suffered severely from bedbuegs, and I killed over fifty on my bed during several occasions when I foreed myself awake enough to rise, ight the lamp, and hunt them down. Chien-chiange (Kienkiane Hsien) is a large town, the largest by far we had seen since Wan Hsien. It is a distriet city, one of the two eastern- most of southeastern Szechuan Provinee, and not only is near the Hupeh border, but is likewise not very far from both Hunan and Kweichow provinces. From here we ascended the south end of a range and traversed the side of a large broad canyon with partly precipitous sides, still of the Vot. XXVIII] GRESSITT: DAWN-REDWOOD EXPEDITION 53 same thick, horizontal limestone pattern. We then crossed some passes and descended into another valley with a small river which we had to ford. Here naked fishermen were casting their weighted throw (umbrella) nets and catching small fish. They would dive down and eateh some of the fish for certain before pulling in the nets. As we watehed after our swim one of them came up with the anterior end of a wigeline nine-ineh fish in his mouth, to the amusement of the erowine audience. We. fol- lowed this river to its headwaters, and had to cross it many times, first removing shoes, and as it became small, taking running jumps where there were some partial stepping stones. The frequent wetting did not help the sores on my feet, which had originated from bedbug bites except for one blister 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, Just at dark. Our host, an aged, somewhat confused man, was not particu- larly hospitable, as the tiny inn was already overcrowded. The food, again, was poor, but the boiled potatoes were weleome once more as a change. The inn consisted of two rooms, one the kitehen. 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 durine the mornine. We seemed to be crossing numerous passes between branches 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 (14% pounds), but had to earry 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 where a braneh of the trail had lead to a ford not lone before. However, the boatman managed to push up- 54 CALIFORNIA ACADEMY OF SCIENCES [ Proc, 4TH SER. stream above the branch and then crossed with us to the northwest bank of the Chen. We now traveled northeast along the west bank of the Chen River. 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 eliff 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 cliff in the middle of the steep slope leading up to the summit of a high ridge. I realized we were approaching the terminus point 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 speak 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 1,600. That 1,500-foot climb 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 eo to Tsung-lo. A man guided us along the trail until there were no further branches to confuse us, but we found the path full of rocks and Vor. XXVIII] GRESSITT: DAWN-REDWOOD EXPEDITION 55 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, eges, 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 fire, 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 we 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, we 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. We 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 lone rows of sharp canine teeth. This eon- tinued, through cornfields, practically all the way after getting out of the valley in which we had spent the night, and until we reached the trail down which we had started the trip, a few miles south of Hsiaoho. Thus ended our somewhat disappointing 165-mile, 51-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 had 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. We never received that bateh of letters. That evening I received two letters forwarded from Chuneking, 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 we 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 T wore straw sandals. I sent the three boys out daily, hiring as much local help in addition as possible, and had more of the people bring me in 56 CALIFORNIA ACADEMY OF SCIENCES [ Proc. 4TH Ser. specimens of any sort they came across. By taking care of the plant drying myself and preparing labels and other materials I was able to keep the boys out in the field longer, but still my trouble was greatly handicapping. 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, codeseriber of the Metasequoia, included Professor K. L. Chu, plant ecolo- gist from Shanghai, and Mr. C. T. Hwa, 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 erude 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 parts alone small precipices. Once I nearly upset the 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 might, 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 we were going to cross to the east side of the Mo-tau-chi River after passing through part of a heavy downpour, the boatman velled 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. Within six minutes the water level below us rose two feet, and was still risine when we gave up and went back into the village to find an inn. Apparently 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 econ- tinued only about three miles, to find that the water here was still too VoL. XXVIII] GRESSITT: DAWN-REDWOOD EXPEDITION | ol swift for the recrossing to the 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 Hsien until the forenoon of the sixth day. I spent three days waiting for a ship to Hankow, staying at the home of the Reverend and Mrs. J. B. Matson. With the welcome, good food, and comfortable rest, after the lone 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, the 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 Hone Kone 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’ N.), 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 reae- tions in different seedlings, some becoming brown when others were still almost entirely ereen. 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 appearing as soon as the temperature began to rise. Actually we had only about five of some thirty seedlings die. The pereentage 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 Hsien 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 Jetasequoia seeds and to take more pho- tographs, with a camera I sent by airmail from Canton to Wan Hsien. 58 CALIFORNIA ACADEMY OF SCIENCES [ Proc, 4rH Ser. They experienced snow while returning over the Chi-o Shan ridge, and returned to Canton in mid-November, after an absence of nearly five months. 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 Nearetie relationships. Some of the material has been reported upon in some of the references cited below. REFERENCES CHANEY, R. W. 1948. Redwoods around the Pacific basin. Pacific Discovery, vol. 1, no. 5, pp. 4-14. 1951. A revision of fossil Sequoia and Taxrodium in western North America based on the recent discovery of Metasequoia. Transactions of the American Philosophical Society, new series, vol. 40, pt. 3, pp. 171— 263, pls. 1-12. FLORIN, RUDOLF 1952. On Metasequoia, living and fossil. Botaniska Notiser, hafte 1, pp. 1-30. GREssITT, J. L. 1950. The hispine beetles of China (Coleoptera: Chrysomelidae). Lingnan Science Journal, vol. 23, pp. 53-142, pls. 4-8. 