bola a, Pye ng geet gy i ha Nt <= ae ee ated rei ee ee eee ee anaes centre te EE et Oe ener ws 0 peta ena cpearge ioe ara ae) ee Ge ae oes eee barnett Sr ees be A oP a oe ie es a Ee A Mees oS ot Cita Tie —_ iy oy (eireecs | ar aah tes, Te 4: 96, pled i eins ie o hee UE ee hs +s OR a art at pe ete j &* is A ven ae Ay i 7 ' } ct - he epee | - | : a 1 ae ee | _ » ee hae ir | ae ; a or if 1 ; a elsNind a a Na sc it Jah nae ry 7 oe ue FG ails i ae SMITHSONIAN MISCELLANEOUS COLLECTIONS VOR. 159 ©cecccese®® “EVERY MAN IS A VALUABLE MEMBER OF SOCIETY WHO, BY HIS OBSERVATIONS, RESEARCHES, AND EXPERIMENTS, PROCURES KNOWLEDGE FOR MEN”’—JAMES SMITHSON (PusiicaTion 4468) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION 1961 PORT CITY PRESS, INC. BALTIMORE, MD., U. S. A. ZA HSOWgs AUG 28 1961 LIBRARY ADVERTISEMENT The Smithsonian Miscellaneous Collections series contains, since the suspension in 1916 of the Smithsonian Contributions to Knowledge, all the publications issued directly by the Institution except the An- nual Report and occasional publications of a special nature. As the name of the series implies, its scope is not limited, and the volumes thus far issued relate to nearly every branch of science. Papers in the fields of biology, geology, anthropology, and astrophysics have predominated. LEONARD CARMICHAEL, Secretary, Smithsonian Institution. (iii) ay ai ie ; al ve Fite ee ee A ; re / i ; K TO. BY. CONTENTS PEABoby, FRANK E. The oldest known reptile, Eosauravus copei Williston. 13 pp., I pl., 3 figs. May 7, 1959. (Publ. 4377.) WETMORE, ALEXANDER. The birds of Isla Escudo de Veraguas, Panama. 27 pp., I pl., 3 figs. July 8, 1959. (Publ. 4378.) GREGOIRE, CHARLES. Further observations on distribution of pat- terns of coagulation of the hemolymph in Neotropical insects. 23 pp. Aug. 18, 1959. (Publ. 4379.) Hanp ey, Cuartes O., Jr. A review of the genus Hoplomys (thick-spined rats), with description of a new form from Isla Escudo de Veraguas, Panama. I0 pp., 1 fig. July 3, 1959. (Publ. 4380.) Cooper, G. ArTHUR. Genera of Tertiary and Recent rhynchonel- loid brachiopods. 9o pp., 22 pls., 1 fig. Nov. 23, 1959. (Publ. 4382.) BoarDMAN, RicHarp S. A revision of the Silurian bryozoan genus Trematopora. 14 pp., 2 pls., 1 fig. Oct. 29, 1959. (Publ. 4383.) Gazin, C. Lewis. Early Tertiary Apheliscus and Phenacodaptes as pantolestid insectivores. 7 pp., 2 pls. Aug. 12, 1959. (Publ. 4385.) Snoperass, R. E. The anatomical life of the mosquito. 87 pp., 30 figs. Nov. 4, 1959. (Publ. 4388.) AssotT, C. G. A long-range forecast of United States precipita- tion. 78 pp., 12 figs., g charts. Mar. 23, 1960. (Publ. 4390.) WILLIAMS, JEROME; JOHNSON, E. R. FENIMorE; and Dyer, AL- BERT C. Water transparency observations along the east coast of North America. 181 pp., 2 pls., 13 figs. Oct. 26, 1960. (Publ. 4391.) WETMoRE, ALEXANDER. A classification for the birds of the world. 37 pp. June 23, 1960. (Publ. 4417.) (v) SMITHSONIAN SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 139, NUMBER 1 Charles D. and Mary Waux Walcott Research Fund THE OLDEST KNOWN REPTILE, EFOSAURAVUS COPEI WILLISTON (WitH 1 Prate) By FRANK E. PEABODY Department of Zoology University of California Los Angeles, Calif. (PuBLicaTIon 4377) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION MAY 7, 1959 yy EXON UDA hp ry Pot oy SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 139, NUMBER 1 Charles D. and Mary Waux Walcott Research Fund THE OLDEST KNOWN REPTILE, EOSAURAVUS COPE! WILLISTON (W1TH 1 PLATE) By FRANK E. PEABODY Department of Zoology University of California Los Angeles, Calif. (PusiicaTion 4377) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION May 7, 1959 SMITHSONIAN INSTITUTION MAY 7 THE LORD BALTIMORE PRESS, INC. BALTIMORE, MD., U. S. A. FOREWORD Dr. Frank E. Peabody died on June 27, 1958, leaving several manu- scripts in various stages of completion. The one published here, based on a specimen in the collections of the United States National Mu- seum, was complete except for final touching up of some of the il- lustrations ; these illustrations have been finished by Miss Madeline M. Peabody with the help of Dr. Theodore H. Eaton, Jr. The manuscript has been edited by Dr. Eaton and myself. Dr. Peabody’s paper presents a welcome clarification of the relationships of an important, and here- tofore much misunderstood, early reptile. PETER P. VAUGHN i a Aire aim Charles D. and Mary Vaux Walcott Research Fund THE OLDEST KNOWN. REPTILE: BOSAURAVUS COPED WILLISTON By FRANK E. PEABODY Department of Zoology University of California Los Angeles, Calif. (WitH ONE PLATE) One of the most tantalizing examples of Carboniferous tetrapods is the posterior part of a small skeleton from Linton, Ohio, described by Cope in 1897 as the earliest known reptile. Some 60 years later, and after many taxonomic vicissitudes, the specimen seems in danger of slipping into obscurity among the microsaur Amphibia. Meanwhile no more reptiles have been found at Linton or in earlier horizons.* Various students have described Cope’s specimen, but most have tended to discount its importance because the anterior part of the skeleton, including the skull, is missing, and have tended to accept the early descriptions with little question. Present high interest in the origin of reptiles during the Carboniferous prompted a restudy of Cope’s historic specimen. It was found that strong lighting from a very low angle, and directed from various positions, revealed much new detail that can be demonstrated by photographic enlargements. The result is a new interpretation, particularly of the vertebrae and tarsus, which reaffirms the reptilian affinities of the specimen and furthermore strongly suggests a captorhinomorph relationship. I am indebted to Dr. Peter P. Vaughn of the United States National Museum for permission to borrow Cope’s specimen, and to Miss Madeline M. Peabody, my sister, for assistance with the illustrations. 1[Cephalerpeton ventriarmatum, apparently a captorhinomorph reptile (see Gregory, 1950), is known from the nodule beds at Mazon Creek, IIl., which represent a somewhat earlier horizon (but still within the Allegheny series). —Ep.] SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 139, NO. 1 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 SYSTEMATIC DESCRIPTION EOSAURAVUS COPEI Williston PLATE I; TEXT FIGURES I-3 Isodectes punctulatus Corr, Proc. Amer. Philos. Soc., vol. 36, pp. 88-90, pl. 3, fig. 3, 1897; WuiLutston, Journ. Geol., vol. 16, pp. 395-400, text fig. 1, pl. 1, 1908; Moonie, Proc. U. S. Nat. Mus., vol. 37, pp. 11-16, pls. 4-5, 1909. Eosauravus cofei WILuisTon, Bull. Geol. Soc. Amer., vol. 21, p. 272, 1910; CASE, Carnegie Inst. Washington Publ. 145, pp. 31-32, text fig. 8, IQII. Tuditanus punctulatus Romer, Bull. Amer. Mus. Nat. Hist., vol. 59, pp. 134-135, 1930; Bull. Mus. Comp. Zool., Harvard Coll., vol. 99, p. 300, 1947; Amer. Journ. Sci., vol. 248, p. 641, 1950; HuENE, Palaontologie und Phylogenie der niederen Tetrapoden, p. 163, 1956. ?Tuditanus Romer, Osteology of the reptiles, p. 483, 1956. Type—uvU.S.N.M. No. 4457; posterior $ of a reptilian skeleton pre- served belly-down on a slab of coal from Linton, Ohio. Horizon.—Allegheny group, Middle Pennsylvanian (Westphalian). Diagnosis.—Small reptile with a minimum of 28 presacral vertebrae of generally captorhinid structure, with broad, swollen neural arches, low neural spines, zygapophyseal facets in horizontal plane, and small intercentra; free ribs on all vertebrae except distal caudals; distal caudal vertebrae with low neural arches and probably without haemal spines, centra occasionally fused forming relatively stiffened axis; one principal and one accessory sacral rib; hind limb with prominent internal trochanter, with relatively short epipodial (=zeugopodial) segment having relatively massive fibula; primitive, well-ossified tarsus of basic captorhinid or pelycosaurian plan with separate median and lateral centrale and with a 6th distal tarsal (=postminimus) ; phalangeal formula 2-3-4-5-4, terminal phalanges blunt-ended. No gastralia present; possibly with body scales, having striae radiating from anterior margin of scale. No obvious aquatic adaptations of well-ossified skeleton. Anterior skeleton unknown. Taxonomic notes.—The taxonomic history of Cope’s specimen is so devious and confusing that a short explanation is necessary to sup- plement the synonymy listed above. Cope (1897) described the posterior skeleton and believed it to be conspecific with another small vertebrate represented by a skull and anterior two-thirds of a skeleton. The latter had been described by Cope (1874, p. 271) as Tuditanus punctulatus, but in his 1897 paper, it was referred along with the posterior skeleton to the genus Jsodectes. Williston (1908) and Moodie (1909) offered new descriptions of the posterior skeleton, treating it as distinct from the anterior skeleton, but tending to over- look the fact that the anterior skeleton is the type of Jsodectes NO. I THE OLDEST KNOWN REPTILE—PEABODY 3 punctulatus. (Moodie’s plate description (p. 28) in fact refers to the posterior skeleton as “the type specimen of Isodectes punctulatus,” which, of course, it is not.) Later, Williston (1910) and then Case (1911) established the posterior skeleton as a new genus and species, Eosauravus cope: Williston. Unfortunately, the European genus Sauravus to which Williston related the posterior skeleton is clearly an amphibian with nectridian vertebrae, so the name Eosauravus is inappropriate morphologically but remains valid taxonomically. Romer (1930) restudied the Linton fauna and, in a commendable attempt to reduce the large number of artificial species, referred Cope’s posterior skeleton again to the anterior skeleton now designated as Tuditanus punctulatus. The synonymy of Tuditanus with Isodectes had proved to be wrong since the latter genus now appears to be a captorhinomorph (Gregory et al., 1956, p. 2), and the former genus is a microsaur. Romer’s decision apparently rested mainly on the im- probability that there might be more than one reptile at Linton, and that there was the distinct possibility that the smaller, less ossified anterior skeleton merely represents a more immature individual than the posterior skeleton. The two specimens were regarded by Romer as reptilian with no recognizable ordinal characters. Later, Romer (1947, p. 300) suggested that the two specimens together represent either a seymouriamorph or cotylosaur on the basis of a stemmed in- terclavicle, seemingly broad-arched vertebrae, and a pes with a pha- langeal formula 2-3-4-5-4. Still later, Romer (1950, p. 641) dis- counted the importance of the stemmed interclavicle and phalangeal formula, and, while noting a presumed high presacral count of verte- brae, long, slender body proportions, apparent lack of caudal chevrons, and long postorbital region of the skull, concluded that Tuditanus punctulatus (based on anterior and posterior skeletons) “is not im- probably a microsaur.” This conclusion, undoubtedly influenced by increased understanding of microsaurs, was followed by both Piveteau (1955) and Huene (1956) in their valuable compendia of vertebrate paleontology. Meanwhile, Romer (1956, p. 483) apparently turned once more toward Williston’s opinion of the posterior skeleton as shown by the lone entry “[ Reptilia] Incertae sedis. ?Seymouriamor- pha. ?Tuditanus Cope 1874 (Eosauravus Williston 1910).” Thus at present, the posterior skeleton designated as Eosauravus copei by Williston, is in an obscure position both taxonomically and phyloge- netically. The anterior skeleton is best considered a probable micro- saur amphibian under the designation Tuditanus punctulatus. In any | case it is difficult to demonstrate distinctive reptilian characteristics in 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 T. punctulatus, and especially difficult to demonstrate any real affini- ties with Eosauravus copet. Description —The specimen consists of the posterior two-thirds of a postcranial skeleton preserved belly-down on a coal stratum. Neither the opposing slab, probably containing a dozen thoracic vertebrae and caudal neural arches, nor adjoining blocks of matrix containing the tip of the tail, some terminal phalanges of the left pes, and the anterior end of the skeleton, were collected. The remaining parts of the skele- ton have undergone very little deterioration since Cope’s time, judging from the excellent photograph presented by Williston (1908) and republished (with inaccurate retouchings) by Moodie (1909). The presacral, sacral, and anterior caudal vertebrae lie on their right sides (as observed by Cope, 1897) in such a manner as to cover the proximal tips of the right ribs while the proximal ends of the left pre- sacral ribs are pressed against the upper (left) surfaces of their cor- responding centra. The outline of successive neural spines is clearly visible on the right side between successive ribs. The caudal vertebrae posterior to the rib-bearing caudals are preserved with ventral side down and have lost their neural arches, thus exposing the neural canal as a longitudinal groove in the dorsal surface of the centra. Un- fortunately, Moodie (1909, pl. 5) illustrated the entire column as though it were oriented with the dorsal side uppermost (figure repro- duced by Case, 1911, fig. 8). The result is an erroneous picture of the vertebrae from anterior caudals forward. Cope’s illustration (1897, pl. 3) shows the correct orientation, but is only slightly suggestive of the true form of the vertebrae. The true form of the presacral and anterior caudal vertebrae may be reconstructed with reasonable accuracy from a composite of details exhibited along the column. Specifically the impression of the anterior presacrals clearly shows the contour of the centrum; the first 5 pre- sacrals and anterior caudals preserve details of swollen neural arches as well as of the centra and intercentra. The position of intervertebral foramina is clearly indicated by a series of circular pits. Figure I is presented as a reconstruction based on composite detail. There seems to be little doubt that the neural arch is low and broad as mentioned by Romer (1947, p. 300), has a low spine, and has a perceptible swelling above the posterior zygapophysis ; also that small intercentra are present. The latter are indicated between the first several presacral centra, between the 1st and 2d caudal centra, and by a haemal wedge between the 3d and 4th caudal centra. In the pre- sacral series the left ribs appear to have been crushed precisely against THE OLDEST KNOWN REPTILE—PEABODY I NO. "(o1) esyUadIOJUT ‘(4IS) Sqit [eroes ‘(su) outds yesnou ‘(9S) g]qissog ‘“wowmdads ad4} Jo Jrejop 9}1sodui09 WoIF pozONns}sUOIOI ‘1agoI SnapinDsoy JO UoTseI sayeos Apoq Jepnes Josajue pue ‘eroes ‘sequiny—'I ‘o1,f 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 the intervertebral area; thus the supposed intercentra here may be parts of the ribs. However, there appears to be a distinct intercentral space coincidental with the position of the ribs, and, in any case, the evidence for intercentra in the anterior caudals is clear and unobscured by ribs. Evidence of 24 presacral ribs on the left side is fairly clear, although the first presacral is difficult to see, and only the distal tip of the 24th presacral is preserved on the edge of the slab. The general pattern and number of the presacral ribs and the far anterior position of the left manus have led to the opinion (Romer, 1950, p. 641) that the presacral count is significantly higher than the 25-27 vertebrae usually found in the most primitive reptiles. However, the 5 most anterior ribs are clearly more massive than following ribs, and the distal ends are slightly spatulate—all indicative of an extreme anterior thoracic position related to serrati muscles of the pectoral girdle. Also, the successive positions of the distal ends of the 5 anterior ribs suggest a progressive shortening in a forward direction as might be expected in a smooth transition to the cervical region. Accordingly, a recon- struction will show that the total number of presacrals may have been as few as 28. The forward position of the left manus as an indicator of a far anterior position of the pectoral girdle is probably misleading. The girdle probably shifted forward or to the right side away from its life position lateral to the 5 anterior ribs. Rib heads are obscured in the presacral series generally, but the 3d to 8th left presacral ribs appear to have a proximal expansion com- mensurate with the elongate diapophysis of the neural arch. Certainly these ribs are not single-headed as in lizards, but bear a general re- semblance to captorhinid ribs. The pelvic girdle and sacral vertebrae are distorted beyond certain recognition of salient features, although the spacing of vertebral seg- ments and disposition of lumbar and caudal ribs suggest the presence of two sacral vertebrae. A short, thick element lying across the ad- ductor fossa of the right femur may be a right second sacral rib; an obscure spatulate structure immediately anterior to the anteriormost left caudal rib may be the first or principal sacral rib. Except for a general outline of the acetabular regions of the girdle, little can be demonstrated here except that the mass lying between the heads of the femora probably constitutes a pelvic girdle and sacrum of primitive reptilian plan. According to my interpretation, Moodie (1909, pl. 5) included the internal trochanter of the left femur in his outline of the left acetabular region, thus giving the left pelvis a more NOL THE OLDEST KNOWN REPTILE—PEABODY 7. distinct outline than is warranted. A thin plate lying anterior to the head of the right femur may represent the left ilium broken over to the right. Although the thin plate may be regarded as a patch of over- lying matrix such as obscures the centrale of the left tarsus (see be- low), there is a definite anterior border that looks much like the an- terior edge of an iliac blade. Nowhere is there evidence of a long posterior process of the ilium like that of Eogyrinus. The anterior 4 or 5 caudal vertebrae are associated with 3 pairs of sharply curved ribs. In addition, there are short structures faintly shown on the left side that are not curved and probably represent short haemal spines nearly in the correct position. Also, there is a distinct haemal wedge between the 3d and 4th caudal centra. Certainly there is enough evidence to question seriously earlier observations (Cope, 1897, p. 89; Romer, 1950, p. 641) that there are no haemal spines in the tail. The caudal series becomes twisted, possibly 180 degrees, at the posi- tion of the 7th vertebra, which appears to be lying on its left side. Posteriorly the series is oriented with ventral side down—an unusual position if neural and haemal arches were at all well developed here, or if there was any lateral compression of the centra. Under these conditions the vertebrae would be almost certainly lying on one side or the other as in the anterior column. However, the caudal centra appear broader than high, and occasional fusion of neighboring centra seems to have occurred. All features of the tail, including the orienta- tion, suggest some specialized function—perhaps a prehensile action in the dorsoventral plane. A special aquatic function does not seem possible, insofar as a lateral sculling motion is concerned, although the fused vertebrae may suggest a stiffened axis serving as the founda- tion for a rudder. Part of the left manus (omitted in Cope’s figure, 1897, pl. 3) lies disarticulated near the anterior end of the vertebral column. Enough is shown to indicate that the carpus was definitely as fully ossified as the tarsus, and less surely that the phalangeal formula was comparable to the reptilian count in the pes. Both limbs are complete except for the loss of some terminal phalanges on the left side. The left femur is preserved with the dorsal surface uppermost—the right femur with the ventral surface upper- most. Thus the whole contour of the bone can be recognized in com- posite. The femoral head, internal trochanter, adductor fossa, and distally the tibial and fibular condyles resemble those of primitive reptiles such as ophiacodonts and captorhinids. The trochanter is 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 especially prominent and extends proximally nearly in line with the femoral head. Ossification is fully developed in the femur as well as in the more distal elements. The tibia and fibula are short, stout bones of generally primitive contour ; the fibula appears relatively more massive than is usually the case in the tetrapod limb. The distal end of the left tibia appears to have slipped slightly upward from the life position and now rests on the neck of the astragalus. Otherwise, the left femur, tibia, and fibula are in normal articulation. The right pes is twisted so as to obscure details of the tarsus, but details of the digits help to complete a restoration of the left pes. The left pes is preserved dorsal side uppermost and exhibits one of the most perfect preservations of tarsal structure known from the Car- boniferous, indeed, from the Paleozoic, as will be demonstrated pres- ently. The pes has been given several superficial descriptions (Cope, 1897; Williston, 1908; Moodie, 1909) which fail to recognize the ex- tent of ossification in the tarsus, but nevertheless establish two proxi- mal elements in the tarsus and a phalangeal formula of 2-3-4-5-4. A main difficulty lies in the interpretation of tarsal elements distal to the presumed astragalus and calcaneum, especially in the medial region of the tarsus where no one has been able to recognize central elements. Moodie’s figure (1909, pl. 5), republished by Case (1911), is particu- larly misleading in that the tarsus appears to have an enigmatic pat- tern, doubtfully reptilian. (Also, in Moodie’s figure a nonexistent element is added distal to the lateralmost distal tarsal, although none is shown in Moodie’s retouched photograph—his pl. 4). My photo- graphs (pl. 1A, B), taken under low-angle light from first one direc- tion and then from the opposite direction, demonstrate the wealth of detail making possible text figure 2. The two proximal bones of the tarsus are clearly the astragalus and calcaneum which enclose between them a perforating foramen, not previously noted. The astragalus has a small but definite tibial facet directed mostly preaxially. There is no evidence of tripartite structure such as exhibited by Captorhinus (Peabody, 1951). The preaxial border between the astragalus and the first metatarsal clearly exhibits two bones that must be a median cen- trale and distal tarsal 1. A thin veneer of matrix obscures part of the dorsal surface of these bones, but the oblique lighting (pl. 1A) clearly brings out their contours in the preaxial border. Lateral to these bones and median to the large distal tarsal are at least 2 and probably 3 separate bones that are identifiable as the lateral centrale and distal tarsals 2 and 3. A slight proximal jamming (see fig. 2) has NO. I THE OLDEST KNOWN REPTILE—PEABODY 9 forced distal tarsals 2 and 3 slightly out of position. The existence of two separate centralia seems certain although the separation between the lateral centrale and distal tarsal 2 is not clear—probably because of a slightly overriding relationship due to jamming. A unique feature of the tarsus is a postminimus or distal tarsal 6 in the postaxial border. Such an element is unknown in reptiles but is found in the tarsal pat- tern of the urodele, Salamandrella, by Holmgren (1933, p. 217). Fic. 2—Left pes of Eosauravus copei showing primitive reptilian pattern with separate median and lateral centrale, and with unique postminimus or distal tarsal 6 on postaxial border. There is no doubt that the tarsal pattern is generally comparable to primitive captorhinids and pelycosaurs. The metatarsals are all well developed as indicated in figure 2. No special features seem to be present except for a generally robust os- sification (like that of more proximal bones) that contrasts markedly with a seemingly delicate ossification of the phalanges. The phalanges may be confidently restored with a 2-3-4-5-4 formula, using the evidence from both feet. The terminal phalanges are not acutely pointed and cannot be considered as definitely bearing claws. The relative length of the 5th digit suggests no obvious aquatic adap- tation—in the obviously aquatic Mesosaurus, the 5th digit is longer than the 4th. This condition may also be noted in nothosaurs. 10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 Fic. 3—Left limb of Eosawravus copei reconstructed in fully extended position. NO. I THE OLDEST KNOWN REPTILE—PEABODY II The surface surrounding the skeleton seems devoid of structures re- sembling gastralia as indicated by early descriptions. However, a problematical object that may be an unidentified bone from the an- terior skeleton lies just to the left of the distal tail. Possibly of more importance, an enlarged view of the surfaces near the skeleton reveals a number of delicate, ovoid areas with fine striae radiating from a point near one border. The striated areas occur only close to the skeleton ; an example of the striated areas can be seen clearly between the right ribs in plate 1C. It is possible that these striated areas repre- sent body scales developed from the epidermis of Eosauravus. No bone is indicated in the delicate impressions Conclusions.—It is concluded that Cope’s historic specimen from Linton, Ohio, is surely a reptile that has evolved beyond the sey- mouriamorph level. The broad-arched, cotylosaurian vertebrae pos- sess small intercentra, and the narrow space between successive pleuro- centra is in decided contrast with the wide, unossified gap seen in seymouriamorphs. Here, the pedicel of the neural arch has a marked overhang above the intercentral gap. The tarsus has a characteristic reptilian astragalus and calcaneum, with enclosed perforating foramen in the usual position. The astragalus is fully developed with no indication of a compound origin as in the relict Captorhinus aguti of Early Permian age (Peabody, 1951). The whole structure of the pes is of basic reptilian pattern except for the 6th distal tarsal or post- minimus. The latter may be considered an amphibian feature rather than a supernumerary element that widens the pes surface in correla- tion with aquatic adaptations—an untenable point of view considering the general lack of characteristics suggesting aquatic habits of Eosauravus. The combination of vertebral and tarsal characteristics is consonant with other features of the skeleton; together they strengthen the evidence that the astragalar bone, originating from a fusion of tibiale, intermedium, and proximal centrale of the amphibian foot, may be regarded as a reliable osteological indication of the at- tainment of the amniote level of organization—at least until conflict- ing evidence is found. If it be granted that Eosauravus is a reptile, there is a question as to its subgroup affiliation. Current evidence strongly suggests that early ophiacodont pelycosaurs and captorhinomorphs are very close to the root of the reptilian stock. The tarsus of Eosauravus is exceed- ingly primitive in the possession of separate median and lateral cen- trale, and of the postminimus. Only early pelycosaurs have separate centralia—they are fused in Captorhinus and Limnoscelis. No reptiles I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 presently known have a postminimus. The nature of the vertebrae of Eosauravus would indicate that its affinities probably lie with the cap- torhinomorphs. No pelycosaur is presently known to possess vertebrae of a pure cotylosaur type such as is evident in Eosauravus. In view of the primitive pattern of the tarsus, a position near the base of the captorhinomorphs is indicated. Establishment of a true reptile of captorhinomorph affinities deep in the Middle Pennsylvanian helps to clear away some of the uncer- tainty surrounding the time of origin of reptiles. The varied reptiles found in the Upper Pennsylvanian of Kansas (Peabody, 1954) and more fragmentary remains from elsewhere indicate that the evolution of pelycosaurs and captorhinomorphs (if petrolacosaurs be considered an offshoot of the captorhinomorphs as suggested by Vaughn, 1955, p. 446) was well advanced. Eosauravus appears to have been at an evolutionary stage which could be ancestral to any known later reptile. The particular adaptations of Eosauravus to life in a coal swamp are difficult to assess. Moodie (1909, p. 12) suggests that the reptile was aquatic or semi-aquatic mainly on the basis of an “expanded foot” similar to the broad foot of the obviously aquatic mesosaurs. However, the foot of Eosauravus and the rest of the preserved skele- ton have little to suggest even semi-aquatic habits, but do allow the possibility that this small reptile spent most of its time in the “upper story” of the coal forest at Linton. REFERENCES Case, E. C. 1911. A revision of the Cotylosauria of North America. Carnegie Inst. Washington Publ. 145, pp. 1-122, 52 figs., 14 pls. Corr, E. D. 1874. Supplement to the extinct Batrachia and Reptilia of North America. I. Catalogue of the air-breathing Vertebrata from the Coal-measures of Linton, Ohio. Trans. Amer. Philos. Soc., vol. 15, pp. 261-278. 1896. The Paleozoic reptilian order Cotylosauria. Amer. Nat., 1896, pp. 301- 304, 1 pl. 1897. On new Paleozoic Vertebrata from Illinois, Ohio and Pennsylvania. Proc. Amer. Philos. Soc., vol. 36, pp. 71-91, 3 pls. Grecory, J. T. 1950. Tetrapods of the Pennsylvanian nodules from Mazon Creek, Illinois. Amer. Journ. Sci., vol. 248, pp. 833-873, 11 figs. Grecory, J. T., PEAzopy, F. E., and Price, L. I. 1956. Revision of the Gymnarthridae, American Permian microsaurs. Pea- body Mus. Nat. Hist., Bull. 10, pp. 1-77, 33 figs., 1 pl. Hotmcren, NILs. 1933. On the origin of the tetrapod limb. Acta Zool., Stockholm, vol. 14, pp. 185-295, 106 figs. NO. I THE OLDEST KNOWN REPTILE—PEABODY nS HUuENnE, F. von. 1956. Palaontologie und Phylogenie der niederen Tetrapoden. xii + 716 pp., 690 figs. Jena. Moopir, Roy L. 1909. Carboniferous air-breathing vertebrates of the United States National Museum. Proc. U. S. Nat. Mus., vol. 37, pp. 11-28, 7 pls. 1916. The Coal Measures Amphibia of North America. Carnegie Inst. Washington Publ. 238, x + 222 pp., 43 figs., 26 pls. Peapopy, F. E. 1951. The origin of the astragalus of reptiles. Evolution, vol. 5, pp. 339-344, 2RigSeT) pie 1952. Petrolacosawrus kansensis Lane, a Pennsylvanian reptile from Kansas. Univ. Kansas Paleont. Contr.: Vertebrata, art. I, pp. I-41, II figs., 3 pls. 1954. Pennsylvanian reptiles of Kansas. Bull. Geol. Soc. Amer., vol. 65, p. 1293 (abstract). PIVETEAU, JEAN (Eb.). 1955. Traité de paléontologie. Vol. V, Amphibiens, reptiles, oiseaux. 1113 pp. Paris. Romer, A. S. 1930. The Pennsylvanian tetrapods of Linton, Ohio. Bull. Amer. Mus. Nat. Hist., vol. 59, pp. 77-147, 26 figs. 1947. Review of the Labyrinthodontia. Bull. Mus. Comp. Zool., Harvard Coll., vol. 99, pp. 1-368, 48 figs. 1950. The nature and relationships of the Paleozoic microsaurs. Amer. Journ. Sci., vol. 248, pp. 628-654, 4 figs. 1956. Osteology of the reptiles. xxi+ 772 pp. 248 figs. Univ. Chicago Press. VAUGHN, PETER P. 1955. The Permian reptile Aracoscelis restudied. Bull. Mus. Comp. Zool., Harvard Coll., vol. 113, pp. 305-467, 15 figs., 2 pls. WILttston, S. W. 1908. The oldest known reptile—Isodectes punctulatus Cope. Journ. Geol., vol. 16, pp. 395-400, I fig., 1 pl. 1910. Cacops, Desmospondylus; new genera of Permian vertebrates. Bull. Geol. Soc. Amer., vol. 21, pp. 249-284, 12 pls. EXPLANATION OF PLATE 1 Left pes and lumbar region of Eosauravus copei seen under low-angle illumination. A. Pes, lighted from distal direction, showing clearly: Two elements—median centrale (c. 1) and 1st distal tarsal (dt. 1)—lying between astragalus (a) and Ist metatarsal; and 6th distal tarsal (dt. 6) lying between calcaneum (cal) and 5th metatarsal (mt. 5). B. Pes, lighted from proximal direction, showing 3 distinct elements (indicated by black dots) lying median to large 4th distal tarsal. C. Presacral vertebrae of lumbar region, lighted from anterior direction, show- ing low neural spine (ns), presence of intercentrum (ic), and striated patches (sc) possibly representing body scales. 14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOUS 139; NOS 1, PEs 2 (For explanation, see p. 14.) Hr SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 139, NUMBER 2 THE BIRDS OF ISLA ESCUDO DE VERAGUAS, PANAMA (With One PLatTE) By ALEXANDER WETMORE Research Associate Smithsonian Institution CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION JULY 8, 1959 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 139, NUMBER 2 He BIRDS OF ISLA ESCUDO DE VERAGUAS, PANAMA (WitrH ONE PLATE) By ALEXANDER WETMORE Research Associate Smithsonian Institution (Pus.icaTion 4378) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION JULY 8, 1959 THE LORD BALTIMORE PRESS, INC. BALTIMORE, MD., U. S. A. iris BIRDS OP ISLA, ESCUDO, DE VERAGUAS, PANAMA By ALEXANDER WETMORE Research Associate, Smithsonian Institution (WitH ONE PLATE) Isla Escudo de Veraguas lies in the southern Caribbean Sea at lat. 9°00’ N., long. 81°34’ W., distant a little more than 18 kilometers from Coco Plum Point on the base of the Valiente Peninsula, Prov- ince of Bocas del Toro. The island is roughly rectangular, with a projecting point at the southeast and a somewhat irregular shoreline on the western and northern sides. It is a little over 4 kilometers long by less than 14 wide, with the long axis running east and west. A sand beach extends along three-fourths of the southern side, around the flat, open southeastern point, and across the eastern side, past the mouth of a small stream, to end against a cliff, 12 meters high, of sandy, indurated clay. Similar bluffs separated by short stretches of beach mark the shoreline along the west and north. The northern side is broken by a small bay with a sand beach at its head. On the west the sea has cut back into the land, leaving several small islets, some of them barren except for grass and other low herbage, and some with a crown of brush and trees. Wave action is steadily erod- ing the low cliffs, forming small caves, and in some cases arches that pass through projecting points to the sea on the opposite side. The shallow bank surrounding the island indicates that this process has served to reduce it in size. The land back of the southern beach, elevated sufficiently above high-tide line to form a flat, is fringed with coconut palms on the sea side. Behind these extends low jungle in which scattered trees rise 15 to 20 meters tall. Toward the center the surface is lower and is swampy, with two or three trickles of fresh water, discolored by swamp peat, that drain to the sea. There is a small stand of mangroves at the mouth of the stream that enters the sea above the southeastern point. Columbus during his fourth voyage sighted the island on Oc- tober 17, 1502, when he came out of the Laguna de Chiriqui through Canal del Tigre (Tiger Channel) (Morison, 1942, vol. 2, p. 350). He gave it the name El Escudo as it appeared to resemble an escudo, SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 139, NO. 2 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 or shield. In the following years the island became a landmark for navigators along this stretch of coast, and is mentioned from time to time in ancient documents, the name being abbreviated often to Scudo, Scuda, or sometimes modified to Skoday (Anderson, 1911, p. 371). Presently it was designated Escudo de Veragua, and finally the latter part of the name became Veraguas. In the last voyage of Sir Francis Drake (Hakluyt, 1904, pp. 239-240) it is related that his ships came to Escudo on January 10, 1596, where they anchored on the southern side, remaining until January 23. The island was described as “not past two leagues long full of wood, and hath great store of fresh water . . . and that very good.” Many of the men soon fell sick, and Drake himself contracted the illness that caused his death on January 28 when they were near Portobello. He was buried at sea off that harbor. In occasional seventeenth-century accounts of buccaneers and other voyagers there is casual reference to Escudo de Veraguas as a place of shelter or a source of water. Dampier’s observation (Dampier, 1697, p. 39) made in 1681 that “We past by Scuda, a small Island (where ’tis said Sir Francis Drake’s Bowels were bury’d)” repeats a tale, apparently of common belief, that cannot concern this island since Drake’s death and burial, off Portobello, came more than 200 kilometers to the east. Escudo was visited by Indians, since Dr. Matthew W. Stirling of the Smithsonian Institution informs me that in the town of Bocas del Toro he was shown artifacts found on the island, proof that aboriginal people had lived there, at least from time to time. But there may be confusion with some larger place in the report (Anderson, p. 272) that records a considerable Indian population, divided under two caciques or chiefs. The land area, with due allowance for a reasonable amount of erosion since these early times, is too small to have permitted permanent residence for many persons. At present men come at intervals to gather the coconuts, or occa- sionally to fish, search for turtles, or to hunt the introduced wild pigs. There is no permanent human resident, and the wildlife, except for the pigs, is tame. I was able to visit Escudo de Veraguas through the kind assistance of George Munch, manager of the Almirante Division of the Chiriqui Land Co., which has its headquarters at Almirante, Province of Bocas del Toro. We left Almirante on February 28, 1958, shortly before midnight, on the diesel launch Talamanca, entered the sea through the pass of Boca del Toro, and before dawn anchored in the lee on the southeastern end of the island. Accompanied by Ziska Hartmann NO. 2 BIRDS OF ISLA ESCUDO DE VERAGUAS—WETMORE 3 and Jorge Burke, I was ashore near the southeastern point shortly after 7 o’clock and during the forenoon worked through the southern, level section parallel to the beach nearly to the western end. As the sun rose higher the humidity and heat of the dense jungle, where no breeze could enter, became oppressive, so that it was pleasant at the end to walk back to our cayuco along the open beach. At dawn the following morning the breeze blew from the mainland to the south, so that waves were breaking on the beach. We went off before 7:00 in a choppy sea, and finally landed near the mouth of the small stream. I crossed first into the ridge area at the northeast, but finding this difficult travel and unproductive I sought more level ground. Through this I crossed again toward the western end parallel to the northern shore. The sky was overcast, one shower of rain came, and at times it was difficult to see birds in the heavy jungle shadows. Though there were no trails, the low jungle was open and easy to penetrate. Where the growth became dense the ground was covered heavily with vines. On the north and west the surface rose Io to 25 meters in broken, steep-sided ridges, separated by little valleys. Here there was much undergrowth of the spiny pita (a plant of the pine- apple family) which, with the steep, slippery slopes, and the swampy floors of the small valleys between, made it difficult to get about. The taller trees that grow along the crests of these ridges from the sea give a misleading appearance of true high forest. On this final day we returned to the launch a little after 11:00 and, as the sea was rising, left for Almirante, returning through Crawl Cay Channel. The only record of any earlier visit of a naturalist to the island is the skin of a white-crowned pigeon in the collections of the University of California at Los Angeles. From the end of February to early in April 1936, Dr. Loye Holmes Miller of the Department of Zoology of that Institution, on sabbatical leave, accompanied by a graduate student, Frank Richardson, as assistant, visited the Laguna de Chiri- qui, living on a barge that served as a base for a Navy Hydrographic Office detail engaged in a survey of the area. Dr. Miller informs me that on March 2 Richardson accompanied a shore party of Navy per- sonnel to Escudo and brought back a white-crowned pigeon. No other specimens were taken. While Escudo de Veraguas lies well offshore, it is located on a bank where the sea is shallow. A narrow trench of 24 to 35 fathoms lies to the west and southwest, but elsewhere the depths are considerably less. Since it is estimated that sea levels dropped from go to 120 meters during the last period of extensive glaciation in Wisconsin 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 time during the Pleistocene, it is apparent that then the island was part of the mainland. A similar connection should have come during part or all of the three preceding periods of maximum glacial ice. Return of warmer temperatures in the interglacial periods, which melted the ice, again raised the water level, placing Escudo once more as an island, remote at sea. It is reasonable to suppose that the resi- dent wren and the manakin, as well as the peculiar spiny rat of the island, were established there during one of the periods of land con- nection, since they are jungle creatures that do not range far from cover, nor are the birds of kinds that would be readily windblown by violent storms. Whether the characters of size and color that now mark them were theirs in whole or in part on their arrival, or whether these are distinctions that have developed during isolation, cannot be said, except that it seems probable that the peculiarity of greater size may have become intensified, since this condition is found regu- larly in populations that seem to have been restricted for long periods to small islands. The manner of development of the differences that mark the blue-gray tanager is not easily understood since in mainland regions these birds appear to roam far. It would appear that they may not cross fairly wide water barriers, since another insular form is found on Isla Coiba off the Pacific coast of Panama (Wetmore, 1957, p. 94). Though there were few species of resident birds on Escudo de Veraguas, individuals were fairly numerous. The songs of the bay wren, joined occasionally by the raucous notes of a small flock of parrots, were regular bird notes of the jungle, aside from which there were only the subdued sounds of the wind in the higher treetops, and of the wash of waves against the shoreline. The smaller birds were encountered mainly in the more level areas, where at times they were detected with difficulty in the dim shadows that prevailed in the thickets when the sky was overcast. Occasionally I noted large spiny rats of the genus Hoplomys. One that I shot on the ground proves to be a form new to science. ANNOTATED LIST Family PELECANIDAE: Pelicans PELECANUS OCCIDENTALIS Linnaeus: Brown Pelican, Alcatraz Pelecanus occidentalis Linnaeus, Systema naturae, ed. 12, vol. 1, 1766, p. 215. (Jamaica. ) Several were fishing around the island on the morning of March 2. NO. 2 BIRDS OF ISLA ESCUDO DE VERAGUAS—WETMORE 5 Family SuLipAE: Boobies SULA LEUCOGASTER LEUCOGASTER (Boddaert): Brown Booby, Piquero Moreno Pelecanus Leucogaster Boddaert, Table des planches enluminées, 1783, p. 57. (Cayenne. ) Scattered groups rested on small islets off the western end of the island, selecting those that were rocky or covered with short herbage. They were nesting here, as I noted several large down-covered young. At sunset adults came in from the open sea, flying low above the water, singly or in groups of three or four. As our launch passed, a number, part of them fully grown young, came flying out from the islets to circle about with evident curiosity. There were no frigate-birds here, and so the boobies were free from molestation. I estimated that about 200 individuals were present. Family CHARADRIIDAE: Plovers, Turnstones CHARADRIUS SEMIPALMATUS Bonaparte: Semipalmated Plover, Chorlito Semipalmado Charadrius semipalmatus Bonaparte, Journ. Acad. Nat. Sci. Philadelphia, vol. 5, August 1825, p. 98. (Coast of New Jersey.) A flock of 14 ranged the beach at the southeastern end of the island. Family ScoropacipaAE: Snipe, Woodcock, Sandpipers ACTITIS MACULARIA (Linnaeus): Spotted Sandpiper, Playerito Coleador Tringa macularia Linnaeus, Systema naturae, ed. 12, vol. 1, 1766, p. 249. (Penn- sylvania. ) One seen on March 1. NUMENIUS PHAEOPUS HUDSONICUS Latham: Whimbrel, Zarapito Trinador Numenius hudsonicus Latham, Index ornithologicus, vol. 2, 1790, p. 712. (Hud- son Bay.) One seen on the beach March 1. Family CoLuMBIDAE: Pigeons, Doves COLUMBA LEUCOCEPHALA: White-crowned Pigeon, Paloma Cabeciblanca Columba leucocephala Linnaeus, Systema naturae, ed. I0, vol. 1, 1758, p. 164. (Bahama Islands.) Two were seen March 1 in the top of a thickly leaved tree. A male in the collection of the University of California at Los Angeles was shot on March 3, 1936, by Frank Richardson, now of the Department 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 of Zoology of the University of Nevada, at the time student assistant with Dr. Loye Holmes Miller (see p. 3). Family PsrrracipAE: Parrots, Macaws AMAZONA AUTUMNALIS SALVINI (Salvadori): Red-fronted Parrot, Loro Frentirrojo Chrysotis salvini Salvadori, Catalogue of the birds in the British Museum, vol. 20, 1801, p. 271. (Lion Hill Station, Canal Zone, Panama.) Three pairs were seen in the early morning of March 1, and a female was collected. The same small group was observed the fol- lowing day. Family TRocHiLipAE: Hummingbirds AMAZILIA TZACATL TZACATL (De la Llave): Rieffer’s Hummingbird, Colibri Colimorena Trochilus Tzacatl, De la Llave, Registro Trimestre, vol. 2, No. 5, 1833, p. 48. (México. ) Several were observed among the lower shrubs back of the beaches. Family ALCEDINIDAE: Kingfishers MEGACERYLE TORQUATA TORQUATA (Linnaeus): Ringed Kingfisher, Martin Pescador Grande Alcedo torquata Linnaeus, Systema naturae, ed. 12, vol. I, 1766, p. 180. (México. ) One was recorded on March 2 near the mouth of the small stream at the southeastern end. Family PipripAE: Manakins MANACUS VITELLINUS (Gould): Gould’s Manakin, Matraco Pipra vitellina Gould, in Hinds, R. B. (editor), Zoology of the Voyage of H.M.S. Sulphur under the command of Captain Sir Edward Belcher, R.N., F.R.G.S., etc., during the years 1836-42, vol. 1, pt. 3 (Birds, pt. 1), October 1843, p. 41, pl. 21. (Panama = Panama City, Panama.) The manakin (fig. 1) was fairly common, ranking next to the wren in abundance. The birds were found among the branches of the smaller trees, where they were quiet, moving about rather slowly, often remaining motionless for several minutes at a time. I regretted that there was no indication of display among the males, as their larger size should make the noises that accompany these activities definitely impressive. The bird of Escudo de Veraguas was so different from the repre- NO. 2 BIRDS OF ISLA ESCUDO DE VERAGUAS—-WETMORE Ai sentative of this species around Almirante Bay that I recognized it as an unknown race when the first specimen came to hand. It is described in the following paragraphs: MANACUS VITELLINUS AMITINUS, subsp. nov. Characters ——Similar to Manacus vitellinus cerritus Peters? but definitely larger; bill distinctly larger and heavier; tarsi and toes Fic. 1—Gould’s manakin, Matraco. heavier ; adult male with lower back, rump, and posterior ventral sur- face, including the sides and under wing coverts, darker green ; female and immature male somewhat darker green throughout, with the abdomen less yellowish. Description.—Type, U.S.N.M. No. 468919, male adult, from Isla Escudo de Veraguas, Prov. Bocas del Toro, Panama, March 2, 1958, collected by Alexander Wetmore (orig. No. 22241). Entire crown 1 Manacus cerritus Peters, Proc. New England Zool. Club, vol. 10, September 22, 1927, p. 9. (Almirante, Bocas del Toro, Panama.) 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 to nape, including the lores, wings (except inner lesser coverts), upper back, and tail black; sides of head, throat and neck, including hind- neck, bright apricot yellow, becoming lemon chrome as the yellow collar meets the black of the back; lesser wing coverts, except the outermost, lemon chrome; lower back, rump, and upper tail coverts yellowish oil green; breast, sides, flanks, abdomen, and under tail coverts between warbler green and olive-green; an indefinite light wash of lemon yellow on center of breast and abdomen; outer under wing coverts Roman green, inner ones yellowish citrine; under sur- face of inner webs of primaries and secondaries, except toward the tips, dull white. Bill dull black; tarsus and toes fuscous; claws dark neutral gray (from dried skin). Measurements——Males (3 specimens), wing 59.3-61.4 (60.6), tail 39.2-42.0 (40.2), culmen from base 14.0-14.8 (14.5), tarsus 23.8-24.5 (24.1) mm. Females (2 specimens), wing 59.5-60.0 (59.7), tail 38.2-38.3 (38.3), culmen from base 14.7-14.7 (14.7), tarsus 21.0-21.5 (21.2) mm. Type, male, wing 59.3, tail 39.4, culmen from base 14.6, tarsus 23.8 mm. Range.—Isla Escudo de Veraguas, at sea off the base of Peninsula Valiente, Bocas del Toro, Panama. Remarks.—The greater size of this handsome bird as compared with mainland forms is evident on comparing the measurements with those listed in succeeding paragraphs. In bulk the island birds appear nearly one-third greater. In drawing the description comparison has been made with cerritus since the shades of yellow on head and neck of these two are more nearly in agreement. In terms of present distri- bution Manacus v. vitellinus is assumed to be the form of the main- land opposite Isla Escudo, since it is the one recorded at Cricamola on the shores of Laguna de Chiriqui, opposite Peninsula Valiente. Manacus v. cerritus is known to range south only to the southern shores of Almirante Bay so that if the water barrier is disregarded, cerritus and amitinus are separated by an intervening population of typical vitellinus. The name is taken from the Latin amitinus, a cousin. To determine clearly the affinities of the manakin from Escudo a survey has been made of the related members of the genus Manacus found in Panama, particularly Manacus vitellinus, of which an excel- lent series is at hand from the entire range including Colombia. It became evident immediately that cerritus, described by James L. Peters as a distinct species, was in fact a geographic race of M. vitellinus, as the supposed specific characters break down when the entire area NO. 2 BIRDS OF ISLA ESCUDO DE VERAGUAS—WETMORE 9 occupied by this bird is given review. It may be noted also that the display of males of cerritus, as I saw it in January and February 1958, was similar to that of typical vitellinus. Following is a summary of the subspecies of vitellinus based on this examination, with the races arranged in geographic sequence from west to east. MANACUS VITELLINUS CERRITUS Peters Manacus cerritus J. L. Peters, Proc. New England Zool. Club, vol. 10, Sep- tember 22, 1927, p. 9. (Almirante, Bocas del Toro, Panama.) Characters.—Similar in color pattern, and in colors in general, to Manacus v. vitellinus. Male, with throat, sides of head, and band across hind neck and upper back more yellow, less orange, varying in some to completely bright yellow; lower breast, abdomen, sides, flanks, and under tail coverts more greenish yellow ; rump and upper tail coverts brighter green; female, and male in immature plumage, darker green throughout. Measurements.—Males (9 specimens), wing 51.8-54.2 (53.3), tail 31.2-35.8 (34.2), culmen from base 11.1-12.3 (11.7), tarsus 20.0-22.6 (21.5) mm. Females (3 specimens), wing 54.0-55.7 (54.9), tail 33.1-34.4 (33.9), culmen from base 11.8-12.5 (12.0), tarsus 20.2-21.4 (20.9) mm. MANACUS VITELLINUS AMITINUS Wetmore Characters—Generally similar to M. v. cerritus, but decidedly larger ; darker green. Range.—Isla Escudo de Veraguas, Province of Bocas del Toro, Panama. Full details of differences, and of measurements, are given in the description above. MANACUS VITELLINUS VITELLINUS (Gould) Pipra vitellina Gould, in Hinds, R. B. (editor), Zoology of the Voyage of H.M.S. Sulphur, under the command of Captain Sir Edward Belcher, R.N., F.R.G.S., etc., during the years 1836-42, vol. 1, pt. 3 (Birds, pt. 1), October 1843, p. 41, pl. 21. (“Panama’= Panama City, Panama.) Characters Similar to M. v. cerritus, but male decidedly orange on foreneck, throat, sides of head, and band across base of neck; posterior under surface more greenish; rump and upper tail coverts grayer green. Measurements.—Males (47 specimens), wing 50.4-55.7 (52.3), tail He) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 25.8-31.5 (28.3), culmen from base 11.0-13.0 (11.8), tarsus 20.4- 22.4 (21.4) mm. Females (46 specimens), wing 50.7-54.9 (53.2), tail 27.3-31.7 (29.3), culmen from base 11.1-12.7 (12.0), tarsus 18.3-20.7 (19.4) mim. Range.—On the Pacific slope from the foothills of eastern Veraguas (Santa Fé) eastward through the western part of the Province of Panama (La Campana, Chorrera), throughout the Canal Zone, and eastern Panama, to extreme eastern Darién (Jaqué, Rio Jaqué, Cana) ; on the Caribbean slope from central Bocas del Toro (Cricamola), through northern Veraguas (Guaval on Rio Calovevora), northern Coclé (El Uracillo), the Province of Col6n (Chilar, Portobello) and the Comarca de San Blas (Mandinga, Permé, Obaldia) ; entering Colombia on the western side of the lower Rio Atrato (Unguia, Chocéd) and along the shores of the Gulf of Uraba at Acandi, Choco, on the western side, and Necocli, Antioquia, on the east. This is the first published report of this race for Colombia. Speci- mens from Acandi and Unguia, both near the Panamanian boundary, are like typical examples from Panama. A series of 7 males from Necocli on the eastern shore of the mouth of the Gulf of Uraba averages faintly paler, more yellowish green below, and very faintly more yellowish orange on the head. They thus show an approach toward the paler milleri of the Sint Valley to the east, but are to be placed with vitellinus. Gould published the description of this manakin twice, first in the Zoology of the Voyage of H.M.S. Sulphur, where it appeared in October 1843 as indicated above. The bird was displayed with 8 other new species from this voyage at a meeting of the Zoological Society in London in July 1843, but publication in the Proceedings did not come until December. In the first publication, in October, Gould states that “The specimen here figured was procured by Mr. Hinds at Panama, and is the only one I have seen.” The introduction to the Voyage of the Sulphur indicates that the vessel made surveys along the entire Pacific coast of the Republic, but it appears clear that the locality “Panama” refers to the vicinity of Panama City, which is the only place mentioned that lies within the range of vitellinus. This is accepted, therefore, as the restricted type locality. MANACUS VITELLINUS VIRIDIVENTRIS Griscom Manacus vitellinus viridiventris Griscom, Bull. Mus. Comp. Zodl., vol. 69, April 1920, p. 179. (Jiménez, near Buenaventura, Valle, Colombia.) Characters.—Similar to M. v. vitellinus, but male with lower breast, abdomen, sides, flanks, under tail coverts, rump, and upper tail coverts NO. 2 BIRDS OF ISLA ESCUDO DE VERAGUAS—WETMORE II definitely darker green ; yellow of anterior part of body, including the neck band, somewhat less orange, more yellow ; female darker green, in this resembling female M. v. cerritus, from which it differs in being somewhat less yellowish on the abdomen, and duller green above. Measurements —Males (14 specimens), wing 50.6-53.7 (52.2), tail 26.3-30.6 (28.7), culmen from base 10.8-12.5 (11.6), tarsus 20.4- 22.7 (21.5) mm. Females (6 specimens), wing 53.0-54.3 (53.5), tail 28.1-30.1 (29.5), culmen from base 11.6-12.4 (11.9), tarsus 19.1-20.0 (19.6) mm. Range.—Western Colombia, from northern Chocd (Rio Jurado, Rio Jurubida, Nuqui) and northwestern Antioquia (Villa Artiaga, Dabeiba) south through western Caldas (Santa Cecilia) and Valle (Puerto Muchimbo, Jiménez), including the upper Cauca Valley (Riofrio, Cali). This race has been supposed to range into extreme eastern Darién at Cana but specimens from that locality agree best with typical vitellinus, MANACUS VITELLINUS MILLERI Chapman Manacus vitellinus milleri Chapman, Bull. Amer. Mus. Nat. Hist., vol. 34, Dec. 30, 1915, p. 645. (Puerto Valdivia, Antioquia, Colombia.) Characters—Much paler than M. v. vitellinus; male with head (except for the black crown) and band across hindneck bright, light yellow, without orange; rest of lower surface much paler, being grayish green with a wash of yellow; rump and upper tail coverts paler ; female, definitely paler below, being whitish on abdomen, and duller, grayer green above. Measurements—Males (11 specimens), wing 49.7-52.9 (51.6), tail 26.8-30.4 (28.6), culmen from base 10.8-12.2 (11.5), tarsus 20.6-22.3 (21.3) mm. Females (6 specimens), wing 52.5-54.5 (53.7), tail 28.8-30.8 (29.6), culmen from base I1.0-12.0 (11.6), tarsus 19.0-20.0 (19.4) mm. Range.—Northwestern Colombia, from the valley of Rio Sint (Nazaret, Socarré) in western Bolivar, south to the middle Cauca Valley in northern Antioquia (Taraza, Puerto Valdivia) ; recorded from Remedios in east central Antioquia at the head of Rio Ité, a tributary of the lower middle Rio Magdalena. In the series at hand this race is typical on the middle Rio Cauca in northern Antioquia. In some specimens from the lower Rio Sint, taken at Nazaret, Tierra Alta, Socarré, and Quebrada Salvajin, most of the males have the head somewhat more orange, and the breast I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 and abdomen somewhat darker, varying in the direction of vitellinus. They are thus somewhat intermediate, but are definitely near mulleri. It has been suggested that Manacus aurantiacus (Salvin) found on the Pacific slope of western Panama would eventually prove to be conspecific with M. vitellinus, but my studies to date do not bear out this supposition. Brighter color, particularly in the male, and smaller size mark aurantiacus uniformly throughout its range from south- western Costa Rica through Chiriqui, southern Veraguas, and both sides of the Azuero Peninsula in Veraguas, Herrera, and Los Santos. Manacus vitellinus vitelinus from near Santa Fé, Veraguas, and La Campana and Chorrera in the western section of the Province of Panama, where intergradation, if present, should occur, show no variation from the normal pattern of that race. From present infor- mation the two groups appear to be separated by a savanna area in which neither is found. The two appear so completely distinct that there is no basis for uniting them. Aldrich (1937, p. 95) separated the population of the western side of the Azuero Peninsula as Manacus aurantiacus flaviventris, as a series from that area appeared brighter colored than those available at the time from western Chiriqui and southwestern Costa Rica. During the course of my own field investigations I have accumulated a considerable series from Veraguas and eastern Chiriqui, and have examined additional material from western Chiriqui and Costa Rica. A study of this extensive material indicates that the supposed differ- ences do not hold. Males in fresh plumage from both areas are strongly orange, but as the season progresses there is fading, particu- larly in the dry months when sun is more intense. The following measurements may be useful for comparison with those of the races of Manacus vitellinus. Males (25 specimens), wing 44.8-47.8 (46.3), tail 26.0-30.2 (28.7), culmen from base 11.2-12.2 (11.7), tarsus 19.5-20.6 (20.1) mm. Females (21 specimens), wing 47.8-50.0 (48.7), tail 209.0-30.9 (30.3), culmen from base 11.3-12.3 (11.8), tarsus 18.2-20.5 (19.1) mm. Family TyrANNIDAE: Tyrant Flycatchers TYRANNUS MELANCHOLICUS CHLORONOTUS Berlepsch: Tropical King- bird, Pechi-amarillo Grande Tyrannus chloronotus Berlepsch, Ornis, vol. 14, 1907, p. 474. (Temax, Yuca- tan.) A female was collected and several others seen along a stretch of sandy beach, where they rested on the open ends of branches, or on the tops of low shrubs. NO. 2 BIRDS OF ISLA ESCUDO DE VERAGUAS—WETMORE 13 Family HirUNDINIDAE: Swallows PROGNE SUBIS (Linnaeus): Purple Martin, Golondrina Turquina Hirundo Subis Linnaeus, Systema naturae, ed. 10, vol. 1, 1758, p. 192. (Hudson Bay.) On the return journey on March 2 I noted an occasional purple martin flying northward, low over the water, near the mainland coast from the vicinity of Plantain Cay to Chiriqui Point. These swallows are known as migrants through México and Central America, but little is reported regarding them in Panama. The only published record that has come to my attention is by Zimmer (1955, pp. 4, 5) of an immature male of the southwestern subspecies, Progne subis hesperia Brewster, taken at Cocoplum, Bocas del Toro, October 27, 1927. At Almirante on February 18, 1958, during a forenoon of nearly constant rain, a band of 8 purple martins came to rest in dead branches of a tall avocado tree beside our house. At intervals others arrived until finally between 35 and 40 were present, resting in close forma- tion. When the rain ceased and the sky became lighter two hours later they disappeared. From then until March 6, I recorded purple martins in northward flight, singly or in scattered, straggling groups, across Almirante Bay, along its shoreline, or over the outer beach near Boca del Drago. Occasionally a few came to rest in the tree beside the house. It appears that there is a regular flight in migration along the Caribbean coast. The female of a pair taken on February 18, in its darker color on the under surface and in wing length of 148 mm., represents typical Progne subis subis. The male, with the wing 149.7 mm., agrees in size with that race. Family TRoGLoDYTIDAE: Wrens THRYOTHORUS NIGRICAPILLUS Sclater: Bay Wren, Cucarachero Castafio Cabecinegro Thryothorus nigricapillus Sclater, Proc. Zool. Soc. London, pt. 28, May 1860, p. 84. (Nanegal, 4,000 feet elevation, Ecuador.) This wren (fig. 2) was the most common land bird on the island, found in pairs scattered through the undergrowth. Though they were encountered most often in low tangles, where creepers were matted and cover was dense, they ranged also out into more open areas, and at times worked up through branches and creepers into the tops of the taller trees. They were quite tame, often appearing within 6 feet or so. On our second day ashore the sky was overcast and it was often difficult to see these birds in the darkly shadowed coverts. We were usually notified of the presence of a pair by the series of repeated I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 notes that made up the clear song. This resembled closely that of Thryothorus nigricapillus costaricensis as heard at Almirante, but seemed to be higher in tone and somewhat less varied in repertoire. One pair worked busily at a nearly completed nest located near the tip of a leafy branch about 6 feet from the ground in heavy under- growth. This was a ball, nearly round, of palm and other slender Fic. 2.—Bay wren, Cucarachero Castaiio Cabecinegro. fibers, with the ends projecting all around as a rough fringe. The entrance was in one side. The larger size and paler color of this island population in com- parison with the birds of the adjacent mainland were easily evident in the field. A description of this previously unknown race follows. THRYOTHORUS NIGRICAPILLUS ODICUS subsp. nov. Characters —Similar to Thryothorus nigricapillus costaricensis (Sharpe)? but larger, with longer, heavier bill; in color paler brown. 2 Thryophilus costaricensis Sharpe, Catalogue of the birds in the British Mu- seum, vol. 6, 1881, p. 217. (Valley of the Rio San Carlos, Alajuela, Costa Rica.) NO. 2 BIRDS OF ISLA ESCUDO DE VERAGUAS—WETMORE 15 Description—Type, U.S.N.M. No. 469015, male adult, from Isla Escudo de Veraguas, Bocas del Toro, Panama, taken March 1, 1958, by Alexander Wetmore (original No. 22230). Throat, upper fore- neck, malar region, loral area, a line on the margin of upper and lower eyelids surrounding the eye, a superciliary line extending back from the center of the eye, and the auricular region white, with some mixture of black on loral area and along upper eyelid; crown, hind- neck, side of neck, side of head, except as noted above, and a line separating the white malar area from the throat, deep black; back, rump, and upper tail coverts auburn, the tail coverts with short central bars of black along the shaft; wing coverts auburn, with irregular shaft lines and subterminal bars of dusky neutral gray; tertials and outer webs of secondaries auburn, barred heavily with dusky neutral gray; outer webs of innermost primaries auburn, changing on the outer ones to hazel, the brighter color finally reduced to a narrow edging on the ninth and tenth; concealed webs of remiges fuscous- black ; rectrices dusky neutral gray, barred narrowly with hazel ; breast and center of abdomen ochraceous-tawny; sides and flanks hazel; under tail coverts ochraceous-tawny, barred heavily with black; axil- lars ochraceous-tawny; under wing coverts ochraceous-buff, mixed with white ; edge of wing white. Maxilla dusky neutral gray ; mandi- ble pale smoke gray, becoming smoke gray at the base; tarsus and toes fuscous-black (from dried skin). Measurements.—Males (5 specimens), wing, 75.2-79.2 (77.0), tail 58.6-62.3 (60.2), culmen from base 21.8-24.2 (23.2), tarsus, 28.4- 21-8 (29.7): mm. Females (6 specimens), wing 70.2-72.8 (71.6), tail 54.5-58.8 (56.8), culmen from base 21.0-22.3 (21.5), tarsus 26.2-28.7 (27.2) mm. Type, male, wing 75.2, tail 58.8, culmen from base 24.0, tarsus 29.1 mm. Range.—Isla Escudo de Veraguas, at sea off the base of the Valiente Peninsula, Bocas del Toro, Panama. Remarks.—tThe actual difference in measurements will be indicated by consulting the summary of a series of Thryothorus nigricapillus costaricensis, the nearest relative, both physically and geographically, that is given in the review of the species that follows. The name of the new race is from the Latin odicus, musical, appro- priate because of the pleasing song. The complete and definite dissimilarity in the lower surfaces found in this group of wrens between the chestnut-breasted, white-throated groups of the Caribbean slope of Nicaragua, Costa Rica, and Bocas 16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 del Toro, and the forms with the anterior under surface barred closely with black and white that range from eastern Darién through western Colombia to Ecuador, long led to their separation under two specific names. The series of specimens now available justifies their union under the specific name nigricapillus, though it may be supposed that the two terminal groups must have been separated for a long period to have become so completely different. In costaricensis, the darkest of the Central American races, and the one farthest removed from those of South America, remote common ancestry with the other is indicated in the rather indistinct black bars found on the breast and sides in the juvenile plumage. This marking may persist in the fol- lowing plumage, especially on the abdomen, but many are plain chest- nut on the posterior lower surface except for the bars on the lower tail coverts that are common to many of the numerous species of the genus Thryothorus. Proceeding eastward along the Caribbean coast of Panama from the valley of the Rio Calovevora, on the boundary between the provinces of Bocas del Toro and Veraguas, the wrens of the species under discussion become paler brown, with sides and flanks barred with black, except for occasional plain individuals. This group —the race castaneus—is found through the lowland Caribbean drain- age of the Canal Zone. Continuing eastward there is an abrupt change near Portobello and in the foothills of the Cerro Azul in which the plain white of the throat extends down on the upper breast, the brown on the sides becomes paler, and there are strongly marked black bars on sides, lower breast, and abdomen in most individuals. This style—the race reditus—crosses to the Pacific slope along the base of the Cerro Azul, and at Chiman has reached the coastal lowlands. On the Caribbean slope it continues almost to the Colombian boundary in the Comarca de San Blas, and on the Pacific side to about the western boundary of Darién near the Golfo de San Miguel. There is then rather abrupt transition to birds with lower surface heavily barred—the race schotiti. Markings on the white throat are faint or absent, and the brown is restricted to the flanks and under tail coverts. In the valley of the Atrato the barring reaches its maximum and here the throat in most specimens is heavily marked. The plainer throat persists to the eastward in Colombia along the Rio Sinu, and on the middle and upper Rio Cauca. In southwestern Colombia, beginning in the Depart- ment of Cauca, the throat bars begin to lighten still more and to disappear, and farther south, in Narino, the upper breast also becomes less heavily marked. This style leads over to typical nigricapillus of Ecuador, in which throat and upper breast are white, without bars, and NO. 2 BIRDS OF ISLA ESCUDO DE VERAGUAS—WETMORE 17. the flanks and under tail coverts are lighter brown. In all the changes that have been described these wrens have remained uniformly chestnut above, with black crowns. One possible explanation of this interesting gradient might be that the plain, chestnut-breasted forms had become established fairly early in the Central American area where they have continued with modifi- cation toward the elimination of barring. In the South American area, on the other hand, the barring became intensified. Through a subse- quent spread of range in the latter population, the two groups have been brought in contact, with resultant hybridization that has caused the mixing that has been described. The races recognized as reditus and castaneus represent two stages in this process. It would appear that the schottii group has been the one in active expansion because of the extensive range that it now occupies. It is interesting that the chestnut-breasted group is not found farther north in Central America, though there would appear to be no ecological barrier to prevent this. Hellmayr (1934, p. 180) includes another group, Thryothorus semibadius Salvin, found in tropical lowlands of the Pacific slope from southwestern Costa Rica to western Chiriqui, also as a race of nigricapillus, but this does not seem justified. The bird in question is more finely barred, with 3 narrow dark bars on the individual feathers of the breast, and the crown chestnut, concolor with the back; also it is smaller. In the schottu-nigricapillus group, which semibadius resembles superficially, the black bars are heavier, there are 2 bars on the individual feathers of the breast, the crown and upper hindneck are deep black, and the size is larger. There is no indication whatever of hybridization between semibadius and the adjacent Thryothorus n. costaricensis. While juveniles of the costaricensis-nigricapillus group show spots or a slight wash of brown on the pileum and hindneck, the crown cap remains plainly defined. Thryothorus semibadius would appear to be an older offshoot of the ancestral stock that has produced the forms with barred breast, and from its limited range one that may be on its way to extinction. The following summary, based on extensive series throughout the entire range of these birds, outlines findings as to their relationships and distribution. It should be noted that museum series of skins almost invariably include immature individuals that are not fully grown, especially in the development of the wings. These are easily detected and have been omitted in the measurements that are given under the different forms. 