1951. The longicorn beetles of China. Longicornia, vol. 2, pp. 1-667, 22 pls., 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, pls. 27-36, 1 text fig., 4 maps. MERRILL, E. D. 1948. Metasequoia, another “living fossil.” Arnoldia, vol. 8, pp. 1-8, 2 pls., 1 map. STEBBINS, G. L., JR., 1948. The chromosomes and relationships of Metasequoia and Sequoia. Science, vol. 108, pp. 95-99. WYMAN, D. 1951. Metasequoia brought up to date. Arnoldia, vol. 11, pp. 25-28. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES FOURTH SERIES Vol. XXVIII, No. 3 July 15, 1953 SOME NEW AND NOTEWORTHY APHIDAE FROM WESTERN AND SOUTHERN SOUTH AMERICA (Hemiptera-Homoptera) BY E. O. ESSIG Department of Entomology and Parasitology University of California, Berkeley During the months of December, 1950, to and ineluding April, 1951, Dr. A. E. Michelbacher and his wite, Martha, and Dr. E. S. Ross and his wite, 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 Franciseo on the Grace Line steam- ship Santa Jwana—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 large 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 [ 59 | 60 CALIFORNIA ACADEMY OF SCIENCES {[Proc. 4TH SrEr. which were preserved in alcohol and turned over to the writer for mount- ing and study, and for the publication of desirable information concerning them. | Little previous investigational work has been done on the aphids of Chile, in which country 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 deseribed 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, Tueuman, Salta, Jujuy and La Quiaea, Argentina; through Camargo, Po- tosi, Oruro, and La Paz, Bolivia; via Lake Titicaca, Puno, Tinta, Abaneay, Ayacucho, Huaneayo, 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. Michelbacher 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 been recorded by. ento- mologists in the South American countries visited: Colombia, Peru, Chile, VoL. XXVIIT| ESSIG: APHIDAE OF SOUTH AMERICA 61 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 Mrs. A. E. Michelbacher, and to Dr. and Mrs. E. S. Ross for the opportunity to mount and study this fine collection of South American aphids. 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 especially 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 Provincia. The total collections of aphids amounted to 25 genera; 55 species, in- eluding 14 new species; and 3,395 specimens mounted on 781 slides. CLASSIFICATION AND DESCRIPTIONS OF SPECIES Order HEMIPTERA Suborder HOMOPTERA Superfamily APHIDOIDEA Family LACHNIDAE Subfamily LACHNINAE (PTEROCHLORINAE) Tuberolachnus saligna (Gmelin) Giant Willow Aphid (Figure 1) Aphis saligna GMELIN, 1788; A. polaris CurTiIs, 1828; A. viminalis B. p. Fonsco- LOMBE, 1840; Lachnus dentatus Lr BARON, 1872; L. fuliginosus Buck Ton, 1891; Tuberolachnus viminalis (B. de Fonscolombe) MorpviLKko, 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: 62 CALIFORNIA ACADEMY OF SCIENCES [|Proc. 4rH Ser. a VIM fyy Oly y, (agen se Com si nae e ER yk eae 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, Provinee of Bio-Bio, January 1, 1951; apterae and alatae. ARGENTINA: On Salix sp., Uspallata, Province of Mendoza, February 6, 1951. Many specimens oe Bee and alatae. Other records by Blanchard 1926, pp. 327-29; 1939, 871-72; 1944, 16) are: On Salix sp., ee Province of Buenos Aires; also from San Juan and Zapala. Vout. XXVIII] iSSIG: APHIDAE OF SOUTH AMERICA 63 On English walnut (nogal or noguera) Juglans regia Linnaeus at San Rafael, Province of Tucuman. PERU: On Salix sp., Barranea, March 15, 1951; all apterae. All collected by Dr. A. E. Michelbacher. Family CHAITOPHORIDAE Subfamily SIPHINAE Sipha flava Forbes Yellow Sugarcane Aphid This is a subtropical yellowish species occurring on Gramineae in warm, temperate and tropical North and South America. Although not taken by Dr. Michelbacher the species has been reported in two of the South American countries where sugareane is grown. ARGENTINA: Blanchard (1944:17) records this species on sugarcane (cana 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. Family CALLAPHIDAE Subfamily CALLAPIDINAE Chileaphis Essig, new genus (Figure 2) Alate 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 truneate 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 michelbacheri Essig. 64 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H SEr. Vat Sie — Pugs CER Vy >) R 4 WY ventral fe See Figure 2. Chilean cypress aphid, Chileaphis michelbacheri 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. VoL. XXVIII] ESSIG: APHIDAE OF SOUTH AMERICA 65 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 eauda, 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 cireular 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, 0.85 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 shghtly curved. Second fork of media arises half-way be- tween first fork and wing margin. Abdomen with six distinet pigmented elandular transverse areas. Cornicles dusky, as illustrated; with few short hairs. Cauda unusually large and peeular, dusky, chitinized and with only a few short hairs. Anal plate bilobed, with many short setae. Genital plate as drawn. Leneth of body, 2.45 mm.; forewing, 2.8 mm. Apterous parthenogenetic female: Whitish-gray; in living condition apparently covered with powdery wax; widely oval with short antennae and lees as delineated. Lengths of antennal segments: III, 0.28 mm.; IV, 0.09 mm.; V, 0.10 mm.; VI, 0.12 mm.; unguis nail-lke. Cornicles truncate cones, the base about twice the diameter of the opening, with few short scattered hairs. Cauda large, globular, 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 1.74 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 position 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. Michelbacher by beating the available limbs and foliage of the Chilean cypress, Pilgerodendron uviferum (Don) Florin. The synonymy of this host plant is given by Record and Hess (1943) as follows: (Juniperus uvifera Don = Thuja tetragona Hooker—Libocedrus tetragona [Hooker] Endl. JL. cupressoides Sargent = L. wifera |Don| Pilg. ) 66 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4tH Serr. 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 cireular and transverse sensoria on the antennae of the alate male. Neuquenaphis' michelbacheri Essig, new species Nothotagus Aphid (Figure 3) 2 7727? PPD PPAIPLALS a 32 > . % Ne ye? Drow ce Figure 3. Nothofagus aphid, Neuquenaphis michelbacheri 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 Neuquen, Argentina. VoL. XXVIII] ESSIG: APHIDAE OF SOUTH AMERICA 67 Type: Alate parthenogenetic female: Color uncertain, the cleared aleo- holie specimen indicates that the head, antennal seements 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 that in life a 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 apterous 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 8-16. Host plant and locality: The species was collected by beating the branches of Nothofagus dombei Blume. It was collected along with Spica- phis michelbachert and Neuquenaphis chilensis on the Niagara Rancho near Temuco, Province of Cautin, Chile. Thirteen apterous specimens and 3 alates were also collected in San Andres, Paranque, Chile. Collector, A. E. Michelbacher. The specimen described was selected from 16 adult alate parthenogentie 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. Neuquenaphis michelbacheri differs from N. edwardsi (Laing) (Jyzo- 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 PX. 4. The presence of a prominent tuberculate seta or spine on the outer surface of the cornicle. 5. Somewhat more elongated cauda. Neuquenaphis chilensis Kssig, new species Dombei Aphid (Figure 4) Type: Alate parthenogenetic female: Color uncertain because speci- 68 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SEr. Figure 4. Dombei aphid, Neuwquenaphis chilensis Essig, new species. a, tip of antenna; b, body tubercle; c, frontal tubercles; d, antenna showing 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 Neuquenaphis michelbacheri. 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, eylindrical 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 wines; 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 144 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 which 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 truneate, with a single lateral marginal spine and somewhat imbricated; the hase is twice the diameter of the apex. The eauda is conspicuous; knobbed, Vout, XXVIII] ESSIG: APHIDAE OF SOUTH AMERICA 69 being widest at the base, thickly beset with small pimple-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 eauda. Leneth of body 2.20; forewing 2.60 mm.; antennae 1.30 mm.; cornicle 0.10 mm.: eauda 0.15 mm. This species differs from Newquenaphis michelbacheri and N. edwardsi in having tubercles on the head and thorax; capitate cauda; very much shorter antennal uneguis; and fewer antennal sensoria. Host plant and locality: This species was collected along with the N. michelbachert by beating the branches of Nothofagus dombei at Hacienda San Andres, near Purranque, Province of Osorno, Chile, January 15, 1951, by Dr. A. E. Michelbacher. The type was selected from three alate specimens, all that were collected. Subfamily SPICAPHINA Spicaphis Essig, new genus (Figure 5) Apterous viviparous female: Robust, the dorsum of the entire body beset with thick, rugose, glandular-like spicules or tuberecules arranged for the most part in longitudinal and transverse rows; each spicule has a terminal glandular spine or seta. Similar spines also oceur on the head, two basal antennal segments and cornicles. Antennae 6-segmented; eyes compound with terate tubercles. Cornicles cone-shaped and with a single tubereulate spine. Cauda knobbed. Anal plate with shallow median con- striction. Genotype: Spicaphis michelbacheri Essig. Spicaphis michelbacheri Essig, new species Michelbacher 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 70 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47H Serr. { €. Abernathy Figure 5. Michelbacher aphid, Spicaphis michelbacheri 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 subequal; 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 by beating the limbs and foliage of Nothofagus dombei Blume, a tree native to southern Chile. It was taken along with Newquenaphis michelbacheri deseribed 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 more nu- merous body tubercles. The true apterous forms of N. chilensis are unknown. VoL. XXVITIT] ESSIG: APHIDAE OF SOUTH AMERICA fiall Family APHIDAE Subfamily APHIDINAE Tribe Rhopalosiphonini Hyalopterus arundinis (Fabricius) Mealy Plum Aphid (Figure 6) Aphis pruni GEOFFROY, 1762; A. arundinis Fapricius, 1775; Hyalopterus pruni, Kocn, 1854. 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, 72 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. prunes, apricots, apples, peaches, and related plants, and usually spends the summer on reed grass, Phragmites phragmites (l.) and eat-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 Aphis maidis (Frrcn) 1856. (Figure 7) Figure 7. Corn-leaf aphid, Rhopalosiphum 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. VoL. XXVIII} ESSIG: APHIDAE OF SOUTH AMERICA 73 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: PERU: On Johnson grass, Sorghum halepense, Callao, November 14, 1950— very abundant. On cultivated corn (maiz), Callao, November 17, 1950—1 alate and 3 apterae. On cultivated corn, Cusco, Provinee 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., Avena 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 Holaretie 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. PERU: On Canna sp. in the Botanical Gardens, Lima, November 15, 1950— many alatae. Rhopalosiphum prunifoliae (Fitch) Apple-Grain Aphid (Figures 9-10) Aphis prunifoliae Fircu, 1855; Rhopalosiphum pseudoavenae Patcu, 1917. This aphid is a widely distributed economic species in North America 74 CALIFORNIA ACADEMY OF SCIENCES [Proc, 411 Ser. I ——_—._ ; i os J ore LOW Pa ERE TP TE POET} DIN VAD 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 infests grains, corn, and grasses. In the colder areas it may overwinter on apple and other deciduous fruit trees. CHILE: Sweeping, Los Muermos, Provinee of Llanquihue, January 20, 1951 —3 alates. PERU: Beatine plants at Andahuavlas, Provinee of Apurimaec, February 7 ? . >] . > 1951—1 alate. On eultivated corn, Zea mays L., Cuseo, Mareh 4, 1951—apterae only. ? ‘ b b} b, « It is a serious pest of sweet corn in California. VoL. XXVIII] ESSIG: APHIDAE OF SOUTH AMERICA 75 Figure 9. Apple-grain aphid, Rhopalosiphum prunifoliae (Fitch). Colonies on an ear of corn. (Photo by Oscar Bacon, August 31, 1951.) ARGENTINA : It is possible that some of the records of Rhopalosiphum pseudoavene (Patch) on rye (centeno), maize (maiz), wheat (trigo), and Bromus sp., 76 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4rH Serr. in Argentina, may refer to this species. Blanchard, 1939: 908-911; 1944:19). Figure 10. Apple-grain aphid, Rhopalosiphum prunifoliae (Fitch). Alate and apterous parthenogenetic females and enlarged antennae, caudas, cornicles, and rostrum. Rhopalosiphum pseudobrassicae (Davis) Turnip Aphid (Figure 11) Aphis pseudobrassicae Davis, 1916. This may be 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. Boulvia: On a native Nasturtium 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 (Raphanus sativus 1..). VoL. XXVIII] ESSIG: APHIDAE OF SOUTH AMERICA ~] ~ PERU: On cultivated stocks, Methiola sp., at Callao, Province of Lima, No- vember 4, 1950—many apterae and alatae. TO eo EG oS een NVTOD DAS cL OHV OPAC Gra ad A707 a a so WICC2 SUIS ACULIOL STOO S/ASILDNS (ares ASESSONIE CTE Figure 11. Turnip aphid, Rhopalosiphum pseudobrassicae (Davis). Alate and apterous parthenogenetic females with enlarged antennae, cornicles, genital plate, anal plate and cauda, and rostrum. Rhopalosiphum splendens (Theobald) Subterranean Aphid (Figure 12) Siphonocoryne splendens THEOBALD, 1914; Rhopalosiphum subterraneum Mason, 1937. This aphid 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 78 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4ruH SER. Figure 12. Subterranean aphid, Rhopalosiphum 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. 