18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 THRYOTHORUS SEMIBADIUS Salvin: Salvin’s Wren, Cucarachero Castafio Cabecimoreno Thryothorus semibadius Salvin, Proc. Zool. Soc. London, November 1870, p. 181. (Bugaba, Chiriqui.) Characters—Crown and hindneck chestnut, concolor with the back; under surface white, barred, except for the throat, narrowly with black, the breast feathers having three black bars; size smaller. Measurements.—Males (8 specimens), wing 61.4-65.5 (63.3), tail 42.0-49.2 (46.3), culmen from base 18.7-21.0 (19.9), tarsus 23.4-24.0 (23.7) mm. Females (4 specimens), wing 61.2-64.7 (63.2), tail 43.7-47.4 (45.8), culmen from base 19.9-20.8 (20.3), tarsus 23.1-24.5 (23.8) mm. Range.—Tropical zone of the Pacific slope from southwestern Costa Rica in the valley of the Rio Pirris to western Panama in the Comarca del Bart (Puerto Armuelles), and the lowlands of extreme western Chiriqui (Divala, Bugaba). THRYOTHORUS NIGRICAPILLUS Sclater: Bay Wren, Cucarachero Castaiio Cabecinegro Thryothorus nigricapillus Sclater, Proc. Zool. Soc. London, pt. 28, May 1860, p. 84. (Nanegal, 4,000 feet elevation, Ecuador.) Characters.—Crown and hindneck deep black, in sharp contrast to the chestnut of the remainder of the upper surface; under surface chestnut, auburn, chestnut-brown, clay color, or white, barred more or less with black ; in the races that are white below, with 2 black bars on each breast feather ; size larger. THRYOTHORUS NIGRICAPILLUS COSTARICENSIS (Sharpe) Thryophilus costaricensis Sharpe, Catalogue of the birds in the British Museum, vol. 6, 1881, p. 217. (Valley of the Rio San Carlos, Alajuela, Costa Rica.) Characters—Throat and upper foreneck white, rest of lower sur- face auburn to hazel; sides in some specimens with a few bars of black, which usually are indistinct. Measurements.——Males (17 specimens), wing 66.5-72.0 (69.3), tail 51.0-56.8 (54.3), culmen from base 20.4-22.7 (21.3), tarsus 24.5-27.8 (25.9) mm. Females (9 specimens), wing 62.5-67.2 (64.6), tail 47.8-54.0 (50.2), culmen from base 19.4-21.7 (20.5), tarsus 23.2-25.6 (24.5) mm. Range.—Caribbean slope from southeastern Nicaragua (Los Saba- los, Rio Escondido, San Juan del Norte) through eastern Costa Rica (Rio Frio, Guayabo, Bonilla, Jiménez, Reventazén) to central Bocas NO. 2 BIRDS OF ISLA ESCUDO DE VERAGUAS—WETMORE 19g del Toro, Panama. Specimens from Cricamola at the eastern end of the Laguna de Chiriqui are intermediate toward castaneus. Sharpe described this bird from a single specimen that he said was collected by Adolphe Boucard in Costa Rica, without giving a more definite locality. Boucard (1878, p. 51) in an account of his collec- tions made in Costa Rica listed this wren as Thryophilus castaneus Lawrence, with the statement “Several specimens, from San Carlos; killed in February.” In his itinerary he says that this locality was in the Valley of the Rio San Carlos, a tributary of the Rio San Juan on the Atlantic slope. I have therefore designated this area as the type locality. THRYOTHORUS NIGRICAPILLUS ODICUS Wetmore Characters —Similar to T. n. costaricensis but larger, with longer, heavier bill; paler brown. Measurements.—Given above. Range.—Confined to Isla Escudo de Veraguas, off the base of the Valiente Peninsula, Bocas del Toro, Panama. THRYOTHORUS NIGRICAPILLUS CASTANEUS Lawrence Thryothorus castaneus Lawrence, Ann. Lyc. Nat. Hist. New York, vol. 7, June 1861, p. 321. (“Atlantic slope near the Panama Railroad’= Lion Hill, Canal Zone.) Characters —Similar to T. n. costaricensis, but paler brown on ventral surface, with the white of the throat extending farther down on the foreneck, in some reaching the upper breast; more definitely barred with black on sides and flanks, in some specimens with the bars extending across the lower breast and abdomen. Measurements.—Males (14 specimens), wing 66.2-70.7 (68.9), tail 49.4-53-7 (51.7), culmen from base 20.3-22.0 (21.0), tarsus 24.9-27.5 (25.8) mm. Females (17 specimens), wing 63.1-67.0 (64.8), tail 46.3-53.4 (49.2), culmen from base 19.3-21.9 (20.2), tarsus 23.4-25.7 (24.7) mm. Range.—Caribbean slope from the valley of the Rio Calovevora in eastern Bocas del Toro, through northern Veraguas, northern Coclé (extending inland on the northern slope in the higher foothills to the headwaters of the Rio Coclé del Norte and the Rio Indio), and western Colon (Chilar, Rio Indio, Colén, Marajal), to the Canal Zone (Gatun, Lion Hill, Barro Colorado Island, Frijoles). Back of Fl Valle, Coclé, I found these birds at 2,000 feet elevation along the upper course of the Rio Mata Ahogada, ranging on its 20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 higher branches to 2,500 feet. The divide here between this stream, which flows into the Pacific, and the Rio Indio of the Caribbean side is low so that rainfall in the heads of the valleys is sufficient to main- tain the type of green-leaved undergrowth that these wrens frequent across for a short distance on the Pacific side. The birds here do not range below 2,000 feet elevation where the scrub growth changes to the semiarid type characteristic of the Pacific lowlands of this area. This is the only point known to me at which the race castaneus crosses to the Pacific slope. Records of Salvin (1867, p. 134) and of Salvin and Godman (1880, p. 88) for Santiago de Veraguas are not sup- ported by specimens in the Salvin and Godman collections now in the British Museum (Natural History) and are certainly in error. The type specimen of castaneus, described by Lawrence, came to him in a collection made by James McLeannan and John R. Galbraith during the winter of 1860-1861. The collectors were located at Lion Hill, but it must be borne in mind that it is certain they covered a considerable area along the line of the railroad in the course of their work. In the present instance Lawrence (1861, pp. 315-316) states that their specimens were taken ‘‘on the Atlantic side of the isthmus” except for half a dozen species that he lists, which do not include the bird here under consideration. Though the type specimen of castaneus is labeled only ““Panama” with the initials of the collectors, the desig- nation “Lion Hill” found in current literature may be accepted as the restricted type locality. THRYOTHORUS NIGRICAPILLUS REDITUS Griscom Thryophilus nigricapillus reditus Griscom, Bull. Mus. Comp. Zool., vol. 72, January 1932, p. 358. (Permé, Comarca de San Blas.) Characters.—Similar to T. n. castaneus but with white of breast more extensive; sides, abdomen, and under tail coverts paler, duller brown; more heavily and extensively barred with black. Measurements——Males (15 specimens), wing 67.0-70.5 (68.9), tail 47.5-54.3 (52.0), culmen from base 19.3-21.9 (20.9), tarsus 24.0-26.5 (25.5) mm. Females (11 specimens), wing 63.2-67.7 (65.4), tail 45.0-51.4 (48.5), culmen from base 19.0-21.5 (20.1), tarsus 23.1-26.3 (24.7) mm. Range.—From eastern Colon (Portobello) eastward on the Carib- bean slope through the Comarca de San Blas (Mandinga, Permé, Puerto Obaldia), crossing through the western Cerro Azul to the head of the Rio Pacora on the Pacific slope, ranging eastward in the Prov- ince of Panama along the Pacific side of the Serrania de Majé NO. 2 BIRDS OF ISLA ESCUDO DE VERAGUAS—WETMORE 21 (Quebrada Cauchero, on the base of Cerro Chucanti), reaching tide- water at Chiman, and on the Rio Majé (Charco del Toro). This race constitutes the definite intergrade between the western group with bright brown breast and little or no barring, and the eastern and southern population with completely barred breast. Transition between castaneus and reditus on the west is fairly abrupt, an intermediate condition being evident in one specimen from near Frijoles in the Chagres drainage. Birds from near Colon are definitely castaneus, while those from near Portobello, 30 kilometers to the east, are reditus. At the eastern end the type locality at Permé is barely within the range, since skins from Puerto Obaldia, about 15 kilometers farther east, are intermediate toward schottii, which is the race found on the coast at Acandi, Chocd, Colombia, 25 kilo- meters beyond Puerto Obaldia. THRYOTHORUS NIGRICAPILLUS SCHOTTII (Baird) Thryophilus schottii Baird, Review of American birds in the Museum of the Smithsonian Institution, vol. 1, August 1864, p. 123 (in Key) ; September 1864, p. 133. (Rio Truand6é, Choco, Colombia.) Thryophilus nigricapillus connectens Chapman, Bull. Amer. Mus. Nat. Hist., vol. 31, July 23, 1912, p. 157. (Cocal, 5,000 feet elevation, Cauca, Colombia.) Characters.—White of throat and foreneck extending down over breast, sides, and center of upper abdomen; lower surface heavily barred with black, in typical form the bars covering the throat, but in intermediate stage the throat partly or wholly plain. Measurements.—Males (16 specimens), wing 64.0-66.9 (67.3), tail 44.6-51.6 (48.2), culmen from base 19.5-21.5 (20.4), tarsus 24.4-26.8 (25.4) mm. Females (10 specimens), wing 59.9-65.6 (63.0), tail 43.0-47.8 (45.6), culmen from base 19.0-20.8 (19.6), tarsus 23.0-25.0 (24.1) mm. Range.—Darién, eastern Panama, from the lower Rio Sambu (Jesusito), and the lower Rio Tuira (Cituro, on Rio Cupe) inland to 600 meters elevation near Cana, and south to the valley of the Rio Jaqué; continuing in Colombia throughout Chocd (from the Pacific coast across to Acandi on the Gulf of Uraba), and western Antioquia in the Atrato valley (Villa Artiaga), and western Valle (Buenaventura and San José), to western Cauca (Cocal) ; east into southern Bolivar in the upper Sint Valley (Socarré, Quebrada Salvajin), and northern Antioquia in the lower Cauca Valley (El Pescado), and the valley of the Rio Nechi (Regeneracion, El Real, 22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 Hacienda Belén), crossing to the Rio Magdalena drainage on the Quebrada Enanea (Volador). Remarks.—The typical form of this race, with the throat and fore- neck distinctly barred with black, is found mainly in the Choco. In southwestern Colombia, through western Cauca, the throat barring disappears, and in Narifio the breast appears whiter as the barring on this area is reduced. The birds of this region are intergrades of unstable character between schottii and nigricapillus. The influence of the reditus style of markings produces similar intergrades on the opposite side of the range, beginning in northern Chocé at Acandi on the Gulf of Uraba, and extending across to the upper Sint Valley and the lower Nechi. Specimens from this area are identical in appear- ance with those of western Cauca which Chapman named connectens. Under these circumstances there is no basis for recognition of such a race, as the supposed characters, unstable at best, are duplicated on the opposite side of the population of typical schotiu. The birds described are allocated as intermediates to schottti, except for those of Narifio which are placed best with typical nigricapillus. THRYOTHORUS NIGRICAPILLUS NIGRICAPILLUS Sclater Thryothorus nigricapillus Sclater, Proc. Zool. Soc. London, pt. 28, May 1860, p. 84. (Nanegal, 4,000 feet elevation, Ecuador.) Characters —Similar to T. n. schottti, but averaging lighter brown on back, flanks, and under tail coverts ; throat, foreneck, and center of upper breast immaculate, with the barring reduced on the sides. Measurements.—Males (13 specimens), wing 62.6-67.1 (65.1), tail 44.2-50.8 (48.1), culmen from base 19.1-20.9 (20.1), tarsus 24.0-25.5 (24.6) mm. Females (6 specimens), wing 63.9-66.8 (65.5), tail 46.6-50.7 (48.7), culmen from base 19.2-21.6 (20.0), tarsus 23.0-25.3 (24.2) mm. Range.—lrom western Narifio (intermediate) in Colombia south through the tropical zone of western Ecuador, nearly to the boundary with Pert. Remarks.—As indicated under schottiu, specimens from Narifio are intermediate. Family MimipaE: Mockingbirds, Thrashers DUMETELLA CAROLINENSIS (Linnaeus): Catbird, Pajaro Gato Muscicapa carolinensis Linnaeus, Systema naturae, ed. 12, vol. 1, 1766, p. 328. (Virginia. ) Three were noted, and one female was collected. NO. 2 BIRDS OF ISLA ESCUDO DE VERAGUAS—WETMORE 23 Family PAruLIDAE: Wood Warblers DENDROICA PETECHIA ERITHACHORIDES Baird: Golden Warbler, Canario Manglero Dendroica erihtachorides (= erithachorides, typographical error, corrected in index) Baird, Report of explorations and surveys ... for a railroad from the Mississippi River to the Pacific Ocean, vol. 9, pt. 2, Birds, 1858, pp. 283, 976. (Cartagena, Colombia.) These warblers (fig. 3) were found scattered through the taller trees where they were fairly common, though each of the four taken appeared to be alone. It should be noted that on Escudo they were not restricted to the limited growths of mangroves found near the sea, as is the case on the mainland, but ranged throughout the forest growth, as appears to be the regular habit of this warbler when found on small islands. On the present island they ranked third in abundance among the smaller land birds. The four taken include three adult males which are similar to a small series from the shores of Almirante Bay on the nearby mainland. A female that had just begun the molt from the gray juvenile dress to the yellow adult plumage had the skull fully ossified, indication that this character as a criterion of age is not re- liable in tropical areas, where the life cycle of an individual bird is not necessarily arranged on a calendar year basis. The series from Escudo and from Almirante Bay agree fully with type material of this race, which is interesting since specimens from Limon, Costa Rica, about too kilometers to the north, are Dendroica p. bryanit. Family THRAUPIDAE: Tanagers THRAUPIS VIRENS (Linnaeus): Blue-gray Tanager, Azulejo Loxia virens Linnaeus, Systema naturae, ed. 12, vol. 1, 1766, p. 303. (Surinam.) Blue-gray tanagers were fairly common in the taller trees, a num- ber being seen and three collected. It has been unexpected to find that they are so different from the widely distributed race of the mainland that they merit description as an additional subspecies. THRAUPIS VIRENS CAESITIA subsp. nov. Characters Similar to Thraupis virens diaconus (Lesson)* but darker, particularly below; central lower surface nearly uniform in shade from throat to under tail coverts; sides definitely darker ; bill longer and heavier. 3 Tanagra (Aglaia) diaconus Lesson, Rev. Zool., June 1842, p. 175. (Realejo, Nicaragua. ) 24 SMITHSONIAN MISCELLANEOUS COLLECTIONS Fic. 3—Golden warbler, Canario Manglero. VOL. 139 NO. 2 BIRDS OF ISLA ESCUDO DE VERAGUAS—WETMORE 25 Description—Type, U.S.N.M. No. 469168, female, Isla Escudo de Veraguas, Bocas del Toro, Panama, March 2, 1958, collected by Alexander Wetmore (original No. 22248). Crown mineral gray, with a faint wash of gnaphalium green, which is stronger on hindneck ; back and scapulars dull greenish glaucous-blue, changing to light glaucous-blue on rump; upper tail coverts bluish gray-green, washed with greenish glaucous-blue at tips; shoulder patch formed by lesser and middle coverts, grayish violaceous blue ; primaries and secondaries dusky neutral gray, with outer webs, except for the tips of the primaries, dull Venetian blue; outer webs of scapulars dark gobelin blue; central rectrices and outer webs of others dark gobelin blue, with inner webs of all but the central pair dark neutral gray ; median under surface between court gray and gnaphalium green, with center of abdomen faintly whitish; sides gnaphalium green; edge of wing glaucous-blue ; under wing coverts light gull gray to white. Bill dull black, except for a wash of hair brown toward base of gonys; tarsus, and toes dusky neutral gray (from dried skin). Measurements—Females (3 specimens), wing 87.5-90.1 (88.4), tail 60.1-62.8 (62.3), culmen from base 16.4-18.0 (17.1), tarsus 20.4- 20.7 (20.6) mm. Type, female, wing 90.1, tail 62.8, culmen from base 18.0, tarsus 20.4 mm. Range.—Isla Escudo de Veraguas, at sea off the base of the Valiente Peninsula, Bocas del Toro, Panama. Remarks.—The fact that this widely distributed tanager was repre- sented by a distinct form on this small island was not detected until I began examination of specimens in the preparation of the present report. The three specimens, all females, were taken merely as a matter of routine during my visit. Comparison has been made with a series of recently collected skins, consisting of 15 females of Thraupis virens diaconus, and 21 of T. v. cana. In none of these is there dupli- cation of the characters on which the race caesitia is based. Attention was first drawn to the island form by the large bill, this measuring 13.8 to 15.7 (14.6) mm. in the 15 diaconus, and 13.7 to 15.7 (14.7) mm, in the 21 cana. Hellmayr (1936, p. 214) expressed doubt as to the validity of the race diaconus, and recently Blake (1958, p. 566) has combined this form with cana. In comparing an extensive series taken throughout the range of the two subspecies in question I find, however, that while the two are similar in general, diaconus is darker on the back, and slightly duller blue on the rump, in addition to averaging somewhat 26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 darker in color below. These characters hold in birds of Central America south through the Isthmus of Panama, with intergradation in extreme northwestern Colombia. In making comparison it is necessary to separate adult from immature birds, since the distinctions listed are masked when this is not done. I believe the confusion regarding the two races has been due to lack of understanding of this fact. The name given to the new race, in connection with its darker coloration, is from the Latin caesitius, meaning bluish. LITERATURE CITED ALpricH, Joun W. 1937. Annotated list of birds, in Aldrich, J. W., and Bole, B. P., Jr., The birds and mammals of the western slope of the Azuero Peninsula [Republic of Panama]. Sci. Publ. Cleveland Mus. Nat. Hist., vol. 7, pp. 27-130, Aug. 31. ANDERSON, C. L. G. tot. Old Panama and Castilla del Oro. Pp. i-xiv, 1-550, 7 maps, 35 pls. Washington. BLakeE, Emmet R. 1958. Birds of Volcan de Chiriqui, Panama. Fieldiana: Zool., vol. 36, No. 5, PP. 499-577, June 25. Boucarp, ADOLPHE. 1878. On birds collected in Costa Rica. Proc. Zool. Soc. London, 1878, PP. 37-71, I pl. DAMPIER, WILLIAM. 1697. A new voyage round the world, describing particularly, the Isthmus of America, several coasts and islands in the West Indies, the Isles of Cape Verd, the passage by Tierra del Fuego, the South Sea coasts of Chili, Peru and Mexico; the Isle of Guam one of the Ladrones, Mindanao, and other Philippine and East India islands near Cambodia, China, Formosa, Luconia, Celebes&c. New Hol- land, Sumatra, Nicobar Islands; the Cape of Good Hope and Saint Hellena. London, pp. (1-8), i-vi, 1-550, 5 maps. Haxtuyt, RIcHarp. 1904. The principal navigations voyages traffiques & discoveries of the English nation made by sea or over-land to the remote and farthest distant quarters of the earth at any time within the compasse of these 1600 years. Vol. X, pp. 226-245. Glasgow. HELLMAYR, CHARLES E. 1934. Catalogue of birds of the Americas and the adjacent islands. Field Mus. Nat. Hist., pt. 7, pp. i-vi, 1-531, Nov. 15. 1936. Idem, pt. 9, pp. i-vi, 1-458, Oct. 6. Mortson, SAMUEL ELroT. 1942. Admiral of the Ocean Sea, A life of Christopher Columbus. Vol. I, pp. i-xlvi, 1-448, 14 maps, 25 ills.; vol. 2, pp. i-viii, 1-445, 18 maps, 2 illus. Boston. NO. 2 BIRDS OF ISLA ESCUDO DE VERAGUAS—WETMORE 27 SALvIN, OSBERT. 1867. On some collections of birds from Veragua. Proc. Zool. Soc. London, 1867, pt. I, pp. 129-161, 1 pl., June. SALVIN, OsBERT, and GopMAN, FREDERICK DUCANE. 1879-1904. Biologia Centrali-Americana, Aves, vol. 1 (text), pp. i-xliv, I-512. WeEtmMorE, ALEXANDER. 1957. The birds of Isla Coiba, Panama. Smithsonian Misc. Coll., vol. 134, No. 9, pp. 1-105, 4 pls., 15 figs., July 8. ZIMMER, JOHN T. 1955. Studies of Peruvian birds. No. 66, The swallows (Hirundinidae). Amer. Mus. Nov. No. 1723, pp. 1-35, Apr. 29. ie “ me Mi SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLE Iso NO 2) (Pia 1. Western end of Isla Escudo de Veraguas, from the south. 2. Southern shore of eastern end of Isla Escudo de Veraguas. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 139, NUMBER 3 PUR THER OBSERVATIONS ON DISTRIBUTION, OF PATTERNS OF COAGULATION: OF (‘THE -HEMORYMPH INSINEOTROPIGAN INSECTS By CHARLES GREGOIRE Department of Biochemistry, Institut Léon Fredericq University of Liége, Belgium CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION AUGUST 18, 1959 THE LORD BALTIMORE PRESS, INC. BALTIMORE, MD., U. S. A. FURTHER OBSERVATIONS ON DISTRIBUTION OR PADIERINS OF COACUEATION OF TEE HEMOLY MPH IN NEOTROPICAL INSECES: By CHARLES GREGOIRE Department of Biochemistry, Institut Léon Fredericq University of Liége, Belgiwm The present paper is a contribution to a long-term inquiry on dis- tribution of patterns of hemolymph coagulation in various arthropods, especially in insects. The reactions of the main elements involved in the process of co- agulation of the hemolymph—a category of unstable hyaline hemo- cytes (coagulocytes: Grégoire and Florkin, 1950) and the plasma— differ in various insects. These differences, appreciated by phase- contrast microscopy, have been classified into four patterns of micro- scopic pictures (Grégoire, 1951). The characters of these patterns may be described as follows: Pattern I. Inception of the plasma coagulation in the shape of islands of coagulation around the hyaline hemocytes.—Selective alter- ations in the unstable hyaline hemocytes (shrinkages of the cell body and occasionally of the nucleus, sudden expansions, bulging of blisters and of blebs) result in exudation or in explosive discharge of cell material into the surrounding fluid. Coagulation of the plasma starts in the shape of circular islands of granular consistency around the altered hyaline hemocytes. The islands of coagulation develop to a certain size; then their increase stops. At the beginning of the process, the islands are scattered and separated by fluid channels. When the coagulation proceeds farther, the plasma in these channels clots into a granular substance in which the islands preserve generally their original size and shape. The mechanism involved in pattern I is identical to one of the types of coagulation described by Hardy (1892), Tait (1rg10, 1911), Tait and Gunn (1918), Numanoi (1938), and Grégoire (1955b) in crus- 1 This is No. 9 in a series of papers entitled “Blood Coagulation in Arthro- pods” published in various journals. SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 139, NO. 3 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 tacean blood, in which a special category of cells, the Hardy’s ex- plosive corpuscles, corresponding to the insect hyaline hemocytes or coagulocytes, plays a selective part in the inception of the coagulation of the plasma. Pattern II, Extrusion of cytoplasmic expansions by hyaline hemo- cytes, with development of cytoplasmic meshworks. Reaction in the plasma in the shape of veils—On contacting the glass, a category of fragile hyaline hemocytes undergoes alterations that differ from those characterizing pattern I. These corpuscles extrude threadlike cyto- plasmic expansions, sometimes of considerable length. These ex- pansions are highly adhesive to solid particles (dust, chitinous debris), other hemocytes, and physical interfaces (bubbles). These alterations result in formation of cytoplasmic meshworks of various complexity, on which the other kinds of hemocytes are passively agglutinated. The reaction in the plasma after these cellular changes occurs in the shape of transparent, elastic, and contractile veils, developed within the cytoplasmic systems built up by the hyaline hemocytes, or in their vicinity. In various insects the alterations in the unstable hemocytes are not followed by changes in the plasma, and the modifications of the hemolymph in vitro consist only of a cellular reaction. Pattern III, Patterns I and II combined.—Association of the re- actions taking place in patterns I and II characterizes the picture in pattern III. In the same film of hemolymph, hyaline hemocytes send out cytoplasmic expansions (pattern I1) while islands of coagulation (pattern I) appear around the body of these corpuscles. When they develop within the veils, which characterize the reaction in the plasma in pattern II, the islands form circular, denser areas centered by the altered unstable corpuscles. Pattern IV. No modification in the hyaline hemocytes, or altera- tions not followed by visible reaction in the plasma, in the optical con- ditions of phase-contrast microscopy.—lIn the pictures of this pattern, hemocytes resembling in their cytological characters the unstable corpuscles involved in the other patterns do not visibly alter. They appear as pale vesicles containing a few dark particles. In several insects, these corpuscles are the remnants of darker refractile, hyaline, frequently oenocytoid-like hemocytes, which undergo clarification after explosive discharge of a part of their cytoplasm. In the vicinity of these inert or altered hyaline hemocytes, no change can be detected under the phase-contrast microscope in the consistency of the plasma. NO. 3 HEMOLYMPH COAGULATION IN INSECTS 5 Specimens from more than 1,000 species of insects and of other arthropods have already been tested about the pattern of coagulation of their hemolymph or blood (Grégoire, 1951, 1953, 1955a, b, 1957, unpublished observations on palearctic insects (1957-1958) ; Grégoire and Jolivet, 1957). Predominance of one of the patterns has been observed in several taxonomic groups. In other groups, owing to the scarcity of the data available, or to large variations in the results, the pattern representative of a species or of a group at a supraspecific level could not be established. The aim of the present study was to fill some gaps in the data. Four hundred Neotropical insects, belonging to 215 species, including 185 species not yet investigated, were collected and studied during visits to Tingo Maria, Peru (Estacion Experimental Agricola), August 1956, and to the Smithsonian Institution’s tropical preserve on Barro Colo- rado Island (Canal Zone Biological Area), October 1956. MATERIAL AND METHODS The samples of hemolymph were mostly thin films prepared as soon as possible after capture. The hemolymph issuing from severed or punctured appendages (antennae, legs, wings, joints of the wing- cases) was placed immediately in contact with the edge of a cover glass lying on a slide and was allowed to spread out into films. A phase-contrast optical equipment WILD M/1o was used for the observations (see Grégoire, 1955a, p. 105, and 1957, pp. I and 3). RESULTS DISTRIBUTION OF THE PATTERNS OF COAGULATION OF THE HEMO- LYMPH IN INSECTS (TABLE I) Detailed descriptions of the four patterns of coagulation of the in- sect hemolymph, used in the present study, have been given elsewhere (Grégoire, 1955a, p. 104; 1957, pp. 4-6 and text figs. I-4). In the table, the names of the species are followed by the numbers of specimens studied (adults, unless otherwise stated) and by the patterns of coagulation provisionally found predominant or repre- sentative on the basis of the study of several samples of hemolymph obtained from these specimens. Incidental findings of other patterns are reported under “Comments.” In order to avoid duplication, the patterns recorded in the present study in 50 insects belonging to Neotropical species previously in- vestigated (Grégoire, 1957) are reported in the notes, preceded by the date “(1957).” 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 The patterns of coagulation have been represented in the table by the following symbols: @: pattern I: inception of the plasma coagulation in the shape of islands of coagulation around the unstable hyaline hemo- cytes. Various degrees of extension of the process in the films. ©: pattern II: development of cytoplasmic meshworks by hyaline hemocytes. Reaction in the plasma in the shape of veils. ©: pattern II incomplete: emission of cytoplasmic expansions, characterizing the reactions of the hyaline hemocytes in pat- tern II, but unaccompanied by formation of veils in the plasma. @): pattern III: patterns I and II combined. —: pattern IV: no visible coagulation by phase-contrast micros- copy. (): pattern incidentally or exceptionally recorded in limited fields of preparations exhibiting predominantly another pattern. (?): microscopical characters of a pattern not clear-cut or equivocal. Artifacts possibly involved. Other abbreviations used: sp., species; spm., specimen; T., speci- men captured and studied at Tingo Maria; B., specimen captured and studied on Barro Colorado Island. Gradations in the intensity of the reactions, especially with regard to pattern I, are indicated by the following symbols: I poor (scarce fringes of clotted plasma around a limited number of altered fragile hyaline hemocytes, without extension of the coagulation; I (scattered islands of coagulation of various sizes, with moderate coagulation of the fluid outside the islands) ; I*, I**, 1*** (islands around all the hyaline hemocytes, substantial and general coagulation of the film). NO. 3 HEMOLYMPH COAGULATION IN INSECTS 5 TABLE 1.—Patierns of coagulation Patterns of coagulation representative Number of or predominant Material specimens in samples Comments Orthopteroid Complex DICTYOPTERA BLATTODEA ? # Periplaneta australasiae (Fabricius) (adulbiand larval CDs) 04 Jk ee 2 @ > Archimandrita tessellata Rehn (B.). 1 @ PHASMATIDAE * Pseudophasma menius Westwood & RCE EPUB ieee HOS Lael Wav Parr'ata aliens 6 I © 7 Prisopus cerosus Westwood (B.)... 1 eS * Prisopus ariadne Hebard (B.)..... I ) uF 3 ulidet.isp. (2 adults, 1 larva) (T.). 3 @ G25) aes) ORTHOPTERA PENTIGONIIDAE + Scudderia paronae (Griffini) (T.)... 1 ® 7 Eupeucestes crassifolius (Haan) 9% ly eae cht accis sletera als Wiehe \annrargs 2 © res) Undet. larva (Phaneropterinae) DD aid aaa Ta ORD Va IU A I ® Acanthodes aquilina (Linnaeus) CIDE) Reret ere ett cman cease aa I 6 x Micracentrum: sp. ©, (B. \lea/sccwe se 33 I @ Neoxtphidion conocephalus saltator Goaussieey OUT) oe ie ealeitias I ge he Moncheca pretiosa (Walker) (T.).. 1 @ ih EUMASTACIDAE PGrOniaSia ees NOD.) sate < qaceess des, s 3 © GRYLLIDAE* Paragryllus temulentus Saussure Cee A sare ieee Meh ION, So coe I @ a GRYLLACRIDAE + Abelona salvini (Saussure and Pictet) eC ITA EPR Nice GRABS op au I @ 1 Det. by Dr. C. Willemse. 2 (Grégoire, 1957) Epilampra azteca Saussure (B.): I ***, L a ies Neoconocephalus affinis (P. de B.) 2 (B.): I (**); Caulopsis microprora He- ‘bar Pisce Ls 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 TABLE 1.—Patterns of coagulation—continued Patterns of coagulation representative Number of or predominant Material specimens insamples Comments ORTHOPTERA (continued) FROSCOPILDAE Apioscelis verrucosa Brunner Von Weattenwyl) OCT) aie) I ACRIDIDAE =+ Orphulella concinnula Walker (T.). Tetrataenia surinama (Linnaeus ) OS it es) pert Seccmnete eee eres Leptysma imsularis (Bruner) (T.).. Wind etespyn( Al BM As cals atu gees mie eee a Dicaearchus (gen. sp. nov.?) (T.).. Kegua crenulata.Stoll (B.) 2.2... DERMAPTERA ? PENG ASD Cleat: )is sftusuaiye. iabemraldied: > 2 Hemipteroid Complex HEMIPTERA REDUVITDAE *6 Saica meridionalis Fracken and PSIMOH (MESS ees ec ars Sims eiepeuete ee eee I -— Stenopoda cinerea Laporte (B.).... poor eS poor poor 22k 2K OK Se eH HW Ci") (O poor or @) = (@ °) ws | Rasahus sulcicollis (Serville) (B.).. Zelusisp:? (nymph) (Ps). cb 2 3 ss Zeus spat (nympi)e Cl Wisc daists sxe Castolus subinermis (Stal) (B.).... Moniuna lobaia'Stal (1.). 2% os 8b: Montna fjumosa (Stal) (T.) 2.025. Brontostoma notatum Stal (B.).... Doldina bicarinata Stal (T.)....... PYRRHOCORIDAE © Largus balieatus Stal (T.)......... Dysdercus incertus Distant (T.).... 12 a (S) Ss Se HN SS Se = 4 (Grégoire, 1957) Copiocera specularis Gerstaecker: I; Osmilia flavolineata (de Geer, (T.): I poor; Xyleus rosulentus Stal, 3 larvae (T.): I (**); Schistocerca paranensis Bur meister CL poor: 5 Det. by Dr. me Catz! 6 (Grégoire, 1957) Saica apicalis Osborn and Drake (B.) : —; Zelurus spinidorsis (Gray, (B.): — (II poor or incomplete) ; Panstrongylus rufotuberculatus (Champion) (B.): — Panstrongylus geniculatus (Latreille), 3 spm. (B.): — (1 NO. 3 HEMOLYMPH COAGULATION IN INSECTS 7 TABLE 1.—Patterns of coagulation—continued Patterns of coagulation representative Number of or predominant Material specimens in samples Comments HEMIPTERA (continued) PYRRHOCORIDAE (continued) Dysdercus ruficeps (Perty) (T.)... 1 —_ (Se) Dysdercus sp.? (nymph) (T.)..... I -- COREIDAE® thine. decorata Stal (T.) ..... 432+ I -— Spartocera fusca (Thunberg) (T.). 1 —~ Plapigus foltaceatus (Blanchard) CODY C Clee ciel. dey wean ere 2 _— Anasa haglunds Stal (T.).......... I — Hypselonotus striatulus (Fabricius) LEA er SYS wie eae atahe dig wate ts I — Paryphes adelphus mutans Horvath CTE) Sore Nea ey, iets wo aiken. S I — Hyalymenus tarsatus (Fabricius) CER Riawoate woke aka sei BAe ied aigok i — Leptocorisa filiformis (Fabricius) (1S Ue URS AR a to ec NE PU ee PaO 2 — Zoreva dentipes (Fabricius) (T.)... 5 - Co) Zoreva spintfera Stal. (T,). 3.2.5 5. 2 -—— GELASTOCORIDAE® Nerthra peruviana (Montandon) CUE Nl Sissel aes iatal ss Save. oscar tiace.s ahs I -- CO) PENTATOMIDAE™ Symphylus deplanatus (Herrich- SACRO ails tlaiaes\'a cm aiatn migra I — Augocoris gomesti Burmeister (T.). I — Macropygium reticulare (Fabricius) Meveyaralaiteh ser lcitelealraileioest arse) stele) s 2 == Euschastus crenator (Fabricius) Reyes Ohne Miser ah Bem Le Ts ie) ica 950 2 —_ Euschistus sp.? (nymph) (T.)..... I — ( ?) Loxa picticornis Horvath (B.)..... I -~ ETEORIEEUS SPI Ess) secu ci u 2 aye erynre = I —_ Edessa .afiuis, Wallas (T.) .. 3.5.1 2 _- Edessa polymita Distant (B.)...... I _ Ba cssa.spe SEU Ce CLs aces aes es I -~ 7 (Grégoire, 1957) Mecistorhinus piceus (Palisot de Beauvois) (T., B.), 2 spm.: —; tdessa rufomarginata De Geer, 4 spm. (B.): —; Acrosternum scutellatum Distant (T.): —; Veodine macraspis (Perty), (B.): —. 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 TABLE 1.—Patterns of coagulation—continued Patterns of coagulation representative Number of or predominant Material specimens in samples Comments HEMIPTERA (continued) PENTATOMIDAE (continued) dessa Spe (Ba) its iniete lea bie erie oe I a Fdessaispa: (oyimph) (Ls)... .8-o-. I — Bidessn spire Gaympl) (Cy cine I — MIRIDAE * Mimoncopeltus, n. sp. (T.).......- I — HOMOPTERA CICADIDATE = Carineta sp., near boliviana Distant A Gea DN oescreee aps ak ciaiiaNonaleiare Fe ete I @ FULGORIDAE*»?? Copidocephala ornanda (Distant) CBG yen des ciste se alse mies I & te Odontoptera sp. (CB iis. seein. ss sae: I @ ses: Diareusa annularis imitatrix (Ossia- UNitTSamy (GBS) frecnceancieyci ctsiaiecee sci I 2) a Gen. and sp. unknown (B.)....... i & + CL DAE ® Gen. and sp. unknown (B.) 0.0.05 i — (?) DICTVOPEAARIDAE * Nera florens Stal (Bi)... oc)s66 << a 2) a Taosa herbida (Walker) (B.)...... I @ . Gen. and sp. unknown (B.)........ I Cs] ree MEMBRACIDAE & Star Oleows spy DiC Tha) jee ase weet = oats I @ o CERCOPIDAE® Cephisus siccifolius Walker 9 (B.).. 1 @ oo AMOS Oops Tors (s)he eta ia elas 2 @ + Ai Gusta Eo: Ot Al Lin) aia ai, scsisin aie 2 © sok Homaspis sp. ae tis (1). .cieae ie eae I @ (@ ?) AROMAS PIS iS: tae 2c (W.)) sais see ss ahs I @ RK ( @) FRX) MOMS PIS ISD... 52 3) Pulled). «greta Oss 014 I @ Kei MOmaspis SD: EAs (UA) cies alee pier I @ he Momaspisison sess hE). ee eee I @ +F MOMS PIS Sv aE Geils e)) seem eee I @ ce 8 Det. by Miss Louise M. Russell. 9 Det. by Dr. D. A. Young; Diareusa by Dr. V. Lallemand. 10 (Grégoire, 1957.) Nae elegans (Olivier), 2 spm. (B.): I ***; Cathedra serrate (Fabricius) (B.): NO. 3 HEMOLYMPH COAGULATION IN INSECTS 9 TABLE 1.—Paiterns of coagulation—continued Patterns of coagulation representative Number of or predominant Material specimens in samples Comments TOMOPTERA (continued) CERCOPIDAE (continued) Monmaspes Span 7 96 Clo) wns. 3. 3s 2 @ ***(@)) MOMISPIS Sp: SEO oy CLs) 8. vad s oes I @) aa omasyis sp. 220 6 CB:). 2.256222. I ~~ (@ ?) CICADELLIDAE? Tettigellinae Diestostemma nigropunctata (Signo- BEE Me elise ticks eka ollclfolsioi'e «ats olde wose I @ **(@) Deestostemma:sp, -O CL...) sai. sie ys ss I 8 4k Baleja flavoguttata (Latreille) (B.) 1 ® re Spatmknow On) sepa esate ha anf, I @ Oncometopia sp. +: I, sex anomaly Ba Alsias Aree lic Ph) es Gallia stare ads I @ poor (@)) Oncometopia sp. #£2, normal? (T.). 1 — SpyeuniknOwan CB.) jars /-! .ieiatesslecs s 08s 6 @ 7 LO Wes MAAR E A Wee Mate fel cire 8/5. oy ol) dats) e\aseven we) s “Gypona”’ decorata Fowler (B.).... 2 @ poor Gypona atitlana Fowler (B.)....... I — (?);@ probable Gypona hebes Fowler (B.)........ 3 © poor to **; (@);— in I spm. ALGIERS IPM (1 a)ip Belarc 12: je) 536, 2%! sh. 0h ove 3 I ~ (?); dry spm. PORENESD Ua) e050 RGN, Licata 3 — (@ °) in I spm. Gyponana sp: Ga B2)\. 3. Assen eain « I @ probable. Negostana’ sp. sete (BSS o2bod cc I © Wegostana' sp. 422 9 (Biche. . fei 2 —(@ ?) @ probable PEATIDAE = Anormelis nigrolimbata (Fowler) CEP Tae aris Otis sixes eiaieie shea 8 — (@ poor) Flatormems sp. (2) (B.).......... 2 — (?) Paradascalia metvi (Distant) (B.).. 6 — (@ ?) ES SIDA? Oroneqia isos (CB)... bith gacretes pe I _ 11 (Grégoire, 1957) Carthaeomorpha rufipes Melichar, 3 spm. (B.): I **; (—?) in 1 spm. IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 TasLe 1.—Patterns of coagulation—continued Patterns of coagulation representative Number of or predominant Material specimens insamples Comments COLEOPTERA ADEPHAGA CARABIDAE * Hurpalinae sp.et (B.) 6s. se. ees I sas Fharpalinae sp. 32°(B.)\.. os os ese ss I — Harpalinae sp. 3£3.(B.) 656.2 ee I G WARE Spt e GE) oe sin oe erecta sek ao I O poor (—) Migr arspume 2 (B.) ssi is aislds selene I () (e) probable PACD IM IST Ala) id a's) 8 wleday a nie ane atare = Ve I — POLY PHAGA PASSALIDAE 1224 Passalus (Neleus) interstitialis ESC lISChIc Es) verve etanelel aeaiiaye'= le inp I o— Vetwrins Sp. (QB. )\.'. derais/saeiciels «5 ais I — (@) SCARABAEIDAE * Coprinae (Scarabaeinae)** Cama sp. \(W.)ia's siclc lal lerals erereie'es I @ Uroxys gorgon Arrow (B.)......-. 2 = (a) Rutelinae ** Mesomerodon spinipenne Ohaus Ca pes oe eoceetine olay sls aoe ayia tava 2 ©) * Pelidnota chlorana Erichson (T.)... = 2 @) Anomala virescens Burmeister (T.). 2 (B) (Ss) Pigonala SSP. TCL) sacle ee ole eel ake ares 3 @) Dynastinae ** 3° Gen? near Bothynus: CU.)i.d..< atarereawe I O CEBRIONIDAE ** Zeta TKO WIISI( LE <))\s ceo she olaheyesedei= I 8 * ELATERIDAE Chalepleprdius ep iC Bi) so exe ose ayoyspues I © (@) SCMUIOTES ‘SP. CT: Dis aisrsse lets ciordieieqinteg I © LY CIDAE *% ME NILEST SDM Wea) ale pnts id tatiess te Allee aes tere I z (Os) LAMPYRIDAE POHAUS |S. MLS) os sats oie wale 0 spe me 2 ? 12 Det. by G. Fagel. 13 Det. by O. L. Cartwright. 14 (Grégoire, 1957) Veturius platyrhinus Westwood (B.): — (III?). 15 (Grégoire, 1957) Aspidolea singularis Bates (B.): II. 16 Det. by T. J. Spilman. 17 Det. by Dr. Ch. Jeuniaux. NO. 3 HEMOLYMPH COAGULATION IN INSECTS LE TaBLe 1.—Patterns of coagulation—continued Patterns of coagulation representative Number of or predominant Material specimens in samples Comments POLY PHAGA (continued) VIVE GY LIDAE A° Mekiomma spr CB: )). seks e6 6s «stn I @ or @ ENDOMYCHIDAE” Probably Amphiz sp. (T.)......... I Oo COCCINELLIDAE 7 Bi palachaa: Sp (ls) (cies wher ale -nai'e!2ys 32) I — Monomeda marginata (Linnaeus) GUEE )Raiacve toy srcvenetavace siciettle pis.«: ates, ste I — PROLEYVLIDAE *° Erotylus, prob. spectrum Thomson CULE Rg ciate ses eiaiate' da erchare a el asen ale I @ (@ ?) Prob. Homoeotelus sp. (T.)......-. I 2 *, (@) probable Gent unknown (CEs) a. 6. beets cases I Co Gene naknowa (Dia si< 4. 5.sr0he.0'6 510: I oO TENEBRIONIDAE * Strongylium auratum Laporte (T).. 1 © probable MELOIDAE CERAMBYCIDAE* Prioninae Sienadontes Sp. (E.) as vias.i3 alg srenis I @ ae ymad es sp, (Wa) whe daceie lence oye wiera o's I & *KE(@) Lamiinae Pe swt POTS. (1D. Viajes :s cae «ss ones I 8 (@ ?) TOMES! MUD )iicia ain s).2 si l'aide a oe) I @ poor Oreodera glauca (Linnaeus) (B.).. 1 @ ¥r Acanthoderes bivitta White (B.)... 1 @) **(@). ihogocherus sp. S21) (Be) ce vcieien'« I @ ge MaGgOCherus Spi:FE2 (OB. ) p)ssieicert -s,« I @ kK Golabotea spe (CEs) a5 sess doce. a 506s I gS a Charoules spit (Es) 2385 ssie cases I e@© tok yaroudes Sp. se 2 CN \leineictiae oe eae I @ ee 18 Det. by Dr. J. G. Rozen, 19 Det. by Dr. J. G. Rozen. 20 Det. by Dr. E. A. Chapin. 21 Det. by T. J. Spilman. (Grégoire, 1957) Zophobas prob. atratus (Fabricius) (B.): III ** probable. 22 Det. by T. J. Spilman. (Grégoire, 1957) Epicauta grammica (Fischer von Waldheim), Secum. (8.): 1 *** (IIT). 23 Det. by George B. Vogt. (Grégoire, 1957) Taeniotes scalaris (Fabricius) (B.): I (III). I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 TABLE 1.—Patterns of coagulation—continued Patterns of coagulation representative Number of or predominant Material specimens in samples Comments POLY PHAGA (continued) CHRYSOMELIDAE br. sense *4 EUMOLPIDAE: poss. near Priono- LOPE Ses (Ho) teen gyno vie wien eMetseke I -— Chrysomelinae Woryenona sp. is) 2 dessa eid eo I — (a) Seal odesi( ey Spee ile) cca ee erect I (@) Gosmogramma isp) \(T.)) ss. hae ee ee I CS?) (—) Galerucinae Drab rotica spe Kis) 5 aioe) ve ielaele ee I ao VAM ME GOON SIO ily) cet ale ove ee Gleate eee I —— Alticinae Gedionychus sp. 420 (0). oo ess 2 _— Oedionychus sp.) Fe2(T))). ui a'ss's I — Hispinae Oediopalpis guerini Baly (B.)...... 2 — Cassidinae Cyclosoma tristis Boheman (T.).... I o— Echoma sp., prob. aulica Boheman CHa vale ele a coletave eiaue itebeeutio rs I 2 poor CURCULIONIDAE Wig Pactusisp! sek (ler ie ve es 4 — Naupactus sp. ge 2: (Us) cn. so 56s I — COM PSUSNSP hea) eve ee eee ee 3 — Pett Pits Spy. eT 1B. )ieus ain eu aco /ssaere I — Hiciipus sp. see (UBS) eo cue es te 2 — MiCiOINGSIUS SPig( D\. \is ac sieves Gets oe I — Panorpoid Complex NEUROPTERA-PLANNIPENNIA MANTIS PID A Ey 26, 24 Climaciella semihyalina (Serville) GEER Che oie! aaivucie wiateueeatianets I = (@ ?) 24 Det. by George B. Vogt. 25 Det. by Miss Rose Ella Warner. (Grégoire, 1957) E-xophthalmus jekelianus (White), 2 spm. (T., B.): —. 26 Det. by Miss Sophy Parfin. 27 (Grégoire, 1957) Mantispa phthisica Gerstaecker (B.): —. NO. 3 HEMOLYMPH COAGULATION IN INSECTS 13 TaBLe 1.—Patterns of coagulation—continued Patterns of coagulation representative Number of or predominant Material YEUROPTERA-SIALODEA *8 CORY DALIDAE Corydalus sp., near armatus Hagen CM oe Ae a cae FRICHOPTERA BY DROPSY CHIDAE ** Prob. Leptonemaisp: ¢(B.)s 2.8: EE PIDOPTERA VAD AE sp, Cadult 2? (Be... SAUNT AE sp. (atva) Ci... « MCI DAE sp. Carva)! (1.) ... -..- JIPTERA LARVAEV ORIDAE *° Ormiophasia bushkiu TNS. TY MENOPTERA ICHNEUMONIDAE * WNetela spe OB sob ooo eet FORMICIDAE Aelecaspyae i) 9) (Bi). 58s sevice fone Asteca spy se2 0 (Ba )isi iets tates seats: Pachycondyla crassinoda (Latreille) SCL SR uiccics, nach diutegs (lope ayn ete Dinoponera sp. (worker) (T.)..... Labidus coecus (Latreille) ¢ (B.).. VESPIDALE 25:2 Poehymnenes, SpoCl..)'. vases os ees Polistes major weyraucht Bequaert (aN ea creer tera. MEM lui da « POMMPILIDAE *2° 28 Det. by Dr. A. B. Gurney. 29 Det. by W. D. Field. 30 Det. by C. W. Sabrosky. 31 Det. by Miss Luella M. Walkley. in samples O1O=| ~@@ | | @ Oo Comments KK very poor (—) * poor or @ possibly @ #4 (©) possibly © 82 Det. by Dr. M. R. Smith. (Grégoire, 1957) Paraponera clavata (Fabricius) ¢ (B.): [ **; Camponotus sericeiventris Guérin, br. sense, 4 workers (B.): possibly III. 33 Det. by K. V. Krombein. 34 (Grégoire, 1957) Polistes canadensis panamensis Holmgren, 4 spm. (B.): I. 35 (Grégoire, 1957) Anoplius a-amethystinus (Fabricius) (B.): III. 14 SMITHSONIAN MISCELLANEOUS COLLECTIONS TABLE 1.—Patterns of coagulation—concluded Patterns of coagulation representative Number of or predominant Material specimens in samples HYMENOPTERA (continued) SEH ECIDAE §8 Sceliphron fistulare (Dahlbom) (B.). 1 B Stictia maculata (Fabricius) (B.).. 1 @ ODONATA AGRIONIDAE * Megaloprepus coeruleatus (Drury) EA eisai ans tienatal a tote eadatataubesitars I -- ARACHNIDA * Araneae THERAPHOSIDAE Eury (Brachypelma) sp. (B.)...... I _ THOMISIDAE Epicadus heterogaster (Guérin) Cees Ai Dieta dice anita abc.e Baier meletetalt I —_ OPRTLIONES * COSMMETID ES | Se 95 2 eho sslinss. sft sekanelererers I — PEDIPALPIDA ** Tarantula palmata barbadensis Po- COGIG MOIS! J Wie ulate slam isuttel pave eo oye I — TAO DEDAE =, Amblyomma humerale Koch § (B.). I —— 86 Det. by Dr. A. B. Gurney. 37 Det. by Dr. J. Cooreman. MICROSCOPY VOL. 139 Comments 2K The microscopical features of the reactions which characterize the coagulation of the hemolymph in several supraspecific groups of insects (Orthopteroid Complex, Heteroptera, Homoptera, Scara- baeidae, Cerambycidae, Hymenoptera, Lepidoptera) have been de- scribed elsewhere (Grégoire, 1955a, pp. 109, III, 115, 118, 123 ; 1957, pp. 7, 27, 28; Grégoire and Jolivet, 1957, pp. 28-33). They were also observed in the corresponding groups of the present material. A few particular reactions will be briefly mentioned below. Phasmoptera.—As repeatedly pointed out (Grégoire, 1951, 1955a, 1957; Grégoire and Jolivet, 1957) the various categories of hemocytes are passively embedded in the coagulum initiated by the alterations NO. 3 HEMOLYMPIL COAGULATION IN INSECTS 15 in the fragile hyaline hemocytes or coagulocytes. Modifications of the plasma induced around the former corpuscles are exceptional. Such modifications, recorded previously in two specimens of Neotropical stick insects (Grégoire, 1957, p. 7), were observed in Prisopus cerosus (table 1) around macronucleocytes of small size (stem cells), sec- ondarily to the typical formation of islands of coagulation around the unstable hyaline hemocytes. Heteroptera——Granular precipitates, unrelated to the presence of hemocytes in the vicinity, recorded previously in the same group of insects, were observed in the present material in Montina lobata, Saica apicalis (Reduviidae), Macropygium reticulare, 3 species of Edessa (Pentatomidae), Anasa haglundi, Zoreva dentipes (Coreidae). A tentative interpretation of these occasional findings has been given elsewhere (Grégoire, 1957, p. 7). Coleoptera—The sequence in the alterations in the fragile hemo- cytes and in the plasma, characterizing pattern III (see Grégoire, 1957, p. 2 and text fig. 3), appeared with great clarity in the two specimens of Elateridae mentioned in table I. In the samples of hemolymph from Compsus sp., Heilipus sp., Exophthalmus jekelianus (Curculionidae), characterized, as shown in the table, by the absence of detectable alteration in the plasma, in the conditions of phase-contrast microscopy, a category of highly labile hemocytes, unrelated to the unstable hyaline hemocytes, underwent considerable modifications in their shape: immediately upon with- drawal and spreading out into films of the hemolymph, these hemo- cytes appeared spindle-shaped, with two straight expansions on both sides of the cell body. The expansions became progressively flexuous and exhibited continuous trepidations and jerks. They reached great lengths, bent suddenly at right angles, and sent out lateral ramifica- tions in various directions. Simultaneous development of such changes in neighboring hemocytes resulted in constitution of loose meshworks in wide areas of the preparations. Similar labile hemocytes have been reported in African weevils (Grégoire and Jolivet, 1957, p. 32) and in Diptera by Grégoire (1955a) and Jones (1956). In the present material they appeared in Ormuophasia bushkui (Diptera). Much smaller bipolar corpuscles, of unknown origin, unrelated to the labile elements described above, developed similar modifications. A detailed study of these corpuscles will be reported later. Arachnida. Araneae—In Epicadus and in Eurypelma, a category of hemocytes with coarse refractile granules scattered in their cyto- plasm and highly sensitive to foreign surfaces underwent disintegra- tion immediately upon shedding of the blood, in contrast to other 16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 categories of more resistant blood cells, such as macronucleocytes of small size (stem cells) and other kinds of granular hemocytes. A similar “differential sensitiveness’” has been formerly observed in extensive material of spiders (see Grégoire, 1955b). DISCUSSION DISTRIBUTION OF THE PATTERNS OF COAGULATION IN THE VARIOUS TAXONOMIC CATEGORIES OF INSECTS Detailed accounts on the relationships between pattern of coagula- tion of the hemolymph and taxonomic category have been given in previous papers (Grégoire 1955a, pp. 132-137; 1957, pp. 28-32; Grégoire and Jolivet, 1957, pp. 34-37). In this respect, the informa- tion obtained in the present material supports our former conclusions. With one exception (Carthaeomorpha rufipes, see below), the pattern detected in the samples of hemolymph collected in the present study (table, notes) from 50 specimens belonging to 30 neotropical species already investigated (1957), were identical to those recorded pre- viously. 1. Orthopteroid Complex. That broad group constitutes a highly homogeneous category with regard to the pattern consistently recorded at the specific and at the supraspecific levels. 2. Hemipteroid Complex. Hemiptera.—With the exception of Nepidae and Belostomatidae, studied previously (Grégoire, 1955a; Grégoire and Jolivet, 1957), all the specimens from 14 other families of Hemiptera investigated, including Reduviidae, Pyrrhocoridae (see 1955a), Coreidae, Gelasto- coridae, Pentatomidae, Miridae of the present (38 species) and of former materials, exhibited consistently the pattern IV. Homoptera.—The present material includes 41 species not investi- gated previously (Grégoire, 1955a, p. 110; 1957, pp. 15 and 16). Pattern I was predominant in Cicadidae, Fulgoridae, Dictyopharidae, Cercopidae, Cicadellidae, and was recorded in the only specimen of Membracidae captured, a family not yet investigated. In a few Cercopidae (see also 1955a, p. 110) and Cicadellidae, pattern I was associated with pattern II (= pattern III). A substantial coagulation of the hemolymph, developing rapidly, sometimes instantaneously, characterized these families, with the NO. 3 HEMOLYMPH COAGULATION IN INSECTS L7 exception of Cicadellidae, and was especially conspicuous in Fulgoridae. In Cicadellidae, the amount of clotted material varied greatly and appeared scarcer than in the other groups listed above. Pattern IV was observed in the samples of Cixiidae, Flatidae, and Issidae. However, in Flatidae, pattern I was found in Carthaeomorpha rufipes (table, note 11), a species in which pattern IV had been recorded previously in the only specimen available (Grégoire, 1957, p. 16). Pattern I appeared also incidentally in Anormelis mgrolim- bata and in Paradascalia nietvi. Pattern IV, observed to occur pre- dominantly in the few samples examined till now, is then questionable as being representative of Flatidae, a family which requires further investigation. 3. Coleoptera. The patterns predominant or representative in several groups formerly investigated were seen again in the present material: pat- tern II in Scarabaeidae (Rutelinae, Dynastinae), pattern III in Elateridae and in Tenebrionidae, pattern I in Meloidae (note 22), Cerambycidae (very substantial coagulation), pattern IV in Cur- culionidae. Pattern I, alone or associated with pattern II (= pattern III) was recorded in specimens of Cebrionidae and of Lymexylidae, two fami- lies not represented in our former data. In the other groups listed in the table, scarcity in the material, large variations at the individual, specific, and generic levels, already noticed previously, do not permit conclusions about the pattern predominant or representative of these groups. In this and in former studies (Grégoire, 1957, p. 22; Grégoire and Jolivet, 1957, pp. 22 and 23), absence or scarcity in clotting substances was observed in several specimens of Eumolpidae and of Cassidinae. In the present material, pattern III was recorded in one (Stilodes) out of 3 specimens of Chrysomelidae s.s., a family involving genera with obviously predominant patterns (see 1955, p. 114: Chrysolina, 7 species: patterns I and III; Timarcha, 5 species: patterns I and TU), 4. Panorpoid Complex. The present results are in agreement with former data with regard to Mantispidae (pattern IV: see Grégoire, 1957, p. 23), Sialodea: Corydalus sp. (pattern I, instantaneous reaction: see 1955a, p. I15: 18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 Sialis flavilatera L.) ; Trichoptera: Leptonema (pattern I: see 1955a, p. 116: Limnophilidae sp. and Anabolia nervosa Leach) ; larvae of Lepidoptera (pattern IT, see 1955a, pp. 116-118; 1957, p. 23; Grégoire and Jolivet, 1957, p. 25), and adult Diptera (Ormiophasia bushku: pattern IV, see Grégoire, 1955a, p. 121). As already pointed out, pattern I frequently characterizes insects belonging to relatively archaic orders (Plecoptera, see 1955a, p. 107; Megaloptera, 1955a, p. 115). 5. Hymenoptera. Patterns I and III are representative in several families of this order (Grégoire, 1955, pp. 122-123; 1957, pp. 24-26; Grégoire and Jolivet, 1957, p. 25). However, individual and specific variations may mask the representative pattern of the genus or of the family when only limited material is available. In the present (note 32) and previous materials (1957, p. 24) of Formicidae, a substantial pattern I characterizes the genus Paraponera. Patterns I and III were also recorded, though not consistently, in several specimens of the genus Camponotus (1955a, p. 123; 1957, p. 24; Grégoire and Jolivet, 1957, p. 25). On the other hand, no coagulation could be observed (pattern IV) in seven females of Azteca sp., from which the films of hemolymph were collected and prepared without interference of any artifact. The present observations on Vespidae (note 34), Pompilidae (note 35) and Sphecidae are in agreement with those made previously (pattern I and/or III: 1955a, p. 123; 1957, pp. 25-26). 6. Odonata. As in former studies (1955a, p. 107; 1957, p. 26), pattern IV was recorded in the only (adult) specimen of this order collected in the present material. 7. Arachnida. Coagulation of the blood was not detected in the present and former specimens of Pedipalpa, Ixodidae (1955b, pp. 497-498). Pattern IV was also recorded, in this and in previous studies, in specimens of Opiliones and of Brachypelma (Theraphosidae, Araneae), while other specimens of the latter genus exhibited pat- tern II, sometimes substantial, sometimes incomplete (see 1955), p. 495). NO. 3 HEMOLYMPH COAGULATION IN INSECTS 19 ON THE DISPARITIES IN THE REACTIONS OF COAGULATION OF THE HEMOLYMPH RECORDED AT THE SUPRASPECIFIC, SPECIFIC, AND INDIVIDUAL LEVELS 1. In contrast to the taxonomic categories characterized by a pat- tern of coagulation representative or predominant, other groups, especially Carabidae (Grégoire, 1955a, p. 111; 1957, p. 16; Grégoire and Jolivet, 1957, p. 12), exhibit such variations that, in spite of increased samplings, a representative pattern did not appear clearly in these groups at the family level, but provisionally at the generic or specific levels. In that respect, incidental coincidences may be deceptive and sug- gest erroneously that a pattern is characteristic of a genus, when it may actually represent an incidental failure of the true pattern to appear with all its particularities in a set of specimens being pro- visionally, at the time of capture, in similar abnormal conditions. For instance, in three specimens belonging to three different species of the genus Agra (Carabidae), pattern II, incomplete in two of these speci- mens, was predominantly observed in the present study, while formerly, in three other species of the same genus, pattern I had been consistently found (Grégoire, 1957, p. 16). Pattern III, possibly dissociated in the individual samplings into its two components (patterns I and II), might be the representative pattern of the genus Agra. Other examples are furnished in Hymenoptera in the genera Eciton (1957, p. 24) and Azteca (table), in which the predominant patterns are possibly not the actual ones. In families such as Lycidae, Lampyridae, Coccinellidae, Chryso- melidae (Cosmogramma), and Cassidinae (Cyclostoma), the observa- tions were handicapped by the presence in the hemolymph of particles floating in considerable numbers, a finding already noticed (1955a, p. 106; 1957, p. 27; Grégoire and Jolivet, 1957, p. 30). 2. Divergences at the specific or individual level recorded in genera characterized by a pattern predominant or representative, appear, for instance, in specimens of Cicadellidae. However, the pattern char- acterizing the group was found incidentally in the samples (see under comments in the table). At the individual level, pictures of another pattern were recorded incidentally in limited fields of preparations exhibiting a predominant pattern (Reduviidae: Stenopoda, Rasahus, Dysdercus; see also 1955a, p. 109; 1957, p. 13; Grégoire and Jolivet, 1957, pp. 10-11). Tentative interpretation of these divergences have been presented elsewhere (1955a, pp. III, 124, 126; 1957, discussion; Grégoire and 20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 Jolivet, pp. 36 and 37). Artifacts of preparation are responsible for a part of the pictures recorded. Nutritional balance of the specimens at the time of capture, seasonal and pathological conditions, able to alter the sensitivity of the unstable hemocytes or the amounts of the coagulable substances in the hemolymph, are among the factors which might explain these discrepancies: change in the pattern of coagula- tion has been observed in infected insects belonging to species or to groups characterized in their normal conditions by another pattern (Acrididae, Dermaptera, Cerambycidae) (see Grégoire and Jolivet, 1957, p. 36). Similarly, in a specimen of Gypona hebes from the pres- ent material, exhibiting pattern IV (table, comments), the unstable hemocytes responsible for the inception of the coagulation contained unusual coarse granules, absent in the other normal specimens in which the pattern representative of the group was observed. The present results support former conclusions (1957, p. 30) that the patterns of coagulation are not individual particularities, but rather characterize species, more frequently supraspecific categories. DIVERGENCES BETWEEN NEOTROPICAL MATERIAL AND INSECTS FROM THE OLD WORLD In 10 specimens belonging to 6 species of Neotropical Passalidae (1957, p. 18, and here, table 1), pattern I was recorded exceptionally in one sample from a single species, while this pattern, unmixed or associated with pattern II (= pattern III), appeared in the 5 African species (25 specimens) available (Grégoire and Jolivet, 1957). Pattern I, absent from the samples of Neotropical Coprinae (4 spe- cies, 8 specimens), was found, alone or associated with pattern II = pattern III), in 12 (29 specimens) out of 17 African species examined (Grégoire and Jolivet, 1957), and was questionable in three other species (5 specimens). These data might suggest the possibility of discrepancies, with regard to these two families, between Neotropical and Old World material. However, as already pointed out (Grégoire, 1957, p. 32), large individual variations characterize these families, especially Passalidae. Numerous samplings from insects of both origines, and belonging to genera and species more closely related than those available, are required before any conclusion might be drawn about the existence of such discrepancies. SUMMARY 1. Coagulation of the hemolymph from 400 (mostly adult) speci- mens, belonging to 215 Neotropical species of insects, and including NO. 3 HEMOLYMPH COAGULATION IN INSECTS 21 185 species not yet investigated, has been observed on films in vitro by phase-contrast microscopy. In that material, the pattern of coagu- lation predominant in the samples or representative for the species or for the supraspecific taxonomic category has been recorded. 2. The material contained insects from 14 families poorly (Dictyo- pharidae, Cercopidae, Cicadellidae, Flatidae) or not (Gelastocoridae, Membracidae, Cixiidae, Issidae, Cebrionidae, Lymexylidae, Erotyli- dae, Hispidae, Corydalidae and Larvaevoridae) represented in pre- vious studies. 3. Additional information obtained for the present paper was con- sistent with former data, with regard to the pattern predominant or representative, in the Orthopteroid Complex, in several families of Heteroptera (Reduviidae, Pyrrhocoridae, Coreidae, Pentatomidae, Miridae), of Homoptera (Cicadidae, Fulgoridae, Dictyopharidae, Cercopidae, Cicadellidae), of Coleoptera (Scarabaeidae, Elateridae, Tenebrionidae, Meloidae, Cerambycidae, Curculionidae), of Hymen- optera (Formicidae, Vespidae, Sphecidae). 4. In the families not represented in former investigations, pat- tern I was recorded in specimens of Cebrionidae and of Lymexylidae (Coleoptera). 5. Pattern I was also observed in specimens of Corydalidae (Sialo- dea) and of Hydropsychidae (Trichoptera), in agreement with pre- vious results on palearctic representatives belonging to these groups. 6. Divergences in the reactions of coagulation observed in the present and in a former study between Neotropical and African Passalidae and Copridae (Coleoptera) require further investigations on more extensive material, owing to the large variations existing in these groups of insects. 7. The reactions of the blood in vitro observed in five specimens of Arachnida (Araneae, Ixodidae, Opiliones, Pedipalpa) are briefly men- tioned in relation to previous results on more extensive material. ACKNOWLEDGMENTS I am grateful to Dr. Carl B. Koford, former resident naturalist of the Canal Zone Biological Area, for invaluable help in supplying me rapidly with adequate equipment. I also wish to express my thanks to Mrs. A. Gomez, administrative assistant at the station, who, as usual, was very cooperative in arranging for living accommodations in the Canal Zone. I am greatly indebted to Dr. Ing® Oswaldo Vargas Gonzales, head of the Seccién Entomologia de la Estacion Experimental Agri- cola en Tingo Maria (Peru), for authorization to carry on my work 22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 in his department. I acknowledge with sincere appreciation the assist- ance of Dr. Ing® Matias Reyes Valdivieso for rapid and clever adjustment of my optical equipment to one of the microscopes of the department. I wish to thank Dr. Remington Kellogg, Director of the United States National Museum; Dr. Waldo L. Schmitt, former head curator of the department of zoology, for authorization to have the material determined in the Museum; and Dr. J. F. Gates Clarke, curator of the division of insects, for collecting and mailing the data. I also wish to express my gratitude to the following specialists for identifica- tion of the specimens: O. L. Cartwright, Dr. E. A. Chapin, Dr. J. Cooreman, G. Fagel, W. D. Field, Dr. A. B. Gurney, Dr. Ch. Jeuni- aux, K. V. Krombein, Dr. V. Lallemand, Dr. J. C. Lutz, Miss S. Par- fin, Dr. J. G. Rozen, Miss Louise M. Russell, C. W. Sabrosky, Dr. M. R. Smith, T. J. Spilman, George B. Vogt, Miss Luella L. Walkley, Miss Rose Ella Warner, Dr. C. Willemse, Dr. D. A. Young. REFERENCES Grécorre, CH. 1951. Blood coagulation in arthropods. II. Phase-contrast microscopic observations on hemolymph coagulation in sixty-one species of insects. Blood, vol. 6, pp. 1173-11098. 1953. Blood coagulation in arthropods. III. Reactions of insect hemolymph to coagulation inhibitors of vertebrate blood. Biol. Bull., vol. 104, PP. 372-393. 1955a. Blood coagulation in arthropods. V. Studies on hemolymph coagu- lation in 420 species of insects. Arch. Biol., vol. 66, pp. 103-148. 1955b. Blood coagulation in arthropods. VI. A study by phase-contrast microscopy of blood reactions in vitro in Onychophora and in various groups of arthropods. Arch. Biol., vol. 66, pp. 489-508. 1957. Studies by phase-contrast microscopy on distribution of patterns of hemolymph coagulation in insects. Smithsonian Misc. Coll., vol. 134, Pp. 1-35. GREGOIRE, CH., and FLorKin, M. 1950. Blood coagulation in arthropods. I. The coagulation of insect blood, as studied with the phase-contrast microscope. Physiol. Comp. et Oecol., vol. 2, pp. 126-139. Grécorre, Cu., and JoLivet, P. 1957. Coagulation du sang chez les arthropodes. VIII. Réactions du sang et de l’hémolymphe in vitro, étudiées chez 210 espéces d’arthropodes africains. Inst. Parcs Nat. Congo Belge. Expl. Parc Nat. Albert, sér. 2, fasc. 4, pp. 1-42. Harpy, W. B. 1892. The blood corpuscles of the Crustacea, together with a suggestion as to the origin of the crustacean fibrin-ferment. Journ. Physiol., vol. 3, pp. 165-190. NO. 3 HEMOLYMPH COAGULATION IN INSECTS 23 Jones, J. C. 1956. The hemocytes of Sarcophaga bullata Parker. Journ. Morphol., vol. 99, pp. 233-258. NuMANOI, H. 1938. On crustacean blood coagulation. Japan. Journ. Zool., vol. 7, pp. 613-641. sear 1910. Crustacean blood coagulation as studied in the Arthrostraca. Quart. Journ, Exper. Physiol., vol. 3, pp. 1-20. 1911. Types of crustacean blood coagulation. Journ. Mar. Biol. Assoc., vol. 9, pp. 191-108. Tait, J., and Gunn, J. D. 1918. The blood of Astacus fluviatilis: a study in crustacean blood, with special reference to coagulation and phagocytosis. Quart. Journ. Exper. Physiol., vol. 12, pp. 35-80. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 139, NUMBER 4 A REVIEW OF THE GENUS HOPLOMYsS (THICK-SPINED RATS), WITH DESCRIP- TION OF A NEW FORM FROM ISLA ESCUDO DE VERAGUAS, PANAMA By CHARLES O. HANDLEY, JR. Associate Curator, Division of Mammals United States National Museum Smithsonian Institution (PUBLICATION 4380) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION JULY 3, 1959 THE LORD BALTIMORE PRESS, INC. BALTIMORE, MD., U. S. A. A REVIEW OF THE GENUS HOPLOMYS (THICK- SPINED RATS), WITH DESCRIPTION OF A NEW FORM FROM ISLA ESCUDO DE VERAGUAS, PANAMA By CHARLES O. HANDLEY, JR. Associate Curator, Division of Mammals United States National Museum Smithsonian Institution A specimen of the thick-spined rat, Hoplomys gymnurus Thomas, that Alexander Wetmore shot in a thicket on Isla Escudo de Veraguas on the morning of March 1, 1958, is probably the only mammal from this Caribbean island that is preserved in a museum. Other rats that Wetmore saw in coconut palms on the same day apparently were of another genus. No other mammals have been reported from this locality except feral hogs. Although Indians once lived on the island, human beings are now only transients there. Escudo de Veraguas is a low island, about 1 mile wide and 2.5 miles long, in the Caribbean Sea, 11 miles off the base of the Valiente Peninsula, Province of Bocas del Toro, north coast of the Republic of Panama. Wetmore (Smithsonian Misc. Coll., vol. 139, No. 2, 1959) has given a detailed account of the history, geography, and zoological position of the island. Other echimyid genera, Diplomys and Proechimys, are known to occur on certain islands in the Gulf of Panama and elsewhere, but no insular populations of Hoplomys have been reported. The Escudo de Veraguas Hoplomys differs in so many respects from other known populations of the thick-spined rat that it has prompted a brief review of the genus. Many of the National Museum (US) specimens reported here were collected in cooperation with the Gorgas Memorial Laboratory, Panama. I express my thanks to Carl Johnson, director, and other members of the laboratory staff for numerous courtesies and assistance in fieldwork. Some of the specimens were collected by C. M. Keenan of the Army Preventive Medicine Survey Detachment, Ft. Clayton, Canal Zone. Richard Van Gelder kindly permitted the study of speci- mens in the American Museum of Natural History (AMNH), New York. SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 139, NO. 4 A «J ¥! f \ mic i |i 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 Genus HOPLOMYS J. A. Allen 1908. Hoplomys J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 24, p. 649. Genotype.—Hoplomys truet J. A. Allen. Distribution—The genus has a limited distribution in Central America and northwestern South America. It is monotypic. Published records of collecting localities are mapped in figure 1. Hoplomys is known to occur at medium elevations (800-3,100 ft.) on the Caribbean slope of the highlands of Nicaragua and Costa Rica; near sea level on the Caribbean coast of Panama; at medium elevations (600-4,000 ft.) on the Pacific slope of eastern Panama, Colombia, and Ecuador ; and near sea level in extreme southwestern Colombia and north- western Ecuador. The distribution of Hoplomys in South America appears to be limited by the Western Andes. J. A. Allen’s record for Puerto Valdivia on the Rio Cauca (Bull. Amer. Mus. Nat. Hist., vol. 35, p. 207, 1916) is erroneous (the specimen is a Proechimys). Proechimys cayennensis hoplomyoides Tate (Bull. Amer. Mus. Nat. Hist., vol. 76, p. 179, 1939) from Mt. Roraima, Venezuela, appears not to be a Hoplomys, although a relationship has been suggested (Moojen, Univ. Kansas Publ., Mus. Nat. Hist., vol. 1, p. 324, 1948). In the Caribbean lowlands of Panama, where Proechimys is abun- dant and Hoplomys seemingly rare, I have trapped individuals of both genera under the same log on successive nights. At medium alti- tudes in the mountains of Panama where Hoplomys is fairly common, Proechimys apparently does not occur. All the Hoplomys that I have collected in Panama were caught in banana-baited live traps under large decaying logs—in fairly open mature rain forest, in grassy clearings and adjacent streamside thickets, and in dense, hillside Heliconia thickets. Goldman (Smith- sonian Misc. Coll., vol. 69, No. 5, p. 124, 1920) found Hoplomys associated with fallen trees and rocks in Panamanian forests. Diagnosis ——Dorsum, flanks, and rump, in both adult and juvenile pelages, with spines 26 to 33 mm. in maximum length and 1.5 to 2.0 mm. in maximum diameter, tending to obscure soft fur. Tail shorter than head and body, scaly, and sparsely haired. Ears scantily haired. Hind feet long and narrow; fifth toe scarcely longer than first; claws relatively straight, but claw of second toe slightly expanded. Skull prominently ridged, and supraorbital shelf beaded ; rostrum relatively broad at tip ; auditory bullae relatively small; and infraorbital foramen without subsidiary canal on floor, and with external wall thin in lateral view. Cheek teeth with oblique folds; counterfold formula normally 4/4-4/4-4/4-4/4, rarely 4/4-4/3-4/3-4/3. NO. 4 HOPLOMYS (THICK-SPINED RATS )—-HANDLEY 3 V aviation Specimens of Hoplomys have never before been availa- ble in series. Fourteen specimens, seven of which are adult, recently collected on Cerro Azul, Panama, now permit a fairly good estimate RiO COCO BIJAGUA SAVALA GATUN RfO INDIO FT. SHERMAN CERRO AZUL RiO CANGANDI” CANA MT. TACARCUNA SUERRE STA. TERESA PERALTA ESCUDO DE VERAGUAS ALTO BONITO BAGADO RIO TAMANA. LA GUAYACANA BARBACOAS BUENAVISTA SAN JAVIER CACHABI- Fic. 1.—Distribution of Hoplomys gymnurus. All known specimen localities are indicated. of individual variation in the genus. Eleven specimens from Darien and nine from the Canal Zone, are also helpful. In addition, random series of up to 75 specimens per sample of the closely related Proechimys semispinosus have been used to evaluate the variations seen in the smaller series of Hoplomys. 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 Size, flatness, and ridging of the skull increase with advancing age in Hoplomys and Proechimys. Tooth wear appears to be a reliable criterion of age. Full adult pelage usually is attained after M3 appears and before it becomes functional. Only juvenile and adult pelages have been distinguished. Specimens in which all cheek teeth are functional are considered to be adults. Generally, the largest, flattest, most heavily ridged skulls have the most worn teeth. Apparently these rodents continue to grow after all teeth are functional. Thus, there is considerable size spread among adult skulls. For this reason only maximum and minimum figures are given in the table of measurements (p. 6). Body sizes appear to be uniform throughout the mainland range of the species, but larger on Escudo de Veraguas. The skull is narrow in the south—Ecuador, Colombia, and Darién—somewhat broader in central Panama, Costa Rica, and Nicaragua, and broadest of all on Escudo de Veraguas. Likewise the nasals and cheek teeth are smaller in the southern populations. Size of the auditory bullae increases northward from Ecuador to Nicaragua, but the bullae are largest and most inflated anterolaterally in the Escudo specimen. Several features of the zygomatic arches vary geographically. The maxillary roots of the zygomata flare less widely and less perpendicu- larly from the longitudinal axis of the skull (so that the zygomata are more convergent anteriorly) from the Canal Zone southward than they do in the north. From Cerro Azul southward the maxillary roots tend to flare up, away from the ventral plane of the skull, rather than paralleling that plane as they do in the north. The jugal has a hooklike posteroventral process in most Canal Zone and Cerru Azul specimens, but not in others. Most of the specimens from Ecuador, Colombia, and Darién, and a smaller percentage of the central Panamanian specimens have a small conical projection on the dorsal edge of the zygoma at the jugal-squamosal suture. I failed to check this character in the Costa Rican and Nicaraguan specimens. There is hardly a trace of it in the Escudo individual. The nasals, broad and posteriorly truncate in the island specimen, are usually narrower and posteriorly acute in mainland populations. Among mainland populations of Hoplomys flatness and ridging of the skulls of mature individuals are similar to these features in mature individuals of Proechimys semispinosus. None of the available Hoplomys or Proechimys closely approaches the Escudo specimen in flatness or ridging, despite the fact that the island specimen, judged by tooth wear, is a prime adult, not as old as many individuals with NO. 4 HOPLOMYS (THICK-SPINED RATS )—-HANDLEY 5 which it was compared. The degree of reduction of dorsal doming and ventral depression of the brain case of the Escudo specimen is reflected in the convergence of greatest and condylobasal lengths of the skull, and in the more posteriorly oriented (as opposed to ventrally oriented) foramen magnum. The thick spines that distinguish Hoplomys are longest and strongest just behind the shoulders on the upper midback, from which point they diminish in size in all directions. The spines possibly vary geographically in size. They appear to be longer and stronger toward the southern part of the range of Hoplomys. The Escudo specimen, although it is larger than any other, has the smallest and weakest spines. Maximum spine length varies as follows (mean, followed by extremes): 6 Ecuador 29 mm. (28-31), 4 Darién 30 (28-33), 11 Cerro: Azul 28) (26-29), 5 Canal Zone’ 28 (27-30), 1 Escudo de Veraguas 26. Coloration of the spines is individually variable. All specimens have all spines proximally white and distally colored. The tips of those of the dorsum are always black, but the flank spines usually are tinged with orange or banded with orange and black distally. Occasionally the flank spines are colored like the dorsal spines. Coloration of the soft hairs of the dorsum is geographically variable. At the southern extreme they are reddish orange, especially on the shoulders. The soft hairs of the Escudo specimen are similar but darker and brighter. Costa Rican and Nicaraguan examples have the hairs more orange, and those from Panama and northern Colombia are more yellowish on the average. The presence or absence of black ocular and crown areas appears to be individually variable throughout the range of Hoplomys, but only the Escudo specimen has the soft hairs blackened to form a distinct middorsal stripe from snout to base of tail. All populations have the underparts dominantly white, and all have some individuals that show encroachment of agouti hairs of the side neck onto the throat, suggesting an incipient collar. This is well marked in the Escudo specimen; one from Rio Indio, Canal Zone, has a complete collar. Nine of the 14 Cerro Azul specimens have clear orange collars, and several of them have a band of clear orange hairs separating the agouti hairs of the flanks from the white hair of the belly. Neither of these features is seen in samples of other popu- lations. Coloration of the forefeet (usually white on the inner side, colored on the outer, occasionally colored throughout), and coloration of the cheeks (clear orange, buff, gray, or agouti) are individually variable. 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Cze Cec g'Sz q33u9] [ese Ny giz g'0z I'1z-£'0z ¥'7Z-Q'0z v1z—-9'0z Lze-0'1z g'1z-9'1z Q'ez-1'1z £77 g'0z r'bvz qypessq [eprloyse yy F'O1 Lol PeiI-LeI ZLEI-1'€1 Pei ei Ivi-S'E1 I°€I-1'f1 ge1-o'f1 I'vi CCI o'vi zSI MIRE t [e}iq1010}UT “pamuyuoj—I ATaVT, ‘sq JO Wooaye jo sursieur 19yno0 usemzog | £92 0'9z bgz—-h' Lz 982-892 1°6z-€°9z 6°0£-L'9z 9'6c-b'6z 0'0£-7'9z Toe £°92 £32 Eze qypessq d1}EUIOBAT “qysuey, AL[OOATY » bette teen ee eeeeeeeees greumay I cee eee ee ee eee ese eee eee ayeul I Jopendy “MN pue erquiojod “A'S eee eee wee wee ew eee SoTeUloy Z see eeeeeeeeeeeesereees sore g eweueg ‘Usieqd ere eee ee ee eee eee eee soyemoy c Sitelole elfeiipib rails terete Valtadslaheteteltenie soyeu S eweueg qnzy O19 "+" SoreUaz Z wee ee ee eee ee eee eee eee sayeur © eureueg ‘ouoZ yeued) reese ee eeeeeeeeeeeeeres QrpUT T BINT eBSOD ‘eyesag esatay, “eS ee weer eee eer eee eee wees od] euloy I “* oyeut I enseiesin ‘enselig Ce ee Pees scene eeeeeeeeeeeees oreUT T eureueg ‘senseiaA op opnosy 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 The large size of the Escudo specimen, the massiveness, broadness, and heavy ridging of its skull, the inflation of its auditory bullae, and its distinctive coloration all seem to be beyond the possibility of individual variation. This suggests that the Escudo animal is taxo- nomically distinct from mainland populations. That it is conspecific with them is indicated by its alignment with some of the morphologi- cal clines observed in the mainland populations. Wetmore (of. cit.) has named three birds (a wren, a manakin, and a tanager) collected on Escudo de Veraguas that differ from their mainland counterparts in greater size, in addition to differences in color. Classification—The genus Hoplomys is represented by one species, which includes four subspecies : HOPLOMYS GYMNURUS GOETHALSI Goldman 1912. Hoplomys goethalsi Goldman, Smithsonian Misc. Coll., vol. 56, No. 36, p. 10 (Rio Indio, near Gatun, Canal Zone, Panama). Characters —Size medium; skull of medium width and ridged; brain case domed and slightly depressed ; foramen magnum ventrally oriented ; cheek teeth large; auditory bullae medium; zygomata con- verging conspicuously anteriorly, and maxillary root tending to flare up slightly from ventral plane of skull; jugal with hooklike postero- ventral process and small conical posterodorsal projection ; nasals long, narrow, and posteriorly acute; dorsal spines long and strong; soft hairs of dorsum appear uniform yellowish orange in mass effect. Specimens examined.—Panama: Cana, 2,000 ft., 5 US; Cerro Azul; 2:100 it:., 14 US; Ft? Sherman,.4 US; Gatun, 3 (2 AMNE, 1 US) ; Cerro Tacarcuna, 2,650 ft.,6 AMNH; Rio Cangandi, 200 ft., t US; Rio Indio, 2 US. Colombia: Alto Bonito, Antioquia, 1,500 ft., 1 AMNH;; Bagado, Choco, 600 ft., 2 AMNH. Additional published records——Colombia: Rio Tamana, branch of the Rio San Juan, Choco (J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 35, p. 207, 1916). HOPLOMYS GYMNURUS GYMNURUS Thomas 1897. Echimys gymnurus Thomas, Ann. Mag. Nat. Hist., ser. 6, vol. 20, p. 550 (Cachabi, N. Ecuador, alt. 560 ft.). Characters.—Size medium or small; skull narrow and ridged ; brain case domed and slightly depressed; foramen magnum ventrally oriented; cheek teeth small; auditory bullae small; zygomata con- verging conspicuously anteriorly, and maxillary root flaring up from ventral plane of skull; jugal lacking posteroventral process, but with NO. 4 HOPLOMYS (THICK-SPINED RATS )—-HANDLEY 9 prominent conical posterodorsal projection; nasals short, narrow, and posteriorly acute; dorsal spines long and strong; soft hairs of dorsum giving reddish-orange mass effect, slightly darkened on shoulders. Specimens examined—Colombia: Barbacoas, Narinmo [75 ft.], 8 AMNH;; Buenavista, Narifio [1,200 ft.], 1 AMNH; La Guayacana, Narino, 800 ft., 2 US. Ecuador: Mindo, Rio Blanco [4,000 ft.], 1 AMNH;; San Javier, 60 ft., 7 (1 AMNH, 6 US). Additional published records —Ecuador : Cachabi, 560 ft. (Thomas, Op Cit pe 551). HOPLOMYS GYMNURUS TRUEI J. A. Allen 1896. Echimys semispinosus Alfaro (not Tomes, 1860, Proc. Zool. Soc. Lon- don, p. 265), Primera Exposicion Centroamericana de Guatemala, Museo Nacional, San José, p. 41 (Suerre, Costa Rica). 1908. Hoplomys truei J. A. Allen, Bull. Amer. Mus. Nat. Hist., vol. 24, p. 650 (Savala, Matagalpa Prov., Nicaragua). Characters —Size medium; skull of medium width and ridged; brain case domed and slightly depressed ; foramen magnum ventrally oriented; cheek teeth large; auditory bullae large; zygomata con- verging less anteriorly than in goethalsi, and maxillary root in plane of ventral surface of skull; jugal without hooklike posteroventral process; nasals long, narrow, and posteriorly acute; dorsal spines relatively short and weak; soft hairs of dorsum giving uniform dark orange mass effect. Specimens examined.—Nicaragua: Lavala [ = Savala, 800 ft., along the inner border of the low east coast region], 2 AMNH; Rio Coco [800 ft.], 2 AMNH; Vijagua [= Bijagua, probably 1,500 to 2,000 ft., on eastern slope of highlands in Matagalpa Prov.], 3 AMNH. Costa Rica: Santa Teresa Peralta [3,100 ft.], 1 AMNH; Suerre, 1,500 ft. [near Jiménez], 1 AMNH. Additional published records.—Tate (Bull. Amer. Mus. Nat. Hist., vol. 68, p. 401, 1935) supposed that True’s record (Proc. U. S. Nat. Mus., vol. 11, p. 467, 1889) of Echinomys semispinosus in Nicaragua was the first published reference to a Hoplomys. The specimens, still in the U. S. National Museum, however, are Proechimys. HOPLOMYS GYMNURUS WETMOREI subsp. nov. Holotype —U.S.N.M. No. 307057; adult male, skin and skull; collected March 1, 1958, by Alexander Wetmore; Isla Escudo de Veraguas, Prov. Bocas del Toro, Panama; original No. 1479. 10 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 Characters.—Size large; skull broad and heavily ridged; brain case flattened dorsally and ventrally; foramen magnum posteriorly oriented ; cheek teeth large ; auditory bullae large and inflated antero- laterally ; zygomata converging less anteriorly than in goethalsi, and maxillary root in plane of ventral surface of skull ; jugal without hook- like posteroventral process, or conical posterodorsal projection ; nasals long, broad, and posteriorly truncate; dorsal spines relatively short and weak; and soft hairs of dorsum giving dark reddish-orange mass effect (between Burnt Sienna and Sanford’s Brown of Ridgway, 1912, Color Standards and Color Nomenclature), with black middorsal stripe from snout to base of tail. For measurements see table 1. Specimen examined.—The holotype. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 139, NUMBER 5 Charles DB. and Mary Waux Walcott Research Fund GENERA OF TERTIARY AND RECENT RHYNCHONELLOID BRACHIOPODS (Wit 22 PLatTEs) By G. ARTHUR COOPER Head Curator, Department of Geology United States National Museum Smithsonian Institution { nov 25 1959 \ AS we dif Pe fi RN CITY OF WASHINGTON : PUBLISHED BY THE SMITHSONIAN INSTITUTION NOVEMBER 23, 1959 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 139, NUMBER 5 Charles D. and Mary Waux CA alcott Research Fund Gene Or. TERTIARY AND: RECENT RHYNCHONELLOID BRACHIOPODS (WitH 22 PLATES) By G. ARTHUR COOPER Head Curator, Department of Geology United States National Museum Smithsonian Institution (PUBLICATION 4382) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION NOVEMBER 23, 1959 SMITHSONIAN , INSTITUTION NOV 2 3 195¢ THE LORD BALTIMORE PRESS, INC. BALTIMORE, MD., U. S. A. CONTENTS Page AEE ORUICELOEN i repare cleats eri ae ware) at oisn-w 6 (a,c) sta dk Sater y wataieranet dveraralcecteratartvarc 5 stig Chee pedestal ge ese eels aks ns OG 64 RR TRVIAECHISDECICS 7 cries oickess ahora reesreist sso cs af nro, oe 0s ty ecaeta eam a Re ee aaeral as 66 REL ETC OSH rere oreo VS eee EPA Ree owes Sears EES ne ors (SPST ecco are ea eI RC aietie ole et a iene 67 Perera AM TREGEN MOE! PALES ogc. oes) S516 cis wre oo eel ave te 3 sos eve waaay ston eer afecaues slahaie 73 ui Wis bes ete Pa ae hak Hey ie Coatin isin 1, UCLA. 7257. Eocene (Domengine formation), from section line, 2,600 feet south of the northeast corner of sec. 20, T. 28 S., R. 19 E., M. D. B. and M., near headwaters of west branch of Agua Media Creek, McKittrick Quadrangle, Temblor Range, California. BeSphenarinastcula (Sestenza)'h. Sakae cee ec weac ates cae ectemeaes eons Figs. 6-15. 6-10, Respectively dorsal, posterior, ventral, side, and ante- rior views of a well-preserved complete specimen, I, showing the strongly triangular outline, hypotype U.S.N.M. 549353a. 11, Dorsal view of the preceding showing the foramen and obscure radial lines, X 2. 12, Posterior of the same specimen, X 4, showing alate conjunct deltidial plates. 13, 14, Two views of the interior of another specimen showing the cardinalia, <4, with their small outer hinge plates, dental plates and lack of a median ridge or septum in the brachial valve, hypotype U.S.N.M. 540353c. 15, Enlargement of the shell surface, <6, showing the delicate radial lines, hypotype U.S.N.M. 549353. Pliocene, Milasso, Messina, Sicily. CG. Cryptopora ‘gnomon \( Jetireys) ec keiak eater ees eases cee ceee keene Fig. 16. Brachial valve tilted to the side, & 8, to show the high median septum and the long maniculifer crura with the distal end handlike and with fingers extended, paratype U.S.N.M. 94367. Recent, 780 fathoms, off Cuba. D. Tegulorhynchia. Teguiorhynchia ‘squamasa’ (CEMUton) ic... ere inte ss oa he's we eictee s tan eee Figs. 17-24. 17-20, Respectively ventral, dorsal, anterior, and side views of a complete specimen, 1, hypotype U.S.N.M. 89855a. 21-23, Respectively dorsal, anterior, and side views of the preceding speci- 30 62 50 NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER Ti Page men, X 2, showing the imbricating ornament. 24, A partially exfoli- ated pedicle valve showing the impression of the muscle scars, X 2, hypotype U.S.N.M. 80855b. Miocene (Ototaran), Broken River, Trelissick Basin, Canterbury, New Zealand. T equlorhynchia, daderlemus (Davidson)... «0000 (Dall) osents assaeiaie: sya oy0in.0 15s caps Seales slaispeigr eel aolnie 35 Figs. I-11. 1-3, Respectively anterior, side, and dorsal views of a com- plete specimen showing anterior costation, 1, lectotype U.S.N.M 111086. 4, 5, Respectively full and tilted views of the interior of the pedicle valve showing pallial sinuses, small conjunct deltidial plates, deeply incised muscle scars, and pedicle collar, X 2. 6, Posterior of the pedicle valve of the lectotype, X 3, showing small foramen, conjunct deltidial plates, and corrugated teeth. 7-9, Respectively anterior, full, and partial side views of the interior of the brachial valve showing cardinalia, deeply impressed adductor field, and pallial sinuses, XX 2. 10, Partial side view of the falcifer crura showing serrate distal ex- tremity and corrugated socket, X 4. 11, Posterior view of the same NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 81 Page brachial valve, X 4, showing curved socket ridges, modestly developed outer hinge plates, incipient inner hinge plates, and falcifer crura. Recent, Sealark Expedition, 1905, Station Cl, 123 to 158 fathoms, south of the Saya de Malha Banks, Indian Ocean. B. Bassitola’ beechers: CD allys v5 sec h cme eee ae eee Caen aks 25 Figs. 12-16. 12-14, Respectively side, anterior, and dorsal views of a complete specimen showing the smooth exterior fold, & 1, hypotype U.S.N.M. 334678. 15, Dorsal view of the same specimen, XX 2. 16, Posterior of the preceding specimen showing conjunct, auriculate deltidial plates, 3. Recent, U. S. Bureau of Fisheries Station 4130, 283 to 309 fathoms, 46.1° F., near Kauai Island, Hawaiian group. Figs. 17-23. 17, 18, Respectively tilted and full views of the pedicle interior of another specimen showing deeply impressed muscle scars and pallial sinuses, X 2, hypotype U.S.N.M. 334679. 19, Posterior of the preceding pedicle valve, X 4, showing corrugated teeth, conjunct and auriculate deltidial plates. 20, 21, Respectively full and anterior views of the brachial valve of the preceding pedicle valve showing the cardinalia and pallial marks, & 2. 22, Posterior view of the cardinalia of the preceding specimen showing socket ridges, corrugated sockets, broad outer hinge plates, and falcifer crural plates, <4. 23, Side view of the preceding specimen showing the broad, distally serrate, falcifer crura and the corrugated sockets, X 4. Recent, 147 to 198 fathoms, 49° F., off west coast of Hawaii. PLATE 12 Basiliola Basiolaiambholos Vallix Saxktal. Ao: Sates Wale Alay wana s tae s 27 Figs. 1-6. 1-5, Respectively dorsal, posterior, anterior, ventral, and side views of the lectotype, 1, showing robust form, smooth exterior and strong dorsal fold, U.S.N.M. 229301b. 6, Beak of the preceding specimen, X 3, showing small round foramen and auriculate deltidial plates. : Recent, U. S. Bureau of Fisheries Station 5592, 305 fathoms, 43.3° F., gravel and mud bottom, Sibuko Bay, south of Silungan Island, Borneo. Figs. 7-15. 7, Posterior of the pedicle valve of another specimen, X 4, showing corrugated teeth, and conjunct and auriculate deltidial plates, hypotype U.S.N.M. 274135. 8, 9, Respectively full and tilted views of the preceding pedicle valve, X 2, showing dental plates, muscle area and pallial sinuses. 10, Interior of the apex of the preceding pedicle valve, X 4, showing pedicle collar, auriculation of deltidial plates, corrugated teeth, and small genital areas. 11, 12, Tilted and full views of the interior of the brachial valve of the preceding specimen showing cardinalia, pallial sinuses, and muscle scars, X 2. 13, 14, Two views of the apex of the preceding brachial valve tilted to show the cardinalia in partial side and partial anterior position, the strongly corrugated sockets, and the broad outer hinge plates, X 4. 15, Poste- 82 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 Page rior view of the cardinalia of the same brachial valve as the preceding, > 4, showing the broad and flat outer hinge plates. The crura are shorter than normal because of slight breakage at the distal extremity. U. S. Bureau of Fisheries Station 5487, 585 fathoms, 52° F., on mud, off Panaon Island, Philippines. PHATE SIG Hispanirhynchia? and Neohemithyris (=Basiliola) A... Hispanirhynéina?.. species ii ss socds nea 9 Bis ooisere Roe OER eee 60 Figs. 1-5. I-3, Respectively ventral, dorsal, and anterior views of a somewhat crushed specimen, X 1, figured specimen U.S.N.M. 540361. 4, Dorsal view of the preceding specimen, X 2, showing foramen. 5, Beak of the preceding, & 4, showing foramen and disjunct deltidial plates. Eocene (lower), 200 meters south of the south side of the River- side Yacht Club, west side Almendares River, in Reparto Kohly, Ha- bana Province, Cuba. B. Bastholas lucida CGould) (0 ee Ree a Re eee 34 Figs. 6-23. 6-10, Respectively posterior, anterior, dorsal, side, and ven- tral views of a complete specimen, 1, showing small size, rounded form, and nearly smooth exterior, hypotype U.S.N.M. 110826a. 11-13, Respectively anterior, side, and dorsal views, x 2, of the preceding specimen showing the same features. 14, Interior of the pedicle valve, 2, showing pallial marks indistinctly, hypotype U.S.N.M. 110826b. 15, Apex of same specimen, X 4, showing corrugated tooth and con- junct deltidial plates. 16, The same specimen tilted to show the pedicle collar and dental plates, X 4. 17, 18, Interior and tilted views of another pedicle valve showing the pallial marks, foramen, teeth, and deltidial plates, & 4, hypotype U.S.N.M. 110826c. 109, Interior of the brachial valve, & 2, hypotype U.S.N.M. 110826b. 20, Posterior part of the same specimen showing the cardinalia with the long falci- fer crura, X 4. 21-23, Respectively side, tilted anterior, and full views of another brachial valve, & 4, showing the long falcifer crura, the small elevated inner hinge plates, corrugated sockets, small genital areas, and pallial marks, hypotype U.S.N.M. 110826c. Recent, U. S. Fish Commission Station 4936, rocky bottom at 103 fathoms, Kagashima Gulf, Kyushu, Japan. PLATE 14 Basiliola and Frieleia A. Bastliola beechers (Rial) eee faite etletonte eisseiarees to tatetertns te ete oiesetterene 6 25 Fig. 1. Interior of the pedicle valve of an obese specimen, X 2, showing thickened marginal rim and pallial marks, hypotype U.S.N.M. 334667. Recent, U. S. Fish Commission Station 3864, 163 to 198 fathoms, 55.9° F., Pailolo Channel, Hawaiian Islands. Fig. 2. Dorsal view of the apex of a pedicle valve showing the deltidial plates with their reflected rim and the anterior smooth area of the NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 83 Page pedicle collar which slides over the umbo of the dorsal valve, X 4, hypotype U.S.N.M. 274136. Recent, U. S. Fish Commission Station 3811, 238 to 252 fathoms, 70.5° F.?, south coast of Oahu, Hawaiian Islands. B. Frieleia? nitens (Conrad) = F.? astoriana (Dall)..........0.0cc0ceee 55 Figs. 3-6. 3-5, Respectively side, dorsal, and ventral views of the holo- type, U.S.N.M. 3487. 6, Ventral view of another specimen showing broad sulcus and radial lines on exfoliated shell, 1, paratype U.S.N.M. 3487a. Miocene, Astoria, Clatsop County, northwest Oregon. C. Basiliola elongata Cooper, new species... 16:2... s.cscccccoccecuceccuce, 20 Figs. 7-21. 7-11, Respectively dorsal, anterior, ventral, posterior, and side views of the holotype, 1, U.S.N.M. 235844a. 12-14, Respec- tively dorsal, side, and anterior views of the holotype, < 2, showing smooth surface, elongate form, and growth lines. 15, Interior of the pedicle valve of the paratype U.S.N.M. 235844b, X 2. 16, Beak region of the same pedicle valve, X 4, showing the fused deltidial plates and the reflected rim around the foramen. 17, Same pedicle valve, x 3, tilted to show the pedicle collar, dental plates, and small genital region. 18, Interior of the brachial valve of the same paratype, X 2, show- ing elongated falcifer crura. 19, 20, Side and anterior views of the preceding showing the broad falcifer crura, concave inward, and with serrate distal extremity, the small reflected inner hinge plates, and the broad outer hinge plates, & 4. 21, Interior of the apex of the same brachial valve, 6, showing the falcifer crura, broad outer hinge plates, small inner plates, and corrugated sockets. Recent, U. S. Bureau of Fisheries Station 5146, 24 fathoms on coral sand, Sulade Island, Tapul Group, Philippines. PLATE I5 Frieleia and Eohemithyris a rrelera mole MANN ate steals Sead ste et heck, Sane Mee la al le ee 53 Figs. 1-5, 12-14. 1-3, Respectively anterior, brachial, and side views, X 1, of a complete specimen showing the narrow sulcus in each valve and the rectimarginate anterior commissure, hypotype U.S.N.M. 110830a. 4, Interior of the pedicle valve tilted to show the strong dental plates and small teeth, < 2, hypotype U.S.N.M. I10830b. 5, Apical region of the preceding, X 4, showing the disjunct deltidial plates and incomplete foramen. 12, Brachial valve tilted to show apical chamber of the cardinalia, X 2, hypotype U.S.N.M. 110830c. 13, Another brachial valve tilted to show the apical chamber, X 2, hypotype U.S.N.M. 110830b. 14, The same, x 4, showing the apical chamber and cardinal process. Recent, U. S. Fish Commission Station 4797, 682 fathoms, off Avacha Bay, Kamchatka. Figs. 6-11. 6-8, Respectively full, partial side, and tilted views of a brachial valve showing cardinalia and median septum, X 2, hypotype U.S.N.M. 540348a. 9, 11, Apical part of another brachial valve in full 84 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 and anterior views showing the cardinalia, x2, hypotype U.S.N.M. 540348b. 10, The same, X 4, showing the large inner hinge plates covering the apical chamber and the small transverse cardinal process. Recent, U. S. Fish Commission Station 2923, 522 fathoms, off San Diego, California. B. Bohemithyris:colurnus | (tiledley )) io steer Gases shen cas Sere lees teases Figs. 15-26. 15-17, Respectively brachial, anterior, and side views of a complete specimen showing the anterior costation and uniplicate ante- rior commissure, I, hypotype U.S.N.M. 333012b. 18, Enlargement, < 2, of the beak of the preceding specimen showing the small sub- mesothyrid foramen and conjunct deltidial plates. 19, 20, Two views of the interior of a pedicle valve, X 2, one in full view, the other anteriorly tilted to show the minute dental plates, small genital areas, and pallial sinuses, hypotype U.S.N.M. 333012a. 21, Apical part of the preceding specimen showing beak, foramen, and conjunct deltidial plates, X 4. 22, 23, Full view and slightly tilted view of the brachial valve of the preceding specimen showing the cardinalia, pallial sinuses, and genital areas, X 2. 24, 25, Partial side and full views of the cardi- nalia showing the distally serrate, falcifer crus, corrugated sockets, and thickening over the crural bases, X 4. 26, Posterior part of the preceding tilted to show the concave ends of the crura, the small genital areas, and the interior thickening, x 4. Recent, 115-135 fathoms, off Gabo Island, Victoria, Australia. PLATE 16 Compsothyris A: Combsothurts racovttage ( Joubina) vcs): citer jas ox sis0.¢ bisa «eile Ore nies Figs. 1-17. 1-4, Respectively anterior, dorsal, ventral, and side views of a complete individual, showing faintly uniplicate commissure, < 1, hypo- type U.S.N.M. 549343. 5, Dorsal view of the preceding specimen showing fine closely crowded costellae, X 2. 6, 7, Interior of the pedicle valve of the same specimen, X 2, showing small foramen and small dental plates. 8, Beak of the preceding valve, X 4, showing small corrugated teeth and small disjunct deltidial plates. 9-11, Respectively full, slightly tilted, and strongly tilted views of the brachial interior of the same specimen showing cardinalia, median septum, and muscle scars, X 2. 12, Same brachial interior tilted to show the socket ridges and distally serrate spinulifer crura, x 4. 13-15, Three views of the cardinalia variously tilted to show socket ridges, narrow outer hinge plates, and crura, X 4. 16, Same brachial valve strongly tilted to show junction of crural supporting plates with median septum, X 4. 17, Exterior of the pedicle valve, X 6, showing the very fine radial costellae. Recent, British Antarctic Expedition 1910, Station 316 of Terra Nova, 190 to 250 fathoms, 30.5° F., off Glacier Tongue, 8 miles north of Hut Point, McMurdo Sound, Antarctic. Page 32 56 NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER PLATE 17 Probolarina Aa Prapowurina saris (COadll pvcn/ctie sess cece pele cmela canis a csoas s.asiias.as Figs. 1-19. 1-5, Respectively, posterior, ventral, side, dorsal, and anterior views of a small but complete individual, <2, hypotype U.S.N.M. 5409357a. 6, 7, Dorsal and ventral views of another hypotype showing variation of costation, X 1, U.S.N.M. 549355. 8-10, Respectively dorsal, ventral, and anterior views of a specimen larger and more strongly costate than the two preceding ones, X 1, hypotype U.S.N.M. 549357b. 11-14, Respectively side, anterior, ventral, and dorsal views of a small specimen with few costae, X 3, holotype U.S.N.M. 109293a. 15, Apical portion of a large specimen showing the small submesothyrid fora- men, conjunct and auriculate deltidial plates, X 4, hypotype U.S.N.M. 549354a. 16, Interior of a brachial valve, X 3, showing cardinalia, hypotype U.S.N.M. 549356d. 17, 18, Partial side and full views of the apical part of the same specimen, X 6, showing the concave falcifer crura and large outer hinge plates. 19, The same tilted anteriorly to show the concave crura and lack of median ridge, X 6. Eocene (Castle Hayne formation), at the city quarry near the cemetery, Wilmington, North Carolina. Ba Progolarina: Doles Call) 55 aesaissnicierainlavess sssto «0.556; o0e: wide Win, 6) Saad gets Os Figs. 20-36. 20-24, Respectively posterior, anterior, dorsal, ventral, and side views of a complete specimen, 2, hypotype U.S.N.M. 549350a. 25, The same, X 3, showing the ornamentation and long beak. 26, 27, Beak of the same specimen, X 5, showing conjunct and strongly auric- ulate deltidial plates. 28, Small specimen showing foramen and con- junct deltidial plates, & 4, hypotype U.S.N.M. 549359b. 20, 30, Apical part of another specimen showing conjunct and auriculate deltidial plates, X 6, and the same tilted to show the dental plates and pedicle collar, X 6, hypotype U.S.N.M. 549350e. 31, The same tilted to the side to show the pedicle collar, X 4. 32, Apex of another pedicle valve showing strongly auriculate deltidial plates, 6, hypotype U.S.N.M. 549359f. 33, Interior of the brachial valve, X 3, showing cardinalia and absence of median ridge, hypotype U.S.N.M. 549350g. 34, 35, Apical part of the preceding tilted to show concave falcifer crura, X 6. 36, Same in full view to show the outer hinge plates, x 6. Horizon and locality same as above. PLATE 18 Erymnaria A. Erymnaria. bolcensis, (Massalongo) qs 422.00) «:cisjstersieidlawie’se «iblasis'qeu e 6 Figs. 1-17. 1-5, Respectively dorsal, side, posterior, anterior, and ventral views of a decorticated specimen showing symmetrical form, X 1, U.S.N.M. 549383b. 6, Posterior view of preceding, X 3, to show dorsal umbo, trace of the crural supporting plates, and the shorter socket ridges outside of them (photographed under water). 7, Anterior of same specimen, X 2, showing symmetrical fold on brachial valve. 86 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 Page 8-12, Respectively posterior, anterior, side, dorsal, and ventral views of another symmetrical specimen, X 1, hypotype U.S.N.M. 549383a. 13-15, Respectively dorsal, anterior, and side views of the preceding, x 2. 16, 17, Two views of cross sections of the beak of the brachial valve U.S.N.M. 549383c, ca. X 8, respectively 2.6 mm. and ca. 6.0 mm. anterior to the beak, showing crura, crural supports, and socket ridges (see pl. 22, fig. 9, and explanation). Lower Eocene (Spilecciano), Spilecco, Verona, Italy. B. Erymnaria polymorpha (Massalongo) ..........0...eeceecenscceecenes 64 Figs. 18-25, 31-34. 18-22, Respectively ventral, dorsal, side, anterior, and posterior views of a large but imperfect specimen showing twisted anterior commissure and traces of pallial marks, 1, hypotype U.S.N.M. 75888a. 23, 24, Dorsal and anterior views of the same speci- men, X 2, showing trace of vascula media and twisted anterior com- missure. 25, Dorsal view of the same specimen taken under water and tilted away from the observer to show the long crural supporting plates as black lines and the shorter socket ridges, & 3. 31-33, Respec- tively side, anterior, and dorsal views of the preceding specimen, X 2, showing costae along the anterior margin. 34, Another specimen tilted away from the observer and taken under water to show the trace of the long crural supporting plates and the shorter socket ridges diverging widely from the beak, x 4, hypotype U.S.N.M. 75888b. Lower Eocene, Spilecco, Verona, Italy. Figs. 26-30, 35, 36. 26-30, Respectively anterior, ventral, side, posterior, and dorsal views of another specimen, not decorticated like the pre- ceding, and showing, in addition to the twisted commissure, short radial costae, X 1, hypotype U.S.N.M. 549384a. 35, Posterior of a brachial valve excavated to show cardinalia and crural supporting plates, X 6, hypotype U.S.N.M. 540384b. 36, Another brachial valve interior showing cardinalia with their fairly broad outer hinge plates, and crural supporting plates, < 6, hypotype U.S.N.M. 5409384c. Lower Eocene (Spilecciano), Spilecco, 400 meters southwest of Purga di Bolca, Monti Lessini, Verona, Italy. PLATE 19 Erymnaria and Streptaria A Eramnaria cubensis Coopenan. Js. cka2taee eins soe cele ales tates ac cee 65 Figs. 1-10. 1-5, Respectively dorsal, anterior, ventral, posterior, and side views of the holotype, X 1, U.S.N.M. 540385. 6-8, Respectively side, dorsal, and anterior views of the same specimen, X 2, showing folding. 9, Posterior of holotype, X 3, showing conjunct deltidial plates and foramen. 10, Holotype tilted and photographed under water to show crural supporting plates and the shorter, more widely divergent socket ridges, X 3. Eocene, 80 meters northeast of school, Chucho Machin, Matanzas Province, Cuba. B. Streptaria streptimorpha Cooper, new species...........-...eeeeeeeeee 40 Figs. 11-21. 11-15, Respectively anterior, ventral, side, dorsal, and pos- NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER 87 Page terior views of the holotype, X 1, showing twisted anterior commis- sure, U.S.N.M. 549386a. 16, Dorsal view of the holotype showing smooth exterior, X 13. 17, 18, Side and anterior views of the holo- type, X 2, showing twisted commissure. 19, Posterior of the holotype, 3, showing deltidial plates and foramen. 20, 21, Posterior of another specimen, < 4, showing short dental plates and cardinalia with falcifer crura (see discussion), paratype U.S.N.M. 549386b. Eocene, deep cut north of Grua 9, Ramal Juan Criollo, Camaguey Province, Cuba. C. Streptaria buchi (Michelotti) ........ cece ee cece eee e cence eer ceeeeees 38 Figs. 22-31. 22-26, Respectively ventral, dorsal, anterior, side, and posterior views of a complete specimen showing twisted anterior com- missure, X 1, hypotype U.S.N.M. 549352a. 27-20, Respectively ante- rior, dorsal, and side views of the preceding specimen, X 2, showing twisted commissure. 30, Posterior of the same specimen, X 3, show- ing large foramen with elevated rim. 31, Posterior of the brachial valve of another specimen prepared to show the strong socket ridges, wide outer hinge plates, and falcifer crura with thickened ventral edge, X 6, hypotype U.S.N.M. 549352b. Middle Miocene, Messina, Sicily. PLATE 20 Eohemithyris A. Eohemithyris alexi Hertlein and Grant..........ccceesecceceseccceees 30 Figs. 1-16. 1-5, Respectively dorsal, posterior, ventral, side, and ante- rior views of a complete and undistorted specimen, showing form and true profile of valves, X 1, hypotype U.C.M.P. 15524. 6-9, Respec- tively, anterior, ventral, dorsal, and side views of the same specimen, <2. 10, Dorsal view of another nearly perfect specimen showing rounded outlines and low fold, 1, hypotype U.C.M.P. 15526. 11, Posterior of the same specimen, X 3, showing small foramen. 12, Posterior of another specimen, X 3, showing small foramen and con- junct deltidial plates, hypotype U.C.M.P. 15541. 13, 14, Dorsal and ventral views of a specimen from which the shell has been scraped to show the muscle fields, & 2, hypotype U.C.M.P. 15542. 15, 16, Full view and partial side view of the cardinalia, x 6, showing narrow outer hinge plates and broad, long falcifer crura, hypotype U.C.M.P. 15545. Eocene (Domengine formation), from just below the 1/4 section marker toward the top of the 25-foot last sandstone “reef” on the ridge on the east side of the North Fork of Media Agua Creek, grid. coord. 142001-139004, McKittrick (15’) Quadrangle, Kern County, California (see text for further information). Bo Baheathres, Gave CNV GOGWALG) i, «ai. 5:5:»/orctors fargrs! oyapeieve a olel atten ©\e abno aislels 33 Figs. 17-23. 17, Dorsal view of the exterior, ca. X 2, showing anterior costae and minute foramen, holotype British Museum (Nat. Hist.) ZB280. Specimen coated by ammonium chloride. 18-20, Respectively 88 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. side, dorsal, and anterior views of the same specimen, ca. X 2. 21, Posterior of the same specimen (coated) showing small foramen, ca. X 24. 22, Interior of the pedicle valve of the holotype, ca. X 23, showing small foramen, muscle field, and pallial marks. 23, Interior of the brachial valve of the holotype showing cardinalia with falcifer crura, thickened inner edges of crural bases, muscle field, and pallial marks, ca. X 4. Recent, Fiji Islands. Photographs by permission of the Trustees of the British Museum (Natural History) through Dr. H. M. Muir-Wood Deputy Keeper, Department of Palaeontology. PLATE 21 Frieleia, Patagorhynchia, Septaliphoria, Cryptopora, Tegulorhynchia, Mannia, and Hispanirhynchia? A. Hemithyris astoriana Dall= Frieleia ? mitens Conrad............2000% Fig. 1. Section of the apex of the brachial valve showing the small chamber and median septum, X 8, paratype U.S.N.M. 3487a. Miocene, Astoria, Clatsop County, Oregon. B. Patagorhynchta patagomeca (Thering) 2. v2.30 sciedea ee selene o ejeeiartee see Figs. 2-5. Respectively anterior, dorsal, side, and ventral views of a fairly well-preserved specimen showing costellate ornamentation, x1, figured specimen U.S.N.M. 549387. Eocene, 1 to 2 miles southwest of Ancha Terriza, Rio de los Cier- vos, Provincia de Magallones, Chile. C, Sepralt phoma species: sec cle aa ae tee ain ea oe oc e ee ack eee ae vtelore eatersie Fig. 6. Interior of the dorsal valve tilted to show septalium and median septum, X 2, figured specimen U.S.N.M. 129896a. Jurassic (Lower Callovian), in the railroad cut 300 meters east of the station at Chatillon-sur-Seine, Department of Cote d’Or, France. D. \Cryptopora gnome C(ICHLEYS )le/.(. x o's< o's creyse esis ais alecals,ss sacnislecce eels syelute Figs. 7-14. 7-9, Respectively anterior, side, and ventral views of a com- plete specimen, X 6, hypotype U.S.N.M. 44911a. 10, Anterior of an- other specimen, <6, showing the sulcate anterior margin, hypotype U.S.N.M. 44911c. 11, Dorsal view of the preceding, < 8, showing the nonauriculate deltidial plates. 12, Posterior of the pedicle valve, <8, showing deltidial plates, apical plate, and teeth, hypotype U.S.N.M. 44911d. 13, 14, Full and partial side views of the brachial valve of the preceding interior showing median septum and maniculifer crura. Recent, U. S. Fish Commission Station 2221, 1,525 fathoms, 36.9° F., south of Marthas Vineyard, Massachusetts, in gray ooze. EB. Tegulorhynchia doderlews ‘((Davyidson)). ws sans. 2 2s ce ccvancsete« Fig. 15. Interior of the pedicle valve, X 1, showing pallial sinuses, after Leidhold, Neues Jahrb. Min., Geol., Palaont., BB. 45, pl. 11, fig. 1b [here reduced 1/2], 1922. Original in Institute of Zoology, Strasburg University. Recent, Sagami Bay, Honshu, Japan. 139 Page 56 44 10 22 51 NO. 5 RHYNCHONELLOID BRACHIOPODS—COOPER PERV C1190) AY SEs sD A VAG SOME oycyercicuekteveleversisjevaictokece le PAV alsietate Wreianal siataretaeks reeset s Figs. 16-20. 16, Drawing of the dorsal side, ca. X I. 17, Exterior of the dorsal side, ca. X 7, showing large foramen and long, elevated delti- dial plates. 18, 19, Interior of two brachial valves showing septum with spoonlike plate and long maniculifer crura. 20, Cross section through a complete individual showing relationship of valves, median septum, spoonlike plate, and crura. All from Davidson, Geol. Mag., dec. 2, vol. 1, No. 4, pl. 7, 10-13, 1874. Miocene (Diestien), 3 miles east of Antwerp, Belgium. G.. Elaspanwhynchia °° craneana: (Dall)... Sear Fe Eee cues Hoe iia we Figs. 21-26. 21-23, Respectively dorsal, anterior, and side views of the holotype showing gently sulcate anterior commissure, X 1, U.S.N.M. 122861. 24, The holotype, 2, showing the beak area. 25, The beak region of the holotype, X 4, showing teeth (the deltidial plates prob- ably have been broken away). 26, Posterior of the brachial valve showing spinulifer crura and small outer hinge plate (these struc- tures have been damaged). Recent, U. S. Fish Commission Station 3362, mud at 117 fathoms and 36.8° F., off Cocos Island, Pacific Ocean off Panama. PLATE 22 Eohemithyris, Erymnaria, Notosaria, Aphelesia A. Bohemihyris: alex: Hertlein' and: Grant... 56 cic ew cln ele cee cceccseens Figs. 1-3. 1, 2, Full and partial side views of the cardinalia to show broad, falcifer crura, ca. X 5, hypotype U.C.M.P. 15545. 3, Drawing of the posterior of a pedicle valve, ca. & 4, showing the conjunct deltidial plates and small, round foramen, hypotype U.C.M.P. 15524. Horizon and locality as in plate 20, figures 1-16. B. Erymnaria polymorpha (Massalongo) ..........cccccscccnseccescccvcs Figs. 4-9. Sections through a slightly distorted individual, ca. X 4, hypotype U.S.N.M. 549384e. Sections measured from pedicle beak respectively: (4) ca. I mm., (5) 1.7 mm., (6) I.95 mm., (7) 2.25 mm., (8) 2.6 mm., and (9) ca. 4.0 mm. Lower Eocene (Spilecciano), 400 meters southwest of Purga di Bolca, Monti Lessini, Verona, Italy. Figs. 10-15. Sections through another individual showing long crural supporting plates, ca. X 4, British Museum (Natural History) B 8088. Sections respectively 0.3 mm. apart except figure 13 which is 0.4 mm. from figure 12. Eocene, Castelvecchio, Vicentin, Italy. C. Tegulorhynchia (= Notosaria) nigricans (Sowerby).............20+: Figs. 16, 17. Diagram of the interior of the pedicle and brachial valves of the adult showing pallial sinuses, ca. & 1.5, after Williams (1956, fig. 7, No. (4) on p. 276). Recent, New Zealand. 61 30 64 48 go SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 DA phelésia biparttia, .GBcoccht ) icine 2% kx teiccmie wiciel Mele eee tatale ele aiie 41 Figs. 18-25. Sections through the valves of a full-grown adult, Xx 2, British Museum (Natural History) not numbered, showing sockets and teeth. Sections respectively from the beak of the pedicle valve: (18) 1.0mm., (19) 1.6mm., (20) 2.0mm., (21) 26mm., (22) 2.8mm., (23) 3.2mm., (24) 3.6mm., and (25) 4.0mm. Miocene, St. Lorenzo, Tuscany, Italy. Photographs of figures 10-15 and 18-25 by permission of the Trustees of the British Museum (Natural History) through Dr. H. M. Muir-Wood, Deputy Keeper, Department of Palaeontology. cH a iyi Dau Pita cri Mine a a, ach ay ie a) hy rh i j iu f ea ' vi ra tS is i ua 7 i ry i ; iy fe int! hate a a ( ia Mab ve ; i vat a Wat i Ca i h ah ae Ne Onn iu ' } i Poe . ; gt Gea Ron at Ht i aa i ny AN Lt ae ne ji ‘ r i At ‘ HLH ‘ | " WV ae it tl) \ : ras i | i } Ne i Ms eh a Hea td . An acae ee a ) ina 7 i / | j j ‘* ’ ‘ i \ s : Pies 7 x t (put : i Fae { a i +H ¥ iit mt oa Ay ret y hal i if { ; Biss Neg 4i'3{) H f i F ’ iv Wie 4 ! ; Ri ‘al i eel | 7 i | Hie x : 1 , i i My } , y Mi a Ae . mT i ‘ { ay i / a ah a i PAWEL LST ea , MT 7 j ; | " A ie , , yt f Pi } f anh f ‘ i My you . r B ; Ma } ur ny, ' Al; \ i. Von i y = 3 veal nt \tn Ny f 4 i i my YA \ Te Se My 1 4 My ii t q Ap, Wn J ‘ N ‘ ; . ‘ l is | 1 t y ‘ : J Sa Pe Pn wa sally i i rl ih | iy ‘ i | \ Me ' i hat 7 . j i} yi | Crd pet 1h 4 ’ 7 ue ‘ i j \ 4 cm. - t ¥ ‘ \ 1 iP 1 ys f i 7 y J ce , 1 fi 7) i ‘ . oy \ Vi leat \ VOL. 139, NO. 5, PL. 1 THSONIAN MISCELLANEOUS COLLECTIONS CRYPTOPORA (SEE EXPLANATION OF PLATES AT END OF TEXT.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOES {39 NO 5. PEs CRYPTOPORA AND NEORHYNCHIA (SEE EXPLANATION OF PLATES AT END OF TEXT.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL. 3 A HEMITHYRIS (SEE EXPLANATION OF PLATES AT END OF TEXT.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL. 4 AETHEIA, FRIELEIA, GRAMMETARIA, CAMAROTOECHIA, AND HEMITHYRIS (SEE EXPLANATION OF PLATES AT END OF TEXT.) THSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL. 5 i Ho x cas Ay re EOHEMITHYRIS, SPHENARINA, CRYPTOPORA, AND TEGULORHYNCHIA (SEE EXPLANATION OF PLATES AT END OF TEXT.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, ‘NO: 5, PIS PATAGORHYNCHIA AND NOTOSARIA (SEE EXPLANATION OF PLATES AT END OF TEXT.) VOL2 139) |NO2 5; Pia THSONIAN MISCELLANEOUS COLLECTIONS PLICIRHYNCHIA AND APHELESIA (SEE EXPLANATION OF PLATES AT END OF TEXT.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL 17 SPHENARINA, APHELESIA, AND EOHEMITHYRIS? (SEE EXPLANATION OF PLATES AT END OF TEXT.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL. 9 GRAMMETARIA AND AETHEIA (SEE EXPLANATION OF PLATES AT END OF TEXT.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL. 10} HISPANIRHYNCHIA (SEE EXPLANATION OF PLATES AT END OF TEXT.) MITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL. 11 RHYTIRHYNCHIA AND BASILIOLA (SEE EXPLANATION OF PLATES AT END OF TEXT.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139; NO: 5, (PES if2 BASILIOLA (SEE EXPLANATION OF PLATES AT END OF TEXT.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL. 13 HISPANIRHYNCHIA? AND NEOHEMITHYRIS = BASILIOLA (SEE EXPLANATION OF PLATES AT END OF TEXT.) VOL. 139, NO. 5, PL. 14 SMITHSONIAN MISCELLANEOUS COLLECTIONS BASILIOLA AND FRIELEIA (SEE EXPLANATION OF PLATES AT END OF TEXT.) MITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL. 15 A FRIELEIA AND EOHEMITHIRIS (SEE EXPLANATION OF PLATES AT END OF TEXT.) VOL. 139, NO. 5, PL. 1 SMITHSONIAN MISCELLANEOUS COLLECTIONS COMPSOTHYRIS (SEE EXPLANATION OF PLATES AT END OF TEXT.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL. 17 PROBOLARINA (SEE EXPLANATION OF PLATES AT END OF TEXT.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL. 18 ERYMNARIA (SEE EXPLANATION OF PLATES AT END OF TEXT.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL. 19 ERYMNARIA AND STREPTARIA (SEE EXPLANATION OF PLATES AT END OF TEXT.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL. 20} EOHEMITHYRIS (SEE EXPLANATION OF PLATES AT END OF TEXT.) MITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL. 21 A FRIELEIA, PATAGORHYNCHIA, SEPTALIPHORIA, CRYPTOPORA, TEGULORHYNCHIA, MANNIA, AND HISPANIRHYNCHIA? (SEE EXPLANATION OF PLATES AT END OF TEXT.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 5, PL. 22 EOHEMITHYRIS, ERYMNARIA, NOTOSARIA, AND APHELESIA (SEE EXPLANATION OF PLATES AT END OF TEXT.) RS Oe Ee pe eee ey SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 139, NUMBER 6 Charles D. and Mary Waux THalcott Research Fund A REVISION OF THE SILURIAN BRYOZOAN GENUS TREMATOPORA (WitTH 2 PLATEs) By RICHARD S. BOARDMAN Associate Curator of Geology United States National Museum Smithsonian Institution (PuBLICATION 4383) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION OCTOBER 29, 1959 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 139, NUMBER 6 Charles D. and Mary Waux Talcott Research Fund A REVISION OF THE SILURIAN BRYOZOAN GENUS TREMATOPORA (WiTH 2 PrateEs) By RICHARD S. BOARDMAN Associate Curator of Geology United States National Museum Smithsonian Institution HSOW SS I ATV Wee tI NGTON 32? ee0ee8® (PuBLIcaTION 4383) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION OCTOBER 29, 1959 THE LORD BALTIMORE PRESS, INC. BALTIMORE, MD., U. S. A. Charles D. and Marp Vaux Walcott Research Fund A REVISION OF THE SILURIAN BRYOZOAN GENUS TREMATOPORA By RICHARD S. BOARDMAN Associate Curator of Geology United States National Museum Smithsonian Institution (WitH Two P ates) INTRODUCTION The genus Trematopora Hall, 1851, is placed in the order Tre- postomata of the Bryozoa and is the type genus of the family Trematoporidae Ulrich in Miller, 1889. The type species of Tre- matopora is T. tuberculosa Hall, 1852, from the Rochester shale in New York (type by subsequent designation, Ulrich, 1882, p. 241). The name Trematopora was established in an article by the editors of the American Journal of Science and Arts (Hall, 1851, p. 400) in which parts of Hall’s manuscript for volume 2 of the Paleontology of New York (1852) were quoted. The species of Trematopora listed ~ following the diagnosis of the genus were nomina nuda and were not published by Hall until the next year in volume 2. The development of the generic concept of Trematopora has been controlled partly by the study and preparation techniques employed by the various authors, each advance in technique adding refinements to the original very generalized description. All the work of Hall and Hall and Simpson (1851-1887) was done on external characters with- out the use of thin sections. In fact, Hall’s primary types of the type species were sectioned for the first time for the present paper. Owing in part to the external homeomorphy common in the Trepostomata, Hall included in the genus many forms now placed in other genera, families, and orders. At various times Hall considered such diverse genera as Tremaiella Hall, Orthopora Hall, Chaetetes Fischer (part), and Callopora Hall (part), as subgenera of Trematopora. Ulrich (1883, p. 257) was the first to section some “authentic speci- mens of Trematopora tuberculosa Hall.” These sections are in the SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 139, NO. 6 x 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 U. S. National Museum collections and are conspecific with Hall’s pri- mary types of the species. After seeing the sections, Ulrich greatly restricted the concept of the genus and indicated the great range of forms that Hall had included in the genus. The concept established by Ulrich in 1883 has remained essentially unchanged to the present time and was the type-genus concept for the family Trematoporidae in 1889. Under Ulrich’s definition of the genus, 12 species and sub- species have been assigned to Trematopora, ranging in age from Middle Ordovician through Middle Silurian. The primary type specimens were made available for sectioning and study by N. D. Newell of the American Museum of Natural History. Helpful suggestions were made by Helen Duncan and W. A. Oliver, Jr., of the U. S. Geological Survey, and N. Spjeldnaes, of the Uni- versity of Oslo. Thin sections were prepared by T. M. Robison of the U. S. Geological Survey. Photography was done by J. Scott, and the text figure was drawn by L. B. Isham, both of the Department of Geology of the U. S. National Museum. INTERPRETATION OF SKELETAL MICROSTRUCTURE The skeletal structures of most trepostomatous Bryozoa are com- posed of finely laminated calcite (fig. 1 and pl. 2). These laminae are assumed to have been deposited parallel to the surface of the secreting tissue (Cumings and Galloway, 1915, p. 361). Therefore, trends of the laminae within skeletal structures such as walls and diaphragms are considered to reveal something of the disposition of the original secreting tissue and the mode of growth of the skeletal structures. In longitudinal thin sections of T. tuberculosa, laminae are com- monly oriented parallel to the zooecial walls (fig. 1) in the endozone (immature or axial region of authors) and to the thinner walls and mesopore diaphragms in the inner region of the exozone (mature region of authors). This type of microstructure is here designated longitudinally laminated structure. Such an orientation of laminae is assumed to indicate that the depositing tissue was parallel to the walls and diaphragms, but it does not indicate whether the laminae were deposited on one or both sides of the structures. Another type of structure is characterized by laminae that are curved or angled transversely to the walls and diaphragms as seen in longitudinal sections. The transverse laminae form V- or U-shaped patterns with apices pointing distally and aligned along the median line of a wall or diaphragm. This type of microstructure is here designated transversely laminated structure. In T. tuberculosa, this structure is No. 6 BRYOZOAN GENUS TREMATOPORA—BOARDMAN 3 found in the walls of zooecia and mesopores in the outer region of the exozone, and in the inner region in some of the thicker mesopore walls and in the vicinity of central pores in the mesopore diaphragms (fig. I). Assuming that secreting tissue was oriented parallel to the laminae, transversely laminated structure indicates that the tissue must have been wrapped around the growing edges of walls and diaphragms Inner Region Outer Region <—- Endozone ——>\< of Exozone ——><— of Exozone ———> 1 ! i ! I I ! 1 y MESOPORE + Fic. 1.—Idealized diagram of T. tuberculosa in longitudinal view illustrating the variety of laminated structures commonly occurring in the species. Two mesopores and an intervening zooecium are shown in profile. Few central pores of the mesopore diaphragms are intersected in a longitudinal section. on both sides of the median lines. Thus, transversely laminated struc- ture presumably indicates deposition from both sides of a wall or diaphragm. Such interpretations applied to the skeletal laminae of T. tuberculosa and correlated with other morphologic characteristics of the species suggest that the exozone is divisible into two parts, an inner and an outer region (fig. 1) based on fundamentally different modes of growth of the mesopores. The physiologic significance of the two modes of growth is a matter for speculation. The taxonomic signifi- cance of these characters must await comparable studies in related genera. 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 In the inner region of the exozone, the mesopores are beaded (series of rounded chambers). This beading is produced by the mesopore walls curving transversely to the axis of the mesopore to form a diaphragm. The succession of longitudinally and transversely lami- nated structures along the walls and diaphragms is interpreted to indicate deposition from both sides of longitudinal as well as trans- verse structure throughout the inner region. Thus, at least a part of the depositing tissue of the mesopore remained behind the diaphragm and within the chamber being formed. The pore in the center of the mesopore diaphragm presumably would be necessary to allow the soft parts within the chamber to communicate with the outside environ- ment. There is no indication as to whether the soft parts in the mesopores during the formation of the inner region consisted of any- thing more than a depositing tissue or mantle, but the distal diaphragm would have acted, temporarily at least, as a covering for any soft parts within the last chamber. Continuity of laminae from the distal side of a diaphragm to the wall of the succeeding mesopore chamber implies that at least the proximal part of the chamber was formed as the dia- phragm developed (see middle diaphragm of the inner region in the upper mesopore of figure 1). For a more complete discussion of figure I see species description. The thick-walled outer region of the exozone contains transversely laminated structure in the walls of zooecia and mesopores and longi- tudinally laminated structure in the thick diaphragms. The mesopores in this region are not beaded. The pattern of continuity of the laminae of walls and adjoining diaphragms (fig. 1) indicates deposition was limited to the outer surface of the walls and diaphragms, con- temporaneous deposition of laminae taking place on the outermost surfaces of the zooecial wall, around the median line or boundary to the mesopore wall, and back to the distal side of the diaphragm. There is no evidence that deposition occurred on the proximal side of the diaphragm within the mesopore chamber. The formation of the diaphragms in the inner and outer regions of the exozone is quite different. Diaphragms in the inner region were formed by continued distal growth of mesopore walls that merely curved through an angle of 90 degrees to form transverse structures. Diaphragms in the outer region were formed by an additional, trans- versely oriented sheet of depositing tissue that was continuous with at least the depositing tissue of the mesopore side of the walls and ac- tively deposited calcite at the same time that the mesopore walls were being formed. This transverse sheet of tissue apparently had no counterpart in the inner region of the exozone. No. 6 BRYOZOAN GENUS TREMATOPORA—BOARDMAN 5 Evidence concerning the position of the soft parts in the outer region of the mesopore is inadequate. Configurations of the laminae give no indications of deposition behind the distalmost diaphragm. The common occurrence of single, centrally located pores that either partly or completely penetrate the thick outer diaphragms suggest some activity within outer chambers. The majority of these central pores appear to have been cut through the laminae of the diaphragms. Their termination within outermost diaphragms suggests that activity within the outermost chambers might have been choked off by the growth of the thickened diaphragms. SYSTEMATIC DESCRIPTION Genus TREMATOPORA Hall, 1851 1851. Trematopora Hall, Amer. Journ. Sci. and Arts, ser. 2, vol. 11, p. 400. 1852. Trematopora Hall, Paleontology of New York, vol. 2, p. 1490. 1881. Trematopora Hall, Nicholson, Genus Monticulipora, pp. 232-234. 1882. Trematopora Hall, Ulrich, Journ. Cincinnati Soc. Nat. Hist., vol. 5, p. 241. 1883. Trematopora Hall, Ulrich, Journ. Cincinnati Soc. Nat Hist., vol. 6, p. 257. 1887. Trematopora Hall, Hall and Simpson, Paleontology of New York, vol. 6, p. Xiv. 1893. Trematopora Hall, Ulrich, Geol. Minnesota, vol. 3, pt. 1, p. 308. 1911. Trematopora Hall, Bassler, U. S. Nat. Mus. Bull. 77, pp. 267, 268. 1882. [non] Trematopora Hall, Ulrich, Journ. Cincinnati Soc. Nat. Hist., vol. 5, DP. 153. Type species —Trematopora tuberculosa Hall, 1852. Emended definition—Zoaria are ramose, conspecific overgrowth is common, and monticules range from level to tuberculated. Externally, zooecia are elliptical to subcircular in cross section and walls are slightly elevated above intervening mesopores. Mesopores form shallow, subpolygonal depressions between zooecia. The exozone is divided into an inner thin-walled region containing the earliest chambers of the mesopores and an outer thick-walled region. In the inner region, both mesopores and zooecia are polygonal to subpolygonal in cross section and mesopores are beaded and contain diaphragms with single central pores. In the outer region of the exozone, zooecia become elliptical to subcircular in cross section and the mesopores contain thickened diaphragms and are not beaded. Diaphragms are thin and few in zooecia. In the outer region of the exozone, walls of adjacent zooecia are divided by sharply defined zooecial boundaries, as seen in longitudinal sections. Laminae on either side of a boundary converge to form a 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 V-shaped pattern that has extremely long limbs trending nearly parallel to the boundary and curving very slightly just before the boundary is intersected. Laminae of walls of zooecia and adjacent mesopores are more broadly curved approaching the median boundary and form a broad U-shaped pattern having limbs of varying lengths. Acanthopores are common in the zooecial walls. Discussion.—Based on an examination of thin sections of primary types of species previously assigned to Trematopora now in the U. S. National Museum collections, the following species are considered cor- rectly assigned to the genus: T. halli Ulrich 1883, Niagaran group, Waldron, Ind. T. whitfieldi Ulrich 1883, Niagaran group, Waldron, Ind. The holotype section of T. spiculata Miller 1877, Niagaran group, Waldron, Ind., is not adequate to determine generic affinities. This species is retained in the genus until a more detailed study of addi- tional material can be made. The following species originally placed in Trematopora do not satisfy the generic definition proposed here and are not considered to belong to the genus. Their proper generic assignments must await restudy of both the species themselves and the available genera. calloporoides Ulrich 1890, Cincinnati group, Alexander County, III. cystata Bassler 1911, Kuckers shale (C2), Reval, Esthonia. This species is the type of Aostipora Vinassa 1920). debilis Ulrich 1890, Girardeau limestone, Alexander County, III. kukersiana Bassler 1911, Kuckers shale (C2), Reval, Esthonia. primigenia Ulrich 1886, Decorah shale, Minneapolis, Minn. primigenia var. ornata Ulrich 1886, Decorah shale, Minneapolis, Minn. None of the Ordovician species investigated displayed the two regions of the exozone or pores in the mesopore diaphragms. Thus, the genus is limited presently to the Middle Silurian. A close taxonomic relationship seems to exist between Trematopora and some or all of the Silurian and Devonian species that have been placed in the genus Leioclema Ulrich. These species of Letoclema are largely incrusting and possess many of the morphologic characters now defining Trematopora. In general they have elliptical zooecia with few thin diaphragms, abundant mesopores with closely spaced thicker diaphragms and an irregularly discontinuous inner region of the exozone containing beaded mesopores. Pores in the diaphragms of the mesopores are rare but do definitely occur in the following species. No. 6 BRYOZOAN GENUS TREMATOPORA—BOARDMAN 7 Leioclema asperum (Hall) 1852, Rochester shale, Lockport, N. Y. (Only Bassler’s plesiotypes of 1906 available.) L. confertiporum (Hall) 1883, Hamilton group, New York. L. decipiens (Hall) 1883, Hamilton group, New York. L. passitabulatum Duncan 1939, Traverse group, Michigan. The region now considered to be the inner region of the exozone in species of Leioclema from the Hamilton group of New York was interpreted as the outer part of the endozone (Boardman, in press) and diaphragm pores were overlooked. TREMATOPORA TUBERCULOSA Hall Pl. 1, figs. 1-4; pl. 2, figs. 1-3 1852. Trematopora tuberculosa Hall, Paleontology of New York, vol. 2, p. 149, pl. 40A, figs. Ia-g. 1883. Trematopora tuberculosa Hall, Ulrich, Journ. Cincinnati Soc. Nat. Hist., vol. 6, p. 259, pl. 13, figs. 2, 2a, 2b. 1906. Trematopora tuberculosa Hall, Bassler, U. S. Geol. Surv. Bull. No. 292, p. 43, pl. 13, figs. 15, 16; pl. 17, figs. 1-3; pl. 25, fig. 8. TYPE DATA Lectotype (Hall, 1852, pl. 40A, fig. 1a) and the two paratype zoaria from syntype suite No. 1747, American Museum of Natural History. MATERIAL STUDIED In addition to the primary types, 55 fragmentary topotype zoaria were studied. The topotypes are from U. S. National Museum col- lection No. 2998 and cat. No. 43618 collected by E. O. Ulrich. U. S. National Museum catalogue numbers of illustrated topotypes are 137847 to 137850. OCCURRENCE Rochester shale member of the Clinton formation, Lockport, N. Y. DESCRIPTION Zoaria.—Zoaria are ramose and branches are circular to elliptical in cross section. Branch arrangement was affected by branches rising from conspecific secondary growth superimposed on the normal bi- furcating pattern. Branches of secondary growth produced irregu- larities in branch arrangement, commonly causing branches to anastomose and form erratic and confused zooecial growth at surfaces of contact. These irregularities in branch arrangement were formed 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 by random bends at ramose extensions of overgrowths beyond the distal tips of primary branches, and irregular branch angles in lateral secondary branches. The zoaria were further complicated by repeti- tions of thin- and thick-walled growth in the outer region of the exozone (mature region) without the formation of intervening basal laminae. This apparently rejuvenated growth formed localized swell- ings on the zoaria, and combined with adjacent patches of overgrowth to form some of the secondary branches. Monticules.—Monticules are prominent tubercles. The apertures of some monticular zooecia are restricted or closed by a distal thicken- ing of the walls, and the walls and outer diaphragms of monticular mesopores are somewhat thicker than those of surrounding mesopores. Monticular mesopores generally contain one to several more dia- phragms than intermonticular mesopores. Longitudinal View: Endozone.—In the endozone (immature or axial region), zooecial walls are longitudinally laminated and do not show the dark granularity that is common in the Trepostomata. The zooecial walls range from undeviating to irregularly undulating. Ina few specimens the endozone is interrupted by a zone arching distally across the branch that is marked by variable thickening of the zooecial walls. Normal thin-walled zooecial growth generally continues distally from the thickened walls of the arched zones with some bifurcating but without other break. At apparently random levels within a colony, some or all of the zooecia within the endozone have been eroded and the tubes filled with mud. Subsequent growth was initiated from adjacent zooecia and the eroded areas were covered by a basal lami- nation of the overgrowth that continued the colony distally. Exozone: Inner region of mesopores—The boundary between the endozone and exozone is defined by the points of origin of the mesopores. The inner region of the exozone extends distally for one to several mesopore diaphragms, but generally not more than four. The mesopores begin proximally with walls and diaphragms that are slightly thicker than the zooecial walls of the endozone. Mesopore walls curve broadly through 90 degrees into transverse positions relative to the length of the mesopores, thereby forming diaphragms. The broad curving results in constrictions of the mesopores at the positions of the diaphragms to form cystlike chambers. In this inner region of the exozone, mesopore walls commonly are longitudinally laminated, but many, especially the thicker ones, develop transversely laminated structure, either intermittently or throughout their length. In the inner region, mesopore diaphragms regularly display centrally No. 6 BRYOZOAN GENUS TREMATOPORA—BOARDMAN 9 located single pores that penetrate the diaphragms at right angles. In longitudinal thin sections that pass through these pores, diaphragms display transversely curved laminae that continue uninterrupted to the pores. The curved laminae immediately adjacent to the pores define the rounded boundaries of the pores. If walls of adjacent mesopore chambers are longitudinally lami- nated, generally the wall of the earlier chamber is connected directly with the curved laminae on the proximal side of the intervening dia- phragm and the wall of the later chamber is connected with the distal side of the diaphragm. If walls of adjacent mesopore chambers are formed by transversely curved laminae, the diaphragm and ad- jacent walls will appear to be a continuous unit, or the diaphragm is a direct continuation of the proximal wall and the wall of the distal chamber is discordantly joined to the distal side of the diaphragm. Rare, isolated mesopore diaphragms display complete continuity with the walls of distal chambers. In longitudinal thin sections, mesopore diaphragms in which the pores were not intersected appear longitudinally laminated. Com- monly the diaphragms are compound; the proximal half of a dia- phragm is continuous with the wall of the preceding mesopore chamber, the distal half is continuous with the wall of the succeeding chamber. Other variations in diaphragm-wall relationships are less common ; the two parts of the compound diaphragm can be unequal in thickness, or in extreme development a diaphragm loses its com- pound appearance and is wholly continuous with the preceding or very rarely the succeeding chamber walls throughout or at either end. Outer region of mesopores.—In the outer region of the exozone, mesopores are not beaded and the walls and diaphragms display ex- treme thickening. This greatly thickened skeletal growth can begin on the distal side of the last thin diaphragm, the laminae covering the central pore of the thinner diaphragm and curving distally into the mesopore walls, or it can begin by an abrupt thickening of the meso- pore walls. Diaphragms in this outer region are extremely variable in thickness and spacing. A single diaphragm, greater in thickness than the diameter of the enclosing mesopore, can correspond in thick- ness and position with a series of irregularly and closely spaced dia- phragms in adjacent mesopores. Most diaphragms are planar, but a few are strongly curved and join adjacent diaphragms before reaching the mesopore wall. The last diaphragms that were formed are in the distal ends of the mesopores so that in external view the walls and diaphragms of the mesopores combine to form the very shallow polyg- onal depressions between the zooecia. Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 Many of the thick diaphragms also display centrally located pores that do not penetrate through to the distal sides of most of the thickest diaphragms. Laminae of the diaphragms generally stop abruptly at the pores without changing direction or flexing, so that the pores have no lining or apparent influence on the structure of the diaphragms. In other thick diaphragms the laminae trend in a proximal direction in varying amounts and there is a noticeable decrease in diaphragm thick- ness approaching the pore. The pores in the outer region also differ from the central pores of the inner region of the exozone by being consistently smaller in diameter. In addition to the pores, mesopore diaphragms and walls in the outer region contain small, dark, sub- spherical to elongated cavities formed by the concentric arrangement of laminae about imaginary centers. These cavities seem to be ar- ranged at random in the walls and diaphragms. Zooecia.—In the outer region of the exozone, undistorted wall structure of adjacent zooecia is rarely seen because of intervening mesopores and acanthopores. Zooecial boundaries are well defined, dark, slightly serrated lines or zones in two dimensions, formed by the abutting ends of laminae from adjacent zooecia. In walls formed by a zooecium and adjacent mesopore, or by adjacent mesopores, boundaries are more coarsely serrated and are commonly discontinuous along their length. Diaphragms are not present in most zooecia and not more than two were seen in any one zooecium. If present, diaphragms are very thin, planar to slightly curved, and extend distally into the zooecial wall. Single, hollow, subspherical cystlike structures occur in the zooecial voids of a very few zooecia, more commonly in the monticules. The cyst walls are thick and are constructed of laminae that merge with the laminae of the zooecial walls. Irregular spinelike proc- esses are common in the zooecial walls in the thick-walled outer region. These mural spines have their origins at or very near the zooecial boundaries and trend in general toward the zooecial voids at a high angle to undisturbed laminae in the walls. Zooecial wall laminae surrounding the spines are flexed about the spines in a series of ir- regular superposed cones and some of the laminae are pierced. The spines extend far enough to cause inflection of the walls but none were observed to break through the wall laminae and stand in relief in the zooecial voids. The cores of the spines appear structureless or hollow. Tangential View.—In tangential sections passing through the outer region of the exozone, zooecia range from irregularly elliptical to subcircular to petaloid in cross section. Major axes of the ellipses are approximately parallel to branch length. The rare petaloid ap- No. 6 BRYOZOAN GENUS TREMATOPORA—BOARDMAN Il pearance is caused by extreme inflection of zooecial walls by adjacent acanthopores and mural spines. Acanthopores are large, laminated, possess well-defined central tubes, and are generally located at points of closest proximity of adjacent zooecia. Mural spines appear to begin outside the broad band of striated-appearing tissue lining each zooecium and project inwardly toward the zooecial void, strongly in- flecting the laminated tissue but not breaking through to the void. Mesopores are numerous, subpolygonal to subcircular. In very shallow sections that pass through the outermost and thickest diaphragms, mesopore boundaries are concealed and interspaces between zooecia appear solid. Many of these solid interspaces do not show the smaller central pores that are the rule in sections that pass through earlier parts of the outer region. In deeper tangential sections that pass through the inner region of the exozone, zooecia are polygonal to subpolygonal and approximately equidimensional. Mesopores are also polygonal to subpolygonal and have fewer sides than the zooecia, merely appearing to fill the spaces between zooecia. Pores in mesopore diaphragms here are several times larger in diameter than those in the outer region. Acanthopores are considerably smaller in diameter than they are in the outer region. QUANTITATIVE DATA The following tables are based on sections of two fragments from the lectotype, three fragments from the two paratype zoaria, and 49 fragments from 33 topotype zoaria. Sections from 55 zoaria of Trematopora tuberculosa were examined. All measurements are in millimeters. The axial ratio is the ratio of the diameter of the endo- zone to the corresponding branch diameter. TABLE 1.