8S. Ross. Beating nightshade (Solanum sp.) at Andahuaylas, Province of Apu- rinac, February 7, 1951—a single alate specimen. Tribe Aphidini Aphis alstroemeriae Hssig, new species Alstroemeria Aphid (Figure 13) Type: Alate parthenogenetic female: Mostly 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} ISSIG: APHIDAE OF SOUTH AMERICA 79 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 0 to 2. (In paratypes the variation is III, 5 to 11; IV, 0 to 2.) Rostrum slender and short, extending to the third coxae. Prothoracie tubercle present. Wings normal, second fork of media arises near the middle of the distance from first fork to margin of wings. Legs with seattered 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. Leneth of body 1.75 mm.; hind tibiae 0.10 mm. ; cornicle 0.17 mm. ; cauda 0.14 mm. ; antennae 1.29 mm. Apterous parthenogenetic female: Robust in form and generally dark in color. The dorsum has a mosaie pattern of lines and dark and lighter usually 5-sided tile-lke 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, eauda 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. Number 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, Alstrameria sp., 40 kilometers east of Los Andes, Province of Aconcagua, Chile, February 4, 1951, by Dr. A. E. Michelbacher. This species differs from Aphis medicaginis (Koch) in having secondary sensoria on antennal segment IV and from A. rwmicis Linn. in having fewer hairs on the cauda. 80 Aphis citricidus (Kirkaldy) Tropieal Citrus Aphid (Figure 14) (5 Myzus citricidus Kirk apy, 190 cola VAN DER Goot, 1912. CALIFORNIA ACADEMY OF SCIENCES [ Proc. 47H SER. 7: Aphis tavaresi DEL Guercio, 1908; Aphis citri- Figure 14. Tropical This is a medium-sized black and injurious to the tender apical shoots of citrus aphid, Aphis citricidus (Kirkaldy). aphid which is often very abundant on its many host plants. It VoL. XXVIII} ESSIG: APHIDAE OF SOUTH AMERICA 81 chiefly occurs in the tropical and warm-temperate regions where it feeds mostly upon Camellia, Citrus, Hibiscus, Mangifera, and other plants in the more tropical areas of Japan, China, Pacifie Islands, Australia, Africa, Southern Brazil, and Argentina, and in ereenhouses throughout the world. CHILE: In Chile it was first reported and deseribed as Aphis citricola by P. Van der Goot in 1912. Loeality and host plant were not given. ARGENTINA: On Citrus spp., at Jujuy, Province of Jujuy, February 15, 1951, by Dr. A. E. Michelbacher. It was abundant on the foliage. Three apterae and 3 alates were taken. It appears to the author that Blanchard’s Paratoroptera 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. Babamondes ecol- lected on Salix babylonica L., Guaymallen, Province of Mendoza, 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 ecauda pale. An- tennae relatively short, with few short hairs; roughly imbriecated. Seg- ments III and IV somewhat swollen, III with 8 to 10 large cireular 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 ITT, all of segments [V—VI, and the ecornicles 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 82 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SEr. AD al a ) iA ayy s) SUNN LAE \ ye MS a ye pps EDU TPE ar} NEALE yee el Te) 1) '2) PE PD UR a phy J)2 ) ry Zz FOL Figure 15. Coreopsis aphid, Aphis coreopsidis Thomas. 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. 18), reports this species on 3idens megapotamica (Speng.) at Missiones, in 1937. This aphid was deseribed by Cyrus Thomas in 1878 and has been reported on a number of Compositae in many parts of the United States from Connecticut to California. Aphis gossypii (Glover) 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 VoL. XXVIII} ESSIG: APHIDAE OF SOUTH AMERICA 83 ge 3 a a Rol fer = So SP aT ee Se et STEM GUL CRED NCCU CAM OE (ICs oro nda co AE arn ig nw fi re a Ss fe S | [: As 5 VNVERRIN Rawal ay \ pede, at pers ) i ok 6 ~_—__—_Y =e = \ . a Ss : a» % Se ie rf ¢ BY SE SS we oa Figure 16. Cotton aphid, Aphis gossypii 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 Ameriea as indi- cated by the reports of the entomologists of that continent. It was collected as follows: CHILE: On squash, calabaza (Cucurbita maxima), Botanical Garden, Lima, November 14, 1950. Alatae and apterae. On Bignonia sp. (2) at Anthofagasta, November 21, 1950. Many apterae. 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- euman, 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. 84 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4rH SER. Figure 17. Cotton aphid, Aphis gossypii Glover. Alate female and rostrum, tarsi, cornicle, cauda, antennae, cauda. On Asclepias sp., Jujuy, Provinee 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, Chenopodium sp., Jacaranda ovalifolia R. Br., Guava (guaya), Lilium sp., (azucena), Lepidiwm sp., Hibiscus sp., Solanum sisymbrufolium Lam., Begonia sp., Citrullus vulgaris Sehrad., Cycloma spp., Dyckhia flori- bunda Girs., Lavatera arborea, Mandevillea suaveolens Lindl., Morrenia odorata (Hook et Arn.), Oleaceae, Orthostemon sellowianus Berg., Persea americana Mill., Pyrus malus li, 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 Ii., Mendoza Capital, October 15, 1948—many specimens. VoL. XXVIIT] ESSIG: APHIDAE OF SOUTH AMERICA 85 On Crataegus sp., Mendoza Capital, December 12, 1950—many specimens. On Parthenium hysterophorus, Mendoza Capital, October 16, 1950— many apterae and alatae. COLOMBIA : On Atriplex sp. (?), Buenaventura, November 4, 1950. Many alatae and apterae. On Baccharis sp. (?), November 4, 1950—a few alatae and apterae. On Composite (7), November 4, 1950—1 alate and 3 apterae. On a shrub, November 4, 1950—many apterae and a few alatae. On Casuarina in Botanical Garden, Lima, November 14, 1950—many apterae and alatae. On Pomegranate, Punica granatum L., Callou, November 14, 1950— a few aptera. On squash, Cucurbita maxima L., Botanical Garden, Lima, November 14, 1950—9 apterae and 6 alatae. On cultivated potato, Solanum tuberosum L., Sieuani, Province of Cuseo, Mareh 2, 1951—1 specimen. On cultivated cotton, Gossypium sp., Chanea, Province of Lima, March 15, 1951—4 apterae. Aphis illinoisensis Shimer Grapevine Aphid (Figure 18) Aphis ampelophila BLANCHARD, 1913. This is a large, black North American species which commonly feeds on grapes and related plants. CHILE: On a wild Berberis sp., 2 km. west of Porto Veras, Provinee 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 deseribed from specimens taken in Argentina by Del Guercio (1913, p. 159) as Aphis ampelophila which is a synonym of A. ilinorsensis. 86 CALIFORNIA ACADEMY OF SCIENCES | Proc. 4TH Ser. 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 figured. Antennae black, shorter than Figure 19. Cabildo aphid, Aphis marthae Essig, new species. A, alate; B, aptera. a, cauda and b, cornicle of alate: ¢, reticulations on dorsum of aptera; d, cornicle and e, antennae of alate. VoL. XXVIII] ESSIG: APHIDAE OF SOUTH AMERICA 87 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: I-14 11-15, 12-15, 112-16, 138-14, 13-16, 15-15, 15=18, 16-18, 16-19, 17-19, 17-21, 18-18, ete.) ; IV with 1-2 sensoria. (Paratypes with ihesollowing numbers: 0-1, 0-2, 0-3, 1-2) 1-3, 9-3, 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.) Leneth 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 parthenogenetic female: Of the same general shape, size, and color of Aphis medicaginis Koch whieh it also resembles in having the mosai¢ 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. Michelbacher. 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 paratypes. This species most resembles Aphis rumicits Linnaeus, but differs in the following respects: The body 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 fewer hairs. It differs from Aphis medicaginis 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 bazzi Blanchard in being more heavily pigmented ; with shorter antenal uneuis; 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. 