—General measurements Lectotype AMNH Paratypes and 1747 topotypes Frag. A. Frag. B. Minimum Maximum Diameter of zoaritim: food. 62 weeks 6.5 4.9 3.0 7.2 Width of endozone............00200. 5.3 3.3 28 5.6 No. zooecia in 2 mm. (longitudinal GIGECHOR)) hee sialon cue te amet a 53 63 6 8 Average major axis of zooecial void Per tragment, jcciwcsvoac soca casteleleas 0.14 0.15 0.14 0.22 Average minor axis of zooecial void Deri tragmenty oasis teihsbo steed eres 0.12 0.12 0.09 0.14 Acanthopores per zooecium .......... 0.73 0.63 0.50 0.5 Mesopores per zooecium............. 1.6 — El 2.0 I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 TABLE 2.—Ontogeny Average No. diaphragms Width of in mesopores exozone Axial ratio 2 0.3-0.6 0.87-0.92 3 0.5-1.0 0.74-0.90 Braswe Acs lectotyperdacsciaicieacice 4 1.2 0.82 4 0.9-1.4 0.71-0.86 5 1.0-1.4 0.66-0.82 6 1.4-1.8 0.75 Fira cmmbcmlectOtyperercieeiciciaele sete a 1.6 0.67 2 I.1-1.6 0.68-0.70 DISCUSSION The number of mesopore diaphragms and the width of the exozone are not particularly sensitive indicators for ontogenetic development of the mesopores and zooecia in T. tuberculosa. The variation in dia- phragm counts and in width of exozone within a longitudinal thin sec- tion is unusually large because of a marked variation in the number of chambers developed in adjacent mesopores in the inner region. Also, the unusual variation in thickness and spacing of mesopore dia- phragms in the outer region of the exozone makes diaphragm counts less reliable. T. tuberculosa differs from both T. halli and T. whitfieldi in having the larger branches, tuberculated mesopores, and a broader exozone in mature specimens. Both T. halli and T. whitfieldi are smooth, rhomboporoid-sized species. REFERENCES Basster, R. S. 1906. The bryozoan fauna of the Rochester shale. U. S. Geol. Surv. Bull. No. 292, 65 pp., 31 pls. 1911. The early Paleozoic Bryozoa of the Baltic Provinces. U. S. Nat. Mus. Bull. 77, 348 pp., 13 pls. BoarpMAN, R. S. Trepostomatous Bryozoa of the Hamilton group of New York State. U. S. Geol. Surv. Prof. Pap. 340, in press. Cumincs, E. R., and GAtLoway, J. J. 1915. Studies of the morphology and histology of the Trepostomata or monticuliporoids. Bull. Geol. Soc. Amer., vol. 26, pp. 349-374, pls. 10-15. HALL, JAMES. 1851. Jn Silliman, B., Silliman, B., Jr., and Dana, J. D., eds. New genera of fossil corals from the report by James Hall on the Paleontology of New York. Amer. Journ. Sci. and Arts, ser. 2, vol. II, pp. 398-401. No. 6 BRYOZOAN GENUS TREMATOPORA—BOARDMAN 13 1852. Paleontology of New York. Vol. 2, 362 pp., 101 pls. HA.L, JAMEs, and Simpson, G. B. 1887. Corals and Bryozoa. Paleontology of New York, vol. 6, 292 pp., 66 pls. Miter, S. A. 1889. North American geology and paleontology. 664 PP., 1,194 figs. in text. Cincinnati. NicuHotson, H. A. 1881. The genus Monticulipora and its sub-genera. 235 pp., 6 pls. UricH, E. O. 1882. American Paleozoic Bryozoa. Journ. Cincinnati Soc. Nat. Hist., vol. 5, pp. 121-175, 232-257. 1883. American Paleozoic Bryozoa. Journ. Cincinnati Soc. Nat. Hist., vol. 6, PP. 245-270. 1893. On Lower Silurian Bryozoa of Minnesota. Geol. Minnesota, vol. 3 pt. 1, chap. IV, pp. 96-332, pls. 1-28. I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 EXPLANATION OF PLATES IPPATE T Figs. 1-4. Trematopora tuberculosa Hall. 1. Longitudinal view of paratype, A.M.N.H. 1747, <5, showing primary branch with growth direction upward and branch from secondary overgrowth with growth direction to lower right. 2a. External view of lectotype zoarium, A.M.N.H. 1747, X 2, showing tuber- culated monticules. zb. Longitudinal view of lectotype, < 20, showing beaded mesopore cham- bers in inner region of exozone. 2c. Tangential view of lectotype, 20, showing aspect of both inner and outer regions of exozone. Note thin-walled polygonal tubes of inner region of monticule in upper left. 3. Tangential view of paratype, A.M.N.H. 1747, X 50, showing the smaller central pores in mesopore diaphragms of outer region of exozone. 4. Longitudinal view of topotype, U.S.N.M. 137847, X 5, showing zooecial growth at surface of contact of anastomosing branches. U.S.N.M. collection 2998. PLATE 2 Figs. 1-3. Trematopora tuberculosa Hall. 1. Longitudinal view of topotype, U.S.N.M. 137848, * 100, showing lam- inated structure of a beaded mesopore and two diaphragm pores in the inner region of the exozone. 2a. Longitudinal view of topotype, U.S.N.M. 137849, X 100, showing con- figuration of laminae of mesopores and small diaphragm pore in outer Tegion of exozone. 2b. Longitudinal view of same specimen, X 100, displaying a mesopore with diaphragm pore of inner region covered by first diaphragm laminae of outer region. Note discontinuous and ragged boundary between mesopore wall and zooecial wall above. 3a. Longitudinal view of topotype, U.S.N.M. 137850, X 100, showing first a diaphragm pore and then a compound diaphragm between beaded chambers in the inner region of the mesopore. 3b. Longitudinal view from same zoarium, X 100, illustrating the structure of the wall of adjacent zooecia. 3c. Tangential view from same zoarium, X 100, showing the general aspect of the outer region of the exozone, including acanthopores, mural spines, and a small pore in the center of a diaphragm of a mesopore. The dark intermediately sized spots are the randomly arranged cavi- ties noted in species description. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139, NO. 6, PL. TREMATOPORA TUBERCULOSA (See explanation of plates at end of text.) IITHSONIAN MISCELLANEOUS COLLECTIONS ‘ VOES 1397 NOn Gy REZ TREMATOPORA TUBERCULOSA (See explanation of plates at end of text.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 139, NO. 7 Charles DB. and fMlary Waux CHalcott Research Fund BARLEY (TERTIARY. APHELISCUS AND PHENACODAPTES' AS PANTOLESTID INSECTIVORES (WitH Two PLatTEs) By C. LEWIS GAZIN Curator, Division of Vertebrate Paleontology United States National Museum Smithsonian Institution eee Ov e ea yr ty \ 4 is { 1 ft L ad te! ¢ (Pustication 4385) \\ / \LBRAaBy eo CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION AUGUST 12, 1959 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 139, NO. 7 Charles D. and Mary Waux Talcott Research Fund EARLY TERTIARY APHELISCUS AND PHENACODAPTES AS PANTOLESTID INSECTIVORES (WitH Two PLatTEs) By C. LEWIS GAZIN Curator, Division of Vertebrate Paleontology United States National Museum Smithsonian Institution (PusiicaTion 4385) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION AUGUST 12, 1959 ITHSONIAN ana i '9 4 STITUTION Aug 1'2 1959 THE LORD BALTIMORE PRESS, INC. BALTIMORE, MD., U. S. A. Charles D. and Mary Waux Walcott Research Fund BARELY TAlERELARY, APE LISOCUS AND RHENACODAP LES cAS PANTOLESTEID INSECTIVORES By C. LEWIS GAZIN Curator, Division of Vertebrate Paleontology Umted States National Museum Smithsonian Institution (WitH Two PtatEs) INTRODUCTION Examination in 1954 of Phenacodaptes material in the Paleocene collections at Princeton University, believed pertinent to a review of Eocene artiodactyls then underway, led to rather inconclusive results. Dr. Jepsen’s tentative suggestion (1930, p. 519) of such a relationship may, nevertheless, have merit.1 More recent studies of the Knight faunas, however, involved certain pantolestids, and comparison of these among a wide range of both Eocene and Paleocene forms has convinced me that Cope’s Apheliscus and Jepsen’s Phenacodaptes are closely related and that both are pantolestids, although perhaps some- what less closely related to the Pantolestinae than to the Pentaco- dontinae. Their relationships would seem possibly best illustrated by placing them both in the Apheliscinae as a subfamily of the Panto- lestidae. I am indebted to Dr. Glenn L. Jepsen for permitting me to borrow and illustrate specimens of Phenacodaptes sabulosus in the Princeton collections, and to Dr. George G. Simpson and Mrs. Rachel H. Nichols for sending me materials of Apheliscus imsidiosus and Pentacodon inversus from the collections of the American Museum. The pencil drawings of specimens shown in the accompanying plates were made by Lawrence B. Isham, scientific illustrator for the Department of Geology in the U. S. National Museum. PREVIOUS INTERPRETATIONS OF RELATIONSHIP Apheliscus insidiosus was described by Cope (1874, p. 14) from the lower Eocene San Jose beds in New Mexico. He described it 1 After this manuscript was submitted for publication, Dr. Jepsen showed me a note that he had placed in the collection drawer some time ago suggesting that Phenacodaptes be compared more carefully with Apheliscus. SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 139, NO. 7 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 first as a species of Prototomus and included it together with “Pro- totomus” jarrovii (=Pelycodus jarrovii) in the carnivores with Prototomus (=Sinopa) viverrinus. In 1875 (p. 16), however, he proposed the name A pheliscus, regarding it as “nearly allied to Panto- lestes,’ although at the same time he thought that the molar teeth suggested a relationship to Anaptomorphus, noting, nevertheless, that the premolars were “totally different.’ Cope’s statement that the last lower molar lacked a heel would seem highly significant, but, if the meaning is here properly interpreted, it is surely an error, as may be seen from his illustration (1877, pl. 45, fig. 18). In addition to the described condition of the talonid of the third molar, Cope noted as distinctive in comparison with Pantolestes only the simplicity of the inner anterior tubercle of the lower molars. Matthew (1918), in naming the family Apheliscidae, was very dubi- ous as to its affinities, and while referring it to the Insectivora, con- sidered that it might well be condylarthran, primate, or creodont. It should be noted, however, that at the time of his writing, such genera as Aphronorus, Bessoecetor, and Phenacodaptes were not known. Only large and comparatively aberrant Pentacodon, which he had recognized as a pantolestid insectivore (1909), and the Eocene mem- bers of the Pantolestinae were available for comparison. Simpson (1937a) demonstrated the most logical arrangement for the pantolestids and pentacodonts, while adding the Paleocene genera Bessoecetor and Aphronorus to their respective subfamilies. He noted, moreover, the resemblance of Apheliscus to the Pentacodonti- nae in characters of the fourth premolars, but considered, however, that the molar structure was widely different. Nevertheless, his sug- gestion that Apheliscus might be an offshoot of the same stock seems particularly pertinent and certain of the lacking evidence for such an hypothesis may lie in Phenacodaptes. The family, however, was re- tained incertae sedis, questionably in the Insectivora in his 1945 classification. Saban (1954), evidently following Simpson’s suggestion, included the Apheliscidae with the Pantolestidae in the superfamily Pantoles- toidea. His including Shikama’s Endotheriidae, created for the Man- churian Jurassic Endotherium, as a subfamily of the Pantolestidae, however, seems surprising. McDowell (1958) rejected certain fea- tures of Saban’s classification and in discussing the family Aphelisci- dae regarded it as incertae sedis, but noted that the teeth are “recon- cilable with those of Mirodectes.’ McKenna, on the other hand, in a field conference guidebook (1955) has the Clark Fork species Aphelis- cus nitidus listed as a condylarth. Older but more recently described Phenacodaptes sabulosus is from NO. 7 TERTIARY APHELISCUS AND PHENACODAPTES—GAZIN 3 the Silver Coulee or Tiffanian horizon of the Polecat Bench forma- tion in the Big Horn Basin. The possibility of a relationship to artio- dactyls was tentatively suggested by Jepsen (1930) because of resem- blances noted to such genera as Diacodexis and Bunophorus, evident in certain features of the molars. Simpson, however, in his classifica- tion of the mammals (1945) cited Phenacodaptes as a condylarth under ?Mioclaeninae incertae sedis. COMPARISON OF APHELISCUS AND PHENACODAPTES A lower jaw of Apheliscus, referred to A. insidiosus, in the Na- tional Museum collection (No. 19162) from the Gray Bull beds in the Big Horn Basin, exhibiting P.-M, inclusive (see pl. 1, fig. 1), shows that the form and relative proportions of the lower premolars are strikingly like those in Phenacodaptes (see pl. 1, figs. 3, 4). The rela- tively small size of P, and particularly of P; in comparison with P, is quite alike in the two. P, is a little more slender in Apheliscus and the distinctive talonid seen in this tooth of Phenacodaptes is more sectorial and essentially better developed or exaggerated in Apheliscus. Both have a strongly developed primary cusp and only slight evidence of a paraconid. There is no metaconid on P, in the known material of Apheliscus. It is usually absent, but may be weakly developed in some specimens of Phenacodaptes. The lower molars differ noticeably in the anteroposteriorly shorter trigonid and more elongate talonid in Apheliscus (see pl. I, fig. 2) ; moreover, they are relatively more slender than in Phenacodaptes. There is, nevertheless, a rather marked similarity in many details, particularly in form of the cusps and crest surrounding the talonid basin, and in the shape of the basin. The compressed trigonid of Apheliscus is rather less like that in Phenacodaptes, although the paraconid is absent or very much re- duced on the posterior two molars of both forms. In M, of Phena- codaptes, however, this cusp is moderately well defined as an anterior crest from the protoconid, whereas in Apheliscus only a slight median cuspule remains. The upper cheek teeth of Apheliscus insidiosus (see pl. 2, fig. 1) may appear a little less like those of Phenacodaptes (see pl. 2, fig. 2) than perhaps do the lower teeth, although both exhibit the compara- tively small and subequal second and third upper premolars and en- larged fourth. The more noticeable differences between the two in upper teeth include less development of the cingulum, particularly on P*, and the transversely narrower molars of Apheliscus. More- over, the hypocone, though distinctive on M, and M, of Phenaco- daptes, is weak or absent in Gray Bull Apheliscus. It is important to 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 note, however, that the upper teeth seen in Clark Fork Apheliscus nitidus seem intermediate in most, if not all, of these respects. A com- parison of Matthew’s figure (1918, fig. 24) for the Clark Fork speci- men, which Simpson (1937b) made the type of A. nitidus, with P* and M?* in Phenacodaptes sabulosus, here shown in plate 2, figure 2, reveals little to distinguish them. The Sand Coulee lower teeth of Apheliscus, figured by Matthew (1918, fig. 24) also seem to show a little less compression of the trigonid than more typical Gray Bull specimens. The foregoing comparisons strongly suggest that Phenacodaptes, or at least a very closely related form, gave rise to Apheliscus. The succession may well have been Phenacodaptes sabulosus—A pheliscus nitidus—A pheliscus insidiosus. In the course of this postulated de- velopment it would seem that the principal tendency was toward the transverse narrowing of the teeth, both upper and lower series ; the loss or weakening of the cingulum in the upper series; the increas- ingly Pentacodon-like development of P*; the relative increase in length of talonid of the lower teeth, Ps as well as the molars ; together with the shortening of the lower molar trigonids. RELATIONSHIPS OF APHELISCUS AND PHENACODAPTES The most nearly comparable development to that illustrated in the Phenacodaptes—A pheliscus line would seem to be among the panto- lestids. The suggested comparison is perhaps not so close to the Bessoecetor—Propalaeosinopa—Palaeosinopa—Pantolestes succession as it is to the middle Paleocene Pentacodontinae. The premolar develop- ment would seem rather like that in both Aphronorus (see pl. 2, figs. 3 and 4) and Pentacodon (see pl. 2, figs. 5 and 6), except that there tends to be no metaconid on P, or tritocone (uncertain for Pentaco- don) on P* in the apheliscids. Aphronorus, moreover, differs in having somewhat higher crowned, more definitely insectivore teeth. The upper molars of Aphronorus show better developed and more laterally directed anteroexternal and posteroexternal styles and the lower molar trigonids are a little higher and show better development of the para- conid. Much larger Pentacodon has a more enlarged fourth premolar, but the upper molars (not previously illustrated) do not show the distinc- tive outer styles seen in Aphronorus. Also the trigonids of the lower molars do not appear to be so elevated, but, like Aphronorus and unlike the apheliscids, show a prominent and forward-placed paraconid. The talonid construction, nevertheless, is much alike in the two subfamilies, except for relative length. NO. 7 TERTIARY APHELISCUS AND PHENACODAPTES—GAZIN 5 The mental foramen, the position of which, as Simpson (1937a, p- 120) notes, has been unduly emphasized, may warrant comment. It exhibits a comparatively small posterior opening somewhat farther forward in the Apheliscinae than in Pentacodontinae or Pantolestinae. It is variable in Phenacodaptes and is observed in positions beneath the anterior part of P, to the posterior part of P;. A larger opening is noted beneath P, or Pz. In a specimen of Apheliscus (U.S.N.M. No. 19162), these foramina were noted beneath posterior portion of P; and beneath P,. In Aphronorus the posterior foramen may be small and varies in position from beneath M, to the posterior part of P,. An equally large or larger anterior opening is seen below P.. In Bessoecetor foramina were noted beneath the posterior part of both M, and Pz, and in one specimen, U.S.N.M. No. 9442, anterior fora- mina were observed below the posterior portions of both P, and Ps. In Bridger Pantolestes I have seen only the well-developed foramen beneath M,. Among the Insectivores outside of the Pantolestidae I find a rather more remote relationship to the mixodectids indicated. There would appear to be rather less to suggest affinities with other orders. Among these, however, perhaps the condylarths should be considered. The relatively low trigonids of the lower molars seem indicative of a pos- sible condylarthran relationship, and a form such as Choeroclaenus among the mioclaenine hyopsodonts is not too different from Phenaco- daptes but there is, nevertheless, a more inflated appearance to the molar cusps and the premolars would appear to have little or nothing to recommend them. The possibility that the Phenacodaptes—A phelis- cus line represents condylarth development rather paralleling that of pentacodonts cannot be entirely disregarded, but the same reasoning might apply equally well were they to be regarded as belonging to such other orders as primates, creodonts, or artiodactyls. Comparison with Pentacodon and Aphronorus appears rather more pertinent and better accounts for a number of minor details of similarity not easily dismissed. REFERENCES Corr, Epwarp D. 1874. Report upon the vertebrate fossils discovered in New Mexico with descriptions of new species. Geogr. Expl. and Surv. West of 1ooth Meridian (Wheeler), Appendix FF, Ann. Rep. Chief of Engineers, 1874, pp. 1-18. 1875. Systematic catalogue of Vertebrata of the Eocene of New Mexico, collected in 1874. Geogr. Expl. and Surv. West of tooth Meridian (Wheeler), pp. I-37. 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 1877. Report upon the extinct Vertebrata obtained in New Mexico by parties of the expedition of 1874. Rep. U. S. Geogr. Surv. West of 1ooth Meridian (Wheeler), vol. 4, pt. 2, pp. i-iv, 1-370, pls. 22-83. JEPSEN, GLENN L. 1930. Stratigraphy and paleontology of the Paleocene of northeastern Park County, Wyoming. Proc. Amer. Philos. Soc., vol. 69, No. 7, pp. 463-528, figs. 1-4, pls. I-10. MatTrHew, WILLIAM D. 1909. The Carnivora and Insectivora of the Bridger Basin, Middle Eocene. Mem. Amer. Mus. Nat. Hist., vol. 9, pt. 6, pp. 201-567, figs. 1-118, pls. 42-52. 1918. A revision of the lower Eocene Wasatch and Wind River faunas. Part IV. Entelonychia, Primates, Insectivora (part). Bull. Amer. Mus. Nat. Hist., vol. 34, pp. 429-483, figs. 1-52, pls. 42-52. McDoweELt, SAMUEL B., Jr. 1958. The Greater Antillean insectivores. Bull. Amer. Mus. Nat. Hist., vol. 115, art. 3, pp. 115-214, figs. 1-46, tables 1-2. McKenna, Matcoim C. 1955. Age of the Four Mile local fauna, northeast Sand Wash Basin, Colo- rado. Wyoming Geol. Assoc. Guidebook, roth Ann. Field Confer- ence, Green River Basin, pp. 105-107, fig. 1. SABAN, ROGER. 1954. Phylogénie des Insectivores. Bull. Mus. Nat. Hist. Nat., ser. 2, vol. 26, No. 3, pp. 419-432. Simpson, GEORGE G. 1937a. The Fort Union of the Crazy Mountain field, Montana, and its mammalian faunas. U. S. Nat. Mus. Bull. 169, pp. 1-287, figs. 1-80, pls. I-10. 1937b. Notes on the Clark Fork, upper Paleocene, fauna. Amer. Mus. Nov., No. 954, pp. 1-24, figs. 1-6. 1945. The principles of classification and the classification of mammals. Bull. Amer. Mus. Nat. Hist., vol. 85, pp. i-xvi, 1-350. NO. 7 TERTIARY APHELISCUS AND PHENACODAPTES—GAZIN 7 EXPLANATION OF PLATES PLATE I Apheliscus and Phenacodaptes from the early Tertiary of Wyoming Figs. 1, 2. Apheliscus insidiosus Cope: 1, Right ramus of mandible (U.S.N.M. No. 19162), lateral and occlusal views. 2, Left ramus of mandible (A.M. No. 15696), lateral and occlusal views. All four times natural size. Gray Bull lower Eocene, Big Horn Basin, Wyoming. Figs. 3, 4. Phenacodaptes sabulosus Jepsen: 3, Right ramus of mandible (P.U. No. 13926), lateral and occlusal views. 4, Left ramus of mandible (P.U. No. 13391), lateral and occlusal views. All four times natural size. Silver Coulee (Tiffanian) upper Paleocene, Big Horn Basin, Wyoming. PLATE 2 Apheliscinae and Pentacodontinae from the early Tertiary of the Rocky Mountain Region Fig. 1. Apheliscus insidiosus Cope: Left upper cheek teeth (A.M. No. 15696), occlusal view. Four times natural size. Gray Bull lower Eocene, Big Horn Basin, Wyoming. Fig. 2. Phenacodaptes sabulosus Jepsen: Right upper cheek teeth (P.U. No. 13977), occlusal view. Four times natural size. Silver Coulee (Tiffanian) upper Paleocene, Big Horn Basin, Wyoming. Figs. 3, 4. Aphronorus fraudator Simpson: 3, Right upper cheek teeth (U.S.N.M. No. 9561, Ps from U.S.N.M. No. 9564), occlusal view. 4, Left ramus of mandible (U.S.N.M. No. 6177, type specimen, with molars restored from U.S.N.M. No. 9289, Pi: to Ps from U.S.N.M. Nos. 9537 and 9201), lateral and occlusal views. All four times natural size. Fort Union middle Paleocene, Crazy Mountain area, Montana. Figs. 5, 6. Pentacodon inversus (Cope): 5, Left upper cheek teeth (U.S.N.M. No. 15502), occlusal view. 6, Left ramus of mandible (A.M. No. 17038), lateral and occlusal views. All twice natural size. Torrejon middle Paleocene, San Juan Basin, New Mexico. hy eee we five. SMITHSONIAN MISCELLANEOUS COLLECTIONS VOE 139; NO: 7, IP, 2 APHELISCUS AND PHENACODAPTES FROM THE EARLY TERTIARY OF WYOMING (See explanation of plates at end of text.) SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL 1397) NOS 7) Pez APHELISCINAE AND PENTACODONTINAE FROM THE EARLY TERTIARY OF THE ROCKY MOUNTAIN REGION (See explanation of plates at end of text.) et J Se ae ae ee Se ee = gy NT Le Te . oo & fs a Ree AIA Sy SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 139, NUMBER 8 THE ANATOMICAL LIFE OF THE MOSQUITO By R. E. SNODGRASS Research Associate Smithsonian Institution ase OOseng, "0 f i CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION NOVEMBER 4, 1959 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 139, NUMBER 8 THE ANATOMICAL LIFE OF THE MOSQUITO By R. E. SNODGRASS Research Associate Smithsonian Institution (Pustication 4388) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION NOVEMBER 4, 1959 THE LORD BALTIMORE PRESS, INC. BALTIMORE, MD., U. S. A. CONTENTS Page Ug AOVaLE TSS OTD RIS EHO GO OGD SO DCO TIERCE Bier ha casio 6 bt a ETAT aoe I Pee mila revere sets ainlafocrererecer a ere och ois nia eraye To ole ee SNA NERD eceteatoaat alate 2 Mhew headin yee eR yaAR ae rh atta eae o's ie eae eiao ete aeeie 4 The teed NSsOLRANS rytecrcte oie sistas. ctovere Gee iwik svejaictomrmoreiarsiere aateterere Il Me MIAMI eos « Sig diee's isis g eire mien lava lo rotemrcteree ta ate psrerershotale 12 he preatal. Cavaty sec vi Noes cone Gels © a acatiomarcie sels maleye t's 15 The epipharyngeal apparatusss si: julc tocar cab eelee © eimcieie 15 AM hekmiandi bles Aves arate wisiersie: aeierel nee ei entterenmen evoke ome tater 16 ARS arias Te hai see eyetey bel otepoueere ceousuaro bie. 3 cfpanseoaraed wate rate ets 17 The labrum and hypopharynx< ws... cc ss cieai epee eas 19 SDE SP MATIV UR |, sisi crt eecicre tase ore ober tresoiers a) ays: eSNG autora iarete sie eles 21 Warvaltteeditigs toss ne oa ceteris tere dd abin teeter he enteric. 22 ADRS T CHOTA Ca cA Ae Se ey ere crea ete alevsicia wla-c suslale ERT Selstene ane tials 24 ible ANdaMieny «5 HIV Tha ete aka les, «ced SRM ere ete ee 26 Intennalganatoriy, Gfevtcs ceeeve feelin ie cies stoi s olaernkenstebas sie eiereiaiece 21 ‘Whew itrachealmsystenns wasn e elo, oe oetee le elaevaats aoe helve ZT ‘phe dorsal bloodiaviessel yemrverecrers asicva eee eeere a cores 32 aPhewalimentarya Catiale were se tawe tale: « th ete ces eerclale ey arre erate 32 MheeMalpightanetupulesescaas te cmice shoo eae tees a2 AbheNsalivatymolatid serra sire cels ino Richtee talet ote erons eet 33 AREY MER VOUSH SY.SECIM csyersiey sie reis oreiks ho ohn ae ane aaieh s 34 PRE} FEPrOGUCtiVe OL GANS) «We iale.././«,« ssa isin, Sedscereme lene soins ohans 34 I OOG PESCE VES a tole satan Iai5 "Lo clay idee ousie s,s o/s ercem arlene ate tetnelalees ovate 34 WIE, CMTE APU Tees core c.ciakare ecole scree e aati d clave wee Oars Meme ate cicaaaaats 34 he mpupal developments... Lee sees cies a os stale oe ee lore eile aati 35 Aheematine ppupareyidc solace setae cs oko ec sltieteteis. oavaveteomro tts 39 GeneralVexternall stature ia acie.sie cia. oni teeta 39 The head and amiauth) Parts ch iciustss + os.0 Ghereake aiucinaen’s clueve 43 PINE MEH O tela eh se eerie caterer ete rar seni ehcee cceaic Ace eRT Re ere ete 44 PBheyabdomenly arene setae wiaciete cis Nace ce Nee eee ore etree 45 Ache pupal’ metamoOnpuOsiss,. sheaves. cess hon be eee en ae ee 46 Tela ul he via ttl tice evreratch Shae Hove Mahe tela 8 a ap aaa ie a Se ad 48 AEM HEA sive crcveoe Such eve ca ciey ciate eek l ale rol Gre soa eases oor ETTORE AITO SI PENENOTSANSIOT ACCUM 21a vies us cies) a) ocho bis eu Aue ice SiG aleie c 55 PUG PU OMOSCIS ala 3 cle cis te anor a Leratienet sere. 270) Sachets tomamtene naire Ss cars 55 a exci Daria DUBIN Ne feces cake Saks es cis eters Mere or 60 hekpharyngeal: punipaces isosceles os ERE eee 63 MUNG PROP AS erate aid erarei te acd ute: RAN. 0 |, UNINC 5 le i, 64 sihrerabd omen srs easter asa aicrsiew io sols oleae RO CRIEC Rae icheee 67 internal anatomiye «sscrec sh as Lae a 63) 52.4.0 :0's geen inetd 72 PHS CALCINACOE We (ORE ANGY (20 eaves! o1sne' sce 3 0 sel ammeto nelle eniaks Mats 73 Dhevalimentany. (canal ours ciers crores iaye soo Ao eles ots hee 73 mher salivary) wands ivy sad Hos) SA ORR A oe 76 Dhe reprodtictiver-sy ster). tie: 4 scsi c'e sea en ae claialeieers.cie its 76 IRGLEL ER COSMET ieee SED crac a aired a tee Khaleel isle cheer Oe oh aieiticlats 80 i iy y, Toe den ' WEY, Fela y [6 ee MR Pier ye ed eae Wa ee Mi Ae PRA NR eo e } ¥FY Shia ae A eAN fa .* Na ee A; aM i COUN ME EWE S44 ‘ i a i) H er, ay esiomd rete 1a ha ‘ Cul) Aaa, 3° i ‘ea, oP ection na VMN 4 Mt a MI ye LR 2! Ot Oe any DORIS 1% erty) 7 Sree SMA Le Reo Me Pm ea ey i me ; ae eh Te ree whe ayia Y's FR i , % ey ie hoe bb! MY f . ‘hy VE A ey Oy, xy de) yay ma eee tail bike! ack dA AW Gt in i ‘ a ih& : Reale Variehk fe ghee , Oy bh : ig ; ht t : i ‘ 1 DF rh. |, hake tobe ODS 6a 8 On LR ea we oe ee ey ob cr j : Ale ; alk powekie take ad Dre - Ns ip ¢ \ 8 ee MCU REPMN NT ad mic dy ey a) a 8 YORE As hy nh 4 ! ee) pa Sik ae ve eayeUN a w eiieramaea 9) eta Best hss PR eg eee Mil W ed 9 So asta ¥ PE DIRS eM A wea ye ie bees a to Reh iin eee Mae yt Ribs eh ‘ Re OP Ree he Rae be a j el y ST AT ee Em eS VOR Ts yy (ot LM Eee , i 6 ae Ea Pee ee ie bk ee Wi are aM eae Pi Saye bh pa MT ale ae A R » ad pay Te TS OR Ree a Te ERO ER te me rey : ; } i FeETER Pa a ee a MO Ae Me ae con) | pe bidi, me be ; ret a ea We aed Pee PNRM ork ae WE OU A t U teL 9 ee Ee Pt Peet Wal wR Fon NO ag a hoy ae Pe CL eh ere Tad eae Ree ee ee SORRY AA hen 1S ee Pel ys eee : Vi MAES AAT ht a8 DAVOS 7 F400 ON ele ORY ets ah Ae ae OR NS 4 ee Sala eH oe + ' uf AS 7 aye ee ra ay 8 eek Phe a ew ie eu eas te ins 4! yikes Hew RN made yo RatamN ey (iF ay ava Nh Pak oaks ALN A ilk i) : ee eS Ty k aly eas i gg bhi oe R2SRS2SSStR se Tre ANATOMICAL’ PIPE OF THE MOSOUTITO By R. E. SNODGRASS Research Associate Smithsonian Institution INTRODUCTION Mosquitoes are not popular with warm-blooded animals, but from their own standpoint they have been highly successful insects, until recently when they have been attacked with poison sprays and have had their larval habitats drained. Success, however, is always to be admired whether in man or an insect, and it is instructive to see how it has been achieved. The mosquitoes have attained their place in the world by the evolution of highly specialized anatomical characters. A study of their anatomy may help some in our war against them, and it will give a most interesting example of how insects have evolved structures fitting them for particular ways of living and of feeding that have made them so successful in the struggle for existence. The family name of the mosquitoes is Culicidae, and they belong to the order Diptera, or two-winged flies, which in turn are members of that large group of insects in which the young, or larvae, are very different from their parents in form, structure, and habits, and must undergo a renewed growth to attain the adult state. We are so ac- customed to seeing young animals grow up gradually into adults that it seems very remarkable that an animal can completely change its shape and structure in the middle of its life. The young mosquito, for example, hatches from the egg as an active larva having no resem- blance to its parents but fully adapted in its structure for living and feeding in the water. During its life the larva sheds its cuticle four times. At each of the first three ecdyses it comes out a little larger than before, but with little change otherwise. On shedding the fourth cuticle, however, a very different creature, the pupa, emerges. The pupa has all the adult organs, though in an incomplete state of de- velopment, and is clearly a preliminary adult. With a final moult and ecdysis the completed mosquito appears, equipped for an entirely dif- ferent life from that of the larva. It is commonly said that the larva is metamorphosed into the adult SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 139, NO. 8 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 during the pupal stage. Actually it simply returns to its parental adult structure after having undergone during its evolutionary history a metamorphosis by which it took on a form and structure suited to a way of living quite different from that of its parents. The embryo, from its very beginning in the egg, develops into a larva. The egg, therefore, contains two distinct hereditary factors, one that first produces the larva, another that later generates the parent adult. When the larva does not differ too much from its parents, the adult may be formed mostly by a new growth of the larval tissues ; but as the differ- ence becomes more extreme, the larval tissues go into a state of dis- solution and the adult is built up of embryonic cells that multiply but do not become organized during the larval stage. The transformation of the mosquito is intermediate between these two conditions. Inasmuch as the word metamorphosis means simply a “change of form,” we may say that the larva in its aberrant evolution has under- gone a divergent metamorphosis, and that as an individual it resumes the parental form by a convergent metamorphosis. Since the egg has the potentiality of developing into both the larval and the adult form, there must be some influence that allows the larva to develop first. The inhibition of adult development is effected by a hormone, known as the juvenile, or status quo, hormone. When the larva is mature and has served its purpose in the life of the insect, this hormone ceases to be effective, and the adult development pro- ceeds under the stimulus of another hormone. This at least is the usual story of endocrinal regulation of insect growth and transforma- tion, but, as will be seen, the mosquito does not comply fully with the rules of hormone control in its growth from larva to adult. Before going on with anatomical descriptions of the larval, pupal, and adult stages of the mosquito, a few terms should be defined as they will be used. An instar is the insect between any two consecutive moults. Moulting is the physiological process of separating the old cuticle from a new cuticle being formed by the epidermis beneath it. The new instar begins its development when the moult is completed, but remains inside the old cuticle until it is fully formed. Then it breaks the cuticle and comes out. The emergence of the insect is its ecdysis (coming out). Moulting and ecdysis, therefore, refer to two different events, and are not synonymous terms, though many en- tomologists have not distinguished them as such. In life-history studies the “instar” is usually regarded as the insect between ecdyses, but since development begins inside the old cuticle, an instar is really the insect between moults. The concealed intracuticular period of the no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 3 instar has been appropriately named by Hinton (1946, 1958a) the pharate, or cloaked, stage of development. The pharate stage of the pupa in the larval skin is commonly called the “prepupal stage of the larva,” but the larva has already ceased to be a larva, so the expres- sion does not conform with the facts. The mosquito will demonstrate a number of other errors commonly made by entomologists. The problem of explaining how an animal in its evolution has be- come adapted structurally to its environment and a special way of living is complicated in an insect such as the mosquito that lives two entirely different lives. If adaptation affects two or more organs separately, the matter is relatively simple, but when it involves coadaptation in all parts of the animal, it is hard to understand how evolution by means of natural selection has brought it about. On the other hand, the technique of “special creation” is entirely incompre- hensible. The writer began this work on a very meager acquaintance with the anatomy of mosquitoes, especially of the larva and pupa. For its completion he is deeply in debt to others, in particular to Dr. Alan Stone and Dr. Richard H. Foote at the U. S. National Museum for literature and the identification of species; to Dr. Paul Woke of the National Institutes of Health at Bethesda, Md., for an abundance of live larvae; to Dr. Ernestine B. Thurman, also of the Institutes of Health, and Dr. Jack Colvard Jones of the University of Maryland for much supplementary information and a critical reading of the manuscript ; to various authors for copied drawings; and to Mrs. R. E. Snodgrass for the typing. For morphological interpretations the writer assumes entire responsibility. T Loe CARVA Mosquito larvae hardly need an introduction. They are the familiar aquatic “wrigglers” or “wigglers” that everybody knows turn into mosquitoes. Anatomically the most specialized parts of them are the head, the feeding organs, and the respiratory system. A number of good papers have been written on the larval anatomy, and the facts of structure have been well-enough described, but the writers, par- ticularly on the head and feeding organs, mostly disagree as to the homologies of the parts, and consequently the different terminologies used must be very confusing to students. Hence, in the following text, the larval head and organs of feeding are given a disproportionate amount of space in an effort to arrive at reasonable interpretations and an acceptable terminology. Otherwise than in the head and feed- 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 ing apparatus, the principal specialization of the larva pertains to the respiratory system. The only functional respiratory apertures are a pair of dorsal spiracles near the end of the abdomen, the lateral spiracles being closed except at ecdysis when the tracheal linings are partly pulled out through them. THE HEAD The head of a mosquito larva projects forward from the thorax in line with the axis of the body, bringing the mouth parts to an anterior position. In most adult insects the head hangs downward on the thorax, so that the face is anterior and the mouth parts ventral. In the prognathous mosquito larva the face becomes dorsal and the mouth parts anterior. In going from adult to larva, therefore, instead of reversing the meaning of “dorsal” and “ventral,” it will be better to speak of the upper and lower surfaces of the larval head, though “anterior” and “posterior” in either larva or adult will be directions relative to the axis of the body. The typical shape of the mosquito larval head is oval or ovate, whether seen from above (fig. 1 A,B,C) or from the side (E), but the upper surface is more rounded than the lower. In some species, however, the head is almost rectangular in form (D). Anteriorly the head bears laterally a pair of large mustachelike brushes, and usually between them a small median brush, the three being supported by the labrum. Shortly behind the lateral brushes arise the slender, tapering, unsegmented antennae (E, Ant). Posteriorly on each side of the head is a large dark spot (£) varying in size with the age of the larva. These spots are the pigmented compound eyes of the adult developing in the epidermis beneath the larval cuticle. Behind or below each compound eye is a small, simple, presumably functional larval eye (O). The lateral area of the head between the antenna and the eye is the gena (Ge), that behind and below the eye the postgena (Pge). Posteriorly the head abruptly narrows to the occipital fora- men, which is rimmed by a darkly sclerotized band, the postocciput, set off by a postoccipital sulcus. The membranous neck is usually cylindrical (fig. 1 A), but in Anopheles (fig. 3 C) it is narrowed where it joins the thorax, evidently to facilitate the turning of the head upside down while feeding. The upper surface of the head (fig. 1) is differentiated into a large, shieldlike central area, narrow lateral areas bearing the antennae and the eyes, and a slender transverse anterior sclerite at the bases of the brushes. This sclerite (A,B, Lm) is the dorsal wall of the labrum, No. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 5 as contended by Cook (1944a), though some writers have regarded it as a “preclypeus.’”’ The groove behind it (A, cls) then is the clypeolabral sulcus. The large central area of the head is bounded by lateral lines (CL) that diverge forward from a very short occipital cleft and become continuous with the clypeolabral sulcus. These lines, commonly called “frontal sutures,” are merely lines of weakness Fic. 1.—The larval head, dorsal and lateral. A, Aedes aegypti. B, Culex sp. C, Anopheles quadrimaculatus. D, Toxo- rhynchites rutilus. E, Culex sp., lateral. F, Anopheles farauti, head exuviae. Ant, antenna; Apt, cephalic apotome; CL, cleavage line; cls, clypeolabral sulcus; E, compound eye; Ge, gena; Hstm, hypostomium; Lm, labrum; Md, mandible; Mx, maxilla; O, group of larval simple eyes; Pge, postgena. in the cuticle where the latter will split at ecdysis (F, CL) to allow the emergence of the next instar, and are best termed the cephalic cleavage lines. In most young insects the cleavage lines take the form of a Y, which has been known as the “epicranial suture.” In the mosquito larva the stem of the Y is the short occipital cleft. The frontal arms follow such very different courses in different insects that they can have no morphological significance (see DuPorte, 1946; Snodgrass, 1947), and hence do not define any specific part of the 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 head. The part of the head wall cut out at ecdysis may be termed the cephalic apotome (F, Apt). The space between the arms of the cleavage lines and the clypeolabral sulcus in the mosquito larva has been variously called the “frons,” the “clypeus,” and the “frontoclypeus.” The respective areas of the frons and the clypeus may be identified in other insects by specific groups of muscles that arise on them. In the mosquito larva the clypeal muscles arise anteriorly, the frontal muscles posteriorly on the central head area, but there is no external demarcation between the two regions. This area, therefore, is frontoclypeal in a limited sense, but it is not the entire clypeus or the entire frons. Ordinarily the clypeus extends laterally to the bases of the mandible, and in adult insects the frons is the facial area between the antennae and the eyes. The whole aspect of the mosquito head is changed at the transfor- mation to the pupa and the adult. The larval antennae are slender, unsegmented shafts bearing vari- ously distributed spines and tufts of long hairs. Each terminates in a small apical papilla. The antenna of the pupa, being eventually much larger than the larval organ, does not develop within the latter but in a pocket extending posteriorly from the base of the larval antenna. The eyes of the larva are each a group of simple lateral eyes; their structure in Culex pipiens has been described by Constantineanu (1930) and by Sato (1951b). According to Sato each eye consists of three parts, each with its own retinular cells. One part is central and has three retinulae, a second part is dorsal and has a single retinula of eight cells, the third part is a long band of about 40 cells surround- ing the other two parts dorsally, anteriorly, and ventrally. Con- stantineanu, on the other hand, describes five parts in the eye of Culex, as in some other nematocerous larva. Probably the three retinulae of Sato’s “central part’? are regarded as three eyes. The larval eyes have no lenses, the ordinary head cuticle being continuous over them. They are present from the beginning of the larval stage and persist into the pupa, or even into the adult. The presence of large, darkly pigmented compound eyes visible on the surface of the head gives the mosquito larva, as also the corethrid larva, a very unusual appearance. The compound eyes of other re- lated Nematocera are developed likewise in the larva, but, because of the absence of pigment until the pupa stage, they are not apparent ex- ternally. The undersurface of the larval head (fig. 3 C,D,E,F) is more dif- ficult to understand than the upper surface. The mandibles and No. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 7 maxillae are articulated on a transverse margin between the bases of the antennae. The long ventral cranial wall behind them is sclerotically closed by the union of the postgenae (C, Pge) along an incomplete median suture (C,D,E, ms). This same condition occurs in certain other insects, and to understand how it has come about we shall have to digress on some comparative studies. The hypognathous position of the insect head in which the mouth parts are ventral (fig. 2A) is clearly primitive, because the mouth parts, being modified legs, thus hang down from the head in the posi- tion of the thoracic legs. The prognathous condition has been attained in some cases by a mere turning forward of the head on the neck, involving a ventral elongation of the occipital foramen on the under- side of the head (fig. 3 B). More commonly, however, the foramen remains in the vertical plane, as in the mosquito larva (fig. I E), and the underside of the head is lengthened by a ventral elongation of the postgenae (Pge). With the elongation of the postgenae the entire labium, as in some beetles (fig. 2 B), may be simply enclosed between them, with a gular addition (Gu) to the submentum. This condition, however, does not occur in the larval mosquito, though some writers have so interpreted the mosquito head structure. More commonly, the postgenae come together medially and displace the labium. A primary stage of this transformation is seen at C, which might represent the head of a caterpillar or an adult honey bee, in which lobes of the hypostomal margins of the postgenae are intruded between the occipital foramen and the base of the labium. In other cases the lobes become united (D), forming a bridge between the foramen and the labium. An elon- gation of the bridge then produces the condition seen in the beetle larva at EF, in which the labium is still fully exposed. Finally, as in the larvae of Chironomidae (F), the labium has become greatly reduced and is hidden from below by a median hypostomal lobe (Hstm) of the united postgenae. This same process of closure and elongation of the postgenae and the reduction of the labium can be traced among nematocerous fly larvae. For example, in the primitive rhyphid larva of Olbiogaster (fig. 3 A) described by Anthon (1943b), a pair of small postgenal lobes are approximated behind the submentum (Smt) of the labium. In others, as in Trichocera and Philosepedon figured by Anthon (1943a, figs. 7, 10) the postgenal lobes are united in a bridge; the labium, though much reduced, is still mostly exposed. In the mosquito larva (fig. 3 C) the united postgenae form the long underwall of the 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 cranium and the greatly reduced labium is concealed above a median postgenal lobe (stm) between the maxillae (Vx). The darkly sclerotized dentate lobe between the maxillae has com- monly been regarded as a part of the labium, “mentum” or ‘“‘submen- Fic. 2—Structure of the posterior or ventral wall of the head in different insects, mostly diagrammatic. A, Generalized structure of the posterior head wall and the mouth part at- tachments. B, The labium enclosed between the postgenae. C, The labium separated from the occipital foramen by intruding hypostomal lobes of the post- genae. D, The labium entirely separated from the foramen by union of postgenal lobes forming a postgenal bridge. E, Postgenal bridge lengthened. F, Larval head of Chironomus, postgenal bridge produced in a toothed lobe, labium dis- placed dorsally. For, occipital foramen; Gu, gula; hs, hypostomal sulcus; Hstm, hypostomium, hypochilum; Lb, labium; Md, mandible; ms, median postgenal suture; Mt, mentum; Mz, maxilla; Pge, postgena; pos, postoccipital sulcus; Prmt, pre- mentum; pf, posterior tentorial pit; Smt, submentum. tum.” To cut a long argument short, however, we have only to look at a tipulid larva (fig. 3 B) to see that the lobe is formed by the union of two processes extended forward from the anterior median angles of the postgenae, which themselves are not united in the tipulid. Above no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 9 this lobe are the united labium and hypopharynx (fig. 7 A, Lb, Hphy). We may, therefore, following Anthon (1943a), Hennig (1948, 1950), and Lawson (1951), appropriately call this lobe the hypostomium (Hstm), as it is termed also by Chiswell (1955) in the tipulid larva. Though Schremmer (1949) called it “mentum” in the mosquito larva, he later (1950) expressed doubt of the correctness of this designa- tion, concluding that the lobe is rather a part of the cranial wall. More recently, Gouin (1959) has termed the dentate lobe the hypochilum (underlip). From the base of the hypostomium there arises in some species a thin fold bearing a fringe of pectinate hairs or blunt teeth (fig. 15 A, Aul). The fold is the aulaeum (curtain) of Cook (1944a), but it has been variously named. Shalaby calls it the “glossa” on the assumption that it is formed by the union of a pair of labial glossae, a highly im- probable interpretation since the hypostomium itself is no part of the labium. However, Shalaby has given detailed illustrations of the pectinate hairs of the lobe in Aedes aegypti (1957a) and Culex quinquefasciatus (1957b), and its armature of eight blunt teeth in Anopheles quadrimaculatus (1956). In Psorophora ciliata (1957c) he says the fold is absent. In most mosquito larvae two dark lines in the ventral wall of the head diverge posteriorly from the basal angles of the hypostomium. In some species the lines are short (fig. 3 C,D, 7), in others (E,F) they extend back to the posterior tentorial pits (pt); in Chironomus (fig. 2 F) they are absent. These lines when present are the external marks of internal ridges; their variable development suggests that the ridges are secondarily formed to strengthen the head wall. The surface area between the lines, however, has commonly been regarded as the basal part of the labium, probably because the structural pattern they produce resembles that of the head shown at B of figure 2. It has been suggested even that the median suture is the line where the two origi- nal labial appendages have united! Cook (1944a), for some obscure theoretical reason, calls the area in question the “maxillary segment,” though the maxillae have no relation to it. That the ventral closure of the head results entirely from the union of the lateral cranial walls is clearly indicated in illustrations by Hennig (1948, figs. 31-37) of larval heads of Sciophilidae, in which are shown various degrees of approximation and union of the postgenal margins. In most adult insects the lower edges of the cranium are reinforced by submarginal internal ridges formed by external grooves known as the subgenal sulci. The part of each groove on the postgena behind Io SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 the mandible is distinguished as the hypostomal sulcus (fig. 2 A, hs). Posteriorly these grooves become continuous with the postoccipital sulcus (pos) that surrounds the occipital foramen. In the mosquito larva the lower ends of the postoccipital sulcus have extended forward in the postgenal region carrying with them the minute rudiments of the posterior tentorial arms, the position of which is marked externally by a pair of pits (fig. 3 E,F, pt). The anterior tentorial arms are extremely slender bars arising from the cranial margins mesad of Fic. 3—The larval head, undersurface. A, Olbiogaster sp., Rhyphidae (from Anthon, 1943b). B, Tipula sp. C, Anopheles quadrimaculatus. D, Toxorhynchites rutilus. E, Culex sp. F, Aedes aegypti. r, r, grooves in lower head wall. Other lettering as on figure 2. the antennae that extend back to the posterior arms. Possibly it is the great lengthening of the postgenal regions of the head that has brought the posterior arms to their forward position. The postgenal bridge is known also as the hypostomal bridge be- cause it is the hypostomal margins of the postgenae that come together to form it. Lawson (1951) contends that the sclerotized ventral wall of the head behind the mouth parts cannot be derived from the post- genae because, he says, “the hypostomal sutures form the ventral No. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS II boundaries of the postgenae.” This is clearly making anatomy con- form with definitions. The lower parts of the postgenae are mechani- cally strengthened by ridges formed by the submarginal hypostomal grooves. The narrow strips below the grooves, therefore, are simply the marginal parts of the postgenae, so it is immaterial whether we call the bridge resulting from their union hypostomal or postgenal. The grooves are sometimes absent, and the ridges may be marginal on the postgenae. In the mosquito larva the anterior edge of the ventral cranial wall on which the mandibles and maxillae are articulated is the united hypostomal margins of the confluent postgenae. From the free cranial margins just mesad of the antennal bases, a slender bar on each side (fig. 7 B, hb) extends mesally, downward, and somewhat posteriorly through the preoral epipharyngeal wall to the base of the hypopharynx (Hphy). Each bar runs close before the mandible of the same side and goes below the narrow lower lip of the mouth (Mth). In Dixa, as shown by Schremmer (1950), similar structures are present but are much wider than in the mosquito larva. The mandibles have their anterior articulation on these rods, a very unusual condition, since the anterior mandibular hinges are typically on the basal angles of the clypeus. The rods have been called “cibarial bars,” but there is no defined cibarium in the culicid larva. Since the rods appear to serve principally as suspensoria of the hypopharynx, they are here termed hypopharyngeal bars. They are the Verbindungs- leisten of Schremmer (1949). Since the hypopharyngeal bars carry the anterior articulations of the mandibles, Menees (1958b) reason- ably argues that the parts of the bars laterad of the articulation are extensions of the clypeus. His identification of the posterior parts with the “hypopharyngeal suspensorial bars of generalized insects,” however, is less convincing, since these bars enter the mouth angles and give attachment to the hypopharyngeal muscles, though each may have a lateral preoral branch. THE FEEDING ORGANS One of the remarkable things about insects is the way their feeding organs are variously adapted to feeding in different ways on different kinds of food. Nothing comparable occurs among the vertebrates, their only adaptation to the nature of their food is in the size, strength, and dentition of their jaws or in the length of the neck. Yet the feed- ing organs of all insects are made up of the same fundamental parts. There is an upper lip known as the labrum, a pair of mandibles, a median tonguelike hypopharynx, a pair of ma-illae, and a lower lip, or I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 labium, composed of a united pair of second maxillae. The mandibles, maxillae, and labium, furthermore, have been fashioned from three pairs of legs, since the original arthropods had no other organs for feeding than their legs. The insect mouth parts, therefore, are all outside the mouth; the space between them may be termed the preoral food cavity, but by a long-perpetuated error it has commonly been called the ‘‘pharynx.” For want of a revised nomenclature we still speak of the upper wall of the preoral cavity as the epipharyngeal surface, and call the tonguelike lobe that projects below the mouth the hypopharynx. This is just a part of our heritage from the early insect anatomists, who had only vertebrate names to draw from, and applied them to insects on a functional rather than a morphological basis. The true pharynx is a part of the stomodaeal section of the alimentary canal behind the mouth. The labrum.—The labrum of the mosquito larva includes the small transverse sclerite on the dorsal wall of the head before the clypeus (fig. 1 A, Lm), and a larger membranous undersurface that bears laterally the two vibratory feeding brushes (fig. 4B), and usually a small median brush. The median brush is the “palatum” of mosquito students, another example of misuse of a borrowed vertebrate name, which in this case properly refers to the roof of the mouth cavity. The lateral brushes of the labrum are the organs by which those larvae that feed on particles create currents in the water directed toward the head, and drive a stream of water back to the mouth along the epipharyngeal surface. The individual hairs of the brushes are finely pectinate and serve also as combs for retaining particles filtered from the water. The vibratory movement of the brushes is produced by a pair of strongly musculated sclerites on the under side of the labrum. Simi- lar sclerites are present in the larvae of Chironomidae (fig. 4 G, Tor), which have no brushes, but the posterior ends of the sclerite are produced into strong pointed processes (Mes) projecting freely from the epipharyngeal surface. These toothed sclerites were therefore called by most earlier writers ‘“premandibles.”’ Chaudonneret (1951), however, has shown that this term is entirely inappropriate. Cook (1944b) named the sclerites “messores’” (harvesters) and carried the term over to the mosquito larvae, in which he has been followed by several recent writers, though the culicid sclerites are unarmed. It must be noted that the insect labrum is commonly equipped with four muscles, one pair dorsal, the other ventral, all of which arise on the frons. The ventral muscles are usually attached on a pair of no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 13 sclerites in the lower labral wall known as the tormae. In a tipulid larva (fig. 4A) the tormae (Tor) are simple sclerites, each giving attachment to a long muscle (mc/) from the frons. There is, there- fore, no apparent reason why the similarly musculated sclerites of the mosquito larval labrum (C,D, Tor) should not be the tormae. On the other hand, Cook (1944b) has contended that the sclerites are Fic. 4.—The larval labrum and tormae. A, Tipula sp., underside of larval labrum. B, Culex sp., labrum of young larva, anterior. C, Anopheles quadrimaculatus, left labral brush and torma, undersurface. D, Aedes aegypti, same. E, Toxorhynchites rutilus, anterior view of larval head. F, Same, labral brush with torma and muscles. G, Chironomus plumosus, underside of larval labrum. Ap, tormal apodeme; B, labral brush; Hstm, hypostomium; Lm, labrum; mcl, mels, tormal muscle or muscles; Md, mandible; Mes, messorial teeth of torma; Myr, maxilla; Tor, torma. a, connective sclerite between torma and brush; b, c, detached sclerites of cena wall; d, e, anterior and posterior articulations of torma; f, epipharyngeal ar. not the tormae because he finds in chironomid larvae another pair of muscles attached more dorsally and laterally on the labrum, which he insists are the true tormal muscles. These muscles, however, would appear to be the usual dorsal muscles of the labrum, which may have a lateral position. Furthermore, Cook adds that the ventral muscles are not tormal muscles because they arise on the clypeus, but what he calls 14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 “clypeus” is the entire frontoclypeal area of the upper head wall be- tween the cleavage lines. The sclerites in question, being in the ventral wall of the labrum and giving attachment to the ventral labral muscles, practically identify themselves as the tormae, and they have been regarded as such by Anthon (1943a), Schremmer (1949, 1950), and Menees (1958b). If it is desirable to keep the term “messor,” it might be restricted to the free prongs of the tormae where they occur (fig. 4G, Mes). The tormae of an Anopheles larva (fig. 4C, Tor) are elongate sclerites lying mesad of the brushes. Each torma is connected by its tapering anterior end with the base of the corresponding brush; posteriorly it is hinged to a small sclerite (c) in the cranial margin. A connective plate (a) lies between the torma and the base of the brush. A single muscle (mcl) from the frontal region is attached by a long tendon to a small point anteriorly on the lateral margin of the torma. Cook (1944a) ascribes a second posterior muscle to the torma of Anopheles, but this muscle, as shown by Farnsworth (1947) and by Schremmer (1949), belongs to a V-shaped sclerite of the epipharyn- geal wall between the posterior ends of the tormae. In the culicine mosquitoes the tormal apparatus is somewhat more complex than in Anopheles. In Aedes aegypti (fig. 4D) the tormae have the same relation to the brushes and the cranial margin as in Anopheles, but each torma is specifically hinged posteriorly (e) to a detached triangular plate (b) of the cranial wall, and anteriorly (d) to the end of a transverse epipharyngeal bar (f). Since both this bar and the connective plate (a) underlap the torma, the anterior part of the latter appears to be sunk into the lower wall of the labrum. Posteriorly a strong apodeme (AP) arises from the dorsal surface of the torma and curves mesally. On this apodeme are attached two large muscles (mcls) from the frontal region of the head. Con- traction of the muscles evidently rocks the torma mesally on its articular points and thus gives a backward and mesal stroke to the connected brush. The reverse movement of the brush, as other writers have noted, results from the elasticity of its basal connections. Ac- cording to Cook (1944a) in specimens of Theobaldia [Culiseta] killed and fixed with the brushes retracted, on cutting the muscles the brushes quickly spring back to the expanded condition. The Aedes tormal mechanism is probably characteristic of the Culicinae. The same structure and musculature is shown to be present in Culex by Thompson (1905) and by Chaudonneret (1951), and in no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 15 species of Theobaldia [Culiseta], Lutzia, and Armigeres by Cook (19442). In the predaceous larvae of To.xorhynchites the brushes are sup- ported on prominently projecting lateral lobes of the labrum (figs. 1D, 4E). The brushes are narrow, stiff, and falciform, and appear to be grasping organs, but as observed by Breland (1949) and by Horsfall (1955) they are not used for obtaining prey. Just mesad of the base of each brush is a small, slender sclerite (fig. 4 E, Tor). Dissection reveals that this sclerite has a connection with the base of the brush (F) and gives attachment to two large muscles, leaving no doubt that it is the torma. The preoral cavity—The undersurface of the labrum is continuous with the so-called epipharyngeal surface below the clypeal region, which extends back to the mouth. In most adult insects the part of the preoral cavity above the base of the hypopharynx becomes a special food pocket, the cibarium, opening directly into the mouth. In the mosquito larva the shortness of both the labium and the hypopharynx leaves the entire preoral cavity open below, but still it serves as a channel for water carrying food particles to the mouth. In the tipulid larva, however, there is a short cibarial pocket (fig. 7A, Cb) above the hypopharyngiolabial lobe just in front of the mouth. In the adult mosquito and other sucking insects the closed cibarium becomes a preoral sucking pump. In the mosquito larva the pharynx assumes the sucking function. The epipharyngeal apparatus—Lying in the epipharyngeal surface between the posterior ends of the tormae is a structure that serves to comb food particles from brushes on the mandibles. Since it is musculated, and hence functions actively instead of passively, this in- strument has been termed by Schremmer (1949) the Epipharynx- apparat. Other writers have called it the “palatal bar,” the “epi- pharynx,” and the “epipharyngeal armature.” It includes a transverse bar and groups of setae or other structures arising in front of the bar. The crossbar is usually bow-shaped or V-shaped with the arms diverging forward to the posterior ends of the labral tormae. The setal accompaniment of the bar is quite different in different species. In Aedes (fig. 8 A) the epipharyngeal apparatus is relatively simple. The bar is slender, gently curved forward, and its ends appear to be connected with the tormal apodemes. Arising in front of the bar are two large brushes of stiff hairs that converge posteriorly beneath the bar. At the sides of the brushes arise a pair of large, tapering, hair-bearing processes directed posteriorly, and at the base of each are 16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 two small clawlike structures. In Culex (B) the bar is strongly de- veloped and angulated, its ends, as in Aedes, appear to be attached to the apodemes of the tormae. In front of the bar are two large oval masses of setae curving inward and posteriorly. From above these setal masses two brushes of long hairs project posteriorly. Medially there arise two pairs of short tapering processes that project beneath the bar, and from each angle of the bar a slender, bladelike, sharp-pointed process extends posteriorly. The epipharyngeal apparatus of Anopheles as described by Schrem- mer (1949) is again quite different from that of either Aedes or Culex. The bar is V-shaped with an acute angle. Several brushes arise in front of the bar, but particularly developed are two long, wide combs of flattened, sharp-pointed bristles that extend posteriorly from a pair of triangular sclerites in front of the bar. These are the Klingenborsten of Schremmer, who says they are used for cleaning the food particles from the combs of the mandibles. In Anopheles maculipennis, as shown by Schremmer and by Farnsworth (1947), a large muscle from the clypeal region of the head is attached on each end of the epipharyngeal bar. These muscles the writer has not been able to find in Aedes and Culex, but the close connection of the bar with the apodemes of the tormae possibly coordinates the movements of the epipharyngeal apparatus with the movements of the labral brushes. In all three genera a pair of very slender, closely adjacent muscles is attached medially on the bar. Contraction of the lateral muscles of Anopheles, according to Schremmer, protracts the ap- paratus from the epipharyngeal wall, the median muscles are retrac- tors. Thompson (1905) makes no mention of lateral muscles attached on the epipharyngeal bar in Culex, but he notes the presence of the median retractors. The mandibles—Both the mandibles and the maxillae lie on the underside of the head, where they are implanted obliquely in the membranous area that turns upward from the hypostomal margins of the postgenae to the hypopharyngeal bars (fig. 7B, Md, Mx), the mandibles being above the maxillae. The typical culicine and anopheline mandibles are flattened lobes (fig. 5 D,E,F) with their mesal ends produced into strongly sclero- tized toothed processes and a lower seta-bearing lobe. The dorsal margins bear large comblike fringes of long setae directed mesally. The tips of the mandibles on opposite sides do not meet when the mandibles are closed, but come against the hypopharynx, which lies between them (fig. 7B, Hphy). Each mandible has a posterior basal no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 17 articular point (fig. 5 E,F, a@) that articulates with a process of the hypostomal margin just laterad of the base of the maxilla (fig. 7 B, a). Its anterior articulation (fig. 5 D, c) is with the hypopharyngeal bar (fig. 7B, hb). The mandibles move in the transverse plane by strong abductor and adductor muscles. The principal function of mandibles of this type is the collection by their setal combs of food particles from Fic. 5.—Larval mandibles. A, Tipula abdominalis, right, ventral. B, Lutzia sp., right, ventral. C, To-xro- rhynchites rutilus, left, dorsal. D, Anopheles quadrimaculatus, left, dorsal. E, Culex, sp., right, ventral. F, Aedes aegypti, right, ventral. a, posterior (ventral) articulation; c, anterior (dorsal) articulation. the labral brushes, but the incisor points are said to break up larger particles that collect on the hypopharynx. The mandibles of predaceous larvae, such as Culex vorax (fig. 5 B) and Toxorhynchites (C), are strongly toothed jaws, the points of which come together in adduction (fig. 4 E). Most larval Nematocera have jawlike mandibles (fig. 5 A), though they present many varieties of structure. C. vorax is a culicine mosquito, and its mandible (B) might be derived from the culicine type, but the mandible of To..o- rhynchites (C) is a typical biting insect jaw. The maxillae—The maxillae of the mosquito larva (fig. 6 B-F) are so greatly simplified that they have lost the appearance and structure of an ordinary insect maxilla. They are borne on the transverse hypostomal margins of the postgenae at the sides of the hypostomium, 18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 where they lie below the mandibles (fig. 3 C-F, Mx). The principal part of each maxilla is a flat lobe (fig. 6 D, St) of different shape in different species, bearing brushes of long setae or combs of shorter ones. Laterad of this lobe is a second cylindrical or fusiform lobe regarded as the palpus (P/p) varying in size relative to that of the mesal lobe. At the base of the palpus is usually a small sclerite (+) in the articular membrane. A ry = be Fic. 6.—Larval maxillae, right, ventral. A, Tipula abdominalis. B, Culex vorax. C, Culex sp. D, Aedes aegypti. E, Anopheles quadrimaculatus. F, Toxorhynchites rutilus. Plp, palpus; St, stipes; x, sclerite at base of palpus. In other nematocerous larvae, as in Tipula (fig. 6 A), the maxillary palpus (PIp) is a small lateral appendage of the main maxillary lobe, as it is also in the culicid Culex vorax (B). In most mosquito larvae, however, the palpus appears to have somehow become separated from the rest of the maxilla (C-F). The main maxillary lobe in some nematocerous larvae, as shown by Anthon (1943a), may bear on its distal margin mesad of the palpus two variously developed out- growths, which are identified as the galea and lacinia. The main maxillarly lobe, therefore, appears to be the stipes (St). The nature of the small sclerite (1) at the base of the palpus is uncertain. Cook no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 19 (1944a) calls it the “palpiger,” but it might be referred to the cardo, though no muscles are attached on it. The principal functional features of the culicid larval maxillae are their setal brushes and combs, which serve to collect food particles from the labral brushes. In the predaceous To.xorhynchites the maxillae (fig. 6 F) are similar to those of other species, but they are greatly reduced in size (fig. 4E, Mx). The palpi are presumably sensory organs, but their disparity in size, as between Culex (fig. 6 C) and Anopheles (E), for example, is difficult to explain. The princi- pal movements of the maxillae are in the transverse plane. The labium and hypopharynx.—In most adult insects the salivary duct opens between the bases of the hypopharynx and the labium. In some larval insects, as in caterpillars and hymenopterous larvae, the labium and hypopharynx are united in a single suboral lobe traversed by the duct of the salivary, or silk, glands, which opens at the tip of the composite lobe. The same is true of some nematocerous fly larvae, as is well seen in the tipulid (fig. 7 A, SIDct). In the mosquito larvae the combined labium and hypopharynx are reduced to a flat or somewhat protruding vertical surface between the mouth and the hypostomium, with the salivary duct opening on it. The salivary orifice, therefore, separates the dorsal hypopharyngeal component from the ventral labial component. The hypopharynx (fig. 7 B, Hphy) is supported by the hypopharyn- geal bars (hb) from the lateral cranial walls; immediately above it is the wide mouth (Mth) opening into the pharynx. The labial area below the hypopharynx (D,E, Lb) is variously developed, usually strongly sclerotized and armed with spines or teeth. Other writers have well illustrated the details of the labial structure in different mosquito species. Some have attempted to analyze the larval labium into the parts of a typical insect prementum, but their results are not fully convincing. At C of figure 7 is shown the labiohypopharyn- ‘geal complex of Toxorhynchites rutilus in dorsal view, in which the salivary duct (S/Dct) is seen opening between the two component parts. Attached laterally on the base of the labium are the tendons of a pair of muscles from the ventral head wall, as in the tipulid larva (A). Inasmuch as all the cranial muscles of the insect labium are inserted on the prementum, the labium of the mosquito larva is evidently the prementum; the hypostomium and the ventral head wall, as already shown, being no part of the labium. Menees (1958a), however, has argued that the ventral head area behind the hypostomial lobe must be the labial submentum because the labial muscles have their origins on 20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 it. He thus assumes that these muscles are the retractors of the pre- mentum. The premental retractors, when present, do arise on the submentum, but they are always median in position. The muscles of the mosquito larval labium are lateral muscles, and therefore should pl i y We nencls- ifs iL FrGne LZ LZ i mi Z/f AG ee ra it fA iit Ati i Fic, 7.—Labiohypopharyngeal complex of larvae, and associated structures. A, Tipula sp., section of larval head. B, Culex sp., posterior part of head, anterior. C, Toxorhynchites rutilus, labiohypopharyngeal complex, dorsal. D, Aedes aegypti, same, anterior. E, Culex sp., same, anterior a, posterior articulation of mandible; Ant, antenna; c, anterior articulation of mandible; Cb, cibarium; Clp, clypeus; dlcb, dilator muscles of cibarium; Fr, irons; FrCon, brain connective of frontal ganglion; FrGug, frontal ganglion; hb, hypopharyngeal bar; Hphy, hypopharynx; Hstm, hypostomium; Lb, labium; lbmcl, muscle of labium; lbrmcl, labral muscle; Lm, labrum; Md, mandible; Mth, mouth; Mx, maxilla; phmcls, pharyngeal muscles; RNv, recurrent nerve; S/Dct, salivary duct; Stom, stomodaeum. be one pair of the usual two pairs of cranial muscles of the prementum, which in other insects commonly arise on the tentorium. The same muscles in the tipulid larva (fig. 7 A, /bmcl) certainly have their origins on the head wall, since there is no sclerotization between the postgenae (fig. 3 B). The labial muscles of the mosquito larva, there- no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 2] fore, do not identify the head area on which they arise as any part of the labium. The interpretation of these parts has been still further confused by Shalaby (1957d), who regards the median ventral head area as the labial submentum and mentum, the toothed hypostomial lobe as the paraglossa, the fringed lobe below it the glossa, and the entire complex above the toothed lobe the hypopharynx. Comparative studies, as already shown, give no basis for any such interpretation. Moreover, the adult labium is formed entirely from the rudiment beneath the cuticle of the larval labium (figs. 9 F, 15 A, pLb) and involves no part of the ventral head wall of the larva. The larval labiohypopharynx is evidently retractile, but it plays no active part in feeding. Its principal function is said to be that of an “anvil” on which the incisor points of the mandibles strike to break up food particles. Elaborate studies of the developmental changes in the mouth parts of larval instars of Anopheles, Aedes, Culex, and Psorophora have been made by Shalaby (1956, 1957a, 1957b, 1957c). The pharynx.—The pharynx of larvae that feed on water-borne particles is a small, flattened, ovate or heart-shaped, thin-walled sac (fig. 8 C, Phy) opening directly from the wide mouth (fig. 7 B, Mth) and tilted upward and posteriorly in the head. From its posterior ventral surface is continued the thick-walled oesophagus (fig. 8C, Oe). The ventral wall has an outer layer of semicircular muscles (E, cmcl) the dorsal wall is crossed by four wide muscle bands (C, tmcl) ; the extrinsic musculature includes dorsal and ventral dilator muscles from the head wall. The lateral margins of the pharynx are strengthened by two narrow, concentric, riblike thickenings on each side, convergent to the narrowed posterior end. Internally each of these ribs bears a long brush of fine hairs (D), suggestive of the brushes in the mouth of a baleen whale, and in fact they serve the same purpose, namely, that of filtering the food matter from the in- gested water. A pharyngeal filter apparatus very similar to that of the mosquito larva is shown by Anthon (1943a) to be present in the larvae of several other nematocerous families. The pharynx of the predaceous culicid larva of Toxorhynchites, however, is a simple funnnel-shaped enlargement of the anterior end of the oesophagus, and has no filter brushes. In any case, the larval pharynx is not to be identified with the sucking pharynx of the adult mosquito, which lies in the posterior part of the head (fig. 24 A, PhP), and the larva has no cibarial pump. 22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 Larval feeding.—tThe process of feeding by nonpredaceous larvae is not a mere matter of having food particles washed into the mouth by streams of water from the vibrating labral brushes. It involves cooperative action on the part of the labrum, the epipharyngeal apparatus, the mandibles, the maxillae, the labiohypopharynx, and the pharynx. The whole feeding process has been minutely described by Schremmer (1949) for the Anopheles larva, in which it is more readily Fic. 8—The larval epipharyngeal apparatus and the pharynx. A, Aedes aegypti, epipharyngeal apparatus. B, Culex sp., same. C, Anopheles maculipennis, pharynx and its muscles, dorsal (from Schremmer, 1949). D, Same, filter-bearing rib of pharynx. E, Same, cross section of pharynx (from Imms, 1907). cmcel, circular muscle; fb, filter brush; Mth, mouth; Oe, oesophagus; Phy, pharynx; tmcl, transverse muscles. observed than in other species because the head is held with its under- side turned upward against the surface of the water. Briefly, Schrem- mer’s account is as follows. The movements of the lateral brushes of the labrum create currents in the water that converge to the front of the head and are directed medially by the middle brush. With the backward stroke of the lateral brushes the mandibles and the maxillae are closed upon them, and as the brushes again go forward particles that may be adhering to No. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 23 them are scraped off by the combs of the mandibles. Accompanying the opening of the mandibles, the epipharyngeal apparatus is pro- truded by action of its muscles and its bristles remove the food particles from the mandible combs. These freed particles and others that may be adhering to the epipharyngeal surface are then collected by the long basal brushes of the mandibles (fig. 5 D) and, with the closure of the mandibles, are pushed into the mouth of the pharynx. Though the mandibles and the maxillae close at the same time, the maxillae open first and the mandibles following remove whatever particles may be adhering to the maxillae, which lodge on the hypopharynx and with the next stroke of the maxillae are thrust into the pharynx. Large particles collected on the hypopharynx are broken up by the toothed lobes of the mandibles, which strike on the hypopharynx like hammers on an anvil. The pharynx, by muscular expansion of its walls, functions as a sucking organ for drawing in a stream of water accompanying the mechanically ingested food particles. A contraction then follows in which the dorsal wall is deeply infolded by the action of the dorsal transverse muscles (fig. 8 E), reducing the pharyngeal lumen to two lateral channels containing the filter brushes (fb). At the same time the water is driven toward the mouth and the food particles are filtered out by the brushes. The water is then discharged through the open angles of the mouth, goes above the mandibles and escapes past the sides of the head. Schremmer made further experiments on a Culex larva by impregnating the water in a dish with carmine particles. After feeding by the larva, the carmine was found massed in the brushes along the sides of the pharynx. When the pharyngeal brushes have worked as filters for some time and have become well loaded, the pharynx makes a strong contraction which suddenly removes the carmine particles from the brushes and lodges them in small clumps at the mouth of the oesophagus, into which they are finally taken. The mosquito larva swallows no appreciable amount of water, its water balance being maintained by the anal lobes. The extreme specialization of the mouth parts and the pharynx in the filter-feeding mosquito larvae gives a striking example of how independent of the adult structure an insect larva may become in its adaptation to a new way of feeding. In various mosquito genera, however, the larvae of some or all species are predaceous on other small aquatic animals, particularly on other mosquito larvae. Notable in this group are members of the subgenus Lutgia among the Culicini, and of the genus Toxorhynchites. In these forms the mandibles are 24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 strongly developed jaws (fig. 5 B,C), the toothed lobes of which come together or overlap for grasping and biting. Yet these larvae have labral brushes and some of the other special features of particle-feed- ing larvae, so it is difficult to say whether they represent a partway stage in the evolution of filter feeding, or have been secondarily adapted for feeding on whole live prey. In some species the larvae are particle feeders in the first instar and become predaceous in their later instars. It would appear, therefore, as said by Bates (1949), “that the predacious habit has developed independently in the larvae of a number of mosquito groups, involving distinct adaptations both of structure and behavior.” THE THORAX The larval thorax has a simple oval form, in which the interseg- mental lines are but faintly marked as grooves of the cuticle, and there is no external trace of appendages. In the fourth instar the thorax becomes conspicuously enlarged (fig.9g A). Beneath the cuticle on the ventral side are now plainly visible the extroverted wings and legs of the future pupa, and on the dorsal side the pupal respiratory trumpets. On removal of the cuticle (C) the legs are seen to be long, fully segmented appendages (E) closely folded in loops against the sides of the thorax. The forewings (W-) are large pads corrugated in their basal parts (D) to allow expansion; the smaller hindwings (Ws) are more slender and tapering free folds of the metanotum. It has been shown by Imms (1908) that the rudiments of the wings, legs, and respiratory trumpets are formed in a young larval instar of Anopheles as integumental folds in pockets of the epidermis (B). Ap- parently they are extruded beneath the cuticle at the beginning of the fourth instar. This early eversion of the wings and legs occurs also in other nematocerous larvae, such as Dixa, Corethra, and Chirono- mus, shown by Miall and Hammond (1900) in Chironomus. On each anterior lateral angle of the thoracic dorsum of Anopheles larvae there is usually to be seen a pair of minute, tapering, trans- parent lobes arising from a common base (fig. 9 A, no). These struc- tures are known as the “notched organs.” They are retractile and hence are not visible on all specimens, or only their tips may project. Between the lobes of each pair is a funnel-shaped depression that ends in a strand, which is said by Chang and Richart (1951) to be at- tached to the neighboring dorsal tracheal trunk. These writers con- tend, therefore, that the organs are the “prepupal respiratory trum- pets.”” However, when the cuticle of a fourth-instar larva is removed, no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 25 the lobes and the funnel come off with it, showing that the organs are larval structures. Furthermore, the trumpets of the “prepupa’’ (i.e., the pharate pupa) are present beneath the larval cuticle. They appear to arise from the pupa just beneath the larval organs, but they project forward or mesally until the pupal ecdysis, when they stand out from the thorax. Fic. 9.—The larva, and developing pupal appendages. A, Anopheles punctipennis, fourth instar larva, dorsal. B, Anopheles maculi- pennis, section of wing and leg buds in early larval instar (from Imms, 1908). C, Aedes aegypti, thorax of mature larva with cuticle removed, exposing extro- verted legs and wings. D, Same, pupal wings of larva. E, Same, third left pupal leg of larva. F, Anopheles maculipennis, pupal labium developing inside larval labium (from Imms, 1908). al, anal lobes; e, dorsal brush of larva; L, leg bud; pLb, pupal labium; s/O, salivary orifice; Sp, spiracle; W, wing bud; We, Ws, pupal wings of larva. The nature of the “notched organs” of the Anopheles larva is not clear. Their position on the dorsum of the thorax suggests that they might be remnants of anterior spiracles such as are present on larva of many other flies, including some Nematocera. Since these spiracles of successive instars are not formed in the usual manner within the 26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 preceding spiracle, but as independent branches from a persisting spiracular atrium, it is perhaps possible that the pupal trumpets are in this manner related to the “notched organs” of the larva. Chang and Richart contend that the latter serve to keep the anterior part of the Anopheles larva afloat while feeding at the surface, but experi- ments have shown that the organs can be cut off without any apparent effect on the suspension of the larva (Jones, unpublished obser- vations). THE ABDOMEN The larval abdomen (fig. 10 G) appears to have only nine segments, and it is usually represented as nine-segmented, with the respiratory apparatus on the eighth segment and the terminal segment enumerated as the ninth. However, there is reason for believing that a true ninth segment is combined with the eighth. Christophers (1922) con- tended that though “much of the apparent eighth segment is actually this structure, the greater part of the spiracular apparatus must be assigned to the tergite of a hitherto unrecognized ninth abdominal segment.” Convincing evidence of this interpretation is the fact that the rudiments of the male genitalia are formed beneath the larval cuticle at the base of the terminal segment, and that in the adult male the genital claspers are carried on the posterior margin of a small but distinct ninth segment (fig. 27 B). Though this segment is not evident as a distinct annulus in the larva, it must be represented by some part of the apparent eighth segment immediately anterior to the genital rudiments. In the pupa, as will be shown (fig. 16 D,E) a small ninth-segment ring (JX) lies behind the eighth segment and carries the tail fins and the small anal lobe. The anal segment of the larva (fig. 10 B) must therefore be the tenth, as it is in the pupa and the adult. The fully segmented abdomen of the mosquito embryo is shown by Telford (1957) in Aedes and by Menees (1958a) in Anopheles to have Io segments. Telford says the tenth segment, or telson, dis- appears with the ingrowth of the proctodaeum, but since a tenth seg- ment is present in the adult, the “telson” must be an eleventh segment. In some larvae, as seen in Mansonia (fig. 11 A) a small lobe (XJ) protrudes from the end of the tenth segment, which would appear to be the evaginated anus-bearing telson. Even in the embryo, then, the ninth segment is not differentiated from the eighth. It appears as a distinct ring first in the pupa and as a definite segment in the adult. The first seven segments of the larval abdomen have no distinctive features, except that in Anophelini (fig. 9 A) the last five or six of no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 27 them bear on the back pairs of small palmate brushes (¢) that suspend the larva from the surface of the water in its usual horizontal feeding position. The respiratory apparatus on the dorsum of the ninth seg- mental region contains a pair of large open spiracles, which are either Fic. 10—Larval respiratory organs. A, Anopheles maculipennis, tracheal system, dorsal (from Imms, 1907). B, Anopheles quadrimaculatus, end of abdomen, spiracular apparatus open. C, Same, spiracular apparatus closed. D, Same, details of spiracular apparatus. E, Culex pipiens, end of abdomen. F, Aedes aegypti, same. G, Culex quinquefasciatus, larva in feeding position. H, Culex pipiens, end of respiratory siphon. al, anal lobes; dTra, dorsal tracheal trunk; Sp, spiracle; VIJI-X, abdominal segments. flush with the surface (fig. 10 A, Sp) or carried out on the end of a respiratory tube (E,F,G). The tenth segment contains the func- tional anus at its end, and bears four lanceolate, thin-walled apical appendages, or anal lobes (A,B,E, al). Flat dorsal and ventral brushes 28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 of long, spreading hairs are usually present on the end of the tenth segment, and perhaps serve as a rudder during swimming. Though mosquito larvae are commonly known as ‘“‘wrigglers’” or “wigglers” they swim by lashing movements of the abdomen, which drive them forward, backward, or sideways. The active larvae of Culex zigzag through the water like tumbling acrobats. Anopheles, however, is a true wriggler ; it swims either on the surface or under the water by quick lateral movements of the abdomen and propels itself backward. The dorsal spiracles of the abdomen are the only breathing aper- tures of the mosquito larva. The lateral spiracles are closed except at the ecdyses, when they are temporarily opened to allow the tracheal linings to be pulled out. Since the dorsal spiracles open into the dorsal trunks of the tracheal system and the lateral spiracles into the lateral trunks (fig. 10 A), the dorsal spiracles cannot be supposed to be a pair of lateral spiracles that have moved up onto the back. It may be conceded that spiracles can change their position, but they cannot change their tracheal connections. The spiracles of anopheline larvae lie in the floor of a shallow, basinlike peritremal structure elevated on the back, the margins of which are variously produced into lobes (fig. 10 B). In Anopheles maculipennis (D) there are two large, thin posterior lobes, a pair of small tapering lateral lobes, and a single anterior lobe supported on a transverse basal bar. The spiracles (Sp) lie anteriorly; behind them is a median V-shaped sclerotization on the floor of the basin, and on each posterior lobe is a weak submarginal sclerotization. As the Anopheles larva feeds stretched out against the surface film of the water the peritremal basin projects just above the water with the spiracles freely exposed to the air. When the larva submerges, the whole apparatus folds up and the lobes clamp tight together (C). Imms (1908) describes three sets of paired muscles that effect the closing of the lobes, which retain a bubble of air between them. When the muscles relax the lobes open. Curving around the end of the ninth segment beneath the ends of the posterior lobes is a narrow semi- circular bar that supports on each side a small plate bearing a comb of strong recurved bristles (D), or in some species is armed with spines or teeth. In the larvae of Culicinae and Toxorhynchitinae the spiracles are carried out on the end of a tube, or siphon, varying in length and thickness in different genera (fig. Io E,F). The spiracles are at the end of the tube and are surrounded by lobes similar to those in Anopheles, but necessarily much smaller (H). When the larva is at no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 29 the surface it hangs from the end of the siphon with the spiracles ex- posed to the air (G). Two strands of slender muscle fibers traverse the tube and converge to attachments on a strong apodeme from the terminal apparatus. An extensive comparative study of the peritremal structure has been made by Montschadsky (1930) from a taxonomic standpoint. His illustrations are not realistic since they appear to have been drawn from flattened specimens, and the sclerotic parts are overemphasized by an unnaturally dark tone, but they show the great specific variation in the pattern of the peritremal lobes. Glands associated with the spiracular apertures have been described by Keilin, Tate, and Vincent (1935). The secretion is oily and serves to give a hydrofuge quality to the peritremal surface, which prevents wetting and the entrance of water into the spiracles. Though the respiratory siphon is primarily constructed for breath- ing air at the surface of the water, in species of Mansonia and a species of Ficalbia it is modified for insertion into the roots of aquatic plants. The siphon tapers distally and the apex is armed with spines, teeth, and hooks, which, operated by the inner muscles of the tube, enable the larva to insert the tip of the organ into the plant. In Mansonia indubitans (fig. 11 A) the siphon is large, conical in shape, and narrowed at the distal end. The apex is not sharp, but is armed with a pair of strongly toothed movable lobes (B), which can be re- tracted and brought together, or protracted with the teeth turned out- ward. The siphon in this case is a cutting and not a piercing instru- ment. It contains only one tracheal trunk, formed by the union of the dorsal body trunks in the eighth abdominal segment, and there is a single median, ventral spiracle between the bases of the toothed lobes. These larvae live entirely submerged and obtain their air from the air channels of the plant, to which they remain attached. According to Iyengar (1935a, 1935b) species of Mansonia in India attach themselves only to the water plant Pistia stratiotes. To insert the siphon the larva moves backward with the siphon held horizon- tally and thrusts the tip against the root. It then wriggles actively backward, while it operates the apical armature with muscles attached on a rodlike apodeme, until the end of the siphon penetrates the root deep enough to enter an air chamber, when apical hooks anchor the larva to the root. The adult female lays her eggs only on submerged leaves of the Pistia plant, thrusting her abdomen into the water to do so, and where Pistia is not present she will lay no eggs. While most other mosquito larvae spend most of their time at the 30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 surface of the water, any of them can stay below without apparent discomfort, and some do so indefinitely. It was formerly supposed that the four thin-walled tracheated lobes borne on the end of the tenth abdominal segment were gills serving for underwater respira- tion. Wigglesworth (1933), however, has produced evidence that Fic, 11.—Respiratory tubes of larvae and pupae that get their air from the roots of aquatic plants. A, Mansonia indubitans, terminal segments of larva. B, Same, apical part of siphon, ventral. C, Mansonia sp., thoracic respiratory horns of pupa, dorsal. D, Same, right horn, mesal. E, Mansonia richiardii, pupal respiratory horns, dorsal (from Wesenberg-Lund, 1920-21). F, Ficalbia hybrida, terminal part of pupal respiratory horn (from Bonne-Wepster, 1932). An, anus; Tra, trachea. these lobes are water-absorbing organs rather than gills. By immers- ing larvae in a water culture of the flagellate protozoon Polytoma, which is highly sensitive to the amount of oxygen in the water, he found that the flagellates first assemble at the posterior end of the larva and then spread all over the body surface. Soon, however, they move away in a mass, indicating that oxygen is being consumed by no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS ie the general integument of the larva as well as by the anal lobes. The submerged mosquito larva, therefore, breathes through its skin, and some other aquatic larvae are known to do the same. From experimental ligaturing of the body of the larva in different places, Wigglesworth furthermore showed that the larva absorbs water from the posterior end of the body, presumably through the thin, permeable anal lobes. During feeding, the larva does not swallow the water taken into the pharynx with its food, this water, as already noted, being discharged from the mouth. The anal lobes thus serve to maintain the physiological balance of water in the larval body. INTERNAL ANATOMY Inasmuch as the principal specializations of the mosquito larva have to do with feeding and breathing, there is little in the internal or- ganization that is essentially different from that of other insects. The tracheal system.—The tracheal system of most insects in- cludes a pair of lateral tracheal trunks running lengthwise through the body, with which the lateral spiracles are connected. Many insects, however, have also a pair of dorsal longitudinal trunks. In dipterous larvae, including the mosquito larva, that breathe through dorsal spiracles, the dorsal trunks are particularly large (fig. 10 A, dTra), and the lateral trunks connected with the closed lateral spiracles are mutch reduced. The dorsal spiracles of the ninth abdominal segment are evidently secondary respiratory apertures to allow the larva to breathe at the surface of the water, since it is hardly to be sup- posed that a primitive lateral spiracle could migrate dorsally and change its tracheal connections. In general the last pair of lateral spiracles is on the eighth segment. In the larvae of higher Diptera there is also a pair of secondary anterior dorsal spiracles on the thorax. The fine end branches of the insect tracheal system in general go to the cells of the body tissues, which are thus directly oxygenated. In the larva of Anopheles, Imms (1907) describes a series of small tubes from the longitudinal trunks in the eighth abdominal segment that break up into fine branches going to the posterior end of the heart. Imms suggested that these branches may oxygenate the blood in the heart, but Jones (1954) says they end on the heart wall. At each larval ecdysis the cuticular intima of the tracheal tubes is shed with the outer cuticle. In the mosquito larva, according to Wig- glesworth (1949), the intima of the main tracheal trunks breaks be- tween the segments, and the pieces attached to the shed cuticle are drawn out through the lateral and the posterior dorsal spiracles of 32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 the new instar. The lateral spiracles are then closed again, since they are not functional in the larva for respiration. In the same manner, at the ecdysis of the pupa the tracheal trunks in Culex are said by Hurst (1890) to break up into segmental pieces, which are pulled out through the temporarily opened spiracles. The soft inner tissue of the respiratory siphon is withdrawn into the body where it is finally absorbed. The siphon itself is shed with the larval cuticle, and its two tracheal trunks break off at the base. The tracheal system of the young larva on hatching is filled with a liquid. According to Frankenberg’s (1937) observation on Culex, air enters the tracheae only when the end of the respiratory siphon comes above the water surface. One of the dorsal longitudinal trunks fills first, and then the other. The air is drawn into the tracheae as the embryonic liquid diffuses through the tracheal walls. The dorsal blood vessel——vThe dorsal blood vessel of the mosquito, particularly in Anopheles quadrimaculatus, has been elaborately de- scribed by Jones (1954). Structurally it differs in no essential respect from the vessel of other insects, except for a dilatation, or sinus, of the aorta in the thorax. The larval organ is a simple muscular tube extending along the midline of the back from the eighth abdominal segment into the head. The part in the abdomen, known specifically as the heart, is perforated along the sides by eight pairs of segmental openings, or ostia. The part in the thorax, called the aorta, is im- perforate. In the head the aorta goes beneath the brain, where it is open ventrally allowing the blood to be freely discharged into the head cavity, whence it flows backward through the body to reenter the heart through the ostia. The larval heart, Jones says, always beats forward at an average of 85.2 pulsations a minute, but it has no nerve connec- tions. Along the sides of the heart are attached the usual fan-shaped segmental groups of muscle fibers, the so-called alary muscles, that support the heart on the body wall. The alimentary canal.—In the mosquito larva the alimentary canal (fig. 12) is a relatively simple tube. It consists of the usual three parts of the arthropod digestive tract, an ectodermal stomodaeum, an endodermal mesenteron, and an ectodermal proctodaeum. The sto- modaeum begins in the head with the pharynx (Phy), which is fol- lowed by a narrow oesophagus (Oe) that goes through the neck into the thorax, where it enters the first part of the mesenteron, known as the cardia (Car). (This term, borrowed from vertebrate anatomy, has no literal significance in the insect.) Within the cardia the oesopha- geal walls are reflected to form the usual entrance funnel of the no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 33 stomodaeum into the mesenteron. The cardia is followed by a long, straight tube, the stomach, or ventriculus (Vent), that extends back into the seventh abdominal segment. The anterior end of the ventricu- lus bears a circle of eight large pouchlike diverticula, the gastric caeca (GCa). The dark mass of food particles in the ventriculus is contained in a thin tubular peritrophic membrane (PMb), shown by Wigglesworth (1930) to be secreted by the cell walls of the cardia surrounding the stomodaeal funnel. The proctodaeum, or intestine, is differentiated into a short anterior part (Alnt), and a longer posterior part, or rectum (Rect). The anterior intestine begins as an expansion against the end of the ventriculus, and then narrows to a tube that makes an S-shaped bend to the saclike anterior enlargement Fic. 12—Lengthwise section of a Culex larva, showing the alimentary canal. Alnt, anterior intestine; An, anus; Car, cardia (anterior part of ventriculus) ; GCa, gastric caeca; Hstm, hypostomium; Mal, Malpighian tubules; Mth, mouth; Oe, oesophagus; Phy, larval pharynx; PMb, peritrophic membrane; Lect, rectum; S/G/d, salivary glands; Vent, ventriculus (stomach). of the rectum, which finally proceeds as a narrow tube to the anus (An). ae For a detailed study of the general structure, histology, and move- ments of the larval alimentary canal of Anopheles the reader is referred to a forthcoming paper by Jones (in press). The Malpighian tubules—The excretory Malpighian tubules of the larva (fig. 12, Mal) are five in number. They arise from the an- terior end of the proctodaeum, first going forward into the sixth abdominal segment, and then turning posteriorly to end in the sub- terminal segment around the rectal sac. The salivary glands——The larva has a pair of small salivary glands of various shapes lying ventrally in the thorax (fig. 12, SIG/d). The ducts unite in a common outlet duct that enters the head and opens on the labiohypopharyngeal surface just below the mouth (fig. 15 A, SIO). The glands usually consist each of two parts of different shape 34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 separated by a constriction. The histology of the glands in Anopheles larvae has been described by Jensen and Jones (1957). In Anopheles albimanus the globular anterior part of each gland consists of 12 to 15 large cells; the pear-shaped posterior part contains 50 to 60 much smaller cells. The glands of opposite sides are connected by a strand of nephrocytes. In other genera the relative size and shape of the two gland parts differ in various ways. The nervous system.—The central nervous system of the larva in- cludes a brain and suboesophageal ganglion in the head, and a ventral chain of segmental ganglia in the abdomen united by paired connec- tives. The last ganglion is that of the eighth abdominal segment. The reproductive organs——Rudiments of the reproductive organs are present in the young larva in a very elementary state; they slowly develop during the larval life. Food reserves.—The insect larva has no idea of the meaning of its life or of what is to become of it. Its hereditary factors automatically determine its destiny by converting it into a pupa and finally into an adult. Yet, physiologically, the larva is loaded with responsibilities. Not only must it maintain its own existence, but at the same time it must provide for the future nutritional needs of the pupa and for its transformation to the adult. In the mosquito pupa there is a minimal breakdown of larval tissues to furnish food for the developing adult organs. The active mosquito pupa, moreover, is not a “resting stage,” and, since it cannot eat, it is dependent upon the larva for everything except the air it breathes. An important function of the larva, there- fore, is the storage of food reserves in its body to maintain the pupa and to insure the development of the adult. Only when the winged adult finally emerges from the pupal skin can the mosquito again take food and become once more an independent, self-sustaining insect. The elaboration and storage of food reserves in the body of the fourth-instar mosquito larva is the subject of a special study by Wigglesworth (1942). The stored materials include principally pro- tein, fat, and glycogen, which are shown by experiments to be rapidly consumed when the larva is subjected to starvation, and replenished on subsequent feeding. Normally, it is to be supposed, the stored prod- ucts are passed on intact to the pupa, but Wigglesworth does not go into this phase of the subject, or follow the utilization of the reserves by the pupa. The matter, however, is well-enough known in other insects. Peewee The active pupa is familiar to all students of mosquitoes after its ecdysis from the larva. The fact, however, that it is already fully no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 35 formed shows that it became a pupa while still within the larval cuticle. It will therefore be of interest to follow the transformation processes that convert the larva into a pupa. THE PUPAL DEVELOPMENT As before noted, the primary buds of the pupal wings, legs, and respiratory trumpets are formed at an early larval period in pockets of the epidermis beneath the cuticle, as are also those of the antennae and the labium, and rudiments of the compound eyes are present in the first instar. Fic. 13.—Pupae in natural floating position against the surface of the water. A, Aedes atropalpus. B, Anopheles punctipennis. The development of the compound eyes of the mosquito has been described by Zavrel (1907), by Constantineanu (1930), and by Sato (1951a, 1953a, 1953b). The eye rudiments are first evident in the first larval instar as thickenings of the epidermis just in front of the larval eyes. With development of the eye pigment, the compound eyes become visible externally in the second instar or the early part of the third instar. From then on they increase in size as the ommatidia are slowly differentiated in the epidermis. During the larval stage the ommatidia are covered by the unmodified cuticle, but in the pupa the cuticle over each ommatidium becomes convex and the corneal facets are thus defined. After emergence of the adult the lenses become bi- convex, and the ommatidia are completed in from 3 to 12 hours, but the lenses may continue to thicken during the first 24 hours of adult life. The early development of the wings and legs in the mosquito larva is nothing unusual. The leg buds are always formed in the embryo, and all immature insects have legs, whether external or internal. Like- 36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 wise the young of all winged insects have wing rudiments developing either externally or internally. The unusual thing about the mosquito and related Diptera is that the legs, wings, and pupal respiratory trumpets are fully extruded beneath the cuticle of the thorax at the third larval moult instead of at the moult to the pupa (fig.9 C). The wings are still in the form of pads (W2, Ws), but the legs (E) are already fully segmented appendages. At a somewhat later period of the fourth instar, the larval cuticle is separated from the abdomen except at the posterior end (fig. 14), and beneath the cuticle on the back of the first segment are now seen the two small suspensory brushes of hairs characteristic of the pupal abdomen. The thorax and the abdomen inside the moulted larval Fic. 14.—Fourth-instar larva of Aedes aegypti with larval cuticle (/Ct) moulted over the thorax and most of the abdomen. The inner cuticle (pCt) is that of the pupa. cuticle, therefore, pertain to the future pupa. The head cuticle of the larva has not yet been moulted, so that the larva in the fourth instar still feeds with its own mouth parts. It breathes with its posterior respiratory apparatus, and uses for locomotion the muscles now in the pupal abdomen. The condition found in the mosquito is simpler than that described by Hinton (1958b) in Simulium. Here the fully formed pupa still within the larval skin is active for several days before ecdysis. Its activity is due to the fact that many of the former larval muscles, in- cluding those of the head, remain attached by tonofibrillae to the moulted cuticle of the larva. The pupa of Simulium is thus able to use the larval organs, and it not only continues to feed, but spins its own cocoon. In the mosquito there is no evidence of muscles retain- ing their attachment on the moulted larval cuticle; the insect feeds with the larval mouth parts until the latter are cast off at the final pupal moult. The larval musculature of the thorax and abdomen is said by Thompson (1905) in Anopheles to go over into the pupa and the no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 7 adult with little alteration. The rudiments of the future wing muscles, however, are present in the thorax during the last larval instar. Hulst (1906), on the other hand, reports that in the larva of Culex there takes place an extensive histolysis and histogenesis of the body mus- culature, beginning when the larva is two-thirds grown. Some imaginal muscles thus appear first in the larva, particularly those of the wings and legs, prior to the advent of the pupal stage. Destruction of the larval muscles of the abdomen, however, Hulst says, is not complete even in a late stage of the pupa. In the Simuliidae, according to Hinton (1959), “the indirect flight muscles and the tergal depressor of the trochanter develop quite independently of the larval muscles in all post-embryonic stages.” Histological changes in the alimentary canal beginning in the larva have been described by Samtleben (1929), by Berger (1938) for Culex, and by Richins (1945) for Aedes. The replacement of func- tional cells from regenerative cells in the ventriculus during larval life is generally in other insects not a metamorphic process but the usual procedure of replacing worn-out digestive cells by new cells. At the fourth ecdysis to the pupa, however, Berger (1938) says, rapid changes take place. The alimentary canal of the pupa, well illustrated by Hurst (1890), differs from that of the larva, but is still not that of the adult. The short pupal stomach is said by Richins to be formed from only the posterior part of the larval stomach. According to Samtleben no specific pupal epithelium is formed for the pupal stomach. Considering the precocious development of so many of the imaginal organs, the fourth instar of the mosquito larva presents the anomalous condition of being part larval and part pupal. In other words, the pupal development begins within the larva long before its completion at the pupal ecdysis. It ends with the formation of the pupal head, mouth parts, and tail fins. In most young insects the endocrinologists find that the larval struc- ture is maintained by the inhibitory action of the corpus allatum hor- mone on the adult development until the end of the larval life. The early origin of pupal organs in the mosquito larva and the continu- ance of their development through the larval period shows, however, that the juvenile, or status quo, hormone does not necessarily function as a complete inhibitor of adult development. In the mosquito it ap- pears to be selective in its action, allowing the growth of pupal parts that do not interfere with the normal activities of the larva, while it maintains to the end of the larval period such parts as the head, feed- 38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 ing organs, and respiratory apparatus that are essential to the life of the larva. The corpora allata of the mosquito larva are described by Boden- stein (1945) as a “corpus allatum complex” composed of two small cellular bodies of elongate form, tapering posteriorly, attached laterally on the aorta just behind or within the neck. Anteriorly they adhere closely to a transverse trachea and are connected with each other by a loose chain of cells. Each body is entered by a slender nerve from the brain. Since the bodies contain different kinds of cells it is possible that they include elements of the usually separate corpora cardiaca. In higher Diptera the aorta is surrounded by a cellular ring, which is thought to include the corpora allata and corpora cardiaca, but accord- ing to Bodenstein the nature of the cells in the mosquito larva is not certain. The larval complex goes over into the adult in reduced form as two small, rounded bodies lying on the sides of the aorta. If the fourth-instar larval mosquito behaves as other larvae have been shown to do when experimentally given an extra dose of juvenile hormone, it should go over into a fifth larval instar. In this case the larva issuing from the fourth-instar cuticle would have external legs and wings! We can only wait the results of some endocrinologist who may make the experiment. When at last the cuticle of the larval head is moulted, taking with it the larval antennae and mouth parts, the corresponding pupal organs are rapidly developed within the still-unshed larval cuticle. The recon- struction of the mouth parts involves an extreme change from the specialized organs of the larva to the equally but differently specialized organs of the adult. The development of the pupal mouth parts has been described by Thompson (1905) for Culex, and by Imms (1908) for Anopheles. The pupal labrum begins its growth as a fold of the epidermis at the anterior end of the dorsal wall of the head that first extends pos- teriorly beneath the cuticle (fig. 15 B, pLm). The fold elongates (C, Lm) and finally turns forward and downward over the other mouth parts. The buds of the new mandibles and maxillae are formed directly from the epidermis retracted into the bases of the larval organs. An early stage of their development still within the larval cuticle is seen at C of the figure taken from Thompson. The labium and the hypopharynx of the larva, as already shown, are greatly re- duced and united in an area between the mouth and the hypostomium, the two components being separated only by the opening of the salivary duct. In Aedes the labiohypopharyngeal complex as shown by no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 39 Salem (1931) forms a distinct lobe below the mouth (fig. 15 A, Hphy, Lb), as it does also in a tipulid larva (fig. 7 A). The rudiment of the pupal labium within the larval labium (fig. 15 A, pLb) is said by Imms (1908) to be a pair of hollow lobes confluent at their bases. There is no separate rudiment of the adult hypopharynx. It is shown by Thompson (1905) that the hypopharynx is still united with the pupal labium (fig. 15 B) when the larval cuticle (/Ct) is moulted. Later, as will be described, the hypopharynx of the adult female is separated from the labium. In their final stage of development the pupal mouth parts have become greatly lengthened and are closely pressed together in a long curved proboscis (D). Rudiments of the pupal tail fins are formed beneath the cuticle of the fourth larval instar behind the respiratory apparatus, and the primary buds of the male external genital organs appear beneath the cuticle of the same instar behind the sternal region of the ninth abdominal segment. THE MATURE PUPA The pupa at ecdysis (fig. 16 A) is fully formed in all its outer parts and thereafter does not change externally. It is clearly a preliminary adult with the appendages in a halfway state of completion. The pupa can hardly represent a former active stage in the life of the mosquito, since its mouth parts are unfitted for any kind of feeding. The pupal thorax has already assumed the approximate size and shape of the adult thorax. In Simuliidae, Hinton (1959) says, the definitive thoracic structure is developed during the pharate stage of the pupa. General external structure.—The head and thorax of the mosquito pupa are combined in a large cephalothorax, from which projects the slender abdomen (fig. 16 A). When at rest the pupa floats at the sur- face of the water (fig. 13), but it does not hang from its respiratory trumpets (as it often does in pictures). The back of the thorax and of the two anterior abdominal segments comes against the water, while the rest of the abdomen hangs downward as ballast. The open ends of the respiratory trumpets project just above the surface of the water, and two small brushes of spreading hairs on the back of the first abdominal segment help keep the pupa suspended. The floating position of the pupa is necessary for the future emergence of the adult, and is maintained by bubbles of air enmeshed in the folds of the legs and beneath the wings. The source of the air that maintains the buoyancy of the pupa, according to Hurst (1890), appears to be a pair of large open spiracles 40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 Mada Mx MxPlp Fic. 15.—Development of pupal mouth parts and an adult leg. A, Vertical median section through labiohypopharyngeal lobe of larva of Aedes, with contained rudiment of pupal labium (from Salem, 1931). B, Section of anterior part of head of Culex larva, with pupal labrum and labium forming inside the unshed larval cuticle (from Thompson, 1905). C, Head of Culex pupa removed from larval cuticle, with pupal mouth parts in early stage of development (from Thompson, 1905). D, Pupal head of Aedes aegypti, lateral. E, Same, anterior. F, Fully developed pupal mouth parts of Aedes aegypti. G, Distal part of a pupal leg with adult leg formed within it. Ant, antenna; Aul, aulaeum; C/p, clypeus; Hphy, hypopharynx; Li, first leg; Lb, labium; /Ct, larval cuticle; Lm, labrum; Md, mandible; Mx, maxilla; MxPlp, maxillary palpus; pLb, pupal labium; pLm, pupal labrum; P/p, palpus; ae salivary duct; S/O, salivary orifice ; SoeGng, suboesophageal ganglion; , tibia, no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 4I on the sides of the first abdominal segment of the pupa covered by the metathoracic wing pads. The tracheal system of the pupa, how- ever, is so weakly developed that it would hardly seem capable of supplying the amount of air carried by the living pupa. Manzelli Fic. 16.—The pupa. A, Aedes aegypti, male pupa, lateral. B, Same, terminal part of female abdomen. C, Culex sp., thorax and base of abdomen, dorsal. D, Same, end of female abdomen, dorsal. E, Same, apical structures of male abdomen. F, Same, phallic organ of male pupa, ventral. br, suspensory brush of first abdominal tergum; H, head; Ns, metanotum; PhL, phallus; Prb, proboscis; Rect, rectum; S, sternum; 7, tergum; #f, tail fin; Tmp, respiratory trumpet; W+2, mesothoracic wing; Ws, metathoracic wing; I-X, abdominal segments. (1941) described and figured the pupa as “enclosed in a sac-like struc- ture,” which he says “has long been seen by all mosquito workers and is usually known to them as the pupal shell.” This is a curious state- ment, since no such structure exists. Furthermore, the “shell” is said 42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 to enclose a large air cavity, but on pressing a pupa in alcohol the air issues as free bubbles from beneath the legs and wings. The pupa has two features that are peculiarly its own. First are the trumpet-shaped respiratory tubes projecting from the back of the thorax (fig. 16C, Tmp), and second, a pair of thin, oval fan- shaped tail fins, or paddles, borne on the end of the abdomen (A, tf). Because it is necessary for the pupa to float with the back of its thorax against the surface of the water, with the abdomen hanging down, it had to discard the posterior spiracles of the larva and have its breathing apertures forward. The trumpets are connected with the anterior ends of the dorsal longitudinal tracheal trunks, and their open ends project just above the surface of the water. It is a curious fact that in species of Mansonia and Ficalbia, the larva of which gets its respiratory air from the roots of aquatic plants, the pupa does the same thing by means of its thoracic trumpets. The trumpets in these species are drawn out into a pair of long horns directed forward from the thorax. In Ficalbia hybrida each horn ends in a pair of tapering blades (fig. 11 F), but in species of Man- sonia each terminates with a strong, curved spine. The spines of Mansonia richiardiu (E) are convergent and are said to be applied close against each other as inserted into the plant. In the species shown at C of the figure the spines are divergent, and, as in other species, each is bordered anteriorly and posteriorly (D) by a very thin, trans- parent, faintly striated flange. A trachea (C,E, Tra) is attached to the base of the organ, but does not penetrate the latter. The cylindrical basal stalk contains a wide lumen, which narrows abruptly where it enters the spine and opens by a minute aperture at the tip. Wesenberg- Lund (1920-21), however, says of M. richiardu that “the trachea runs through the whole tube,” and Grossbeck (1908) figures a tube of Culex perturbans with a trachea going through it to the tip of the spine. It seems very unlikely, however, that the thoracic respiratory tubes in any case contain tracheae. They are merely elongated trum- pets, and a typical trumpet is an open funnel with the trachea opening into its base (fig. 17 C, Tra). As the pupa of Mansonia emerges from the larval skin, according to Galliard (1934) as quoted by Marshall (1938), it brings the tips of its horns together and searches for a neighboring root. Then it violently works its way out of the anchored larval skin and at the same time inserts its horns. When the adult is ready to emerge, the pupa breaks away from the plant and comes to the surface where it floats by reason of two tracheal air sacs in the thorax. The winged mosquito thus escapes into the air in the usual manner. No. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 43 It is truly remarkable that the same kind of structural adaptation for the same purpose has occurred twice in the life of the same indi- vidual, affecting two different organs. Furthermore, with the acquisi- tion of a new structure designed for a new use, the insect must be twice endowed with a new instinct for using the modified organs. It is enough to make us wonder if we really understand the nature of biological adaptation. Though pupae that breathe free air ordinarily float at the surface of the water, they can escape danger by darting around on the surface or submerging quite as actively as the larvae by snapping movements of the flexible and well-musculated abdomen. The large tail fins are organs for increasing the motor efficiency of the abdomen. Func- tionally they are comparable to the tail fan of a crayfish. The pupa when swimming progressively on or below the surface kicks back- ward with its abdomen and propels itself forward, but the crayfish does just the opposite. When the pupa swims downward in the water, however, it goes tail first, and thus maintains its floating position. If it remains inactive it passively rises to the surface, otherwise it swims up by abdominal movements. The head and mouth parts ——The head of the pupa (fig. 16 A, H) is closely attached to the lower anterior angle of the thorax, with its true dorsal surface directed anteriorly. It retains nothing of the structure of the larval head. The long, many-jointed antennae curve upward and backward beneath the lower edges of the wings. The large, black compound eyes (fig. 15 D, E) are conspicuous beneath the cuticle, and between them the clypeal region (E, Clp) makes a prominent bulge on the face. Posteriorly the head is produced into a long, tapering proboscis that lies beneath the thorax with its end upcurved behind the lower legs (fig. 16 A, Prb). The component elements of the proboscis are closely adherent (fig. 15 E), but are easily separated (F). Along the lower side is the relatively thick labrum (E, Lm) which is continuous from the clypeus. Flanking the labrum are the very delicate slender mandibles (Md), and bordering the mandibles are the maxillae (Mx). The wide base of each maxilla bears a free, tapering palpus (F. Plp). On the posterior (upper) side of the proboscis is the soft, slender, tubular labium ending in a bifid tip (F. Lb). There is no free hypopharynx in the young pupa. As we have seen, the hypopharynx is not separated from the labium in the larva, and the two parts go over still united into the pupa, with the salivary duct enclosed between them. In most adult insects the hypopharynx is an independent suboral lobe, and the salivary duct 44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 opens behind its base in front of the labium (fig. 23, SIO). The fe- male of the mosquito and other adult Diptera possesses a free hypo- pharyngeal stylet, but it is traversed by the salivary duct. According to Thompson (1905) the hypopharynx of the female mosquito is differentiated by cellular growth from the median line of the anterior (lower) surface of the labium during the pupal stage. Since the hypopharynx, when it becomes a free stylet, contains the salivary duct, it would seem that in its separation from the labium it must take a part of the labium with it. In the male the hypopharynx is not separated from the labium, and the salivary channel remains in the labium. Dimmock (1881) says that in the male of Culex “the hypo- pharynx is, throughout its whole length, joined to the labium,” and Hurst (1890) observes that it is “inseparable from the labium.” The fact that the hypopharynx of Diptera contains the salivary duct has given rise to the idea that this stylet is a new formation not homologous with the hypopharynx of other insects (see Demerec, 1950, pp. 375, 376). Yet the stylet in Diptera has all the usual rela- tions of the hypopharynx to surrounding parts, and its base forms the floor of the preoral cibarial pump (fig. 24 E), just as in the cockroach (fig. 23) and other generalized insects. The cuticle of the pupal mouth parts represents the organs as they are developed in the pupa. Inside the cuticular sheaths a renewed growth of the epidermis produces the final adult form of the stylets, just as the adult legs are formed within the cuticle of the pupal legs (figs. 15 G, 17 A). The segmented maxillary palpus of the adult, for example, is clearly seen inside the simple palpal sheath of the pupa (fig. 15 F, Plp), and within the end of the pupal labium (Lb) are visible the labellar lobes of the adult. The thorax.—The large thorax of the pupa is indistinctly seg- mented, but it bears the legs and wings, and carries on its back the respiratory trumpets (fig. 16 A). The legs and the wings of the pupa have been taken over directly from the larva. The legs have increased in length and their joints are more distinct (fig. 17 A), but they are closely folded in loops against the sides of the thorax as in the larva. The mesothoracic wings are much larger and more winglike in shape; the hind wings are still triangular lobes of the metanotum. Within the cuticle of the pupal appendages are plainly seen the developing appendages of the adult. The venation of the forewing is already laid out (D). Within the hindwing may be seen the club-shaped halter (E, Hit), which, whatever may be its evolutionary history, is not formed in ontogeny by a gradual modification of the wing. no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 45 The abdomen.—The abdomen of the pupa (fig. 16 A) resembles that of the larva except for the lack of the respiratory apparatus, the presence of the tail fins (tf), and the reduction of the tenth segment (X) to a small anus-bearing lobe. The dorsum of the first segment has a special pattern of sclerotization (figs. 16 C, 17 F) and bears the two brushes of spreading hairs that keep the base of the abdomen sus- pended at the surface of the water. It is suggested by Hurst (1890) that these brushes, besides serving as suspensoria, probably also are Fic. 17.—Pupal characters and an adult leg of Aedes aegypti. A, Left third leg of pupa with adult leg formed inside the cuticle. B, Same leg of adult on emergence, same magnification. C, Right respiratory trumpet, mesal. D, Left mesothoracic wing with adult wing inside the cuticle. E, Left metathoracic wing with halter forming inside it. F, Metanotum and first two ab- dominal segments. G, End of abdomen with tail fins. Hilt, halter; Ns, metanotum; Tra, trachea; Ws, metathoracic wing; I, IJ, first and second abdominal segments. sensory organs responding to vibrations in the water. The pupa be- comes immediately active on any disturbance of the water, even to a tap on the containing vessel. The pupal tail fins, as usually drawn in illustration, appear to be attached to the end of the eighth abdominal segment (fig. 17 G). If they are pulled away from the eighth segment, however, they are seen to be carried by a transverse dorsal bar entirely separated from the tergum of the eighth segment (fig. 16 D,E, XT), which, in fact, is the tergum of the ninth segment. On it is supported also the small tenth segment (X). In the male pupa (E) the ninth segment is a 46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 complete narrow annulus (JXT, XS) as in the adult male (fig. 27 B), and below the small tenth segment projects a pair of large lobes (fig. 16 E, PhL) ona common base arising from the sternal arc of the ninth segment (F). These lobes are the genital appendages of the male as far as they are developed in the pupa. Male and female pupae, there- fore, can be distinguished by the presence (A) or absence (B) of the genital lobes (PHL), though in the male the lobes might be mistaken for the tenth segment, since the latter is mostly concealed above them (Ay Xx). THE PUPAL METAMORPHOSIS The pupal life of most mosquitoes is very short, two or three days or less, though with some species it is much longer. During this time the contour of the adult is modeled by new growth of the epidermis beneath the pupal cuticle, while the mouth parts, wings, halteres, and legs take on the adult structure within their pupal sheaths. At the same time reconstruction of internal organs takes place inside the body. The degree of reconstruction necessary to change the larval organs into those of the adult, however, is much less in the mosquito than in many other insects, especially in the higher Diptera. The mosquito pupa breaks with the tradition that a pupa is a “rest- ing stage” in the life of the insect. When an ordinary pupa is broken open it is seen to be full of a creamy mass of soft material resulting from the disintegration of the larval tissues. The inside of a mosquito pupa is as clean as that of the larva or the adult, and its organs appear to be intact. Whatever reorganization is going on takes place mostly inside the alimentary canal and the refuse is not thrown into the body cavity. The abdominal muscles are so well preserved that the pupa is an extremely active stage of the mosquito, and the thoracic muscles are so well developed that the pupa might be expected to fly if its wings were more mature. As already noted, Hulst (1906) has described the process of muscle histolysis and histogenesis as beginning in the larva, but he is not explicit as to what larval muscles are destroyed or when the imaginal musculature is completed. In Culex, according to Hurst (1890), the muscles of the pupa are those of the imago; the principal muscles are present in the young pupa, but they increase greatly in size. A casual examination of the abdominal musculature in the larva, pupa, and adult shows little difference between the stages, except for the greater size of imaginal muscles. However, we need a more de- tailed comparative study of the muscle pattern and more information on the replacement of individual muscles. no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 47 The larval head musculature appears to be largely replaced by an imaginal musculature. According to Thompson (1905) there is an extensive histolysis of the larval head muscles, accompanied by a regeneration of muscles appropriate to the adult, which takes place in the eighth to tenth hour of pupal life. The pupal tracheal system is weakly developed and is difficult to see in dissections. According to Hurst (1890) tracheae go from the base of each thoracic trumpet to various parts of the head and body, and a transverse trunk connects the two trumpets. A pair of longi- tudinal trunks runs back to the rear end of the body, giving off branches to the internal organs and to the site of each spiracle. Only the spiracles of the first abdominal segment remain open. In his study of the heart of Anopheles quadrimaculatus, Jones (1954) reports that no evidence was found that the heart is “de- stroyed, reconstructed, or otherwise drastically modified during meta- morphosis.” In young pupae, according to Jones, the heart beats in a forward direction as in the larva, but later it may cease beating for prolonged periods of time. Circulation of the blood, therefore, ap- pears to be unessential for the regenerative changes taking place in the pupa. The alimentary canal of a young pupa, as described and illustrated by Hurst (1890) in Culex, might be supposed to be a functional organ if the pupa could feed. It more resembles the digestive tract of the larva than that of the adult, but since the adult feeds on a very dif- ferent kind of food from that of the larva, the alimentary canal under- goes a complete reconstruction in the pupa, details of which have been described by Hurst (1890), Thompson (1905), Samtleben (1929), and Richins (1938). The oesophagus is least affected insofar as its epithelium goes over intact from larva to adult, but the larval pharynx is lost, and an enlargement in the back of the head forms the post- cerebral sucking pump of the adult. In the thorax the dorsal and ventral diverticula of the adult grow out from the oesophageal wall. The larval gastric caeca are absorbed and not replaced in the adult. The larval epithelium of the stomach, according to Richins, de- generates completely and is cast off into the stomach lumen, as a new epithelium is formed by permanent regenerative cells. Transforma- tion in the proctodaeum is brought about partly by histolysis and histogenesis of the epithelium and partly by regrowth. The five Malpighian tubules of the larva go over into the adult without change. In the rectal sac of the pupa are formed six invaginations of the wall that become the rectal papillae of the adult. The salivary glands of 48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 the larva degenerate and each is replaced by three slender tubules generated from cells in the neck of the larval gland. The central nervous system undergoes little change in the pupa other than growth and union of some of the ganglia. The first ab- dominal ganglion of the larva is drawn into the thorax, where it fuses with the metathoracic ganglion, and later the four ganglia now in the thorax condense into a single mass. The last abdominal ganglion of the larva unites with the ganglion of the seventh segment. In the adult mosquito, therefore, there are only six separate ganglia in the abdomen (fig. 30 C). In the head, as described by Woolley (1943) for Aedes, the brain and the optic lobes grow rapidly by peripheral formation of new cells. The circumoesophageal connectives shorten and the suboesophageal ganglion unites with the brain around the oesophagus. Though the visible changes that take place in the nervous system are slight, there must be a considerable reorganization of the internal structure. The behavior and instincts of the adult mosquito are en- tirely different from those of the larva. Since the activities of the in- sect resulting from sensory stimuli are determined by established neuromuscular pathways and synapses in the central nervous system, the system that serves the larva must be entirely reorganized into one appropriate for the activities of the adult. Of this, however, we know little or nothing in any insect. Ill. THE ADULT The adult mosquito fully formed within the pupa has now only to cast off its pupal mold to gain its freedom in the garb of a mature winged insect. But this is not easily done since the confined mosquito has no instruments for cutting or breaking the pupal cuticle. More- over, the wings, legs, antennae, and mouth parts are enclosed in tight- fitting sheaths, from which they must be slowly extracted. However, much as we might wish that the mosquito should remain a prisoner in the pupal skin, nature has made provision for its liberation. As noted by several observers, the first evidence that the adult is about to emerge is the appearance of a film of air beneath the pupal cuticle on the back of the thorax. A slight retraction of the adult ap- parently breaks the connections of the pupal trumpets with the tracheal system and thus allows air to escape beneath the cuticle. Usually a short piece of trachea remains attached to the base of each trumpet. According to Marshall and Staley (1932) rhythmical movements now begin in the sucking pump of the adult which draw the air forward, no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 49 forming a bubble at the base of the proboscis. This air is then pumped into the stomach as a long narrow bubble that extends back to the fourth abdominal segment. Pressure by the distended abdomen now pushes the thorax forward until it ruptures the pupal cuticle in a median slit along the back from the neck to the end of the meso- thorax. Outside air then enters the cleft and is rapidly swallowed, going back in the stomach as far as the sixth abdominal segment and greatly distending the abdomen. Knab (1909), in describing the role of air in the ecdysis of insects, says of the mosquito that on emergence from the pupa it is distended with air far beyond its natural size, the integument being stretched to its utmost. According to the writer’s observations on emerging mosquitoes the degree of distention is highly variable, even with individuals of the same species. Pupae of Aedes aegypti, before the adult ecdysis, are observed to have the abdomen extended straight back from the thorax, and during the emergence it is held, or floats, in this position with the tail fins against the water surface (fig. 18 A). When the pupal cuticle splits on the back of the thorax, the thorax of the adult bulges out and pushes apart the lips of the cleft. This produces a transverse split over the back of the pupal head, so that the pupal skin can now be widely opened anteriorly (B) to allow the egress of the adult. At the same time the cuticle on top of the pupal head between the eyes breaks out and folds forward as a free flap beyond the antennal bases (A,B). Behind the antennae the anterior tentorial arms project internally as a pair of slender tapering rods (B). Inasmuch as the legs of the adult, as well as the wings and mouth parts, are enclosed in tight-fitting pupal sheaths, the mosquito cannot use its appendages for freeing itself. Yet, when the head and thorax are free, the abdomen follows and the entire adult slowly rises verti- cally from the pupal skin as if pushed out from below. The legs and wings are at first closely pressed against the body, but as the legs are freed they at once become active, and appear to be reaching for the surface of the water. The mosquito seems to know instinctively that now and henceforth it must support itself on its legs. It will be noted that the legs of the emerged adult are greatly longer than their pupal sheaths ; the hindleg of an Aedes (fig. 17 B), for example, may lengthen to two and a half times the length of the corresponding folded leg of the pupa (A). When the end of the abdomen and the wings are finally out of the pupal thorax and the legs are all free, the new insect confidently steps out onto the surface of the water and calmly walks away from the discarded pupal skin. It may come to rest on 50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 some nearby floating object (as a bit of cardboard in the aquarium), but usually in a very short time it is able to fly, and immediately is gone. Sometimes, however, mosquitoes in culture appear to have much difficulty in finally extracting their legs; often they fall over on the surface of the water, and some perish in this position with their tarsi still held in the pupal sheaths. It is probable that in such cases the larvae were not properly nourished. ic. 18—An adult female of Aedes aegypti emerging from the submerged pupal cuticle; and the open thorax of the discarded pupal cuticle of Anopheles quadrimaculatus. A remarkable thing about the mosquito is that, after its whole previous life spent in the water, on emergence from the pupa it is at once at home in the air. Without a flutter of the wings or any practice trial, it makes a perfect takeoff, flight, and distant landing. During the pupal stage, therefore, the mosquito has not only been equipped with a complete mechanical apparatus of flight, but in its nervous system a mechanism of control has been fully elaborated. Compare this with the difficulty the young human has in learning even to walk, but of course his ancestors did not always walk upright on two legs. no. 8 ANATOMICAL LIFE OF THE MOSQUITO—-SNODGRASS 51 The newly emerged mosquito (fig. 19) is really an elegant insect as it stands high on its long slender legs, the abdomen held straight back beneath the neatly folded wings, and the long proboscis extended from the head. The sexes of most species are readily distinguished at once by the antennae, those of the female having usually circles of short hairs, those of the male being large spreading plumes. Fic. 19.