88 CALIFORNIA ACADEMY OF SCIENCES [Proc. 471 Serr. It was named for Mrs. Martha Michelbacher, one of its collectors, in recognition for her contributions to the South American aphid survey. Aphis medicaginis Koch,* 1854 Cowpea Aphid (Figure 20) This aphid is shiny black in life, especially the apterous forms with the distal portions of the legs distinetly 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. 7), Valparaiso, November 17, 1950; many apterae, On Cassia sp. Pedegua, Province of Aconcagua, November 28, 1950. Many apterae and 1 alate. On thorny legume, Ilapel, Province of Coquimbo, November 28, 1950 —10 apterae. On Artenusia sp. ?, 90 km. east of end of tunnel, Hlapel, November 28, 1950 On Composite (Artemisia sp. ?), 90 km. east of end of tunnel, Ilapel, November 28, 1950—6 apterae and 2 alatae. 5 apterae and 1 alate. On Cereus sp., Las Palmas, Provinee of Aconcagua, November 29, 1950—apterae and alatae. On Cassia sp. ?, Illapel, Provinee of Coquimbo, November 30, 1950— apterae only. On vellow-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 medicaginis 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 Visé 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. VoL. XXVIII} ESSIG: APHIDAE OF SOUTH AMERICA 89 we | RAR IBDS Seo, RRO ew bn . e ops dite Qe Figure 20. Cowpea aphid, Aphis medicaginis Koch. Adults and antenna, cauda and cornicle of aptera. (After Zimmerman, 1948.) On leguminous tree, Valparaiso, December 17, 1950—apterae and alatae. On Baccharis sp. ?, Chillan, Province of Nuble, December 27, 1950— many apterae and 1 alate. Very small specimens. 90 CALIFORNIA ACADEMY OF SCIENCES |Proc. 4rH Ser. On thorny legume, Antuco, 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 Solanum sp., Los Andes, Province of Aconcagua, February 4, 1951, many apterae and alatae. On wild nasturtium (nasturcia), Portillo (3,000 feet), Provinee of Aconcagua, February 4, 1951. Many apterae and alatae. ARGENTINA: On Astragalus sp. ?, La Quiaea, Province of Jujuy, February 18, 1951 —many specimens apterae and alatae. In addition, Blanchard (1939, pp. 914-916; pp. 922-9238) lists the following host plants (as Aphis fabae Scopoli and A. laburni Kaltenbach) : Acacia sp., Arachis hypogaea l., Cicuta sp., Dyckia floribunda Gris., Eryngium spp., Euonymus sp., Eucalyptus spp., Foeniculum vul- gare, Gladiolus sp., Cirsium sp., Chrysanthemum sp., Cosmos sp., Dahlia sp., Phaseolus lunatus L., Phoenix sp., Gerbera jamesona, Senecio bonariensis H. & A., Sesbania punicea (DC), Spiraea chamaedrifolia li., Vicia 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 Gloxinia sp., Mendoza Capital, Mareh 10, 1948—a few apterae. (COLOMBIA : On lecume, Buenaventura, November 4, 1950. Collection consisted of z 4) b] 21 apterous females. PERU: On broad bean, Vicia faba, Callao, November 4, 1950—5 alates and 10 apterae. On Astragalus sp., Sicuana, Province of Cuseo, Mareh 1, 1951—many specimens, all apterae. On Cassia sp., Cuseo, Province of Cuseo, Mareh 2, 1951—many speci- mens—all apterae. On thorny legume, Cuseo, Provinee of Cusco, Mareh 2, 1951—all apterae. VoL. XXVIII] ESSIG: APHIDAE OF SOUTH AMERICA 91 Aphis nerii Boyer de Fonscolombe, 1841 Oleander Aphid (Figure 21) ears > Drshees a Figure 21. Oleander aphid, Aphis nerii (Boyer de Fonscolombe). Alate and apterous parthenogenetic females with enlarged antenna of alate and cornicles and caudas of alate and apterous forms. A bright yellow and black species. Cerosipha nerii (Boyer de Fonscolombe) BORNER, 1952. This is a widely distributed aphid occurring on Asclepias spp., Caltro- phis spp., Gonolobus 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- eagua, December 17, 1950—1 alate. Beating (plants not ascertained) near the mouth of Maullin River, Provinee of Llanquihue, January 22, 1951—a large number of apterae and alatae. 92 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4rH SER. BOLiviA: On Asclepias sp., Cumergo, February 20, 1951—many apterae and alatae. ARGENTINA: On Nerium oleander L., Araujia sericofera Bert., Asclepias curas- savica L., and Clematis sp. (Ornamental) by Blanchard (1923, pp. 39-41; 1939, pp. 925-26). PERU: On a vine-like Asclepias (?), 65 miles west of Cuseo, 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) > _ = : Se a e205 DEAS Ee OVesesaecn Ree rassanulcainada gal (om SPER ETOWEGI TC EAP SSESECLLE Gens G(G Gt ( eta ca ‘ ‘J Taye (COO) Ces CC 6 * FASS Saran COT (eT PE COaee RENIN MA Ze GUAR OLR Figure 22. Bean or dock aphid, Aphis rumicis Linnaeus. Apterous partheno- genetic female and enlargements of a, anal plate and cauda; b, tarsus; ¢, lateral abdominal tubercle; d, mosaic and glandular areas; e, cornicle; f, rostrum; g, antenna. VoL. XXVIII} ESSIG: APHIDAE OF SOUTH AMERICA 93 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 only. 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 Lianquihue, 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. eae DEO cas ee (CE endck 2 Figure 23. Bean or dock aphid, Aphis rumicis Linnaeus. Alate parthenogenetic female and enlarged cauda, tarsus, cornicle, rostrum (venter), and antenna. 94 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. On wild currant ? (grosellers), Hacienda San Andres, Parranque, Provinee of Llanquihue, January 15, 1951—apterae and alatae. ARGENTINA: On wild potato (papa), Salta, Provinee of Salta, February 15, 1951— 1 single alate. On Solanum sp., Salta, February 15, 1951—many specimens; many apterae and 1 alate. On thorny legume, Jujuy, Province of Jujuy, February 15, 1951— many specimens, all apterae. x OOS EEC Sy (fas oe ys ay, & ee vi Figure 24. Bean or dock aphid, Aphis rumicis Linnaeus. Sexual female and enlarged anal plate and cauda, antenna, cornicle, rostrum (venter) and hind tibia showing sensoria. a2 * TOSS eo “ 6 °00%0 2% 8 ANY, = St C Qe ) O° <4 O8 ae 959 oe ‘. YP y fe 3S ~ G' @) oo ae) one 2. On Solanum sp., Jujuy, February 16, 1951—many apterae and alatae. Blanehard (1935: 373-375; 1939: 929-930) records it On Rumesx sp., at Tueuman. On Rumex sp., Cestrum parqu L.’Herit, Solanum lycopersicum L., S. nodiflorum Jaeq., 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 Tulipa sp., Las Heras, Mendoza, October 10, 1950—apterae and alatae. VoL. XXVIIT} ESSIG: APHIDAE OF SOUTH AMERICA 95 On Rumex sp., Mendoza, Guaymallen y Capital, November 1, 1950— apterae only. On Eugenia sp., Mendoza Capital, May 10, 1948—apterae. On Solanum sp. (beating), Ando Huylas, Province of Mendoza, Feb- ruary 15, 1951—1 alate. ee a a ae Aye Cae ORNS at WEE ORD) Ker T IGS CEL ae aneet Gisse 6 OL SEQo'R oe loge SS yor~ et? (62 Ola elu (Sia SAL OVS Figure 25. Bean or dock aphid, Aphis rumicis Linnaeus. Alate and enlarged tarus, cornicle, anal plate and cauda, rostrum, and antenna of same. PERU: On Agave sp., Rio Pampas, Province of Purimac, March 8, 1951— many apterae and alatae. About 60 per cent killed by a fungus (hongo). On Composite (?), on road 65 miles west of Cuseo, Mareh 5, 1951. Only apterae. On Composite, Cuseo, March 5, 1951—1 alate. Toxoptera aurantii (Boyer de Fonscolombe) Black Citrus Aphid (Figure 26) Aphis aurantii BoyER pre FoNSCOLOMBE, 1841; Aphis camelliae KALTENBACH, 1843; Toxoptera aurantiae Kocu, 1856. 