—Aedes aegypti, adult male. THE HEAD The head of an adult mosquito has little likeness to that of the larva. It is an oval capsule (fig. 20D) with the facial region carrying the antennae directed forward, and the long proboscis extended from its lower end. The sides are largely occupied by the great compound eyes, which almost meet dorsally and ventrally (A,B, E). The large bases of the antennae arise so close together on the face that the frons is reduced to a narrow verticle bar between them (A, Fr), but its lower end forks into diverging arms that support the clypeus (Clip). A median coronal sulcus (cs) on the vertex extends down- ward on the face through the frons. The strongly convex clypeus (A,C, Clp) forms a prominent lobe just above the base of the proboscis. The undersurface of the head (B) resembles that of the 52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 larva in that it is completely closed from the occipital foramen to the base of the proboscis. The head is attached to the thorax by a slender membranous neck and is supported by a pair of lateral cervical sclerites (D,E). The head of the male is similar to that of the female, but is a little smaller. The internal head skeleton consists of a pair of simple tentorial arms extending from anterior pits above the lateral angles of the clypeus (A, at) to posterior pits (B, pt) on the ventral margin of the occipital foramen. From the front of the face arise the long antennae (fig. 20 D,E). The hairy flagellum of each organ is borne on a large globose base (A, Pdc), which is the pedicel, or so-called torus, but when the pedicel is removed (right) it is seen to be itself supported on a narrow ring (Scp) that represents the usually much longer scape of other insects. The slender shaft of the flagellum is divided into 14 sections (errone- ously called “segments”), 13 of which carry each a whorl of hairs. In general the sexes are readily distinguished by the number and length of the flagellar hairs, which in the male (fig. 22 A) give the antennae a plumose appearance in contrast to the short-haired female antennae (D,E). The two types, however, intergrade, females of some species having bushy antennae, and some males short-haired antennae. In the female the hairs arise from clear areas near the bases of the flagellar units (B) ; in the male (C) they are borne on promi- nent, darkly sclerotized, subapical expansions of the units. Tulloch and Shapiro (1951) have shown from electron microscope studies that the flagellar hairs are armed with rows of minute teeth; in Culex quinquefasciatus they estimate there are at least 16 rows along each hair. These writers, however, are in error where they say the hairs “arise at the junctions of the flagellar segments.” The large globose pedicel of the antenna in each sex contains a highly developed sclopophorous sense organ, present also, though usually much smaller, in the antennal pedicel of most insects. The organ was first described in Culex as an auditory organ by Johnston (1855), who did not at all understand the nature of the structure in the pedicel, but it has since been known as Johnston's organ. Sub- sequently Child (1894) made good histological studies of the organ in various insects, including the mosquito, and his illustrations are now famliar in most entomological texts. A more recent comparative study of the organ in Culex, Aedes, and Anopheles is given by Risler (1955). The component sensory elements in the pedicel are attached to a plate or prongs on the base of the flagellum, and thus evidently register movements of the flagellum. no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 53 7 A i A4YTT Ay y we SMM \| Lm re Hphy Md Mx 1 Fic, 20.—Head and mouth parts of an adult female mosquito, Aedes aegypti except G. A, Head and base of proboscis, anterior, B, Same, posterior. C, Clypeus and base of proboscis, lateral. D, Head, lateral. E, Same, with mouth parts separated. F, End of labium, dorsal. G, Cross section of proboscis of Anopheles (from Vogel, 1921). H, Distal ends of mouth part stylets. I, Basal parts of maxilla. J, Proximal parts of right maxilla and labium, posterior. at, anterior tentorial pit; Cd, cardo; Clp, clypeus; cs, median cranial sulcus; E, compound eye; fc, food canal; For, occipital foramen; Fr, frons; Hphy, hypopharynx; Hst, hypostome; Lb, labium; LObI/, labellum; Le, lacinia; LG, labial gutter; Lig, ligula; Lm, labrum; mel, muscle; Md,Mds, mandible, mandibles; Mx,M-xae, maxilla, maxillae; MaPlp, maxillary palpus; Nv, nerve; Pdc, antennal pedicel; P/p, palpus; Prb, proboscis; pt, posterior tentorail pit; sc, salivary canal; Scp, antennal scape; St, stipes; Thc, theca; Tnt, anterior tentorial arm; Tra, trachea; Vx, vertex. 54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 While it is probable that the organ of Johnston in the antenna of most insects registers the movements of the flagellum, the elaborate experimental work of Roth (1948) leaves no doubt that the highly developed organ in the male mosquito is responsive to the effect of sound waves on the flagellum. This, of course, does not imply that the mosquito has an auditory “sense” ; mechanical reaction to stimuli is all that we can attribute to the insects. Male mosquitoes are at- tracted to the females in flight by the tone produced by their wings. Roth showed that males with intact antennae, when subjected to the sound of a tuning fork at 480 vibrations a second held behind a suspended piece of cloth, fly to the source of the sound where they | exhibit typical mating activities though no females are present. Even after complete removal of the flagellar hairs, males still respond to more intense sounds apparently by vibrations of the shaft alone, but on complete removal of the flagella they give no reaction. Roth’s tests were made particularly on Aedes, but males of other genera were found to react similarly. Females of Aedes aegypti gave no evidence of being attracted to sounds, “though they may give shock-reaction to certain intensities.” Further experimental work of Roth (1951) on females of Aedes seems to show that the antennae function as directional distance thermoreceptors and probably also as chemoreceptors. Females de- prived of their antennae are unable to locate a host from a distance. The antennae and the palpi are said to be the chief organs responding to stimuli that induce probing by the proboscis. The receptor organs of the antennae, however, are not described, but along the shaft of the female antennae (fig. 22 B) are numerous hairs, and on the male antenna (C) a ring of very short hairs encircles the distal end of each flagellar section. The antennae of insects in general are known to be the principal seat of chemoreception. The compound eyes of the mosquito are so large that they almost encircle the head. Sato (1950, 1953a, 1953b) reports that by actual count there are from 440 to 462 facets in the eye of a male Culex pipiens, and 503 to 566 in the female; and that in Aedes japonicus the male eye contains 440 to 462 facets, the female eye 504 to 527. The surface area of the eye in each genus is larger in the female than in the male. The internal structure of the compound eye in Culex is described by Constantineanu (1930) and by Sato (1950). An extensive experimental study of the visual responses of flying mosquitoes made by Kennedy (1939) on unfed females of Aedes aegypti shows that the mosquitoes react negatively to light, and are No. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 55 attracted to dark objects. Experimentally they orient toward black stripes on a white background, and continue to do so when the stripes are rotated about them. When confronted by two black stripes, they face one or the other and not the intervening space. In a wind tunnel freely flying mosquitoes move against the current. THE ORGANS OF FEEDING The feeding organs of the adult mosquito include the proboscis and two sucking pumps, One of the latter is a preoral cibarial pump beneath the clypeus, the other is a pharyngeal pump, being a part of the alimentary canal behind the brain in the back of the head. In describing the feeding organs of the adult it will be better to take the female first, because in most mosquitoes she is the biting and bloodsucking member of the species and has the mouth parts fully developed. In the nectar-feeding male some of the parts are much reduced or absent. The proboscis—The slender, rodlike proboscis in the female mosquito is usually composed of all the mouth parts possessed by in- sects that feed on solid food, namely, a labrum, a pair of mandibles, a hypopharynx, a pair of maxillae, and a labium, but the parts are all structurally modified in adaptation to the mosquito’s way of feeding. The relation of the parts in the undisturbed proboscis is best seen in a cross section (fig. 20G). In the deeply channeled upper side of the labium (LD) are enclosed the labrum (Lm), the mandibles (Md), the hypopharynx (Hphy), and the maxillae (Mx). The labrum itself is practically an inverted tube, since its margins are curved downward and may overlap. The enclosed labral canal (fc) is the food conduit. The hypopharynx contains the salivary canal (sc). By careful manipulation with a dissecting needle all these parts can be separated as shown at E. The labrum (fig. 20 H, Lm) is the thickest and the strongest of the stylets. It is movable by muscles from the clypeus attached on its base (fig. 24D), but the muscles simply elevate and depress the labrum, which is firmly hinged on the clypeus. The term “labrum- epipharynx” often applied to the labrum is quite unnecessary, since in its general form the labrum is a flat lobe of the head and therefore has an upper and lower surface. In the mosquito the decurvature of the lateral parts converts the labrum into a tube through which the ingested liquid food is drawn up by the sucking apparatus at its base. At the sharp-pointed distal end (fig. 20 H, Lm) the walls of the channel diverge to make an opening like that of a hypodermic needle. 56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 The mandibles are the slenderest of the stylets, but they vary some- what in thickness and shape in different species. In Aedes here illustrated (fig. 20 H, Md) each is slightly enlarged toward the taper- ing distal end. The base of each mandible is movably connected with the lower part of the cranial wall by a small suspensory sclerite, and a slender muscle from the tentorium is inserted on the mandibular base. The mandibles are thus retractile for a short distance, and, when retracted, their withdrawn tips give free entrance to liquid into the open end of the labral food canal. Protraction results from the elas- ticity of the suspensory mechanism on relaxation of the muscles. The single, median hypopharynx, present as an independent stylet only in the female, is a simple, flattened rod (fig. 20H, Hphy) traversed by the salivary outlet canal (sc), which opens on its acute tip. The hypopharynx is not individually movable; its anterior wall is continued basally into the floor of the cibarial pump. The maxillae are less reduced than the other mouth parts, and are well equipped with muscles. The principal part of each maxilla (fig. 201) is a long, flattened, sharp-pointed blade (Lc) armed with re- curved teeth near the end of its outer margin (H, Mx). From the base of the blade projects a usually short four-segmented palpus (1, Pip). The maxillary blade has been regarded as the galea by some writers (Robinson, 1939; Snodgrass, 1944), but it is more reasonably interpreted by Schiemenz (1957) as the lacinia, which is usually the operative part of a generalized maxilla. From its base a long, strongly sclerotized, apodemelike rod extends backward in the head and gives attachment to muscles (J). This rod is evidently the stipes, or more probably stipes (St) and cardo (Cd) combined, sunk into the head, since in some related flies, such as Phlebotomus (fig. 22 G), it is superficial on the back of the head and articulates on the cranial margin. The maxillary musculature of Aedes (fig. 20]J) includes a long retractor arising on the head wall close to the posterior end of the tentorial arm (Tnt) inserted on the distal end of the stipes, and two protractors attached proximally on the stipito-cardinal rod. One of these muscles arises on the tentorium, the other, very curiously, on the base of the labium. A lateral muscle from the tentorium and a short mesal muscle both attached on the base of the lacinia are regarded by Schiemenz (1957) in Theobaldia [Culiseta] as an abductor and adductor respectively of the maxilla. A short muscle from the stipes is inserted on the base of the palpus, and each palpal segment contains a small muscle inserted on the segment distal to it. no. 8 ANATOMICAL LIFE OF THE MOSQUITO—-SNODGRASS 57 The long, gutterlike labium of the mosquito is the so-called pre- mentum of a generalized labium, the usual basal part of the labium being absent, though a small postmental sclerite may be present in other Nematocera (fig. 22 G, Pmt). The prementum in Diptera is known as the theca because it ensheaths the other mouth parts. Apically it bears two small movable lobes, the Jabella (fig. 20 F, Lol), and ends between them in a slender median projection, or ligula (Lig). The labella appear to be two-segmented, and evidently represent the labial palpi because each is provided with an abductor and an adductor muscle from the prementum. The only muscles attached on the base of the labium are the two already noted that arise on the maxillary stipites (J) and probably act as protractors of the maxilla, since the labium is firmly fixed to the head. The styliform mouth parts within the labial theca adhere to one another in a compact fascicle. They are usually said to be held to- gether by an oil liquid, but Bhatia and Wattal (1957) have described rings issuing from the margins of the labrum that surround the hypopharynx, mandibles, and maxillae and bind these stylets to the labrum. However, no other investigator has reported the presence of any such structures, and the writer has failed to see them in Aedes, Culex, or Anopheles. The incurved lower edges of the labrum enclose only the food canal. When the female mosquito is about to take a meal of blood, she places the tip of the proboscis against the skin of the victim (fig. 21 A), closely holding the end of the stylet fascicle between the labial labella. The movable maxillary stylets are the active piercing organs. Acting alternately, first one is protracted and holds its position in the flesh by means of its recurved teeth, then the other is forced in beyond the first and takes a deeper hold. The labrum, mandibles, and hypopharynx penetrate along with the maxillae. The retractor muscles of the maxillae, instead of pulling the stylets out of the wound, where they are held by the maxillary teeth, bring the head down closer to the feeding surface. The labrum, still holding the stylet fascicle between the labella, is thus forced to bend backward (B) and the bend becomes greater the deeper the stylets penetrate (C). When finally the stylets pierce and enter a small blood vessel, or let out a pool of blood, the mandibles are drawn back from the end of the labrum to allow the blood to enter the food canal in response to the suction of the cibarial pump. Saliva discharged from the hypopharynx in some species serves to prevent coagulation of the blood. A more detailed account of the feeding act and of accompanying movements by the maxillary palpi is 58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 given by Robinson (1939). After feeding, the maxillary stylets are retracted, the female braces herself against the skin of the victim with her legs, and forcibly pulls out the fascicle of stylets, which again is ensheathed in the straightened labium. In discussing the feeding of mosquitoes, we must not overlook the fact that not all females are bloodsuckers. A prominent exception to the rule are species of Toxorhynchites, in which both sexes feed on Fic. 21.—Successive stages in the penetration of the stylets of a female mosquito feeding on blood. (B, C, from Gordon and Lumsden, 1939, with neck plate added.) nectar or other plant juices. In this genus (fig. 22 D) the proboscis is very long, tapering, and strongly decurved. The maxillary palpi pro- jecting from the base of the proboscis are long and four-segmented. The laciniae by contrast are weak and taper into filaments reaching only a little beyond the end of the first palpal segment ; evidently they play no part in feeding. A slender labrum extends to the tip of the proboscis, but mandibles appear to be absent. Then there are species of Malaya (=Harpagomyia) that get their no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 59 food from ants. In these the proboscis is curved forward at its lower end (fig. 22 E) ; the distal part is thickened and armed with long hairs. The elongate labella terminate with a pair of small transparent lobes. The species of Malaya are minute mosquitoes, much smaller than ordinary ants. As described by Jacobson (1911) they sit on branches AE \ Wi \y YY l Fic. 22.—Various types of antennae, palpi, proboscides, and labia of adult mosquitoes. A, Culex sp., head of male. B, Same, part of female antenna. C, Same, part of male antenna. D, Toxorhynchites rutilus, head and proboscis of female. E, Malaya jacobsom, head and proboscis of female. F, Culex, distal end of male labium, showing salivary duct. G, Phlebotomus verrucarum (Ceratopogonidae), head and proboscis, posterior. Cd, cardo; For, occipital foramen; LObI, labellum; Pmt, postmentum; S/Dct, salivary duct; St, stipes; Thc, theca (prementum). inhabited by ants, and when an ant runs between the legs of one of them the mosquito thrusts the end of its proboscis between the open mandibles of the ant, which accommodatingly gives up its dinner to the mosquito. The proboscis of the adult Malaya lacks mandibles and maxillae. According to de Meijere (1911) these members are present 60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 in the pupa, but the imaginal parts formed inside of them are short and disappear. The mouth parts of the male mosquito are much simplified by the great reduction of the mandibular and maxillary stylets and the entire absence of a hypopharyngeal stylet. The male proboscis, therefore, consists principally of only the labrum and the labium, but the maxil- lary palpi are usually highly developed and may be much longer than the proboscis (fig. 22 A). Mandibular stylets when present are seldom longer than half the length of the proboscis and are usually much shorter. Marshall and Staley (1935) report that they are present in all genera examined except “Aedes and Ochlerotatus.” These writers found maxillary stylets to be present in representatives of all genera examined, but the length is highly variable, even in species of the same genus. The labium is a deep trough, as in the female, and ends with a tapering median ligular lobe between the labella (fig. 22 F). It will be recalled that the hypopharynx of the male mosquito is not separated from the labium, as in the female. The hypopharynx thus retains in the adult male the larval condition of union with the labium. The male “labium” is, therefore, really a labiohypopharynx. The hypo- pharyngeal component in Anopheles is identified by Vizzi (1953) as a sclerotic plate on the floor of the labial gutter. In sectional figures he shows the salivary canal in an apparent median thickening of the plate. In Culex (fig. 22 F) the salivary duct (S7Dct) is a thread- like tube that traverses internally the floor of the labial gutter and opens on the tip of the ligula, but it appears to be free in the labial lumen, The cibarial pump.—The structure here termed the cibarial pump lies just beneath the clypeus at the base of the proboscis, and is the organ that sucks the liquid food up through the canal of the labrum. The same pump is present in all Diptera and is the sucking apparatus of other liquid-feeding insects, such as the Hemiptera. It has long been erroneously called the “pharynx,” and even some recent writers continue to call it such on the pretext of not wishing to confuse stu- dents. It is possible, however, that some students might prefer to know the facts. The organ in question is entirely outside the mouth, as no true pharynx could be, but admittedly it is difficult to understand its anatomical status in the mosquito. We must therefore turn to some other more generalized insect for light on the nature of the preoral sucking organ, and for this purpose the cockroach will be par- ticularly illuminating. In a vertical lengthwise section of the head of a cockroach (fig. no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 61 23) the mouth (Vth) is seen to lie beneath the upper end of the clypeal region (C/p) of the cranial wall. Below the mouth projects the large tonguelike lobe commonly termed the hypopharynx (Hphy), which has a long base sloping up to the mouth. On this basal part of the hypopharynx is a depression that forms the floor of a pocketlike space (Cb) in front of the mouth beneath the inner wall of the clypeus. The masticated food passed back from the mandibles is stored in this pocket before it is swallowed. The pocket, therefore, is named the Th / Lib S10 Fic. 23.—Vertical section through the left side of the head of a cockroach, exposing the preoral cavity. Cb, cibarium; Clp, clypeus; Fr, frons; hf, hypopharyngeal fulcrum; Hphy, hypopharynx; Lb, labium; Lm, labrum; Mth, mouth; Phy, pharynx; PrC, preoral cavity; S/Dct, salivary duct; S/O, salivary orifice; y, oral suspensory arm of hypopharynx. 5a,5b, dilator muscles of cibarium; 6,7, frontal muscles of stomodaeum; 13, adductor of hypopharynx; 14, abductor of hypopharynx. cibarium (food container). On its dorsal wall are attached strong muscles (5a, 5b) from the clypeus. The hypopharynx can be pressed against the inner clypeal wall by muscles (13) attached to arms (y) from its base. The cibarium then becomes a closed chamber that can be dilated by the clypeal muscles, and probably serves as a sucking organ when the cockroach drinks liquids. In insects that habitually feed on liquid food, the cibarium becomes elaborated to form a permanent sucking pump. 62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 When we turn now to the mosquito, a section of the head (fig. 24 A) will show beneath the bulging clypeus (C/p) a small elongate capsule (COP), which is the primary sucking pump. The basinlike lower wall is strongly sclerotized and, in the female, is directly con- Fic. 24.—The sucking apparatus of an adult mosquito. A, Diagrammatic section of female head. B, Culex sp., section of pharyngeal pump (from Thompson, 1905). C, Aedes aegypti, pharyngeal pump exposed by removal of anterior head wall. D, Same, muscles of labrum. E, Same, cibarial and pharyngeal pumps, left, cibarial pump opened to show lumen and dilator muscles. at, anterior tentorial pit; Br, brain; CbP, cibarial pump; For, occipital foramen; FrG, frontal ganglion; lvr, labral lever; MaPIlp, maxillary palpus; Oe, oesophagus; PhP, pharyngeal pump; sc, salivary canal; S/P, salivary pump; SoeG, suboesophageal ganglion; Tt, tentorial arm; y, oral arm of cibarial pump. Other lettering as on figure 23. tinuous with the supper surface of the hypopharynx (Hphy). The dorsal wall (E) is the so-called epipharyngeal surface from the labrum (Lim) to the mouth (Mth), and is thin and flexible. On it is attached a great mass of dilator muscles (5) from the clypeus. The NO. 8 ANATOMICAL LIFE OF TITE MOSQUITO—SNODGRASS 63 inner end of the organ opens through the mouth (Mth) into the narrow first part of the alimentary canal, and at each side of the mouth projects a small process (y) on which are attached two antagonistic muscles (A, 13, 14), as in the cockroach. All these features so closely duplicate those of the cibarium in the cockroach as to leave no doubt that the preoral sucking pump of the mosquito is the ciba- rium. In the mosquito, however, the organ has been made into a much more efficient sucking apparatus than that of the cockroach by the union of the edges of its lower hypopharyngeal wall with the epi- pharyngeal wall, thereby converting the lumen into a closed cavity. The clypeal muscles on contraction lift the flexible dorsal wall and expand the lumen, drawing in the liquid food from the canal of the labrum. On relaxation of the muscles the dorsal wall snaps back by its own elasticity and drives the liquid from the pump back through the mouth. On the epipharyngeal wall of the cibarial pump are small spines and papillae of various kinds, some of which are sense organs. A com- parative study of these structures and an armature of ventral teeth at the mouth entrance has been made by Sinton and Covell (1927), and Chwatt and Major (1945) in the anophelines, and by Barraud and Covell (1928) in anopheline and culicine species. The epipharyngeal sense organs are described by Day (1954). The pharyngeal pump.—From the mouth at the inner end of the cibarial pump the stomodaeal section of the alimentary canal begins as a narrow tube (fig. 24 A,E) that curves upward and backward in the head, going between the brain (A, Br) and the suboesophageal ganglion (SoeG). Behind the brain it expands into a large, bulblike structure, which is the pharyngeal pump (PAP). The walls of the organ when relaxed are deeply concave above and on each side, as seen in cross section at B. Into the concavity of the dorsal wall is inserted a pair of large muscles (A,B,E, S) from the dorsal wall of the head behind the brain, and into each lateral concavity a large flat muscle (zr) from the side of the cranium. Contraction of the muscles dilates the lumen of the pump; on their relaxation the walls spring together again by their own elasticity. From the rear end of the pump, the narrow oesophagus (Oe) proceeds through the neck into the thorax. A cibarial and a pharyngeal pump like those of the mosquito are common to bloodsucking nematocerous flies. Presumably the two pumps work in alternate phases to keep the ingested blood flowing freely back into the stomach. In the nectar-feeding male mosquito the sucking apparatus is less strongly developed than in the female. 64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 THE THORAX The thorax of a winged insect may truly be said to be the most re- markable anatomical mechanism developed anywhere in the animal kingdom. It is remarkable both for its efficiency as a flight mechanism and for its structural simplicity. In insects with two pairs of wings the two wing-bearing segments have essentially the same structure, and are equipped with duplicating sets of muscles. In the Diptera, however, in which the flight function has been taken over entirely by the first pair of wings, the mesothoracic wing muscles have to do the work of the muscles of both winged segments in four-winged insects. Consequently, the mesothorax of the flies has been greatly enlarged and the metathorax much reduced. The knobbed stalks known as halteres borne on the metathorax are undoubtedly reduced wings, since, as seen in the mosquito pupa (fig. 17 E), they are developed in flat wing lobes of the metanotum. They are still important accessories of flight, being vibratory organs for maintaining the equilibrium of the flying insect, but their musculature is very simple, and the usual wing musculature of the segment has been eliminated. In the adult mosquito (fig. 25) the mesothorax appears as a great wedge inserted between the narrow prothorax and metathorax. It alone retains the structure typical of a thoracic segment. Two princi- pal plates, an anterior notum (AN,) and a posterior postnotum (PNz), cover almost the entire dorsum of the thorax. The strongly convex postnotum, furthermore, is deeply infolded posteriorly be- neath the narrow metanotum (Ns) and extends into the first ab- dominal segment as a bilobed phragma (fig. 27D, Ph). A narrow paranotal fold (pf) borders the edge of the notum between the first spiracle and the wing. The pleural area tapers downward and becomes continuous with the sternum (S,) between the first and second legs. A typical pleural sulcus (PIS,) extends from the base of the middle leg to the wing fulcrum at the base of the wing (W). The area before the groove is episternal, that behind it epimeral. The episternal area includes a major episternal plate (Eps,) continuous below with the sternum, and a smaller preepisternum (eps,). The epimeron (Epme) is a simple quadrate plate. Below it is a small triangular plate (Ss), which in the mosquito appears to be a postcoxal lobe of the sternum ; but a plate in the same position in higher flies is the detached meron of the coxa. In some species the episternum is divided into an upper and a lower part (fig. 27 A). The prothorax is so reduced and modified that it is difficult to interpret its parts. The notum (fig. 25 Nz) includes a narrow plate No. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 65 across the back beneath the overhanging front end of the mesonotum, and apparently a larger posterior plate on each side. This posterior plate, however, tapers narrowly down to the coxa so that its lower part must be epimeral. The episternum then is represented by a short plate (Eps,) between the first notal plate and the coxa. A plate in the side of the neck (CvP1) that supports the head is unquestionably a cervical sclerite. Fic. 25——Thorax of Psorophora, with head and base of abdomen (from the author’s illustration in Howard, Dyar, and Knab, 1912). AN, wing-bearing notal plate; CvPl, cervical plate; Epm, epimeron; Eps, episternum; eps, preepisternum; H/t, halter; N, notum; PI, pleuron; PIS, pleural sulcus; PN, postnotum; puf, paranotal fold; S, sternum; Scl, scutellum; Sct, scutum; W, wing. Subnumbers 1,2,3 designate parts of prothorax, mesothorax, and metathorax. The metathorax is even more simplified than the prothorax. The notum (fig. 25, Ns) is much narrowed across the back, but it expands on the sides where it carries the halteres (H/t). From the notum the pleural region continues downward on the side, tapering to the hind coxa. Close to its posterior margin is a faint line that perhaps repre- sents the pleural sulcus. A narrow strip (PNs;) between the meta- notum and the first abdominal segment, more plainly seen in Aedes (fig. 27 C,D, PNs), is clearly the metapostnotum, since it gives at- tachment to the first abdominal muscles (G). The wings of the mosquito have a simple pattern of venation, shown at A of figure 26, in which the veins are named according to the Com- 66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 stock-Needham system. Mosquito taxonomists, however, usually designate the veins behind the subcosta by numbers. In this scheme R, is vein 1, Rg and Rs are branches of vein 2, R,,5 is vein 3, M and its two branches are vein 4, Cu and its two branches are vein 5, and A is vein 6. The veins are densely clothed on both sides of the wing with long, slender, fusiform, or scalelike setae (omitted in the figure). While the simple venation of the mosquito wing is of a fairly generalized pattern, the basal wing structure has little resemblance R Se \ i ‘ R Cup Cu, M3 My+2 ae Fic. 26.—The wing and halter. A, Culex, left wing and halter, wing partly flexed on basal lobe, scales re- moved to show venation. B, Anopheles, base of wing, flattened. C, Culex, halter enlarged, posterior. A, anal vein; C, costa; Cu, cubitus; M, media; R, radius; Sc, subcosta. to that of most other insects, and would appear to be specialized by elimination of the usual axillary sclerites. When the wing is flexed (fig. 26 A) a fold near the base sets off a triangular basal lobe by which the wing is attached to the thorax. During flexion the wing turns posteriorly over the basal lobe, which is then covered from above by the fully flexed wing, and gives the wing the appearance of being supported on a lobe of the thorax. The principal sclerotization of the wing base is a long, anterior jointed bar (B, r) that supports the radial vein, and bends at the joint when the wing is flexed (A). No. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 67 Otherwise the membrane of the whole basal area is occupied by irregu- lar thickenings or weak sclerotizations that are hardly sclerites and seem to have no mechanical significance. They are better developed in Anopheles (B) than in Culex (A). The same structure in modified form is present also in some related Nematocera, but not in Tipulidae. The wing mechanism of extension and flexion is not understood, but all the direct muscles of flight appear to be attached on the basal lobe. The indirect flight muscles are as fully developed as in any other fly. They include great masses of dorsal longitudinal fibers and lateral vertical fibers that almost completely fill the thorax. The weight of the flight muscles of Aedes has been calculated by Hocking (1953) as from 16.5 to 18.7 percent of the total body weight, which, however, is small as compared with Tabanus in which the flight muscles are 23 to 35 percent of the body weight. The rate of the wing vibration in flight, measured in wing beats per second, is given by Sotavalta (1947) for females as 165 to 196 for Culex pipiens, 165 to 247 for Anopheles maculipennis, 241 to 311 for Aedes cantans and Aedes punctor. With males the rate is consistently higher, from 330 to 587 beats per second by Anopheles and Aedes. Hocking (1953) has measured the flying speed of five species of Aedes. In ordinary cruising flight they go from 75 to 110 centimeters per second, but for short distances they can make 220 to 252 centi- meters in a second. The legs of the mosquito have no unusual features, except for their length and relative slenderness. Each leg (fig. 17 B) has the usual six segments of an insect leg, a coxa, trochanter, femur, tibia, tarsus, and pretarsus. The long tarsus is subdivided into five tarsomeres. The pretarsus has two decurved claws but no arolium. In some species, as in Culex, the foot is provided with a pair of small padlike pulvilli; in others there is only a heel-like hairy swelling at the bases of the claws. Most mosquitoes, however, whether they have foot pads or not, are able to cling to smooth vertical surfaces, such as window panes or the walls of a glass jar. THE ABDOMEN The abdomen of the adult mosquito (fig. 27 A) is broadly joined to the thorax and tapers posteriorly. The tergal and sternal plates are separated on the sides by membranous areas containing the spiracles, which are present on segments I to VII. In each sex the abdomen has Io segments, as in the pupa, but in the females of some species the eighth segment is ordinarily retracted into the seventh, and in the male the ninth segment is concealed within the eighth. 68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 Fic. 27.—Details of the adult thorax and abdomen. A, Aedes aegypti, male thorax and abdomen. B, Same, end of abdomen ex- tended. C, Same, thorax and base of abdomen, dorsal. D, Same, postnotum of metathorax extended as a phragma into base of abdomen, ventral. E, Same, tenth abdominal segment of male, lateral. F, Same, undersurface (dorsal). G, Same, showing dorsal muscles of first abdominal segment. H, Culex, end of female abdomen, ventral. I, Same,. lateral. J, Composite diagram of female terminalia, lateral (from Gerry 1932). K, Same, ventral, with ventral arc of sigma (c) turned forward (from Gerry 1932). a, cowl; An, anus; b, dorsal arc of sigma (ninth sternum?) ; c, ventral arc of sigma; Cer, cercus; Gir, gonotreme (opening of genital chamber) ; H/t, halter; N, notum; pgpl, postgenital plate; Ph, phragma; Pmr, paramere; PN, post- notum; s, lateroventral prong of tenth segment; S, sternum; Scl, scutellum; Sct, scutum; ¢, tergum of tenth segment; 7, tergum. Subnumbers 1-3, thoracic segments; J-X, abdominal segments. No. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 69 The male mosquito is readily distinguished from the female by the presence of a pair of large, two-segmented genital claspers, or parameres, projecting from the end of the abdomen (fig. 27 A, Pmr). Though the ninth segment is ordinarily concealed by retraction into the eighth, on pulling out the end of the abdomen (B), it is seen to be a small sclerotic ring (JX) carrying the parameres. The anus- bearing tenth segment, or proctiger (X), is mostly hidden between the bases of the parameres, and is apparently ventral in position. In fact, the whole terminal part of the male abdomen beyond the seventh segment, except in newly emerged individuals, is turned upside down, so that the tergal plates are ventral and the sternal plates dorsal. The inversion takes place slowly during the first 24 to 48 hours after emergence from the pupa. The tenth abdominal segment of the male is a flattened anal lobe with an expanded base projecting from above the inverted tergum of the ninth segment (fig. 27 E,F). In its base are two dorsolateral sclerites (¢) that may be regarded as tergites. On the ventral (upper) surface are two marginal bars (s), the ends of which project as a pair of free, toothed prongs. These bars have commonly been re- garded as sternites, but Christophers (1923) says they are the cerci united with the anal lobe. The external genital organs of the male insect, because of their generic and specific variations, are important diagnostic features for taxonomists. In the mosquito they include primarily the paired lateral claspers and a median intromittent organ, carried by the ninth ab- dominal segment. Various names are given to these parts by different specialists, but the organs have essentially the same origin in all insects, and there is no need for special terms in the several orders, and certainly there is no excuse for specialists in one order to use different names for the same parts in different species. For simplicity the claspers are here termed the parameres, and the intromittent organ the aedeagus. Various secondarily developed accessory parts, of course, must have more specific names. In the insects in general the male genitalia take their origin from a pair of primary phallic lobes that develop in a late instar of the nymph or larva on the posterior part of the ninth abdominal segment at the sides of the future gonopore. Later, each lobe divides into two parts, a mesal mesomere and a lateral paramere. Eventually the mesomeres unite around the gonopore to form the aedeagus, and the parameres become the claspers. The development of the genital organs in the male mosquito has been shown by Christophers (1922) to proceed in the usual manner. 7O SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 Early in the fourth instar of the larva paired thickenings of the epi- dermis appear behind the region of the ninth sternum. These “genital plaques” soon take on the form of budlike outgrowths, which are the primary phallic lobes (fig. 28 A, PhL). With further development the lobes elongate and unite at their bases, forming the genital ap- Fic. 28.—External genital organs of the adult male, and their development. (A,B,C, from Christophers, 1922.) A, The primary phallic lobes that appear in a late instar larva behind the sternal region of the ninth abdominal segment. B, Later stage of same, each primary lobe divided into a mesomere and a paramere. C, Still later stage, mesomeres united around the gonopore to form the aedeagus. D, Adult genital apparatus of Anopheles quadrimaculatus, lower surface (dorsal). E, Parameres and claspettes of Aedes pullatus. Aed, aedeagus ; Bmr, basimere; bp, basal plate; c/sp, claspette; Gpr, gonopore; IXT, ninth abdominal tergum; Mmr, mesomere; PhL, primary phallic lobes; Pmr, paramere; Tmr, telomere. pendages as they appear in the pupa (fig. 16F). At this stage the lobes are termed ‘‘proandropodites” by Christophers (1922), but this term literally translated would mean “primitive male parts of legs” (as “coxopodite” means the “coxal part of a leg’). Since there is no real evidence that the male genital organs of insects represent primitive legs, the genital organs of the pupa are simply the developed phallic lobes. Within them are formed the definitive genitalia of the no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 71 adult. From the base of each lobe inside the pupal cuticle, as de- scribed by Christophers, is cut off a small median lobe (fig. 28 B, Mmr), and the lateral part becomes the rudiment of the clasper (Pmr). Finally, the two median lobes unite around the gonopore to form the aedeagus (C, Aed), while the lateral parameral lobes elongate to become the two-segmented claspers of the adult (D,E). In the mature condition the genitalia take on a great variety of forms and are complicated by the development of accessory parts. All this is a great boon to taxonomists, but it often creates difficulty for the morphologist. Anopheles quadrimaculatus (fig. 28D) gives a good example of one type of structure. Each paramere is divided into a large basimere (Bmr) and a long slender telomere (Tmr). The telomere is movable on the basimere by strong antagonistic muscles arising in the latter. The slender aedeagus (Aed) lies between the bases of the parameres and is connected with the basimeres by a pair of small basal plates (bp). The basimeres are equipped with long spines, and proximally each bears a membranous median lobe (clsp) united with the one from the opposite side. Each lobe is armed with strong spines and is known as a claspette, or claspette lobe. In other genera the claspettes are more commonly independent ap- pendages of the parameres, as seen in Aedes (E). The claspettes, according to Christophers (1922), are cut out from the parameres by secondary incisions of the latter. For illustrations of generic and specific variations in the male genital structure the student must consult taxonomic papers, but the nomenclature will be confusing. In the current terminology of mosquito specialists, the aedeagus is called the “mesosome” or “phallo- some,” the basal plates (bp) that connect it with the claspers are the “parameres,” and the claspers are the “side pieces.” In this scheme the term “paramere” is entirely misapplied, since it was first given to the claspers, and moreover, “side piece” is a direct English translation of “paramere.” The segments of the claspers are known also as the “basistyles” and “‘dististyles,’ but as shown by their development the claspers have no relation whatever to legs or abdominal styli. The terminology given on figure 28 is recommended for its simplicity and because it can be applied, on the basis of development, to the male genitalia in all the principal orders of insects (see Snodgrass, 1957). The terminal parts of the female abdomen are much simpler than those of the male, but their homologies are more difficult to under- stand. Beyond the eighth segment projects a small lobe (fig. 27 1) representing the combined ninth and tenth segments. The dorsum of 72 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 the ninth segment is a transverse basal arc (JX) usually containing a small tergal sclerite. Beyond it is the tenth segment (X) bearing a pair of lateral cerci (Cer) and the terminal anus (4m). Ventrally is a lobe known as the postgenital plate (pgpl) because the gonotreme (H, Gtr), or opening of the genital atrium, is situated at its base above the sternum of the eighth segment (VJJ/). The nature of the postgenital plate is doubtful ; it looks as if it should be the projecting sternum of the ninth segment. On its base there is generally a trans- verse fold known as the cowl (K, a) because it is sometimes reflected to form a hoodlike pocket. Surrounding the gonotreme above the end of the eighth sternum is a sclerotized ring (b, c) named the sigma by Christophers (1923). In figure K the ventral arc of the sigma (c) is turned forward; normally it is directed posteriorly (J, c). The sigma thus, as described by Christophers, resembles the lips of a half- opened clasp purse, in which it is represented by the metal frame- work of the purse. Some writers, however, without adducing specific evidence, regard the dorsal arc of the sigma as the ninth sternum. According to Christophers the whole structure is formed as a scleroti- zation in the intersegmental membrane of the gonotreme. All parts of the female terminalia are subject to much variation, as shown in comparative studies by Macfie and Ingram (1922), Christo- phers (1923), Davis (1926), Gerry (1932), Gjullin (1937), Roth (1946), Rees and Onishi (1951), and Hara (1957). The student, however, will be somewhat confused by the different ways the parts are represented and named. The drawings J and K on figure 27, taken from Gerry, are composite diagrams showing all the parts that have been described, but they probably do not present the exact structure in any one species. The gonotreme surrounded by the sigma above the eighth abdominal sternum leads into a small infolded pouch, the genital chamber, or atrium. In its anterior wall is the female gonopore, which is the open- ing of the median oviduct. Behind the gonopore the globular sperma- thecae (one, two, or three in number) open through the dorsal wall of the atrium, and into a posterior pouch of the dorsal wall, the caecus, opens the single accessory gland, called the “mucus gland,” but the nature of its secretion is not known (fig. 30 B). INTERNAL ANATOMY A thorough study of the internal anatomy of the mosquito has not been made, but the parts of principal interest will be the alimentary canal and the reproductive organs. The muscular and tracheal systems no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 73 have no features peculiar to the mosquito, and even the unusual char- acters of the reproductive organs are common to other Diptera. The simple nervous system is that of the larva with an elaboration of the brain and the optic lobes in the head, a transposition of the first abdominal ganglion to the thorax, and the union of the eighth ab- dominal ganglion with the ganglion of the seventh segment. In the abdomen of the adult, therefore, the first ganglion is in the second segment (fig. 30C, Gngl/), and the last is a composite ganglion (Gng VII+VIII) in the seventh segment. The tracheal system has lost the large dorsal trunks of the larva, and the lateral trunks along the spiracles have been enlarged. The circulatory organs.—In the adult mosquito, as described by Jones (1954) in Anopheles, the dorsal blood vessel has in general the same structure as that of the larva. The part in the abdomen, how- ever, is more distinctly “chambered” because of segmental swellings before the ostia. An aortic sinus is said by Jones (1952) to be present in the adult as in the larva and pupa of Anopheles, Culex, and Aedes. The sinus is a dilatation of the aorta in the dorsal part of the thorax, with the corpora allata-cardiaca attached to it laterally. Anteriorly the sinus is continued into the cephalic aorta. The adult heart, ac- cording to Jones, beats predominantly forward, but periodically re- verses the direction of the beat. The heart has no innervation from any source and therefore its pulsations are myogenic, that is, engen- dered by the muscles themselves of the heart wall. Lateral alary muscles support the heart, but they do not vibrate, and when cut the heart keeps on beating. A vibratile muscular membrane across the cavity of the mesothoracic scutellum appears to be an accessory pulsatile organ, as in some other insects. A frontal bulblike organ between the bases of the antennae has been described by Day (1955) as a sense organ, and by Clements (1956) as a pulsating organ for driving blood into the antennae. If it is a sense organ, it is a newly discovered one as Day claims; if it is a pulsating organ it is not unique since a pulsatile organ in the same place is present in various other insects. The alimentary canal——The alimentary canal of the adult mosquito (fig. 29 A) in its general form is quite different from that of the larva. From the pharyngeal pump in the head (PhP) a short, narrow oesophagus (Oe) extends into the front of the thorax, where it joins a wider tube, which is the beginning of the stomach, or ventriculus (Vent). Shortly before its junction with the stomach the oesophagus gives off three pouches, known as the oesophageal diverticula, two 74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 of which are dorsal and one ventral. In Aedes aegypti the dorsal diverticula (A, ddv) are small, flat, elongate sacs with slender necks diverging forward and laterally from the oesophagus (C). The single ventral diverticulum (4, vdv) has a long, slender neck which ex- pands into a large sac in the anterior half of the abdomen. This ventral diverticulum corresponds with the usual ‘“‘crop” of other Diptera. Fic. 29.—Alimentary canal and salivary glands of the adult female of Aedes aegypti. A, Sectional view of body showing alimentary canal and salivary system (glands on left displaced). B, The salivary glands. C, The oesophageal diverticula, ventral. D, Rectal sac and papillae. Alnt, anterior intestine; CbP, cibarial pump; ddv, dorsal diverticulum; Hphy, hypopharynx; Lb, labium; Lm, labrum; Mal, Malpighian tubules; Oe, oesophagus; PhP, pharyngeal pump; Rect, rectum; rp, rectal papillae; S/Dct, salivary duct; S/GI/d, salivary glands; S/P, salivary pump; vdv, ventral di- verticulum; Vent, ventriculus. The ventriculus (fig. 29 A, Vent), which is the functional stomach of the insect, for most of its length in the female mosquito is a narrow tube that extends upward through the thorax and then turns backward into the abdomen where it ends in a saclike enlargement that joins the intestine. The first part of the latter, or anterior intestine (AlInt), is a short, slender tube thrown into a small loop. Its anterior end, the pyloric region, joins the ventriculus by a funnel-shaped ex- pansion. At the other end the anterior intestine is continued into the no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 75 posterior intestine, or rectum (Rect), which is much enlarged an- teriorly and tapers back to the anus. The inner wall of the pyloric funnel is armed in some species with numerous small spines directed posteriorly. These pyloric spines have been described and well il- lustrated by Trembley (1951) in species of Anopheles, Aedes, and Culex. In the anterior end of the rectum are six small, soft, conical rectal papillae (D, rp) projecting inward from the rectal wall. Five Malpighian tubules (A, Mal) arise from the pyloric region of the intestine as in the larva. The oesophageal diverticula are said to be empty on emergence of the mosquito from the pupa. Within an hour after ecdysis, however, according to Marshall and Staley (1932), the air that was pumped into the stomach begins to pass forward into the diverticula, and in 12 to 22 hours the stomach is empty. The function of the oesophageal diverticula in relation to food intake has been studied by a number of investigators, but, though using the same experimental methods of feeding, the latter have not all come to the same conclusions. The subject has recently been well re- viewed by Trembley (1952) and by Megahed (1958), and good bibliographies are given by both these writers. In general it is found that ingested blood goes directly to the stomach, while fruit juices and sugar solutions go first into the diverticula, to be later delivered to the stomach. According to Trembley, blood in small amounts may occasionally go to the diverticula, and sugar solutions sometimes go direct to the stomach. The work of Megahed on Culicoides gives essentially the same results, the stomach being ordinarily the receptacle for blood, the diverticula for concentrated sugar solutions, but water and dilute sugar solutions go direct to the stomach. Most observations seem to apply to the female insect. Day (1954), however, in experi- ments on male mosquitoes, found that the sexes react similarly : “blood went to the mid-gut and sugar to the diverticulum in the male in spite of the fact that males do not ingest blood under natural conditions.” The “switching mechanism” that determines whether the ingested food goes into the stomach or the diverticula, Day (1954) has pro- posed, is governed by the different kinds of sense organs in the wall of the cibarial pump (buccal cavity). If receptors of one type are stimulated by sugar it may be supposed that they cause a relaxation of sphincter muscles of the diverticula; if others are sensitive to blood components, they may effect a relaxation of the cardiac sphincter of the stomach. In the neck of the ventral diverticulum, Day notes the presence of a group of spines, which would appear to assist in keeping 76 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 blood corpuscles out of the diverticulum when the circular muscles in the neck of the diverticulum are contracted. The salivary glands.—The salivary glands of the mosquito consist each of three lobes (fig. 29 B), of which the middle lobe is shorter than the other two. The glands lie at the sides of the anterior end of the ventriculus (A, S/G/d; the left gland is displaced in the figure). The two ducts extend into the back of the head, where they unite in a single outlet tube (fig. 24 A, SIDct), which ends at the base of the hypopharynx in a small syringelike swelling that acts as a salivary ejection pump (S/P). On the elastic dorsal wall of the pump is in- serted a dilator muscle (18) from the floor of the cibarial pump. The salivary pump discharges through the salivary canal (sc) of the hypo- pharynx in the female; in the male the duct traverses the labium (fig. 22 F). The salivary secretion in species of Anopheles, according to Metcalf (1945), contains both an anticoagulin and an agglutinin, but in other pest species neither appears to be present. The salivary glands are of particular interest in connection with the transmission of disease by mosquitoes. They offer the only avenue of escape for disease organisms from the body cavity of the mosquito into the blood of an alternate host. The sporozoites of malaria, for ex- ample, that penetrate into the salivary glands are carried in the saliva of the biting mosquito directly into the vertebrate host, which is necessary for the completion of the complex life history of the malaria parasite, Plasmodium. This suggests the question of how it became obligatory for some parasites to divide their developmental history between two different animals, but the known facts give no answer. Mosquitoes do not bite each other, and there is no way by which the malaria parasite can be normally transferred from one vertebrate to another. The reproductive system.—The organs of reproduction in the Diptera include the parts common to all insects, but their structure in two respects is exceptional. Each testis appears to correspond with a single testicular tube in other insects; the egg tubes of each ovary are extremely small, and all are enclosed in a cellular sheath. The male organs of the mosquito include a pair of testes (fig. 30 E, Tes), a pair of testicular ducts, or vasa deferentia (Vd), which enlarge posteriorly to form a pair of seminal vesicles (SV) that in some species are united (D). The vesicles end in a very short common ductus ejaculatorius (Dej), which receives a pair of large accessory glands (AcG/d) and then opens directly into the base of the aedeagus (Aed). In the normal condition the reproductive organs lie beneath the alimentary canal, but, with the inversion of the terminal segments of the abdomen, the relation is reversed (fig. 30 A)—the ejaculatory no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS TG. Fic. 30.—Reproductive organs and the abdominal nerve cord of the adult mosquito. A, Culex quinquefasciatus, end of male abdomen, left side removed, exposing the inverted reproductive organs and rectum. B, Diagram of female reproductive organs, dorsal (adapted from Christophers, 1901). C, Aedes aegypti, male, ventral nerve cord of abdomen and genital outlets, dorsal. D, Same, male re- productive organs, ventral. E, Culex quinquefasciatus, male reproductive organs after inversion, lower side (dorsal). F, Same, with part of intestine, upper side (ventral). AcGld, accessory gland; Aed, aedeagus; Atr, atrium, genital chamber; Bmr, basimere; Dej, ductus ejaculatorius; Gng, ganglion; Gtr, gonotreme; Int, intestine; Odc, oviductus communis; Odl, oviductus lateralis; Ov, ovary; Ovl, ovariole; Pmr, paramere; Ptgr, proctiger; Rect, rectum; S, sternum; Sh, sheath of ovary; Spt, spermatheca; SV, seminal vesicle; T, tergum; Tes, testis; tf, terminal filament of ovary; Tmr, telomere; Vd, vas deferens. 78 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 duct, the seminal vesicles, and the accessory glands now lie above the intestine. Since the testes are not affected by the inversion, the long vasa deferentia as in Culex (E) cross each other, but when the ducts are united as in Aedes (D) a simple twist takes place at the junction of the ducts. The testis of most insects consists of a number of individual tubes in which the spermatozoa are formed as are the eggs in the ovarian tubes, and, except in the apterygotes, the tubes of each testis are enclosed in an investing sheath. The testes of the mosquito are elongate, pear-shaped bodies (fig. 30 D,E,F, Tes) continuous with the ducts. Each testis, however, appears in its entirety to be a single testicular tube. The same is true of the testes in other Diptera. In the narrowed upper end of each organ is a mass of undifferentiated cells ; the rest of the lumen is filled with spermatocytes and spermatozoa in various stages of development. The mature spermatozoa are ex- tremely long and threadlike; when liberated from the testis they exhibit active undulatory movements. The spermatozoa are stored in the seminal vesicles preliminary to mating, and the accessory glands probably have a prostate function, giving the spermatozoa a liquid medium in which they are discharged. The reproductive organs of the female mosquito, represented diagrammatically at B of figure 30, include the parts characteristic of the female organs of insects in general. These are a pair of ovaries (Ov), the lateral oviducts (Odl) from the ovaries, and a median common oviduct (Odc) with which the lateral ducts are joined. The common duct opens by the primary genital aperture, or gonopore, into a small pocket above the end of the eighth abdominal sternum. This pocket, the genital chamber, or atrium, being a secondary inflection of the body wall between the eighth and ninth abdominal segments, is therefore not a part of the primary genital passage. The external opening of the atrium may be designated the gonotreme (Gtr). Into the dorsal wall of the atrium just behind the gonopore open the ducts of the spermathecae (Spt), which are usually three in number, though in Anopheles there is only a single spermatheca. Behind the spermathecal openings arises an accessory gland (AcGld), the func- tion of which is not known in the mosquito. In other insects accessory glands usually secrete a cement for attaching the eggs to a support, or a material to form an egg covering. The atrium serves as a copulatory pouch at the time of mating, and the spermatozoa from the male are stored in the spermathecae. Then when the eggs leave the oviduct they are received in the atrium and no. 8 ANATOMICAL LIFE OF THE MOSQUITO—SNODGRASS 79 are here fertilized by sperm discharged from the spermathecae. Finally the eggs are passed out through the gonotreme at the time of laying. The ovaries of the mosquito differ in several respects from the usual structure of these organs in other insects. A typical insect ovary consists of a group of slender tubes known as ovarioles opening into the end of a lateral oviduct. The ovarioles taper upward and end in filaments that unite in a common strand attached to tissues in the neighborhood of the heart. A mature ovariole contains a series of ripening egg cells of successively larger size, with the mature egg in its lower end. Each egg is accompanied by a number of nutritive cells, or so-called nurse cells, which are absorbed by the egg as it matures. Each egg and its nurse cells are contained in a compartment of the ovariole known as a follicle. The follicles appear as swellings along the ovariole, increasing in size with the growth of the egg. The egg cell and the nurse cells are formed by division of the undif- ferentiated cells in a chamber, the germarium, in the upper end of the follicle. The eggs do not pass down the ovarioles; each ovariole grows from the germarium as an egg leaves the lowermost follicle and the latter disintegrates. In the mosquito ovary (fig. 30 B) the ovarioles (Ovl) are very short and are arranged in rows along an axial cavity of the ovary. As in other Diptera, each ovary is invested in a thin membranous sheath (Sh) in which there are fine muscle fibers, and the sheath itself ends in a terminal filament (tf) attached to tissues along the sides of the heart. The muscle fibers of the ovarian sheath in Anopheles are said by Nicholson (1921) to be striated, but Jones (1958) finds that those of Aedes do not show a distinct striation in live, unstained whole mounts at 1,000 magnification under phase optics. Each ovariole consists of a large egg-containing follicle with a small projection on its free end representing the germarium and one or two minute undeveloped follicles, The structure of the egg follicle of Culex has been described by Nath (1924), and an account of the de- velopment of the ovary and the development and nutrition of the eggs in the ovary of Anopheles is given by Nicholson (1921), by Christo- phers, Sinton, and Covell (1928), and by Mer (1936). The develop- mental processes described in the mosquito differ little from those in insects generally. Many female mosquitoes need a meal of blood for the production of eggs. The eggs of Anopheles and Aedes are fully developed in two to three days after the female has fed. 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ABBOT Research Associate, Smithsonian Institution (Pusication 4390) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION MARCH 23, 1960 THE LORD BALTIMORE PRESS, INC. BALTIMORE, MD., U. S. A. Roebling Fund A LONG-RANGE FORECAST OF UNITED STATES PRECIPIFATION By C. G. AxBBor Research Associate, Smithsonian Institution FOREWORD A hidden family of harmonic regular periods exists in weather. The periodic members of this family persist with unchanged lengths for scores of years. By determining their average forms and ampli- tudes for intervals of a thousand months, successful forecasts may be made for years to come; or backcasts may be made for former years and compared to former events. Agreement of such backcasts with the records warrants confidence in future forecasts. These claims seem preposterous to most meteorologists. Therefore, before proceeding to explain the method and to give forecasts to 1967 for 32 cities of the United States, illustrative forecasts for the years 1950 to 1958 will now be shown and compared to the records of that interval graphically. Figures 1, 2, and 3 show forecasts (dotted) and the observed march of precipitation, 1950-1958. These curves represent 3-month running means, and are expressed in percentages of normal precipitation. Fig- ure I represents precipitation at Madison, Wis., and figure 2 at Nash- ville, Tenn. The curve at the top of figure 2 will be described later. Figure 3 shows forecast and observation for Sacramento, Calif. I have computed for several cities coefficients of correlation be- tween my forecasts and the observed precipitation for the years 1950 through 1958. They are as follows: Washington, D. C., 52.3 percent ; Cincinnati, Ohio, 57.3 percent ; Nashville, Tenn., 59.0 percent ; Inde- pendence, Kans., 52.0 percent; Madison, Wis., 56.6 percent; Sacra- mento, Calif., 69.0 percent. These coefficients indicate that my forecasts are over halfway to- ward perfect long-range prediction of weather. There still remain undisclosed variables that produce the discrepancy of about 40 percent between my coefficients and perfect correlation. SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 139, NO. 9 VOL. 139 SMITHSONIAN MISCELLANEOUS COLLECTIONS ‘suvall SuIUUNI YUOW-L£ WOIJ [[Y ‘2AINd [[N} Guede four] peyop ‘\svoo10} foul] [eWOozti0y AAvay ‘feuION “gS61-O0S61 “uoHepdIoeI1d [eIOU WOIF soInjIedap pee JO JUOAD puUv JSBIVIO “SIM “UOSIPEIY—I “DIy] a 1S Fs ag a eee Pict as rT eee Le — + VAI be on ae boise fr bo |p Very pou [ere ba FORECAST OF U. S. PRECIPITATION—ABBOT 9 NO. ‘suvaL SuIUUNI YJUOUI-£ WOIJ [TY ‘dAIND [[NJ YusAs foul] po}Op YseoaroF f aUT] [e}UOZIIOY AAvOY ‘eur -ION *9S61-0S61 ‘uorepdi901d JeuIOU Wor, sainjiedsp A][YyJWOU Jo Jada pure jsvoIIO “WUaT, ‘o]]IAYSeN—z “SI 6561 £15) PER eS6t C A 7161 2561 9S61 SS6I vS6l £s6l zS61 1S6I pa EM ES: eee) ere ce et lb afb ATUL VA pT PURE ae Aer 16! O16! 6061 g06! 2061 SECA UESE SAV ra Et A 906) S061 Ab ie | CHEETA “CANE ROMA CEE CECI BORE eR EE) Me epi PU EE COCCI CTD TAT NECA CS ONS PAT NENT SICA ' ACCC NPA CREE ECR Ce EP COLOR EEE EEE GO Pre a MM NS og vo6l SMITHSONIAN MISCELLANEOUS COLLECTIONS foul] yewwozioy AAvay as "gS61-0S61 ‘uonepdIoaid jeutoU Woy sammjziedop A]Yy}UOUW # JUDAD PUL JSBIVION “Jed ‘ojuswIVIseGS—E ‘91.7 SS osét, La i} t 238 ai aa / ea eA Ee seen HY of SHE ain? NO. 9 FORECAST OF U. S. PRECIPITATION—ABBOT 5 FORECASTS OF PRECIPITATION FOR 32 CITIES, 1950-1967 This project was sponsored by the Association for Applied Solar Energy of Phoenix, Ariz., and the Smithsonian Institution of Wash- ington, D. C. Funds for the costs of electronic computations were supplied to the Association by the Valley National Bank and the Arizona Public Service Company. About 7,000 tables of precipitation were electronically computed by Jonathan Wexler, a student at the Arizona State College at Tempe. He ingeniously programmed the machine for this special purpose. Monthly records of precipitation at 32 stations from about the year 1870 were taken from publications generously furnished by the United States Weather Bureau. TABLE 1.—List of stations 1. Abilene, Tex. 17. Nashville, Tenn. 2. Albany, N. Y. 18. Natural Bridge, Ariz. 3. Albany, Oreg. 19. Omaha, Nebr. 4. Augusta, Ga. 20. Peoria, Ill. 5. Bismarck, N. Dak. 21. Port Gibson, Miss. 6. Charleston, S. C. 22. Rochester, N. Y. 7, Cincinnati, Ohio 23. Sacramento, Calif. 8. Denver, Colo. 24. Salisbury, N. C. 9. Detroit, Mich. 25. Salt Lake City, Utah 10. Eastport, Me. 26. San Bernardino, Calif. 11. El Paso, Tex. 27. Santa Fe, N. Mex. 12. Helena, Mont. 28. Spokane, Wash. 13. Independence, Kans. 29. St. Louis, Mo. 14. Little Rock, Ark. 30. St. Paul, Minn. 15. Madison, Wis. 31. Thomasville, Ga. 16. Montgomery, Ala. 32. Washington, D. C. Secretary Leonard Carmichael of the Smithsonian Institution as- signed Mrs. Lena Hill and Mrs. Isobel Windom to assist me in preparing forecasts. He approved grants from funds given for the study of solar radiation and weather by the late John A. Roebling. I am greatly indebted to Miss M. A. Neill for careful preparation of my manuscript. I selected 32 cities distributed with approximate uniformity over the United States. The cities chosen are listed in table 1. THE METHOD As I suppose no one hitherto has ventured to predict values of precipitation, at definite places, for as much as 8 years in advance, I now indicate briefly how it is done. I quote apposite passages from 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 my former papers,’ with slight changes dictated by later experience. Periods in sun and weather.—The sun’s radiation which we see and feel, like that of many other stars, is variable. Solar output of radiation seldom exceeds 2 percent in its variation. However, its variation comprises as many as 60 regular periodic pulses, ranging from 1 month or less to 273 months or more. All are exact submul- tiples (or aliquot parts) of 273 months, as 91, 39, 7 months, and many more. They range in amplitude from 1/50 to 1/4 percent. All go on simultaneously, like overtones of a musical note. As many as 30 of these exact periods have been found in monthly weather records which have been kept from 1870 and earlier. They occur in records both of precipitation and temperature. Far from being confined to fractions of 1 percent, as in solar radiation, in pre- cipitation they individually range from 5 to 35 percent of the normal average. In temperature they range from 1° to 3° F., and these limits refer to 3-month smoothed records. Owing to the large number of these weather periods, some in plus, some in minus phases at any one time, their combined influence is not usually startlingly great. Normals.—Long records of weather ordinarily state “normal” monthly values found by taking the monthly averages of all the years tabulated. I have found considerable differences in normals if com- puted separately for years of high and low sunspot frequencies, re- spectively. I therefore compute separate monthly normals for years above and below an average of 20 Wolf numbers in sunspot fre- quency. From these normals I tabulate the departures in tempera- ture, and the percentages of normal precipitation. The monthly values have too wide jumps to be most useful. I smooth the record by 3-month consecutive means. Thus for February I use (January+February+ March) X1/3, and similarly for other months. Lags.—Supposing, contrary to meteorologists’ opinion, that the variation of the sun is the real cause of the variation of the weather, since it has identically the same periods, I point out that well-known variations of insolation suffer variable lags in their weather influence, depending on place and time. Lags of solar effects, as they differ with locality, indicate that the state of the atmosphere is an important factor. The atmospheric 1a, Journal of Solar Energy, Sci. and Eng., vol. 1, No. 1, January 1957; b, ibid., vol. 2, No. 1, January 1958; c, Smithsonian Misc. Coll., vol. 122, No. 4, August 1953; d, ibid., vol. 128, No. 3, April 1955; e, ibid, vol. 128, No. 4, June 1955; f, ibid., vol. 134, No. 1, September 1956; g, ibid., vol. 138, No. 3, February 1950. NO. 9 FORECAST OF U. S. PRECIPITATION—ABBOT 7 condition varies not only with locality but with time of the year, prevalence of sunspots, and march of population. To partially meet these difficulties, I tabulate separately for three periods of the year: January-April; May-August ; September-December ; also with Wolf sunspot numbers above and below 20; also with lapse of time before and after the midpoint of the record. These divisions of the available monthly data lead to computing 220 tables at each station before undertaking a forecast. Forecasts by periods —My forecasts are made by adding the effects of 27 regular periodic cycles in precipitation. These cycles, like the harmonics of musical sounds, proceed simultaneously, and are in- tegrally related to a fundamental cycle. This fundamental is 273 months. The harmonics employed are as follows: TABLE 2.—Periods used for forecasting Fraction Months Fraction Months Fraction Months 1/3 gl 1/12 22-3/4 1/27 10-1/9 1/4 68-1/4 1/14 19-1/2 1/28 9-3/4 1/5 54-3/5 1/15 18-1/5 1/30 9-1/10 1/6 45-1/2 1/18 15-1/6 1/33 8-3/1 1/7 39 1/20 13-13/20 1/36 7-7/12 1/8 34-1/8 1/21 13 1/39 7 1/9 30-1/3 1/22 12-9/22 1/45 6-1/15 1/10 27-3/10 1/24 11-3/8 1/54 5-1/18 1/1! 24-9/11 1/26 10-1/2 1/63 4-1/3 The harmonic family referred to was discovered in the variation of the measures of the solar constant of radiation. Figure 4 shows 26 of over 60 periods discovered in solar variation.” Identical cycles were later found in precipitation and temperature by study of long- continued weather records. While the periods of the harmonics are invariable, both in the sun and weather, and their phases are invari- able in solar radiation, their phases shift in weather, depending on atmospheric influences, as will be described below. On account of these phase changes, depending on several variables discovered in my studies of precipitation begun with Peoria, Ill., about 10 years ago, the harmonic family in weather is obscured and hidden, and is as yet unrecognized by most meteorologists. Nevertheless it is verified by an enormous mass of evidence, as will appear below. No observations required—Many meteorologists and others sup- pose that my method of long-range weather forecasting depends on solar observations, but this is not so. The harmonic family referred 2 See in reference, footnote 1, e, above, figure 3 and table 3. VOL. 139 SMITHSONIAN MISCELLANEOUS COLLECTIONS ‘cS61-0Z61 JO SuOT}eAIIS -qO JUP}sUOI-Ie[OS URIUOSY WIS WoIZ [PY ‘sporsod poyefot A][e1sojUT SulsoJsojUL o}yeurpsroqns Jo poseo[o [[@ ‘yjSuey ur syjuoW ve - Zo/¢Ze wor Bursuer ‘UOTyeLIeA IejOsS Ul sported xIs-AyUOMT—V “Oy est Mant Pe ist ak SOMA TI Nae Naar TP Me CPR elie tet er Te ep SE EPA eS ane EbS ahh re ee ere al test | SZ A Baar Ws at As are ae Pee | a eae AP PCa pie ee EN Ce TE a el Cod Pi a Ee NT CEPT TCE a PSA Ti PW TAA Da’ NO. Q FORECAST OF U. S. PRECIPITATION—-ABBOT 9 to was indeed discovered by the study of over 30 years of daily “solar- constant” observations of the Smithsonian Astrophysical Observatory. But now that the harmonic family has been found in weather, no observations of any kind are required. It is only necessary to employ a long record of monthly mean values of precipitation, or temperature, to make long-range predictions. These are approximately verified if no unusual alterations of atmospheric conditions make the averages from long records inapplicable. Sports—As my forecasts depend on the assumption that the av- erage conditions of the periods over a thousand months will be pro- jected into the future, it is important not to include wild “sport” values of precipitation in the thousand-month basis. Hence I have diminished sporadic very high values to about two times normal, and have raised sporadic drought values of less than 4o percent of normal to exceed that limiting low value. These limits refer to 3-month smoothed records. For most of my 32 stations these changes are very rare. But in two or three of the desert stations possibly one value in ten was changed to avoid spoiling the representative character of the basis. The considerable measure of success of my forecasts is the main defense of the method used to produce them. If the degree of success is found to be valuable, no doubt those who in future will use the method will greatly improve it by modifications dictated by reason and experience. Backcasts—Since my forecasts are made by adding the average effects of 27 harmonic periods over an interval of about 1,000 months, the 12 months of record for any one year can produce only about 1 percent of influence on the forecast for that year, even if those 12 months are among the thousand months employed as a basis. There- fore all forecasts or backcasts are equally sound, whether they relate to time before, within, or after the thousand months of record.