96 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. — FO RO 7 a a te Senet <= Pe es * Bre © Figure 26. Black citrus aphid, Toroptera 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, Coccoloba, Coffea, Ficus, Gar- denia, Hibiscus, Ilex, Mangifera, Persea, Straussia, Theobroma, 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 buxifolia Reiss., and Vibur- num tinus Li. COLOMBIA: On mango (Persea sp.), Buenaventura, November 5, 1950—many apterae. VoL. XXVIII] ESSIG: APHIDAE OF SOUTH AMERICA 97 Subfamily ANURAPHINAE Tribe Anuraphidini Subtribe Anuraphidina Brachycaudus helichrysi (Kaltenbach) Leaf-eurl Plum Aphid (Figure 27) == Ss. S— Se = , = Es 2 & Gis 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 BUCKTON, 1879; A. verbenae MAccutati, 1883; A. leonopedii SCHOUTEDEN, 1903; Anura- phis cyani THropaLp, 1915; A. abrotaniella THEOBALD, 1919; A. cantauriella THEOBALD, 1921; A. sherardiae, THEOBALD, 1926. 98 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SEr. This small, pale aphid has conspicuous dark body markings. It is widely distributed throughout the temperate regions of the world and attacks many kinds of plants. CHILE: Very abundant on peach (Prunus 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 many apterae. ARGENTINA : Blanchard (1922, pp. 54-56) 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 lL. 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 Goort, 1915. VoL. XXVIII] ESSIG: APHIDAE OF SOUTH AMERICA 99 5 Oo. F =) = 19 00°) BG) , on\ 0 20’o 2 2010.0) — ery AO ar TOO KO IIL ne, 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 Holarctie 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. 100 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SEr. < —S NOY nn on 2S 8 S02 o SS ARS Sat SSE [See Willa Ee 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., Raphanus 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 spinosa, Botanical Garden, Lima, November 14, 1950. VoL. XXVIIT] ESSIG: APHIDAE OF SOUTH AMERICA 101 Hyadaphis conii (Davidson) Honeysuckle Aphid (Figure 30) OBES SLEEVE TOD Rs PS SOS os AEE = WORT eo. Ae seo a 8 fT ey Won ) - ro : ia: 35 ] Miws Lieu ggasane: yore. Figure 30. Honeysuckle aphid, Hyadaphis conti (Davidson). Enlarged drawings of antennae, cornicles, rostrum, caudas, of aptera and alate and wing of alate. Siphocoryne conii Davipson, 1909; Aphis xylostei ScCHRANK, 1801; Hyadaphis co- niella THEOBALD, 1925; H. sii BORNER, 1931-32; Rhopalosiphum melliferum Horres, 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 particularly infests Lonicera spp. and members of the large family Umbelliferae. CHILE: A single collection of many apterae and alatae was made on Caruwm 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 Scoprori, 1763; Cavariella capreae (Fabricius), 1775 (Many authors). This is probably a Holarctie species commonly occurring in Europe and North America. It has been introduced into many parts of the world. 102 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. SUPEADT RWS MADE Cn ia 6) Figure 31. Parsnip and willow aphid, Cavariella aegopodii (Scopoli). Alate and apterous parthenogenetic females and enlarged antennae, cornicles, caudas, and caudal tubercle of female (lateral aspect). (After Zimmerman, 1948.) CHILE: In Chile it appears to be widely distributed on umbelliferous 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, Anethum 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. VoL. XXVIIT] ESSIG: APHIDAE OF SOUTH AMERICA 103 ARGENTINA : Under the above specifie name and also as C. capreae (Fabricius) this species has been recorded: On Apium sp., Daucus sp., Carum sp., Pastinaca sp., Pimpinella ani- sum L., and Salix sp. (Blanchard 1925, pp. 14-16; 1944, pp. 31-32). Specimens have been received from L. A. Bahamondes, collected on Salix babylonica 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 circumflera BUCKTON, 1876; S. callae HEINRICH, 1909: Myzus vincae GILLETTE, 1908. This is a European species now widely distributed on lilaceous 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., Tulipa sp., and other Lilaceae, Orchi- daceae, and Vinca major L. Myzus ornatus 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 104 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Ser. ee 4s Saarinen RRA TENE Laer (ET J f aA 7 Da saan ; v7 Se oPal oes 6 65.9 6 Syl A) oe Ss ALE a aOiane CR gee p Figure 32. Lily aphid, Myzus circumflecus (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 ¢, 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. VoL. XXVIII] ESSIG: APHIDAE OF SOUTH AMERICA 105 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 partheno- genetic females with enlarged antennae, caudas, cornicles, and rostrum. Myzus persicae (Sulzer) Green Peach Aphid (Figure 34) Aphis persicae Suuzer, 1776; Aphis dianthi ScHRANK, 1801; A. persicecola Bots- DUVAL, 1867; Myzus malvae OESTLUND, 1886; Phorodon cynoglossum WILLIAMS, 1910; Rhopalosiphum betae THEOBALD, 1913. 106 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4rH SER. “Crise eo 068s oo aa. PRE GaN ry LD Ne a: Dh LN 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 ot host plants and is the most important aphid vector of plant virus diseases (Essig 1948b). CHILE: On Datura sp., Antofagasta, November 21, 1950—10 apterae. On Cereus sp., Las Palmas, Province of Aconeagua, 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: Abutilon sp., Antirrhinum sp., Bellis sp., Beta sp., Brassica sp., Can- nabis sativa L., Capsicum sp., Chenopodium sp., Cineraria sp., Cosmos sp., Cuminum cyminum L., Cynara sp., Descurainia ap- pendiculata (Gris.), Dianthus sp., Lonicera sp., Malva sp., Pim- pinella anisum L., Prunus spp., Pyrus sp., Senecio sp., Solanum VoL. XXVIII] ESSIG: APHIDAE OF SOUTH AMERICA 107 spp., Tulipa sp., Vinca sp. (Blanchard 1922, pp. 211-213, fig. 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 ., Mendoza Capital, October 3, 1948—4 alatae. On Begonia, Mendoza Capital, March 7, 1949—many specimens. On nettle, ortiga, Urtica wrens, Godoy Cruz, Mendoza Capital, Novem- ber 20, 1950. On Tulipa sp., at Mendoza, October 13, 1950—1 aptera and 1 alate. On Rumer sp., Mendoza, Guaymallen y Capital, November 1, 1950— several specimens On Barnadesia odorato (2?) Mendoza Capital, November 1, 1950—many specimens. Bouivia : On cultivated potato (patata, papa), at Camargo, February 22, 1951 —2 apterae. PERU: On eaper-bush, Capparts 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. By beating nightshade, Solanum sp., Andahuaylas, Province of Apuri- mac, Mareh 7, 1951—6 apterae and 1 alate. On cultivated potatoes, Sicuani, Province of Cuseo, Mareh 2, 1951— 4 alates. On thorny tree by beating, Abaneay, Province of Apurimac, Mareh 6, 1951—1 aptera. By beating, Rio Pampas, Province of Ayacucho, March 8, 1951—3 apterae. 108 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SEr. Tribe Cryptomyzini Capitophorus braggi (Gillette) Artichoke Aphid (Figure 35) Myzus braggti GILLETTE, 1908. This pale yellow and greenish aphid often appears in overwhelming numbers on artichoke and other hosts. CHILE: On artichoke (aleaucil) Cynara scolymus, 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. 8. Ross. ARGENTINA: On artichoke (aleaucil), Cynara scolymus, in the provinees 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, polipodio) (Polypodiaceae) in tropical and subtropical areas. CHILE: A single alate specimen was taken by sweeping 40 km. east of San Carlos, Nuble Province, December 24, 1950, by Dr. A. E. Michel- bacher. ARGENTINA: Blanchard (1939, pp. 41-42) records this species on (Acrostichum), Adiantum, Nephrolepis, and other cultivated ferns in the Province of Buenos Aires. VoL. XXVIII] GiSSIG: APHIDAE OF SOUTH AMERICA 109 Figure 35. Artichoke aphid, Capitophorus braggi (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.) 110 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH Serr. Figure 36. Fern aphid, /diopterus nephrelepides Davis. Alate and apterous females; antennae and cornicles of each; cauda of alate; and body tubercle of ap- terous female. (After Zimmerman, 1948.) Subfamily DACTYNOTINAE 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, pl. 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. VoL. XXVIII] ESSIG: APHIDAE OF SOUTH AMERICA 111 Figure 37. Pea aphid, Acyrthosiphon onobrachis (Fonscolombe) [Macrosiphum pisi (Kaltenbach)]. Alate parthenogenetic 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. 112 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. Aphis pisum M. Harris, 1776; A. onobrychis BoyER DE FONSCOLOMBE, 1841; A. lathyri Mostey, 1841; A. pisi KALTENBACH, 1843; Siphonophora ononis Kocn, 1855; Nectarophora destructor JOHNSON, 1900;Macrosiphum trifolii PERGANDE, 1904; Macrosiphum pisi Parcu, 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. PERU: Sweeping legumes at Mollendo, Province of Arequipa, November 19, 1950—a single apterous female. Subtribe Metopolophiina Aulacorthum pelargonii (Kaltenbach) Geranium Aphid Aphis pelargonii KALTENBACH, 1943; A. malvae WALKER, 1848-9. This species was collected only twice as follows: PERU: 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) Aphis solani KALYENBACH, 1843; Macrosiphum solani TuHroBaLp, 1913, Myzus ve- ronica THEOBALD, 1913; M. hydroctylei TreoBaLp, 1925; M. pseudosolani THEO- BALD, 1926; M. veronicellus THEOBALD, 1926. This aphid is widely distributed and has many host plants, but there were only two collections: CHILE: On a wild umbelliferous plant and by beating various plants at Pur- ranque, Provinee of Osorno, January 15, 1951—1 alate and 7 apterae. On wild potato at the same place on January 26, 1951—1 alate and 8 apterae. VoL. XXVIIT] ESSIG: APHIDAE OF SOUTH AMERICA 113 Figure 38. Solanum or foxglove aphid, Aulacorthum 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. Tribe Macrosiphonini Subtribe Macrosiphonina Macrosiphum® ambrosiae (Thomas) Ambrosia Aphid (Figure 39) Siphonophora ambrosiae THOMAS, 1877. 5. The spelling of the generic name Macrosiphum Passerini, 1860, was corrected to Macrosiphon to conform to the Latin word siphon 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- clature. 14 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SrEr. See (Cy. = OnO-0 Figure 39. Ambrosia aphid, Macrosiphum ambrosiae (Thomas). Alate and ap- terous parthenogenetic females with enlarged antennae, caudas, cornicles and ros- trum. Dark red and black. VoL. XXVIII] ESSIG: APHIDAE OF SOUTH AMERICA 115 This 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 mueh in- vestigational work. CHILE: On Artemisia sp. ?, Papudo, Province of Aconcagua, November 28, 1950—5 apterae. On Baccharis sp. ?, Rivadavia, Provinee 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 —) apterae. On a grindelia-like plant, 40 km. east of San Carlos, Provinee of Nuble, December 23, 1950—11 apterae. On Baccharis sp. ?, Tueapel, 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 Maullin 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 specimens, all apterae. On Artemisia sp. ?, Salta, February 14, 1951—6 apterae. PERU: On Chrysanthemum sp., Botanical Garden, Lima, November 13, 1950 —many apterae. On Encelia sp., Botanical Garden, Lima, November 13, 1950—many alatae and apterae. On tobacco, Nicotiana sp., Botanical Garden, Lima, November 14, 1950—2 alatae. 116 CALIFORNIA ACADEMY OF SCIENCES [Proc. 41H Ser. On Gerbera jamesonu, Botanical Garden, Lima, November 14, 1950— 2 alatae and 12 apterae. On Picris sp. ?, Botanical Garden, Lima, November 13, 1950—many apterae. Macrosiphum bonariensis Blanchard Composite Aphid This fine species was collected on Tanacetum vulgare L. at Flores and Lomas de Zamora, Argentina, by Blanchard (1912, pp. 205-6, fig. 10). It was raised to species status by the same author (1932, pp. 19-21, fig. 10) and more completely deseribed by him from additional specimens collected on eultivated Pyrethrum, Buenos Aires; and again on Tanacetum vulgare L., from the Provinee of Buenos Aires y Cordoba, Argentina (1939, pp. 953-55). CHILE: A fine series of this species was collected by the Michelbachers on a member of the Family Composite near San Carlos, Province of Nuble, December 23, 1950. It appears to fall in the genus Pharalis Leach accord- ing to the elassifieation by Lambers (1929). Macrosiphum chilensis Hssig, new species Chilean Bacecharis 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 gibbous and much larger than II; IIT 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 ; VIT 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 see- ond eoxae. Wines as drawn. Lees 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 ambrosiae (Thomas). Cornicles eylindrieal, 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 VoL. XXVIII] ESSIG: APHIDAE OF SOUTH AMERICA 117 Figure 40. Chilean baccharis aphid, Macrosiphum 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. solanifolia (Ashmead), but the constriction is not as noticeable; dusky except the basal one fourth. The anal plate is very small. Cauda 0.8 mm. in length and tapering as shown. Length of body 3.00 mm.; antennae 0.33 mm.; forewing 4 mm.; cornicles 0.8 mm. ; eauda 0.5 mm. Apterous 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 mm.; width 1.6 mm.; antennae 3.5 mm.; cornicles 0.8 mm.; cauda 0.6 mm. This species somewhat resembles VW. solanifolii (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. (2), at La Serena, Province of Coquimbo, Chile, December 8, 1950, by Dr. A. E. Michelbacher. 118 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. In all, 4 alates, 25 apterae, and 5 immature individuals as well as one east skin were taken. These are mounted on 5 aluminum microscope slides. The alate type is mounted on a slide with 4 mature apterous paratypes. Macrosiphum cordobensis Blanchard Cordobens Aphid (Figure 41) Figure 41. Cordobens aphid, Macrosiphum cordobensis Blanchard. Upper—aA, antenna, cornicle and cauda of alate parthenogenetic female: B, antenna, cornicle cauda, and rostrum of aptera. ARGENTINA : On Baccharis sp. (2), at 4,000 feet on a ridge west of Tucuman, Feb- ruary 11, 1951—many apterae and alatae. On EFrigeron sp., Sierras de Cordobe, February 18, 1931. Original de- seription. (Blanchard, 1932, pp. 24-86, 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 body, slender, with short knobbed hairs, and segment ITI with 15-17 large and small circular secon- dary sensoria arranged in a row along the underside throughout the full VoL. XXVIII] ESSIG: APHIDAE OF SOUTH AMERICA 119 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, ete.) 