® The preceding paragraph is important. The forecasts for 32 cities all extend from 1950 to 1967. The degree of similarity between the forecasts and what happened up to 1958 is the index of their prob- able agreement from 1959 to 1967. The 273-month period.—Daily solar-constant observations pro- ceeded from 1920 to 1952 at Montezuma, Chile. This interval is not long enough to determine the master period accurately. But the 10-1/9-month period in weather is a strong one and has long been followed in Washington precipitation. I determined its amplitude for several periods differing slightly from 10-1/9 months. For this pur- 8 See discussion of backcasts at a later page. IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 pose I used about 790 monthly mean values of Washington precipita- tion, all observed when Wolf sunspot numbers exceeded 20. These values were smoothed by 3-month consecutive means, which of course reduces the ranges of percentage departures from normal to about two-thirds of their actual monthly values. Table 3 and figure 5 show the results. Figure 5 clearly shows that a value of the master period between 273 and 275 months is definitely indicated. I have preferred 273 months rather than 275 months because it is an integral multiple of the strong periods 7, 13, 39, and 91 months. It cannot be much more than 1/3 percent from the true master period. TABLE 3.—Percentage amplitudes of proposed periods Period Ranges Months Percent 2752 \NO5.7. STOS:4| O25" 100.7. 166.9) 00.3 \, 07.3.1, 107.0, Gk.0U pon 9.4 27 273.0 05:7. 95.8 \ 03.4... 06.1, 00.3) 102.0, 103.7 108.0). 104:8 {tOL.1, (34.6 27 275.0 109.8 102.4 103.3 99.3 95.4 929 962 97.6 988 104.5 16.9 27 277.0 94.6 104.4. 106.2. 10T.3 105.8 105.5 (946.. 97.5 » 96.0. 03:3. 12.90 27 The subordinate periods—Of the 27 periods used in forecasting, 12 exceed 15-1/6 months in length. Owing to arrangements used to treat changes of phase, which will be described, 42 tabulations for each city are made of these 12 periodicities. Almost without excep- tion the curves representing these 42 tables betray overriding har- monics of the period in question, from two to eight in number. These overriders must be evaluated and eliminated before the period in question stands free. I show in table 4 and figure 6 the treatment of one only of the four tables representing the 39-month period in precipitation at Helena, Mont. Eight tabulations of successive runs of this period over the interval of years 1891 to 1917 give the mean values and average de- viations from the mean in percentages of normal precipitation. Then five harmonics of 39 months are successively removed, yielding the smooth-curve deviations from 100 percent given in column S, and its deviations from what remains after the five removals of harmonics. In the final column of table 4, and the final smooth curve of figure 6, we see the real periodicity of 39 months. The average deviation from NO. 9 FORECAST OF U. S. PRECIPITATION—ABBOT II curve a is 29.6 percent, and that from curve 0 is 2.1 percent. The reduction of 93 percent in deviation is due to removing exact har- monics of 39 months. Overriding periods—As another example I quote from footnote I, g, cited above, showing figure 4 of that reference (here figure 7). T NORMAL PRECIPITATION AMPLITUDE yp PERCEN Fic. 5.—Demonstration of 273-month master period in weather. From the mean of 16 repetitions of the periodicity of 45-1/2 months in Natural Bridge precipitation, the true 45-1/2-month period is cleared of four overriding harmonics.* The reader will note what similarity to true sine curves is attained in both the above examples, 4 Refer also to the clearing of overriders from the period of 68-1/4 months at St. Louis. Note 1, g, figure 3. 12 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 when overriding harmonics are computed and removed. From the examples given (out of about 10,000 cases available in my files) the following 10 exact harmonics of 273 months are exposed as follows: 1/4, 1/7, 1/8, 1/12, 1/14, 1/21, 1/28, 1/35, 1/49, 1/56. I 3 5 13 17 33 37 Fic. 6.—Helena, Mont. Thirty-nine-month period in precipitation as cleared of over- riding subordinate integrally related periods. Original tabulation, a; cleared curve, b, with smoothed curve above. Note approximate sine form. Range, 27 percent of nor- mal precipitation. While most removals of harmonic riders are done to clear periods exceeding 15-1/6 months, many curves representing periods between g-1/10 and 15-1/6 months required removal of harmonics of 1/2 or 1/3 of their length. An algebraic theorem affords a check on mistakes of computation when clearing half periods. Let a periodic curve be represented by equally spaced ordinates a,b,c .*.': k,l, mi, and, proceedimp tuttuer, 1, Orgy... ok yee: NO. 9 FORECAST OF U. S. PRECIPITATION—-ABBOT 13 The mean form of the supposed overriding period of one-half length is: Gari bite O Cita) sie rai 2 2 2 nea 2 2 2 Eo & 2 = 3 t When this half-length curve is written twice, and subtracted, we have: a—n BS ay c—p k—-sx I—y m—zZ ) ? Ya ar ,’ d 2 2 2 2 2 ? b and following that: z= | Qa ° | Ss > | a 8 a K= ~ t —m eee ) ) - 2 2 2 2 2 2 So the last half of the long curve, when cleared of the period of one- half of its length, is exactly like the first half, but with reversed signs. Grouping of periods——All weather influences caused by changes in solar rays are subject to lags. For instance, June and noonday are times of highest solar altitudes, but the warmest months and hours occur later. The lag is longer the longer the period of the solar radi- ation change. These lags are due to atmospheric conditions, and vary from locality to locality, from month to month, from times of great sunspot activity to quiet solar times, and as population and foresta- tion change. Hence, though the family of periods integrally related to 273 months proceeds with perfect regularity in measures of the solar constant, in weather the same family of periods is affected by changes of phase, depending on the locality, the population, the sun- spot frequency, and the time of the year. The periods are the same in weather that they are in solar radiation, but owing to complex atmospheric influences on the lags the weather phases are so altered from time to time that these periods are unrecognizable without a segregation of the data, governed by consideration of these modifying influences. It is not possible to anticipate and allow for these phase changes precisely. I content myself as follows: (a) The year divided: January to April; May to August; September to December. (b) Solar activity divided: Wolf numbers > 20; Wolf numbers < 20. (c) Secular time divided: first half of tabulated records; second half thereof. All these divisions of data hold for periods up to 15-1/6 months, or 15 groupings for these periods. The segregation according to the Wolf numbers holds from 18-1/5 months up to 39 months, but not the segregation for times of the year. 14 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 4HoKA papas L\ A ‘ ori ea Sa i A AL Ar cag oS Een ae [Aaa ko Sue ee eS ad al) M. babsad” Pct lh ot Nes cit aP cle | MeN a NU oes a Ha WES A Sei Lael etd Va So fh a tat} | pao a TION ey, oil aa EN sala 4a De hy EOMNIIN CE eo YS A" SE SH |) YI I aa on NE VAM CTT eNom pot eal iv Tg ay a ee alee YH pec eNIOAIMOD TS e eesleeer ee ee re yi Pe 0 eh Sa a si — A Ki gi Aid an ae Sa Wk NA a a a ff apap ERS a aan HPA A TE Pen Vee ad opal che /\ ena a TPR al al ede Fic. 7.—Natural Bridge, Ariz. Forty-five-and-one-half-month period in pre- cipitation cleared of overriding subordinate integrally related periods. Range reduced ninefold by clearing. NO. 9 FORECAST OF U. S. PRECIPITATION—-ABBOT 15 TABLE 4.—From three-month running means of precipitation, Helena, Mont. 39-month period = p Elimination of subordinate periods. Interval 1891-1917 Mean percentages of the normal. Original mean and departures after removing subordinate periods Average Original devia- mean of __ tions. 8deter- Percent X 4 mina- ° Removed periods Final tions normal (ooo, Smooth ~/8—S mean a ppt D2 PS iP se ue) 7a PLS 15) A cleared 102 30 o +2 -—9 —I3 —I7 —I2 —5 88 99 40 —8 —8 —I0 —II —II —13 +2 87 92 29 —24 —26 —18 —17 —I7 —I4 —3 86 82 35 —35 —32 —20 —I5 —I2 —I4 +2 86 82 29 —37 —27 —I5 —I5 —I7 —I4 —3 86 89 32 —16 —9 —9 —II —I5 —I3 —2 87 112 34 —2 o —9 —I3 —I3 —I3 0 87 III 31 +12 —2 —I12 —I3 —I3 —I2 —I 88 III 29 —6 —4 —I5 —I4 —II —I10 —I 90 98 38 —9 —II —13 —8 —I0 —8 —2 92 109 23 Onis Ee tele 3 —6 cao 94 109 30 —8 —12 o -—-4 —-4 —4 oO 96 112 40 —I —Ir -+I1 oO O —I +1 99 110 40 —9 —7 —7 -6 -—3 0 —3 100 107 18 Oo Gata Said +2 +1 102 III 19 +11 +9 +44 44 oO +4 +4 104 112 34 +16 +19 +8 +6 +46 +6 oO 106 119 26 +7 +17 +15 +11 +11 +7 3 107 97 34 —13 —6 +2 +1 +4 +9 —5 109 100 23 oO) Ay) 1-0) Ob 7 +9 ae 109 102 27 o —4 +8 +13 +9 +11 —2 III 116 20 +o +11 +11 +11 +411 +11 0 Ill 146 16 +25 +13 +14 +10 +10 +12 —2 112 152 20 +35 +28 +18 +17 +20 aig +7 113 156 33 +37 +33 +22 +23 +19 +13 +6 113 122) Jk 34 +17. +7 +5 +10 +10 +13 —3 113 117 21 +3 +5 +13 +13 +13 +12 +1 112 108 16 —I —I +11 +9 +12 +11 +1 III 123 27 +6 +4 +16 +12 +10 +9 +1 109 117 21 t+i1o +13 +13 +12 48 +8 0 108 109 28 o +10 +1 +2 +42 +4 —2 104 125 37 +8 +15 —3 +2 +2 +2 oO 102 115 26 Spe EA mez) ang a —I —3 99 128 32 ea 5 ft) 97 89 32 =O 7a hyd oO) 4 —4 0 96 90 33 —I0 —I2 o +1 +1 —6 +7 04 81 30 —I5 —24 —I2 -7 —7 —7 oO 93 106 50 —6 —10 —I0 —I0 —7 —9 +2 QI 123 30 +13 +3 —6 —8 —1I0 —I0 oO 90 Mean da 29.6 percent. Mean A 2.1 percent. Average deviation before clearance 29.6 percent. After clearance 2.1 percent. Norz.—Thus the removal of overriding harmonics reduces the average deviation by 93 percent. Of about 10,000 such removals of overriding harmonic periods, probably 4,000 gave fully as satisfactory end results as the 39-month curve at Helena did for the years 1891 to 1917. 16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 Hence for these longer periods there are about four divisions to a period. The secular time segregation holds beyond 39 months, two divisions each for four periods. The grouping just indicated leads to computing many tables for each station: Up to 15-1/6 months, 15 X 12 = 180 tables Thence to 39 months, 8X 4= 32 tables Thence to 91 months, 4X 2= 8 tables otal, i seiacacis ss eauyeasisice 220 tables Shifts of phases—The numerous groups used for the shortest 15 periods leads to tabulations with so few columns that the mean values of individual periods are of little weight. To remedy this defect, I assume that the forms and amplitudes of periods up to 15-1/6 months in length, and in the same grouping as regards Wolf numbers, will be similar, though in different phase relations. I therefore make superposed graphs of the six tables of one period for each of the two stated conditions of sunspot activity. From inspection checked nu- merically I am then able to shift the individual curves of the graphs to the same phase relations. Then I take a mean for all six tables and use that generalized mean in forecasting. But when using it in forecasting, I must shift back the generalized mean to the proper phase, as will appear by an example later. Figure 8 gives an example of these shiftings in phase. NOMENCLATURE, SYMBOLS, AND TIME As stated above, 27 periods, all aliquot parts of 273 months, are to be used in the forecasts. But, as just stated, these are used in several groups, depending on the length of the periods. Lags, depend- ing on atmospheric conditions, dictated tabulations of 12 independent groupings for the periods of shortest length, that is a1, D1, C3, @2, De, Co, as tabulated for the period of 9-1/10 months of SS>2o in tables 5 and 6. Besides these, there are six tables a’, b's, c's, a2, b's, c's, for SS<2o0. However, for periods above 15-1/6 months this extended grouping brings too few columns into the tables to be capable of yield- ing satisfactory mean values. Hence for periods 273/15 to 273/7, the distinction between months of the year is dropped, thus reducing the number of groupings from 12 to 4 for these 8 periods. For the re- maining 4 periods, 45-1/2 to gt months, the distinction SS>2o0 or SS<20 is also dropped, reducing their groupings to 2. So there are three different arrangements of assembly, as just explained (12X15) NO. 9 FORECAST OF U. S. PRECIPITATION—ABBOT 17 = 180+ (8X4) =32+(4x2)=8, making 220 separate tabulations in all. In tables of periods 1/18 to 1/63 of 273 months, there are many cases when the number of columns for @,, b1, ¢1, @2, D2, C2, and a’, b’1, ae ean | Mean Es if a AS o vee Pl cade i aa heebre eee” SE Sy : \ i a on ays deg eter he agate RIAs EON tpi Cree ia repent A= oe Cicer \ Fic. 8.—Sixfold grouping of periods to form generalized means. c’,, a2, b's, c’, are too few to give a trustworthy mean. Accordingly, as I stated above, I have made the assumption that in form and amplitude groups of SS>2o0 will be fairly similar, though of different phases, and in form and amplitude groups for SS<2o will also be 18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 fairly similar, though differing in phases. Making this assumption, I combine into one table a,b,c,d2b2c2, merely changing the phases to give best accord, and similarly I combine a’,b’;c’,a’,b’c’s into a single TABLE 5.—Grouping of six tables when SS > 20. Period of 9-1/10 months. Eastport, Me. First half of the records, 1891 to 1920 7 aia aay ee en Na 2 lg eee PR Pe gy gael am 64 92 55 62 130 158 79 84 90. 92 87 71 47 56 92 164 127 QI 66 85 80 88 79) 54> OB S04 ERO Ce OZ 27" Eo 7e igo gG 08 103 62 98 179 156 Sr 320 69 95 109 108 103 SO) 106} Lr Bivc E10. 80 120 96 = 108 93 108 OI 78 E22 111 TOO 640133 2: 123)... 10s... TOT 108 51 Ja TAA) 20S 55 120 66 127 107 86 94 G0) 47 AEIO 88-5 70 134) 54 TOO Fel A 90 88 59 80 61 81 78 78 79 70 «4116 79 75 Teee clade ke Maced Mises: Recess Tehek- fede d odee soak ts 86 115 72 160 81 132 70 ~=+167 64 105 II4 120 67h 62 PTO rst 76 04 70 105 85 04 134 103 90 128 82." 175 88 102 86 04 164 66 102 126) 103 77 85 100 SOOO SOU 77) 75.) ETA 97) 80. 770 92 G5 ' 04 IIs. 93)) 94. 128 108. | 72/75 94 69 3865 04 82 83°°"926"" "530" ‘100 84 92 75 70 109 Baie tOO. | TIO. (56S 06 86 08 ZU 92 BO 57 AP 05) SE Ta) 178") 04. W203 97 75 78 1891 1894 1807 1903. 1906 1916 191 Means Sept. Oct. Oct. Nov. ov. Oct. Oct. C1 126). "67 80 06 123 Wg.) VATS 08 100 72 85 04 131 88 104 96 99 7it20)) 1OAy | 1148 95 87 105 105 68 E46. 130 FSH TOO ETO 115 104 So) 150 142) 143) IO TAO 123 117 Sa 147) 128) tI6) Ose 172 124 Bi ii, Zonas es) 80. OOD Onn TIO 95 89 63). \112 70 96 150 92 06 90 73 73>) LES Ae IOT 7134 55 92 table. I give samples of this simplification here in tables 5 and 6 for SS>20. Figure 9 shows the matter graphically. The final combinations of two sets of six tables each, with phases shifted to harmonize, is given in figure 9 and table 7, both from East- port data. NO. 9 FORECAST OF U. S. PRECIPITATION—ABBOT 19 TABLE 6.—Grouping of six tables when SS > 20. Period of 9-1/10 months. Eastport, Me. 1926 Aug. 63 136 145 150 92 85 70 70 1927 May 95 135 185 189 180 178 167 119 oI 1925 1928 1937 1940 1941 1947 1950 1956 Feb. F Mar. Apr. Jan. Feb. Feb. ar. apes. Adil 7G. AS Tae Tie. Tas 68) 72) (166: 7 so 126 9206) 129 G4, |, 462). (89) 63)). 50) "196 887 (a9 Go 166, (7 75. 68 | TOG 107 83). 109) 63) 426.) 54 3651107, 459 Of, 129 "74, S17)" Ok 134 162) ¥05 03 1201 G5, 128 03) 88" a4. “SS 1240 TIO) 102") oss e1OLN 508) 102) 193 122-87 —. 67-844 | 105° 69). (FT, TOL 1929 1930 1936 1939 1945 1946 1948 1949 1951 Aug. ay June July July May Aug. May Aug. 92) 55° 97 S72).400" 134, 9104: (90 “150 OS -95. 161) 7824 66. 83 0 GAR O77 ah rss S7ert07) 77.) 73h) WMO: (568-104). 54 130 VA SETOO) “IOT!” WGagets7, (80) 130) 104, 50 20) SS Oa TOSS e7ON 12) 0138) ln 200 TOS, Ate Ite, GhioEsO Fe Tez eT 5s. MES TOS Sh O04) Ol 127. nila lez Ter i TAS TOK, 05 0) Oly 1 50k) DO UTIOW eLTOM 1366 107, bony 76. 52. )79 166 145 ‘108! 13. 66 84 154 Mie itor, Dee Sek Be. Gee; Gee or tne. TION 972. S25.) FTA" S60 NEO) “ZO | FAC WISE Ore 76>-nIP—So4-456> 73 | S420 84 Oo S4: “Tee - 66° 950 GS as GE. c08, 965 $E2/7).03. OS 550) 7O 7A, SQu i TOR Oye 965093 54 Gooy 76) 62, S87" 88 "1Z0 Sa )a12. G2 102) 104. 025 «9 8r). 14r sas0 SI Os Ga 10s" "IS1. Tog) | Shiro eras oP BO ek! | TIOM BST) OS) GOR Io ae 7 OS BO. ETF... TAG), 04, ST 22 TAG 112 Second half of the records, 1925 to 1956 Means aa 84 86 86 104 112 103 97 95 1952 May 130 12I 131 99 105 81 95 110 136 1955 June 60 80 87 65 113 102 126 Means 89 86 87 107 107 106 103 The meaning of the symbols on figure 9 is as follows: ok, no shift. T , shift backward. ) , shift forward. Subscripts, number of months shifted. Means ba 97 105 116 115 108 108 101 102 20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 TasLe 7.—Phase adjustment. The 6-1/15-month period Division = Time before and after 1900. Category = Records when Wolf sunspot numbers Phase shifts indicated: Basis of forecast, over 1,000 monthly records smoothed by 3-month running means. Forecasts employ 27 periods all exact submultiples of 273 months. = 20. ok, 4.n, YN, drawn dotted below. Phases shift with changing atmospheric conditions, but periods remain, and are of the exact lengths found i solar variation. It requires 220 tables electronically computed to make a forecast for one station. a1 105 102 100 106 105 103 biok 95 95 94 92 103 114 CatTEecory 2 ASSEMBLY cifs 108 106 95 azok beok cook Mean 100 93 92 97 106 105 96 95 97 88 93 80 94 88 100 91 101 99 Fic. 9.—Phase shifts 100 97 92 97 101 105 6/593 99 A +1 —2 az. —I +2 +6 CaTEGoRY 1 ASSEMBLY as biyi ciok azok bef2 cook Mean ZZ 120 95 100 105 96 117 107. + 108 86 ~=100 gI 940) 110 98 110 86 100 90 92 100 95 — 92 97 102 87 84 88 gt — 105 gt 104 gI 90 93 93 = 110 98 103 102 94 97 IoI + 6|586 98 2 in sixfold grouping of periods. NO. Q FORECAST OF U. S. PRECIPITATION—-ABBOT Zi Times.—The growth of population, destruction of forests, multi- plying of oil engines, automobiles, and airplanes alter the properties of the atmosphere and thereby shift phases of periods. Hence, as stated above, I divide the thousand months of records into first and second halves and compute the phases and amplitudes within the two parts separately. TABLE 7a.—The sixfold groupings.* The 9-1/10-month period. Eastport, Me. Values in percentages of normal precipitation A. WoLF SUNSPOT NUMBERS BELOW 20 a4 b1 ok ci 4 a2 ok boY3 c2 ok = =+6 A 101 83 94 90 63 106 537 89 —9 88 88 oI IOI 66 107 547 90 —8 88 79 98 107 73 107 552 92 ae 75 82 81 108 78 100 509 88 —I0 97 97 93 110 80 106 583 97 —I 104 95 III II5 QI 110 626 104 6 106 III 112 105 109 122 685 114 16 106 107 109 118 98 116 654 109 II IOI 114 97 107 76 109 604 IOI 2 Mean 98 B. WoLF SUNSPOT NUMBERS EXCEED 20 a ok bil 3 car az ok bz ok cof2 = =+6 A 87 92 92 84 97 89 541 oo 10 88 98 98 86 06 86 552 92 —8 08 97 96 86 105 87 569 95 aad 108 105 105 98 116 107 639 106 6 108 105 II5 104 II5 107 654 109 9 108 102 123 112 108 106 659 110 10 04 100 124 103 108 100 629 105 5 90 92 95 97 101 98 573 96 4 79 94 96 95 102 99 565 94 —6 Mean 100 * The shifting of phases is indicated by arrows as in figure 9 and table 7. The accompany- ing subscripts indicate the number of months shifted up or down. Not only so, but considerable differences of amplitude between the two halves are sometimes found. As forecasts are for present and future time, weights, as 2/1, 3/1, or 4/1, are given to favor the second half when considerable differences in amplitude of periods between the two halves appear. It matters not whether the later amplitudes are the less or the greater, the larger weight is ascribed to amplitudes 22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 of the second half. If a backcast were to be made to long ago, the weights would of course.be reversed. At some chosen date all periods must be in the same phase and preferably in zero phase. I chose 1957-0 as this zero date. To insure that any particular period will be in zero phase with 1957-0 it is necessary to compute ahead from the start at about the year 1870. This may be done as follows. Take the period 8-3/11 months for example. From 1870 to 1957, 87 years, there are 1,044 months. About 126 periods of 8-3/11 months would cover this interval. But a date must be chosen which is an exact integral multiple of 8-3/11. The nearest is that which gives 121 periods in the interim. Multiplying, we find that 121 periods require 1,001 months, or 83 years 5 months. Sub- TABLE 8.—Repeated 8-3/11 months and round numbers I 8.2737 8 7 57.9089 8 2 16.5454 9 8 66.1816 8 3 24.8181 8 9 74.4543 8 4 33.0908 8 10 82.7270 9 5 41.3635 8 II 90.9997 8 6 49.6362 9 tracting these figures from 1957-0 we find 1873-7. Thus a suitable starting point is August 1873. But it was assumed that the record begins about 1870-0. If so, 43 months would be lost. One therefore counts backward from 1873-7 five periods, and therefore begins with March 1870. We now come to considering periods ending in fractions of a month. We may make tables of accumulation for them. Again using the period 8-3/11 months, table 8 results. For most of the periods of inexact months, tables to 91 months suffice. But for such as 12-9/22, 13-13/20, 24-9/11 and 27-3/10 the tables must be carried on to 273 months. RESULTS OF FORECASTS Having treated of most of the features of the method, the re- mainder of this paper will disclose the results of these forecasts of precipitation. As I have stated, I discovered discrepancies sometimes as great as 10 percent between the published monthly normals and new normals obtained by separating years when Wolf sunspot num- bers are respectively above and below 20. As my new normals may NO. Q FORECAST OF U. S. PRECIPITATION—ABBOT 23 be of value to other investigators of periodicity I first give in table 9 the two sets of normals for the 32 cities I have investigated. The cities are in alphabetical order. The months in the first column apply for all cities. Precipitation is given in inches. Columns A and B give monthly normals for times when Wolf sunspot numbers are respectively Jess and more than 20. Departures; observation minus forecast 1950-1958.—There are 20 cities showing (1950-1958) departures in level of 4 percent or more from the values given in table 9. This is to be expected. One could not suppose the mean precipitation, 1950-1958, would be identical with the average precipitation, 1870-1958. Table 9a gives all the cities where such differences of 4 percent or more occurred. When I come to give tables and maps of forecasts, 1959-1967, I shall not use table ga to correct the maps, but shall quote the results as they are determined from table 9. Persons interested may apply the values of A, table ga, as corrections in level to the forecasts, using them in reverse of the signs given in table ga. Sunspot effect on normals.—Lest readers think the differences between mean precipitation values attending high and low sunspot frequency are merely due to the sparsity of evidence, considering the irregularity of precipitation, I call attention to the numbers of months entering into the mean values of table 9. For nearly all of the stations approximately a thousand months participated. That indicates about 600 for high sunspot frequency, about 400 for the low. Dividing by 12, there were about 50 values per monthly mean for sunspots ex- ceeding 20 Wolf sunspot numbers, and about 33 per month for the low sunspot frequencies. Referring to table 9, the yearly sums show seven cities where sun- spot frequency makes no more than 1 percent difference in the totals. For seven other cities low sunspot activity brings more precipitation, with an average difference of 5 percent. For the remaining 18 cities precipitation averages 5-1/2 percent higher at high sunspot frequency. While the discovery and elimination of these differences by computing new normals was of importance in my forecasting, seasonal differ- ences made the elimination of the sunspot effect imperative. Thus at Salisbury, N. C., precipitation averages 17 percent higher with low Wolf numbers, January-April; 9 percent lower, May-August; and I1 percent higher, September-December, for Wolf numbers below 20 than for those above 20. Credibility of forecasts.—It is difficult to compress within the limits of a paper, aimed to be available at moderate price to all who desire VOL. 139 SMITHSONIAN MISCELLANEOUS COLLECTIONS o'oS 1°08 GV ea Te SeeOles Oe eee OF Stee. 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Lo Oe (Ot A of VE OS ae Oe Os SOW. Sts OS LBL es Oi) Vy, O;S) cis Tay Os = O17 ee —_ ‘uuay, ‘aT[LAqseN a Vv . tes eax Joquiss9q] JIqUIdAO NT * raqgopo Jaquis}das "* ysneény eens Ain[ eeeee aunt eeee ARW eree Judy -e° OIC Areniqo 4 * Arenuel Sire tea Jaquiao9q JaquisAoN * sraqopoO Jaquie}das ‘* 4snsny ee eee An eeeee sunt eens AeW eeee Judy eee eee Yoel eeeee Aren1qa.y eeeeee Avenue f 26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 it, the results and comments representing this project. Even with 32 stations, the United States is so vast in area and so varied in con- trasting conditions that with the fullest use of my results no adequate country-wide coverage of the expected precipitation to 1967 can be made. As stated above, confidence in the forecasts must depend largely on the fidelity with which the first half of the forecast, 1950- 1958, inclusive, fits the observed record. Table 10 presents in parallel columns for all 32 stations the monthly percentage departures of forecasts and observed records, 1950-1958, from the normals given in table 9. That readers may see from a graphical standpoint to what degree the forecasts represent the events, I present figure 10. It gives the march of forecasts and events from 1950 to 1958 for Cincinnati, one of the best, and Denver, a less favorable station. TABLE Qa.—Percentage departures (O-F) 1950-1958, from table 9 GHEY Festi ate fea e's oo Abilene Augusta Bismarck Charleston Cincinnati TA. aes claie atayele Hass —I2 —I7 —6 —Il +4 CIE Yair eciorers oieiale Detroit Eastport Helena Independence Little Rock Gi UA arora musics doe oce —4 +23 —II —I7 +4 Gaby ice vals se arta Natural Bridge Peoria Sacramento Salisbury Salt Lake Te NW sre etcintateieavere —7 —6 —4 —5 —7 CH ere a elec nid ais San Bernardino SantaFe St. Louis St. Paul Thomasville GN ie wana oniermss +10 —I17 —8 —II —9 Figure 10 shows for a more favorable and a less favorable station a graphic view of data taken from table Io. A glance at figure 10 shows for both cities an obvious similarity of the features of the forecasts and of the events for the majority of months covered. There are, to be sure, differences in amplitude of features observed and forecasted. In many cases the forecast, built on average conditions of about 1,000 previous months, hits the fea- tures found in the observed record from 1950 to 1958 on the exact months. But in the better station, as well as in the worse, there occur relative displacements of features common to both forecast and event. These displacements are rarely as great as 5 months for any station, but may extend through durations sometimes as great as several years before returning to agreement. Displacements of features.—Several years ago I published the ac- count of a forecast for 104 years of St. Louis precipitation, including a comparison with the observed records. I quote from my discussion * * Text continued on page 44. NO. Q FORECAST OF U. S. PRECIPITATION—-ABBOT 27 SRS Reeeees Sees HA SO EERTn Ko wt LA A Fae ee CICA HEIN. ACC UANT He “ATTEN RC CECT ASN a tots “ARECACEAE PT ie My a) gis Wiech Ny ella ale (0 a la a i HHA-R ET --L eee LaLa a ACEC CREEE EE FLAY LUT ET OG Be Ca SETA IP SEEEEPCECEP AVE eee al ele Walaa ti fi 1 ens a HeLa oele ee PN ol AS] 2 RPT Ea ee feet ttt ji SUNG a Wa dt ae lai ell 1952, ps7 Fic. 10—Comparison of forecasts and events, 1950- Le Upper curves, Denver, Ooi: lower curves, Cincinnati, Ohio. Forecasts, dotted lines; events, full curves. oi a Pe nS ag ECO THiS POPPE EF Percent NorRMAL Precipitation § = + ir nr al armel aa Sel LS eee f1— _ vir fer f+ ovr Z1r w— Z1r ght “28+ 41 “1 ci— 66 III oO1+ 6g £Z gz— 9. 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IzI gol vel gel 101 v6 62 OII cz1 €or 66 ool £6 to ey cv +6 Czr bri f+ zI— ty— or— 1Z— 16— €c+ €11+ zoI-+ ev or+ bri+ 4$-+- bi gh— €z+ 6z¢+ o¢+ z1I+ €11+ 16+ e+ oZ $6 QzI oS! gIl v6 ge 14 SzI 991 SSI 96 vel ZO0z £61 Z91 Cor 88 ZO grI 6rI gSI Cor £9 Lol 1Z1 €Z1 Sgr 6Z1 ZIl ) es es 88 Qf1 eS1 LY1 QZ ZII £s of £9 zol IvI— 02 ofI— £1 101+ zz Ccib-++ tor Sof+ 96f O8t+ géI ss— of oS+ ger 601+ 9goz 16+ ebz oI+ oI €1+ ri z— SOI ozi-+- ozz So+ zogl be+ FEI ZII— 0 vS+ Z&1 oviI-+ g1z QgI+ tz 9I+ 36 gz— 9S 9— 4 Z+ 16 191 £1 IfI 16 26 58 / aX L6 zSI PSI Ifl LZ ool £6 OOI ZII £o1 / ~ gs €or rg £3— c6— IlI— SzI— IlI— ZQ— 61I— Sr++ 9Z+ z+ er— eS— 1g— z+ SZ-+- L£L+- IzI+ 49+ ol— Se Vil CII 140) z6 £9 OOI IOI ecr gr €gi LSI £11 z6 SEI Cer 6£1 Lz1 gII Lh 06 +6 36 gor 69 29 06 €z1 SSI gil 43 44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 of discrepancies from pages 2-3 of my paper cited in footnote 1, d, above: 8. Of 100 years of St. Louis precipitation forecasted, 70 seem fairly satisfactory and yield high correlation coefficients with the events. The failure of the other 30 is reasonably explained. 9. As shown by Dr. W. J. Humphreys in his “Physics of the Air,” figure 227, great volcanic eruptions, which throw high columns of vapor and dust, pro- foundly modify weather. He cites the first four cases in the following list [here my table 11], and I add several more. TABLE 11.—Great atmospheric disturbing causes Approximate dates Volcanic eruptions TOHOMeeials Gale er aciale levelalote Cotopaxi and others. PGSG= TOO: oes: a ers ovale Krakatoa and others. TOOT=1OO4) oh iat- ers eeciets Pele, Santa Maria, Colima, and others. LOUD Hye epe sree este steta ciate Katmai. TOZATANAILOZB xenon Many great eruptions. TOZOM pe rheres ae se cic Great eruptions. TOAT OU eee eye par eNotes Niuafoo Island. 10. Of 30 unsatisfactory years, in 100 years of synthesis of St. Louis precipi- tation, these lie in groups as follows: 1854 first half; 1856 to 1860; 1887 to 1889; 1900; 1901; 1905 to 1907; 1912 last half; 1913 first half; 1915 to 1917; 1020; 1923 to 1926; 1930; 1940 to 1950. It will be seen that many of these unsatis- factory intervals fall either soon after tremendous volcanic eruptions occurred or there was tremendous use of explosives in war or explosions of atomic bombs. As has been pointed out, atmospheric changes alter the lags in the weather effects of all solar impulses, and of course unequal periods have unequal lags. These unusual atmospheric disturbances may very well have mixed up the timing of terrestrial responses to the 23 periods so as to cause the events to differ from the predictions. At some future time it may be possible to connect theoretically the displacements found in my forecasts with causes producing atmos- pheric alterations of importance in weather. As yet I have been unable to name with certainty causes operative to produce these occa- sional displacements. For the practical inquiries of farmers, however, it is of importance to estimate the magnitude of forecasting error rather than the cause attending such discrepancies. As a step toward that, I cite the case of Spokane, Wash., figure 11. A computation made in 1957 derived a “correlation coefficient” of 59+5 percent over the interval March 1950 through October 1956 between forecast and event in Spokane precipitation. In simple lan- guage this means that my forecast represented the observed precipi- tation 59 percent perfectly for almost 7 years. 45 ABBOT S. PRECIPITATION FORECAST OF U. NO. 9 *S9AIND Po}JOp ‘sjudAa { sSaAInd [[NF ‘sjseovIOY “D *q ‘UO}SuIYse AA ‘SAAIND JOMOT f*yseEAA ‘oUeYOdS ‘saasnd Jadd~, “gS61-OS61 ‘syuaAa pue s}sedaIOF Jo UOSIIedUIOQ—TI “OIA a3Ad2SaC} Ntvzj 40 SHINO ByOW 2% OOO! 49 sqgaoosay NO S1S3y NoO1L9I10d3aaq S,HINO\|W/ wy7 ° SNWA\J PNINNNA HINO S fF WAON Wous SIUNLavd3g LNIIAAg 46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 When records through 1958 became available two considerable dis- crepancies between forecast and event were noted. In the months January and March 1950 heavy precipitation (over twice the normal even in 3-month running means) raised the February observed curve far above the predicted curve. Both curves, as has been said, are smoothed by using 3-month running means in all computations, hence the February effect. Not until April did the two curves come close together. Yet there was a difference of only 6 percent of normal precipitation between the averages of their heights, January-April, 1950. Beginning October 1956, and extending through August 1957, there was a shift of 5 months, leaving the predicted curve in the rear, and exposing opposed high and low values of the prediction and event. When the two curves were averaged over this interval of 11 months, the predicted curve was 116 percent of the normal and the observed curve 96 percent of the normal. To sum up: At Spokane, in the 9-year interval, my forecast gives for over 7 years a correlation with observations of 59 percent. Two intervals of marked discrepancy occurred. The first, of 4 months, culminating with February 1950, was obviously caused by extraordi- nary precipitation in two almost adjacent months. It produced a difference of only 6 percent between the averages over these 4 months. The second discrepancy, extending 11 months, was of unknown cause. It involved a 5-month shift of phases and produced 20 percent dif- ference between forecast and event in average precipitation over those II months. Having set forth those discrepancies I remark that this is in the infancy of my method of forecasting, before any help has come to me from theoretical meteorologists. It may be that some of them will discover the causes of occasional displacements of features be- tween forecast and event. If so, it may reduce error of forecasts greatly. Then, too, my method assumes that the average behavior of periods in weather in a thousand months that are past will be followed in the months to come. It perforce neglects changed conditions which may arise from unpredictable storms, volcanoes, or even from man’s interposition, as from forest destruction, invention of new powerful devices, wars and the like. Even a minor atmospheric change may alter the time of a feature in precipitation by a month. All these factors tend to lower the coefficient of correlation. The 273-month period in weather features——It will have occurred to some readers that if one were backcasting from April 1927 or from July 1904 he would employ the same tabular data that I have used NO. 9 FORECAST OF U. S. PRECIPITATION—ABBOT 47 in forecasting from January 1950. Hence one might infer that the precipitation following these earlier dates should parallel that follow- ing January 1950. There is indeed a partial similarity, as I pointed out many years ago, between the march of weather at successive intervals of 273 months. But the correspondence is very imperfect. This appears in figure 2, where the precipitation at Nashville following July 1904 is compared to that following January 1950. However, I call attention to the close agreement of the two curves for the last three years of the comparison. I have computed for several cities, including Nash- ville, the coefficients of correlation of the observed precipitation fol- lowing April 1927 and July 1904, and compared with the forecast made to follow January 1950. These coefficients have fallen between 18 and 22 percent, while, as stated in my foreword, the correlation following January 1950 ranges from 52 to 59 percent. This difference is easily explained. Over 40 percent of perfect 100 percent correlation is unpredictable as yet. There are several causes. (a) There is occasional unusual precipitation, as occurred in January and March 1950 at Spokane. (b) There are displacements of fea- tures as yet unexplainable. (c) The graphs I have published show large discrepancies in amplitude between forecast and event of ob- viously identical features. (d) Unpredictable events occur to alter weather from the averages of 1,000 months. In the march of precipitation from April 1927 and from July 1904, the vicissitudes of the later years up to January 1950 cannot have affected the observed precipitation of the earlier times as they have done that following 1950. As such vicissitudes account for 40 per- cent and more in coefficients of correlation, the tabulation suited to January 1950 can only roughly forecast what follows these earlier dates. FORECASTS, 1959 TO 1967 Table 12 gives for 32 stations for the interval 1959-1967 the ex- pected monthly mean percentages of the normal precipitation tabu- lated in table 9. The reader will recall that all forecasts are made from 3-month running means taken from published monthly mean values, and expressed in percentages of the normal values of table 9. Expressing these forecasts in a more usable form, table 13 gives average percentages of the normal for the intervals January-April, May-August, September-December, of each year, 1959 to 1967, inclusive. VOL. 139 SMITHSONIAN MISCELLANEOUS COLLECTIONS G+ $+ Open gh+ b+ 9961 1961 pi+ e+ pS— 1€+ v6— + Si+ cl—- v— oSs— 6 — So se— 6z+ 6I— 19-- €c— Zy+ e£e— os+ I — €r+ €e-+ 1+ 2+ $96I og6r De Ee €+ 6r+ 9o4- 2 Oo-- co— O03 = se oI+ zI— lZz+ £€S-+- era scr “E+ 39+ bee) 58 6z— 6f-+ to6r 66x ‘OOD ‘1aAUueqd Teo 6Z+ €— vl+ ce— 996r r96r tit+ tbh— ZI— += Iv— f+ Sze-+- eve; = ¥O=- Of-- _ &2-- VGa- = east Baap ies 6Z-- ~2— POs Ve 12-99 lz+ 0o9— $961 og6r c+ 7z— 4Z£=-- 1%— 1+ r= ev—= —of-- a= 0a €c— 6¢+ vS— SE gy— vi— Te A00— Sze— sr— ie Qe =Sis- $96r =66S6r Enea orto ‘IyeuUToUID Ze+ “E+ QS-- O1-+ 49+ S1— 9968 I961 o9+ Le— 7st+ gr— 12+ Sy— OI-=>-Si-- Z£— 1+ SI— oz+ t= “go-- of Sb z+ b+ 1f-- de-- OI-+ I1+ $96r og6r vit 6+ oO 1+ b— Se+ ¢— i+ €c-+- €&-+ w+ ce+ Ze £9 4 9§— Ze— z~— 2— I+ 6z— ¢— tv— yz— 2S— voor 661 ee ‘O'S ‘uojsalieqa 6 a[qeL ‘q pue Y suWNjod Woy soinjiedsp sSejuIdINg a fee g— 4+ e+ ¢1— 9961 1961 £1-- <3 — or+ 9 — VO=vi=t 9 — OI— te+ Lz— 1+ SI— Z+ e+ 1I— 2 — O¢— ~62-- gze+ €& — I1v+ = ge+ gI+ I — $96r og6r C= als oz+ ozc+ “Z1-- ~ LZy-- QZ— ~2c-|- pS— y+ 99- 2+ £e— 9c— z+ So— SOnr a= fo+ ze— QZ+ 9f— w+ 2+ bo6r 86561 OT dN yp1eUsig Og TI=F of gf A (0, 996% 1961 €+ m+ 4— i1z2— ZE— 6¢— Oo 0g 61— cb— Z+ zeb— ge- bve— op+ 24+ w+ v— zZt 2@— gr+ = =FI— ov+ oO $961 0961 z— 9 — oz— f+ GI" "892-|= gf— gt+ vVI— ZiI+ e+ 61+ Si-- ge-- ¢+ 12+ cz— viI— fc— oF — lz— vhr— SOLS $961 6561 ee "ep ‘ejsnany 41+ 6— oS y-+- ce+ 996% I961 4i+ 61+ IlI-+ giI-+ Zl ~SE-- Zo-+- oS-+- Zz-+- 9S-- Iz+ 99+ Si 50s iz— 61-- Eos SLea. Soa eae ZI viI— tO SQ6I o9g61 te+ of-+ 8 te <984- €1— o6ob+ g— 6r+ ZE— 6 + oI— I— 4s— ov— Ii— _—- 1Z— vo zQ— 990 Se= Iv 2 — w+ LIi— bO6I 6f6r —— “2310 ‘Aueqry uoynji1gi9a4g¢ JSVIaAOJ—TI AVL ot csIi+ +¥— ve+ c+ 9961 I96r i+ 72+ ozc+ e+ 6f+- 6 — fc+ o7c— €— ti2e+ I— Z£+ Qi se 81s i a il oc+ 61+ e+ ve+ £-+ 92-+- 2— 9giI+ $961 og6r ied ae eee 4Ir 9+ gi— £ + z— 6 — 42— GS + vb— e+ ge— —s: TI— ZI— + gI— ¢+ 1&— ai—— te £- oc— bo6r 8666561 en ae ACN ueqly gE+ 98+ 99+ b+ o£ ev+ 9961 I961 61+ 9gI+ 61— or+ zvy— o9g+ 6rb— 9S+ e+ ¢+ gofe+ z2— cs+ oI— gh+ oO goI+ oz— gor-+ 9€— 6g-+ Si— e+ +4 — $96r og6r mee eae Z2— oc— 6S— €z— Zo— 6¢— Zs— 6b— I— 6+ 6S-+ S — O0-ee Sha Zy+ ce— e¢+ Z4— b+ 7z— iv— ti-+ b96r 6S6r J *xoL “aualiqv scinces “SBN e@eeveee ‘qoJ eeeeeee ‘uel eeeeeece ‘29q eeeeee “AON eeereeee ‘po eeeeee *ydaS sq}UOyy 49 FORECAST OF U. S. PRECIPITATION—ABBOT NO. 9 zI+ zi+ Bt be+ of-+ ze+ €1-+ Bort Sis zI— Cc1+- ge-+ 1961 99+ bl+- 1v+ re— oe 6z— Ib— oI— oo be-+- oI— II— of— zZI— f1— cz— c= £961 Ii-— cz— re— 61— zl— 16— 6£— yl— ¢l— e961 re— 11+ 6 + €z+ Zz+ zb+- tear gI-+ gé+ Or oe Go 1h 95— fgo— £961 ce— II— or— — Zz— ios ZI— gi+ Zz+ gt+ Sec cse+ $Z+ 99+ Z4¢-+- £961 gI-+- vi— Lz— € + vI— y+ fe-+- £ + II— to Sr— 2961 1z— gI— git zs+ wy+ zs+ Si 8 —_— f9+ bh Se git L£=— oI— 2961 C=: Soci 9f— ss— 2961 Zz+ bi-+ ge-+ gi+ ce-+ of-++ Zz+ Z1+ o1+ bose ae Vey L961 Scc+ zc+ Or fo oI— zi+ 1v+ 1b-+- Soe Ib— Sie eo— £961 gI+ Ib-+- €c-+- 11+ (panuijzuo?) I- €+ Sce+ ce+ Si+ bi-+ QF Tost gi— O-- £961 €1+ 2 = bi— zo— 3e— ZI— Qe g1+ rp — €c— re— 61— 61— of-+ gi+ Z1-+ Ic+ £961 2961 e1+ or-+ 6z-+- oc— 2961 61+ f1-+- €S-- gb+- o1-+- or bi+ 7— gt— gr— 6z— 61+ 61— Vase €1-+ o1— Ii-_ £961 61+ Z1+ z+ L+ eat ort Et vi-+ Le-+ a oi1+ zi+ 6 + be+ gz+- o1-++ Ic+ c¢— gz— 2961 oar or— 8i— zI— gI— Or SiS Zi+ LI— SF c1— ZI— 6 —F Vick £961 oI-+ €1-+ 61+ ci+ II— a — £1— Woo Lz— 11+ Si+ bet vi-+ obv-++ €r-+ Z1+ 2961 61+ Sp w— va oI+ 17+ feet c1— y— 61+ 1€-+- 63+ 6¢-+ £961 ae LI— be+ Si+ 8e— Sze— of— c= Sc+- £¢+ 2901 1g+ gI-+ eeeeee eee eeee -99q7 + “AON pO "yd2S ‘any inf oun Kew “Idy "IVI ‘Pa ‘uel ‘20 "AON pO *yd9S ‘sny An oun Ae “Idy ‘ILI Fees gout VOL. 139 SMITHSONIAN MISCELLANEOUS COLLECTIONS 50 g€+ 9+ Bele ea OV-— 2S-F 9961 I961 I— gt Sc Ol— tc—) Qi Go Gear ee Ce e+ + + Olea {poeta le cet+ = te+ 69+ 1+ Oar 19ea 09-- vi— gt of— $96r og6r 0) ie po aes: Oe hea Of=— 0451 te==- <20-- Bien em Ovaie Ol Ons Cte a Ota to ea een oa Zoo Le ic Ze bo6r 6561 rs "ely ‘£1903 UOT 19—— seo gr— f+ AEE) oe z— o-+ 9961 1961 ot Oo el to Oe 99+ 91+ dea 6Qeat € — c+ fci— v— LZ— gi— Te a fea Gr teal ae — QI Zn $96r og61 te+ tvi— £4 te+ es Sv ~— se+ Soop. Gar Bie Olle VI— 9c-- css— gI-+ of— 9 — Ol Ol ae a= zi— 3s— b96r 86661 — RSENS ‘UOSIpe 6z+ € — Ze+ =SI— GRaj= GE— or-+ of— 9961 IQ6r €1-+ 61+ of— E+ vo— cet vo— LZy+ fg9— Ss + €y— ogi+ 4— 9+ QU Bea vo+ o1+ 69+ VI-+ O97 = Gua Wamp Ole $o6r 0961 sz— gI— gr— e+ og— 62+ Qh Vest OS —- ou Ole sera: 9e-- hI— e+ ZI— Lo-- ve— or-- o1— ¢+ tvi+ Sr bo6r =: OS OT a “2V ‘90y eT vi— gi-+ Se+ 1¢— e+ vI— 9961 I961 a 11+ 9+ Caan Ova of+ vr-+ ge+ 6€+ Ze-+- = 19-- of-+- 6€-- re+ or+ 6c+ 6+ Ol siec— oi+ ¢e— Lia $96% og6r es— 6 — 1~— 91+ Gea 00a: Zz+ ofZ+ ee+ 7+ vi GS-+ g+ or+ bi+ ci+ 6+ 9gI— €— os— te— Sl— orp— Ll— bo6r 6S6r ee ‘suey ‘gouapuadapuy panuyuoI—e1 ATAVY, y— vS+ Cleo. fc-- ZE=-- 9961 I96r ~¢— oi+ ev+ 8 €1+ 6€-+- 09+ b€+ 69+ Obs elvae Of sect $o6r og6r 6+ oz— bi— e+ 2+ vS+ Sit gst Vil c ieee cO ate Z-+ oe+ Ly-+- €z2-- w+ 9gI— 6I— I19— es— zg— zs— 1f— 1v— tvi— to6r 8666S 6r . “‘qUOyW ‘eualaH 46— 9e-- Qo set Sy— S€+ CO oe 996I 1961 iit Ol 6vI— 3g — gr+ = 1I— vor+ 1So— gori+ zi— v6+ 69+ S6+ tvor+ 96+ gét Zo-- O04. e+ Wt OS4> *Zb-- $96I og6r Lyle lave of + + a2-- Ot zo+- SI— O24 ee esate G7 eee rae bz f9+ OC aia oie 6— z+ 6+ 65+ €-+ ght hoor §=66S OT sexay, ‘oseg IH OI o1-+ ti— oz+ ef— $i + 9961 IQ6I 6+ 6r— 6r+ vo— gy+ re— of+ £1— oi+ tv — b-— oc+ g— e+ 9I— VvI-- ef ON £9— 0z— OQ eee Ga GO VS — $96r og6r So) Og oI+ 7— ce+ g— f+ gI+ Ze+ obv+ €I- z+ Se+ o1+ OIF = Ors: y+ 6— emg es L— oz— oz~— 41— b961 OS6r ET oureyW ‘ya0d\seq I— Iv— of— re— Ce ee 4i+ SiI— 9961 I961 i+ € — of+ gc-+ get ght ie ee eet Sit nO tae oft Oe fe) or — Lice ge 625 ic — of— 2 — €e— ze+ $96r og6r gs— ss-+ 89——~ sha £S— 1+ €1— vet Z4I+ “Zi+ Z+ 1I— eSt+ €1— r+ g— pe+ 2z~— (A Se— ¢I— 6r— te— bO6r 6S6r uu, —Y “QoL ‘plorjaq eeeeeee “idy veers ey teeeeee cggg veseeee sues eeeoese ‘29 eeeeee “AON ereesee ‘po eeeeee *¥d9S eeesee “ony eoeeeee Ajn( eeoseee oun eeeeee Ae see iON eeeees “Ie eoeeeee ‘qoyy eevee eee ‘uel teseeee Q9q reese “AONT reeeees 199 reese adag eeeeee ‘sny eooeeee Ajnf oun eeevre AeW susevekened’ Jen Cy, teeeee pW eoeeeee ‘qa eeeoeeee ‘uel sqquOW 51 FORECAST OF U. S. PRECIPITATION—ABBOT NO. 9 ze+ Ze+ €S+ 1Z+ zb+ 1Z7— 7— vi— Os oI— 2961 oc+ gt+ oc-+- Sls zi+ t1i+ S9— Ly— 9 — gi+ gi 2961 €r+ ci+ ob-+ oz-+ ge+ £b— or1+ fz+ ci+ fet 61+ be+ z+ 6¢-++ Lz-+- 2961 Gra ci+ 1+ Ti+ aie [ie zI— or+ be-+ 6€-+- ve-+- ei1+ oz+ 2961 eet gz+ Scot oz+ ei1-+ 0z-++- ge+ Ib-+ s+ Z1+ 8e— 2961 gi-+ 1g-+-+ €1+ 8e— Lz— z+ €z+ Iz+ vo+ €z+ si— 61+ I—_— Done bi+ 2961 6 + 2s ci+ Teste 1I— zZg— gi-+ bi+ Sr+ to+ Zz+ 9e-F [o— SI 4o— 2961 Li coat bi+ ci+ (panuijuo2) Coa zf+ Zy+ 6r-+ 34° oz— b= ci+ 1g¢+ or+ Zi+ ze+ £961 zb+ 2o-+ ze Z1— eo Co: ve— gi Ls ve— fects 41+ Qeat be-+ 1c— est Tp 1+ £961 (0) oe gi— €c+ cp t1+ ce+ of-++ ce— Si— e1— Ve ov— 2961 oI— 1zZ-- 6 + (oe ots Tea 9 + 1Z7— or— Le— 6z— 9 — ge+ 1v-+ € + c1— 6z— 4961 61— Ip— g4— SQ++ ze-+ or1+ f9— [7— ysS— og— or— g1+ 1 + ve+ Zi-- 6z-+ 1I— ss— £961 ZQ— [9— ve— 61+ ge+ or+ be-+- 6r— or— 6r— ss— 2961 c1— be— 7e— v= ve— I—_ tact Gat 6z-+ 1g-+- oz+ Z— 0 Lz— osS— g5— ev— £061 9I— 61— cit v— Z-+- 21+ 0 A ee— 9+ Zy— oS+ of— 9+ Ol) se 6— 4£-— 6i= Ze Le a iu— €-+4 496% 2961 9S— zi+ gof&— zS-+ 1+ ei+ GO-- aia ooI-- 9g — zoi+ ti+ 61— ci— zi+ €c+ 4961 gz-+ of+ LZz-+ z1i+ 66-++ zg+ aa re— II— Gees ga Sp gz+ 8 _— be— 89-— sy gh— £961 ci— zI— zI— C= LE = i ZI— Z1I— WP os 2961 6z+ gi-+ Ze+ 0z-++ 9z+- gz+ eeereee 20, eeeses "AON erreree ‘po eoeeee *yd9S eeeeee ‘sny eee eeee Ayn[ eeeere oun eeeeee ACW oioteiele:ss eet eereee “Ie e@eeeeeve Qu eoeveeee ‘uel eeeeee “any eeeeeee An VOL. 139 SMITHSONIAN MISCELLANEOUS COLLECTIONS 52 aA —a —_- — — 4 pada -_= oI+ 6 — Zt e+ gp+ 4— QI— Sz-+ ¢— ort gI— gf+ gI+ zw— 4£— e+ zo+ ¢i— 6€-+- SZ+ i6-- 2— Of-+- 97+ gi— + =$Z+ 4+ 1€+ vi+ 61— bi-- 12+ 99+ SI— o6+ 11+ 99+ Le— zI+ € — gz— 39+ Sy+ zit Z— z— 6S+ 6€-+ z+ oO 411+ vii+ gv+ gz— 6+ i+ ez— $+ gI+ Zi+ 9I+ gg— 6r-+ c+ 9961 I961 9961 I961 996% 1961 9961 I961 996I I961 996r I961 9961 I961 996r 196% ZI— + 9f— £6+ 66+ Ly+- 3t— z+ s+ Si— 4+ Zze+ 0 ¢— — oO Z— z— tf— IZI-+ 06+ 41+ sz— ie—- -“Vi—= Set 200-1 zo— I+ OI— of€+ gf— gI— gz— vI— L£Z+ 9+ 1I-+- e+ e— Si Oe 29-1 v6— E-+- Zs— £0-- f— I— f€+ 3-— 8— 92+ t-+ Sbt-+ €+ ov— fz— gI+ f9— St-- zl— 6¢-+ 9—- gt See Oe fe= 2e— 2 -- ~S9-+- ve-+- So— 61— ¢£— os— 41-1 €Z— -62-+ b+ z+ g+ re— fo == 492+ PSE ape— ig) le Zt— <0 gg—- iI— iit 0 Seo Sia S45 — 1+ 11+ Iz+ 93c— vI— gI— v— Q- 4L— ¢v-+ gI- vI— Sea eae ia — =F Sia 16.-F oh— vI— QI ae gI— gI+ e— L/¢— fZe+ zi— gr-- 9 + Ege cee Rea =8e-/ C-- e— Lie ee oO ate of or— zl+ ve— HC am WS zo+ 62+ 6 1 — 6-4. 6c-- oO gi+ Ze Oz-|- €9-- 65— SE-- =O. of-+- o1-+ g1i+ &¢+ Si Si ~— I€-+ + tvI— ge+ Sze+ w+ €z— 6-+- 1+ 41-- Ti+ Ori-+ 9obv+ I— g— of— 1+ f1-+- 6e— S1-- fe= fi-- —& -- g@+ z+ $96r 0961 S$Q6I og61 $96Ir og6r $96I o9g6r [96% og6r $96r og6r SO6r 0961 $96I og6r et pvt ezi+ ¢o+ e+ e1+ a 0) oit+ P— ov-+- =1b— Iiz— O1-+- fzc— b+ y— €2+ 6s+ 6+ 4— oI+ cse— go— zI— Ss 6 + ss+ + S1-- S€-- b— gf+ “Zz— ¢S+ 2+ e+ ¢— 61+ 2+ 9 — 61- +F¥— 1+ oz+ gb+ =€s+ Iz— €9+ yE— 19+ goI— rr— 6— 9+ vi— zI— I+ 12— 6I— o6of+ zo+ SP wzo— bl+ SI— 0Z+ f6— £6— gi— 66+ vyr— e2h— ve+ 12— zi+ 61+ 0 Gre sf— oz+ oO 1€-++ gr— tvzi— f1— 3+ Srp— £so— ce+ oz+ €€-+- fe+ b+ lz— gI+ 9 — zi— 2+ ¥g— of— oI— _-95— oz+ 6r— €r+ of+ co+ f+ g9— oI-+ 6z+ P+ 6z— vze+ gz— 6b— €1— 7— ¢— tve— re+ 6c-+ Q— e+ vi+ Zy+ z— se— zw— 12— Se=- -se— of— Lr— Sgo— zI— Iz+ 12— g+ si+ 46+ 9b+ cc— ch— Iz— £e— ob+ + — gz— ef — Z4Z— LI— 9+ tr 6— 9gz+ 46+ z+ 6b— + og— 6— 72— vS+ gz+ 6s— 0oz— zS— 7+ vi+ ob— oO $2+5 83+ sr— SZ— ~7— te— £1— gg-+ 1S-++ zeZ— 6+ “i+ «1+ “2+ ¢e— + 4— vo+ Sh— e+ + — bo6r §=66S6r bo6r 86661 boor 6961 boor 6S6r to6r 6961 bo6r 86961 bo6r 6f6r to6r 661 ‘O(N “yreO ‘A'N “SST “HII “Iq9N ‘wy “‘auay, panuyjuor—el AAV], e's Sa — a | fa ~ G's ——_ Pine —- eeeeee eeeeee eoeeree ee eeee "IeW "qq "Ie * “qoq . ‘uel sqUO WY 53 ABBOT S. PRECIPITATION FORECAST OF U. NO. 9 11+ 61+ €z+ 61+ ¢— oI— Oe ce of-+- Sel ce+ gs+ 4961 c¢e+ €z-+ 6z-++ ¥g— 95— Z2s— Q7— gi— - fe-+ Zz-+- Oi 61I— gt— Seis bi+ Ou II— ceo £961 zz+ ¢ + 8e— er— ee— 1Z7— ¢i— 61— gf — — 9h— 2961 gI— zS-+ og+ gh+ gz+ Ze+ b+ yy— gt— ve— vi— 1g— c1— er 41+ vE-+- 62Z+ SZ 4961 g-+ 92+ v2— v6— ZI— 1yi— zhi— 76+ g5+ se— 88— gzI— 66— 2L— ¢s— 1g— LE-- 9f+ Ta-|- £061 99+ er-+- zs-+ e+ IZ— ce— g — z6— So— ceo yfZ+ fo+ 2961 gz-+ 9— Zo— o£— of— ¢{— ¢o— Pest Ip+ Se+ €e+ oI— zI— 4 ae ¢s— 1S— be— oI— 4961 8 — gI-+- gi+ zb— II— oils c= Fi— 6¢-+- oS+ or-+ cai (ia ig 1z+ co £061 oI— acl 11+ CiL— 6z— 8e— gi— oI+ Ons 2961 9— gi+ 2+ 1p+ oI— ZI— oI— oz— co— Zo g — €1-+ eo— ce— Ly— zI— c+ 2961 bi-+ ce— ei— goI— €zI— gl— 02— oI+ 9e+ 6z-++ 2961 ze+ gI-+ eb Lo+ 6¢-++ f1— Le— (panuyuor) 9 — 6z-+ ob+ ge-+ o¢+ Si+ ee+ ey+ os$-++ ze+ Cr+ zs-+ 4961 TZ ri+ S-= 61+ be+ oz-+ 6e+ 6¢-+ fe+ OF=- cn e+ vi— lz— ceo— ois De se Cre f96r 41+ ei 6z— 2v— 9z2— 61I— Tot eos 9 — Beale 1v+ 2961 vi+ Bok cc— ov— gi— Sz— ¢ — 6z-+ ge+ ee o1+ OF 1¢— 8e— ¢ + er of-+ e+ e+ 2961 Lz— of-+ ze+ I—_— € + zeo— 61+ gb— Ze— co— ¢1-+ zo+ gf-+ ve— ¢f— 6g— so— oI— £961 gs+ ce Leet Di 8e— be— — Zz+ z—_— 1Z7— 1s— 2961 c1— Lo o1+ 1¢-+ 2961 vI— one gist zI+ 8e— So— zeo+ Lo zs— act oz— “lees 61-++ Sce+ ge ci+ £961 ee Of c6— ¥g— 88— 61I— of — S9— 99— viI— 2961 of— oh— or+ gS-+ 84+ 98+ z+ 1Z-++ £9+ bS-+- bi-- fc+ orI+ 296r of-+- gz-+ Z1-+ oo €c— eee eee Gis.6 6 a) e)s. 2) eee eee eee eee VOL. 139 SMITHSONIAN MISCELLANEOUS COLLECTIONS 54 aes p= 20 9— e— Oe ee Oe Go eel 9961 I961 zI— oOI+ SI— € -+- or— bvy— 6+ 6s— Oba aa ee-- Vet Tea Ava gs+ 42+ ce+ z+ re+ f4i— Gece = Sse One ver: $961 og61 ert git Tee Vie te f= Seq: esa) 2 Zo+ gI— Z4i+ vi— co—- - Of-- zeZ+ Ie+ Oe ea 1f-- Qe— 9961 1961 ei €c— €e-- ame a Og ais 6zc— + OS arte Gc aaa eb ete o9+ ti+ 6v+ o1+ Veo) aes =f gc— vI— Bea = eal $o6r og61 Talo ce tr+ 0 10-5 (Ol cr+ 6I1— 6g€— £— €y— vi-+ cIi— —-¥ + [= oO -+- I+ e¢I+ gs— 4E+ CVeeei la 1 VB Glee too6r 8=66S61 — OR ‘smoy “3S papnjIu0J—zZI ATaV C= ea ia Ope Bee OR gz+ 92— St-+ ie-+ 9961 1961 e+ €-+ tof L+ Sipe eaeele Zit mI — ve— Sv-+ of— 7cS+ €Ss— zr+ vo— Zb-- vVE— ge+ OTs = Sie g9+ sé+ 09+ 91+ SQO6I ogor 2 OFF vo-+- 06-- ef = €gr a ey T= Oe 0 S05 cit GS + z+ 9f— glt+ 9e— 69-- 1S— z6+ of— 6Z+ 1m6— vOOr OS6r Sa - "qse MM ‘guexods CL AO 96— oc— gi teat le— t+ Oi hes 9961 I9Q61 62-}-— “SE zgt+ gh Sv-- 16— €z— zg+ QgI— Ib+ Le Le 4i+ ¢f— 6h+ Lg— sy eal g6-- SZ— gor Or SQ6r 0961 ob+ 92+ Vi== 102=- o7— £1-- VO—=— *2e-- WV eS See c= 2.55 €z— gz— eI— —st— git 9€— gy =e to-- 1-{- v96r 6S6r Se ae “HHO ‘ouIpieulag ueS Or. ih oe oer Z2— S$ — goI— c— O1I— 61— 9961 I9Q6I 14— o9f— 6&—" or — zw— ve— te— ce— i oc— €Z+ tv — io 9c— ob+ lz— 1z+ tr— gz— rs— ev— + ob— eS— ~— SQ6r O96! £e€— of-+ 66— 6S-+- SS— €€-+ Z— ob+ Ze+ Set 4y+ €1+ 61+ 41+ OI— SI— Le— ~— oI— Sz— t~e— ¢y— or— oS— b96r OS61 ees nena ‘xO ‘N ‘aq eyueS re wo 61— + ss— O 6S— i, Se 9961 I9Q6I oI+ i+ 1S-- 62-1 Vos St - €S+ of€+ O14: — GVA Zt cet tr— 61+ ce— =ei1— 9+ gi— Ov+ f+ So gi-- SQ6I ogor e+ of€+ II— ¢b+ gi— 64-4 Ors Sach re+ b+ 6r+ iI + Se Or 11+ ¢e— e+ o6of— le—- +F¥— b961 © 6S 61 yen ‘aye’] aes eevee idy RIS Sse TET eeveeee ‘Qt eeeeevee ‘uel ‘20q ree ees AON teeeeee 9G eveeee *ydaS eeeeee “ony eeereee Ayne oun eeeeee ACTIN eee ‘Idy reese rey eeereees ‘qoyy eeerereee ‘uel . . . sq}uOy 55 ABBOT PRECIPITATION FORECAST OF U. S. 9 NO. bi— zi+ e¢o-+- zg+ vs co+ g — ze— or bi+ L961 61+ co— e1— ZI— 61— ze+ 9 — gI+ ¢o+- es-+ 1€+ a — br— 9 — 6z— re— 69-++ £961 fe -1= zi+ ce— or 6e— Le— of— €z+ gr+ zo 296r gI-+ Zo+ 9 — 8e— 2) — te— 9+ f9o— yS— 1b— eae 1 hs Ct €e-+- Ip+ 19+ vS+ 61+ L961 DUS £o— ga— £o— 6z— LE of— oz— 11+ al e1— e1+ Set Sie zI— a zI+ £961 9$-+ f9+ get Die ziI— 2961 ez— of— ae Ly— gf— ci— ¢+ 6e— cz— er+ gI-+- €c+ zI-+ I-_ oles lI— = £961 of— Fri— ¢f— f9— IbF— Mele Si-+ 2961 oS-+ ig— 61— eat 6z+ 1s-- oc Zi+ Iv-+- cr ZS++ Oc-+ or1-+ 6; Or oP oz~— 1g¢— 49061 gz-+ 09+ zo+ € =F zQ— 1g— oI— Iv+ 92+ 1S-+- 61+ Oar Le— 19— = TO £061 Sr— 1z— 8e— ZI— €I— OC cc— of— cI— Ibp— 296r cL gf+ er-+ 1¢+ zo+ Ze-+ Si+ ci+ 61— Zi+ €1+ ci+ 8e— Ze— Gat 9 + (ee 29061 0) 61+ zg+ 69— oz— biI— gt eet of Wp €96r 1s— 61+ Qe-F gb oh (0) tc 1s— f9— 1Q— 6z— 2Q6r 6-3 me QI Oats 9 == O=- cIi— c=—_ OF bi+ cet ge+ £+ 61+ cot acts Cla £961 ot oo Ze— Zi— ge bor CoA Zi 61+ 61+ c= £961 or1+ Sc+ Le-- Scc+ O0c— 2961 co— ol cet Zz+ i Sle eo+ ge opt pS-- ge+ Ze+ 41+ cc+ 7 — Sc— c= £961 I~-_— ae 9) gI-+ gI-+ Yoo ss— cc— 1s— Olias oI— ze+ One 6e+ 9e+ Si+ 6 =- fe+ £961 e+ be+ S =f ge of-+ Qe— LS— 89— 1f— 61— 296r ® cls 98+ £S+ 09-+- ev+ be+ br Z1+ erp OE ohe Sete Ch 1g¢— Ly— zb— 6e— ci— 4061 Scot Zy-+ s+ cL 6z— lo— br— oz~— II— oI+ oI— oz+- crt 11+ 8e— cz— £961 zi+ €c+ oi-+ Ze+ gl+ os+ eo+ 95— ¢/— bre— Or 61— 2961 €c+ r+ pe 1g— of— Fi— eooeeeee ‘Ie ‘Pu ‘uel ‘20 "AON pO *yd9S “ony Ang oun Ae eee “Idy oe “Ie . "qo.7 uel 20 “AON 0 *¥daS “ony Anf oun 56 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 At the end of text are 27 maps of the United States with the 32 cities as listed above, and each accompanied above the circle by a number identical with the appropriate number in the column headings in table 13. Below is the predicted departure from normal. Each group of three maps covers the three intervals per year of four months each named with table 13. Large areas of approximately equal departures from normal precipitation are clearly noticeable on the maps. These area similarities may aid farmers remote from the 32 cities to estimate the precipitation probable in their locations. APPLICATIONS Periods control long-range weather.—I have sought to present to meteorologists evidence of two important propositions. First, that there exists in weather a family of periods, all exact submultiples of 273 months. These periods are hidden from immediate recognition because their phases are shifted according to the state of the atmos- phere. When, however, the long monthly records are grouped and reduced with reference to time of the year, sunspot activity, and march of population, the family of periods is clearly disclosed with constant length, and with approximate sine-curve forms. Second, long-range precipitation is to nearly 60 percent governed by this family of periods. By evaluating the average forms and amplitudes of these periods from thousand-month records, precipita- tion and temperature may be forecasted for years in advance, with considerable approximation to the event. Whether these forecasts will appear to interested parties as trust- worthy guides to help in planning their future operations must depend on the agreement attained between forecasts and events, 1950-1958. I therefore prepared table 14 which gives for 32 cities the 4-month forecasts and observations, 1950-1958, and the differences in per- centages of normal precipitation, A, in the sense observed minus fore- casted. Their means are given disregarding signs. Agricultural requirements.—For agricultural purposes a foreknowl- edge of seasons rather than of individual months is most desired. Hence I give in table 14 4-month mean values computed from table ro. But it is the difference between forecast and event which would be the controlling factor in estimating the value of the forecasts.* The average differences, A (observed minus scale-corrected forecasts) are 5 As differences in level of observed precipitation, 1950-1958, from the averages of 1,000 months, are disclosed in table ga, I refer to that table for possible cor- rections of level which might be applied to values for some stations in table 14. NO. 9 FORECAST OF U. S. PRECIPITATION—ABBOT 57 entered at the bottom of the columns of A in table 14. These averages will be needful to the use of table 15 which is to follow. Assuming that the degree of success attained in the forecasts from 1950 through 1958 will be attained from 1959 through 1967, I have prepared table 15 from which the probable sizes and numbers of discrepancies between forecasts and events in 4-month mean values over the entire interval of 9 years, 1959-1967, may be estimated. Selected from table 14, four groups of cities, 25 in all, are tabulated in table 15. The first group of I1 cities have average 4-month mean discrepancies, 1950-1958, of about 20 percent between forecasts and events. The second group of six cities have mean 4-month discrepancies of about 26 percent, the third group of five cities, 30 percent, and the fourth group of three cities, 4o percent. All the percentages relate to normal precipitation given in table 9, with the scale correetions from table ga used in table 14. The six columns of table 15 give, respectively, the numbers of cases in table 14 when the discrepancies between forecast and event, 1950- 1958, are (a) less than one-fourth, (b) one-fourth to one-half, (c) one-half to one times, (d) one to one and one-half, (e) one and one- half to two, and (f) over two times the average discrepancy of the group. If the same degree of success is reached 1959-1967 as was reached 1950-1958, the interested person of a city in Group I would expect the numbers of discrepancies (O-F) among the 4-month means stated in the mean values at the bottom of the columns of table 15 to occur in the entire interval of 9 years with magnitudes in percent of the normals as stated at the top of the columns of the first group. If he were located at a city of Group 4, the percentages would be twice as large, because the numbers heading Group 4 are twice those head- ing Group I. But the numbers of cases would be the same. Stated numerically, a person residing where the mean departure of forecast from observation, given in table 14 for 4-month intervals from 1950 through 1958, was about 20 percent of normal precipita- tion, may expect the following numbers and magnitudes of departure from the forecast of 4-month means during the entire 9 years, 1959- 1967, given in table 14. Numbers of departures..... 4.6 4.5 6.1 6.0 2.8 3.0 Magnitudes in percent...... 0-5 6-10 II-20 21-30 31-40 >40 If he resided where the mean departure given in table 14 was greater, the numbers of departures as just given would be unchanged, but 7 + Text continued on page 67. VOL. 139 SMITHSONIAN MISCELLANEOUS COLLECTIONS 58 bi-+ oz+ e+ e+ ze— 9I+ zs— zi+ 6+ 6€+ vi+ gI— 1I— SI— Zo— og— e+ gI+ €r+ 17+ 1z— Sr gi+ ie 19+ “ojo ‘raAuoq €z+ es-+ LZ— b+ 61— g+ ae-t= — oc— €S-+ 6b— o= be+ zb— ogo ‘1yeuUToUIZD Z vi— IZ— of— 9+ ee— zI— 1I— of-+ Ze+ be— gis 1g€-+ zi+ Se+ 6f— MES: ‘uojso[ieyaD 9 2961-6561 uwounpidisacd jousou wosf Saanjangap pajaipa4q—E€l ATV], ‘% 92+ 6z-+ ci+ S-- ee— Cr+ Zi+ 1z+ Ze— ci+ oI— ci— oI— er+ Ze+ II— 82— cee s 9+ z1+ Si— O= 99+ 61— bi+ g9+ tz— e+ €z— c— oI— zi+ tp— of— tr— I1+ zI+ gi+ 61— Sc— c= €e-+ zo+ ze— 29 ‘ejsnony v UvoW YJUOW-INOY c1— 61— co— Iz+ 11+ ci+ £g-+- e+ ZI— e+ 6z— es+ 1I— 6I— Tt 61+ Sa: of-+- gh+ e+ Iv-+- LZe— ge— "3310 ‘Aueqiy £ £+ Sie ci— ‘A'N ‘Aueqry z or— Ze+ ez— ce+ gz+ Lp— bi+ £+ ZI— ie Zo+ oz— 9 8e— 7S ev+ 61I— ZI— ° cI— SI— "xOL, eusTiqy eevee “AMUSMOUSMOSMOSAMOAMOAMOAMOAMYO 2961 9961 S961 to61 £961 zg61 1961 0961 6S61 59 ABBOT FORECAST OF U. S. PRECIPITATION NO. 9 gr+ £1— oI— Si+ ge-+ Zo+ gt+ — er == gz+ z+ 1s— ‘ey ‘£19UI0ZUO TT gt Vote Iz+ 11+ oz+ gz-+ 6+ oO z+ 6g— z+ ee+ of— ‘SIM ‘UOSIPeIT SI SS ee 6= oO o=— Sr bh— Iz— 19-++ os— £+ ge+ ce— 6e— £¢-+- 61+ 1z— gI— 1€+ gI— Iz— ye+ 61+ I-P or1+ Z—- S— Vv ‘yooy 2107 VI ze+ oI+ cbr— ci+ €1-++ ete 6I— Z1+ Fi— Ge+ £1— 1v— ve— €+ osS— i= oz+- O=— ce Cr+ 61— 9+ Z¢-+- 95— “sue ‘gouepusdapuy 1 (panuruos) 9s+ z+ 0z+- fe+ f= 6= ge+ lo+ y— e+ ge+ 1y— get gt+ zI+ bbh— 1¢+ cz— zb— *qUOW!W ‘eua[eH zi 6s— oo1+ 1$+ £9+- gt+ p— 89— e+ o= cr— €1+ oy lz— zI— oz-+ £1— Zz+ or+ G2 €S+ of— “xo ‘ostd IF II 1g¢— oO 61+ 9i+ Z or— gi+ 6z— oI— 7+ £I1— aUIeyy ‘yaodyseq or gr— €€-++ ie Scc— of+- 6r— oa oe II— €z-+- bi+ 6z— ze+ 9+ z— ce+ y— bi— ° “Qo ‘yorqd 6 eee ee wens £961 9961 S961 Fo61 £961 zgo1 1961 0961 6S61 VOL. 139 SMITHSONIAN MISCELLANEOUS COLLECTIONS gi+ s+ cf+ Pate ge— or-+ c1— £+ vh+ 1I— vI— 61— 8+ ¢1— e+ 1I— 6c— 1v— of+ bi-+ 6= tz— €1— or+ ce+ Ze+ e— ‘O'N *Aangsies ve gt— vi— 1+ gli— £9+ 1Z+ zg+ 61+ zs+ gs— 98+ 69+ 1f¢— gt+ oi+ Soa zi+ “FHED ‘oyUIWIEIIES €z €z-+- zeo— ce— 9+ zS-+- 61+ 4I— Pe ger I+ be+ == ¢z— “A 'N ‘J9}SaOOY ze t1— lz+ 61— of-+ gI— cr+ cc— y+ og— Se+ 6+ gi— 9= Sin 41+ gi— e— b+ Z— S— gI— be+ br— 61+ oz+ of-+ Zze+ oI— oI+ SI— 91+ ziI— Z— I7— €1+ 6+ ze+ II— oO sI— ci+ 09+ gI— b+ Ly— Z1I— bi+ zI— L— c— zb— vi+ Ca of — “SSI, IIL ‘UOSqID) 210g ‘el1oag Iz oz pamyuor—El FIV L ci+ SI— 9z+ Se Si— €1+ gy Lz— oz+- le+ 41+ cts Se— z+ 6s— Iz+ oI— gi— I+ p— be+ 17+ 9+ 6+ £+ Cc+ o1+ “Iq2N ‘eqeuo 61 61+ p— e+ 9+ cv— 11+ ce— 89— 6v-++ z+ t+ 98+ y— cr— Ie+ 9s— ZQ— Ly— ~— Iz+ ¢s— 12+ 8+ ve— ZE+- gi c— ‘ZuYy ‘asplig [einjen gi gr+ ee ceceeee [a= cece 6z-++ oe Si+ ereee og+ ° ZI— ° c+ ° 61 a Iz+ ° II— °*° pa ee 6+ eee ¢o— ° Dia = oz-+ ° oI— * get 1 Q— ttre ip ce. s° p— crteeeeee 4f—° ‘uudL ‘ay[raqse N “I GMOUAMOMTMOSTMOSMOAMOAMOAKMOAAMDYO £961 9961 S961 to61 £961 zg61 1961 0961 6S61 61 FORECAST OF U. S. PRECIPITATION—ABBOT NO. 9 9+ ce+ ZI— ozc— oI— 61— Ze-+ oc+ L+ zi+ gh+ be+ 6+ ys— == gf — 17+ z+ II— cco— tc— gz+ I— ze-- viI— cc— Oa RTROe ze ge— 9+ ve-+ ch— I-- ve or— oS-++- lz+ o=— vo+ Oo oS e+ 3== Se— af ce+ Lo— ol LE+ se— ge S++ 9z— o1+ e+ "ex ‘OT [Asemoy T, 1f bi— zi+ oI— of— 8e— gI+ Le— SI— Zy-- 91+ be+- ats 9I— bi+ 6z-++ I— 9+ 12+ II— Oca If-+ Sz— “UUIPL “(neq 4S of gz-++ ge+ 1I— gf+- ge— gf+- 6f— aoe oI— fe bz— Ly— git+ 9+ yo— ge— 1z— Ib— Z1-+- 6b-+- 9+ 11+ f+ z+ 6— Pafe oI+ ow ‘sInoy 39 oz f+ o— oa e+ 8I— 11+ Zy+- er— Ze+ 19+ S1+ 6b-+- cc+ fe— SI— o1-+- gi-+- 9s— £+ oe: 9e— + ov+ €z-+- s9+ c1— 6b— “GSeM ‘gueyods gz Z€+- Ze+ SI— e+ gt+ 69— be— 6S-+- Sz— gr €z-- Sz— Ze— Z1+ 61+ c1+ 11+ £5-+ 8z+- cl— of— IZ~@— 1v— (0) o1+ se— ‘xOTW 'N ‘QT eyes le Tete €z+ b+ f1— rg— Sr— obv-+- 3 $Z+ II— 6e— ge+ ZI— €1+ zi+ ce+ of— SI— oI-+ gI— 61-+ 99+ 3i— Z9— Sc+ oz+- oI— “HHED ‘OuIpIeusiag UPS ge QI-+- eereeee ve— ° of— ° 6c+ ° we— ° —— tr-+-° vi— ° te+ ° 91— °° ze-+ c— . 6z— ° ci+ ° SI— * oz-+ ° e+ ° ce— ° ziI-+ oe OC = or— ° z+ ° e+ ° fi— ° Qorr or-++ 61— ° qe ‘aye eS Sz H“MOUSMUSMOKTMNOSAMOKMNOAMOAMOSAOYO 2961 9961 S961 vo61 £961 zgO1 1961 0961 6S61 992 “gi Sp— eS1 Zor gi— 601 W-- og- Zer _11-— ei o9+ og ofr €&— SZ gI— cei VII 6c+ Fil vit Sor 611 2S+ gor gr-+ €6 rr SE of Si— oir 26 gz— OO! gz+ 2g o11 6— AZII 1z+ 99 68 €1-+ vol ov— O6£1 66 II+ gl Sc+ Zor zr Sb— 9gg1 Q— fil col —¥zZ——— Zo ee+ ob €2Z I1z— IVI f— 72 69 gy— gSI cc— 6£ OF I— €Z ge— cli vor gf+ OI ce+ 2g Zor e+ 2S Zz— ghi izI1 oz— 601 S— 16 98 6— 901 gz— 66 14 1v— vv1 II— obi 621 f+ 121 0 OOr oor fi— £1 vi— £1 ozi Sz— 16 Olsj- os slie Tél — si = 09 QI— ZOI 98 zi— ZV 1S+ gS 601 c+ SII sss Cite eee 1s+ 624 yseo =paAtoS Vv £0 “2107 -4O TO 40g ‘o]OD oro ‘19AUuIq ‘yeuUuloUlD yseo = psAtas 16 ¢S1 zZ vi og Sor vZ gol ZII 6fI IzI gel ozl OII ZZ gr 6L 68 46 €o1 vel gzi 99 Col SEI OZI Of! -4O oe g'0z oi+ zZ ZQ z+ 01 gol Iz— II £6 gs— Szi 69 €e-- C11 Soft. €— 101 96 cst+ 1g QZI gz— I€1t Sor vi-+ gir gzi SiI-+ &4 88 of-+ 9S 98 cf€— 101 99 ¢€— S6 z9 zi+ $6 Zo1 c—— 10 98 z+ 88 OIl o-— 2g 8022 VI-- 59> 60 9+ 44 Coes ve Orie val v~— QOI 2g 61+ g6 Lit €ir+ 142 9 =%bv— v6 06 vL+ gf ZII 9og— Zor Iv I1f— or 62 Zt Qel = gel ZZ— QB zg Zz+ gv $2 6I— $6 94 zI1+ z2f F898 4i+ of & €— gor Sor S+ gf Coe (g— Crre Ser ee+ zy %8 =Qe— PS ge t~e— zi 66 LZi— ZL Sa of— If 111 gi+ Zor SzI oz+ o£ 06 1z— Szi- v01 e— £6 1g ©=6bS+ 399 zz1 II— zg 1Z i= 40 £8 €1— oor 4g 6+ gf 48 lI— ZII “101 €1— 4g ZZ gz— 3 ZS 19+ gir 621 Vv 4yseo peAtos V yse0) = pa Aas £0 7301 GO, FO 70f -40 aus, "xd “N uojsapieyy ‘yoreusig £°9z FA0 v+ €c+ 14+ zI+ Fi— ¢s— oI— 6z+ zI-+ £3— Si+ 1v+ Iz+ zI— e+ — 6I1— ge— (Os ge— oI— oI— ce— ce+- zb+ Rene ie 96 ve Sg 6g SZ gd zg est bl 98 Sg 1Z cet £9 69 6S 19 06 gs o£ ror ZZ O41 6g eS 89 ev tg eg bg 96 «#8 $6 Z6 VII Ill 88 69 Zit 6 06 66 Zi1,- 16 gf 89 0g o£ ozt -50 ZZ Ill eZ v9 yseo = pe Atos 2107 -4O sonyeA ULOU YUOW-InN0F 0'9z Tate ceo— gz+ er— Zo+ z+ 6f— ti+ or Date ae o— vi— of+ erp tz— cS-- 6¢+- ol— p— viI+ be+ ez— 1z+ Qo ZI— Ly+ S'1z LYve £6 +6 S1+ 22 £8 gt-+ gor &Z gz— O11 72 zS+ g9 98 atte S-— -Olipeesile=s0-j-7 = OVT ver, 10 vi— gor v6 os+ +6 1Q gor of— 6o1 64 fv+ 1g 601, el te ZV —= 7 ie OF fo+= 10 SzI 98 g— 306 06 o>. €S vS So O6b— O6zl og 6&— 99 Cll] ghee Ii (Slim vel roc Met Orr oSi 65+ 6g gbr 6z— 98 IZi O11 zez— 06 89 gi+ 66 vor g6 f— 66 96 19— SEI igi) ZOE i4- 00 46 g9— eri ovr ofr of-+ 12 rom s9Z— Tel 69 lit 6— O0ZI- tir 7— $9 cH 161i. 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Mean (O-F)=20 percent <5 6-10 10-20 20-30 30-40 > 40 Bismarck As ota eta. 6 3 8 5 I 4 Charleston join! see eb 4 oO 10 8 I 4 CCITICHATTA CTY chee clap als ais a's 4 4 7 3 4 5 Independence .......... 4 5 4 8 2 4 Madison acisjtaccnkicrse tvs 3 5 8 4 4 3 INa‘Shivilleteemcr crete 2 z, 2 8 5 3 PortGibsonuac cece 8 2 6 5 4 2 Rochester y auc aisle sietiele's 5 5 5 7 3 2 SMOkane oes selciss aheeaie 3 6 7 6 2 3 Spr Eos ie eid ae ovelete 6 4 3 6 4 4 Washington! 2.330.056 6 5 8 4 2 2 Group 2. Mean (O-F)=26 percent <6 7-13 14-26 27-40 41-52 > 52 Albany? Oreg.) 4.6 ei sa 4 5 7 5 4 2 POUISE A ele aches mais I 7 9 6 I 3 Denver. oh aah eae 3 2 10 7 3 2 METEGlemROGI jaysrreecd rears 5 6 4 6 I 5 Peoria. ss Heidislesusdnses 4 3 6 7 4 3 SAUISEUIBY tera larns c eers wi bie 5 5 B 8 3 3 Group 3. Mean (O-F)=30 percent <7 8-15 16-31 32-46 47-62 >62 DEtROItE cis ae eit cteke wove 5 5 4 8 2 3 Natural Bridge, Ariz.... 6 4 4 9 2 2 Salt Wake os cja.0 sictesore's ss 6 2 4 6 4 5 Santan Mel sivas. cs iiets astonsis 5 5 7 4 2 4 Sti (Paul iijccrcsae,crrsinarevats 3 5 8 6 4 I Group 4. Mean (O-F)= 40 percent <10 II-20 21-40 41-60 61-80 > 80 FGINBASO eecorebetas uieisvaeieies 3 5 5 3 7 4 SacramentOn iescsiicts oO 4 II 6 2 3 San Bernardino ........ 3 2 9 7 2 4 Sumsvohiesuenneaces 104 106 159 152 74 80 MEATS iehsjcleersteie eres 4.2 4.2 6.4 6.1 3.0 3.2 amitsyeeieeseree ce 98° Fic. 13. 27 NO. 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P. ees = - 7 #- ee = 2 = E eet = Se sere ao — ae aSF-et- > 1 J c3c-se ted , er SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 139, NUMBER I1 (Enp oF VoLUME) A CLASSIFICATION FOR THE BIRDS OF THE WORLD By ALEXANDER WETMORE Research Associate, Smithsonian Institution (Pusiication 4417) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION JUNE 23, 1960 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLUME 139, NUMBER 11 (Env oF VoLuME) & CLASSIFICATION FOR, THE BIRDS OF THE WORLD By ALEXANDER WETMORE Research Associate, Smithsonian Institution SOeCOCeeg (Pustication 4417) CITY OF WASHINGTON PUBLISHED BY THE SMITHSONIAN INSTITUTION JUNE 23, 1960 SMITHSONIAN ,..0. \ INSTITUTION YUNA & ttf, THE LORD BALTIMORE PRESS, INC. BALTIMORE, MD., U. S. A. AV CLASSIBIGATION: KOR THE BIRDS; OF) THE WORLD By ALEXANDER WETMORE Research Associate, Smithsonian Institution The principal additions to current information that affect the ar- rangement of the family and higher groups in birds since the previous paper on this subject by the author was published (1951, pp. 1-22) have come in the fossil field and deal in part with the earliest known forms of the Jurassic and Cretaceous periods. While there has been much discussion of family limits among the Passeriformes, with considerable spread of opinion as to family limitations, in the main these have been expressions of individual viewpoint, without com- pletely firm support in the new information offered. Valuable new data that are accumulating from many sources relative to this order, where they are completely decisive, in the main suggest better align- ment of existing families through shift of genera from one group to another. The great majority of the many species still require detailed anatomical study. Under the revision of the International Code of Zoological Nomen- clature as adopted at the Fifteenth International Zoological Congress held in London in July 1958, now in press, a new rule provides that family names are to be based on strict priority in publication. There is no attempt to follow this requirement in the classification presented herewith since the final draft of the Code was not yet in print when the paper was under preparation. It is apparent, however, that ac- ceptance of this new proviso, while intended to establish stability, in the beginning will bring many changes in current family and higher group designations in the class Aves. The following notes that discuss the more important changes are added to material from the introductory section of the revision of 1951 where this remains pertinent. In the classification at the end of the text the fossil groups are enclosed in brackets to enable their ready recognition on the part of students familiar mainly with the family and other categories of living kinds. Archaeornithes—The recent careful study of the specimen of Archaeopteryx in the British Museum (Natural History) by Sir SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 139, NO. 11 2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 Gavin de Beer has added greatly to knowledge of this bird through application of modern methods of examination. De Beer (1954, pp. 39-41) has outlined clearly the resemblances found in the two nearly complete specimens preserved in London and in Berlin and has shown that most of the differences between them that have been described either have been misinterpreted or do not rate the value that has been assigned to them. His conclusion is that “proposed generic and even specific distinction between them calls for very critical examination.” In his final statement on this part of his study (lLc., pp. 50, 57) he unites both under the name “Archaeopteryx lithographica Meyer.” In brief review, formal recognition of the two specimens as repre- sentative of separate species came when Dames (1897, p. 829) named the one in Berlin Archaeopteryx siemensii. Petronievics (in Petro- nievics and Woodward, 1917, p. 5) considered that differences be- tween the two were of sufficient weight to separate siemensii tenta- tively as the type of a new genus, Archacornis. In a later study Petronievics (1921, p. 10), after further consideration, was definite in establishing the two in distinct genera and added that they might “vielleicht sogar zu zwei verschiedenen Familien gehoren.” In a more detailed account (1925, pp. 67-69) he placed the two in separate families, which he maintained later in a further review (1950, pp. 118-120). The major points on which Petronievics based his two families have disappeared through the information supplied by de Beer. There re- main, however, distinctions of size and relative proportion, the London specimen being about Io percent larger in general dimension, with the foot about 25 percent greater. De Beer regards these size char- acters as individual, to be attributed either to age or to sex. Steiner (1938, p. 292), who also has considered the two identical, says that in his opinion the Berlin specimen was a young individual and a female, in contrast to the London example which he believes was a mature male. While my personal study of this problem has been confined to views of the London fossil and the nearby cast from Berlin in the British Museum, additional comparisons of casts of the two in the U.S. Na- tional Museum, and examination of published figures, it appears to me that the foot of the Berlin bird not only is smaller but also has the toes of different proportion in relation to one another and to the tarsometatarsus. The wing elements in the two specimens appear quite similar, but the entire leg in the Berlin bird seems more slender. It is possible that these ancient birds, like some reptiles, continued to grow in size for a longer period than is true with modern species, NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 3 a factor, however, which must remain hypothetical. Steiner’s supposi- tion that the London specimen is male and the Berlin fossil female is equally speculative, since if sex is assumed, the reverse might be true. While the male is larger than the female in most living birds, this is not the universal rule, and as reptilian characteristics persist in these earliest known avian forms it must be remembered that in reptiles it is common for the female to be larger than the male. As a further contribution to available information there should be noted the analysis of the primary wing feathers by Savile (1957, pp. 99-101), which points out an apparent difference in wing formula between the London and the Berlin birds. This recent observation if accepted would indicate rather wide separation, but, on the other hand, if denied would serve to bolster the conclusions of de Beer. A third specimen found in 1956 near the point where the first ex- ample was discovered shows mainly wing and leg bones and vertebrae, in addition to feather impressions. It has been described in detail by Heller (1959, pp. 1-25), who finds that it agrees in size and char- acters with the one in London, so that there are now two of the larger form known. | It is important to have a modern study, like that of de Beer, of the Berlin specimen, to add to the data assembled by Dames. As matters stand, the three known skeletons present an appearance of differences sufficient to mark them as two distinct species on the basis of criteria found in the osteology of living birds. These data, for the present, appear to warrant recognition of two genera, Archaeopteryx represented by two specimens and Archaeornis by one, which, how- ever, should be united in one family, the Archaeopterygidae. Ichthyornithes——A recent study by Gregory (1952, pp. 73-88) has severed the long-standing association of Hesperornis and Ichthyornis in a superorder separated from all other birds known from the New World through the possession of teeth. In brief, Dr. Gregory has shown that the toothed lower jaw fragments allocated to the skeleton of Ichthyornis dispar Marsh, unduly large in proportion to the rest of the skull and the skeleton with which they have been associated, in reality are not avian but are those of a small mosasaur. Two other jaw fragments placed by Marsh with Ichthyornis anceps and I. victor are similar, so that all these specimens, which have the teeth in sockets, are identified as reptilian. This leaves Hesperornis as the only group of Cretaceous age in which teeth are known. To give a balanced treatment that will emphasize the important characters of the birds concerned it has seemed appropriate to establish a suborder Ichthy- ornithes for the Ichthyornithiformes, separated from all other birds 4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 by the possession of biconcave vertebrae. I have given a somewhat more detailed discussion of this matter elsewhere (Wetmore, 1956, pe 2)i The penguins.