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 lone. The cornicles are black apically, long and slender, shehtly 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 differs from most members of the genus. The apical flaring is some- what after the pattern of the pea aphid, Acyrthostphon onobrachis (Boyer de Fonscolombe) (Macrosiphum), but in the pea aphid the reticulations are absent; length 0.7 mm. The cauda is rather robust, flaring at base and then slightly constrieted ; 0.85 mm. in length, with six hairs. The anal plate is very small and erescent-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 IIT 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, ete. The cauda is more robust 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 what appears to be a nettle (ortiga), Urtica sp., at Rio Pampas, Peru, March 8, 1951, by Dr. Michel- bacher. Type: An alate form mounted on a slide with 4 apterous females. The 120 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4rH SER. remaining 6 alates and all the 24 apterous forms are mounted on 6 other slides and are designated as paratypes. This species differs 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 (aleachofa, arcacil, aguaturma), Cynara scolymus L., Jujuy and Buenos Aires, in 1924, and on Vernonia sp., Sierra de Cordoba. CHILE: 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 Malleco, January 29, 1951. Macrosiphum huantana Essig, new species Huanta Aphid (Figure 43) 0 OO 833096 26 Ge *o CS) Figure 43. Huanta aphid, Macrosiphum 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 VoL. XXVIII] ESSIG: APHIDAE OF SOUTH AMERICA 121 knobbed hairs: secondary sensoria on segment III variable in size and seat- tered over the basal half or two thirds; varying in number from 17 to 19. (In paratypes they vary in number after this pattern: 13-13, 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 mm. long; clear; venation as drawn. Legs slen- der. Cornicles dark, long and cylindrical; widening basally and apieally ; 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 eauda, with many long spines. Apterous parthenogenetic female: Dark in color with all appendages brownish in cleared specimens; without definite pigmented areas. Secon- dary sensoria on antennal segment ITI 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 (Guaita), Province of Coquimbo, Chile, December 7, 1950, by Dr. A. E. Michelbacher. This species is close to M. littoralis Blanchard, but has more sensoria and hairs on rostrum and eauda 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: 3 alatae and 27 apterae. The type specimen 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 lizerianum Blanchard Lizer Aphid A single apterous female of what appears to be this aphid 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 glauca, and species of Com- posite at Punta Chiea, San Isidro, Flores and Canuelas, Province of Buenos Aires, and at Potrerillos and Cachueta, Province of Mendoza. He states it is close to M. solidaginis (Fabricius). 122 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SEr. Macrosiphum macolai Blanchard Macola Aphid (Figure 44) Figure 44. Macaloi aphid, Macrosiphum macaloi 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. 29-80, fig. 6. Original description: 1939, 966-7.) On Erigeron sp. (echilea), Provinee of Mendoza, by Blanchard (1944, p. 42). CHILE: On the common dandelion, Taraxicum vulgare, at El Abanico, Provinee of Bio-Bio, December 30, 1950—1 aptera. On Baccharis sp. ?, at Negrete, Province of Bio-Bio, January 30, 195] —27 apterae. Macrosiphum muermosa Essig, new species Muermos Aphid (Figure 45) VoL. XXVIII] ESSIG: APHIDAE OF SOUTH AMERICA 123 ve as ot os RIAA eA ee | Figure 45. Muermos aphid, Macrosiphum muermosa 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. Tyre: Alate parthenogenetic female: Probably deep red, brown, and black—the aleoholic preserving fluid 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; approximately the length of the body and extending to the tip ot the eauda; dark to black in color; secondary sensoria only on segment ITT, eireular to oval, rather small, quite 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 inconspicuous hairs. Legs slender; hind tibiae strongly curved; tarsi very small. Cornicles black; wider at base, nearly eylindrieal, shightly eurved outwardly ; apical one fourth plainly reticulated; 1 mm. in length. Cauda 124 CALIFORNIA ACADEMY OF SCIENCES [Proc. 47TH SER. black, long, and well developed; with many hairs; 0.5 to 0.7 mm. long. Anal plate small and erescent shaped; with few rather long hairs. Length of body 4 mm.; antennae 4.2 mm.; forewing 4 mm.; cornicles 0.75 mm.; eauda 0.50 mm. Apterous parthenogenetic 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, ete. 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 oceur in Macrosiphum ambrosiae (Thomas), Dactynotus picridis (Fabricius), D. taraxact (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 both 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 WM. lizerianum Blanchard, but differs in being larger; darker; with many more sensoria on antennal segment IIT 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: Apterous parthenogenetic 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 beset with rows of short spines arising from basal cones and eurved (drooping) and knobbed. The an- tennae are long and slender with 13 to 16 small cireular 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] ESSIG: APHIDAE OF SOUTH AMERICA 125 Figure 46. Nuble aphid, Macrosiphum nuble 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; evlindrical 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. Leneth of body 3.30 mm. Host plant and locality: The species was collected on a composite plant in the Andes Mountains near Fabian, Province of Nuble, Chile, Decem- ber 15, 1950, by Dr. A. E. Michelbacher. The specimens consist of seven apterous females which are mounted on three slides. One individual is indicated as the type. All the others are designated as paratypes. No alates were collected. 126 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4TH SER. This species resembles Macrosiphum solanifoli (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 LINNAEUS, 1758; A. scabiosae Scopout, 1763; A. dipsaci SCHRANK, 1801; Siphonophora 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 Camellia japonica, Dip- sacus fullonum, D. sylvestris, Rudbeckia lirta, 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 apterae. On Sonchus, sp., Valparaiso, November 26, 1950—1 alate and 17 apterae. On cultivated rose, Vieuna, Province of Coquimbo, December 4, 1950 —apterae and alatae. On beans, Phaseolus sp., San Carlos, Province of Nuble, December 23, 1950—1 alate—casual ?. On rose, Temuco, Province of Cautin, January 6, 1951—apterae and alatae. 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. VoL. XXVIIT} ESSIG: APHIDAE OF SOUTH AMERICA 127 G use HA B. oO. Keer, Figure 47. Rose aphid, Macrosiphum rosae (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. 128 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4rH SER. \3 r z mae y ¥ OP ee A Y / en 2 Gy e@aec Iesetos SB SAS ROS a Om : = ee Z a = == VA aaa ——— by ———— a Cae. ae = ——— Z Op >= a = : SS nly Zo PA Se ee 2 ZZ a “ Z ZA gee Te) ig Figure 48. Potato aphid, Macrosiphum solanifolii (Ashmead). A, apterous par- thenogenetic female with enlarged: a, cornicle; b, dorsum of rostrum; ¢, 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, Chiclayo, Province of Lima, March 21, 1951—many specimens; all apterae. ARGENTINA: Common on cultivated roses throughout Argentina and also on loquat (nispero) Eriobotrya japonica. (Blanchard 1922, pp. 187-190, fig. 2; 1939, pp. 967-9). Macrosiphum solanifolii (Ashmead) Potato Aphid (Figures 48—49) Siphonophora solanifolii ASHMEAD, 1882; S. euphorbiae THoMas, 1878; Macrosi- phum cucurbitae pet GuERCIO, 1913; Macrosiphum gei (Koch) Horres ANnpD Frison, 1931; Macrosiphon solanifolii (Ashmead) BO6OrRNER, 1952. This is a very widely distributed Holaretie species and appears to Vor. XXVIII] ESSIG: APHIDAE OF SOUTH AMERICA 129 \\ \\ Ny \ , 2A ee Oe ais SS = re SS aera) i wWWOS5 oo s ——— \ 7 ae) 14 ( ) ; Mae. Soa ee ¥ \ 3 Q< = S :