—The question of the weight to be given the peculiari- ties of uniform pterylosis, extreme specialization of the wing as a flipper for submarine progression, and incomplete fusion in the meta- tarsal elements, as well as such other details as erect posture in stand- ing and walking and the anatomical adjustments involved, found in the penguins, is one that has merited careful review. It seems reasonable after this examination to retain the Impennes as a superorder, at least until we have further evidence through fossils as to their line of evolution. It is necessary, however, to remove the fossil family Cladornithidae, since Simpson (1946, pp. 24-25) has found that the two genera Cruschedula and Cladornis placed in this family have no apparent relationship to the Sphenisciformes. These two, described by Ameghino from the Deseado formation of Patagonia, now placed in the Oligocene, are based on fragmentary, considerably flattened metatarsi. The descriptions and figures that have appeared thus far are not sufficiently definite to demonstrate characters of importance in classification. However, from what we now know these ancient birds cannot be considered as ancestral penguins of terrestrial habit, as has been supposed. The only suggestion that has come to me is that pos- sibly they may belong in the order Pelecaniformes, in which I have placed the family tentatively in a suborder Cladornithes (see p. 25). The Neognathae.—One important result of recent studies has been the allocation to the Neognathae of the orders formerly separated as the Palaeognathae. For years I have felt that recognition of the Palaeognathae, as a separate group apart from other birds, on the basis of a supposed peculiarity in the palate, stood on flimsy ground. The studies of McDowell (1948, pp. 520-549) demonstrate that the structure of the palaeognathous palate, in which the palatine and pterygoid bones are articulated by a squamous suture, is variable from order to order and that in fact the details of this union differ con- siderably in the several groups. For example, McDowell points out that in Dromiceius the palatine and pterygoid are not in contact, while in a number of families placed in the Neognathae, as in the Anatidae, to name only one, the two bones are in articulation. As there is no clear-cut separation, the former Palaeognathae must be combined with the Neognathae. The supposed bird Caenagnathus collinsi described by R. M. Stern- berg (1940, p. 81) from the Belly River series of beds of Upper Cretaceous age in Alberta has been carried tentatively in our avian NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 5 classification, though it has been my belief from the beginning that it was reptilian. It is known from a lower jaw, beautifully preserved, without appreciable deformation and practically complete except for part of the lower section of one ramus. The resemblance to birds is found in the lack of teeth, fused symphysis, and the considerable size of the mandibular foramen. While these are characters found in birds, there is nothing peculiar included since all are duplicated in some of the groups of the Reptilia. The fossil resembles Reptilia in the form of the articular surface, the forward position of the coronoid area, the conformation at the symphysis, especially on the upper surface, the upward curvature in that area, and in the general texture of the bone. In none of these is there exact duplication in Aves, except partially in the form of the symphyseal region. The whole appearance of the bone strongly suggests a species related to the Ornithomimidae among the therapod dinosaurs. In view of this the “Order Caenagnathiformes” is now omitted from the avian classifica- tion, since it is felt that its continued tentative inclusion may promote misunderstanding as to its status. The family Eleutherornithidae is introduced for the fossil Eleu- therornis helveticus Schaub, from the Eocene of Switzerland, de- scribed from a fairly well preserved pelvis. Apparently this is repre- sentative of an ancestral group from which the living ostriches may have come. Its greatest importance is found in its indication of re- lationship with carinate groups though of unquestioned ratite stock. It is thus important as definite indication that the struthious birds are descended from flying ancestors, not from some distinct cursorial line that always has been flightless, as some have contended. The genus Podiceps.—The differences of opinion that prevailed for years as to the application of the generic name Colymbus have been adjusted currently by an arrangement under which Gavia has been accepted for the loons and Podiceps for the grebes. There is, how- ever, discussion still as to the proper spelling of the ordinal and familial names for which Podiceps is the base. The uncertainty arises from misunderstanding of the derivation of this generic term. The colloquial name applied to these diving birds in the English of the 16th to the 18th centuries (and later) was “arse foot,” or “arsfoot,” from the posterior position of the leg. The term is found in the early dictionaries of Johnson, was carried in the later editions of Todd and Walker, and is still found in a footnote in Webster’s 1953 volume, with indication there that the word now is obsolete. Some early authors who wrote in Latin rendered this term appropriately as “‘Podicipes,” as for example Willughby (1676, p. 258), and Ray (1713, 6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 pp. 125, 190), where the horned grebe is listed as “Colymbus sive Podicipes minor.” Catesby (1731, p. 91) wrote of the pied-billed grebe under the heading “Prodicipes Minor Rostro vario,” but he corrected the spelling of the first word in the legend for the plate that faces the text, which is labeled “Podicipes &c.” This account by Catesby was the sole basis on which Linnaeus (1758, p. 136) estab- lished his specific name for the pied-billed grebe. And it is here that present-day confusion has its beginning, since Linnaeus called the bird “Colymbus Podiceps,’ and in citing the reference to Catesby wrote it ‘“Podiceps minor, rostro vario.” While he corrected Catesby’s error in spelling he thus made another of his own, which remains in our current name Podilymbus podiceps (Linnaeus) for the pied- billed grebe. Following Linnaeus, John Latham (1787, p. 244) pro- posed the genus Podiceps, in which he included several species of grebes, with basis for the name on Linnaeus, as he makes reference to “Colymbus Lin.” The error in spelling was recognized by several early authors, as in a note attributed to Oken (1839, p. 674) and one by Gloger (1854, p. 430). Correct usage for a family name based on Podiceps (=Podicipes) was indicated by Newton (1896, p. 381). That this history, well known up to 40 years or so ago, has been for- gotten by many is shown by recent action of the International Com- mission on Zoological Nomenclature (1957, pp. 300-304) which it appears should have further review. The data supplied by the Com- mittee to Dr. Grensted, as classical adviser, were misleading, as there was no indication for his information that “Podiceps” had been derived from “Podicipes.” As the terminal root in Podiceps is a contraction of the Latin pes, pedis, it would appear that the correct form for the family name is Podicipedidae (not Podicipidae or Podicipitidae), and for the order Podicipediformes (not Podicipitiformes or Podicipidiformes). The Procellartiformes.—Family segregation in this order has been oversimplified in some recent discussions, probably through misunder- standing of the group characters, possibly also through somewhat con- fusing names that have been applied to familial and generic categories. Verheyen (1958, pp. 11-14) has placed the Pelecanoididae in an order with the Alcidae, as indicative that the auk group is allied rather closely to the Procellariiformes. The resemblances that he cites appear due to convergence, as the basic form of the diving petrels is definitely that of the shearwater-petrel group. Aside from this, the Diomedeidae and the Pelecanoididae have been accepted without apparent question, but the remaining species have been combined by some under a single family name. Lowe (1925, pp. 1436-1443) has shown that the genera NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE Ti included in the Hydrobatidae have a simplified condition in the quadrato-tympanic region of the skull in which the opening of the upper tympanic recess is small, and is so located that it separates the squamosal and opisthotic facets. In addition, the posterior border of the sternum is truncated and entire, and basipterygoids are absent or are represented only by small spines. In the Procellariidae, on the other hand, the foramen of the upper tympanic recess is greatly enlarged and lies anterior to the two facets for the quadrate, which are joined by a bridge of bone; the posterior border of the sternum is notched ; and basipterygoid processes are present. These constitute distinctive characters at the family level. The Pelecaniformes.—lIn the arrangement of suborders in the order Pelecaniformes we encounter in marked degree the standard difficulty of logical placement in linear alignment of groups that really stand in three-dimensional relationship. Lanham (1947, pp. 65-70) has made a summary of the major anatomical characters of the group in which he points out the differences that set off the Phaéthontes and the Fregatae from the Pelecani. There is no question that the first two carry primitive characters, which may be presumed to be similar to those found in ancient ancestral stocks, since in these resemblances they are more like other types of birds, notably the Procellariiformes. From this style the families of the suborder Pelecani have become widely divergent. Although the tropicbirds and the frigate-birds both have retained a part of what may be regarded as a basic pattern, they are so distinct in other respects that it appears to be more reasonable to relate them individually as branches from a common stock rather than to combine the two on one line, separate from the Pelecani. The Phaéthontes possibly may have separated earlier than the Fregatae. Among interesting differences other than those of internal anatomy, it may be noted that the tropicbirds have the young covered with down at birth and that the adults possess series of air cells under the skin on the forepart of the body like those found in pelicans and boobies. The frigate-birds have young almost naked at hatching, and the emphy- sematous condition is mainly lacking. In view of this I prefer to continue to align these groups on either side of the Pelecani. Though there is no question that the cormorants and snake-birds are closely allied, they differ in such degree that they should be retained in separate family status. The snake-birds are marked by a peculiar conformation of the cervical vertebrae through which the beak be- comes a triggered spear in feeding. The bridge of Donitz on the ninth vertebra is an important part of this arrangement. The stomach also is unusual in possessing a curious pyloric lobe, lined with a mat of hair- 8 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 like processes. And there is only one carotid artery while in cor- morants there are two. The description of Osteodontormis orri by Hildegarde Howard (1957a, pp. 1-23) from the Monterey formation in the Miocene of California adds a third species to the strange Odontopteryges, whose common character is found in the sharply pointed, dentate projections developed on the margins of upper and lower mandibles as continuous parts of the bony structure of the jaws. This suborder was placed tentatively (Wetmore, 1930, p. 3), following Lydekker (1891, pp. 57-58), in the Pelecaniformes, but this was not definite, as the char- acters of Odontopteryx have been interpreted by some as indicating closer alliance to the petrel-albatross group. In July 1956, at the British Museum (Natural History), through the kind attention of Dr. W. E. Swinton, I had the privilege of studying the type skull of Odontopteryx toliapica Owen, which came from the London clay of the lower Eocene, on the Isle of Sheppey, Kent, England. It was possible thus to ascertain certain details not clear from the published accounts. Asa result of this study it is my opinion that the characters clearly indicate relationship with the Pelecaniformes. Without repeating unnecessary detail, available in Lambrecht’s great volume (1933, pp. 304-307), it was interesting to note the strongly marked craniofacial hinge at the base of the bill, like that of gannets and cormorants, and also the impressed line along the side of the premaxilla, and the definite closure of the external narial opening, as in the Sulidae. The distal articular end of the quadrate suggests that of Phaéthon, though somewhat more flattened, with the whole articular surface narrower, and the separate segments more nearly in line than in any living species of the various pelecaniform families. The lachrymal appears to have been slender and is firmly anchylosed on its upper margin to the frontal as in Phalacrocorax. The rounded cranium suggests that of pelicans, rather than the more flattened form of other families of the order. The sum of the char- acters indicates a bird of gannetlike diving habit that, when slippery aquatic prey was seized, could hold it firmly in the sharp dentations of the mouth. Dr. Howard in her interesting study of Osteodontornis has elevated the group to the rank of an order, on the consideration that it “may represent an early connection with procellariiform—pelecaniform stock” (1957a, p. 22). It has seemed to me appropriate to emphasize the evident pelecaniform character by retaining the two families recog- nized in subordinal status in that group, since the resemblances that point toward the Procellariiformes appear to be much less definite and NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 9 possibly may be subject to other interpretation. It is desirable now to place the Odontopteryges at the beginning of the order because of their antiquity. The known history of the group, which begins in the early Eocene, indicates probable ancestry in Paleocene time. The pointed projections on the jaws, assumed to have been sheathed in the in- tegument of the bill, were without question used in seizing prey. The disappearance during Miocene time of such a holding apparatus may indicate that the bony projections were not completely successful for their purpose, perhaps because of their hollow centers, as accidental breakage in them would not be restored. The fine serrations restricted entirely to the ramphotheca, found in the straight-billed species of the pelecaniform order (tropicbirds, gannets, boobies, and anhingas), may be regarded as a functional replacement. The change in position made to the beginning of the order covers only the Odontopterygidae and the Pseudodontornithidae and leaves Cladornis and Cruschedula still unsettled as to relationship. As ex- plained above (p. 4), Ameghino described both as forms of penguins, but Simpson says that they have no connection with this group. As the suborder Cladornithes, they are located in their former uncertain position at the end of the Pelecaniformes. Suborder Ardeae—The general resemblance of the boat-billed heron (Cochlearius cochlearius) to the night herons has been the occasion of differences in allocation of its rank in classification from that of a subgenus of Nycticorax to full family status. In a recent review of the Ardeidae, Bock (1956, pp. 31-35) has treated it as a separate genus in a “Tribe Nycticoracini” allied to Nycticorax. Super- ficially the boatbill is like a black-crowned night heron, but in detail there are outstanding differences. The enlarged bill is obvious, and there are four pairs of powder-down patches, instead of the three found in the other herons. In the skull, the bill has been changed from the spear point usual in herons to a broad scoop with the roof of the mouth smoothly arched. The lower jaw is widely bowed to fit this change, and the symphysis is greatly reduced in length. The palatines are so greatly broadened, and so inflated on the outer posterior margin, that they have little resemblance to the ordinary heron form. The quadrate has the orbital process shorter and thicker and the mandibular articulation narrowed ; the lachrymal is small ; the eye opening considerably enlarged to house the exceptionally large eye; and the external nasal opening considerably reduced. The palatal musculature is decidedly stronger than in the true herons. In life boatbills act like night herons, as they roost and nest in groups and are mainly nocturnal. When hunting at night, I have IO SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 found them feeding in shallow waters, often in riffles where they scoop at their living prey, rather than spear at it as is the custom with the typical herons. The eyes, wood brown by day, at night reflect the jacklight with a faint orange sheen, which I have not observed in other herons. The eggs are pale, nearly white, and often are lightly speckled with brown, so that they resemble those of the tiger bittern, Tigrisoma lineatum, rather than those of the night herons, which are deep blue. While there is no fossil record for the boatbill, I regard it as an ancient sideline from the typical herons that, judged from its present restricted range in the American Tropics, has not been too successful. It may seem attractive to unite Cochlearius with the true herons, but from long acquaintance I regard their characters, briefly outlined above, sufficient to maintain a separate family status. In view of the fact that the structural characters of the Balaenicipit- idae have been summarized clearly by Stresemann (1934, p. 809), it seems strange that the status of this family has been a matter of ques- tion. The single species shows affinity both with storks and with herons, in addition to outstanding peculiarities of its own. Miss Cottam (1957, pp. 51-71) has made a careful summary of the osteol- ogy from which she deduces a pelecaniform relationship, but this appears to be due to convergence rather than to actual relationship. The great enlargement of the skull has occasioned superficial re- semblances to pelicans, but these, and others seen elsewhere in the skeleton, are subordinate to the general sum of all characters, which is ciconiiform. Phoenicopteri—The position of the modern flamingos, which show characters that point on one hand to the Ciconiiformes and on the other to the Anseriformes, has been a matter of some variance in al- location, Mayr and Amadon (1951, pp. 7, 33), with only brief discussion, have set them up as a distinct order, but general opinion has carried them as a suborder allied to the herons, storks, and their relatives. The latter course remains justified when the fossil genera Palaelodus and Elornis of the upper Eocene to Miocene of western Europe are considered (Wetmore, 1956, p. 3). This group of flamingo relatives was identified in North America when Alden Miller (1944, p. 86) described Megapaloelodus connectens from the lower Miocene of South Dakota, a species to which remains from the upper Miocene of California also are referred (Loye Miller, 1950, pp. 69-73; 1952; pp. 296-298). The group may be recognized as the family Palaelodidae, on the generic name Palaelodus Milne-Edwards (1863, pp. 157, 158). (There has been confusion relative to the proper spelling, since Milne- NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE Dt Edwards in his important later work [1868, p. 58] used the form Paloelodus.) Howard (1955, pp. 3-23) has described a still different form of the flamingo group as Telmabates antiquus from the lower Eocene (Casamayor formation) of Chubut in Patagonia. While this species resembles the Palaelodidae in shortness and other details of form in the leg, it may prove to be representative of a separate family on characters found in the vertebrae and wing, as suggested in the original description. It is regarded for the present as of subfamily status in the Palaelodidae. Suborder Cathartae.—The superfamily Neocathartoidea, and family Neocathartidae, for the curious vulture Neocathartes grallator (Wet- more), discovered in the Upper Eocene fossil beds of Wyoming, in- troduced a new element in our known avifauna in the form of a small-winged, strong-legged vulture that evidently was terrestrial with limited powers of flight. It had about the same relation to the other American vultures that the secretarybird has to the hawks and falcons. Its inclusion also requires a separate superfamily, the Cathartoidea, for the previously known cathartine families. Galliformes.—The Numididae, which have been placed by some as a subfamily of the Phasianidae, differ in completely lacking the tuberosity or plate on the inner side of the second metacarpal that is sO prominent in pheasants and grouse. It should be recorded, however, that Hudson, Lanzilloti, and Edwards (1959, p. 64) note that Numida shows no peculiarities in the leg musculature when compared with the Phasianidae. The Tetraonidae, in contrast with the Phasianidae, have the pelvis relatively much broader and different in proportion, and the tarsus relatively shorter in relation to the length of the tibiotarsus. With these differences in mind it seems reasonable to retain the three groups in family status, at least until more detailed knowledge of their anatomy as a whole warrants change. Gruiformes.—In the Turnices the two genera of bustardquails, Turnix and Ortyxelus, have no hind toe, the wing is eutaxic, only the left carotid is present, and the eggs are rounded oval. The plain- wanderer of Australia, Pedionomus, has a small hind toe, the wing is diastataxic, right and left carotids are found, and the large eggs are pyriform. It seems desirable to continue these as separate families, rather than as subfamilies of one group, an arrangement that Strese- mann (1933, p. 760) has accepted. It has long been known that Mesites Geoffroy for the curious roatelos of Madagascar is antedated by the same name used by Schon- herr for a group of beetles. It has been in error, however, to replace I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 this with M/esoenas Reichenbach 1862, since the conflict had been noted seven years earlier by Prince Bonaparte who gave the group the name Mesitornis (Bonaparte, 1855, p. 484). The suborder becomes Mesi- tornithides and the family Mesitornithidae. In the course of study of the fossil Andrewsornis abbotti from the Oligocene of Patagonia, Bryan Patterson (1941, pp. 50-53) has re- viewed related groups to the end that he has added the family Psilopteridae for the South American fossil genera Psilopterus and Smuiliorms. Further, he has placed Phororhacos and its allies as a superfamily Phororhacoidea under the suborder Cariamae. His further observations on these matters are to appear later in a more comprehensive paper. The family Cunampaiidae, for the fossil Cunampaia simplex, named by Rusconi (1946, p. 1) from the Oligocene of western Argentina, while placed in the Cariamae, still remains of uncertain status. The allocation of the phororhacid group to its new position and its demotion from subordinal status requires recognition of a super- family Cariamoidea for the living Cariamidae and the fossil group Hermosiornithidae. The common name for the Cariamidae in most English writings has been “Cariama,” being the form instituted by Marcgrave in 1648 in his Historiae rerum naturalium Brasiliae, when he rendered the Tupi name “cariama” as cariama. This was copied by subsequent authors, including Linnaeus in his twelfth edition, and so came finally into English usage, beginning with Ray’s translation of Willughby’s Ornithologiae in 1678. Seriema, a modification of the Indian word gariama, is used in Brazil, and with that spelling has come into the Engish language, where it should replace the other form. Charadruformes.—Differences of treatment at present are found mainly in the superfamily Charadrioidea and the suborder Lari, in which the groups have been regarded by some as of family value and by others have been allocated to the rank of subfamilies. The var- ious studies that have been made have not been complete from a taxonomic point of view except for part of the species, and the con- clusions derived from the data available appear in the main more philosophical than concrete. The picture therefore still remains confused. In view of the diverse specializations that are apparent, and the obvious long evolutionary history, it appears better to me to continue to acknowledge the main segregations as families, at least until the subjects involved have been more thoroughly investigated. A family, Rhegminornithidae, covers the fossil Rhegminornis calobates Wet- more, described from the lower Miocene of Florida. This was as NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 13 large as a medium-sized curlew, of peculiar form as regards the foot, the only part of the skeleton known, which shows certain char- acters that seem to point toward the jacanas, though the bird is to be placed in the Charadrioidea. It should be noted that the family affinity of the turnstones and the surfbird, long considered members of the plover family, is not certain as some studies (Lowe, 1931, pp. 747-750) place them in the Scolo- pacidae. (See also Bock, 1958, pp. 85-86.) In the Lari the terns and the gulls are regarded as one family, though there are some reasons that make further examination of this treatment desirable. The Stercorariidae possess a 2-notched sternum, large caeca, a cere, and a complex rhamphotheca. In the Laridae ambiens and biceps slip are present, the sternum is 4-notched, there is no cere, and the rhamphotheca is simple in form. In further discussion of proposals relative to this group it is pertinent to observe that a logical scheme of classification should attempt to outline relationships in living and fossil species through examination of all available data, considerations in which modern studies of behavior find increasingly useful part. There are pitfalls and hidden traps, however, when attempt is made to establish affiliation through any single method of approach, as inevitably inconsistencies appear. I fully agree with Martin Moynihan (1959, pp. 22-23, 35-38) that the skimmers (Rynchops) represent an early separation in the ancestry of the gull-like birds and find it pertinent that this is shown in their behavior pattern. At the same time these birds present out- standing peculiarities that should be considered in assigning them ap- propriate status in relation to their relatives. The bill, compressed to knifelike form, with great elongation of the ramphotheca of the lower jaw, is unique, and the method of feeding, where the lower mandible cuts the water surface with the bird in flight, is equally strange. The structural modifications in the form of the skull from that found in skuas, gulls, and terns also are too extensive to be ignored. The elongated blade of the lower mandible anterior to the symphysis of the rami is intriguing but less important than the pro- found changes elsewhere. The palatine bones are greatly expanded, the orbital process of the quadrate is reduced to a short, pointed spine, the impression for the nasal gland is much reduced, the frontal area is inflated and produced posteriorly, with compression of the lachrymal, and consequent reduction in size of the cavity for the eye, to enumerate the most outstanding differences in the osteology. Ex- ternally, the pupil of the eye is a vertical slit similar to that of a cat, and thus unlike that of any other group of birds (Wetmore, 1919, I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 p. 195). Other peculiarities have been described in the musculature. The sum of these characters justifies treatment of the Rynchopidae as a distinct family in their suborder. The fossil humerus, type of Mancalla californiensis Lucas, that was the first intimation of a flightless auk on the west coast, while unique for many years, now has been supplemented by abundant ma- terial from which an additional, smaller species, Mancalla diegense (L. H. Miller), is recognized. It has been possible also to construct a composite skeleton of the larger one that is sufficiently complete to give a clear picture of its form and characters. The evident peculiari- ties of the genus Mancalla are found in the wing, as elsewhere the skeleton resembles that of other alcids, except for differences of a generic and specific nature. In comparison of the wing with that of the great auk, now extinct, that formerly ranged the coasts of the North Atlantic, the humerus of Mancalla is generally similar, the forearm appears proportionately shorter, and the hand more elongated. Ulna, radius, metacarpal, and phalanges so far as present are more slender. The head of the humerus in Mancalla differs decidedly in the relative angles of different elements, and also in the conformation of the distal articular surface. The general indication in the west-coast bird is of a proportionately longer wing, with the slighter bones to be ex- pected in a form of lesser bulk. Loye Miller (1946, pp. 34-36) and Loye Miller and Howard (1949, pp. 222, 225) have likened the specialization seen in the wing to that found in penguins and explain any similarity to the great auk, Pinguinis impennis, as due to con- vergence. On this basis they have separated Mancalla from the other auks in the family Mancallidae. While I followed this, with some reservation, in the last revision of the fossil list (Wetmore, 1956, pp. 3, 80-81), a further review of the subject raises definite doubt, since, except for some specialization in the wing, Mancalla, as said above, is like other alcids. The change in the wing is no greater than that of Pinguinis, though the divergence is in a different direction. It would seem sufficient to place Mancalla in a well-marked subfamily, rather than in a separate family. Finally, the proposals of several authors to separate the auks in a distinct order appear to require further study. Strigiformes.—Old World ornithologists in the main regard the owls as belonging to a single family, but while all are deceivingly similar in general aspect, Ridgway (1914, p. 598) years ago sum- marized the considerable structural characters that separate the Tytonidae and the Strigidae. It is necessary here only to point out the more outstanding differences of the barn owls in lack of the manu- NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 15 brium, the different form of the posterior margin of the sternum, which is entire or 2-notched, the straight outline of the palatines, and in the ventral pteryla where the outer branch joins posteriorly to the main tract. The Strigidae possess a manubrium, the sternum is 4-notched, the palatines are greatly expanded posteriorly, and the posterior end of the ventral pteryla does not join the main tract at the posterior end. Apodiformes.—Lucas (1889, pp. 8-13; 1895, pp. 155-157) long ago demonstrated the differences between the true swifts and the crested swifts, though his work seems latterly to have been overlooked, in view of the recent inclusion of the two in one group, as by Stresemann and by Mayr and Amadon. The skull in the Hemiprocnidae is quite dis- tinct in the general form of the cranium and in the development of the nasals, vomer, and palatines. The hypotarsus has a tendinal foramen (like that found in hummingbirds), and the plantar tendons have the flexor longus hallucis connected with the branch of the flexor perforans digitorum, which extends to the fourth digit. Coupled with this there may be noted the curious nest, which, fastened to the side of a branch, is barely large enough to contain one egg, and the further fact that these birds perch regularly on branches and twigs in trees. As Apus Scopoli, published in 1777, is recognized now in place of Micropus Meyer and Wolf, 1810, for the type genus of the swifts, the terms in the classification change to order Apodiformes, suborder Apodi, and family Apodidae, which replace the former terms Mi- cropodiformes, Micropodi, and Micropodidae, respectively. Coraciiformes.—The proposal of Mayr and Amadon (1951, p. 35) to include the rollers in one family, the Coraciidae, with three sub- families, goes back to the arrangement of Dresser in his monograph of the group (1893, pp. xviii, 85, 101). Sclater (1865, pp. 682-688), however, many years ago, pointed out the pelvic powder-down tracts, the small manubrium, and other peculiarities of Leptosoma, and set it apart in a distinct family. The anatomy of the syrinx and feet was further elaborated by Forbes (1880, pp. 464-475). The family Leptosomatidae therefore should be recognized. The groundrollers, Brachypteracias, Atelornis, and Uratelornis, usually have been included as a subfamily of the Leptosomatidae, but Stresemann (1934, p. 829) places them in a separate family, the Brachypteraciidae. There seems to be reason for this in their general appearance, though their anatomy is not well known. Brachypteracias, in its skeleton, differs from Coracias and Eurystomus in the much greater depth of the outer notch on the posterior border of the sternum, in the much broader and stronger pelvis, the heavier femur, 16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 and the greater curvature of the shaft and reduction of the crista superior of the humerus. I have not seen the skull. The habit of life is markedly different. Although anatomical material of the other genera is not presently available, it seems reasonable to accept Strese- mann’s proposal. These peculiar birds certainly are not closely allied to Leptosoma. Lack of information on the anatomy of the woodhoopoes must be the reason for recent nonrecognition of the Phoeniculidae as a family separate from the Upupidae, since the two are quite distinct and have been so recognized for many years. The external differences are readily apparent. In the skeleton in Phoeniculus (of which I have seen several examples) the posterior part of the nasal area is ossified, there being only a small, narrow, elongated nasal opening ; the ecteth- moid is much reduced; the anterior end of the pterygoid is broadly expanded ; the sphenoidal rostrum is swollen at the anterior end, where the expanded ends of the pterygoids join it; the quadrates are de- cidedly larger; the keel of the sternum is greatly reduced, being only half as high as in Upupa; the furculum is broader ; the pelvis is nar- rowed, and considerably enlongated posterior to the acetabulum, with the ischio-pubic fenestra greatly enlarged; and the tarsus is heavier and broader, with two definite fenestra below the head. There are other minor details. In all of the above the characters of Upupa are directly opposite. The two groups appear to me to be sharply set off as distinct families. Passeriformes.—This order, with more living species than all the others combined, and far fewer fossil forms known, presents many difficult problems in logical arrangement. The major groups are clear, whether we rank them as suborders or superfamilies being a matter of opinion. But the limits and status of numerous families contained in these larger categories are uncertain since the internal anatomy is known for so few kinds that details of difference are poorly under- stood. Superficial resemblances, on the other hand, are so obvious in many cases that they cause confusion. Under the circumstances it continues to seem appropriate to me to accept the family grouping that has been current for many years, except in those cases where acceptable studies clearly indicate change. Supposition in these matters has led to various proposals for changes, some part of which un- doubtedly will prove correct. It is equally probable that a part, possibly the considerably larger part, may prove to be unfounded when details are more clearly known. If change is accepted under these cir- cumstances it may prove unwarranted, necessitating further shift, perhaps a return to the original status. Since this can only prove NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 17 confusing I prefer the conservative course. In the remarks that follow I shall discuss only a few matters on which I have more or less concrete ideas. In the superfamily Furnarioidea, von Ihering (1915, pp. 145-153) united the Furnariidae and the Dendrocolaptidae, since he was unable to separate two groups on the basis of the form of the posterior border of the nasal opening. The variation that he showed seems valid, but there are numbers of other points of supposed difference in the osteol- ogy and other structural details, so that his suggestion is far from established. Pycraft (1906, pp. 133-159), though seemingly uncer- tain in the beginning, finally retained the two families. It may prove that some genera are wrongfully allocated at present between the two groups, so that their shift, when we have sufficient information, will clear our understanding. In the Tyrannoidea, the family Oxyruncidae is known through ex- ternal characters that seem to warrant separation. If the sharpbills have other affinities it is doubtful that these are within the family Tyrannidae, where some have placed them. In the family Cracticidae, recognized by Australian ornithologists, the skull, according to Pycraft (1907, pp. 355-365), mainly from examination of Gymnorhina, has the zygomatic process of the squamosal bifurcate, the postorbital process large, the orbitosphenoid ossified, the interorbital septum with a single opening, the prefrontals unusually large, and the form of the palate peculiar. In his phylo- genetic tree Pycraft places the group on a common stem with the Artamidae, and not far from the Paradisaeidae. His account is diffi- cult to summarize in concrete form. The family Grallinidae is likewise recognized officially by Austra- lian ornithologists for Grallina cyanoleuca, the magpie-lark. The principal study of the osteology is that of Shufeldt (1923, pp. 16-19, pl. 6) but his account is mainly descriptive and without definite con- clusion. Amadon (1950, pp. 123-127) has placed Corcorax and Struthidea here tentatively, though this seems subject to further proof. Stonor (1937, pp. 475-490) has outlined excellent reasons for recog- nition of the Ptilonorhynchidae, finding that they differ from Para- disaeidae, with which they have been united, in having an apterium in the center of the dorsal feather tract, the tip of the vomer convex, larger, more developed maxillo-palatines, the margin of the palatines angular, smaller ectethmoid, much larger lachrymal, and _ slender, greatly elongated orbital ramus of the quadrate. The genera Loria and Loboparadisea, usually included here, he transfers to the Para- disaeidae. His conclusion is that “the Ptilonorhynchidae constitute 18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 a singularly complete and isolated family of the acromyodian passerine birds and show no special relationship to any other, being sharply marked off by the structure of the skull, the colour-pattern, and the bower-building habit.” (It should be noted that the names on Stonor’s figs. 6 and 8 have been transposed, fig. 6 being Semioptera wallacei, and fig. 8 Amblyornis subalaris, not the reverse as printed on pp. 481 and 483.) Oberholser (1917, pp. 537-539) has set up a distinct family Irenidae for the fairy bluebirds (Jrena), and Delacour (1946b, p. 3) a family Aegithinidae for the leafbirds, which would cover Irena, Aegithina, and Chloropsis. The proper allocation of the genus Chamaea for the wrentits, at present accepted by the A. O. U. Committee on Classification and No- menclature as a separate family, the Chamaeidae, is one of consider- able uncertainty. Delacour (1946a, pp. 18, 25, 35) has suggested that the group be located in the family Timaliidae in a special sub- family in which he includes also such diverse genera as Chrysomma (Moupinia), Panurus, Conostoma, and Paradoxornis (combining un- der this name Suthora, Psittiparus, Neosuthora, and Cholornis). This is an obviously heterogeneous assemblance, in which Chamaea has slight resemblances to the first only. From Moupinia poecilotis (placed in Chrysomma by Delacour) the wrentit differs definitely in weaker, less arched bill and in differently proportioned feet. It has no close similarity to any of the others that are mentioned. Although the relationships of Chamaea are obviously uncertain, it is retained as a family pending other information. In consultation with Herbert Deignan, expert in matters that relate to the birds of eastern Asia, the Campephagidae have been placed near the Pycnonotidae, an arrangement that agrees with that adopted by Charles Vaurie in his recent volume on the palearctic region (1959, p. 181), and the Paradoxornithidae are brought nearer the Timeliidae. The fossil family Palaeoscinidae, proposed by Hildegarde Howard (1957b, p. 15) for the species Palaeoscinis turdirostris, has been in- serted provisionally near the Pycnonotidae. The specimen on which this name is based is a skeleton found in Santa Barbara County, Calif., compressed in a slab of Miocene limestone of the Monterey forma- tion. The type, in which most of the bones are outlined, is one of those attractive silhouette impressions that delight the eye but that often pose difficulties in classification through lack of clear-cut characters on which to judge relationship. In the present instance Dr. Howard has concluded that “affinities of the Palaeoscinidae lie with the Pycnonotidae, Bombycillidae, Corvidae and Cinclidae” of the suborder NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 19 Passeres. Affinity with the Bombycillidae may be queried, as the fossil differs from Bombycilla in the proportions found in the hind limb, where both metatarsus and femur are longer in comparison with the tibiotarsus, and the toes appear longer, as well as of different propor- tion. The corvid affiliation also seems uncertain because of the slender form of Palaeoscints, since the skeleton of the crows and their relatives is strong and robust. Separation of the two genera of leafbirds, Aegithina and Chloropsis, in a family distinct from the Pycnonotidae is justified on the basis of characters found in the skull. The entire palatal structure is slighter than in Pycnonotus and allied genera, with the central plate of the palatine reduced in area, and the transpalatine produced posteriorly. The sphenoidal rostrum is slender, as is the orbital process of the quadrate. In Pycnonotus the palatine is broad, the transpalatine proc- ess distally is only slightly angular without posterior projection, and both the rostrum and the orbital process of the quadrate are strong and heavy. Herbert Deignan informs me that the group, recognized by several authors, seems to have been first separated by Cabanis (1847, p. 326), who designated it as the subfamily “Phyllornithinae” based on Phyllornis Temminck, 1829. This generic term is antedated by Chloropsis Jardine and Selby, 1826, so the family name based on this genus will be Chloropseidae, rather than Aegithinidae which dates from G. R. Gray in 1869 (p. 312). The fairy bluebirds, genus Jrena, often have been placed with the leafbirds but have no close connection with that group. The main external peculiarity of Jrena is found in the smooth, enamel-like tip- ping found in adult males on the feathers of the central dorsal area from the center of the crown back over hindneck, back, rump, and upper tail coverts, and on the elongated under tail coverts. As this is a secondary sexual character, not present in females, it has no value at the family level. In the osteology, the skull differs from Chloropsis and Aegithina in the completely open external narial opening, the ossification of the vertical plate between the nares, the more inflated lachrymal, and the more elongate maxillo-palatines. In the sternum the depth of the notch on either side of the posterior margin relatively is decidedly less, and in the pelvis the antitrochanter has the dorsal margin much produced laterally. The general resemblance in these matters is to species of the genus Oriolus. It may be observed further that the feathers of breast and back in the aberrant species Oriolus traillii and O. mellianus have smooth exposed ends that suggest the condition found in male Jrena. In view of these resemblances, and in lack of important differences, it seems sufficient to include the fairy 20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 bluebirds in the family Oriolidae, as the subfamily Ireninae, which incidentally dates from G. R. Gray (1869, p. 288) and not from the name Irenidae set up later by Oberholser (1917, pp. 537-539). Suggestions for the union of the Bombycillidae, Ptilogonatidae, and the Dulidae in one family are not substantiated by examination of the skeleton. Dulus, the palmchat, is widely different from the other two, a structural distinction that is further emphasized by its curious communal nesting habits. The first two seem more closely related but are separated clearly by characters found in the ectethmoid region of the skull, and in the manubrium, to mention only two points that are easily apparent. Delacour and Amadon (1949, pp. 427-429) con- sider Hypocolius closely allied to Ptilogonys. While Zimmer (1942, p. 10) believed that the family Vireolaniidae should be included in the Vireonidae, separate family rank in my opinion is definitely justified. In addition to characters assigned by Pycraft (1907, pp. 378-379) for the shrike-vireos I have found that in the pterylosis the dorsal tract on the lower back is divided, the arms being broad at the ends, and separated from the narrowed line that continues onto the caudal area. This is completely different from the usual rhomboid in the vireos, and may indicate that the family eventu- ally should be removed from the vicinity of the Vireonidae. The family characters of the peppershrikes, likewise outlined by Pycraft in the reference given above, are easily apparent on examina- tion of the skeleton. The family Callaeidae has been separated by Stonor (1942, pp. 1-18) on the weakened keel of the sternum, the great development of the lower limb coupled with reduced powers of flight, and the presence of a mouth wattle, for three peculiar genera, Callaeus, Heterolocha, and Philesturnus of New Zealand. Continuing discussion relative to the group of families to be placed in elevated position at the end of the list has led to publication of several useful studies and interesting statements. Beecher (1953, pp. 270-333) from examination of the musculature of the jaw, aided by other anatomical features, has proposed two major divisions of the suborder of the song birds, within which he has diagramed radiating lines of family and subfamily relationship. While he shows a variety of connections that in many cases vary widely from ideas current at present, he places the crow group in the assemblage with simpler muscle development in the area of the jaw, in contrast to those of higher status with a more complicated arrangement. Tordoff (1954a, 1954b) in a study of the skull, particularly the palatal structure, of species allied to the Fringillidae, has proposed the NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 2I union of part of the honeycreepers and the wood warblers in one family, the tanagers, with part of the coerebine assemblage with some of the fringillids in the Fringillidae, and removal of the cardueline finches to the Ploceidae, placing that family at the end of his list. His detailed studies afford much valuable information. I agree with him that shifting of certain genera to families in which they are not classi- fied at present will lead to better alignment, but I am not prepared from present information to completely dismember the Coerebidae without further study. Coereba, for example, has a stomach peculiar in its small size; Diglossa differs in the form of the bill, in which the gonys is extended posteriorly behind the level of the nostril, so that it differs from all other oscinine species, to cite only two easily seen characters. Mayr and Greenway (1956, pp. 2-5, 8-9) discuss problems of sequence in some detail and cite the approval of a committee appointed at the International Ornithological Congress held in Basel in 1954 to allocation of the Corvidae at the higher end of the list, as has been long customary among most ornithologists of Europe. In further consideration of these matters, I published a note on the humerus of the Corvidae (Wetmore, 1957, pp. 207-209), which called attention particularly to the proximal end of the bone, where the pneumatic fossa in Corvus, for example, has a form not only generally similar to that of the New World flycatchers and their allies, which are recog- nized as low down in the linear classification, but also to the wood- peckers, the Coraciiformes, and the trogons. There is transition from this simpler form to the style found in such groups as the Icteridae, Thraupidae, and Fringillidae, where the fossa is enlarged, and is more complex, as it is partly divided by a bladelike process pro- jecting from the internal tuberosity. (In the paper cited I neglected to refer to an earlier study by James T. Ashley [1941] on the humerus of the Corvidae, which outlined the same differences, and on which Ashley considered the crow group to have more primitive status. ) Amadon (1957) recently has outlined the three major groups of oscinine families, with the conclusion that the one most highly ad- vanced includes the 9-primaried New World groups, while the section containing the crows is placed low at the beginning. There is general agreement with this in the classification outlined by Delacour and Vaurie (1957). Storer (1959) in a clearly stated summary of these recent contri- butions, in which he includes a more recent statement by Mayr (1958), writes that in a classification for a text on the biology of birds now in preparation he has placed the 9-primaried groups in the highest place, 22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 and indicates that this is the procedure that is gaining in acceptance in parts of the world other than America. The former family Melithreptidae becomes the family Melipha- gidae, since the name of the type genus is now accepted as Meliphaga Lewin, 1808. In a similar way the family Compsothlypidae for the wood warblers becomes the family Parulidae, since the former Compsothlypis Cabanis, 1851, is replaced by the older Parula Bonaparte, described in 1838. The order of arrangement in the Passeriformes as said above is in part necessarily arbitrary, through the easily perceptible and often- remarked fact that we are required to list the groups in linear order in a two-dimensional alignment when actually they stand in three-dimen- sional relationship to one another. A further element that may be re- garded almost as a fourth dimension is found in some of the extinct groups known only as fossils that have no close relatives alive today. The sequence in the following pages is the one that best represents my present understanding, based on personal studies over a period of more than 50 years. I continue to place the Fringillidae at the end of the list, because of my feeling that this group is the modern expres- sion of a main core or stem that through the earlier Tertiary periods has given rise to more specialized assemblages that we now recognize as distinct families. Further specialization is apparent in some parts of the existing fringilline assemblage that, if undisturbed, may lead to further differentiation, should these variants be able to persist for the necessary millenniums in our rapidly changing world. Adjacent to the Fringillidae I place the other groups that obviously are closely allied to them. Attempts to arrange the avian families with the Cor- vidae and their allies in the terminal position, because of supposed more advanced development of the brain, appear to me quite uncertain, particularly in view of our decidedly limited information in this field. Should this idea be coupled with belief in superior mental reactions in the corvine assemblage, I would consider this more an anthropo- morphic interpretation than one supported by scientific fact. In the formation of group names the suffixes -idae and -inae for families and subfamilies are accepted rather universally so that they do not require examination. In view of the limited number of species covered in ornithology I see no point in the introduction of tribes as another category between the subfamily and the genus. This may be useful to entomologists with their tens of thousands of species but seems unnecessary and cumbersome with birds. In some of the more comprehensive avian genera there are groups of species more closely NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 23 allied to one another than to their fellows, but the taxonomist may discuss these at need as groups without imposing another burden on a classification that now is highly divided. For the group names above the family level, I believe it preferable to use suffixes that allow im- mediate identification of the rank, coupled with a stem that, like the family name, is based on a current generic term. Where ordinal and subordinal names are both formed as Latin plurals there is possibility of confusion. SYSTEMATIC LIST Fossil groups in brackets Class Aves, Birds. [Subclass Archaeornithes, Ancestral Birds (fossil). ] [Order Archaeopterygiformes, Archaeopteryx, Archaeornis (fossil) .] [Family Archaeopterygidae, Archaeopteryx, Archaeor- mis (fossil).] Subclass Neornithes, True Birds. [Superorder Odontognathae, New World Toothed Birds (fos- sil).] [Order Hesperornithiformes, Hesperornithes (fossil).] [Family Hesperornithidae, Hesperornis (fossil) .] [Enaliornithidae,! Enaliornis (fossil) .] [Baptornithidae, Baptornis (fossil).] [Superorder Ichthyornithes, Ichthyornis and Allies (fossil).] [Order Ichthyornithiformes, Ichthyornithes (fossil). ] [Family Ichthyornithidae, Ichthyornis (fossil).] [Apatornithidae, Apatornis (fossil).] Superorder Impennes, Penguins. Order Sphenisciformes, Penguins. Family Spheniscidae, Penguins. Superorder Neognathae, Typical Birds. Order Struthioniformes, Ostriches. [Family Eleutherornithidae, Eleutherornis (fossil).] Struthionidae, Ostriches. Order Rheiformes, Rheas. Family Rheidae, Rheas. Order Casuariiformes, Cassowaries, Emus. Family Casuariidae, Cassowaries. Dromiceidae, Emus. [ Dromornithidae, Dromornis (fossil).] 1 Position provisional. 24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 [Order Aepyornithiformes, Elephantbirds (fossil and sub- fossil) .] [Family Aepyornithidae, Aepyornis (fossil and sub- fossil).] [Order Dinornithiformes, Moas (fossil and subfossil).] [Family Dinornithidae, Dinormis (fossil and subfos- sil).] [ Anomalopterygidae, Anomalopteryx, Emeus, and Allies (fossil and subfossil).] Order Apterygiformes, Kiwis. Family Apterygidae, Kiwis. Order Tinamiformes, Tinamous. Family Tinamidae, Tinamous. Order Gaviiformes, Loons. Family Gaviidae, Loons. Order Podicipediformes, Grebes. Family Podicipedidae, Grebes. Order Procellariiformes, Albatrosses, Shearwaters, Petrels, and Allies. Family Diomedeidae, Albatrosses. Procellariidae, Shearwaters, Fulmars. Hydrobatidae, Storm Petrels. Pelecanoididae, Diving Petrels. Order Pelecaniformes, Tropicbirds, Pelicans, Frigate-birds, and Allies. [Suborder Odontopteryges, Odontopteryx, and Allies (fos- sil).] [Family Odontopterygidae, Odontopteryx (fossil).] | Pseudodontornithidae, Pseudodontornis, Os- teodontornis (fossil) .] Suborder Phaethontes, Tropicbirds. Family Phaéthontidae, Tropicbirds. Suborder Pelecani, Pelicans, Boobies, Cormorants, Snake- birds. Superfamily Pelecanoidea, Pelicans and Allies. Family Pelecanidae, Pelicans. [Cyphornithidae, Cyphornis, Palaeochendides (fossil) .] Superfamily Suloidea, Boobies, Cormorants, and Allies. Family [Pelagornithidae, Pelagornis (fossil).] Sulidae, Boobies, Gannets. NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 25 [Elopterygidae, Elopteryx, Eostega, Actiornis (fossil) .] Phalacrocoracidae, Cormorants. Anhingidae, Snake-birds. Suborder Fregatae, Frigate-birds. Family Fregatidae, Frigate-birds. [Suborder Cladornithes, Cladornis and Cruschedula (fos- sil) .] [Family Cladornithidae, Cladornis, Cruschedula (fos- sil).] Order Ciconiiformes, Herons, Storks, and Allies. Suborder Ardeae, Herons, Bitterns. Family Ardeidae, Herons, Bitterns. Cochleariidae, Boatbilled Herons. Suborder Balaenicipites, Whale-headed Storks. Family Balaenicipitidae, Whale-headed Storks. Suborder Ciconiae, Storks, Ibises, Spoonbills. Superfamily Scopoidea, Hammerheads. Family Scopidae, Hammerheads. Superfamily Ciconioidea, Storks. Family Ciconiidae, Storks, Jabirus. Superfamily Threskiornithoidea, Ibises. Family Threskiornithidae, Ibises, Spoonbills. Suborder Phoenicopteri, Flamingos. [Family Agnopteridae, Agnopterus (fossil).] [Scaniornithidae, Scaniornis, Parascaniornis (fossil).] Phoenicopteridae, Flamingos. [Palaelodidae, Palaelodus, Megapaloelodus, Telmabates (fossil).] Order Anseriformes, Screamers, Ducks, Geese, Swans. Suborder Anhimae, Screamers. Family Anhimidae, Screamers. Suborder Anseres, Ducks, Geese, Swans. [Family Paranyrocidae, Paranyroca (fossil).] Anatidae, Ducks, Geese, Swans. Order Falconiformes, Vultures, Hawks, Falcons. Suborder Cathartae, New World Vultures. [Superfamily Neocathartoidea, Neocathartes (fossil).] [Family Neocathartidae, Neocathartes (fossil). ] 26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 Superfamily Cathartoidea, New World Vultures. Family Cathartidae, New World Vultures. [Teratornithidae, Teratornis, Cathartornis (fos- sil).] Suborder Falcones, Secretarybirds, Hawks, Falcons. Superfamily Sagittarioidea, Secretarybirds. Family Sagittariidae, Secretarybirds. Superfamily Falconoidea, Hawks, Falcons, and Allies. Family Accipitridae, Hawks, Old World Vultures, Harriers. Pandionidae, Ospreys. Falconidae, Falcons, Caracaras. Order Galliformes, Megapodes, Curassows, Pheasants, Hoat- zins. Suborder Galli, Megapodes, Curassows, Grouse, Pheasants. Superfamily Cracoidea, Megapodes, Curassows. Family Megapodiidae, Megapodes. [Gallinuloididae, Gallinuloides (fossil).] Cracidae, Curassows, Guans, Chachalacas. Superfamily Phasianoidea, Grouse, Pheasants, Turkeys. Family Tetraonidae, Grouse. Phasianidae, Quails, Pheasants, Peacocks. Numididae, Guineafowl. Meleagrididae, Turkeys. Suborder Opisthocomi, Hoatzins. Family Opisthocomidae, Hoatzins. Order Gruiformes, Cranes, Rails, and Allies. Suborder Mesitornithides, Roatelos, Monias. Family Mesitornithidae, Roatelos, Monias. Suborder Turnices, Bustardquails, Hemipodes. Family Turnicidae, Bustardquails. Pedionomidae, Plainwanderers. Suborder Grues, Cranes, Limpkins, Trumpeters, Rails. Superfamily Gruoidea, Cranes, Limpkins, Trumpeters. [Family Geranoididae, Geranoides (fossil).] [Eogruidae, Eogrus (fossil).] Gruidae, Cranes. Aramidae, Limpkins. Psophiidae, Trumpeters. NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 27 Superfamily Ralloidea, Rails. [Family Orthocnemidae,? Orthocnemus, Elaphrocne- mus (fossil). ] Rallidae, Rails, Coots, Gallinules. Suborder Heliornithes, Sungrebes. Family Heliornithidae, Sungrebes. Suborder Rhynocheti, Kagus. Family Rhynochetidae, Kagus. Suborder Eurypygae, Sunbitterns. Family Eurypygidae, Sunbitterns. Suborder Cariamae, Seriemas and Allies. [Superfamily Phororhacoidea, Phororhacos and Allies (fossil).] [Family Phororhacidae, Phororhacos and Allies (fos- sil).] [Psilopteridae, Psilopterus and Allies (fos- sil).] [Brontornithidae, Brontornis, Liornis, and Allies (fossil).] [ Opisthodactylidae, Opisthodactylus (fossil) .] [Cunampaiidae, Cunampaia (fossil).] Superfamily Cariamoidea, Seriemas and Allies. [Family Bathornithidae, Bathornis (fossil).] [Hermosiornithidae, Hermosiornis, Procari- ama (fossil).] Cariamidae, Seriemas. Suborder Otides, Bustards. Family Otididae, Bustards. [Order Diatrymiformes, Diatryma, Omorhamphus, and Allies (fossil).] [Family Diatrymidae, Diatryma (fossil).] [Gastornithidae, Gastornis, Remiornis (fos- sil).] Order Charadriiformes, Shore Birds, Gulls, Auks. Suborder Charadrii, Shore Birds. Superfamily Jacanoidea, Jacanas. Family Jacanidae, Jacanas. Superfamily Charadrioidea, Plovers, Sandpipers, and Al- lies. 2 Position provisional. 28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 [Family Rhegminornithidae, Rhegminornis (fossil).] Rostratulidae, Painted Snipe. Haematopodidae, Oystercatchers. Charadriidae, Plovers, Turnstones, Surfbirds. Scolopacidae, Snipe, Woodcock, Sandpipers. Recurvirostridae, Avocets, Stilts. [| Presbyornithidae, Presbyornis (fossil).] Phalaropodidae, Phalaropes. Superfamily Dromadoidea, Crabplovers. Family Dromadidae, Crabplovers. Superfamily Burhinoidea, Thick-knees. Family Burhinidae, Thick-knees. Superfamily Glareoloidea, Pratincoles, Coursers. Family Glareolidae, Pratincoles, Coursers. Superfamily Thinocoroidea, Seedsnipe. Family Thinocoridae, Seedsnipe. Superfamily Chionidoidea, Sheathbills. Family Chionididae, Sheathbills. Suborder Lari, Gulls, Terns, Skimmers. Family Stercorariidae, Skuas, Jaegers. Laridae, Gulls, Terns. Rynchopidae, Skimmers. Suborder Alcae, Auks. Family Alcidae, Auks, Auklets, Murres. Order Columbiformes, Sandgrouse, Pigeons, Doves. Suborder Pterocletes, Sandgrouse. Family Pteroclidae, Sandgrouse. Suborder Columbae, Pigeons, Doves. Family Raphidae, Dodos, Solitaires. Columbidae, Pigeons, Doves. Order Psittaciformes, Lories, Parrots, Macaws. Family Psittacidae, Lories, Parrots, Macaws. Order Cuculiformes, Plantain-eaters, Cuckoos. Suborder Musophagi, Plantain-eaters. Family Musophagidae, Plantain-eaters, Touracos. Suborder Cuculi, Cuckoos, Roadrunners, Anis. Family Cuculidae, Cuckoos, Roadrunners, Anis. Order Strigiformes, Owls. [Family Protostrigidae, Protostrix (fossil).] Tytonidae, Barn Owls. Strigidae, Typical Owls. NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 29 Order Caprimulgiformes, Oilbirds, Goatsuckers. Suborder Steatornithes, Oilbirds, Family Steatornithidae, Oilbirds. Suborder Caprimulgi, Frogmouths, Goatsuckers. Family Podargidae, Frogmouths. Nyctibiidae, Potoos. Aegothelidae, Owlet-frogmouths. Caprimulgidae, Goatsuckers, Order Apodiformes, Swifts, Hummingbirds. Suborder Apodi, Swifts. [Family Aegialornithidae,? Aegialornis (fossil).] Apodidae, Swifts. Hemiprocnidae, Crested Swifts. Suborder Trochili, Hummingbirds. Family Trochilidae, Hummingbirds. Order Coliiformes, Colies. Family Coliidae, Colies. Order Trogoniformes, Trogons. Family Trogonidae, Trogons. Order Coraciiformes, Kingfishers, Bee-eaters, Rollers, Horn- bills. Suborder Alcedines, Kingfishers, Todies, Motmots. Superfamily Alcedinoidea, Kingfishers. Family Alcedinidae, Kingfishers. Superfamily Todoidea, Todies. Family Todidae, Todies. Superfamily Momotoidea, Motmots. Family Momotidae, Motmots. Suborder Meropes, Bee-eaters. Family Meropidae, Bee-eaters. Suborder Coracii, Rollers, Hoopoes. Family Coraciidae, Rollers. Brachypteraciidae, Groundrollers. Leptosomatidae, Cuckoo-rollers. Upupidae, Hoopoes. Phoeniculidae, Woodhoopoes. Suborder Bucerotes, Hornbills. Family Bucerotidae, Hornbills. 3 Position provisional. 30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 Order Piciformes, Jacamars, Barbets, Toucans, Woodpeckers. Suborder Galbulae, Jacamars, Barbets, Toucans. Superfamily Galbuloidea, Jacamars, Puffbirds. Family Galbulidae, Jacamars. Bucconidae, Puffbirds. Superfamily Capitonoidea, Barbets, Honeyguides. Family Capitonidae, Barbets. Indicatoridae, Honeyguides. Superfamily Ramphastoidea, Toucans. Family Ramphastidae, Toucans. Suborder Pici, Woodpeckers. Family Picidae, Woodpeckers, Piculets. Order Passeriformes, Perching Birds. Suborder Eurylaimi, Broadbills. Family Eurylaimidae, Broadbills. Suborder Tyranni, Ovenbirds, Tyrant Flycatchers, and Al- lies Superfamily Furnarioidea, Ovenbirds, Woodhewers, and Allies. Family Dendrocolaptidae, Woodhewers. Furnariidae, Ovenbirds. Formicariidae, Ant-thrushes. Conopophagidae, Antpipits. Rhinocryptidae, Tapaculos. Superfamily Tyrannoidea, Tyrant Flycatchers, Pittas, and Allies. Family Cotingidae, Cotingas. Pipridae, Manakins. Tyrannidae, Tyrant Flycatchers, Oxyruncidae, Sharpbills. Phytotomidae, Plantcutters. Pittidae, Pittas. Acanthisittidae, New Zealand Wrens. Philepittidae, Asities, False Sunbirds. Suborder Menurae, Lyrebirds. Family Menuridae, Lyrebirds. Atrichornithidae, Scrubbirds. Suborder Passeres, Songbirds. Family Alaudidae, Larks. [ Palaeospizidae, Palaeospiza (fossil) .] Hirundinidae, Swallows. Dicruridae, Drongos. No. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 31 Oriolidae, Old World Orioles. Corvidae, Crows, Magpies, Jays. Cracticidae, Bell Magpies, Australian Butcher- birds, Grallinidae, Magpie-larks. Ptilonorhynchidae, Bowerbirds. Paradisaeidae, Birds of Paradise. Paridae, Titmice. Sittidae, Nuthatches. Hyposittidae, Coralbilled Nuthatches. Certhiidae, Creepers. Paradoxornithidae, Parrotbills, Suthoras. Chamaeidae, Wrentits. Timaliidae, Babblers. Campephagidae, Cuckoo-shrikes. Pycnonotidae, Bulbuls. [Palaeoscinidae,* Palaeoscinis (fossil).] Chloropseidae, Leafbirds. Cinclidae, Dippers. Troglodytidae, Wrens. Mimidae, Thrashers, Mockingbirds. Turdidae, Thrushes. Zeledoniidae, Wrenthrushes. Sylviidae, Old World Warblers. Regulidae, Kinglets. Muscicapidae, Old World Flycatchers. Prunellidae, Accentors. Motacillidae, Wagtails, Pipits. Bombycillidae, Waxwings. Ptilogonatidae, Silky Flycatchers. Dulidae, Palmchats. Artamidae, Woodswallows. Vangidae, Vanga Shrikes. Laniidae, Shrikes. Prionopidae, Woodshrikes. Cyclarhidae, Peppershrikes. Vireolaniidae, Shrike-vireos. Callaeidae, Wattled Crows, Huias, Saddlebacks. Sturnidae, Starlings. Meliphagidae, Honey-eaters. 4 Allocation to this position is tentative. 32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 139 Nectariniidae, Sunbirds. Dicaeidae, Flowerpeckers. Zosteropidae, White-eyes. Vireonidae, Vireos. Coerebidae, Honeycreepers. Drepanididae, Hawaiian Honeycreepers. Parulidae, Wood Warblers. Ploceidae, Weaverbirds. Icteridae, Blackbirds, Troupials. Tersinidae, Swallowtanagers. Thraupidae, Tanagers. Catamblyrhynchidae, Plushcapped Finches. Fringillidae, Grosbeaks, Finches, Buntings. December 31, 1959. LITERATURE CITED AMADON, DEAN. 1950. Australian mud builders. Emu, vol. 50, pt. 2, Oct., pp. 123-127. 1957. Remarks on the classification of the perching birds [order Passeri- formes]. Proc. Zool. Soc. Calcutta, Mookerjee Mem. Vol., May 1, pp. 259-268. ASHLEY, JAMES F. 1941. A study of the structure of the humerus in the Corvidae. Condor, vol. 43, No. 4, July 15, pp. 184-195, figs. 50-56. BEECHER, WILLIAM J. 1953. A phylogeny of the Oscines. Auk, vol. 70, No. 3, July, pp. 270-333, 18 figs. Bock, WALTER J. 1956. A generic review of the family Ardeidae (Aves). Amer. Mus. Nov., No. 1779, July 27, pp. 1-49. 1958. A generic review of the plovers (Charadriinae, Aves). Bull. Mus. Comp. Zool., vol. 118, No. 2, March, pp. 27-97, 6 figs. BONAPARTE, PRINCE. 1855. (Remarks, in proceedings of Académie des Sciences de Paris.) Rev. et Mag. Zool., ser. 2, vol. 7, Oct., pp. 482-485. CaBANIS, J. 1847. Ornithologische Notizen. II. Arch. Naturg., vol. 1, pp. 308-352. CATESBY, MARK. 1731. The natural history of Carolina, Florida and the Bahama Islands: Containing the figures of birds, beasts, fishes, serpents, insects and plants: Particularly the forest-trees, shrubs, and other plants, not hitherto described, or very incorrectly figured by authors. Vol. 1, Pp. i-xii, I-100, 100 pls. London. CotTAM, Patricia A. 1957. The pelecaniform characters of the skeleton of the shoe-bill stork, Balaeniceps rex. Bull. Brit. Mus. (Nat. Hist.), Zool., vol. 5, No. 3, July, pp. 1-71, 1 pl., 4 figs. NO. II CLASSIFICATION, BIRDS OF THE WORLD—WETMORE 33 DaAMEs, W. 1897. Uber Brustbein, Schulter- und Beckengiirtel der Archaeopteryx. Sitzb, preuss. Akad. Wiss. Berlin, vol. 38, Aug. 9, pp. 818-834. DE BEER, SiR GAVIN. 1954. Archaeopteryx lithographica, a study based upon the British Museum specimen. Pp. i-xii, 1-68, 16 pls., 9 text figs. British Museum (Natural History), London. DELACOUR, JEAN. 1946a. Les timeliines. Ois. Rev. Frang. Orn., n.s., vol. 16, pp. 7-36. 1946b. Notes on the taxonomy of the birds of Malaysia. Zoologica, vol. 31, pt. 1, Apr. 29, pp. 1-8. DELACcoUR, JEAN, and AMADON, DEAN. 1949. The relationships of Hypocolius. Ibis, vol. 91, No. 3, July, pp. 427- 429, pls. 19-20. DELACOUR, JEAN, and VAURIE, CHARLES. 1957. A classification of the Oscines (Aves). Los Angeles County Mus. Contr. Sci., No. 16, Oct. 31, pp. 1-6. DRESSER, HENRY EELES. 1893. A monograph of the Coraciidae, or family of the rollers. Pp. i-xx, I-111, 27 pls. London. Forpes, WILLIAM ALEXANDER. 1880. On the anatomy of Leptosoma discolor. Proc. Zool. Soc. London, 1880, pp. 464-475, 5 figs. GLoGER, CONSTANTIN LAMBERT. 1854. In J. Reinhardt, Bemerkungen zur Ornithologie Gronlands. 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